Rhodora

Hodova

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by

ALBION REED HODGDON, Editor-in-Chief
ROLLA MILTON TRYON )
RADCLIFFE BARNES PIKE
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
ALFRED LINN BOGLE

f Associate Editors

RICHARD EDWIN WEAVER S
Vol. 76 March, 1974 No. 805
CONTENTS:

The Phytogeography and Systematics of Xanthisma tex-

anum D. C. (Asteraceae); Proper Usage of Intraspecific
Categories

John C. Semple EMEND 1
A White-Flowered Form of Utricularia purpurea

C. Barre Hellquist eet 19
On the Scientific Name of the Longleaf Pine

BEE Ey Ward T R in 20
The Occurrence of Bald Cypress (Taxodium distichum (L.)

Richard) in Suffolk County, Long Island, New York

Edward J. Hehre and Peter M. Bolid Leeds: 25

The Nef England Botanical Club, Ine.
Botanical Museum. Oxford St.. Cambridge, Mass. 02188

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Rhodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 76 March, 1974 No. 805

THE PHYTOGEOGRAPHY AND SYSTEMATICS OF
XANTHISMA TEXANUM DC. (ASTERACEAE):
PROPER USAGE OF INFRASPECIFIC CATEGORIES?

JOHN C. SEMPLE?

The monotypic genus Xanthisma DC. (Asteraceae) in-
cludes sufficient variation in its single species X. texanum
DC. to illustrate the three levels of taxonomically recog-
nized infraspecific variation: subspecifie, varietal and for-
mal. Proper usage of the three taxonomic ranks requires
consideration of morphological and distributiona] data ac-
cording to a consensus of international opinion. Although
much has been published on the B-chromosome of the spe-
cies (see Semple, 1972a), no modern systematic treatment
of X. texanum has been published. Examination of some
five hundred herbarium collections and experimental data
indicate that a revision of Gray's 1852 treatment, which
was based on only a few specimens, is necessary.

A detailed diseussion of the distribution of the species
as a whole and each of the infraspecific taxa is warranted
for two reasons: first, to document the allopatric condition
of the two subspecies, and second, to show that the distri-

"The work was part of an NDEA Title IV Fellowship sponsored
doctoral dissertation submitted to Washington University, St. Louis,
Mo. Aug. 1972: Dr. Walter H. Lewis advisor.

2Current Address: Dept. of Biology, University of Missouri-St.
Louis, St. Louis, Mo. 63121.

2 Rhodora [Vol. 76

bution of the species is not predictable from the general-
ized vegetation distributions given by Gould (1962). The
Edwards Plateau and its eroded portions are basic con-
siderations in both cases. A taxonomic treatment consider-
ing all aspects of the problem is given at the end of the
discussion of distribution and morphological variation.

All populations of a species ought to be interfertile to
some degree. The validity of treating all the morphological
forms of Xanthisma as a single species was tested by hy-
bridization experiments involving greenhouse plants grown
from fruiting material collected at wild populations. No
barriers to fertilization were found and normal chromosome
homologue pairing in meiotic prophase was observed in F,
generation hybrids, which also set viable seed. Cross pol-
lination was accomplished following the procedure of Smith
& Parker (1971) and was facilitated by self-incompatibil-
ity. Selfing could be induced, but yielded progeny of low
fitness. Clearly, Gray's combining all taxa of Xanthisma
into a single species is acceptable.

DISTRIBUTION OF THE SPECIES

Xanthisma texanum grows in a portion of the central
southwest of the United States: Texas, Oklahoma and east-
ern New Mexico. The known distribution of the species is
illustrated in Figure 1 and is based on herbarium collections
from GH, KSU, MO, NY, OKLA, SMU, TEX, TTC, US, WIS, and
Arizona State University-Tempe.’ Collections of subsp.
drummondii (T. & G.) Semple and subsp. texanum are rep-
resented by dots and stars respectively; var. texanum and
var. orientalis Semple of subsp. texanum are represented
by encircled and unenclosed solid stars; putative hybrids
between the two subspecies are represented by open stars.
Treatment of Gray's (1852) varieties as subspecies, a term
he did not use (Fernald, 1940), is based upon involucral
bract morphology and the nearly allopatric distribution of
the two taxa, while the distribution and bract morphology

"The assistance of Frederick Utech and Timothy Love at the last
two herbaria, respectively, is gratefully acknowledged.

1974] Xanthisma — Semple 3

Distribution of
Xanthisma texanum DC
ye subspeci(ic hybrids
subsp drummondii
@® subsp texanum var texanum

* subsp texanum var. orientalis

atc

Figure 1. The distribution of Xanthisma texanum DC. Locations
of collections have been determined from herbarium collections from
eleven herbaria cited in the text. A key to the symbols used is given.
Collections are plotted on a county outline map of Texas and Okla-
homa.

4 Rhodora [Vol. 76

variation within subsp. texanum is properly treated at the
varietal level. Variation in environmental factors accounts
for the species distribution.

Climatic factors determine the extent of the species’
range in the east, northwest and south. Too much rain in
the east and too little in the north and west are the ap-
parent limiting factors, while increasingly tropical condi-
tions become limiting to the south (Visher, 1954). Local
variation in geological factors accounts for the distribution
within the range and secondarily along the margins of the
range. The western margin of the eastern deciduous forest
(Bray, 1906; Tharp, 1926; Braun, 1950; and Gould, 1962)
forms the eastern margin of the range of X. texanum,
which is a member of the ecotone flora sensu latu between
the forests and grasslands. Reasons for the present loca-
tion of the forest-prairie ecotone are not appropriate for
discussion in this paper.

A detailed treatment of geological influences on the spe-
cies’ range is given by Semple (1972b) and summarized
for each taxon under each appropriate heading. The Rio
Grande Valley supports a noticeably distinct vegetation
community (Foscue, 1932). Xanthisma texanum is excluded
from the community, which apparently forms a barrier to
southward expansion of the range of X. texanum, since no
collections have yet been made in Mexico. Either the Rio
Grand Valley community is an impassible barrier or no
inhabitable locations exist in Mexico for X. texanum.

DIAGNOSTIC MORPHOLOGY AND DISTRIBUTION
OF THE TWO SUBSPECIES

A subspecies is characterized by all members exhibiting
a particular morphology distinct from other individuals in
the species and by the allopatric distribution of these mem-
bers from the rest of the species (Du Reiz, 1930; Clausen,
1941; van Steenis, 1957; Valentine & Lóve, 1958; and
Ehrendorfer, 1968). Those morphologically intermediate
individuals between the subspecies and the rest of the spe-

1974] Xanthisma — Semple 5

$ €

Figure 2. Geographie distribution of involucral bract variation of
Xanthisma texanum DC. Bracts from plants from 68 populations are
illustrated in outline on a map of Texas and Oklahoma. Bracts 1-48
are from individuals of subsp. drummondii (T. & G.) Semple; bracts
52-68 are from individuals of subsp. texanwm; and bracts 49-51 are
from putative hybrids between the two subspecies.

6 Rhodora [Vol. 76

cies only occur in populations geographically located be-
tween the subspecies and the remainder of the species.
Within X. texanum all members of subsp. drummondit
(T. & G.) Semple are distinguished by involucral bracts
that are lanceolate and taper to an acute apex. Subspecies
texanum includes other members of the species and is char-
acterized by involucral bracts that are depressed-ovate in
shape and shorter than those of subsp. drummondii.
Diagrammatic representations (Figure 2) of involucral
bracts typical of those occurring in the middle of the series
forming the involucre are superimposed on an outline map
of Texas and Oklahoma and were taken from plants occur-
ring in wild populations at the locations indicated. Bracts
1-48 show the similarity that occurs within subsp. drum-
mondii (see Figure 3.C for a detailed illustration of a typi-
cal involucral bract). Bracts 52-68 illustrate the range of
variation in involucral bract shape found in subsp. texanum
(Figures 3.A and 3.B), Bracts 49-51 illustrate involucral
bracts from putative hybrids between the two subspecies
and are the only bracts that are intermediate in shape.
The allopatric distribution of the two subspecies is due
to the presence of the Edwards Plateau and the Blacklands
Prairies between the two taxa. Geologically and topo-
graphically the Edwards Plateau is broken up into five
regions (Fenneman, 1931; Oetking, 1959; and Raisz, 1957),
although Gould (1962) included all the varied vegetation
types within a single vegetation region. With a few excep-
tions, in part due to man's influences, subsp. drummondii
does not grow in areas overlying uneroded portions of the
Edwards Plateau, since the upper strata limit the amount
of water available to the shallow rooted plants. Depending
upon the extent of erosion and the nature of the underlying
bedrock, subsp. drwmmondii grows in varying degrees of
commonness in the areas of the eastern portion of the
plateau (colloquial sense) in the following manner: com-
mon at the lower elevations of the Central Mining District,
common throughout the Lampasas Cut Plains and rare on
the more northeastern Comanche Plateau. Both subspecies

1974] Xanthisma — Semple 7

are absent in the southeastern portion of the Edwards
Plateau (the Texas Hill Country). Thus differences in ge-
ology account for the different numbers of collections in this
well botanized area and also explain the means by which
the two subspecies are isolated north and west of San
Antonio, Texas.

The presence of the Blacklands Prairies isolates the two
subspecies north and east of San Antonio. The prairies,
which run from San Antonio to Dallas and further north-
eastward, are characterized by dark, high clay content soils
(Sellards et al., 1966) and are dry and rock hard in the
summer. Typically, Xanthisma occurs on sandy, light
brown or reddish soils. Field work in the central Texas
region showed that X. texanum is very rare throughout
the entire length of the prairies, except for a few individu-
als surviving on disturbed or introduced soils along high-
ways and river banks. Clearly, Gray's varieties are exem-
plary of what are today treated as subspecies.

A narrow zone of isolated subsp. drummondii populations
occurs in Caldwell, Gonzales and Wilson Counties, but the
small populations only thrive in favorably moist years.
Interestingly, Heterotheca pilosa (Nutt.) Shinners and
Bradburia hirtella T. & G. are common in these counties
and further east, the former being the most prevalent yel-
low flowered member of the Astereae. Similarity in super-
ficial appearance of these two species with X. texanum
undoubtedly explains confusion about the distribution of
X. texanum in this critical region between the two sub-
Species.

DIAGNOSTIC MORPHOLOGY AND DISTRIBUTION OF THE
TWO VARIETIES OF SUBSP. TEXANUM

A variety is characterized by all members of a population
exhibiting a particular morphology distinct from other in-
dividuals in the species. The distribution of these popula-
tions is sympatric with populations whose members are not
within the same variety, and also many populations of

8 Rhodora [Vol. 76

morphological intermediates exist (Du Reiz, 1930; Clausen,
1941; van Steenis, 1957; Valentine & Lóve, 1958; and
Ehrendorfer, 1968). By this definition varieties can be
recognized within a species, which does not have subspecies.
Furthermore, variation that is mot known to be confined
to an isolated group of populations can not be given sub-
specific status. Van Steenis (loc. cit.) described varietal
level variation as being continuous with other varieties,
although the continuum would have pronounced modes.
The variation between subspecies can be described as dis-
continuous, except for the few hybrids. Figure 1 illustrates
the sympatric condition of the two varieties of subsp.
texanum: var. texanum with obtuse involucral bract apices
(Figure 3.A) and var. orientale Semple with cuspidate
involucral bract apices (Figure 3.B).

FORMAL LEVEL VARIATION IN VAR. ORIENTALIS

Formal level morphological variation is generally re-
garded as some phenotype possessed by only some members
of a population, whereas the subspecific and varietal level
phenotypes are possessed by all members of a population.
Those individuals in populations of subsp. texanum var.
orientale which exhibit red pigmentation along the lateral
margins of the involucral bracts forming the middle and
inner series of the involucre have been designated as forma
rubrum Semple (Figure 3.B). Such individuals are rare,
but do occur in populations throughout the range of var.
orientalis.

GENETIC BASIS OF DIAGNOSTIC CHARACTERISTICS

Experiments were conducted to show that all diagnostic
characters have a genetic basis. Size but not general shape
of the involucral bracts was found to be variable. Lack of
sufficient light, available water, amount of soil or a high
degree of crowding all resulted in smaller bracts being
formed than those bracts formed by the individual when
sufficient quantities of soil, light and water were provided.

1974] Xanthisma — Semple 9

The maximum size was determined by heredity and not the
environment. Therefore size should be considered second-
arily to shape in aetermining the proper identity of a par-
ticular collection.

The red pigment of f. rubrum was found to be passed
from one generation to the next in var. orientale, but was
never seen in var. texanum or in subsp. drummondii. Al-
though the pigment is not always present in detectable
quantities, its importance warrants some taxonomic status,
since Xanthisma has always been previously described as
monochromous. However, in some experimental plants
heads were occasionally produced which manifested the
red pigment in the distal portion of the ray straps on the
adaxial surface and on the lobes of the disc florets, but only
on the last day or two of disc anthesis. Most individuals
produced enough pigment to give the rays an orange hue,
but one plant produced rays that were deep rust red. Such
variation in intensity of color does not occur on bract mar-
£ins. No documentation of bicolor rays in wild plants
exists.

HABIT AS AN EXAMPLE OF VARIATION NOT WARRANTING
TAXONOMIC STATUS

Despite the great range of variation in habit occurring
in the species, no one form was found to warrant taxonomic
recognition. Within a single population long-stemmed and
short-stemmed individuals were found in nature. The dis-
tribution patterns of the short-stemmed plants (Figure
4.C) and the stemless plants (Figure 4.D) suggest that
these forms are ecotypes, since they have a genetic basis.
The extremes of short-stemmedness and stemlessness only
occur in populations of var. orientale and western popula-
tions of subsp. drummondii, respectively. The existence of
à full range of intermediates between the extremes and the
long-stemmed forms (Figures 4.B, 4.F and 4.G) in most
populations makes taxonomic recognition unjustified.

[Vol. 76

Rhodora

10

$a!

npuowwnjp "esans

Sijejuaio “ea WAVEIA) "fSqns

whuera) “eA = whueray ULL

1974] Xanthisma — Semple 11

A COMMENT ON CHEMICAL STUDIES

The taxonomic treatment that follows is based on mor-
phology and distribution. Two dimensional paper chroma-
tograms were run on leaf and flower extracts of specimens
of each taxon. In general, subspecies texanum had fewer
compounds than subspecies drummondii, and var. orientale
has fewer compounds than var. texanum. In var. orientale
individuals of forma rubrum had one compound, an uniden-
tified anthocyanin (Semple, 1972b), not found in other
individuals of the species. The full significance of the
chemical studies can only be known upon complete identifi-
cation of each compound. The preliminary results show
that chemical data support the taxonomy. The chemical
studies will have meaning in work on determining the evo-
lution of the taxa described.

TAXONOMIC TREATMENT

The most recently published treatment of the genus Xan-
thisma is in Correll & Johnston (1970). The following
detailed description of the genus is more complete and ac-

Figure 3. Involucral Bracts of Infraspecific Taxa. Each bract
shown. is typical of those in the middle of the series forming the
involucre: (A) subsp. texanum var. texanum; (B) subsp. texanum
var. orientale Semple (dashed lines indicate the extent of red pigment
on the margins of bracts of f. rubrum Semple); (C) subsp. drum-
mondii (T. & G.) Semple.

Figure 4. Habit Variation of Xanthisma texanum DC. The base-
line under each silhouette equals one decimeter. Each form shown il-
lustrates the habits of experimental plants and represents naturally
occurring forms. Plants were grown under similar conditions from
fruiting material collected at wild populations. All plants were from
different populations, except (B) and (C). See the text for a full
discussion.

Figure 5. Heterocarpic Fruit of Xanthisma texanum DC. Ray and
disc florets produce different kinds of fruit: (A), (B) and (C) are
radial, tangential and diagrammatic cross-sectional views of a ray
fruit; (D) and (E) are radial and diagrammatic cross-sectional
views of a central disc fruit. Peripheral disc fruit are larger and
angular ovate in cross-section.

12 Rhodora [Vol. 76

curate in light of the findings of my investigations. The
previously unreported heterocarpic fruit condition is also
described (Figure 5). A complete list of specimens exam-
ined in the study is given in Semple (1972b) and only rep-
resentative specimens are cited here.

Xanthisma DC., Prodr. 5:94-95. 1836.
Centauridium T. & G., Fl. N. Amer. 2:246. 1842.

Taprooted annuals, rarely biennial and flowering the
second season; branches ascending, few to many. Leaves
alternate, the upper sessile, the lower terminally serrate;
basal leaves petioled, lobed or pinnatifid to bipinnatifid.
Heads solitary; involuere turbinate to hemispheric or cam-
panulate; involucral bracts broad in 3-4 series, imbricate,
stiff, the margins pale; ray florets pistillate, fertile, yellow;
dise florets perfect, fertile, yellow. Fruit heterocarpic with
pappus bristly and margins finely serrate; both fruit kinds
with fine ascending white hairs, straw-colored when mature
and darkening with age. Receptacle slightly convex, with
subulate chaff reticulately dispersed between the fruits.

The genus is monotypic and is endemic to the central
southwestern areas of the United States.

Xanthisma texanum DC., Prod, 5:95. 1836.
Lectotype: Texas; Bexar County, in woods near Medina,
May 1828, Berlandier 2039d (G!). Isolectotypes; G! (4),
GH! (1), NY! (1), and us! (2).

Herbs 2-7 (rarely more) dm tall, branching near the
middle to ground level from a main stem 0.1-3 dm long, or
rarely acaulescent. Upper leaves 1-2 cm long, linear, entire;
basal leaves 5-8 cm long, ovate to obovate in outline, inter-
grading above. Heads solitary, not crowded; involucre
5-10 mm high, involucral bracts 1.5-4 mm broad, rarely
recurved, margins pale (the lateral margins very rarely
red), the central portion green, reticulate venation pro-
nounced, the inner-most bracts chartaceous, the outer-most
linear grading into the peduncle bracts; ray florets 10-30,
the straps 8-15 mm long, yellow adaxially, pale yellow

1974] Xanthisma — Semple 13

abaxially (rarely red-tinted abaxially) ; disc florets many
40-150, slightly ampliate to tapering, the lobes short. Ray
floret cypselas curved obpyramidal, triangulate in cross-
section, the pappus grading from short linear to long
basally broad scales; disc floret cypselas obovate radially
and narrowly-so tangentially, the central fruits rhombic
in ercss-section, the outer obtrullate, the pappus in two
whorls, the inner members ca. 6 mm long, linear, 8-15, the
outer members 3 mm long or less many.

The species is characterized by the following combination
of characters: heterocarpic fruit with a pappus of bristly
seales only, fruit pubescence of long white ascending hairs,
receptacle slightly convex with a persistent reticulate net-
work of subulate scales.

The lectotype of the species was chosen from five speci-
mens seen on microfilm only (1DC micro edition of Candolle
Prodromi Herbarium). Since all five specimens are similar
in quality, the specimen with the complete label giving col-
lection number, date and location, and floral color (fourth
from the left on the microfilm) is designated lectotype.
Torrey (1859) first noted the typographic error in the Pro-
dromus, which cited the type collection as Berlandier 2639
rather than Berlandier 2039.

Key to Infraspecific Taxa

I. Involucral bracts shorter than wide above the widest
part, 1.5-3.5 mm wide and 1-2 mm long above the widest
We eS, ur. spe Feats d yr g e raa subsp. texanum (1)
II. Involucral bracts rounded with little or no flare

1 mm from the apex, occasionally recurved.
i gu ck na ease esse es ewer’ var. texanum (2)
II. Involueral bracts obtusely pointed, the deltoid por-
tion 2-3 mm wide and 1.5-2 mm long above the
widest part. ............-...-- var. orientale (3)
III. Lateral margins of bracts whitish. ..........
JI NRORURUIOPNBA TO ROS .... f. orientale: (38)
III. Lateral margins of bracts red. f. rubrum (3b)

14 Rhodora [Vol. 76

I Involucral bracts longer than wide above the widest
part; deltoid portion 2.5-5 mm wide and 3.5-8 mm long
in bracts of the middle series. .. ssp. drummondii (4)

1. Xanthisma texanum spp. texanum

Xanthisma texanum var. berlandieri Gray, Smiths.
Contrib. Knowl. 3(5) : 98. 1852.

Xanthisma berlandieri (Gray) Small, Fl. SE. U.S. p.
1184. 1903.

This subspecies includes the type collection of X. texanum
and is distinguished by small involucral bracts with rounded
or obtusely pointed apices. Two varieties are recognized
within the subspecies.

2. Xanthisma texanum ssp, texanum var. texanum

The type variety of the species is distinguished by its
small apically rounded involucral bracts 1.5-2 mm wide,
appressed or rarely recurved; generally 30-35 bracts are
visible without removing the outer series. The variety is
endemic to the Nueces Plains of southern Texas from San
Antonio to the south and west. East of San Antonio this
variety forms hybrids with subspecies drummondit.

SOME REPRESENTATIVE SPECIMENS EXAMINED. TEXAS: Atascosa Co.,
Charlotte, 4 May 1919, Schulz 39 (Us). Bexar Co., San Antonio,
4 June 1948, Burr 21 (NY, TEX), in woods near the Medina River,
July 1829, Berlandier 629 (GH, MO, NY, US). Dimmit Co., Big Wells,
21 April 1945, Shinners 7,400 (NY, OKLA, SMU, TEX), in front of
cemetery, 6 May 1964, Turner 4,996 (TEX). Frio Co., Dilley, 6 mi.
W, 8 July 1970 Lewis 7,634 (Mo). La Salle Co., Artesia Wells, 3 mi.
W., 7 April 1963, Dickey 170 (SMU, TEX). Medina Co., Farm Rd.
1343, 4.4 mi S of U.S. Hwy. 90, 29 June 1972, Semple & Shea 720
(MO) ; Devine, 3 mi. SW, 1 May 1954, Tharp & Turner 3,439 (OKLA,
TEX). Zavala Co., Batesville, 6 mi. E, 6 May 1964, Turner 5,002
(TEX).

9. Xanthisma texanum ssp. texanum var. orientale Semple,
var. nov.
Involucri bracteae parvae manifeste depresse ovatae,
apice obtuse, cuspidatae, interdum marginibus lateralibus
rubris.

1974] Xanthisma — Semple 15

HOLOTYPE: Texas: Hidalgo Co.: 13.5 mi. N. of Edinburg on
U. S. Hwy. 183, 8 July 1970, W. Lewis 1642 (Mo). Iso-
types: four, to be distributed.

The involucral bracts are small and pronouncedly shield-
shaped, occasionally with red lateral margins. The apices
are obtusely cuspidate.

This variety is distinguished by the 2-3.5 mm wide in-
volucral bracts having obtuse, cuspidate apices. The short
stem habit is also frequently encountered. Hybridization
with var. texanum is common, but is unknown with ssp.
drummondii. The name is taken from its distribution in
the eastern half of the range of ssp. texanum. Two forms
are recognized.

3a. Xanthisma texanum var. orientale Semple f. orientale
The type form is the most common form of the variety.
SOME REPRESENTATIVE SPECIMENS EXAMINED. TEXAS: Aransas Co.,

Aransas Refuge, 29 Sept. 1944, Cory 45, 895 (Ny, TEX). Bee Co.,

Beeville, 21 June 1935, Drushel 9,941 (NY). Brooks Co., Hebbron-

ville, 15 mi. E, 25 Nov. 1962, Dohnke T (sMU). Calhoun Co., Port

Lavaca, 14 June 1953, Johnston 53.280.157 (OKLA, TEX). Goliad Co.,

Goliad, 15.9 mi. S, 9 July 1970, Lewis 7,655 (Mo). Hidalgo Co., Rio

Grande Valley, 8 Aug. 1942, Walker 70 (KSU, TEX). Jim Hogg Co.,

Hebbronville, 8 mi. E, 8 July 1970, Lewis 7,638 (MO). Jim Wells Co.,

Premont, 12 mi. N, 24 Nov. 1954, Johnston 542,144 (TEX). Kenedy

Co., King Ranch, Norias Div. 24 Nov. 1953 Johnston 53.280.154

(OKLA, TEX.) Kleberg Co., Kingville, 1940, Sinclair s.n. (TEX).

Refugio Co., 5 mi. S of county line on U. S. Hwy. 77A, 12 June

1971, Semple 581 (Mo). San Patricio Co., Aransas Pass, 24 May

1922, Schulz 868 (TEX, US). Near San Patricio, May 1834, Berlandier

2578 (GH, MO, NY). Webb Co., Mirando City, 31 Aug. 1936, Baird

s.n. (NY). Willacy Co. Yturria Station, 6 Aug. 1924, Runyon 664

(TEX, US).

3b. Xanthisma texanum var. orientale f. rubrum Semple,
forma nov. Involucri bracteae cum marginibus later-
alibus rubris.

HOLOTYPE: Texas: San Patricio Co., 0.5 mi. E. of Ingleside
on Texas Hwy. 361, 13 June 1971, Semple 602 (Mo).
Isotype: one to be distributed.

This form is distinguished from the type form by having
bracts with red lateral margins.

16 Rhodora [Vol. 76

REPRESENTATIVE SPECIMENS EXAMINED. TEXAS: Aransas Co.,
Aransas Refuge, 29 Sept. 1944, Cory 45,895 (Mo). Goliad Co.,
General Zaragoza State Park, 1 mi. S, 12 June 1971, Semple 579
(mo). Kenedy Co., Raymondville, 16 mi. N, 2 Dec. 1945, Cory
51,493 (us). Willacy Co., Port Mansfield, 0.5 mi. W of beach by
garbage dump, 9 July 1970, Lewis 7,644 (MO).

4. Xanthisma texanum DC. ssp. drummondii (T. & G.)
Semple, stat. nov.
Centauridium drummondii T. & G., Fl. N. Amer. 2: 462.
1842.
SYNTYPES: Texas, north of San Antonio, Riddell s.n. (NY) ;
Texas, exact locality unknown, Nov. 1835, Drummond
227 (GH!, NY!).
LECTOTYPE: Drummond 227 (GH). Isolectotypes: GH, NY.

Xanthisma texanum var. drummondii (T. & G.) Gray,
Smiths. Contrib. Knowl. 3(5) : 98. 1852.

Xanthisma drummondii (T. & G.) Hooker f., Curtis’
Bot. Mag. 33: t. 6275. 1877.

Subspecies drummondii is distinguished by its involucral
bracts, which are wider than those of ssp. texanwm and are
distinctly lanceolate. Populations occur in much of Texas
north of San Antonio, and in Oklahoma and eastern New
Mexico.

Drummond 227 (GH) was selected lectotype because it is
the largest and most complete specimen. The Riddell col-
lection cited by Torrey & Gray (1842) is fragmentary and
a putative hybrid between the two subspecies.

SOME REPRESENTATIVE SPECIMENS EXAMINED. NEW MEXICO: Roose-
velt Co., Elida, 5 mi. NE, 28 Sept. 1965, Anderson 3,004 (KSU). Lingo,
1.2 mi. N, Semple & Shea 705 (MO). OKLAHOMA: Beckham Co.,
Carter, 5 mi. S, 5 July 1965, Thomas 79 (OKLA). Caddo Co., Cement,
27 June 1936, Demaree 18, 133 (NY). Comanche Co., Cache, June
1927, Ortenburger s.n. (U.S.) Creek Co., Drumright, 10 June 1935,
Fry 8 (OKLA). Custer Co., Clinton, 28 June 1966, Ganz s.n. (ARIZ.
st. U.). Grady Co., Verdun, 25 June 1963, Pearce 885 (OKLA, SMU).
Greer Co., Granite, 17 June 1939, Grawmanns 85 (OKLA). Harmon
Co., Vinson, Shinners 31,644 (sMU). Jackson Co., Duke, 1.7 mi W.
15 July 1970, Flyr s.n. (MO). Kay Co.; 26 July 1898, White s.n.
(us). Kiowa Co. Mt. Park, 23 June 1913, Stevens 1,293 (OKLA).

1974] Xanthisma — Semple 17

Lincoln, 23 Aug. 1895, Blankenship 17,334 (wis). Logan Co., Guthrie,
12 July 1916, Keyser 6,036 (Ny). Osage Co., 8 mi. S of Kans. state
line on Okla. Hwy. 99, 24 June 1972, Semple & Shea 667 (MO).
Oklahoma Co., Oklahoma City, 27 May 1937, Waterfall 582 (OKLA,
NY). Pawnee Co., Pawnee, 8 mi. E, 8 June 1934, Brodell s.n. (TEX).
Payne Co., Mulhall, 1.1 mi. N, 25 June 1972, Semple & Shea 674
(MO). Stephens Co., Duncan, 8 mi. N, 27 May 1960, Waterfall 15,911
(OKLA). Washita Co., 2.2 mi. E of county line on Okla. Hwy. 152,
20 Aug. 1970, Semple & Love 264 (Mo). TEXAS: Andrews Co.,
Andrews, 4 mi. W, 28 May 1956, Turner 3,957 (TEX). Archer Co.,
Mankins, 2 mi. NE, 8 June 1956, Shinners 23,758 (SMU). Baylor Co.,
Seymour, 4 Aug. 1955, Shinners 20,776 (SMU). Blanco Co., Silverton,
Farm Rd. 145, 26 June 1972, Semple & Shea 699 (M0). Burleson Co.,
Caldwell, 14 June 1971, Semple 625 (Mo). Burnet, 8 mi. W, 19 May
1955 Turner & Johnston 2,480 (TEX). Callahan Co., Baird, Aug. 1882,
Letterman 26 (Mo, US). Childress Co., Childress, Biology Class C.
High School 12 (TEX). Coke Co., Robert Lee, 3 mi. S, 4 Oct. 1968.
Gary 192 (TrC). Comanche Co. Comyn, Theney School s.n. (TEX),
Dallas Co., Dallas, near Comanche Peak, Aug. 1877, Reverchon s.n.
(vs). Dewitt Co., western part of Co., 20 July 1941, Reidel s.n.
(TEX). Erath Co., 1922, Gaugh 29 (vs). Fisher Co., Rotan, 10 Sept.
1933, Brooker s.n. (TEX). Gaines Co., Lamesa, 15.1 mi. W, 14 Sept.
1946, Whitehouse 16,788 (NY, SMU). Gillespie Co., Cherry Spring,
18??, Jermy 639 (Mo). Gonzales Co., Cost, 11.5 mi. SW, 29 June
1972, Semple & Shea 730 (Mo). Hall Co., Turkey, 5 mi. S, 12 June
1950, Tilton T.550.604 (TTC). Hood Co., 6 Aug. 1877, Reverchon
1368 (KsU. NY, US). Howard Co. Big Spring, 22 mi. S, 27 April
1963. Dallas 64 (OKLA). Kerr Co.. 6 June 1929, Whitehouse 7.056
(TEX). Limestone Co.. Kosse, 3.5 mi. N, 14 June 1971, Semple 626
(MO). Lubbock Co., Lubbock, Demaree 7,715 (MO, TEX, TTC. US).
McCullogh Co., 1 mi. S of Colorado R., 28 June 1972, Semple & Shea
716 (MO). Navarro Co., Corsicana, 21 June 1872, Hall 311 (GH, MO,
NY, Us). Robertson Co., Calvert, 8 May 1904, Tyler s.n. (US). Tar-
rant Co. Lake Fort Worth, 16 July 1923, Ruth 742 (GH, KSU, US,
wis). Travis Co. Austin, 9 June 1908, Biltmore s.n. (US). Van
Zandt Co., Edom, 3 July 1939, Ball 7 (sMv). Washington Co., 20
May 1938, Brackett s.n. (TEX). Young Co. Belknap, 6 May 1858,
Hayes 397 (NY).

LITERATURE CITED

Braun, E. L. 1950. Deciduous Forests of Eastern North America.
Blakiston Co. Philadelphia.

Bray, W. L. 1906. Distribution and Adaptation of the Vegetation
of Texas. Bull. of U. of Tex. No. 82, Sci. Ser. No. 10.

18 Rhodora [Vol. 76

CANDOLLE, A. P. DE. 1836. Prodromus 5: 94-95.

CLAUSEN, R. T. 1941. On the terms “subspecies” and “variety.”
Rhodora 43: 157-167.

CORRELL, D. S. & M. C. JoHNSTON. 1970. Manual of the Vascular
Plants of Texas. Texas Research Foundation Press. Austin.

Du RErTZ, G. E. 1930. The fundamental units of biological tax-
onomy. Svensk. Bot. Tidskr. 24: 333-428.

EHRENDORFER, F. 1968. Geographical and ecological aspects of
infraspecific differentiation. In V. H. Heywood, ed. Modern
Methods of Plant Taxonomy. Academic Press. New York.

FENNEMAN, N. M. 1931. Physiography of the Western United
States. MeGraw-Hill Book Co. New York.

FERNALD, M. L. 1940. Some spermatophytes of eastern North
America. Rhodora 42: 239-278.

FoscuE, E. J. 1932. The natural vegetation of the lower Rio
Grande Valley, Texas. Field & Lab. 1: 25-30.

GOULD, F. W. 1962. Texas Plants— a checklist and ecological
summary. Texas A. & M. — Tex. Agr. Ex. Sta. College Station,
Texas.

GRAY, A. 1852. Plantae Wrightiana Texano — Neo-Mexicanae.
Part I. Smiths. Contrib. Knowl. 3: 98.

Hooker, J. D. 1877. Curtis’ Bot. Mag. 33: t.6275.

OETKING, P. F. 1959. Geological Highway Map of Texas. Dallas
Geological Soc. Dallas.

Rarsz, E. 1957. Land forms of the United States. Jn Oetking,
P.F. 1959. Geological Highway Map of Texas. Dallas Geological
Soc. Dallas.

SELLARD, E. H., W. S. ADKINS & R. B. PLUMMER. 1966. Geology of
Texas. Vol. I. Stratigraphy. U. of Tex. Bull. No. 3232.

SEMPLE, J. C. 1972a. Behavior of B-chromosomes in Xanthisma
texanum DC.: a non-random phenomenon. Science 175: 666.

1972b. Cytology, Flavonoid Chemistry and System-
aties of the Texas Sleepy Daisy Xanthisma texanum DC. (As-
teraceae). Ph.D. Dissertation. Washington University. St. Louis.

SMALL, J. K. 1903. Flora of the Southeastern United States of
America. Pub. by Small. New York. p. 1184.

SMITH, E. B. & H. M. Parker. 1971. A biosystematic study of
Coreopsis tinctoria and C. cardaminefolia. Brittonia 23: 161-170.

STEENIS, C. G. G. J. VAN. 1957. Specific and infraspecific delimina-
tion. Flora Malesiana Ser. 1, 5: clxvii-eexxxix.

THARP, B. C. 1926. Structure of the Texas Vegetation East of the
98th Meridian. U. of Tex. Bull. No. 2606.

Torrey, J. 1859. In Emory, W. H., ed. Report on the United States
and Mexican Boundary Survey. Nicholson Press. Washington.
Vol. II. pp. 80-81.

1974] Xanthisma — Semple 19

. & A. GRAY. 1842. A Flora of North America. Vol. II.
facsimile ed., 1969. Hafner Pub. Co. New York. p. 246.
VALENTINE, D. H. & A. LóvE. 1958. Taxonomic and biosystematic
categories. Brittonia 10: 153-166.
VISHER, S. S. 1954. Climatic Atlas of the United States. Harvard

U. Press. Cambridge.

MISSOURI BOTANICAL GARDEN
ST. LOUIS, MO. 63110

A WHITE-FLOWERED FORM OF UTRICULARIA
PURPUREA FROM NEW HAMPSHIRE. A white-
flowered form of Utricularia purpurea Walter was found
while conducting research in a small pond in New Hamp-
shire. This form was extremely abundant forming exten-
sive mats on the surface of this shallow pond. None of
the normal purple-flowered plants were observed. Plants
commonly associated with the white form were Utricularia
vulgaris L., Utricularia intermedia Hayne and Najas flexilis
Rostk. & Schmidt. The pond had a pH of 6.5 and methyl
orange alkalinity reading of 8.0 mg/1.

The following form is described. Utricularia purpurea
Walter forma alba Hellquist, forma nova. differt forma
purpurea quantum flores albos non purpureos habet. TYPE:
NEW HAMPSHIRE: CARROLL COUNTY: small unnamed pond
southeast of Dorr Pond on N.H. Route 153 at the settlement
of Woodman, Town of Wakefield (U.S.G.S. quadrangle
Newfieid, Me.-N.H.), C. B. Hellquist 8935 (NHA).

The holotype has been placed at the University of New
Hampshire Herbarium (NHA). Isotypes are in the her-
barium of the New England Botanical Club (NEBC) and
Boston State College Herbarium.

C. BARRE HELLQUIST
BIOLOGY DEPARTMENT

BOSTON STATE COLLEGE
BOSTON, MASSACHUSETTS 02115

ON THE SCIENTIFIC NAME OF THE
LONGLEAF PINE'

DANIEL B. WARD

The Longleaf Pine of the southeastern United States has
been spared the numerous changes in scientific name that
have cast confusion over many of our other tree species.
From pioneer days the dominant practice, both among
botanists and foresters, has been to apply the name Pinus
palustris Mill. to the Longleaf. Prior to the 1940's the
only author of significance to do otherwise was J. K. Small
(1933) who interpreted P. palustris Mill. to be the northern
variety of slash pine, P. elliottii Engelm. Small’s Manual
of the Southeastern Flora, however, achieved only regional
impact, and his use of P. australis Michx. f. for the Long-
leaf did not gain a wide following.

A much stronger impetus for associating the Longleaf
with Pinus australis was generated in 1948 by M. L. Fer-
nald who advocated this name in two detailed and strongly
worded articles (Fernald & Schubert, 1948; Fernald,
1948). Fernald's position was reinforced by the publica-
tion of his monumental reworking of Gray's Manual of
Botany (1950), and was further entrenched when the inde-
pendent H. A. Gleason adopted the same name in his New
Britton and Brown Illustrated Flora (1952) as did A.
Cronquist in their companion field guide, the Manual of
Vascular Plants of Northeastern United States and Adja-
cent Canada (Gleason & Cronquist, 1963). These three
publications totally dominate the field of regional floras in
the Northeast, and their consistent use of Pinus australis
inevitably suggests that this name is supported by general
professional agreement and by nomenclatural legitimacy.

Yet in publications other than these or their derivatives,
the Longleaf remains known by the name Pinus palustris

"This paper is Florida Agricultural Experiment Station Journal
Series No. 4775.

20

1974] Longleaf Pine — Ward 21

Mill. This is the name found in state or district floras
such as the Manual of the Vascular Flora. of the Carolinas
(Radford et al., 1968), the Manual of the Vascular Plants
of Texas (Correll & Johnston, 1970), and A Flora of
Tropical Florida (Long & Lakela, 1971), as well as in the
more basic modern treatments of the genus by Mirov
(1967) and by Little & Critchfield (1969). The U. S. Forest
Service, following the guidance of its dendrologist, E. L.
Little, has remained steadfast with P. palustris, and the
state forest services have done likewise.

The rationale behind Fernald's heterodox displacement

of Pinus palustris was his view that Philip Miller (1768),
superintendent of the Chelsea Physic Garden and British
author of a series of widely used and authoritative horti-
cultural dictionaries, did not have the Longleaf when he
described an American tree under this name. The original
description contained phrases that do not apply to the
stately and versatile Longleaf: “I have been informed that
they grow to a height of twenty-five or thirty feet.
I have not heard the wood was of any use but for fuel."
However, Miller reported these statements as hearsay, not
as observed fact, while his description also contained such
passages as ‘“‘Pine-tree with the longest leaves growing by
threes out of each sheath" and “Their leaves are a foot or
more in length, growing in tufts at the end of the branches,
so have a singular appearance." These phrases were given
without qualification and form a brief but unmistakable
description of the Longleaf,

Fernald further relied on the habitat of Pinus palustris
reported by Miller, “swamps in many parts of North
America," as excluding the Longleaf. He proposed instead
that Miller may have had the Loblolly Pine, P. taeda L.
Actually however, although the Longleaf usually occurs on
dry well-drained soils, it not uncommonly is found associated
with the Slash Pine in the extensive level undrained swamps
known as flatwoods and throughout its range probably
occurs as frequently in wet soils as Fernald's proposed
substitute, the Loblolly. Of couse, Miller's use of an epithet

22 Rhodora [Vol. 76

meaning “swampy”, even if it were totally unsuitable,
would not be justification for discarding his P. palustris;
Article 62 of the International Code of Botanical Nomen-
clature (Stafleu, 1972) provides that: “a legitimate name
or epithet must not be rejected merely because it is in-
appropriate . . ."

No information is available from specimens preserved
by Miller; Dr. Schubert examined the Miller collections of
the British Museum and was unable to find a specimen that
could be considered a type. Fernald, by correspondence,
did locate a specimen he presumed to be of P. taeda that
Miller may have seen and that someone, possibly Miller,
had noted as “palustris.” It seems improbable that Miller
would have so misidentified P. taeda since this species was
well known to him and was treated in some detail in the
several editions of his “Dictionary.”

Fernald did demonstrate at some length that the name
Pinus palustris was not clearly understood and was even
misused by Miller's contemporaries and other early workers,
but this again does not provide justification for discarding
the name. There seems to be essentially no sound rationale
for interpreting this name to mean anything other than,
in the conventional usage, the Longleaf Pine.

In the event, moreover, that Miller's brief but clear
diagnosis should be considered ambiguous, no alternative
name is available. Pinus lutea Walter, whimsically sug-
gested by Fernald, is probably P. taeda. Pinus australis
Michx. f. is unmistakably the Longleaf, and was selected
as the correct name for this species by Small and by
Fernald. Little (1948), however, accurately pointed out
prior to the publication of Fernald's first paper on the
subject, that Article 63 (then Article 60) of the Inter-
national Code makes this name illegitimate since it was
superfluous when published. Not only did Michaux (1810)
cite P. palustris as a synonym, but he specifically explained
that he was replacing this name with P. australis, using
an epithet (“southern”) that he thought more appropriate.
It is precisely such actions as this that Article 63 is de-

1974] Longleaf Pine — Ward 23

signed to prevent, and Fernald's protestations that Michaux
was “definitely defining a new species" are contradicted by
Michaux' own words, in translation: *I have thought like-
wise that the specific name ‘australis’ was preferable to
that of ‘palustris’, under which this species has been de-
scribed by botanists; for this last gives an absolutely false
idea of the nature of the soil where this tree grows.”

The only other specific name apparently ever given the
Longleaf was Pinus longifolia Salisb., but here again the
epithet used was superfluous, for Salisbury (1796) was un-
abashedly providing a substitute name for P. palustris and
thereby formed an illegitimate and unusable combination.

The person who would apply a scientific name to the
Longleaf Pine, therefore, has the choice, should he wish to
follow the Code, of either accepting Miller’s brief descrip-
tion of Pinus palustris as adequate, or of discarding it as
confused and coining and publishing a name of his own
creation. Neither P. australis Michx. f. nor P. longifolia
Salisb. may be legitimately used for this tree, and prudence
and practicality, as well as historical precedent, indicate
strongly the advisability of retaining the Longleaf Pine
under the name Pinus palustris Mill.

LITERATURE CITED

CORRELL, D. S. & M. C. JOHNSTON. 1970. Manual of the Vascular
Plants of Texas. Texas Research Found. 1881 p.

FERNALD, M. L. 1948. The confused bases of the name Pinus
palustris. Rhodora 50: 241-249.

. 1950. Gray's Manual of Botany, 8th ed. Ameri-

can Book Co. 1632 p.

, & B. G. SCHUBERT. 1948. Studies of American
types in British herbaria. Rhodora 50: 181-190.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated
Flora. Lancaster Press. Vol. 1.

,& A. CRONQUIST. 1963. Manual of Vascular Plants
of Northeastern United States and Adjacent Canada. Van Nos-
trand Co. 810 p.

LITTLE, E. L. 1948. Notes on nomenclature of trees. Phytologia
2: 457-458.

24 Rhodora [Vol. 76

, & W. B. CRITCHFIELD. 1969. Subdivisions of the
genus Pinus (pines). U. S. Dept. of Agric. For. Ser. misc. publ.
1144. 51 p.

Lonc, R. W., & O. LAKELA. 1971. A Flora of Tropical Florida.
Univ. of Miami Press. 962 p.

MicHAUX, F. A. 1810. Histoire des Arbres Forestiers de l'Amerique
Septentrionale. 1: 64.

MiLLER, P. 1768. The Gardeners Dictionary, 8th ed. London.

Mirov, N. T. 1967. The Genus Pinus. Ronald Press. 602 p.

RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1968. Manual of the
Vascular Flora of the Carolinas. Univ. N. Car. Press. 1183 p.

SALISBURY, R. A. 1796. Prodromus Stirpium in Horto ad Chapel
Allerton Vigentium. London.

SMALL, J. K. 1933. Manual of the Southeastern Flora. The
author, New York. 1554 p.

STAFLEU, F. A. 1972. International Code of Botanical Nomencla-
ture. Utrecht. 426 p.

DEPARTMENT OF BOTANY
AGRICULTURAL EXPERIMENT STATION
UNIVERSITY OF FLORIDA
GAINESVILLE, FLORIDA

THE OCCURRENCE OF BALD CYPRESS (TAXO-
DIUM DISTICHUM (L) RICHARD) IN SUFFOLK
COUNTY, LONG ISLAND, NEW YORK. In October,
1972, Mr. Richard Fackovec, a student in an aquatic
botany course at Southampton College, brought to our
attention a specimen of Taxodium distichum that he col-
lected along the shore of Long Pond in Sag Harbor, New
York. We returned to investigate the region in November
and found eight bald cypress seedlings, ranging in height
from one to 214 meters and growing close to shore in
0.5-0.7 meters of water. The four smallest trees (approxi-
mately one meter tall) are growing close together at the
southern end of the pond at the edge of a large marsh. The
other cypress trees are located at the southern end of the
pond also, but along the western shore. The two largest
trees (215 and 114 meters tall) are at the edge of a large
stand of Typha latifolia. Most of the remaining shoreline
of Long Pond consists of very dense vegetation composed
primarily of shrubs and small trees (e.g. Myrica gale,
Decodon verticillata, Cassandra calyculata, Vaccinium
corymbosum, Nyssa sylvatica, Acer rubrum). It is within
this shrub zone that we found two additional bald cypress
seedlings. One had become encompassed so completely by
encroaching shoreline vegetation that we removed it; sub-
sequently, it has been replanted along the shore of a pond
in the Morton Wildlife Refuge, a Federal Wildlife Preserve
in Noyac, New York.

We have communicated with Dr. Richard Stalter, plant
taxonomist at St. Johns University in Jamaica, New York,
regarding the northern distribution of Taxodium distichum.
As far as we have been able to determine, this is the first
record of a natural population of bald cypress in New
York State. The closest naturally-occurring populations
of Taxodium are in Delaware, where there are a few large
colonies (Stalter, personal communication). There was,
at one time, a colony in New Jersey, but it has been un-
known in that state for several years (Stalter, personal
communication).

25

26 Rhodora [Vol. 76

Accompanied by Dr. Stalter, we returned to Long Pond
in December in an unsuccessful effort to locate mature
bald cypress trees. We share the opinion with Dr. Statler
that this colony of Taxodium distichum represents a natu-
ralized population. The presence of a few bald cypress
seedlings in relatively inaccessible portions of the shore-
line strongly suggests that they were not planted. The
apparent absence of large bald cypress in the area leaves
unanswered the question as to how the seedlings became
established; hopefully, further investigation will afford us
an answer.

The voucher specimen collected by Mr. Fackovec is de-
posited in the Southampton College Herbarium.

EDWARD J. HEHRE

PETER M. BOLLARD

DEPARTMENT OF BIOLOGY
SOUTHAMPTON COLLEGE
SOUTHAMPTON, NEW YORK 11968

STUDIES ON THE BRYOPHYTES
OF SOUTHERN MANITOBA
III. COLLECTIONS FROM GRAND BEACH
PROVINCIAL PARK

PAUL W. STRINGER AND MURIEL H. L. STRINGER

Grand Beach Provincial Park lies approximately 57 miles
north of Winnipeg. The entire park, which covers 59,000
acres, is in the Manitoba Lowlands Section of the Boreal
Forest Region (Rowe, 1959). Three miles of the area
fronts on Lake Winnipeg. 'The underlying bedrock is
palaeozoic limestone, and the overlying beds are modified
till, or lacustrine clays deposited in glacial Lake Agassiz.
Most soils have a high lime content.

On the flat, poorly drained areas, black spruce (Picea
mariana (Mill. BSP.) and tamarack (Larix laricina (Du
Roi) K. Kich) forest prevail, while on the better drained
alluvial strips bordering the rivers and creeks, white spruce
(Picea glauca (Moench) Voss), aspen (Populus tremuloides
Michx.) and balsam poplar (Populus balsamifera L.) pre-
dominate. On the low, well-drained ridges, jack pine (Pinus
banksiana Lamb.) is dominant, with aspen, paper birch
(Betula papyrifera Marsh.) and some bur oak (Quercus
macrocarpa Michx.) present.

No bryophytes have previously been reported for the
park. In the present study, several sites were visited,
mostly in the vicinity of 96°35’W and 50°33’N. The sites
were chosen as representative of the major vegetation types
in the area, and the following seven, visited in July, 1972,
are reported as representative of the area studied. Several
of the vascular collections could be determined to genus
only, because of lack of flowering or fruiting material.

1. Sand dunes and dune slacks behind the East Beach.
Dominant shrubs were Salix spp., especially in the slacks,
with Prunus pumila L. conspicuous on the drier areas.
Other prominent vascular plants on the dry areas included
Carex spp. Rhus radicans L. var. rydbergii (Small)

27

28 Rhodora [Vol. 76

Rehder, and Artemisia spp. Total plant cover was rather
thin (about 20%) except in the slacks, and most bryophyte
cover was on wet litter in the slacks. Small turf mosses
such as Bryum spp. were, however, common in dry habi-
tats.

2. Aspen-dominated forest just s. of the parking lot at the
East Beach. Most mosses were from tree bases.

3. Mixed mature aspen — jack pine forest on w. side of
the road n. of main park entrance.

4. Black spruce bog with some tamarack and white cedar
(Thuja occidentalis L.). The understory was dominated by
Ledum groenlandicum Oeder and the terrain consisted of
pools and Carex hummocks. Just s. of Highway 12, 14 mile
from its junction with Highway 59.

5. Open jack pine with some birch, on s. side of Highway
12, 2 miles from its junction with Highway 59.

6. Tamarack — black spruce bog on e. side of Highway 59,
2 miles n. of its junction with Highway 12. Drier than
site 4, and with a tall shrub layer of Alnus sp., as well as a
short shrub layer dominated by Ledum groenlandicum.
Deadfall was abundant.

7. Mature, open jack pine forest on w. side of Highway 59
4 miles n. of its junction with Highway 12. There was an
open short shrub and herb layer dominated by Vaccinium
angustifolium Ait., Chimaphila umbellata (L.) Bart. and
Amelanchier alnifolia Nutt., with Linnaea borealis L. and
Vicia americana Muhl. prominent. The forest floor was
dominated by Plewrozium schreberi and Dicranum spp.,
especially D. polysetum. The many large humus-covered
boulders present all had an extensive, species-rich bryo-
phyte cover. Numbers in the species list refer to the sites
described above.

Taxonomic and field work were shared equally by both
authors. Voucher specimens have been deposited in the
authors’ own herbarium and at the University of Winnipeg.
Nomenclature for Sphagnobrya and Eubrya follows Crum,
Steere, and Anderson (1965) with modification according
to Crum (1971). Hepatic nomenclature follows Schuster

1974] Bryophytes — Stringer & Stringer 29

(1969), supplemented by Schuster (1953) with abbrevia-
tions of authorities amended to conform with the list of
Sayre, Bonner and Culberson (1964). Vascular plant no-
menclature follows Scoggan (1957).

Hepaticae

Calypogeia muelleriana (Schiffn.) K. Miill. (6). Three
collections, two on humus mixed with well-rotted wood,
and one on a rotten stump with Lophocolea heterophylla,
Cephalozia media and Hypnum pallescens. Only once pre-
viously recorded for the province (Longton, 1972).

Cephalozia media Lindb. (6). One collection, on a rotten
stump. |

Cephaloziella rubella (Nees) Douin (4). On humus over
rotten wood, with Amblystegium juratzkanum, Chiloscy-
phus pallescens and Thuidium recognitum.

Chiloscyphus pallescens (Ehrh.) Dum. (4). Only once pre-
viously recorded for the province, in Spruce Woods Pro-
vincial Park (Stringer & Stringer, a, in preparation).

Chiloscyphus polyanthus (L.) Corda (3, 4). On bark at the
base of an aspen tree (3) with Hypnum pallescens, and
on humus with Distichium inclinatum. Recorded previ-
ously only in Spruce Woods Provincial Park (Stringer &
Stringer, in preparation).

Lophocolea heterophylla (Schrad.) Dum. (3, 6, 7,). On
humus or rotten wood, often with Pohlia nutans or Hyp-
num pallescens. Previously recorded only in the Spruce
Woods area by Bird (1969) and by Stringer & Stringer
(in preparation).

Lophocolea minor Nees (6). On thick humus (dead Aula-
comnium palustre), with Pohlia nutans and Eurhynchium
pulchellum. Not previously recorded for the province.

Ptilidium ciliare (L.) Nees (5, 6, 7,). On loose humus or
litter with Dicranum spp. and Pleurozium schreberi. One
collection (7) on loose conifer needles over rock, with
Hedwigia ciliata.

Ptilidium pulcherrimum (Web.) Hampe (7). One collec-
tion, on fallen birch bark.

30 Rhodora [Vol. 76

Radula complanata (L.) Dum. (7) One collection, on packed
humus at the base of a jack pine tree.

Riccardia pinguis (L.) S. Gray (4). On thick, moist humus
mixed with silt, with Myurella julacea, Amblystegium
juratzkanum and Platydictya jungermannoides. Previ-
ously recorded only in the Spruce Woods area by Bird
(1969) and Stringer & Stringer (in preparation).

Sphagnobrya
Sphagnum capillaceum (Weiss) Schrank (6). In deep hum-
mocks, with Pleurozium schreberi.

Eubrya

Amblystegium juratzkanum Schimp. (1, 2, 4, 5, 6, 7). On
rotten wood or humus, especially at tree bases, where it
was often found with Pylaisiella polyantha and Brachy-
thecium salebrosum.

Amblystegium varium (Hedw.) Lindb. (1, 2, 6). On thick
humus and rotten wood, with Brachythecium spp. and
Mnium cuspidatum.

Aulacomnium palustre (Hedw.) Schwaegr. (4, 5, 6). On
thick humus, in deep pure sods, or with Pleurozium
schreberi. One collection (5) on rotting birch bark.

Brachythecium campestre (C. Müll.) B. S. G. (5, 6, 7). On
humus, usually over well-drained sandy soil. Previously
recorded in Manitoba only in the Winnipeg and Spruce
Woods areas by Stringer & Stringer (1973) and Stringer
& Stringer (in preparation).

Brachythecium rutabulum (Hedw.) B. S. G. (1, 2, 5, 6, 7).
On humus or rotten wood in moist habitats. Previously
recorded only by Mueller-Dombois (1964), Longton
(1972), and by Stringer & Stringer (in preparation), in
the Spruce Woods area.

Brachythecium salebrosum (Web. and Mohr) B. S. G. (1,
2, 3, 5, 6, 7). Common in many habitats, especially on
tree bases.

Bryoerythrophyllum recurvirostrum (Hedw.) Chen (4). On
thick humus, with Amblystegium juratzkanum, Myurella
julacea and Platydictya jungermannoides.

1974] Bryophytes — Stringer & Stringer 31

Bryum angustirete Kindb. ex Mac. (1). On humus over
sand, alone, and with Encalypta procera.

Bryum argenteum Hedw. (1, 3). On dry humus and sand.
Often with Funaria hygrometrica.

Bryum creberrimum Tayl. (3, 5). On humus mixed with
sand. Often with Ceratodon purpureus.

Bryum pallescens Schleich. ex Schwaegr. (I, 2,3). Oh
humus in moist habitats.

Bryum pseudotriquetrum (Hedw.) Gaertn. Meyer and
Scherb. (2, 4). On thick, moist humus with Campylium
stellatum and Brachythecium rutabulum.

Callicladium haldanianum (Grev.) Crum (5, 6, 7). On
humus and moist rotten wood. Previously reported only
in the Winnipeg and Spruce Woods areas by Bird (1969)
and by Stringer & Stringer (a, in preparation).

Calliergon giganteum (Schimp.) Kindb. (4). On water-
logged litter, in large masses mixed with Campylium
stellatum.

Campylium chrysophyllum (Brid.) J. Lange (4). On well-
rotted wood and humus.

Campylium hispidulum (Brid.) Mitt. (4). One collection,
on humus. with Amblystegium juratzkanum.

Campylium stellatum (Hedw.) C. Jens. (4, 6). On wet
humus or litter, often with Mnium affine or Thuidium
delicatulum var. radicans.

Ceratodon purpureus (Hedw.) Brid. (1, 3, 4, 5, 7). On
humus over dry sand, with Bryum spp., Funaria hygro-
metrica, or Polytrichum juniperinum.

Dicranum bonjeanii De Not. ex Lisa (7). On thick humus
over rotten wood, with D. polysetum and Ptilidium cili-
are.

Dicranum drummondii C. Müll. (6, 7). In thick, deep, pure
clumps on humus, or as small plants on rotten wood with
such typical species of decayed wood as Pohlia nutans
and Tetraphis pellucida.

Dicranum fuscescens Turn. (5, 7). On humus or rotten
wood, usually in pure tufts.

32 Rhodora [Vol. 76

Dicranun polysetum Sw. (4, 5, 6, 7). Common, especially
in jack pine forests of the area. Forms deep pure tufts.
Often found also with Pleurozium schreberi.

Distichium inclinatum (Hedw.) B. S. G. (4). Common in
this collecting area and most often found with Myurella
julacea, Amblystegium juratzkanum, and Platydictya
jungermannoides. Several of the collections had mature
capsules, and the large warty spores (354-404) distin-
guish this species from the closely related Distichium
capillaceum (Grout, 1936).

Drepanocladus aduncus (Hedw.) Warnst. (1). On wet lit-
ter under willows.

Drepanocladus aduncus (Hedw.) Warnst. var. polycarpus

. (Bland. ex Voit) Roth (1, 2). More commonly found

. than the preceding, this variety occurred on moist humus
or litter, often with Leptodictyum riparium.

Drepanocladus revolvens (Sw.) Warnst. (4). Several col-
lections made on wet litter or humus, in pure mats or
with Pleurozium schreberi and Thuidium recognitum.

Drepanocladus uncinatus (Hedw.) Warnst. (4, 5, 6, 7). On
rotten wood or humus, in drier habitats than other mem-
bers of the genus. With Brachythecium spp. and Bryum
Spp.

Encalypta procera Bruch (1). One collection, on thin hu-
mus over dry sand.

Eurhynchium pulchellum (Hedw.) Jenn. (4, 6, 7). On
thick humus or litter in both wet and dry habitats.

Fissidens adianthoides Hedw. (4). One small collection,
with Campylium stellatum, on loose, moist humus. Not
previously recorded for the province.

Funaria hygrometrica Hedw. (1, 4, 5, 7). Frequent on
sandy soil, often mixed with ashes of old fires. Com-
monly with Bryum spp., Ceratodon purpureus, and Lepto-
bryum pyriforme.

Haplocladium microphyllum (Hedw.) Broth. (2, 3, 6, 7).
Almost all collections on rotten wood but one collection
(7) on humus.

1974] Bryophytes — Stringer & Stringer 33

Hedwigia ciliata (Hedw.) P. Beauv. (7). Several collec«
tions, all on humus over rock or loose conifer needles over
rock.

Hylocomium splendens (Hedw.) B. S. G. (4, 6, 7). On litter
and loose humus, often with Pleurozium schreberi.

Hypnum lindbergii Mitt. (4, 5, 6). Usually on wet, well-
rotted wood or wet humus.

Hypnum pallescens (Hedw.) P. Beauv. (3, 5, 6, 7). Com-
mon on live and fallen bark of both conifers and hárd-
woods. Often associated with Pylaisiella polyantha.

Hypnum pratense Koch ex Spruce (1). One collection, on
wet rotten bark with Drepanocladus aduncus var. poly-
carpus.

Isopterygium turfaceum (Lindb.) Lindb. (7). One collec-
tion, on thick humus over granite rock, with Pohlia nu-
tans and Brachythecium rutabulum.

Leptobryum pyriforme (Hedw.) Wils. (1, 2, 4, 7). Com-
mon on a wide variety of substrates and with a wide
range of associated species.

Leptodictyum riparium (Hedw.) Warnst. (1, 2). On wet
humus or litter, often with Drepanocladus aduncus or
Amblystegium varium.

Leptodictyum trichopodium (Schultz) Warnst. var. kochii
(B. S. G.) Broth. (1, 2). On deadfall or humus in moist
habitats.

Leskea polycarpa Hedw. (1, 2,5). On bark or humus, found
most often with Amblystegium varium or Leptodictyum
trichopodium var. kochii.

Mnium affine Bland. ex Funck (4, 6). On wet humus or
disintegrated rotten wood.

Mnium cuspidatum Hedw. (1, 2, 3, 5, 6). On humus, rotten
wood or litter in drier habitats than M. affine, often alone,
or with Brachythecium salebrosum.

Myurella julacea (Schwaegr.) B. S. G. (4). Common in
wet habitats in this collecting area and often found with
Campylium stellatum and Aulacomnium palustre.

Orthotrichum obtusifolium Brid. (3). One collection, on
poplar bark, with Hypnum pallescens.

34 Rhodora [Vol. 76

Orthotrichum speciosum Nees ex Sturm (3, 7). On live
poplar bark, often with Pylaisiella polyantha, but per-
sisting on deadfall even in advanced stages of decay
where it often occurs with Haplocladium microphyllum.

Platydictya jungermannioides (Brid.) Crum (4). Fairly
common in this collecting area, with Myuwrella and Dis-
tichium. Recorded only once previously for the province,
as Amblystegiella sprucei (Bruch) Loeske, by Crum and
Schofield (1959).

Pleurozium schreberi (Brid.) Mitt. (3, 4, 5, 6, 7). Com-
mon, and often forming large pure colonies in both wet
and dry coniferous forests.

Pohlia nutans (Hedw.) Lindb. (5, 6, 7). Usually on well-
rotted wood but sometimes (7) on humus.

Polytrichum juniperinum Hedw. (5, 6, 7). Usually on hu-
mus over sand in drier habitats.

Polytrichum piliferum Hedw. (7). One collection on humus
over sand.

Ptilium crista-castrensis (Hedw.) De Not. (5, 7). Usually
a forest floor species of dry coniferous forests in the
area, occurring on loose humus or needle litter over sand.

Pylaisiella polyantha (Hedw.) Grout (2, 3, 5, 6, 7). Com-
mon with Orthotrichum speciosum, especially on bark of
live hardwoods, but also on humus (6, 7) or at the base
of jack pine (5).

Tetraphis pellucida Hedw. (6). On well-rotted wood, with
Pohlia nutans and Drepanocladus uncinatus.

Thuidium delicatulum (Hedw.) B. S. G. var. radicans
Crum, Steere & Anderson (4). On loose wet humus and
litter with Hypnum lindbergii, Campylium stellatum and
Tomenthypnum nitens.

Thuidium recognitum (Hedw.) Lindb. (4, 6). On loose hu-
mus and needle litter in wet or moist habitats.

Tomenthypnum nitens (Hedw.) Loeske (4). On loose, wet
humus and litter.

Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb. (7). On
thin humus over sand.

Stringer & Stringer 35

1974] Bryophytes

DISCUSSION

A search of the literature reveals that little work has
been done on the bryophytes of Manitoba. Bird (1966),
who gives a comprehensive list of the recordings for the
province, lists only 289 taxa of bryophytes for the whole of
Manitoba. This list has been somewhat augmented by Bird
(1969) and Longton (1972), but published work on the
bryophyte flora of the province is still greatly lacking. This
situation is especially unfortunate as Manitoba is particu-
larly interesting from a phytogeographic point of view, ly-
ing as it does at a “crossroads of floral elements, from both
north and south, east and west" (Lóve, 1959).

Approximately half of the species recorded in the present
study are pan-North American in distribution, according to
the designations given by Bird (1969) and Bird and Won
Shic Hong (1969). However, the Boreal element is repre-
sented by species such as Dicranum fuscescens, D. polyse-
twm, Tetraphis pellucida, Myurella julacea, Thuidium deli-
catulum var. radicans, Thuidium recognitum, Pleurozium
schreberi, Ptilium crista-castrensis, and Pylaisiella poly-
antha. The Arctic Boreal element is represented by Di-
cranum | bonjeanüi, Bryoerythrophyllum | recurvirostrum,
Bryum pallescens, Orthotrichwm speciosum, Calliergon gi-
ganteum, Campylium stellatum, Eurhynchium pulchellum,
Tomenthypnum nitens, Hylocomium splendens, Polytrichum
juniperinum, and Polytrichum piliferum. A few Eastern
Boreal species are also present, i.e. Dicranum drummondii,
Leptodictyum trichopodium var. kochii and Callicladium
haldanianum.

The hepatics are largely pan-North American species.
Cephaloziella rubella, however, is Boreal and Ptilidium
ciliare is Arctic Boreal (Bird & Won Shic Hong, 1969).
Calypogeia . muelleriana and Cephalozia media have been
previously recorded only from the north-east part of the
province, from Churchill and York Factory respectively
(Longton, 1972). The only previous record of Cephaloziella
rubella was from Spruce Woods Provincial Park ( Stringer

36 Rhodora [Vol. 76

& Stringer, in preparation). Lophocolea minor, recorded
from Grand Beach as new to the province, has since been
found at several stations in Birds Hill and Whiteshell Pro-
vineial Parks, where Radula complanata is also quite fre-
quent. Riccardia pinguis has only two previous records,
both from Spruce Woods Provincial Park, although subse-
quent work by the authors shows that it is common on the
calcareous clays of the southern part of the province. All
of these liverworts are probably much more widely dis-
tributed throughout the province than the scant records
indicate.

Several moss species which, according to the literature,
should have a wide distribution have few records in Mani-
toba. This is especially true of Brachythecium spp., per-
haps because of the apparent scarcity of fertile material
and the consequent difficulty in making a positive identifi-
cation. Brachythecium rutabulum, common in the Grand
Beach aréa studied, was reported as new to the province
by Longton (1972), although Mueller-Dombois (1964) de-
scribed it as one of the mosses forming the typically con-
tinuous carpet in one of the jack pine — black spruce forest
types in southeastern Manitoba. The only other records of
Brachythecium campestre are also from the southern part
of the province (Stringer & Stringer, in preparation, 1973)
although this species appears to have a pan-North Ameri-
can distribution.

Callicladium haldanianum, reported for the first time in
the province by Bird (1969) as Heterophyllium haldani-
anum (Grev.) Kindb., reaches its western limit in Mani-
toba. The Grand Beach record is further north than either
of the Bird records.

Fissidens adianthoides, a new record for Manitoba, ap-
pears to be rare in western Canada (Bird 1966), although
Grout (1936) gives its distribution as from Cape Breton
Island to Vancouver Island and south to California and
Florida.

It is intended that the information presented in this
study should, in conjunction with information presented in

1974] Bryophytes — Stringer & Stringer 37

further papers in this series, assist in describing the bryo-
phyte flora of Manitoba and its relationships with the floras
of adjacent regions.

ACKNOWLEDGEMENTS

The field work was supported in part by Grant No. 140-
118 from the University of Winnipeg. The authors also
wish to express their gratitude to Dr. Dale H. Vitt, Univer-
sity of Alberta, for his prompt assistance in identification
of some of the Brachythecium material.

LITERATURE CITED

Birp, C. D. 1966. A catalogue of the bryophytes reported from
Alberta, Saskatchewan and Manitoba. Processed publication,
University of Calgary, Calgary, Alberta.

. 1969. Bryophytes of the aspen parkland of west-
central Canada. Can. Jour. Bot. 47: 187-212.

. & Won Suic Hone. 1969. Hepaticae of southwestern
Alberta, Can. Jour. Bot. 47: 1727-1746.

Crum, H. 1971. Nomenclatural changes in the Musci. The Bryol-
ogist 74: 165-174.

. & W. B. SCHOFIELD. 1959. The mosses of Gillam, Mani-
toba. Bull. No. 160, Nat. Mus. Can., Ottawa,

.„ W. C. STEERE & L. E. ANDERSON. 1965. A list of the
mosses of North America. The Bryologist 68: 377-434.

Grout, A. J. 1936. Moss flora of North America. Vol. I, Part I.
Newfane, Vermont. Published by the author.

LoNGTON, R. E. 1972. Nineteen bryophytes new to Manitoba. The
Bryologist 75: 348-350.

LóvE, D. 1959. The postglacial development of the flora of Mani-
toba: a discussion. Can. Jour. Bot. 37: 547-585.

MUELLER-DoMBors, D. 1964. The forest habitat types in south-
eastern Manitoba and their application to forest management.
Can. Jour. Bot. 42: 1417-1444.

RowE, J. S. 1959. The forest regions of Canada. Bull. 125, Dept.
Northern Affairs and Nat. Resources, Forestry Branch. Ottawa.

SAYRE, G., C. E. B. BONNER, & W. L. CULBERSON. 1964. The authori-
ties for the epithets of mosses, hepatics, and liehens. The Bry-
ologist 67: 113-135.

ScHUSTER, R. M. 1969. The Hepaticae and Anthocerotae of North
America east of the hundredth meridian. Vol. II. 1062 pp. Colum-
bia University Press, New York.

38 Rhodora [Vol. 76

1953. Boreal Hepaticae, a manual of the liver-
worts of Minnesota and adjacent regions. Amer. Midland. Nat.
49(2) : i-v, 257-684,

SCOGGAN, H. J. 1957. Flora of Manitoba. Bull. 140, Biol. Ser. No.
47, Nat. Mus. Can., Ottawa.

STRINGER, M. H. L. & P. W. STRINGER. Studies on the bryophytes
of southern Manitoba. I. Collections from Spruce Woods Pro-
vincial Park, In preparation.

& . 1973. Studies on the bryo-
phytes of southern Manitoba. II. Collections from the Winnepeg
area. Can. Field Nat. 87: 141-144.

DEPARTMENT OF BIOLOGY
UNIVERSITY OF WINNIPEG

515 PORTAGE AVENUE

WINNIPEG 2, MANITOBA, CANADA

SOME NEW OR NOTEWORTHY
VASCULAR PLANT RECORDS FROM
NORTHWESTERN SASKATCHEWAN

VERNON L. HARMS AND JOHN H. HUDSON

During the course of recent botanical studies in north-
western Saskatchewan, several new or otherwise interest-
ing plant records were collected. Most of these were col-
lected from along the Green Lake-La Loche Road during a
study supported by the Institute for Northern Studies, Uni-
versity of Saskatchewan, or from the Lake Athabasca area.
The purpose of this paper is to comment on these records
in order to make this information generally available to
taxonomists and phytogeographers. The collection numbers
are those of the senior author unless otherwise indicated.
All specimens cited have been deposited in the Fraser Her-
barium, University of Saskatchewan (SASK).

SPARGANIUM CHLOROCARPUM Rydb. 7 mi. s.-s.e. of La
Loche, Mile 60 Buffalo Narrows-La Loche Road, Harms
17635; McAneely Creek, 2 mi. s.w. of Turnor Lake village,
Harms 18390; Bear Creek, Mile 41 Buffalo Narrows-La
Loche Road, Harms 19661.

This species has never been listed for Saskatchewan by
Fraser and Russell (1937), Fraser and Russel] (revised by
Russell, Ledingham and Coupland (1954)), or Breitung
(1957). The specimens of the species were for the most
part confused with Sparganium multipedunculatum. While
a taxonomic problem exists between the two taxa, and even-
ually both of them, together with S. angustifolium, may be
merged as varieties under an enlarged S. simplex, there is
no question that many of our specimens best fit S. chloro-
carpum as it is now recognized. Other specimens in the
Fraser Herbarium now identified as S. chlorocarpum in-
clude the following:

Melfort, J. B. Millar 64-163, 64-12; Saskatoon, J. H.
Hudson 2488, 2548, J. B. Millar 65-217, W. P. Fraser 1930;

39

40 Rhodora [Vol. 76

Pike Lake, 20 mi. s. of Saskatoon; W. P. Fraser 1 Sept.
1937, R. C. Russell, et al., 4 August 1937; St. Denis, J. B.
Millar 62-72; W. of Whitefox, W. P. Fraser & G. F. Leding-
ham 28 July 1936; Swift Current, J. L. Bolton 8 July 1936;
Dahlton, W. P. Fraser and. G. F. Ledingham 20 July 1937;
Egg Creek, s. of Cumberland House, G. H. Townsend 55;
Waskesiu, W. P. Fraser 24 July 1938; Lake Waskesiu Road,
W. P. Fraser July 1932; Mile 23 Waskesiu Road to Prince
Albert, W. P. Fraser 24 August 1934; Meadow Lake Pro-
vincial Park, Waterhen River, J. H. Hudson 2427; Sylvan
Lake, Candle Lake Road, J. K. Jeglum 18 July 1964; 1.7 mi.
n. of forks at Candle Lake, J. K. Jeglum 9 July 1964.

CALAMAGROSTIS LAPPONICA (Wahl.) Hartm. 3 mi. s. of
La Loche, Mile 64 Buffalo Narrows-La Loche Road, Harms
17532, 17565; 2 mi. s. of La Loche, Mile 65 Buffalo Nar-
rows-La Loche Road, Harms 17547, 17543; Island in Wil-
liam River, south shore Lake Athabasca, R. Hermesh 485;
Thompson Bay, south shore of Lake Athabasca, R. Hermesh
582.

This northern species has never been previously reported
for Saskatchewan. There seems to be intergradation be-
tween Calamagrostis lapponica and C. neglecta (Ehrh.)
Gaertn. as indicated by an intermediate collection from
Taylor Lake, Mile 19 Buffalo Narrows-LaLoche Road
(18160), and between C. lapponica and both C. neglecta
and C. inexpansa A. Gray as indicated by intermediate
specimens from the same area (18142, 18147). The whole
C. lapponica, C. neglecta, and C. inexpansa complex appears
in need of careful taxonomic study.

CAREX ADUSTA Boott. Turnor Lake, Harms 18364-B,
18361-A.

In the Fraser Herbarium, there are three specimens from
the Waskesiu area or the road to Montreal Lake, Prince
Albert National Park (W. P. Fraser July 8, 1939, June 25,
1940, June 22, 1941). This species has been collected by
J. H. Hudson at Flin Flon, Manitoba, near the Saskatche-
wan border. Breitung (1957) reports it also from Meadow

1974] Saskatchewan Plants — Harms & Hudson 41

Lake but a substantiating voucher is missing from the
Fraser Herbarium. The above record represents a con-
siderable northwestward range extension in Saskatchewan.

CAREX RUFINA Drej. Beach at Thomson Bay, south shore
Lake Athabasca, R. Hermesh 421.

This represents the first report for this species in Sas-
katchewan. Characteristic material of this species comes
from Greenland and Iceland. Scoggan (1957, p. 179) re-
lates that A. E. Porsild collected typical material from
southern Keewatin, Northwest Territories, while Baldwin
found abberrant material at Nueltin Lake, northern Mani-
toba, about 125 miles south of Porsild's station. Porsild
has annotated Baldwin’s sheet as follows: “Differs from
typical C. rufina by paler scales having more prominent
midveins, paler and more prominently nerved perigynia,
and by terminal spikelets being not so strictly gynaecan-
drous (in some even androgynous). Perigynia only rarely
with a few teeth in the upper part." The present Lake
Athabasca material agrees well with the characters men-
tioned in these statements. This collection represents about
a 350 mile westward range extension for the species.

HABENARIA ORBICULATA (Pursh) Torr. Lac La Plonge, 5 mi.
e. of Beauval, Harms 17959-B.

In the Fraser Herbarium there are specimens of this
species from Big Sandy Lake, Hansen Lake Road (G. W.
Argus 4237), from La Ronge (J. S. Maini 611), and from
Pinkney Lake, 35 mi. n.e. of Candle Lake (J. M. A. Swan
65-45). Breitung (1957) also reports it from Torch River
and Amisk Lake, the latter based on a collection by J. H.
Hudson. The present record is the first one from the west-
ern part of the province and represents a considerable west-
ward extension of the main species range. It is disjunctly
present in British Columbia.

SPIRANTHES GRACILIS (Bigel) Beck. (S. lacera Raf.).
Little Amyot Lake area, Mile 69 Green Lake-Buffalo Nar-
rows Road, Harms 17736; 2 mi. e. of Beauval, Harms

42 Rhodora [Vol. 76

17776; Taylor Lake, Mile 19 Buffalo Narrows-La Loche
Road, Harms 18139-B.

There are specimens of this species in the Fraser Her-
barium from Lake Waskesiu, Prince Albert National Park
(Anonymous July 1932) and from 20 miles south of Mea-
dow Lake (A. J. Breitung 8164). The above records am-
plify the known range of the species and extend it to north-
western Saskatchewan.

SPERGULA ARVENSIS L. 2 mi. w. of Beauval, Harms 17846.

Breitung (1957) does not report this plant for Saskatche-
wan, and Scoggan (1957) does not report it for Manitoba.
Boivin (1968) listed this species as doubtful for Saskatche-
wan. Hudson (1972) reported it as new to Saskatchewan
on the basis of a collection from Gronlid (B. Zuk 1971,
DAO). Boivin, in personal correspondence to J. H. Hudson,
indicated the presence of an earlier collection from Sylvania
(K. Drake Juiy 26, 1966, DAS). Mrs. H. D. Bobier recently
sent a collection of the species from Rapid View (west of
Meadow Lake) to J. H. Hudson. A map by J. F. Alex indi-
cates two infestations of this weed near Outlook, Saskatche-
wan but these are not substantiated by voucher specimens.
Thus the above record represents only the fourth report for
Saskatchewan and the only one from this far northwest in
the province.

CALTHA NATANS Pallas. 7 mi. s.-s.e. of La Loche, Mile 60
Buffalo Narrows-La Loche Road, Harms 17633.

There are no Saskatchewan specimens of this species in
the Fraser Herbarium, Breitung (1957) reported it from
Windrum Lake (56°02’N, 104°W) and Amisk Lake, the
latter based on a collection by J. H. Hudson. The present
collection presumably represents the third report for Sas-
katchewan.

RANUNCULUS HYPERBOREUS Rottb. Open beach, William's
Point, south shore Lake Athabasca, R. Hermesh 415.

This represents the first record of this northern butter-
cup species in Saskatchewan.

1974] Saskatchewan Plants — Harms & Hudson 43

MYRIOPHYLLUM ALTERNIFLORUM DC. Little Amyot Lake,
Mile 70 Green Lake-Buffalo Narrows Road.

Breitung (1957) omits this species from his list of Sas-
katchewan plants. Boivin (1968) reports only 3 collections
from the prairie provinces: Cochrane River, Reindeer
Lake, and Lake Axis. The present collection would there-
fore represent only the second Saskatchewan record for the
species.

EUPHRASIA ARCTICA Lane var. DOLOSA Boivin (E. subarctica
Raup, E. disjuncta Fern. & Wieg. var. dolosa Boivin). 2 mi.
s. of La Loche, Mile 65 Buffalo Narrows-La Loche Road,
Harms 17550.

In the Fraser Herbarium, the only Saskatchewan speci-
men of this species is from Stony Rapids, north of Fond-du-
lac River (J. S. Maini & M. Swan Aug. 3, 1961). Breitung
(1957) cites a Raup report from Lake Athabasca. The
above record represents about a 200 mile southward range
extension for the species in Saskatchewan.

LOBELIA DORTMANNA L. Little Amyot Lake, Mile 70 Green
Lake-La Loche Road, Harms 16972.

In the Fraser Herbarium we have specimens of this spe-
cies from Little Gull Lake, south shore Lake Athabasca
(G. W. Argus 553-63) and Carswell Lake, 40 mi. south of
Lake Athabasca (G. W. Argus 628-62). Breitung (1957)
reports the species also from Windrum Lake, north of the
Churchill River, 56°02’N, 104°W. Therefore, the above rec-
ord represents the third general locality for the species in
Saskatchewan.

ADOXA MOSCHATELLINA L. 17 mi. n. of Meadow Lake, J. H.
Hudson 2729 (SASK).

Boivin (1972, p. 5) reports this species in Saskatchewan
from Pasquia Hills and Candle Lake; the latter specimen,
which he collected, was the first record for Saskatchewan.
The present material is evidently the third record of the
species for the province and represents more than a 100
mile westward range extension in the province.

44 Rhodora [Vol. 76

ANAPHALIS MARGARITACEA (L.) B. & H. 3 mi. s. of Buffalo
Narrows, Mile 124 Green Lake-Buffalo Narrows Road,
Harms 18244.

There are various specimens of this species in the Fraser
Herbarium from the Cypress Hills area of Saskachewan.
Breitung (1957) reports it also from Cut Knife but a sub-
stantiating voucher has not been found. Therefore the
above record represents at least a 200 mile, if not a 300
mile northward range extension.

BIDENS BECKII Torr. Little Amyot Lake, Mile 70 Green
Lake-Buffalo Narrows Road, Harms 16978.

There are two specimens of this species in the Fraser
Herbarium from the Cumberland House area on the Sas-
katchewan River (A. E. Etter Aug. 8, 1934; D. Dabbs 126-
66) in east-central Saskatchewan. The above report there-
fore represents the second known Saskatchewan locality for
the species and is a considerable westward range extension
in the province.

LITERATURE CITED

BorviN, B. 1968. Flora of the Prairie Provinces, Part II. Reprinted
from Phytologia 16-18.

1972. Flora of the Prairie Provinces, Part III. Re-
printed from Phytologia 22-23.

BREITUNG, A. J. 1957. Annotated Catalogue of the Vascular Flora
of Saskatchewan. The American Midland Naturalist 58: 1-72.

FRASER, W. P. & R. C. RUSSELL. 1937. An Annotated List of the
Plants of Saskatchewan. University of Saskatchewan, Saska-
toon.

HUDSON, J. H. 1972. 1971 Botanical Records for Saskatchewan.
The Blue Jay 30: 122-124.

RUSSELL, R. C., G. F. LEDINGHAM, & R. T. COUPLAND. 1953. Re-
vision of Fraser, W. P. and R. C. Russell, An Annotated List
of the Plants of Saskatchewan. University of Saskatchewan,
Saskatoon.

SCOGGAN, H. J. 1957. Flora of Manitoba. National Museum of
Canada Bulletin No. 140, Ottawa.

FRASER HERBARIUM
UNIVERSITY OF SASKATCHEWAN
SASKATOON, SASKATCHEWAN, CANADA

PARASITIC WITCHWEED: STRIGA ASIATICA
VERSUS S. LUTEA (SCROPHULARIACEAE)

F. N. HEPPER

When plants of a witchweed were discovered in 1956
parasitizing Zea mays L. in North and South Carolina,
specimens were sent to Kew for identification. I determined
them as Striga asiatica (L.) O. Kuntze and communicated
the name to Dr. S. F. Blake and other authorities in the
United States, who used it until a paper by Dr. C. J.
Saldanha (1963) proposed the rejection of that name in
favour of S. lutea Lour. (Smith 1966). The purpose of this
paper is to reinstate S. asiatica on the basis of typification
of the Linnean basionym Buchnera asiatica, rather than
on the opinions of other taxonomists.

Linnaeus’ description of Buchnera asiatica in ‘Species
Plantarum' (1753) is unusually full for that work and
entirely devoid of any references to literature. This indi-
cates that he had before him a specimen, or specimens,
from which he drew up his description. Examination of
his herbarium at the Linnean Society, London, revealed
six sheets bearing the name ‘Buchnera asiatica! in Linnaeus’
hand and now numbered according to Savage's Catalogue
as 790/10, 11, 12, 13, 14, & 15.

Sheets 13, 14 and 15 can be discounted, since they are
annotated as having been collected by Thunberg (13) some-
time after 1772, and by Koenig (14 & 15) in 1777 according
to the date on the sheet. They are referable to Striga
bilabiata (Thunb.) O. Kuntze (syn. S. thunbergii Benth.)
and S. euphrasioides Benth., respectively.

Sheet 12 bears a single simple plant with pale flowers
that would be known in India as Striga lutea. The only
annotation on the sheet is ‘Indica,’ indicating its country
of origin. The specimen does not fit Linnaeus' description,
and it can be ruled out as the type.

Five specimens are glued to sheet 10, including a well-
branched plant with dark-coloured flowers that fits the

45

16 Rhodora [Vol. 76

original description admirably. Two other specimens match
it except for their simple habit. The remaining two are
only the inflorescences of Striga densiflora Benth. The
specimen on sheet 11 is very similar to the large branched
one on sheet 10, but there is no annotation apart from the
name. On sheet 10, however, Linnaeus has added the
locality “ins. Johan.," and although the ‘Species Plantarum’
cites the distribution as “Habitat in Zeylona, China” this
reference to “ins. Johan.” provides a clear indication of the
provenance and history of what must be regarded as the
type material of Buchnera asiatica. Where, then, is “ins.
Johan.” and who collected the specimen?

Several voyages to China were undertaken before 1753.
including one by Peter Osbeck who wrote an account of
it in Swedish (1757) which was translated into German
(1765) and from the German edition into English, and
published in 1771 under the title ‘A voyage to China and the
East Indies, together with a voyage to Suratte by Olof
Toreen in a series of letters to Doctor Linnaeus.’ Toreen
(or Torén), in fact, continued his voyage as far as China
and it is his letters, and not Osbeck’s text, which are of
particular interest since he tells (p.166) how, on 16 August
1750, he landed at the island of “St. Joanna” in the Comoros
near Madagascar. It is evident that Linnaeus worked out
the collections made by Toreen and Osbeck, attributing
them al] to China in the appendix to Osbeck and Toreen’s
travels entitled ‘Flora Sinensis.’ There, on p. 356, is listed
Buchnera asiatica with the reference to ‘Species Plantarum.’
It is interesting to note that the same island was visited by
Koenig in 1768 where he gathered the plant Linnaeus
(1771) described as Baccharis arborea (which Brenan
(1968) identified as Vernonia grandis (DC.) Humbert var.
comorensis Humbert), yet Linnaeus gave the occurrence as
the “East Indies.”

Striga asiatica (L.) O. Kuntze, Rev. Gen. Pl, 2: 466. 1891.
Type. Comoro Islands, St. Joanna, Toreen LINN. (Sav-
age Cat. No. 790/10 partly).

1974] Striga — Hepper AT

Buchnera asiatica L., Sp. Pl. 630. 1753.

Striga lutea Lour., Fl. Cochinch. 22. 1740.

Buchnera hirsuta Benth., Scroph. Ind. 41. 1835.

Striga hirsuta, (Benth.) Benth. in Hook., Comp. Bot. Mag.
1: 363. 1836.

Bentham's work was completed long before an interna-
tional code of botanical nomenclature had been agreed, and
certainly his concepts of typification were not in accord
with modern attitudes. Hence, he coined the new names
to replace Buchnera asiatica which seemed to him, as to
Saldanha, to apply to a number of different species. The
more so since he considered S. lutea Lour. to be a distinct
species.

Saldanha considered Striga asiatica and its basionym as
nomena rejicienda under Art. 69 since “any attempt to
decide a lectotype is bound to be arbitrary and open to
question." On the contrary, this species is one of the few
Linnean species that may be typified and localized with
certainty.

I am grateful to my colleague Mr. R. D. Meikle for his
helpful advice and assistance.

LITERATURE CITED

BRENAN, J. P. M. 1968. The identity of Baccharis arborea L.
Kew Bull. 21: 427.

LiNNAEUS, C. 1753. Species Plantarum, 630.

1771. Mantissa Plantarum, 284.

OsBECK, P. 1771. A voyage to China and the East Indies. (English
ed.).

SALDANHA, C. J. 1963. The genus Striga Lour. in Western India.
Bull. Bot. Surv. India 5 (1) : 67-70.

SMITH, C. EARLE. 1966. Identity of witchweed in the southeastern
United States. Rhodora 68 (774) : 167.

THE HERBARIUM
ROYAL BOTANIC GARDENS
KEW, ENGLAND

CACALIA RUGELIA: A NEW COMBINATION FOR
A NORTH AMERICAN SENECIONOID*

T. M. BARKLEY AND ARTHUR CRONQUIST

The problematic taxon long-called Senecio rugelia (Shuttl.
ex Chapman) A. Gray (Compositae) is a distinctive species
occurring in coniferous associations along the higher ridges
of the Great Smokies astride the North Carolina-Tennessee
border. It is a tall, coarse herb with several conspicuous
discoid heads and dull, dirty-white corollas. The species
was introduced to science as Rugelia nudicaulis Shuttl. ex
Chapman in 1860, but in 1883 Asa Gray transferred it to
Senecio, where it has remained.

Many botanists have recognized the incongruity of in-
cluding this entity in Senecio, but the morphological toler-
ances of Senecio have been usually treated as very wide,
and so the matter has not been pursued. However, Green-
man (1901) and Alexander (1937) noted the possible
relationship to Cacalia.

The species referable to Cacalia have had their systematic
woes, being variously regarded as constituting several gen-
era, or as members of the single genus Cacalia, or as a part
of Senecio. Compounding these difficulties has been a dis-
agreement concerning the typification of the name Cacalia.
A review of these matters is beyond the scope of this paper,
but we are treating Cacalia in the expanded sense, and we
accept Cacalia hastata L. as the generic type (Vuilleumier
& Wood, 1969).

In general, Cacalia differs from Senecio by the two floral
features of discoid heads and white to creamy corollas.
Both are features which occur independently as evolution-
ary tendencies in other alliances within Senecio. In Cacalia,
however, the two occur together, presumably defining a

natural assemblage.

*Contribution no. 1193. Division of Biology, Kansas Agr. Expt.
Station, Manhattan. Financial support from the National Science
Foundation is gratefully acknowledged.

48

1974] Caealia — Barkley and Cronquist 49

Cacalia is here regarded as a derivative of Senecio section
Palmatinervii of Mexico, a group of coarse herbs strongly
resembling Cacalia in aspect. Nearly all species of the
Palmatinervii for which chromosome counts are reported
have n=30 (Gibson, 1969). Cacalia has n—30, and a re-
duction series of n=28, 27, 26, 25, and 20 (Ornduff et al.,
1963, 1967). The species here offered as Cacalia rugelia
has »—28, a number unreported for Senecio (Fedorov,
1969).

In a recent study of Mexican cacalioids, Pippen (1968)
recognizes four genera, but he offers a table summarizing
the distinctions between these ‘“‘Cacalioid genera” and Sene-
cio (table 2, p. 371). Incorporated into the table are the
cacalioid features of discoid heads and white or creamy
florets. The table also shows that the *Cacalioid genera"
characteristically have deeply lobed disk corollas, a feature
which we find in Cacalia rugelia as well. Furthermore, in
Pippen's key the present species falls into the assemblage
treated as the Mexican segregate-genus Odontotrichum, and
it is morphologically compatible with that group. The
floristic relationships between the Southern Appalachian
region and some of the mountains of Mexico have been
noted by other authors (e.g. Sharp, 1946).

In the light of the foregoing considerations, the following
transfer is proposed:

Cacalia rugelia (Shuttl. ex Chapman) Barkley & Cronquist,

comb. nov.

Type: “Smoky Mountains, Tennessee. Rugel, Buckley."
Paratype: Buckley, NY!

Senecio rugelia A. Gray, Proc. Am. Acad. 19: 54. 1885.

Rugelia nudicaulis Shuttleworth ex Chapman, FI. S. U. S.
246. 1860.
non Cacalia nudicaulis (Less.) O. Kuntze, 1891.
non Senecio nudicaulis Buck.-Ham. ex D. Don, 1825.

LITERATURE CITED
ALEXANDER, E. J. 1937. Senecio rugelia. Addisonia 20: 29-30,
pl. 655.

50 Rhodora [Vol. 76

FEDOROV, AN. A., ed. 1969. Chromosome numbers of flowering
plants. Soviet Sciences Press. Leningrad.

GIBSON, E. S. 1969. A revision of the section Palmatinervii of the
genus Senecio (Compositae) and its allies. Unpub. doctoral dis-
sertation, Kansas State Univ. Manhattan. pp. 1-178 + VI.

GREENMAN, J. M. 1901. Monogr. nord-und centralam. Arten der
Gattung Senecio. Leipzig. (Reprinted in Englers Bot. Jahrb. 32:
1-33. 1902.)

ORNDUFF, R. P. H. Raven, D. W. KvHos & A. R. KINCHEBERG.
1963. Chromosome numbers in Compositae III. Senecioneae.
Am. Jour. Bot. 50: 131-139.

OnNDUFF, R., T. Mosquin, D. W. KvHos & P. H. RAVEN. 1967.
Chromosome numbers in Compositae VI. Senecioneae II. Am.
Jour. Bot. 54: 205-213.

PIPPEN, R. W. 1968. Mexican “Cacalioid” genera allied to Senecio
(Compositae). Contr. U. S. Natl. Herb. 34: 365-447.

SHARP, A. J. 1946. Informe preliminar sobre alqunos estudios
fitogeograficos efectuados en Mexico y Guatemala. Rev. Soc.
Mexicana Hist. Nat. 7: 35-39. figs. 1-5.

VUILLEUMIER, B. S. & C. E. Woop. 1969. Lectotypification of Caca-
lia L. (Compositae-Senecioneae). Jour. Arnold Arb. 50: 268-275.

HERBARIUM, DIVISION OF BIOLOGY,
KANSAS STATE UNIVERSITY, MANHATTAN, KANSAS

THE NEW YORK BOTANICAL GARDEN,
BRONX, NEW YORK

TYPIFICATION AND APPLICATION OF THE
NAMES SCIRPUS AMERICANUS PERS.,
S. OLNEYI GRAY, AND S. PUNGENS VAHL

ALFRED E. SCHUYLER

The names Scirpus americanus Pers. (Syn. Pl. 1: 68.
1805) and Scirpus pungens Vahl (Enum. Pl. 2: 255. 1805)
are usually applied to the same species by Cyperologists
(Beetle, 1947; Koyama, 1963). The former name for plants
commonly known as Three-square in North America pres-
ently receives widespread usage because of priority. Data
from type specimens and original descriptions now make it
apparent that these names should be applied to different
species. The type of S. americanus (Hab. in Carolina in-
feriore, Herb. Michx., P) is conspecific with plants usually
treated (Beetle, 1947; Koyama, 1963) as S. olneyi Gray
(Bost. Journ. Nat. Hist. 5: 238. 1845). The type of S.
pungens (Erhart misit, Herb. Vahl, C) is conspecific with
plants usually treated (op. cit.) as S. americanus.

In his original description of Scirpus americanus, Per-
soon cites Michaux's description of S. triqueter L. (Fl. Bor.-
Am. 1: 30. 1803.) and gives the habitat as "in Carolina
inferiore." The specimens on the sheet thus named and
labeled in Michaux's herbarium at P have strongly winged
culms, short involucral bracts, shallowly cleft scales, small
achenes, and broad bristles characteristic of plants resem-
bling the type of S. olneyi (Rhode Island, Providence, Olney,
GH). Thus the correct name for plants usually treated as
S. olneyi is S. americanus.

In the original description of Scirpus pungens, Martin
Vahl gave the habitat as, *in Europa, in insula Borboniae.
Commerson." The specimen from Vahl's herbarium desig-
nated as the type probably was collected in Europe and also
contains the word “pungens Vahl" presumably in the hand-
writing of Jens Vahl, son of Martin Vahl. No specimen
labeled, *in insula Borboniae," or collected by Commerson
has been located. The triangular culms, long involucral

51

52 Rhodora [Vol. 76

bracts, deeply cleft scales, and narrow bristles are char-
acteristics of plants usually but incorrectly treated as S.
americanus. Scirpus pungens is the earliest available name
for these plants.

The literature cited by Vahl is also in accordance with his
description and the designated type specimen. The cited
illustrations of Morrison (Pl. Hist. Univ. Oxon. s. 8, t. 10,
f. 20. 1699.) and Plukenet (Algm. Bot. t. 40, f. 1. 1696.)
closely resemble the type specimen of Scirpus pungens.
Roth's descriptions of S. mucronatus L. (Tent. Fl. Germ.
1: 28. 1788, and 2: 60. 1789.) and S. triqueter L. (Neue
Beytr. 1: 91. 1802.) cited by Vahl best apply to plants
resembling the type specimen of S. pungens. Apparently
Roth was confused about the identity of S. mucronatus and
S. triqueter, and incorrectly applied the names to plants of
S. pungens.

ACKNOWLEDGMENTS

I am grateful to H. Heine (P), J. Raynal (P), and A.
Skovsted (C) for courtesies enabling me to examine critical
specimens and data at their respective institutions, and to
John Braxton (PH) and Helena Greenwood (PH) for their
assistance. Abbreviations of institutions are those given
by Lanjouw and Stafleu (1964).

LITERATURE CITED

BEETLE, A. A. 1947. Scirpus. N. Am. Fl. 18: 481-504.

KovAMA, T. 1963. The genus Scirpus Linn., critical species of the
section Pterolepis. Canad. Jour. Bot. 41: 1107-1131.

LANJOUW, J. and F. A. STAFLEU. 1964. Index Herbariorum. Part 1.
The herbaria of the world, International Association for Plant
Taxonomy, Utrecht. 251 pp.

DEPARTMENT OF BOTANY
ACADEMY OF NATURAL SCIENCES OF PHILADELPHIA
PHILADELPHIA, PENNSYLVANIA 19103

THE CORRECT AUTHORITY FOR
CARDAMINE CLEMATITIS (CRUCIFERAE)

T. R. DUDLEY!

The most recent entry in the literature of Cardamine
clematitis appears on page 508 of the Manual of the Vas-
cular Flora of the Carolinas by Radford, Ahles and Bell
(1968). They cite “Shuttlew.” as the authority for this
taxon. This authority refers to Robert James Shuttle-
worth, a famed naturalist and collector who was the spon-
sor of Ferdinand Rugel, a prolific plant collector in eastern
North America from 1840 and for many years thereafter
until his death in 1879.

The various references in the literature to Cardamine
clematitis have consistently overlooked the fact that Shut-
tleworth cannot be cited as the sole publishing authority for
the specific epithet, clematitis. In addition to Radford,
Ahles and Bell (1968), several other earlier North Ameri-
can references also provided descriptions and occasionally
illustrations of C. clematitis, and all accepted Shuttleworth
as the sole publishing authority. These include: Chapman,
Flora of the Southeastern United States, ed. 2, Supplement
p. 605. 1887; ibid., ed. 3, p. 25. 1897; Britton & Brown,
Illust. Flora of the Northern United States and Canada, ed.
1. 2: 130. fig. 1730. 1897; ibid., ed. 2. 2: 185. fig. 2088.
1913; Small, Flora of the Southeastern United. States, ed. 1,
p. 482. 1903; ibid., ed. 2, p. 568. 1933; Fernald in Gray's
Manual of Botany, p. 721. 1950; Gleason and Cronquist in
Britton and Brown, Illust. Flora of the Northeastern United
States and Adjacent Canada 2: 230, fig. p. 231. 1952;
Gleason & Cronquist, Manual of Vascular Plants of North-
eastern United States and Adjacent Canada, p. 340. 1963;
and Radford, Ahles and Bell, Guide to the Vascular Flora
of the Carolinas, p. 173. 1964.

‘Research Botanist, Herbarium, U.S. National Arboretum, Agricul-

tural Research Service, Northeastern Region, U.S. Department of
Agriculture, Washington, D.C. 20002.

53

54 Rhodora [Vol. 76

In the course of evaluating the Ferdinand Rugel collec-
tions in the Isaac C. Martindale Herbarium at the U.S.
National Arboretum, F. G. Meyer called my attention to an
unidentified specimen of Cardamine collected by Rugel and
accompanied by a signed holograph label that reads “Top
of Smoky Mts. on the first Branch Nord Carolina Side.
May 1866." Although this Rugel specimen clearly repre-
sents C. clematitis, it is not a part of the type.

The first published reference to Cardamine clematitis is
on page 53 of Sereno Watson's Bibliographic Index to
North American Botany (Smithsonian Miscellaneous Col-
lections No, 258, 1878) that reads as follows: “C. clematitis
Shuttl. in Herb. Gray, ined." It is evident that Watson took
the name from Shuttleworth’s printed herbarium exsiccata
label which reads “Cardamine clematitis Shuttle. n. sp.”
and that name was used only as a nomen nudum. Watson’s
reference does not incorporate a validating description. On
this basis alone, clematitis cannot be used as a valid specific
epithet. It is listed, however, in Index Kewensis 1: 421.
1895 as: “Clematitis Shuttle. ex S. Watson" !

Later, in a paper by Asa Gray entitled “Some New
North American Genera, Species, &c." (Proc, Am. Acad.
Arts Sci. n.s., 7:45. 1880, a valid Latin description is pro-
vided. Gray precisely identified the plant as “Cardamine
clematitis Shuttleworth in coll. distrib. Rugel," and explains
that the original collection was from “wet ground along
streamlets in the higher Iron or Smoky Mountains of North
Carolina and Tennessee, collected in 1844 by Rugel .. .”

When dealing with nomenclatural problems of this na-
ture, we are guided by Article 46 and its accompanying
recommendations (particularly Recommendation 46C) in
the International Code of Botanical Nomenclature, p. 46.
1972. 'The correct authority citation for Cardamine clema-
titis is C. clematitis Shuttleworth ex A. Gray, although it
may be shortened, particularly in floristic treatments, to
cite only the publishing author. Under no circumstances
can Shuttleworth be assigned solitary authorship. The cor-
rect citation of the name is:

1974] Cardamine clematitis — Dudley 55

Cardamine clematitis Shuttleworth ex A. Gray, Proc.

Amer. Acad. Arts Sci, new series, 7: 45. 1880 — non

Shuttl. ex Watson, Bibliog. Index North American Bot.,

p. 53, 1878.

The original collection and designated type of Cardamine
clematitis is Rugel No. 19 collected in 1844 in the Smoky
Mts. of Tennessee. The holotype is deposited at GH, and
cited duplicates (isotypes) are to be found at G-Herb. Bois-
sier and W.

TYPE: Tennessee, Smoky Mountains [‘‘in locis humidis
et ad regionis super. montium Smoky Ms., Tennessee, Mai
1884.”] May 1844, Ferdinand Rugel No. 19 (holotype, GH;
isotypes BM, G, W).

The most complete set of Rugel’s material, purchased from
Shuttleworth, is at the British Museum; however, reference
to the existence of such a specimen has not previously ap-
peared in the literature. N. K. B. Robson of the British
Museum (Natural History) assures me that a duplicate is
retained at BM.

The one Rugel collection of Cardamine clematitis not
previously reported in the literature is a specimen collected,
probably a unicate, after Rugel’s professional contacts with
Shuttleworth had ceased. The specimen was maintained in
Rugel’s personal herbarium until that herbarium was pur-
chased in 1881 from Rugel’s heirs by Isaac C. Martindale:
North Carolina: “Top of Smoky Mts. on first Branch, Nord
Carolina Side", Ferdinand Rugel, May 1866 (NA — from
herbarium of Isaac C. Martindale).

The astute monographer of Cardamine and indefatigable
expert on the Cruciferae, D. E. Schulz, in Monographie der
Gattung Cardamine” (Bot. Jahrb. 32: 440. 1903.) recog-
nized Shuttleworth as responsible for the epithet clematitis,
but also recorded that Asa Gray provided a description of
the species in Proc. Am. Acad. Arts Sci. n.s. 7: 45. 1880.
Schulz examined the original 1844 material of C. clematitis
collected by Rugel (GH, G & w — but not BM), which was
annotated and distributed by Shuttleworth; he also cited
numerous additional collections made by J. K. Small, N. L.

56 Rhodora [Vol. 76

& E. G. Britton & A. M. Vail, S. B. Buckley, J. K. Small &
A. A. Heller, W. M. Canby, M. E. Hyams and J. W. Chicker-

Ing.

SYNONYMS OF CARDAMINE CLEMATITIS

A point should also be discussed that concerns the synon-
omy of Cardamine clematitis Shuttleworth ex A. Gray as
presented by the Radford, Ahles and Bell publications
(1964 & 1968). These works refer “C. flagellaris” (= C.
flagellifera Schulz, Bot. Jahrb. 32: 405. 1903.) into synon-
omy under C. clematitis. Although the type specimen of
C. flagellifera (Biltmore Herbarium 7756) was originally
annotated as C. clematitis, it is not conspecific with C.
clematitis. Not only does C. flagellifera stand morphologi-
cally and ecologically distinct from C. clematitis, but it was
also assigned by Schulz within Cardamine to Sect. Macro-
phyllum, whereas C. clematitis was referred by Schulz to
the typical section. Small (1903) described C. hugeri that
Radford, Ahles & Bell (1964 and 1968) sank under C.
clematitis. However, if referrable into synonomy at all,
C. hugeri, a low altitude plant, more logically pertains to
C. flagellifera.

In Watson’s reference (loc. cit.) to Cardamine clematitis,
he cites as a synonym a “Nasturtium officinale” that ap-
peared in the Supplement to Torrey and Gray, A Flora of
North America 1: 666. 1843. In addition to validating,
describing and typifying C. clematitis, Gray (loc. cit.)
explains Watson’s confusion in incorrectly citing this
“Nasturtium officinale” as a synonym of C. clematitis. The
original Rugel material, annotated and distributed from
Switzerland by Shuttleworth, was a mixed collection. A
part representing C. clematitis “was mixed up with a Flor-
ida species intermediate between Cardamine and Nastur-
tium, first received from Leavenworth without fruit, and
referred in the supplement to the first volume of Torrey and
Gray's Flora to N. officinale.” This element of “Nasturtium
officinale”, according to Gray, was later received from S. B.

1974] Digitaria sanguinalis — Ebinger 57

Buckley; again later received from Shuttleworth’s dis-
tributed collections of Rugel as Cardamine curvisiliqua
Shuttleworth; and yet again received from Shuttleworth as
Nasturtium stylosum Shuttleworth! The current fate of
Leavenworth’s “Nasturtium officinale’ is not within the
scope of this paper; however, both Chapman (1887) and
Small (1903) equate Leavenworth’s “Nasturtium officinale”
with Cardamine curvisiliqua Shuttleworth.

U.S. NATIONAL ARBORETUM
WASHINGTON, D.C. 20002

A NEW FORM OF DIGITARIA SANGUINALIS. As
a result of extensive field work in east-central Illinois an
unusual form of the common crabgrass was found. This
form differs from typical Digitaria sanguinalis (L.) Scop.
in that long, spreading, papillose-based hairs are found
scattered along both sides of the flattened rachis. These
colorless hairs are 3-6 mm long (rarely 1 cm long) and on
the specimens examined usually 1 to 5 hairs are found on
each cm of rachis length. As a result of this difference the
following form is described.

Digitaria sanguinalis (L.) Scop. forma illinoensis Ebinger
forma nova.

A forma sanguinalis differt pilis base papillosis in rhachidi.
TYPE: ILLINOIS: DOUGLAS CO.: 4 miles east of Hindsboro,
Sargent Twp. (NE!4, Sect. 3, RIOE, T14N), in open field
at edge of road, J. E. Ebinger 6845 (EIU). This area was
revisited on 9 October 1969 and a second collection (J. E.
Ebinger 9282) was made at that time (EIU, ISM).

Long, papillose-based hairs are known in a few species
of Digitaria. The presence of these hairs is an important
diagnostic characteristic used by Hitchcock (1935), Hen-
rard (1950), and many others to separate the tropical Digi-
taria horizontalis Willd. from other members of the genus.
The hairs in this species are similar in all respects to those

58 Rhodora [Vol. 76

found in D. sanguinalis f. illinoensis. The two taxa are
easily separated however, since in D. horizontalis the nearly
glabrous spikelets are usually less than 2.4 mm long, ex-
tremely narrow and with an acuminate apex while D. san-
guinalis has pubescent spikelets that are longer (2.5-3.5 mm
long) and broader and with an acute apex. The only other
closely related taxon in which these hairs have been re-
ported is D. adscendens (HBK) Henrard var. rhachiseta
Henrard. The differences between this taxon and D. san-
guinalis have been studied by Ebinger (1962, 1965).

LITERATURE CITED

EBINGER, J. E. 1962. Validity of the grass species Digitaria
adscendens. Brittonia 14: 248-253.

. 1965. Digitaria sanguinalis in South America.
Trans. Ill. St. Acad. Sei. 58: 255-258.

HENRARD, J. T. 1950. Monograph of the genus Digitaria. xii + 999
pp. Leyden, Universitare Pers Leiden.

Hitcucock, A. S. 1935. Manual of the grasses of the United
States. U. S. Dept. of Agric. Misc. Publ. 200. 1040 pp. (Ed. 2,
revised by Agnes Chase, 1950).

JOHN E. EBINGER
EASTERN ILLINOIS UNIVERSITY
CHARLESTON, ILLINOIS

THE GROWTH OF SOME NEW ENGLAND
PERENNIAL SEAWEEDS

NORMAN B. REYNOLDS

Growth studies on Fucus vesiculosus L., Ascophyllum
nodosum (L.) LeJolis, Rhodymenia palmata (L.) Greville,
Laminaria digitata (L.) Lamouroux and Laminaria sac-
charina (L.) Lamouroux were initiated at Dover Point,
Dover, New Hampshire and Jaffrey Point (Fort Stark),
Newcastle Island, Portsmouth, New Hampshire in July
1968. Dover Point is an estuarine tidal rapid area whereas
Jaffrey Point is a semi-exposed open coastallocation. Both
areas have been studied extensively and much is known
of their flora and ecology (Mathieson, Hehre & Reynolds,
in press; Mathieson, Reynolds & Hehre, in press; Reynolds,
1971).

Approximately 20 plants of each species were tagged at
both locations with plastic forester's tape and small white
tags. The monthly growth of F. vesiculosus, A. nodosum
and R. palmata was recorded by measuring their total
length, while in L. digitata and L. saccharina growth was
recorded by determining the monthly increase in distance
between the intercalary meristem and paired holes (Sun-
dene, 1964). The holes were made with a cork borer 1 inch
above the meristem. New holes were made every 2 months,
because the terminal portions of the lamina sloughed off
continuously. Specimens of L. saccharina, L. digitata and
R. palmata were transplanted from the high to the low
current areas at Dover Point, and their growth and
longevity was recorded. During the 16 months of the ex-
periment, nearly all the initial plants were lost due to
human interference, grazing of snails, wave action or
degeneration. Consequently, it was necessary to continu-
ously tag and measure new plants.

The growth of Rhodymenia palmata was sporadic in the
areas of low and high currents at Dover Point. Frag-
mentation occurred extensively during the late winter, The

59

60 Rhodora [Vol. 76

area of low currents was not suitable for the sustained
growth of R. palmata, for when plants were transplanted
there they died in a few months. All specimens of R.
palmata grew at least one inch during June 1968.

The best growth of Fucus vesiculosus and Ascophyllum
nodosum occurred during the summer while the poorest
growth occurred during cold weather. The growth of F.
vesiculosus and A. nodosum at Jaffrey Point was consistently
lower than at Dover Point. The maximum growth of
F. vesiculosus was 2 inches per month, and it occurred
from May 15 to August 15, 1968, in the area of low cur-
rents at Dover Point.

Figures 1 and 2 show the growth of Laminaria digitata
and L. saccharina. Because of the extreme variability in
growth rates between individual plants, these rates are
described only in relative terms as follows:

(1) good growth — over 3 inches/month
(2) average growth — 2-3 inches/month
(3) poor growth — up to 2 inches/month.

The growth of Laminaria digitata in the low current
areas at Dover Point was quite variable. When trans-
plants were made from the high to the low current areas,
the growth was initially high, but the plants eventually
degenerated. The probable cause is siltation and the physi-
ological decrease of oxygen and nutrients due to the de-
creased currents. Good growth of L. digitata occurred in
the spring and summer at Jaffrey Point and in the area
of high currents at Dover Point. Poor growth was particu-
larly evident in the winter at the same locations. In general,
L. digitata had a higher rate of growth and it was sustained
for a longer time in the high current areas at Dover Point
than at Jaffrey Point.

An interesting phenomenon was observed when L. digi-
tata was transplanted from the high to the low current
areas at Dover Point. No dissection of the newly formed
parts of the blades took place, and in a few months it was
difficult to distinguish the plants from L. saccharina, ex-
cept for the identification tags. The mechanical force of

1974] Seaweeds — Reynolds
H è
5 3
B & 3 z £
3 ii ii

RELATIVE GROWTH OF LAMINARIA DIGITATA

61

1- 1- | ee
s d ;
v 2 o 3 E $ 3 $
$ i d oi i $ d
H1MO39
Figure 1
RELATIVE GROWTH OF Laminaria digitata
LEGEND:
Poor Growth 0-2 inches per month
Average Growth 2-3 inches per month

Good Growth 38-54% inches per month

poor

62

RELATIVE GROWTH OF LAMINARIA SACCHARINA

Rhodora

high current area
low current area

=
à
b
[j]
»
o
O

Jaffrey tt
Dover Pt.

[Vol.

T T T "CT |
EN . , ¢ ,
i1 : i i | i o: OB
HLMOYS
Figure 2

RELATIVE GROWTH OF Laminaria saccharina
LEGEND:
Poor Growth 0-2 inches per month
Average Growth 2-3 inches per month
Good Growth 3-5 % inches per month

6

1974] Seaweed — Reynolds 63

the current was not high enough to cause the final separa-
tion of the frond segments.

The growth rate of L. saccharina was good for most of
the year in the low current areas at Dover Point. A slight
decrease in growth occurred during the winter. Good
growth of L. saccharina occurred during the summer and
early fall at Jaffrey Point and in the high current areas
at Dover Point. Poor growth was evident during the late
fall and early winter.

Laminaria saccharina exhibited good growth at all areas
during the summer. Laminaria saccharina dominated the
kelp beds at Dover Point during the winter, but L. digitata
was dominant the rest of the year. It can be concluded
that L. saccharina is better adapted to the estuarine en-
vironment than is L. digitata. The penetration of L. sac-
charina into the estuary is dependent on the presence of
currents to compensate for the reduced salinities.

LITERATURE CITED

MATHIESON, A. C., E. HEHRE & N. B. REYNOLDS. Investigations of
New England marine algae I. A floristic and descriptive eco-
logical study of the marine algae of Jaffrey Point, New Hamp-
shire. Nova Hedwigia, In press.

MATHIESON, A. C., N. B. REYNOLDS & E. HEHRE. Investigations of
New England marine algae. II. Distribution of benthonic marine
algae in the Great Bay Estuary System. Nova Hedwigia, In
press.

REYNOLDS, N. B. 1971. The ecology of a New Hampshire estuarine
tidal rapid. Ph.D. Thesis, University of New Hampshire Li-
brary, 101 pp.

SUNDENE, O. 1964. The ecology of Laminaria digitata in Norway
in view of transplant experiments. Nytt Mag. Bot. 11: 83-107.

DEPARTMENT OF BIOLOGICAL SCIENCES
STATE UNIVERSITY OF NEW YORK
COLLEGE AT CORTLAND

CORTLAND, NEW YORK 13045

CHROMOSOME NUMBERS IN KOSTELETZKYA
PRESL (MALVACEAE)

ORLAND J. BLANCHARD, JR.

Heretofore, knowledge of the cytology of Kosteletzkya
Presl has consisted of a single chromosome count from
K. hastata Presl (Skovsted, 1941). This note corrects the
number erroneously reported by Skovsted and adds counts
from two African species, K. adoensis (Hochst. ex A. Rich.)
Mast., and K. buettneri Gürke, and four American species,
K. coulteri A. Gray, K. pentasperma (Bertero ex DC.)
Griseb. K. paniculata Benth., and K. virginica (L.) Presl
ex A. Gray (Table 1).

Counts were made from pollen mother cells of buds col-
lected in the wild or taken from plants grown from seeds
collected in the wild. Methods of fixation, preparation, and
documentation are those reported by Bates and Blanchard
(1970). Voucher specimens have been deposited in the
L. H. Bailey Hortorium (BH).

The species counted represent a rather wide range of
morphological diversity in the genus as well as a con-
siderable geographical range. Kosteletzkya paniculata be-
longs to the specialized sect. Orthopetalum Benth. of
western Mexico, which is notable in its convolute, tubular
corolla and its exserted staminal column. The remaining
species, both African and American, belong to the wide-
spread sect. Kosteletzkya, but within that section repre-
sent a diversity of forms. The fact that counts made from
all these species are the same, n = 19, suggests that the
number in the genus may be constant, At variance with
this conclusion is Skovsted's report of 2n = 34 in K. has-
tata, a species closely related to K. pentasperma. How-
ever, an examination of the illustration by Skovsted (1941,
fig. 99) shows 38 chromosomes rather than the reported
34. It may therefore be assumed that the gametic chromo-
some number of K. hastata is also » — 19.

64

1974] Kosteletzkya — Blanchard 65

The error that entered Skovsted's paper is of further
interest because he used the erroneous count to support
Hochreutiner's suggestion (1900) that Kosteletzkya could
be related to Hibiscus sect. Pterocarpus Garcke ex
Hochreutiner, a section known cytologically only from
H. vitifolius L. with a chromosome number of n = 17
(Skovsted, 1935, 1941). There are ample reasons on
morphological grounds for relating Kosteletzkya and
Hibiscus sect. Pterocarpus, e.g., they share depressed,
pentagonal, 5-crested capsules, the valves of which fall
away completely from the floral axis at maturity, as well
as similarities in calyx, involucre, and vegetative char-
acters. Now, however, any proposal to unite the two taxa
must take into account the difference in chromosome
numbers.

Acknowledgments

Field collections were aided by funds from National
Science Foundation Grant GB-8759 to David M. Bates.
I also wish to thank Paul A. Fryxell for providing trans-
portation in Mexico.

LITERATURE CITED

Bates, D. M., & O. J. BLANCHARD, JR. 1970. Chromosome numbers
in the Malvales. II. New or otherwise noteworthy counts rele-
vant to classification in the Malvaceae, tribe Malveae. Amer.
J. Bot. 57: 927-934.

HOCHREUTINER, B. P. G. 1900. Revision du genre Hibiscus, An-
nuaire Conserv. Jard. Bot. Genéve 4: 23-191.

SKOVSTED, A. 1935. Chromosome numbers in the Malvaceae. I.
J. Genet. 31: 263-296.

1941. Chromosome numbers in the Malvaceae. II.

C. R. Lab. Carlsb. Physiol. 23: 195-242.

66 Rhodora [Vol. 76

Table 1. Chromosome counts in Kosteletzkya

Species n-number Locality and Collector
K. adoensis 19 Cultivated, BH-69:555. Seed
source — Angola: Huambo

Distr., Instituto de Investigação
Agronómica. de Angola

K. buettneri 19 Cultivated, BH-72:100. Seed
source — Zambia: “C Pro-
vince," Robinson 6706
K. coulteri 19 Mexico: Sinaloa, Fryxell, Bates
& Blanchard 1544
K. pentasperma 19 Mexico: Jalisco, Fryxell, Bates
& Blanchard 1610
19 Mexico: Michoacan, Fryzell,

Bates & Blanchard 1650
19 Mexico: Nayarit, Fryxell, Bates
& Blanchard 1563

K. paniculata 19 Cultivated, BH-71:119. Seed
source — Mexico: Jalisco,
Fryxell, Bates & Blanchard
1590

K. virginica 19 USA: Florida, Collier Co.,

Blanchard & Blanchard 303
19 USA: New Jersey, Ocean Co.,
Blanchard & Blanchard 284
19 Cultivated, BH-69:430. Seed
source — USA: Texas, Jeffer-
son Co., Knight 88

L. H. BAILEY HORTORIUM
CORNELL UNIVERSITY
ITHACA, NEW YORK 14850

NAME CHANGES FOR THE SEED PLANTS
IN THE BAHAMA FLORA

WILLIAM T. GILLIS

The author has been working in collaboration with
Richard A. Howard of the Arnold Arboretum and George
R. Proctor of the Institute of Jamaica towards a revision
of Britton and Millspaugh’s “Bahama Flora," published
first in 1920 and reprinted without changes in 1962. A
recent paper by the three of us (Gillis, Howard, and
Proctor, 1973) has reported additions to the flora of the
Bahamas based on personal collections, new collections
available to us, and reports from the literature.

The present paper offers updated annotations on the
correct scientific names to be applied to the species listed
by Britton and Millspaugh whose work was completed
under the provisions of the American Code of Botanical
Nomenclature. We encourage others to use this list and its
supporting bibliography, and to report additional changes
or additions to me. We would particularly value contribu-
tions which specialists in non-vascular plants could make
to the listings in Britton and Millspaugh for mosses and
liverworts, algae, fungi, slime molds, and lichens. Our
efforts will consider only the flowering plants and pteri-
dophytes. We therefore encourage workers in these other
fields of botany to update knowledge of their disciplines
in the Bahamas too.

The value of lists of nomenclatural corrections has
been evident to us in the work of Alain (1965) on the flora
of Puerto Rico and the Virgin Islands, Additional nomen-
clatural changes have been obtained from monographs or
the recent floras of Jamaica (Adams, 1971) and Barbados
(Gooding et al., 1965). In all of these, however, the rea-
sons for the changes of names are not always explained,
requiring the curious or careful worker to reinvestigate
each and every problem. I believe the explanations which
aecompany the changes reported here will be of value to

67

68 Rhodora [Vol. 76

other workers in subtropical New World floras and par-
ticularly in the West Indies. Reasons given for changes are
usually: use of an earlier epithet, avoidance of a tautonym,
replacement of a later homonym, correct application of a
name, etc. When name changes have been made by others
than myself, I have cited the publication upon which this
information was based. Conversely, when I have had
reason not to accept a particular treatment, then reasons
have been stated.

The listing is by no means complete. Certain groups
within the Boraginaceae, Rubiaceae, Loranthaceae, and
Asclepiadaceae need intensive study and many more obser-
vations in the field, especially in such genera as Helio-
tropium, Borreria, Dendropemon, Phthirusa, and Cynan-
chum. A new look must be given to the genus Agave for
the West Indies, inasmuch as the most recent treatment
is now 60 years old (Trelease, 1913). Other groups in-
cluding the ferns are being studied by others at the present
time and we await publication of the conclusions of these
investigators.

The Turks and Caicos Islands are politically a separate
Crown Colony from the newly independent Commonwealth
of the Bahama Islands. Britton and Millspaugh included
all within the Bahama Flora and such is the geographical
sense employed here in that it includes the Turks and
Caicos Islands as well.

For ease of reference this paper will follow the order of
species presented in Britton and Millspaugh’s Bahama
Flora, which will be Monocotyledons first, followed by the
Dicotyledons, and then the Gymnosperms. Again for
brevity, Britton and Millspaugh’s flora is referred to by
the expression B&M in the text of this paper. The figure
in the left margin refers to the page in B&M on which the
taxon in question is discussed. The name following the
arrow (—) is the name considered to be correct. These
names are not necessarily nomenclatural or taxonomic
equivalents, hence the reason for not employing an equals

1974] Bahama Flora — Gillis 69

sign (—). In some instances, B&M misidentified the plant
or used a binomial incorrectly. Where monographs or
significant studies support the use of names as they were
employed in B&M, such references are cited under the
generic name, e.g., Guaiacum (Porter, 1972).

In a few instances where the name changes have involved
closely allied genera, or where the existing keys in B&M
are wholly inadequate, particularly for use in the field, I
have provided new keys. They are designed to employ
characters of the floral, fruiting, and vegetative portions
of the plant so that they can be of optimum use in working
with specimens, whether living or in the herbarium, and
whether reproductive or vegetative.

ACKNOWLEDGEMENTS

I should like to acknowledge the help of my associates Dr.
Howard and Mr. Proctor in drawing up this list, of Dr.
Carroll E. Wood, Jr. and Mr. George N. Avery, who read
the manuscript and made valuable suggestions, of my col-
league Dr. Kenneth R. Robertson who gave advice, and of
the many workers who have studied genera that have
representatives in the Bahamas. Special appreciation is
due to the librarians at the Arnold Arboretum and Gray
Herbarium of Harvard University and for the use of the
bibliographic files of the Generic Flora of the Southeastern
United States project, made available by Dr. Wood. I should
also like to acknowledge with thanks the continuing co-
operation of the personnel of the Ministry of Agriculture
and Fisheries, Commonwealth of the Bahama Islands.

This study was made while the author was a Research
Fellow at the Arnold Arboretum and I express my grati-
tude for this opportunity. A generous gift from an anony-
mous donor interested in the work on the Bahama flora.
made this study possible.

70

PAGE
IN
B&M

4

Rhodora [Vol. 76

TYPHACEAE
Typha angustifolia > Typha domingensis (Pers.)
Kunth, These are two different valid species accord-
ing to Hotchkiss and Dozier (1949) with only Typha
domingensis occurring in the Bahamas.

POTAMOGETONACEAE (ZANNICHELLIACEAE)

Potamogeton heterophyllus > Potamogeton gramineus
L. var. gramineus. On the basis of both herbarium
specimens and abundant fresh material from a pond
on San Salvador Island (Gillis 8864), I have con-
sidered all the Potamogeton material on the Islands
to be the same (see Ogden, 1948).

The question of good characters, reliable under all
conditions, for separating Ruppia maritima from
R. cirrhosa, is still open. Various treatments have
been consulted (Fernald and Wiegand, 1914; Hag-
strom, 1916; McCann, 1945; Setchell, 1946; Reese,
1962a and b; Gamerro, 1968; and Schwanitz, 1967)
for specimens collected in the Bahamas. For the
moment, it appears that both species seem to be
present. Therefore, R. cirrhosa is an addition to the
flora, rather than merely a different name for the
species in B&M. Collectors are urged to look for R.
cirrhosa inasmuch as flowering and fruiting material
are as yet unknown in the Bahamas. An adaptation
of Gamerro's key follows:

Peduncle of inflorescence more than 10 cm. (8-60
em.), coiled until anthesis; anther sacs reniform,
1.7 mm. long; fruits rounded at apex; leaves 1 mm.
broad; leaf epidermal cells 16-194; 2n = 40 ....
TEN Ruppia cirrhosa (Petagna) Grande.
Peduncle of inflorescence less than 5 cm., not coiled
in spiral until anthesis; anther sacs elliptic, 0.6-0.7
mm. long; fruits irregularly pointed; leaves 0.5

1974]

6

13

14

Bahama Flora — Gillis 71

mm. broad; epidermal cells of leaves 12.5-16.0,5
Oe ee et Mert a tad Ruppia maritima L.

ZANNICHELLIACEAE (CYMODOCEACEAE)

Cymodocea manatorum —> Syringodium filiforme Kütz.
This change of name follows Dandy and Tandy (1939)
and den Hartog (1970).
Halodule wrightii Aschers. is indeed in the Bahamas,
but so is H. beaudettei (den Hartog) den Hartog
(1964). These two species can be separated as fol-
lows:

Leaf tips bicuspidate; leaf blades 1/3-4/5 mm.
wide; leaves with linear teeth, concave on the inner

side ... Halodule wrightii Aschers.
Leaf tips tricuspidate; leaf blades 2/3-114 mm.
wide; leaves with linear lateral teeth ... Halodule

beaudettei (den Hartog) den Hartog.

HYDROCHARITACEAE (ELODEACEAE)
Halophila — consult Hartog (1959).

ALISMATACEAE

Echinodorus cordifolius — Echinodorus berteroi
(Spreng.) Fass. (See Fassett, 1955).

GRAMINEAE

There is some question among agrostologists whether
Schizachyrium should be segregated from Andropogon
as it was in B&M. Chase (1951) joins the two genera.
For the moment, however, I shall follow treatments
by Gould in maintaining them as separate genera
(see Gould, 1967, 1968, and 1969).

Nazia aliena > Tragus berteronianus Schult. Tragus
is conserved over Nazia. The species epithet changed
from aliena to berteronianus because the type of
Lappago alienus Spreng., the basionym, is referred
to Pseudechinolaena. polystachya (H.B.K.) Stapf, and
not to this Bahamian plant (Chase, 1951).

72

15

16

16

16

18

18

18

19

20

Rhodora [Vol. 76

Syntherisma filiformis > Digitaria panicea (Sw.)
Urb. The species on which the epithet filiformis is
based is a different species from the one of concern
here. The earliest name for this species is Milium
panicea Sw. which was transferred to Digitaria by
Urban. The combination was made twice in the
same year (1920), by Urban in February and by
Fernald in June, The Urban combination obviously
has priority. Digitaria is an earlier generic name
than Syntherisma.

Syntherisma digitata — Digitaria horizontalis Willd.
Milium digitatum Sw. (1788) is the earliest basionym,
but is is preoccupied in Digitaria by D. digitata Buse
(1854). The next earliest name available is D. kori-
zontalis Willd. (1809).

Syntherisma sanguinalis > Digitaria sanguinalis (L.)
Scop.

Valota insularis > Trichachne insularis (L.) Nees.
Valota is inadequately published (Chase, 1951).
Paspalum poiretii R. et S. ^ Paspalum sagetii Chase.
Paspalum caespitosum — Paspalum poiretii R. et S.
The work of Chase (1929) sorts out the names of
these Paspalum species by typifying them. It is thus
apparent that, although the name P. poiretii is still
used in the Bahama flora, it must be applied to a
different species from the one to which it was as-
signed by B&M.

Paspalum portoricense ^ Paspalum molle Poir. in
Lam. The change is required because of an earlier
epithet (Chase, 1951).

Paspalum simpsonii > Paspalum blodgettii Chapm.
Paspalum blodgetti is an earlier name for P. simp-
sonii Nash, and not a synonym of P. caespitosum
Chase, 1951).

Paspalum glabrum — Paspalum laxum Lam. The
change is required because of an earlier epithet
(Chase, 1951).

Paspalum distichum L. is attributed by B&M to the

1974]

22

22

22

24

24

27

29

Bahama Flora — Gillis T8

Pugil. Pl. Jam., one of the dissertations of Linnaeus
(1759). It actually appeared earlier in the same
year in the Systema Naturae, ed 10, and should be
so attributed.

Panicum barbinode > Panicum purpurascens Raddi.
The change is required because of an earlier epithet
(Hitchcock, 1936).

Panicum distantiflorum — Setaria distantiflorum (A.
Rich. in Sagra) Pilger.

Panicum chapmani — Setaria chapmanii (Vasey)
Pilger. Rominger (1962) has shown that these two
species properly belong in Setaria.

Panicum dichotomiflorum — Panicum _ bartowense
Scribn. et Merr. These names were treated as synony-
mous by B&M. Hitchcock (1935) and Chase (1951)
felt that they are separate species and that the Baha-
mian plant is P. bartowense, Fernald (1934) treated
the taxon as P. dichotomiflorum var. bartowense
Schribn. et Merr.) Fern.

Panicum coerulescens > Panicum coerulescens Hack.
ex Hitehe. This change is simply a corrected spelling.
Oplismenus hirtellus (L.) R. & S. 1817 — Oplismenus
hirtellus (L.) Beauv., Ess. Agrost. 54, 168. 1812. An
author change is needed for this earlier publication
of the binomial.

Chaetochloa geniculata — Setaria geniculata (Lam.)
Beauv.

Chaetochloa setosa — Setaria setosa (Sw.) Beauv.
Chaetochloa macrosperma — Setaria macrosperma
(Scribn. et Merr.) K. Schum. Setaria is the older
and preferred name over Chaetochloa. Setaria Beauv.
is conserved over Setaria Michx.

Cenchropsis mysuroides — Cenchrus mysuroides
H.B.K. Cenchropsis is not now considered distinct
from Cenchrus (Chase, 1936 and Delisle, 1963).
Cenchrus carolinianus ^ Cenchrus incertus M. A.
Curtis. The use of Cenchrus carolinianus in B&M
was a misapplication of the name. That name is

74

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30
31

31

32
38

33

Rhodora [Vol. 76

now considered a synonym of C. longispinus, but is
correctly used for a plant not found in the Bahamas.
The plant in question is C. incertus which is wide-
spread in the Bahamas (DeLisle, 1963).

Cenchrus microcephalus > Cenchrus incertus M. A.
Curtis. DeLisle (ibid.) felt that there was so little
difference between the *endemic" race of Cenchrus
with small fruits and the more widespread species
that he treated them as conspecific. The small fruited
form should be considered an insular form.
Cenchrus viridis —^ Cenchrus brownii R. et S. This
name change is required because of an earlier epithet
(DeLisle, ibid.).

Stenotaphrum — Consult Sauer, 1972.

Aristida gyrans > Aristida vilfifolia Henr. After
publication of B&M, Aristida vilfifolia was segregated
as a distinct species from A. gyrans which was the
name in B&M (Henrard, 1926-33; 1929-33). It is
the segregate which indeed is the one in the Bahama
flora.

Aristida scabra > Aristida ternipes Cav. This change
is required because of an earlier epithet.

Sporobolus indicus — Sporobolus jacquemontii Kunth.
Sporobolus berteroanus — Sporobolus indicus (L.) R.
Br. In the Sporobolus indicus complex, there are two
Caribbean species which are often confused: S. jac-
quemontii, which has an obtuse upper glume and is
a fully tropical species, and S. indicus, which has an
acute upper glume and is a more subtropical species.
According to the description in B&M, these taxa have
been confused (Clayton, 1964 and personal communi-
cation).

Sporobolus atrovirens — Sporobolus bahamensis Hack.
Sporobolus atrovirens is a different plant from the one
in the Bahamas; it is a Mexican plant, is a perennial,
and has smaller spikelets and glumes than S. baha-
mensis. The latter is a Bahamian endemic and an
annual.

1974]

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36

39

39

41

Bahama Flora — Gillis 75

Sporobolus argutus > Sporobolus pyramidatus (Lam.)
Hitehe. This name change is required because of an
earlier basionym. The citation for Domingan drop-
seed-grass should be Sporobolus domingensis (Trin.)
Kunth, Rév. Gram. 1, Suppl.; xvii. 1830, a citation
three years earlier than that cited in B&M.
Capriola dactylon > Cynodon dactylon (L.) Pers.
Cynodon is conserved over Capriola,

Chloris polydactyla > Chloris dandyana Adams.
Adams (1971) published a new name for this plant
without giving the background reasons for the illegiti-
macy of the original name. The background is as
follows: There is a Jamaican plant with the legiti-
mate name Andropogon barbatus L. Some time after
describing this Jamaican grass, Linnaeus applied the
same binomial to an East Indian grass. The epithet
barbatus (referring to the East Indian species) was
transferred to Chloris by Swartz, an act which pre-
vents the transfer of the Jamaican species epithet to
Chloris. In the meantime, Linnaeus proposed a super-
fluous name for the first Andropogon barbatus, i.e.,
A. polydactylos. Swartz’s transfer of this super-
fluous name to Chloris is illegitimate; hence Adams’s
publication of a new name for the species in the West
Indies. (I should like to acknowledge personal com-
munication from C. D. Adams and J. E. Dandy for
the historical background on this matter.)
Phragmites phragmites ^5 Phragmites australis (Cav.)
Trin. ex Steud. Clayton (1968) pointed out that the
correct name for the pan-tropical reed is the com-
bination used above.

Diplachne fascicularis > Leptochloa fascicularis (Lam.)
A. Gray. Current views of agrostologists suggest that
Diplachne should be contained within Leptochloa.
Eragrostis cilaris (L.) Link 1827 Eragrostis cili-
aris (L.) R.Br. in Tuckey, 1818. Chase (1951) re-
corded the earlier publication of this combination.

76

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44

45

46

AT

49

Rhodora [Vol. 76 -

Uniola virgata > Leptochloópsis virgata — (Poir.)
Yates. Yates (1966) has subdivided Uniola into
Uniola, Chasmanthium, and Leptochlodpsis on the
basis of a number of anatomical, cytological, morpho-
logical, and embryological characters. Because this
species of spike-grass is considered close to Lepto-
chloa, Yates erected Leptochlodpsis to contain it and
another species from Ecuador.

CYPERACEAE

Cyperus paniculatus > Cyperus polydactylos Rottb.
These names originate from the same date and are
considered to be synonymous. Corcoran (1941) chose
C. polydactylos to be the inclusive name. The Bahama
populations might be designated as C. polydactylos
var. texensis (Torr.) Fern. (McLaughlin, 1944).

Cyperus cuspidatus ^ Cyperus aristatus Rottb. True
C. cuspidatus does not occur in the Bahamas, but in
Cuba. (See Horvat, 1941; McLaughlin, 1944).
Cyperus pseudovegetus > Cyperus distinctus Steud.
Cyperus pseudovegetus was a name misapplied to the
plant of the Bahamas (O’Neill, 1939).

Cyperus brunneus — Cyperus planifolius L. C. Rich.
This name change is required by an earlier epithet
(McGivney, 1938; McLaughlin, 1944).

Cyperus ferax — Cyperus odoratus L. var. odoratus.
O'Neill (1940) demonstrated that these two names
apply to the same species, and that C. odoratus is the
earlier name. He further thought that the typical
variety was the one found in the Bahamas.

According to the treatment by McLaughlin (1944),
the populations of Cyperus filiformis Sw. in the
Bahamas are of the typical variety (var. filiformis).
Stenophyllus wilsonii > Bulbostylis floccosa (Griseb.)
Clark in Urb. Stenophyllus is merged with Bulbo-
stylis in the treatment by Kral (1971). Bulbostylis
floccosa is the earliest name. It is not endemic to the

1974]

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51

52

53

54

54

55

55

56

Bahama Flora — Gillis T4

Bahamas as stated in B&M, but also is found in Cuba
and Hispaniola.

Fimbristylis diphylla — Fimbristylis dichotoma (L.)
Vahl. This change is required because of an earlier
basionym.

According to B&M, the record of Fimbristylis hirta
in the Bahamas is very tenuous. Neither Kral nor
I have seen any material from the Bahamas. If in-
deed it does occur there, its more appropriate name
should be F. squarrosa Vahl (see Kral, 1971).
Abildgaardia monostachya > Abildgaardia ovata
(Burm. f.) Kral. This change is required because of
an earlier epithet (Kral, ibid.).

There are cyperologists who wish to merge Dichro-
mena with Rhynchospora. On the basis of the striking
white bracts and insect pollination characteristic of
Dichromena, I am, for the moment at least, retaining
it as distinct.

Rhynchospora cyperoides > R. cyperoides (Sw.)
Mart. var. cyperoides.

Rhynchospora tracyi —^ Rhynchospora cyperoides var.
triceps (Vahl) Bock. This treatment follows that of
Kükenthal (1949).

Rhynchospora perplexa > Rhynchospora microcarpa
Baldw. ex Gray. Gale (1944)maintained that true
R. perplexa was not in the Bahamas and that all the
material which Britton and Millspaugh called by this
name was misdetermined.

Rhynchospora bahamensis Britt. ~ Rhynchospora lin-
deniana var. bahamensis (Britt.) Gale. Gale's treat-
ment (ibid.) of the West Indian Rhynchospora species
reduced Britton’s species to a variety of the more
widespread R. lindeniana.

Rhynchospora marisculus > Rhynchospora odorata
C. Wright ex Griseb. Gale (ibid.), in examining the
same material which Britton and Millspaugh did,
determined that the use of R. marisculus in B&M was
a misapplication of the name.

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56

60

61

67
67

Rhodora [Vol. 76

Mariscus jamaicensis > Cladium jamaicense Crantz.
Most treatments now segregate Cladium from
Mariscus.

Scleria. Consult Jackson, 1949.

PALMAE (ARECACEAE)
Thrinax parviflora > Thrinax floridana Sarg. Thri-
nax parviflora is an endemic species of Jamaica.
Furthurmore, the treatment of Thrinax by Read in
Adams (1971) will be superseded now that additional
type material has been examined. The correct name
for the small thatch of South Florida and the northern
West Indies with leaf segments broadest at the point
of fusion, with scattered, fimbriate, centrally trans-
lucent seales, and glabrous axes should be T. floridana
(Read, personal communication).
Thrinax parviflora > Thrinax floridana Sarg. Thri-
(Jacq.) L. H. Bailey. True C. argentea may exist in
the southern Bahamas, but this has yet to be demon-
strated. It is now treated as being indigenous to
Hispaniola. (See Moore, 1963).
Paurotis wrightii > Acoelorrhaphe wrightii (Griseb.
ex H. Wendl.) H. Wendl. ex Becc. The preferred
name for the Paurotis palm, Spanish-top, or Ever-
glades palm is now Acoelorrhaphe (Moore, 1963).
Pseudophoenix vinifera > Pseudophoenix sargentii
subsp. saonae (Cook) Read. True P. vinifera is in-
digenous to Hispaniola and is not known from the
Bahamas. (Read, 1968).

COMMELINACEAE
Commelina longicaulis ^ Commelina diffusa Burm. f.
Commelina elegans —^ Commelina virginica L. Name
changes are made here in accord with the treatment
by Brashier (1969). The following key may be more
easily used than that in B&M:

Plant annual, trailing; spathes not united at the
base, margins ciliate; internodes glabrous; sta-

1974]

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69

71

78
78

79

83

83

84

84

84

Bahama Flora — Gillis 79

minodes Qoi m Commelina diffusa Burm. f.
Plant perennial, tufted and erect; spathes united at
the base, margins membranous; internodes puberu-
lent; staminodes 3 ...... Commelina virginica L.

Rhoeo discolor ^ Rhoeo spathacea (Sw.) Stearn. This
name change follows Stearn (1957) who discovered
that the earlier epithet of Swartz applies to this
plant.

AGAVACEAE (LILIACEAE)

Cordyline guineensis 5 Sansevieria hyacinthoides
(L.) Druce. This change follows Wijnands (1973).

SMILACACEAE
Smilax. Consult Coker, 1944.

AMARYLLIDACEAE

Atamosco rosea — Zephranthes rosea Lindl.
Atamosco cardinalis > Zephyranthes cardinalis C.
Wright. Zephyranthes is conserved over Atamosco.
Hymenocallis caymanensis ^ Hymenocallis latifolia
(Mill.) Roemer. This change follows the treatment
by Sealy (1954).

ORCHIDACEAE

Carteria corallicola — Basiphyllaea corallicola (Small)
Ames. Carteria is a later homonym for a genus of
algae. Basiphyllaea was erected as a genus to correct
this situation.

Vanilla eggersii — Vanilla dilloniana Correll.
Vanilla articulata — Vanilla barbellata Reichb. f. This
treatment of Vanilla in the Bahamas follows Cor-
rell (1950).

Pelexia adnata — Spiranthes adnata (Sw.) Benth.
Although Pelexia is a conserved name, it is also a
generic synonym of Spiranthes (Garay and Sweet,
1972).

Pelexia setacea > Eltroplectris calcarata (Sw.) Garay

80

85

85

86

87

87

89

89

90

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Rhodora [Vol. 76

et Sweet. Luer (1972) changed Pelexia setacea to
Centrogenium setaceum but Garay and Sweet (1972)
maintained that this combination is based on an ille-
gitimate name and therefore should be Eltroplectris
calcarata (Sw.) Garay et Sweet.

Ibidium tortile > Spiranthes tortilis (Sw.) L. C.
Rich. Jbidiwm is considered to be a generic synonym
of Spiranthes. (Garay & Sweet, 1972).

Ibidium lucayanum — Spiranthes polyantha Reichb. f.
These names are considered synonymous by Luer
(1972).

Stenorrhynchus lanceolatus — Spiranthes lanceolata
(Aubl) Léon. Stenorrhynchus is now treated as a
generic synonym of Spiranthes. (See Dunsterville
and Garay, 1965).

Ponthieva brittonae > Ponthieva racemosa var. brit-
tonae (Ames) Luer. Two varieties of Ponthieva
racemosa occur in the Bahamas. One is the typical
variety (var. racemosa) as treated by Luer (1972)
and reported by him as new to the flora. Var. brit-
tonae is the reduction made by Luer of the name in
B&M.

Physurus querceticola —^ Erythodes querceticola
(Lindl.) Ames. Physurus is considered to be a gene-
ric synonym of Erythodes (Garay and Sweet, 1972).
Polystachya minuta — Polystachya flavescens (Lindl.)
J. J. Smith. Although the epithet minuta (from
Epidendrum minutum Aubl.) is the oldest name
available, it is preoccupied in Polystachya, and hence,
P. flavescens must be used.

Spathiger | rigidus > Epidendrum rigidum Jacq.
Spathiger is considered a generic synonym of Epi-
dendrum.

Auliza nocturna > Epidendrum nocturnum Jacq.
Auliza is considered a generic synonymn of Epiden-
drum (Dunsterville and Garay, 1965; Luer, 1972).

There is much discussion among orchid taxonomists
whether Encyclia should be segregated from Epiden-

1974]

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93

93

95

95

103

104

Bahama Flora — Gillis 81

drum or not. For the moment, I accept the treat-
ment of Dressler (1961; 1966) and Beckner (1970)
in maintaining Encyclia at the generic level as in
B&M.

Encyclia rufa > Encyclia bahamensis (Griseb.) Britt.
et Millsp. Specimens labeled E. rufa from the Ba-
hamas are now all best considered E. bahamensis.
Encyclia rufa is, as B&M suspected, still not known
from the archipelago.

Encyclia diurna — Encyclia hodgeana (A. D. Hawkes)
Beckner. This name change is required because of
more complete understanding of the taxa involved.
The species previously called E. diurna or E. gracilis
is, in fact, a hybrid between E. hodgeana and E.
tampensis. (See Beckner, 1970).

Epicladium boothianum — Encyclia boothiana var.
erythronioides (Small) Luer. Epicladiwm is a generic
synonym of Encyclia.

Anacheilium cochleatum — Encyclia cochleata var.
triandra (Ames) Dressler. Anacheiliwm is a generic
synonym of Encyclia.

Laeliopsis domingensis ^ Broughtonia lindenii (Lindl.)
Dressler. This change is made in accord with the
treatment by Dressler (1961).

Limodorum simpsoni — Calopogon tuberosus (L.)
Britten, Stearns, and Poggenberg. This treatment
follows Luer, 1972.

PICRODENDRACEAE
Picrodendron macrocarpum — Picrodendron baccatum
(L.) Krug et Urb. in Engl. I presently have Picro-
dendron under study. It appears that this genus
should be placed within the Euphorbiaceae. There
is likely only one species, which would be the typical
one, P. baccatum.

ULMACEAE
Trema lamarckiana — Trema lamarckianum (R. et S.)
Blume. The gender of Trema should be neuter.

82

Rhodora [Vol. 76

MORACEAE

105 Ficus jacquinifolia > Ficus perforata L.

Fieus brevifolia — Ficus citrifolia Mill. Recent treat-
ments of Ficus (DeWolf, 1969 and Condit, 1969) have
equated F. perforata with F. jacquinifolia A. Rich.
in Sagra. Moreover, they have also equated F. brevi-
folia and F. citrifolia. B&M’s F. brevifolia Nutt.
(lectotype at BM) must become F. citrifolia Mill., the
earlier name (lectotype at BM, chosen by Dandy). I
have further studied F. perforata which DeWolf and
Condit have given as the correct name for F. jac-
quinifolia (which itself should be written jacquinii-
folia). The Linnaean name must be typified by an
illustration in Plumier associated with the disserta-
tion on the plants of Surinam (1775). Although there
is a specimen of F. perforata at LINN (Savage Cata-
log No. 1240: 9), the only reference in the Disserta-
tion (and the later Amoenitates) in this context is
that to Plumier. The specimen at LINN appears to
be of the tree of Central America and northern South
America, F. pertusa L. f. I should like to acknowledge
the consultation and discussion with Dr. Gordon De-
Wolf which proved very valuable in application of
names in Ficus. A key to the Bahama material fol-
lows:

1. Syconia sessile .......... Ficus aurea Nutt.
1. Sycania stalked
2. Syconia 7-10 mm. in diameter; petioles less
than 1 cm.; blades obovate; lateral veins

flush with lower surface; leaf base cuneate
Lee eee eee eee ee eee Ficus perforata L.

2. Syconia 8-18 mm. in diameter; petioles
longer than 1 cm.; blades ovate; lateral veins

+ prominent, somewhat elevated on lower
surface; leaf blade base rounded .........
Lee eee ee eee eee eee Ficus citrifolia Mill.

1974]

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113

114

114
115

115

117
TIT
DT

121
121
121

Bahama Flora — Gillis 83

LORANTHACEAE

Generic study of Dendropemon, Phthirusa, and Phora-
dendron is needed before good species concepts and
correct nomenclature emerge.

ARISTOLOCHIACEAE
Aristolochia passifloraefolia — Aristolochia passiflori-
folia A. Rich. in Sagra. With the exception of the
orthograhic change (above) made in accord with
rules adopted at the Seattle Congress, the names for
Aristolochia species in the Bahamas remain as they
were in B&M (Pfeifer, 1966; 1970).

POLYGONACEAE
Persicaria hydropiperoides > Polygonum hydropipe-
roides Michx.
Persicaria punctata — Polygonum punctatum Ell.
Persicaria portoricensis Polygonum densiflorum
Meissn. Persicaria is treated by Graham and Wood
(1955) as a section of Polygonum.
Fagopyrum fagopyrum —> Fagopyrum esculentum
Moench. This change is necessary in order to avoid a
tautonym.
Coccolobis laurifolia — Coccoloba diversifolia Jacq.
Coccolobis bahamensis — Coccoloba tenuifolia L.
Coccolobis diversifolia —^ Coccoloba swartzii Meisn.
Coccoloba is conserved over Coccolobis. All Coccoloba
species therefore should change spelling to conform.
The 3 species mentioned above are changed further
in accord with Howard (1957), From Howard's
treatment also comes the change of citation for
Coccoloba uvifera (L.) L., Syst. Nat., ed. 10, p. 1007.
1759.

CHENOPODIACEAE
Dondia linearis > Suaeda linearis (Ell.) Moq.
Dondia fruticosa —> Suaeda fruticosa (L.) Forsk.
Dondia insularis ^ Suaeda insularis (Britt.) Urb. et
Ekm. Dondia is considered to be a taxonomic synonym

84

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126

126

126

127

128

128

130

131
131
132

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Rhodora [Vol. 76

of Suaeda, Suaeda Forsk. ex Scopoli remains in the
list of conserved names even though the reason for
its inclusion is no longer valid.

AMARANTHACEAE

Amaranthus gracilis > Amaranthus viridis L. Reed
(1968) and Merrill (1936) considered these species
to be taxonomic synonyms. Amaranthus viridis L.
is the earlier name.

Centrostachys indica — Achyranthes aspera var. in-
dica L. Centrostachys seems better treated as a
generic synonym Achyranthes.

Achyranthes maritima — Alternanthera maritima St.
Hil.

Achyranthes polygonoides — Alternanthera polygo-
noides (L.) R. Br.

Achyranthes repens —> Alternanthera pungens H.B.K.
This treatment follows Reed, 1968. The epithet
pungens is used for the last species rather than the
earlier epithet repens because Achyranthes repens is
a nomen confusum according to Reed.

Philoxerus remains the same except for citing an
earlier date of publication: Philoxerus vermicularis
(L.) R. Br. Prodr. Fl. Nov. Holl. 416. 1810.

Iresine celosia — Iresine diffusa Humb. et Bonpl. ex
Willd. (Sp. Pl., ed. 4, 4: 765. 1806). Iresine celosia
is an illegitimate name.

NYCTAGINACEAE
Boerhaavea coccinea Mill. > Boerhavia coccinea Mill.
Boerhaavea erecta L.— Boerhavia erecta L. Boer-
havia is the original spelling.

Torrubia obtusata > Guapira obtusata (Jacq.) Little.
Torrubia cokeri > Guapira obtusata (Jacq.) Little.
Torrubia longifolia > Guapira longifolia (Heimerl)
Little.

Torrubia bracei > Guapira bracei (Britt.) Little. If
one considers that blollies are similar to Pisonia

1974]

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139

143

144

144

146

147

Bahama Flora — Gillis 85

species, then Torrubia merges with Pisonia. If one
wishes to recognize both as distinct genera, then one
must take up Guapira, an earlier name than Torrubia.
The Committee for Spermatophytes rejected a pro-
posal to conserve Torrubia, hence a paper by Little
(1968) which made the appropriate transfers to
Guapira, It is currently felt that Torrubia cokeri is
synonymous with Guapira obtusata; it has been so
treated here. This is an exceedingly variable species
with different forms having been named in the past.

PORTULACACEAE
Portulaca phaeosperma —> Portulaca rubricaulis H.B.K.
Legrand (1952) chose the earlier name for this
species.

NYMPHAEACEAE
Castalia pulchella > Nymphaea pulchella DC. Wood
(1959) has treated Castalia as a subgenus of Nym-
phaea.

LAURACEAE
Ocotea coriacea > Nectandra coriacea (Sw.) Griseb.
Recent treatments have merged Ocotea and Nectandra
(Wood, 1958; Allen, 1966).
Persea pubescens Persea palustris (Raf.) Sarg.
Kopp (1966) pointed out that palustris is an earlier
epithet than pubescens for this species.

CASSYTHACEAE
Cassytha americana —> Cassytha filiformis L. This
change is necessitated because filiformis is the earlier
epithet.

CRUCIFERAE (BRASSICACEAE)
Hitchcock (1945) recognized several varieties of
Lepidium virginicum. If this treatment is followed,
the Bahama one is var. virginicum.
Sinapis arvensis Brassica kaber (DC.) L. C.
Wheeler. Wheeler (1938) thought that Sinapis should

86

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148

151

152

153

154

Rhodora [Vol. 76

properly be considered a part of Brassica. Sinapis
arvensis of B&M cannot be transferred to Brassica
because there is already a B. arvensis. Brassica kaber
appears to be an older combination than B. will-
denovii Boiss., as attributed to the Bahamas in
Adams, 1972.

Radicula brevipes —> Rorippa portoricensis var. pu-
mila (O. E. Schulz) Stuckey. The name used in B&M
and its synonyms are misapplied to the Bahamian
taxon, according to Stuckey (1972). Stuckey inad-
vertently maintained a neuter ending for the varietal
epithet in his treatment; this is corrected above.
Canara didyma — Coronopus didymus J. E. Smith.
Coronopus (1757) is an older name than Canara
(1792). Coronopus Zinn. (1757) is conserved over
Coronopus Miller (1754), a genus of Plantaginaceae.

MORINGACEAE
Moringa moringa — Moringa oleifera Lam. In avoid-
ing the tautonym, Moringa moringa, Ernst (1963)
has picked up Moringa oleifera for the horseradish-
tree.

CRASSULACEAE
Bryophyllum pinnatum — Kalanchoë pinnata (Lam.)
Pers. Treatments of Kalanchoë and Bryophyllum
(Baldwin, 1938; Hamet, 1907) have recognized
Kalanchoë as the inclusive genus.

ROSACEAE (AMYGDALACEAE)
Laurocerasus myrtifolia > Prunus myrtifolia (L.)
Urb. The cherry-laurel is best considered in the larger
genus Prunus especially in view of other recent floras
which have so placed it.

CHRYSOBALANACEAE (AMYGDALACEAE)
Chrysobalanus pellocarpus ^ Chrysobalanus icaco L.
Prance (1972) thought that there were insufficient
means to separate C. pellocarpus from C. icaco and
has treated them as taxonomic synonyms.

1974]

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158

159

159

Bahama Flora — Gillis 87

LEGUMINOSAE (MIMOSACEAE)
Pithecellobium is conserved over Pithecolobium, and
hence all species change to this orthography. The
ram’s horn has been a source of confusion for a
number of years. Pithecellobium keyense Britt. ex
Coker in Shattuck was proposed to replace P. guada-
lupense because the latter name was not applicable.
Britton did not state why he considered the name
inapplicable. In creating the new name, Britton
(actually Coker) had no description and hence,
created a nomen nudum. This situation was rectified
in 1928 by Britton and Rose. Although the type of
Mimosa guadalupensis Pers., the basionym, cannot
be found at present in the Jussieu Herbarium at P,
there appears to be no valid reason why this name
cannot be taken up, pending proof that this plant
does not represent the concept of the species known
as “ram’s horn" in the West Indies and South Florida.
Isely (1972) referred to specimens at the Delessert
Herbarium and at Leiden which should be “critical,”
but inasmuch as the basionym is linked by Persoon
with “Hab ad Guadalupam (Herb. Juss.)," the lecto-
type material should be sought in the Jussieu Her-
barium. Why Britton and Rose indicated that this
species was “not Inga guadalupensis Desv." (which
is based on the same basionym) is unknown. The
only Pithecellobium which occurs on Guadaloupe is
this one. I am therefore retaining the name in B&M,
but adding the parenthetical reference to the author
of the basionym which was omitted in B&M: Pithe-
cellobium guadalupense (Pers.) Chapm.

The Lysiloma species remain as in B&M (Gillis and
Stearn, in press), except for correcting gender to
neuter.

Anneslia haematostoma — Calliandra haematomma
(Bert.) Benth.

Anneslia formosa > Calliandra formosa (Kunth)
Benth. Calliandra is conserved over Anneslia. The

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Rhodora [Vol. 76

specific epithet haematostoma in B&M appears to be
a misprint, judging from the basionym they cite.
Despite efforts of Britton and Rose (1928) to frac-
tionate Acacia into Bahamia and Lucaya, it seems
best to maintain the three species of Acacia in B&M
under the same names as in the flora: Acacia acuifera,
A. choriophylla, and A. macracantha.

Vachellia farnesiana — Acacia farnesiana (L.) Willd.
Vachellia is best considered a generic synonym of
Acacia.

Leucaena glauca — Leucaena latisiliqua (L.) Gillis et
Stearn. Based on typification of an earlier name than
either the epithet glauca or leucocephala (deWit,
1961). Gillis and Stearn (in press) have shown that
latisiliqua should be taken up.

Acuan virgatum — Desmanthus virgatus (L.) Willd.
var. virgatus.

Acuan depressum —> Desmanthus virgatus var. de-
pressus (Willd.) Turner. Desmanthus is conserved
over Acuan. The two Bahama plants probably are
best considered distinct only at the infraspecific
level; therefore the treatment of Isely (1970) is fol-
lowed,

Neptunia — consult Windler (1966).

Prosopis — consult Johnston (1962).

LEGUMINOSAE (CAESALPINIACEAE)

Cassia. tora, —^ Cassia obtusifolia L. B&M suggest that
Cassia tora and C. obtusifolia are synonyms and chose
the former as the name to be used in the Bahama
flora. Typification shows that such is not the case.
DeWit (1955) selected a specimen in the Linnaean
Herbarium as the type of C. tora L., but Brenan
(1958) objected to this choice and selected a speci-
men in the Hermann Herbarium (at BM) as the
type. Based on the position of the petiolar gland,
length of flowering and fruiting pedicels, anther
shape, and presence of a 1.2-2 mm. areole on both

1974]

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Bahama Flora — Gillis 89

sides of the seed, Brenan distinguished these two as
distinct species. There is probably no Cassia tora
in the New World according to Brenan.
Chamaecrista chamaecrista — Cassia nictitans L. De-
spite the efforts of Britton and Rose to fractionate
Cassia into segregate genera Chamaecrista, Ditre-
mexa, Peiranisia, ete., I maintain Cassia as an in-
clusive genus. This necessitates different names for
the six species placed in Chamaecrista in B&M. Cassia
chamaecrista L. is based on a composite collection
according to Pennell (1917). Therefore the correct
name for the annual plant that is distributed through-
out Eastern United States is generally accepted as
C. fasciculata Michx. “Chamaecrista chamaecrista"
of B&M should be rather the perennial species, Cassia
nictitans L. I believe it is necessary to retypify all
the names used in this complex in order to apply
these names properly.

Chamaecrista riparia > Cassia caymanensis C. D.
Adams. Adams (1970) has shown that Cassia riparia
is an illegitimate name, and therefore this plant must
have a new epithet in Cassia.

Chamaecrista lucayana — Cassia lucayana Britt.
Chamaecrista caribaea — Cassia caribaea Northrop.
Chamaecrista inaguensis > Cassia inaguensis Britt.
Chamaecrista lineata > Cassia lineata Sw. These
species revert to their original names in Cassia.
Guilandina crista > Caesalpinia bonduc (L.) R. Br.
Gray nickerbean has been shown by Dandy and Exell
(1938) to be Caesalpinia (or Guilandina) bonduc.
The epithet crista as used by Linnaeus refers to an
Old World species.

Guilandina bonduc — Caesalpinia divergens Urb.
Guilandina ovalifolia — Caesalpinia ovalifolia Urb.
Current thought among those working with this
group of scrambling shrubs of the coastal regions of
the West Indies is to treat them as a section of
Caesalpinia, rather than a separate genus Guilandina.

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Rhodora [Vol. 76

For some long period of time, G. bonduc was used to
refer to the yellow-seeded nickerbean. As we have
seen, proper typification by Dandy and Exell has
shown that the name bonduc is misapplied when
used thus. Among the synonyms given by B&M for
this species, most apply to still other species. Caesal-
pinia divergens seems to be the earliest name available
for this species. The yellow-seeded nickerbeans in
the Bahamas (with the exception of a new species
to be described by Gillis and Proctor from Inagua)
can be separated as follows:

1. Seeds spherical, yellow; leaflets elliptic to sub-
orbicular, obtuse or emarginate .............
Lee ee eee Caesalpinia ovalifolia (Urb.) Britt.

1. Seeds oblong, bronze-brown; leaflets ovate,
acute or acuminate ............. nee

I accept B&M’s treatment of 1920 instead of the later
treatment of Britton and Rose (1928) which recog-
nized the segregate genus Vicarago. I see no reason
to separate this taxon from Caesalpinia.

LEGUMINOSAE (FABACEAE)

Ateleia cubensis > Ateleia gummifera (Bert. ex DC.)
Dietr. The epithet gummifera is an earlier epithet
for this species (Rudd, 1968).

Yakovlev (1967 a and b) has recognized a number of
subspecific units in Sophora tomentosa L. For the
moment, these seem difficult to accept as other than
insular forms, and therefore I consider the taxon to
be the single, variable S. tomentosa throughout the
archipelago as in B&M.

Cracca cinerea > Tephrosia cinerea (L.) Pers.
Cracca cathartica — Tephrosia senna H.B.K. Although
Cracca Benth. in Benth. et Oersted is conserved over
Cracca L., Tephrosia Pers. is conserved over Cracca
L., Needhamia Scopoli, and Reineria Moench. The

1974]

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182
182

183
184

184

184

185

Bahama Flora — Gillis 91

correct generic name for the plants in the Bahamas
is Tephrosia. Because the key to species in B&M is
somewhat inadequate, a revised one follows:

1. Leaflets 9-15; racemes opposite the leaves; pod
2.5-4 mm. broad .. Tephrosia cinerea (L.) Pers.
1. Leaflets 5-9; racemes terminal and opposite the
leaves; pod 4-5 mm. broad .................
Tephrosia senna H.B.K.

e «.4* 9» 9 6,8 he 6 6 8 à 5 € 8 9.0, 9

Sesban sericea — Sesbania sericea (Willd.) DC.
Sesban occidentalis —^ Sesbania emerus (Aubl.) Urb.
Agati grandiflora — Sesbania grandiflora (L.) Pers.
Sesbania is conserved over Sesban and Agati. The
second of the species mentioned above was known
only from fruit at the time of the writing of B&M.
From collections made recently from the same popu-
lation, I have determined it to be S. emerus.
Stylosanthes — consult Mohlenbrock (1957).
Meibomia supina —^ Desmodium canum (Gmel.) Schinz
et Thell.

Meibomia tortuosa —^ Desmodium tortuosum ( Mill.)
DC.

Meibomia mollis ^À Desmodium glabrum (Mill.) DC.
Desmodium is conserved over Meibomia. Supina
cannot be used as an epithet for the first species above
because it is based on a name which is a later
homonym (Hedysarum supinum Sw. non H. supinum
Chaix ex Villars). Incanum has often been taken
up as a specific epithet for this plant also (based on
Hedysarum incanum Sw.), but the basionym is a
later homonym for H. incanum Thunb. Desmodium
canum is based on the earliest available legitimate
epithet. Desmodium glabrum is based on an earlier
basionym than Meibomia mollis of B&M.
Alysicarpus nummularifolius > Alysicarpus vaginalis
(L.) DC. Alysicarpus vaginalis has been used for the
false moneywort rather than a combination based on
Hedysarum nummularifolium L. Schindler (1926)

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188

188

191
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Rhodora [Vol. 76

interprets the Linnaean material as being Indigofera
echinata, in part, and Alysicarpus monilifer, in part,
hence the choice of Hedysarum vaginale L. as the
basionym rather than the earlier one,
Ecastophyllum ecastophyllum > Dalbergia ecastophyl-
lum (L.) Taub. Dalbergia’s having been conserved
over Ecastophyllum eliminates the problem of the
tautonym in B&M.

Ichthyomethia | piscipula — Piscidia piscipula (L.)
Sarg. Piscidia has been conserved over I[chthyome-
thia. (See Rudd, 1969).

Abrus abrus — Abrus precatorius L. The later epithet
of Linnaeus is used to avoid the tautonym.

Bradburya floridana — Centrosema floridanum (Britt.)
Lakela.

Bradburya virginiana — Centrosema virginianum (L.)
Benth. Bradburia has been conserved for a genus of
Compositae over Bradburya of the Leguminosae. See
also Lakela (1963).

Canavali lineata — Canavalia rosea (Sw.) DC.

Canavali bahamensis > Canavalia nitida (Cav.) Piper.
Canavalia is conserved over Canavali. Sauer (1964)
revised species of Canavalia, but apparently over-
looked Johnston’s note (1949b) on publication of
names referring to C. maritima. The basionym of
this oft-used binomial has been treated as Dolichos
maritima Aubl., but Thouars, in making the combina-
tion Canavalia maritima, did not in fact cite the
Aublet name; hence it is not a new combination, but
a new name dating from 1813. Aublet’s name cannot
then be transferred to Canavalia because it is pre-
émpted by the Thouars combination, albeit for the
same species. Alas, another name proposed in the in-
terval is the oldest which must be used, Canavalia
rosea (Sw.) DC. Species 2 of B&M (Canavali gladi-
ata) is probably not in the Bahamas. All specimens
bearing this determination by either Britton or Mills-

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193

193

193

194

195

195

197
197

Bahama Flora — Gillis 93

paugh have been annotated by Sauer as C. maritima
(=C. rosea).

Cajan cajan —> Cajanus cajan (L.) Millsp. Cajanus
is conserved over Cajan. The tautonym of B&M can
thus be avoided.

Dolicholus reticulatus ^ Rhynchosia reticulata (Sw.)
DC.

Dolicholus swartzii > Rhynchosia swartzii (Vail)
Urb.

Dolicholus minima > Rhynchosia minima (L.) DC.
Rhynchosia is conserved over Dolicholus. Dr. John
Grear indicates (personal communication) that
Rhynchosia caribaea based on Glycine caribaea Jacq.
is not in the flora, thus agreeing with B&M. The
latter is ironically not native to the New World, but
is restricted to South Africa.

Phaseolus lathyroides ^ Macroptilium lathyroides (L.)
Urb. On the basis of having five calyx teeth rather
than four, Macroptilium is segregated from the large
genus Phaseolus.

Dolichos lablab > Lablab purpureus L. Many workers
(including Adams, 1972) use the generic segregate
Lablab in preference to Dolichos. If one follows this
trend, then the change above is indicated. Otherwise,
Dolichos lablab may continue to be used.

Dolichos insularis ^ Oxyrhynchus volubilis Brande-
gee. This change follows studies by Rudd (1967).
Vigna repens > Vigna luteola (Jacq.) Benth. in Mart.
Although the epithet repens is the oldest name, it is
invalidated in Vigna by V. repens Baker, published
in 1876 (Merrill, 1910).

OXALIDACEAE
Inoxalis intermedia > Oxalis intermedia A. Rich.

Xanthoxalis corniculata > Oxalis corniculata L. Mod-
ern treatment of Oxalidaceae would merge /noxalis
and Xanthoxalis into Oxalis (Eiten, 1963).

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Rhodora [Vol. 76

LINACEAE
Cathartolinum curtissii ^À Linum medium var. texa-
num (Planch.) Fern.
Cathartolinum corallicola —^ Linum bahamense var.
corallicola (Small) Rogers.
Cathartolinum bahamense > Linum bahamense North-
rop var. bahamense.
Cathartolinum bracei —^ Linum bahamense var. bracei
(Small) Rogers.
Cathartolinum lignosum — Linum bahamense North-
rop. var. bahamense. Rogers (1963; 1968) has restored
Linum as the correct name for the yellow-flowered
species in the West Indies. I have followed his 1963
treatment chiefly.

ZYGOPHYLLACEAE
Guaiacum — consult Porter (1972).

MALPIGHIACEAE

Byrsonima cuneata — Byrsonima lucida (Mill.) DC.
It is generally agreed that the plum-berry or locust-
berry should be Byrsonima lucida, based on Malpighia
lucida, an earlier epithet than cuneata. But upon
whose M. lucida? Most authors attribute the name
to Swartz, but his M. lucida is a later homonym for
M. lucida Mill. Both names can be typified by material
at BM. DeCandolle (1824) specifically indicated the
Swartz material as basionym, but it is illegitimate as
a later homonym. Kunth (1923) indicated that he
thought that M. lucida should be a Byrsonima but did
not make the transfer in accord with the Code. Rather
than presume to make a new combination at this date
myself, I interpret this situation in the light of his-
tory. Swartz undoubtedly knew of Miller’s name and
probably even examined herbarium material from the
Chelsea Physic Garden upon which the lectotype of
M. lucida Mill. (chosen by Gillis) has been selected.
That he (Swartz) did not specifically indicate that

1974]

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219

220

Bahama Flora — Gillis 95

he was taking up Miller's name is not important, it
seems to me. There is ample precedent with Jacquin
and Linnaeus, for instance, in assuming that the later
author knew of the earlier work but did not, under
existing convention, always cite the author of the
earlier binomial, or place of publication. With this
assumption, I shall follow Robertson (1971), in at-
tributing the combination to DeCandolle with a lecto-
type of Miller's name at BM.

SIMAROUBACEAE
Cronquist's treatment of Alvaradoa (1944) would
make the Bahamian populations Alvaradoa amorpho-
ides subp. psilophylla (Urb.) Cronq.

BURSERACEAE
Elaphrium inaguense > Bursera inaguensis Britt.
Elaphrium simaruba — Bursera simaruba (L.) Sarg.
Bursera is conserved over Elaphrium.

POLYGALACEAE
Badiera oblongata > Polygala oblongata — (Britt.)
Blake. It seems desirable not to segregate the woody
species of Polygala into the genus Badiera.

EUPHORBIACEAE

Cicca disticha — Phyllanthus acidus (L.) Skeels. This
name change follows Webster (1956-58).
Margaritaria bahamensis > Margaritaria tetracocca
(Baill) Webster. Webster (1956-58: J. Arnold
Arbor. 38: 66) found that an earlier epithet applied
to this species.

Xylophylla epiphyllanthus —^ Phyllanthus epiphyllan-
thus L. This treatment follows Webster (1967).
Phyllanthus pruinosus > Phyllanthus caroliniensis
subsp. saxicola (Small) Webster.

Phyllanthus pentaphyllus > Phyllanthus pentaphyllus
C. Wright subsp. pentaphyllus. This treatment fol-
lows Webster (1970).

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225

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229

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Rhodora [Vol. 76

Croton flocculosus Geisl. > Croton flavens var. bal-
samiferus (Jacq.) Muell.-Arg. It appears to me that
Croton flocculosus is best treated as a variety of the
widespread Croton flavens. It matches well the lecto-
type of Croton balsamiferus Jacq. (lectotype from
Martinique at BM), the basionym.

Curcas curcas > Jatropha curcas L. Placement of the
physie-nut in Jatropha seems to be the best place-
ment taxonomically, and also avoids a tautonym.
Adenoropiwm gossypifolium — Jatropha gossypifolia
L. Adenoropium is best treated as a subdivision of
Jatropha.

Argythamnia — consult Ingram (1967).

The place of publication of Acalypha alopecuroidea
Jacq. should be Collect. 3: 196. 1790.

Acalypha ostryaefolia > Acalypha ostryifolia Ridd.
Pera bumeliaefolia > Pera bumeliifolia Griseb. These
orthographic changes are required by the Seattle
Code.

Manihot manihot > Manihot esculenta Crantz. This
name is the earliest available to replace the tautonym
used in B&M.

Gymnanthes lucida — Ateramnus lucidus (Sw.) Rothm.
Rothmaler (1944) and Dandy (1967) have resur-
rected Ateramnus P. Browne to replace the later
Gymnanthes Sw. Although I am of the opinion that
Gymnanthes should be conserved over Ateramnus,
conservation was already rejected once by the Special
Committee for Pteridophyta and Phanerogamae
(Taxon 3: 241. 1954). 'There appears to be no
choice but to adopt the new combination in
Ateramnus.

Adenorima gymnonota > Euphorbia gymnonota Urb.
Adenorima is treated as a section of Huphorbia by
Webster (1967) and that treatment is followed here.
Arthrothammus | cassythoides > Euphorbia cassytho-
ides Boiss. Arthrothamnus differs too little from
other forms of Euphorbia to be segregated from it.

1974]

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238

238

238

238

243

244

247

248

248

248

249

Bahama Flora — Gillis 97

Tithymalus | trichotomus > Euphorbia trichotoma
H.B.K. Tithymalus is treated by Webster (1967) as
a section of Euphorbia.

Aklema petiolaris > Euphorbia petiolaris Sims.
Aklema differs too little from other forms of Eu-
phorbia to be segregated from it.

Chamaesyce wilsonit > Chamaesyce lecheoides var.
wilsonii (Millsp.) Burch.

Chamaesyce | lecheoides > Chamaesyce — lecheoides
Millsp. var. lecheoides.

Chamaesyce insulae-salis ^ Chamaesyce centunculo-
ides (HBK.) Millsp.

Chamaesyce exumensis —^ Chamaesyce lecheoides var.
exumensis (Millsp.) Burch. These name changes fol-
low the treatment by Burch (1966).

BUXACEAE
Tricera bahamensis ^ Buxus bahamensis Baker in
Hook. Tricera is best treated by the earlier name
Buxus (Howard, 1962).

ANACARDIACEAE

Poison-ivy in the Bahamas is the typical subspecies:
Toxicodendron radicans (L.) Kuntze subsp. radicans
(see Gillis, 1971).

CELASTRACEAE
Rhacoma coriacea, ^ Crossopetalum coriaceum North-
rop.
Rhacoma — crossopetalum — Crossopetalum rhacoma
(Sw.) Hitchc.
Rhacoma aquifolia + Crossopetalum aquifolium
(Griseb.) Hitchc.
Rhacoma ilicifolia > Crossopetalum ilicifolium (Poir.)
Kuntze. Crossopetalum was revived by Brizicky
(1964) as the earliest name for this genus.
Although several recent rorkers have accepted Cas-
sine as a widespread genus that would include
Elaeodendron (Adams, 1972 and Ding Hou, 1963),

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253

257

257

258

258

263

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Rhodora [Vol. 76

I have chosen to follow Robson (1965) to recognize
the segregate genus Elaeodendron for the Bahamas,
thus leaving Cassine sensu stricto for a small African
genus.

SAPINDACEAE

Thyana discolor > Thouinia discolor Griseb. Thou-
inia Poit. is an earlier name than Thyana Hamilt.
and is conserved over Thouinia Thunb. ex L.f.
Allophylus cominia — Allophylus cobbe (L.) Raeusch.
Leenhouts (1967) makes a strong case for recognizing
only one worldwide species in this genus. He has
found no morphological gaps which can be used to
distinguish separate populations in any geographical
sense. Having chosen to recognize only one species,
Leenhouts chose A. cobbe as the all-encompassing
name from two Linnaean names of the same date.
Melicocca bijuga — Melicoccus bijugatus (L.) Jacq.
The correct spelling of this name was worked out by
Brizicky (1963).

RHAMNACEAE
Sarcomphalus taylori — Ziziphus taylori (Britt.)
M. C. Johnston. Johnston’s treatment (1964) merges
Sarcomphalus with Ziziphus.
Colubrina reclinata > Colubrina elliptica (Sw.) Briz.
et Stern.
Colubrina cubensis > Colubrina cubensis var. flori-
dana M. C. Johnston.
Colubrina colubrina > Colubrina arborescens (Mill.)
Sarg. These changes follow the monographic treat-
ment by Johnston (1971).

TILIACEAE
Triumfetta — consult Ko Ko Lay, 1950.

MALVACEAE
Phymosia — consult Fryxell (1971).
Gayoides crispum — Herissantia crispa (L.) Brizicky.

1974]

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268

268

271
271

273

274

Bahama Flora — Gillis 99

The earliest generic name available for this segre-
gate of Abutilon is Herissantia (See Brizicky, 1968).
Sida ciliaris — Sida ciliaris L. var. ciliaris.

Sida carpinifolia —^ Sida acuta subsp. carpinifolia
(L.f.) Borss.

Sida, spinosa, — Sida spinosa var. angustifolia (Lam.)
Griseb.

Sida hederaefolia > Sida javanensis Cav. emend.
Borss. Recent treatments of Sida (Kearney, 1954b;
Clement, 1957; Borssum Waalkes, 1966) do not all
agree on treatment of the species in the Bahamas. I
have followed Clement and Borssum Waalkes here.
Malache scabra — Pavonia spicata Cav. var, spicata.
Malache bahamensis > Pavonia bahamensis Hitchc.
Pavonia is conserved over Malache. Kearney’s
(1954a) treatment is followed here except for one
point. He overlooked the fact that Pavonia scabra
(B. Vogel) Juble et Quentin is a later homonym of
P. scabra Presl. It must be replaced by P. spicata
var. spicata.

Malvaviscus sagraeanus > Malvaviscus arboreus var.
mexicanus Schlecht. Schery (1942) has found an
older name than the one given in B&M.

Hibiscus bahamensis — Hibiscus brittonianus Kear-
ney. Because H. bahamensis Britt. is a later homo-
nym, Kearney (1954c) published a new name.
Although a number of recent publications place okra
in the genus Hibiscus as H. esculentus, Bates (1965)
and Borssum Waalkes (1966) retained it in Abel-
moschus, as in B&M, on the basis of the spathelike
calyces which are basically adnate to the staminal
tube and corolla, and which are circumscissilely de-
ciduous.

Pariti tiliaceus — Hibiscus tiliaceus L. subsp. elatus
(Hochr.) Borss. Borssum Waalkes (1966) is followed
in his treatment of this taxon.

Gossypium barbadense — Gossypium arborescens var.
nadam (Watt.) Prockh.

100

Rhodora [Vol. 76

214 Gossypium punctatum > Gossypium hirsutum var.

276

punctatum (Schum.) Hutch. Hutchinson (1943) is
followed for the treatment of indigenous cotton

STERCULIACEAE

The use of names in Melochia has been thoroughly
considered by Goldberg (1967) who considered Mo-
luchia and Melochia synonymous. The only changes
from B&M therefore are orthographic, Because the
keys in B&M are inadequate, a new key to the Bahama
species is given here:

1. Flowers in sessile glomerules, many per cluster;
sinus between calyx teeth acute; fruit a 5-parted
coccus less than 3.5 mm. in diameter ........
SNMP Melochia nodiflora Sw.

. Flowers pedicelled, 3-10 per cluster; sinus be-
tween calyx teeth rounded to truncate; fruit
a 5-angled capsule, greater than 5 mm. in
diameter .......... ee RII 2.
2. Capsule with rounded angles and unbranched

hairs; inflorescences mostly in axillary cymes
or appearing terminal; foliage leaves
glabrous or with scattered, simple hairs;
leaves only slightly lighter in color on lower
surface than on upper ........... enne
IM Melochia pyramidata L.
Capsule sharply angled, with dense stellate
hairs; inflorescences mostly opposite the
leaves, never terminal; foliage leaves densely
stellate; leaves tending toward being dis-
colorous, much lighter on the lower surface
than upper due to dense pubescence ......
2.3232 *2»*«***3** Melochia tomentosa L.

278 Waltheria americana —^ Waltheria indica L. An ex-

amination of Linnaean material and resultant lecto-
typification has resolved the question of a name for

1974]

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280

284
284

Bahama Flora — Gillis 101

the common Waltheria in accord with Brizicky (1966)
and the example used in the International Code (Art.
57). The lectotype (chosen by Gillis) of Waltheria
indica L. is in the Hermann Herbarium at BM. The
lectotype of Waltheria americana L. is at LINN
(Savage Catalog No. 852: 1). They represent the
same taxon; both names originate from the same
date (1753). Robert Brown (in Tuckey, Narr. Exp.
River Zaire 484. 1818) appears to be the first to
adopt the name W. indica L. for the combined species,
and therefore should be followed.

The descriptions and separation of the two species
of Waltheria in B&M are ambiguous. The foliage of
W. bahamensis is decidedly bronze in the field; her-
barium specimens do not show this character well,
but it is useful as a field character. The following
key should assist in making determinations:

Foliage green; branches of stellate hairs 2-4 mm.
Ing 550500 Lot Waltheria indica L.
Foliage bronze; pubescence in small tufts, branches
of stellate hairs less than 1 MIN ...... ers
Waltheria bahamensis Britt.

Ayenia pusilla > Ayenia insulicola Cristobal. Except
for an orthographic change necessitated by the Seattle
Code, the Ayenia species should be named according
to Cristobal (1960).

HYPERICACEAE
Ascyrum linifolium > Hypericum hypericoides (L.)
Crantz. The nomenclature of Ascyrum has under-
gone much discussion (Adams, 1957; Adams and
Robson, 1961). The treatment of Adams and Robson
(ibid.) has been followed here,

FLACOURTIACEAE

Myroxylon ilicifolium > Xylosma ilicifolia Northrop.
Myroxylon bahamense —^ Xylosma bahamensis (Britt.)
Standl. Xylosma G. Forst. has been conserved over

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Rhodora [Vol. 76

Myroxylon J. R. et G. Forst. as a genus of Flacourtia-
ceae (Myroxylon L. has been conserved in the Legu-
minosae). The question of whether these two species
are, in fact, distinct still needs to be explored further.

PASSIFLORACEAE
Passiflora pallida > Passiflora suberosa L. The two
names above are synonyms of the same date; they
were united by Killip (1938). See also Brizicky, 1961.
Passiflora species in the Bahamas will be evaluated
in a separate paper.

LOASACEAE
Mentzelia — consult Darlington (1934).

LYTHRACEAE
Parsonsia parsonsia — Cuphea parsonsia (L.) R. Br.
Parsonsia is conserved for a genus of Apocynaceae;
it therefore cannot be used for a genus of Lythraceae.

COMBRETACEAE (TERMINALIACEAE)
Conocarpus erecta > Conocarpus erectus L. The
generic name Conocarpus should be treated as mascu-
line in accordance with the International Code
(Recommendation 75A). In Stearn (1966) the sub-
stantive termination -carpus is consistently treated
as masculine. Stearn (personal communication) sug-
gests that Linnaeus may have treated Conocarpus as
feminine by analogy with many classical second-
declension names of trees.

MYRTACEAE

Eugenia buxifolia > Eugenia foetida Pers. The
Myrtaceae need to be reworked in the light of Mc-
Vaugh’s generic guidelines and definitions (1956;
1968). The change indicated above is made in accord
with his analysis of types (McVaugh, 1973).
Pimenta pimenta — Pimenta dioica Merr. This change
is made to avoid a tautonym.

1974]

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306

306

310

313

315

316

316

Bahama Flora — Gillis 103

Anamomis longipes > Psidium longipes (Berg) Mc-
Vaugh var. longipes.

Anamomis bahamensis — Psidium longipes var. orbi-
culare (Berg) McVaugh.

Anamomis lucayana — Myrcianthes fragrans (Sw.)
McVaugh. The first two of these changes is in accord
with the interpretation of these species by MeVaugh
(1973). Psidium longipes var. orbiculare is further
elaborated in the same paper by McVaugh by de-
scribing unnamed races which can be distinguished.
Myrcianthes fragrans is defined in an earlier paper
by MeVaugh (1963).

ONAGRACEAE

Jussiaea suffruticosa > Ludwigia octovalvis subsp.
sessiliflora (Micheli) Raven. This name change fol-
lows the treatments of Ludwigia by Raven (1962;
1968).

UMBELLIFERAE (AMMIACEAE)

Foeniculum foeniculum — Foeniculum vulgare Gaertn.
This change is made to avoid a tautonym.

MYRSINACEAE

Icacorea paniculata — Ardisia escallonioides Cham.
et Schl.

Icacorea guadalupensis > Ardisia obovata Desv.
Ardisia is conserved over Icacorea. The Bahama
plants need further study and interpretation, but they
appear to be named best as stated above.

Rapanea guianensis > Myrsine floridana A. DC.
Rapanea guianensis Aubl. may be typified by a speci-
men of Aublet's in P (lectotype of Gillis). It repre-
sents a plant with lustrous, large leaves from northern
South America. The northern West Indian and Flo-
ridian species has dull, smaller leaves. Futhermore,
it represents the group of species with a globose
stigma, a character which has been used to separate
Myrsine from Rapanea. Stearn (1969) made the

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323

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Rhodora [Vol. 76

combination Myrsine punctata (Lam.) Stearn for this
species, but the name is a later homonym for a species
of the Pacifie region which had been published several
years earlier. The earliest available name for this
plant is Myrsine floridana A. DC. If one prefers
this species in Rapanea, then its name should be
R. punctata (Lam.) Lundell.

PRIMULACEAE

Samolus floribundus — Samolus parviflorus Raf. This
change is made in aecordance with Channell and Wood
(1959) ; it is based on an earlier epithet.

SAPOTACEAE

Chrysophyllum — consult Cronquist (1945).
Sideroxylon foetidissimum — Mastichodendron foeti-
dissimum (Jacq.) Cronq. This change is made in ac-
cord with the treatment by Cronquist (1946a).
Lucuma serpentaria > Pouteria domingensis (Gaertn.)
Cronq. var. domingensis. This change follows the
treatment by Cronquist (19462).

Dipholis salicifolia > Bumelia salicifolia (L.) Sw.
Although Cronquist (1945) retained Dipholis as a
segregate genus, Stearn (1968) merged it with
Bumelia. It is the latter treatment which is followed
here.

Bumelia angustifolia > Bumelia celastrina H.B.K.
Bumelia loranthifolia > Bumelia americana (Mill.)
Stearn subsp. americana.

Bumelia bahamensis > Bumelia americana (Mill.)
Stearn subsp. americana. The Bumelia complex in
the West Indies has been treated by Stearn (1968),
a treatment followed here. The following key should
help clarify the species in the Bahamas:

1. Ovary glabrous; endosperm copious; leaves
longer than 7 cm.; terminal shoots never be-
coming thorns .. Bumelia salicifolia (L.) Sw.

1. Ovary pubescent; endosperm little or none;

1974] Bahama Flora — Gillis 105

leaves shorter than 6 em. ; terminal shoots often
thorny a C ok oie CE ELE 2.
2. Leaves narrow, linear to spathulate, 2-10
mm. broad (juveniles may be larger) ; fruit
oblong; plant inhabiting shores, mangal,
and SUNG areas. va ck ee ts re

2. Leaves obovate to oblanceolate, 1-4 mm.
broad; fruits globose; inhabiting uplands ..
A cR Bumelia americana, (Mill.) Stearn
subsp. americana.

324 Mimusops emarginata — Manilkara bahamensis
(Baker) Lam. et Meeuse. Probably no species in the
Bahamas is easier to identify and yet harder to name
than the wild dilly! In addition to the long list of
synonyms given in B&M, it has been called Achras
emarginata (L.) Little, Mimusops jamaiqui (Wright)
Dubard, M. jamaiqui subsp. emarginata, (L.) Cronq.,
and Manilkara parvifolia (Nutt.) Dubard, to name
a few. For the moment, I have selected Manilkara
bahamensis (Baker) Lam. et Meeuse. Thus, it re-
sides in the same genus preferred by Moore and
Stearn (1967) for its close relative, the sapodilla,
and also preferred by Cronquist (1945).

324 Sapota achras — Manilkara zapota (L.) P. van Royen.
By the same token as with the wild dilly (above), the
cultivated sapodilla has had a variety of names,
many of which are permutations of each other. Moore
and Stearn (ibid.) analyzed the problem thoroughly
and leave a choice of names depending upon the
breadth of one's generic concepts. Along with them,
I choose the one indicated above.

EBENACEAE
325 Maba crassinervis > Diospyros crassinervis (Krug
et Urb.) Standl. Maba is treated as being a synonym
of Diospyros. The name given above in Diospyros is
temporary. A new combination will soon be pub-
lished by Mr. Frank White (Oxford).

106

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331

331
332

333

334
334

334

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Rhodora [Vol. 76

OLEACEAE

The Mayepea bumelioides of B&M will be placed in
Chionanthus by Stearn at a later date, at which time
a new combination will be published.

GENTIANACEAE
Based on Shinners’s typification (1957), the name of
the marsh gentian remains the same, but the author
of the binomial and place and date of publication
change from what is given in B&M: Eustoma exalta-
tum (L.) Salisb., Parad. Lond. t. 34. 1806.
Sabbatia campanulata — Sabatia stellaris Pursh.
Sabbatia simulata — Sabatia stellaris Pursh, Wilbur
(1955) and Perry (1971) have studied the taxonomy
of Sabatia in detail. As a consequence, one species
only is considered to be in the Bahamas. The correct
spelling of the generic name should also be noted.

MEN YANTHACEAE
Nymphoides aureum — Nymphoides grayana (Griseb.)
Kuntze. This name change is made in accord with
the treatment by Ornduff (1969). It should be noted
that this so-called endemic Nymphoides (according
to B&M) is also found in Cuba.

APOCYNACEAE
Plumiera obtusa — Plumeria obtusa L. var. obtusa.
Plumiera inaguensis > Plumeria obtusa L. var. ob-
tusa.
Plumiera bahamensis —^ Plumeria obtusa L. var. ob-
tusa.
Plumiera rubra — Plumeria rubra L.
Plumiera sericifolia, + Plumeria obtusa var. sericifolia
(C. Wright) Woodson, Woodson (1938) reworked
the species of Plumeria, taking up the correct spelling
of the generic name. He considered several of the
"species" of B&M to be insular races of P. obtusa
var. obtusa.

1974]

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336

336

337

337

538

Bahama Flora — Gillis 107

Woodson (1936) recognized Neobracea bahamensis
as endemic, but also noted that there are three other
species in Cuba, i.e, it is not a monotypic genus as
stated in B&M.

Stearn (1964) confirmed the name of the periwinkle
as Catharanthus roseus (L.) G. Don,

Echites echites > Echites umbellata Jacq. var. um-
bellata.

Rhabdadenia paludosa > Rhabdadenia biflora (Jacq.)
Muell.-Arg.

Rhabdadenia sagraei —^ Angadenia berteri (A. DC.)
Miers.

Urechites lutea — Urechites lutea var. serica Long.
Woodson (1936) redefined the generic limits of a
number of American Apocynaceous plants. His treat-
ment is followed here. Long (1970a), in studying
populations in South Florida, treated Urechites such
that the Bahamian populations are recognized as a
distinct pubescent population, U. lutea var. sericea.
Because the Apocynaceous vines are not adequately
keyed in B&M, and because the names of all of
them have been changed, the following key has been
prepared for use in separating them:

KEY TO BAHAMIAN APOCYNACEOUS VINES

1. Corolla white or ivory, usually more than 5 cm.
long; leaves remaining flat or folding along the
midrib when pressed ; follicles of pair divergent,
not tending to curve back like pincers, 4-10 mm.
ihiek sou ily doe rer EDITI 2.
2. Follicles of a pair widely divergent (greater

than 180?) ; tube of corolla salverform with
rotate lobes; cylindric part of corolla 10-15
times the length of the sepals; calyx lobes
free nearly to base, linear to linear-lanceolate

. Echites umbellata Jacq. var. umbellata.

2. Follicles of a pair divergent less than 90°;
tube of corolla funnelform; cylindric part of

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Rhodora [Vol. 76

corola 2-3 times the length of the sepals;
calyx lobes free scarcely more than 2/3 the
length, ovate, to 2.5 mm. broad ..........

.. Rhabdadenia biflora (Jacq.) Muell.-Arg.

1. Corolla yellow, usually less than 5 cm. long;
leaves tending to curl under at the edges when
pressed ; follicles of a pair diverging at an angle
of less than 60°, curving back to touch near their
tips, resembling a pair of pincers, 1.5-3 mm.
thick .............eleele eh 3.
3. Vine scarcely 1 m. long; corolla 2-4 cm.
long; leaves oblong, usually less than 1 cm.
broad; pedicels glabrous; calyx lobes free
less than half their length, 1-1.5 mm., acumi-
nate; cylindric part of corolla visible most

of its length (i.e, not hidden by sepals);
petiole 2 mm. long .....................
TEE Angadenia berteri (A.DC.) Miers

3. Vine up to 3.5 mm. long; corolla 4-5 cm.
long; leaves obovate or elliptic to 3 cm.
broad; pedicels densely pubescent; calyx
lobes free nearly to base, 8-12 mm. long,
linear; cylindric portion of corolla hidden
by sepals; petiole 1 em. long .............
ZEE Urechites lutea var. sericea Long

Rauwolfia ` tetraphylla > Rauvolfia nitida Jacq. Rao
(1956) thought that Rauvolfia tetraphylla is indige-
nous to the Greater Antilles, Central America, and
South America as far as Peru and Venezuela, The
Bahamian species is R. nitida, which is not a synonym
of R. tetraphylla as suggested in B&M.

Vallesia glabra — Vallesia antillana Woodson. Wood-
son (1937, 1938b) found that the true Vallesia glabra
is a plant of Mexico and the Pacific Coast of South
America. The species of southern peninsular Florida
and the West Indies that has corollas twice the length
of those of V. glabra needed a name. The name V.
glabra of B&M is thus misapplied.

1974]

342

342
343

343

343

343

344

347

B47

347

Bahama Flora — Gillis 109

ASCLEPIADACEAE

Metastelma | northropiae > Cynanchum northropiae
(Schltr.) Alain,

Metastelma hamatum > Cynanchum caribaeum Alain.
Metastelma inaguense > Cynanchum inaguense (Vail)
Howard et Dunbar.

Metastelma linearifolium > Cynanchum savannarum
Alain.

Metastelma eggersii > Cynanchum eggersii (Schltr.)
Alain.

Metastelma palustre > Cynanchum angustifolium
Pers. Metastelma species are considered synonymous
with Cynanchum. (See Woodson, 1941; Alain, 1955).
Much work needs to be done on the Bahamian species,
but almost certainly several will be united. I shall
refrain from making transfers to Cynanchum of
several names which have not already been trans-
ferred because I feel certain at this point that these
names are synonyms of other names in the flora.
(See also Howard and Dunbar, 1964). Merrill and
Hu (1949) discovered that Muhlenberg had found
Cynanchum angustifolium to be the eariest legitimate
name for the last species listed above. They indicated
the extensive synonyms for this species, a few of
which had never ben included in the standard indices.
Philibertella clausa > Sarcostemma clausum (Jacq.)
R. & S. Sarcostemma is an earlier name than Phili-
bertella (Woodson, 1941).

CONVOLVULACEAE

Evolvulus glaber —^ Evolvulus convolvuloides ( Willd.)
Stearn.

Evolvulus alsinoides > Evolvulus alsinoides var. gri-
sebachianus Meissn. in Mart.

Evolvulus linifolius > Evolvulus alsinoides var. lini-
folius (L.) Baker. Stearn (1972) has modified the
nomenclature of several species of Evolvulus in the
flora in the course of typification of their names. The

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Rhodora [Vol. 76

last taxon listed above is now believed not to occur
in the flora.

Jacquemontia jamaicensis ^ Jacquemontia havanensis
(Jacq.) Urb. Dr. Kenneth Robertson (personal com-
munication) has pointed out that this name change
is required because of an earlier basionym.
Calonyction aculeatum — Ipomoea alba L.
Calonyction tuba — Ipomoea macrantha R. et S. Both
Ooststroom (1940) and Gunn (1972) have worked
with Calonyction. Gunn’s nomenclature is followed
here.

Quamoclit quamoclit ^ Ipomoea quamoclit L.
Quamoclit coccinea > Ipomoea hederifolia L. Quamo-
clit, like Calonyction, is treated as a section of Ipo-
moea. (see Ooststroom, 1953).

Ipomoea cathartica — Ipomoea acuminata (Vahl) R.
et S.

Ipomoea pes-caprae > Ipomoea pes-caprae subsp. bra-
Siliensis (L.) Ooststr. This treatment of Ipomoea
follows that of Ooststroom (1940) and St. John
(1970). The citation for Ipomoea stolonifera in B&M
is incorrect; there is an earlier publication date of the
combination: Ipomoea stolonifera (Cyrill.) Gmel.,
Syst. Veg. I: 345. 1796.

Ipomoea dissecta + Merremia dissecta (Jacq.) Hall. f.
Merremia is segregated from Ipomoea on the basis of
its smooth pollen and its corolla without a red or
purple eye (O'Donell, 1941; Ooststroom with Hoog-
land, 1953; Verdcourt, 1963).

Dichondra. Consult Tharp and Johnston (1961).

CUSCUTACEAE
Cuscuta pentagona — Cuscuta campestris Yuncker.
This species needed a new name because of confusion
in the old one (Yuncker, 1932).
HYDROPHYLLACEAE
Marilaunidium jamaicense > Nama jamaicensis L.

1974]

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358

358
359

359

360

361

362

363

365

366

Bahama Flora — Gillis 111

Nama is an earlier name than Marilaunidium. Fur-
thermore, Nama jamaicensis L. is the conserved type
species.

BORAGINACEAE (EHRETIACEAE)

Sebesten sebestena — Cordia sebestena L.

Varronia globosa > Cordia globosa var. humilis
(Jaeq.) Johnston.

Varronia bahamensis > Cordia bahamensis Urb.
Varronia brittonii > Cordia brittonii (Millsp.) Mac-
bride.

Varronia lucayana — Cordia lucayana (Millsp.) Mac-
bride. Sebesten and Varronia should be treated as
synonyms of Cordia (Macbride, 1916; Johnston,
1949).

Rochefortia bahamensis — Rochefortia spinosa (Jacq.)
Urb. According to Lefor (1968), the Rochefortia in
the Bahamas is not an endemic species, but the same
as that found in Cuba.

There is debate over the generic selection for bay
lavender. For the moment, I maintain Mallotonia,
as in several other recent floras (Gooding, Loveless,
and Proctor, 1965; Adams, 1972).

Heliotropium parviflorum — Heliotropium angiosper-
mum Murray. The change of name is required because
H. angiospermum is an earlier name.

Heliotropium inundatum — Heliotropium procumbens
Mill. Heliotropium procumbens is an earier name for
the same species. Both names can be typified by
specimens at BM; the lectotypes have been selected
by me.

VERBENACEAE

Valerianoides fruticosa > Stachytarpheta fruticosa
(Millsp.) B. L. Robinson.

Valerianoides jamaicensis —> Stachytarpheta jamai-
censis (L.) Vahl. Stachytarpheta is conserved over
Valerianoides.

112

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367
368

368

369

369

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Rhodora [Vol. 76

Bouchea prismatica > Bouchea prismatica var. longi-
rostra Grenzen. This treatment follows the revision
of Bouchea by Grenzenbach (1926). Moldenke (1971)
believes that the typical variety is also present in
the Bahamas.

Priva — consult Kobuski (1926).

Lippia stoechadifolia > Phyla stoechadifolia — (L.)
Small.

Lippia nodiflora — Phyla nodiflora (L.) Greene var.
nodiflora.

Lippia reptans — Phyla nodiflora var. reptans (H.B.K.)
Moldenke. These changes are made in accord with
the treatment of Phyla by Moldenke (1965). Lippia
geminata of B&M is probably not found within the
archipelago. The appropriate treatment of this bi-
nomial would be Lantana microcephala A. Rich. in
any event.

According to treatment by Long (1970a) our variety
of Lantana ovatifolia would be the typical variety
(var. ovatifolia).

Using Moldenke's treatments (1958a, b, and c) of
Citharexylum, I have prepared the following key to
distinguish the Bahamian species:

Pedicels less than 1.2 mm. long during anthesis;
pyremes 2-loculate, fruit not shiny; leaves + pilose,
closely reticulate-veined, the reticulation prominent
above and below ... Citharexylum fruticosum L.

Pedicels 2 mm. long or more during anthesis;
pyremes 1-loculate, fruit shiny; leaves glabrous,
widely reticulate-veined, reticulation obscure above
Citharexylum caudatum L.

Moldenke (1958a; 1971) has considered 5 infra-
specific taxa of C. fruticosum to exist in the Bahamas:
var. fruticosum, var. smallii Moldenke, var. subvil-
losum Moldenke, var. villosum (Jaeq.) O. E. Schultz,
and forma bahamense Moldenke, Having made field

1974]

373
373
374

375

376

377
378

378

Bahama Flora — Gillis 113

studies on the characters upon which these taxa are
based, I consider these to be insular forms or popula-
tions without names. For those who wish to name
these variations, an appropriate key is found in
Moldenke (1958a).

Callicarpa — consult Moldenke (1936).

Petitia — consult Moldenke (1937).

Volkameria aculeata —^ Clerodendrum aculeatum (L.)
Schlecht. var. aculeatum. Volkameria is treated by
Moldenke (1971) as a section of Clerodendrum. The
place of citation for C. aculeatum is incorrect in
B&M. The author of the combination is Schlechten-
dahl, and the citation is Linnaea 6: 750. 1831.
Clerodendrum fragrans > Clerodendrum philippinum
Schauer. Howard and Powell (1968) found that C.
fragrans was an illegitimate name. Moldenke (1971)
considers the Bahamian population to be var. pleni-
florum Schauer.

Avicennia nitida — Avicennia germinans (L.) L.
Stearn (1958) found this earlier epithet which ap-
plies to the species of black mangrove in the West
Indies, but failed to note that its earliest combination
was by Linnaeus himself.

LABIATAE (LAMIACEAE)

Melosmon cubense — Teucrium cubense Jacq. Epling
(1925) united Melosmon and Teucrium.

Leonurus — consult Epling (1925).

The place of publication for Leonotis nepetifolia (L.)
R.Br. is the Prodromus florae Novae Hollandiae,
p. 504. 1810, and not as given in B&M (Epling,
1925).

In Epling’s treatment (1938-39), Salvia serotina L.
and S. micrantha Vahl (indicated as synonyms in
B&M) were separated on rather minor characters.
Through the courtesy of Dr. Mildred Mathias and
Dr. Raymond Harley, I have seen an unpublished
manuscript of Epling and Carlos Jativa in which

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Rhodora [Vol. 76

these species were reunited. It seems best to continue
to treat them as synonymous. (Permission of Carlos
Jativa to refer to this unpublished material is ac-
knowledged with appreciation.)

On the basis of rather minor characters of the calyx,
Shinners (1962) reworked the Micromeria brownei
complex, recognizing the Bahamian populations as
M. bahamensis Shinners. Because my own collections
from Inagua more closely resemble the taxon which
Shinners cites from Hispaniola, I believe more work
needs to be done before the matter is resolved. Chiefly
because of the minor nature of the characters which
are used in defining Shinners's species, I continue to
recognize a broadly treated M. brownei (Sw.) Benth.

SOLANACEAE

Physalis angulata > Physalis angulata L. var. angu-
lata.

Physalis turbinata — Physalis cordata Mill.
Physalis pubescens > Physalis barbadensis Jacq.
These changes are in accord with the treatment by
Waterfall (1967).

Solanum verbascifoltum — Solanum erianthum D.
Don. Roe (1968) found that the old name for this
plant did not apply to the species in our flora.
Lycium spathulifolium > Lycium tweedianum var.
chrysocarpum (Urb. et Ekm.) Hitche. In the mono-
graph of Lycium (Hitchcock, 1932), the Bahamian
plant required a name change. Although it is possible
that the species of the Florida Keys (L. carolinianum
Walt.) is in the Bahamas, it has yet to be found.

SCROPHULARIACEAE
Maurandya antirrhinaeflora > Maurandya antirrhini-
flora H. et B. ex Willd. This orthographic change is
required by the Seattle Code. I can now report this
species to be on the far side of the archipelago from
New Providence as recorded in B&M: Gillis 11780
from Grand Turk.

1974]

389

390

392

393

394
395

399

401

Bahama Flora — Gillis 115

Bramia monnieri > Bacopa monnieri (L.) Pennell.
Bacopa has been conserved over Brami and Moniera.
It is an earlier name than Bramia.

Although Pennell (1935) segregated Erinus from
Mecardonia, I believe that Mecardonia is best treated
as an inclusive genus, thus maintaining Mecardonia
procumbens (Mill.) Small as in B&M.

Afzelia cassioides > Seymeria cassioides (Walt.)
Blake. Seymeria is conserved over Afzelia.
Buchnera elongata —> Buchnera floridana Gandoger.
Philcox (1965) thinks that B. elongata is a mis-
applied name for the Florida and Bahama popula-
tions.

LENTIBULARIACEAE
Setiscapella subulata — Utricularia subulata L.
Stomoisia cornuta — Utricularia cornuta Michx. Tay-

lor (1967) includes Setiscapella and Stomoisia within
Utricularia.

BIGNONIACEAE

Macrocatalpa punctata — Catalpa punctata Griseb.
var. punctata. This change is made in accord with
the treatment of Catalpa by Paclt (1952).

PEDALIACEAE

Sesamum orientale —^ Sesamum indicum L. These
names are synonymous binomials of the same publica-
tion date. Sesamum indicum L. has been used more
commonly than the other name. Hill (1939) pointed
out that DeCandolle (Pl. Rar. Jard. Genève 18, t.5 —
1889) was the first to unite the two names under
Sesamum indicum.

ACANTHACEAE

Blechum brownei should be retained despite the at-
tempt of some to change the name of this species to
Blechum pyramidatum. The Linnaean binomial is
Ruellia bltechum, Lamarck renamed the species Bar-

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403

404

407

408

409

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Rhodora [Vol. 76

laria pyramidata, but this name is illegitimate, being
superfluous when published. Jussieu later published a
new name in Blechum; this is accepted because the
tautonym which would be created by transferring the
Linnaean epithet is illegitimate (Bremekamp, 1938;
Long, 1970b).

Gerardia droseroides > Stenandrium droseroides Nees
in DC.

Gerardia bracteosa — Stenandrium bracteosum (Britt.
et Millsp. Britt. ex Leonard). Stenandrium is con-
served over Gerardia.

Anthacanthus spinosus — Oplonia spinosa (Jacq.) Raf.
Stearn (1971) has studied the generic complex of
which this species is a part and has recognized our
plant to be an Oplonia.

Diapedium assurgens Dicliptera assurgens (L.)
Juss. Dicliptera is conserved over Diapedium (see
also Long, 1970b). I have chosen the lectotype to be
at LINN (No. 28:23 according to the Savage Catalog,
1945).

RUBIACEAE

The Rubiaceae are in need of in-depth treatment as
a whole. Names accepted here may be subject to re-
vision as more work is done.

Oldenlandia callitrichoides > Hedyotis callitrichoides
(Griseb.) Lewis. Merrill and Metcalf (1942) and
Lewis (1961) have united Oldenlandia and Houstonia
with Hedyotis. Lewis’s treatment is followed here.
Rachicallis americana — Rhachicallis americana (Jacq.)
Hitche. The original spelling of the generic name has
been restored. Despite the fact that it has been
written without the first “h” ever since the original
publieation, there seems no reason not to accept the
original orthography considering the Greek origin.
Nickerson and Tripp (1973) have reported this species
to have either male or perfect flowers, never female.
Randia mitis — Randia aculeata L. These two names,

1974]

411

412

413
413

414

415

Bahama Flora — Gillis 117

synonyms of the same date of publication, have been
united under R. aculeata.

Catesbaea, parviflora — Catesbaea parviflora var. sep-
tentrionalis Krug et Urb. ex Urb. The Bahamian
populations fit this variety better than do the typical
species form.

Hamelia erecta > Hamelia patens Jacq. These names
are synonyms of the same date of publication. Thomas
Elias (personal communication) has informed me
that L’Héritier was the first to unite them, and should
be followed.

Guettarda taylori —^ Guettarda nashii Britt. et Millsp.
Guettarda inaguensis > Guettarda nashii Britt. et
Millsp. There does not seem to be sufficient difference
to maintain three distinct species of Guettarda sup-
posedly endemic to the vicinity of Matthew Town,
Inagua. Having studied both the type specimens (at
NY) and material in the field, I have concluded that
Britton and Millspaugh named specimens, not species.
Guettarda inaguensis is represented by a type that
appears to have come from an aberrant form with
long lengths of bare branches and leaves only near
the tips. Guettarda taylori is represented by a type
that has no flowers left and for which no fruits were
ever known. Because of the inadequancy of these two
specimens, it seems best to select the name for the
united species by the type possessing the most ade-
quate material. Inasmuch as these three species
names originate from the same data, I am choosing
G. nashii as the name to be used when all three are
united. The populations of Guettarda in the vicinity
of Matthew Town, although variable, appear to me
to be best treated as a single species.

Stenostomum lucidum — Antirhea lucida (Sw.) Hook.
f. in Benth. et Hook. f.

Stenostomum myrtifolium — Antirhea myrtifolia
(Griseb.) Urb.

118 Rhodora [Vol. 76

415 Stenostomum densiflorum — Terebraria resinosa
(Vahl) Sprague. Stenostomum as used by B&M is
better divided into Antirhea and Terebraria. Antirhea
is an older name than Stenostomum and hence must
be the name used for those species traditionally placed
in Stenostomum. The tortuous history of the name
of Terebraria resinosa is recounted thoroughly by
Sprague (1932). Sprague recognized two West In-
dian species of Terebraria, differing only in shape
of the areoles of the leaf venation. In his treatment,
the Bahamian population would be T. densiflora. I
consider these differences to represent only insular
forms, not worthy of nomenclatural separation. A
key to the species in the Bahamas of the old Stenos-
tomum follows:

1. Ovary 2 (-4) loculate; flowers without bracts;
anthers half-exserted; stipules deciduous; foli-
age without resinous secretions .... Antirhea
lucida (Sw.) Hook. f. in Benth. et Hook. f.

1. Ovary 4-6 loculate; flowers subtended by minute
bracts; anthers included; stipules persistent;
foliage resinous-viscid ................0-. 2.

2. Leaves 3 cm. long or less, elliptic to obovate-
oblong; diffusely-branched shrub with leaves
crowded near ends of branches; midvein very
pronounced, often white; 2-4 flowers per
branch of inflorescence; stipules ovate, not
forming a collar around stem; inflorescences
paired on either side of stem, but not
branched ................ eee

2. Leaves 4-9 cm. long, oblong-lanceolate ;
coarsely-branched shrub or small tree with
leaves widely scattered; midvein only slightly
more pronounced than lateral veins; more
than 6 flowers per branch of inflorescence;
stipules forming a collar 2-4 mm. broad

1974]

419

420
421

Bahama Flora — Gillis 119

around stem; inflorescence bifid with flowers
only on inside (adaxial surface) of branches
Terebraria resinosa (Vahl) Sprague.

Psychotria undata > Psychotria nervosa Sw. Jacquin
described P. undata in the Plantarum rariorum horti
caesarei schoenbrunnensis. According to Stafleu
(1967), the title page date of 1798 is probably in
error, and actual date of publication may have been
as late as 1803. In any event, Swartz in his Prodro-
mus (1788) published the name Psychotria nervosa
for the same species, clearly the earliest date of pub-
lication. Psychotria nervosa Benth, (1841) and P.
nervosa D. Don (1825) are later homonyms.

Ernodea cokeri > Ernodea taylori Britton.

Ernodea nashii o Ernodea millspaughii Britton. It
is difficult to make hard-and-fast decisions on the
Ernodea problems in the Bahamas at this juncture.
Whether E. littoralis and E. angusta are truly distinct
is questionable. Long (1970a) has treated E. angusta
as a variety of E. littoralis which may be the best
way to handle these perplexing taxa. Until intensive
study is made of these two species with comparisons
to the other Ernodea taxa in the Bahamas, I shall
continue to recognize these two as species. Distinc-
tions among the other four species do not seem to
hold up when examined under field conditions. Al-
though more study will be carried out on Ernodea
in the Bahamas, under a tentative arrangement which
seems reasonable at this stage, E. cokeri is considered
to be a synonym of E. taylori, and E. nashii is merged
with E. millspaughii. The latter name was chosen
from the two originating at the same date of publi-
cation because it is represented by the type specimen
possessing both flowers and fruits. The type of E.
taylori has no flowers at all; the type of E. nashii
must have had flowers at the time Britton described
the species, but they are missing from the type

120

422

425

426

Rhodora [Vol. 76

specimens now (types are at NY). The differences
as observed in the type specimens of these micro-
species seem inadequate for maintenance in this ex-
ceedingly variable genus (also see Britton, 1908, for
descriptions of “races”? within this complex). A key
to identify the Bahamian species of Ernodea within
my species concept at the moment follows:

1. Calyx lobes nearly as long as the fruit, or
longer .............. RR RRRRRRI 2.

2. Corolla white to very pale pink ..........
SENE Ernodea littoralis Sw.

2. Corolla red ...... Ernodea angusta. Small.
1. Calyx lobes less than half as long as the
fruits 2.0... 0 eee 3.

3. Leaves 6-8 mm. wide ...................
MEIN Ernodea millspaughii Britton.
3. Leaves 1-9 mm. wide ............... less
ZEE Ernodea taylori Britton.
Borreria saxicola > Borreria brittonii Standl. Un-
like Ernodea, the eight species of Borreria in the
Bahamas — or, at least most of them — seem to be
good biological species with a high degree of ende-
mism in the southern islands, especially Inagua and the
Caicos group. Whereas actual treatment of this genus
is deferred for the moment, one name change needs
to be cited. Following Standley (1931), the name
B. saxicola Britt. is replaced because it is a later
homonym.
Spermacoce — consult Bacigalupo (1972).

CUCURBITACEAE

Anguria pedata > Psiguria pedata (Jacq.) Howard.
In his treatment of modern names for plants dis-
cussed in Jacquin’s Selectarum, Howard (1978)
found that the name Anguria Jacq. was a later
homonym for Anguria Mil. The next available
generic name is Psiguria; Howard made the neces-
sary new combinations for this species,

1974]

428

429

431

432

432
436

440

441

443

Bahama Flora — Gillis 121
LOBELIACEAE
Lobelia — consult MeVaugh (1943).
GOODENIACEAE
Scaevola plumierii > Scaevola plumieri (L.) Vahl.

This orthographic change is made to conform to the
present International Code.

COMPOSITAE (AMBROSIACEAE
AND CARDUACEAE)

Xanthium chinense — Xanthium strumarium L.
Xanthium strumarium is an earlier name than X.
chinense.
Ambrosia paniculata —^ Ambrosia artemisiifolia L. I
have selected a lectotype of Ambrosia paniculata
from material in the Michaux Herbarium at Paris.
Furthermore, I have examined the lectotype of A.
artemisiifolia which was selected by Payne (1970)
at LINN, and concur with Payne that these plants
are conspecific. The Linnaean name is obviously the
older one and should be used.
Iva — consult Jackson (1960).
Ageratum latifolium —> Ageratum conyzoides subsp.
latifolium (Cav.) M. F. Johnson. This name change
has been made in accord with the treatment by John-
son (1971). The typical subspecies has also been
reported from the Bahamas, thus adding a taxon
to the flora.
Chrysopsis graminifolia > Heterotheca graminifolia
(Michx.) Shinners. Shinners (1951), Wagenknecht
(1960), and Harms (1964) have argued for the
merger of Heterotheca and Chrysopsis.
Aster bracei — Aster tenuifolius var. aphyllus Long.
Long (1970a) has treated the Bahamian populations
in connection with those of South Florida which he
was studying. This taxon seems best treated as a
variety of the more widespread A. tenuifolius.
Leptilon linifolium — Conyza floribunda H.B.K.

124 Rhodora [Vol. 76

CUPRESSACEAE (PINACEAE)

462 Juniperus lucayana — Juniperus bermudiana L. More
work needs to be done on typification and on popula-
tion studies in Juniperus to place the Bahamian plant
properly. For the moment, it is treated in accord
with Moore, 1966.

CYCADACEAE

463 A thorough monograph of Zamia is needed. In a
manuscript by Chamberlain, left unpublished at his
death, Z. lucayana has been referred to the Cuban
Z. guttierezii Sauv. So little material of this plant,
as well as of Z. angustifolia and Z. tenuis, is avail-
able at this time, that it is not appropriate to make
a nomenclatural judgment now. The recent redis-
covery of the Long Island population of “Z. lucayana"
by S. R. Hill (in press) and its subsequent cultiva-
tion will help to make material available for further
study.

LITERATURE CITED

ACUNA, J. and J. T. Rorc. 1962, Psychotriae Cubenses Novae
(Rubiaceae). Brittonia 14: 224-229.

ADAMS, C. D. 1970. Miscellaneous additions and revisions to the
flowering plants of Jamaica. Phytologia 20: 309-14.

1971. Miscellaneous additions and revisions to the

flowering plants of Jamaica. III. Phytologia 21: 405-10.

1972. Flowering plants of Jamaica. University of
the West Indies, Mona, Jamaica. 848 pp.

ADAMS, WILLIAM P. 1957. A revision of the genus Ascyrum (Hy-
pericaceae). Rhodora 59: 73-95.

ADAMS, WILLIAM P. and NoRMAN K. B. RoBSON. 1961. A re-evalua-
tion of the generic status of Ascyrum and Crookea (Gutttiferae).
Rhodora 63: 10-16.

ALAIN, HERMANO. 1962. Flora de Cuba, Vol. 5 — Rubiales - Valeri-
anales - Cucurbitales - Campanulales - Asterales. Editorial Uni-
versitaria, Universidad de Puerto Rico, Rio Piedras.

ALLEN, CAROLINE K. 1966. Notes on Lauraceae of tropical America.
I. The generic status of Nectandra, Ocotea, and Pleurothyrium.
Phytologia 13: 221-27.

1974] Bahama Flora — Gillis 125

AMES, OAKES. 1910. A new Ponthieva from the Bahamas. Torreya
10: 90-91.

BACIGALUPO, NELINDA M. 1972. Observaciones sobre algunas es-
pecies de los géneros Spermacoce L. y Spermacoceodes O.K.
(Rubiaceae). Darwiniana 17: 341-57.

BAEHNI, C. 1965. Mémoires sur les Sapotacées. III. Inventaire des
genres. Boissiera 11: 1-262.

BALDWIN, J. T., JR. 1938. Kalanchoé: the genus and its chromo-
somes. Amer. J. Bot. 25: 572-79.

Bates, DAvID M. 1965. Notes on the cultivated Malvaceae. I. Hi-
biscus. Baileya 13: 56-130.

BEAUVOIS, A. M. F. J. PaLisoT DE. 1812. Essai d'une nouvelle
Agrostrographie. Paris.

BECKNER, JOHN. 1970. Notes on Florida and West Indian orchids.
Phytologia 20: 217.

BELCHER, R. O. 1956. A revision of the genus Erechtites (Compos-
itae) with inquiries into Senecio and Arrhenechthites. Ann. Mis-
souri Bot. Gard. 43: 1-85.

BoGLE, A. LINN. 1970. The genera of Molluginaceae and Aizoaceae
in the southeastern United States, J. Arnold Arbor. 51: 431-62.

BoRSSUM, WAALKES, J. VAN. 1966. Malesian Malvaceae revised.
Blumea 14: 1-213.

BRASHIER, CLYDE K. 1966. A revision of Commelina (Plum.) L.
in the U.S.A. Bull. Torrey Bot. Club 93: 1-19.

BREMEKAMP, C. E. B. 1938. Notes on the Acanthaceae of Surinam.
Rec. Trav. Bot. Neerl. 35: 130-170. + 4 pl.

BRENAN, J. P. M. 1958. New and noteworthy Cassias from Tropical
Africa. Kew Bull. 13: 231-252.

BRITTON, N. L. 1908. The genus Ernodea Swartz: a study of
species and races. Bull. Torrey Bot. Club 35: 203-208.

1911. The botanical name of the wild sapodilla.

Torreya 11: 128-29.
1918. Flora of Bermuda. Charles Scribner's Sons.

New York.

and CHARLES FREDERICK MILLSPAUGH. 1920. The

Bahama flora. Published by the authors. Reprinted 1962, Hafner

Publishing Co., New York.

and JOSEPH NELSON RosE. 1928. Rosales - Mimo-

saceae. North American Flora. Vol. 23, Pt. 1.

1923-30. Scientific survey of Puerto Rico and the
Virgin Islands. Botany of Puerto Rico and the Virgin Islands.
Descriptive flora. Part 1 — New York Acad. Sci. 5 (1): 1-158;
5 (2): 159-316; 5 (3): 317-474; 5 (4): 475-626; 6 (1): 1-158;
6 (2): 159-316; 6 (3): 317-521; 6 (4): 523-663.

BRIZICKY, GEORGE K. 1963. The genera of Sapindales in the south-
eastern United States. J. Arnold Arbor. 44: 462-501.

126 Rhodora [Vol. 76

. 1964. The genera of Celastrales in the south-

eastern United States. J. Arnold Arbor. 45: 206-34.

1966. The genera of Sterculiaceae in the

southeastern United States. J. Arnold Arbor. 47: 60-74.

1968. Herissantia, Bogenhardia, and Gayo-
ides (Malvaceae). J. Arnold Arbor. 49: 278-79.

BuncH, DEREK. 1966. Two new species of Chamaesyce (Euphorbia-
ceae), new combinations, and a key to the Caribbean members
of the genus. Ann. Missouri Bot. Gard. 53: 90-9.

CABRERA, ANGEL L. 1971. Revision del género Gochnatia (Com-
positae). Revista Mus. de la Plata (N.S.— Botanica) 12: 1-160.

CANDOLLE, A. P. DE. 1825. Prodromus. Vol. 2. Paris.

CHANNELL, R. B. and C. E. Woop, Jr. 1959. The genera of the
Primulales of the southeastern United States. J. Arnold Arbor.
40: 268-88.

CHASE, AGNES. 1929. The North American species of Paspalum.
Contr. U.S. Natl. Herb. 28: 1-310 + xvii.

1951. (revision of A. S. Hitchcock) Manual of the
grasses of the Unied States, ed. 2. U.S.D.A. Misc. Publ. 200.
1051 pp.

CLAYTON, W. D. 1964. Studies in the Gramineae: vi. Sporoboleae.
The Sporobolus indicus complex. Kew Bull. 19: 287-96.

1968. The correct name of the common reed.
Taxon 17: 168-69.

CLEMENT, I. D. 1957. Studies in Sida (Malvaceae) 1. A review
of the genus and monograph of the Section Malacroideae, Phy-
salodes, Pseudomalvastrum, Incanifolia, Oligandrae, Pseudona-
paea, Hookeria, and Steninda. Contr. Gray Herb. 180: 5-91.

CoKER, W. C. 1944. The woody Smilaxes of the United States. J.
Elisha Mitchell Soc. 60: 27-69.

COLEMAN, JAMES R. 1966. A taxonomic revision of Section Ximene-
sia of the genus Verbesina L. (Compositae). Am. Midl. Nat.
76: 475-81.

ConpiT, IRA J. 1969. Ficus, the exotic species. Division of Agri-
cultural Sciences, University of California, Berkeley. 363 pp.

CORCORAN, SISTER Mary Lucy. 1941. Revision of the Subgenus
Pycreus in North and South America. Contr. Biol. Lab. Catholic
Univ. Amer. 37: 1-68 + xvi.

CORRELL, DONOVAN STEWART. 1950. Native orchids of North
America, north of Mexico. Chronica Botanica Co., Waltham,
Mass.

CRISTOBAL, CARMEN L. 1960. Revision del género Ayenia (Ster-
culiaceae). Opera Lilloana iv. Tucuman, Argentina, pp. 5-230.

CRONQUIST, ARTHUR. 1943. "The separation of Erigeron from Conyza.
Bull. Torrey Bot. Club 70: 629-632.

1974] Bahama Flora — Gillis 127

1944. Studies in the Simaroubaceae. IV. Ré-
sumé of the American genera. Brittonia 5: 128-47.

1945a. Studies in the Sapotaceae. I. The
North American species of Chrysophyllum. Bull. Torrey Bot.
Club 72: 192-205.

. 1945b. Studies in the Sapotaceae III. Dipholis
and Bumelia. J. Arnold Arbor. 26: 435-71.

1945c. Studies in the Sapotaceae. IV. The
North American species of Manilkara. Bull. Torrey Bot. Club
72: 550-62.

1946. Studies in the Sapotaceae. II. Survey of

the North American genera. Lloydia 9: 241-92.

1947. Notes on the Compositae of the north-
eastern United States. V. Astereae. Bull. Torrey Bot. Club
74: 142-50.

DANDY, J. E. 1967. Index of generic names of vascular plants,
1753-1774. Reg. Veg. 51: 1-130.

and A. W. EXELL. 1938. On the nomenclature of the

species of Caesalpinia. J. Bot. 76: 175-80.

and Geoffrey Tandy. 1939. On the identity of Syrin-
godium Kütz. J. Bot. 77: 114-16.

DARLINGTON, JOSEPHINE. 1934. A monograph of the genus Mentze-
lia. Ann. Missouri Bot. Gard. 21: 103-226 + 6 pl.

DELISLE, DONALD B. 1963. Taxonomy and distribution of the genus
Cenchrus. Iowa State Univ. J. Sci. 37: 259-351.

DEVALL, WILBUR B. 1941. The taxonomic status of Pinus caribaea
Mor. Florida Acad. Sci. Proc. 5 (1940) : 121-32.

DEWIT, H. C. D. 1955. A revision of the genus “Cassia” (Caesalp.)
as occurring in Malaysia. Webbia 11: 197-292.

. 1961. Typification and correct names of Acacia
villosa Willd. and Leucaena glauca (L.) Bth. Taxon 10: 50-54.

DEWOLF, GORDON P. 1960. Ficus. In: Flora of Panama. Ann.
Missouri Bot. Gard. 47: 146-65.

DRESSLER, ROBERT L. 1957. The genus Pedilanthus (Euphorbia-
ceae). Contr. Gray Herb. 182: 3-188.

. 1961. A reconsideration of Encyclia (Orchi-
daceae). Brittonia 13: 253-66.

DUNSTERVILLE, G. C. K. and LESLIE A. GARAY. 1965. Venezuelan
orchids illustrated. Vol. 3. Andre Deutsch Ltd., London.

EITEN, GEORGE 1963. Taxonomy and regional variation of Oxalis,
section Corniculatae. Introduction, keys, and synopsis of the
species. Amer. Midl, Nat. 69: 257-309.

EPLING, CARL CLAWSON. 1925. Studies on South American Labia-
tae. I. Synopsis of the genera Teucrium, Rosmarinus, Marru-
bium, Prunella, Lamium, Leonurus, and Leonotis. Ann. Missouri
Bot. Gard. 12: 107-129.

128 Rhodora [Vol. 76

1938-39. A revision of Salvia: Subgenus
Calosphace. Fedde, Repert. Sp. Nov. 110: 1-380.

ERNST, WALLACE R. 1963. The genera of Capparaceae and Morin-
gaceae in the southeastern United States. J. Arnold Arbor. 44:
81-95.

FassETT, NORMAN C. 1955. Echinodorus in the American Tropics.
Rhodora 57: 133-56; 174-88; 202-12.

FERNALD, M. L. 1934. Reallignments in the genus Panicum. Rho-
dora 36: 61-87. (also Contr. Gray Herb. 103).

and K. M. WIEGAND. 1914. The genus Ruppia in
eastern North America. Rhodora 16: 119-27.

FRYXELL, PAUL S. 1971. A revision of Phymosia (Malvaceae).
Madrono 31: 153-74.

GALE, SHIRLEY. 1944. Rhynchospora, section Eurhynchospora, in
Canada, the United States, and the West Indies. Rhodora 46:
89-134; 159-97; 207-49; 255-78. (Contr. Gray Herb. 151).

GAMERRO, JUAN CARLOS. 1968. Observaciones sobre la biología floral
y morfología de la Potamogetonácea Ruppia cirrhosa (Petag.)
Grande (R. Spiralis L. ex Dum.) Darwiniana 14: 575-608.

GARAY, LESLIE A. and HERMAN R. SWEET. 1972. Notes on West
Indian orchids III. J. Arnold Arbor. 53: 515-30.

GILLIS, WILLIAM T. 1971. The systematics and ecology of poison-
ivy and the poison-oaks (Toxicodendron, Anacardiaceae). Rho-
dora 73: 72-237; 370-448; 465-540.

, RICHARD A. HOWARD, and GEORGE R. PROCTOR.

1973. Additions to the Bahama flora since Britton and Mills-

paugh — I. Rhodora 75: 411-425.

and WILLIAM T. STEARN. in press. Typification
and correct names of Leucaena and Lysiloma species in the Ba-
hama flora. Taxon.

GODFREY, R. K. 1952. Pluchea, section Stylimnus, in North Amer-
ica, J. Elisha Mitchell Sci. Soc. 68: 238-71 + 23 pl.

GOLDBERG, AARON. 1967. The genus Melochia L. (Sterculiaceae).
Contr. U.S. Natl. Herb. 34: 191-363.

GooDprNG, E. G. B., A. R. LovELESS, and G. R. PRocTOR. 1964. Flora
of Barbados. Overseas Research Publication No. 7 (Ministry of
Overseas Development). Her Majesty's Stationery Office. Lon-
don. 486 pp.

GouLD, FRANK W. 1967. The grass genus Andropogon in the
United States. Brittonia 19: 70-76.

- . 1968. Grass systematies. McGraw-Hill Co., New
York. 382 pp.

1969. The genus Schizachyrium in the Americas.
XI. Int. Bot. Cong. Proc. p. 75.

1974] Bahama Flora — Gillis 129

GRAHAM, SHIRLEY A. and C. E. Woop, JR. 1965. The genera of
Polygonaceae in the southeastern United States. J. Arnold Arbor.
46: 91-121.

GRENZENBACH, MYRTLE. 1926. A revision of the genus Bouchea
(exclusive of the Chascanwm). Ann. Missouri Bot. Gard. 13:
71-100 + 12 pl.

GUNN, CHARLES R. 1972. Moonflowers, Ipomoea section Calonyc-
tiom, in temperate North America. Brittonia 24: 150-68.

HAGSTRÖM, J. O. 1916. Critical researches on the Potamogetons.
Kungl. Sven. Vetens. Akad. Handl. Stockholms 55: 1-281.

HAMET, R. 1907. Monographie du genre Kalanchoé. Bull. Herb.
Boissier 7 (Sér. 2) : 869-900.

HARMS, VERNON L. 1965. A second character distinguishing Het-
erotheca s. str. from Chrysopsis (Compositae: Astereae). Rho-
dora 67: 86-8.

HARTOG, C. DEN. 1959. A key to the species of Halophila (Hydro-
charitaceae), with descriptions of the American species. Acta
Bot. Neerl. 8: 484-89.

1964. An approach to the taxonomy of the sea-

grass genus Halodule Endl. (Potamogetonaceae). Blumea 12:

289-312.

1970. The sea-grasses of the world. North-Holland
Publishing Co., Amsterdam,

HENRARD, J. TH. 1926-1933. A critical revision of the genus Aris-
tida. Meded. Rijks. Herb. Leiden 54: 1-220 (1926); 54A: 221-464
(1927); 54B: 465-701 (1928); 54C: 703-747 (1933).

1929-1932. A monograph of the genus Aristida.

Meded. Rijks. Herb. Leiden 58: 1-153 + Ix pl. (1929); 58A:

157-325 + pl. Ixi-clix (1932).

1950. Monograph of the genus Digitaria. Uni-
versitaire Pers Leiden. 999 pp.

HILL, STEVEN R. in press. Additions to the Bahama flora. Rhodora.

HiTCHCOCK, A. S. 1936. Manual of the grasses of the West Indies.
U.S.D.A. Misc. Publ. 243: 1-439.

HiTrCHCOCK, C. LEO. 1932. A monographic study of the genus Ly-
cium of the Western Hemisphere. Ann. Missouri Bot. Gard. 19:
179-374 + 24 pl.

. 1945. The Mexican, Central American, and
West Indian Lepidia. Mandrofio 8: 118-43.

HorLM, RicHARD W. 1950. The American species of Sarcostemma
R. Br. (Asclepiadaceae). Ann. Missouri Bot. Gard. 37: 447-560.

Horvat, SISTER MARY LiGOURI. 1941. A revision of the subgenus
Mariscus found in the United States. Contr. Biol. Lab. Catholic
Univ. Amer. 33: 1-147 + ix.

130 Rhodora [Vol. 76

Horcukiss, NEIL and HERBERT L. DoziER. 1949. Taxonomy and
distribution of North American cat-tails. Amer. Mid. Nat. 41:
237-54.

HOWARD, RICHARD A. 1957. Studies in the genus Coccoloba. IV.
The species from Puerto Rico and the Virgin Islands and from
the Bahama Islands. J. Arnold Arbor. 38: 211-42.

1962. Notes on Buxus in the Lesser Antilles

and on Mathou's overlooked publication. J. Arnold Arbor. 44:

96-100.

1973. The Enumeratio and Selectarum of

Nicolaus von Jacquin. J. Arnold Arbor. 54: 435-470.

and HENRY F. DUNBAR. 1964. Additions to

the flora of Inagua, the Bahamas. Rhodora 66: 6-15.

and D. A. POWELL. 1968. Clerodendrum phil-
ippinum Schauer replaces “Clerodendrum fragrans.” Taxon 17:
53-55.

HUTCHINSON, J. B. 1943. The cottons of Jamaica. Trop. Agr. 20:
56-8.

INGRAM, JoHN. 1967. A revisional study of Argythamnia, subgenus
Argythamnia (Euphorbiaceae). Gentes Herb. 10: 1-38.

IsELY, DUANE. 1970. Legumes of the United States. II. Desman-
thus and Neptunia. Iowa State J. Sci. 44: 495-511.

1972. Legumes of the United States. VI. Calliandra,
Pithecellobium, Prosopis. Madrofio 21: 273-98.

JACKSON, Curtis R. 1949. A key to the genus Scleria Berg in
South Florida. Quart. J. Florida Acad. Sci. 12: 220-22.

Jackson, R. C. 1960. A revision of the genus Iva L. Univ. Kansas
Sci. Bull. 41: 793-876.

JERVIS, Roy NEWELL. 1954. A summary of the genus Gochnatia
including a revision of the West Indian species which comprise
the Section Amastraphioides. Unpublished Ph.D. Dissertation.
University of Michigan. Ann Arbor, Michigan, 228 pp.

JoHNSON, Mites F. 1971. A monograph of the genus Ageratum L.
(Compositae — Eupatorieae). Ann. Missouri Bot. Gard. 58: 6-88.

JOHNSTON, IvAN M. 1949a. Studies in the Boraginaceae xvii. A.
Cordia section Varronia in Mexico and Central America. J.
Arnold Arbor. 30: 85-110.

1949b. The botany of San José Island (Gulf
of Panama). Sargentia 8: 1-306.

JOHNSTON, MARSHALL. 1962. The North American mesquites, Pro-
sopis section Algarobia (Leguminosae). Brittonia 14: 72-90.
1964. The fourteen species of Ziziphus in-
cluding Sarcomphalus (Rhamnaceae) indigenous to the West

Indies. Amer. J. Bot. 51: 1113-15.

1971. Revision of Colubrina (Rhamnaceae).

Brittonia 23: 2-53.

1974] Bahama Flora — Gillis 131

KEARNEY, THOMAS H. 1954a. A tentative key to the North Amer-
ican species of Pavonia Cav. Leafl. West. Bot. 7: 122-30.

1954b. A tentative key to the North Amer-

ican species of Sida L. Leafl. West. Bot. 7: 138-50.

1954c. Notes on Malvaceae V. Leafl. West.

Bot. 7: 118-21.

KILLIP, ELLSWORTH P. 1938. The American species of Passiflora-
ceae. Field Mus. Nat. Hist. Publ. Bot. Ser. 19 (1), Publ. 407.

KOBUSKI, CLARENCE EMMEREN. 1926. A revision of the genus Priva.
Ann. Missouri Bot. Gard. 13: 1-344- 5 pl.

Kopp, LUCILLE E. 1966. A taxonomic revision of the genus Persea
in the Western Hemisphere (Perseae — Lauraceae). Mem. N.Y.
Bot. Gard. 14: 1-120.

KRAL, RoBERT. 1971. A treatment of Abildgaardia, Bulbostylis, and
Fimbristylis (Cyperaceae) for North America. Sida 4: 57-227.

KUKENTHAL, G. 1949. Vorarbeiten zu einer Monographie der Rhyn-
chosporideae. Bot. Jahrb. 74: 375-509.

KunTH, C. S. 1824. Synopsis Plantarum aequinctialium orbis novi
(Voyage de Humboldt et Bonpland), Vol. 3. Paris.

LAKELA, OLGA. 1963. Centrosema floridanum (Britton) Lakela,
comb. nov. (Leguminosae). Sida 1: 182.

Lay, Ko Ko. 1950. The American species of Triumfetta L. Ann.
Missouri Bot. Gard. 37: 315-95.

LEENHOUTS, P. W. 1967. A conspectus of the genus Allophylus
(Sapindaceae). Blumea 15: 301-358.

LEFOR, MICHAEL WILLIAM. 1968. A revision of the genus Roche-
fortia Sw. (Boraginaceae). Unpublished M.S. thesis, University
of Connecticut. Storrs.

LEGRAND, C. D. 1952. Revisando tipos de Portulaca. Communic.
Bot. Mus. Hist. Nat. Montevideo 2 (24): 1-10 + 2 pl.

LEON, HERMANO. 1946. Flora de Cuba. Vol. 1 Gimnospermas y
Monocotiledoneas. Contr. Ocas. Mus. Hist. Nat. Colegio de la
Salle. No. 8 Habana.

LEON, HERMANO and HERMANO ALAIN. 1951-57. Flora de Cuba.
Contribucion No. 10 — Dicotiledoneas: Casuarinaceas a Meliaceas.
Vol. 2. Contr. Ocas. Mus. Hist. Nat. Colegio de la Salle. Habana.
Contribucion No. 13 — Dicotiledoneas: Malpighiaceae a Myrta-
ceae. Contribucion No. 16 — Dicotiledoneas: Malastomaeaceae a
Plantaginaceae. Vol. 4.

LEWis, WALTER H. 1961. Merger of the North American Houstonia
and Oldenlandia under Hedyotis. Rhodora 63: 216-23.

LINNAEUS, C. 1769. Plantarum Jamaicensium Pugillus. Disserta-
tion 102.

1775. Plantae Surinamensis. Jac. Alm. Dissertation

177.

132 Rhodora [Vol. 76

LITTLE, E. L., JR. 1947. The name of the wild dilly of Florida.
Rhodora 49: 289-93.

1968. Transfers to Guapira from Torrubia
(Nyctaginaceae). Phytologia 17: 367-71.

Lowc, RoBERT W. 1970a. Additions and nomenclatural changes in
the flora of southern Florida — I. Rhodora 72: 17-46.

1970b. The genera of Acanthaceae in the south-
eastern United States. J. Arnold Arbor. 51: 257-309.

LÜCKHOFF, H. A. 1964. The natural distribution, growth, and bo-
tanical variation of Pinus caribaea and its cultivation in South
Africa. Ann. Univ. Stellenbosch, Ser. A: 39: 1-60.

LUER, CARLYLE A. 1972. The native orchids of Florida. New York
Botanical Garden. New York. 293 pp.

MCCANN, CHARLES. 1945. Notes on the genus Ruppiaceae. J. Bom-
bay Nat. Hist. Soc. 45: 396-402.

McGivney, SISTER M. VINCENT DE PAUL. 1938. Revision of the
subgenus Eucyperus found in the United States. Contr. Biol.
Lab. Catholic Univ. Amer. 26: 1-74 + xvii.

MCLAUGHLIN, ANNE DesmMonp. 1944. The genus Cyperus in the
West Indies. Catholic Univ. Amer. Biol. Studies No. 5 pp 1-108
+ viii.

McVaucu, Rocers. 1943. Campanulales — Campanulaceae (Lobeli-
oideae). North American Flora 32A, Pt. 1: 1-124.

1963. Tropical American Myrtaceae. II. Notes

on generic concepts and descriptions of previously unrecognized

species. Fieldiana 29: 393-532.

1973. Notes on West Indian Myrtaceae. J.
Arnold Arbor. 54: 309-14.

MACBRIDE, J. FRANCIS. 1916. Further notes on the Boraginaceae.
Contr. Gray Herb. 49: 16-22.

1930. Other Peruvian plants, chiefly new
species. Publ. Field Mus. Nat. Hist, Bot. Ser. (Publ. 278) 8:
113-29.

MERRILL, E. D. 1910. An enumeration of Philippine Leguminosae
with keys to genera and species. (Concluded). Philip. J. Sci.
(Botany) 5: 95-136.

1936. On the application of the binomial Amaran-

thus viridis Linnaeus. Amer. J. Bot. 23: 609-12.

and Surv-YiNa Hu. 1949. Work and publications of

Henry Muhlenberg, with special attention to unrecorded or in-

correctly recorded binomials. Bartonia 25: 1-66.

and F. P. METCALF. 1942. Hedyotis Linnaeus versus

Oldenlandia Linnaeus and the status of Hedyotis lancea Thun-

berg in relation to H. consangwinea Hance. J. Arnold Arbor. 23:

226-30.

1974] Bahama Flora — Gillis 133

MOHLENBROCK, ROBERT H. 1957. A revision of the genus Stylosan-
thes. Ann. Missouri Bot. Gard. 44: 299-355.

MoLDENKE, HaRoLD N. 1936. A monograph of the genus Callicarpa
as it occurs in America and in cultivation. ii. Fedde, Repert. Sp.
Nov. 40: 38-131.

1937. A monograph of the genus Petitia.

Fedde, Repert. Sp. Nov. 42: 229-51.

1958a. Materials toward a monograph of the

genus Citharexylum II. Phytologia 6: 262-320.

1958b. Materials toward a monograph of the

genus Citharexylum III. Phytologia 6: 322-368.

1958c. Materials toward a monograph of the

genus Citharexylum IV. Phytologia 6: 383-432.

1965. Materials toward a monograph of the

genus Lippia. I. Phytologia 12: 6-71.

1971. A fifth summary of the Verbenaceae,
Avicenniaceae, Stilbaceae, Dicrastylidaceae, Symphoremaceae,
Nyctanthaceae, and Eriocaulaceae of the world as to valid taxa,
geographic distribution, and synonymy. Privately published,
Wayne, NJ.

Moore, HanoLD E., Jr. 1963. An annotated checklist of cultivated
palms. Principes 7: 119-84.

1966. Nomenclatural notes on the cultivated

conifers. Baileya 14: 1-11.

and WILLIAM T. STEARN. 1967. The identity
of Achras zapota L. and the names for the sapodilla and the
sapote. Taxon 16: 382-95.

MoonE, R. J. and C. FRANKTON. 1969. Cytotaxonomy of some Cir-
sium. species of the eastern United States, with a key to eastern
species. Canad. J. Bot. 47: 1257-75.

NICKERSON, Norton H. and JoskPH W. Tripp. 1973. Floral di-
morphism in Rachicallis americana (Jacq.) Hitchc. (Saltwater-
bush). Rhodora 75: 111-13.

NIKLES, DoNALD GARTH. 1966. Comparative variability and rela-
tionship of Caribbean Pine (Pinus caribaea Mor.) and slash pine
(Pinus elliottii Engelm.). Unpublished Ph.D. dissertation. North
Carolina State University. Raleigh, N.C.

O'DoNELL, C. A. 1941. Revision de las especies americanas de
Merremia (Convolvulaceae). Lilloa 6: 467-554.

OGDEN, E. C. 1943. The broad-leaved species of Potamogeton of
North America north of Mexico. Rhodora 45: 57-105; 119-63;
171-214 + pl. 146-48.

O'NEILL, HucH T. 1939. The status and distribution of Cyperus
distinctus Steud. Rhodora 41: 131-33.

134 Rhodora [Vol. 76

1940. The sedges of the Yucatan Peninsula. xix.
in Botany of the Maya area. Carnegie Inst. Washington Publ.
522: 248-322.

OosTSTROOM, S. J. VAN. 1934. A monograph of the genus Evol-
vulus. Meded. Bot. Mus. Herb. Rijks. Univ. Utrecht No. 14:
1-267.

1940. The Convolvulaceae of Malaysia III.

The genus Ipomoea. Blumea 3: 481-582.

in collaboration with R. D. HooGLanp. 1953.
Convolvulaceae in: Flora Malesiana I: 4: 388-512.

ORNDUFF, RoBERT. 1969. Neotropical Nymphoides (Menyanthaceae):
Meso-American and West Indian species. Brittonia 21: 346-52.

Pacut, JiRí. 1952. Synopsis of the genus Catalpa (Bignoniaceae).
III. Candollea 13: 241-85.

PARKS, JAMES C. 1973. A revision of North American and Carib-
bean Melanthera (Compositae). Rhodora 75: 169-210.

PAYNE, WILLARD W. 1964. A re-evaluation of the genus Ambrosia
(Compositae). J. Arnold Arbor. 45: 401-38.

PENNELL, FRANCIS W. 1917. Notes on plants of the southern United
States — III. Bull. Torrey Bot. Club 44: 337-62.

Perry, JAMES D. 1971. Biosystematic studies in the North Amer-
ican genus Sabatia (Gentianaceae). Rhodora 73: 309-69.

PFEIFER, Howarp W. 1966. Revision of the North and Central
American hexandrous species of Aristolochia (Aristolochiaceae).
Ann. Missouri Bot. Gard. 53: 115-96.

1970. Taxonomic revision of pentandrous
species of Aristolochia. University of Connecticut Publication
Series. Storrs, Connecticut.

PuiLcox, D. 1965. Revision of the New World species of Buchnera
L. (Serophulariaceae). Kew Bull. 18: 275-315.

PoRTER, DUNCAN M. 1972. The genera of Zygophyllaceae in the
southeastern United States. J. Arnold Arbor. 53: 531-52.

POWELL, A. MICHAEL. 1965. Taxonomy of Tridax (Compositae).
Brittonia 17: 47-96.

PRANCE, GHILLEAN T. 1972. Chrysobalanaceae. Flora Neotropica
Monograph No. 9. Hafner Publishing Co. New York.

PRESL, C. B. 1835. Reliquiae Haenkeanae, Vol. 2. Prague.

PURSEGLOVE, J. W. 1968. Tropical crops. Dicotyledons. 2 vol.
Longmans, London. 719 pp.

RAO, ARAGULA SATHYANARAYANA. 1956. A revision of Rauvolfia,
with particular reference to the American species. Ann, Missouri
Bot. Gard. 43: 253-354.

RAVEN, PETER H. 1962. New combinations in Ludwigia. Kew Bull.
15: 476.

1974] Bahama Flora — Gillis 135

1968. The Old World species of Ludwigia (In-
cluding Jussiaea) with a synopsis of the genus (Onagraceae).
Reinwardtia 6: 327-427.

Reap, RoBERT W. 1968. A study of Pseudophoenix (Palmae).
Gentes Herb. 10: 169-213.

REED, CLYDE F. 1968. The nomenclature and synonymy of the
Amaranthaceae, Chenopodiaceae, and Nyctaginaceae of the flora
of Texas. Phytologia 18: 1-44.

REESE, G. 1962. Zur intragenerischen Taxonomie der Gattung Rup-
pia L. Zeit. für Bot. 50: 237-64.

ROBERTSON, KENNETH R. 1971. The Malpighiaceae in the south-
eastern United States. J. Arnold Arbor. 52: 101-12.

Rosson, N. 1965. New and little known species from the Flora
Zambesiaca area. XVI. Taxonomic and nomenclatural notes on
Celastraceae. Bol. Soc. Broter. 39: 5-55.

RoE, KEITH E. 1968. Solanum verbascifolium L., misidentification
and misapplication. Taxon 17: 176-78.

Rogers, C. MARVIN. 1963. Yellow flowered species of Linum in
eastern North America. Brittonia 15: 97-122.

1968. The Linwm bahamense complex. Rhodora

70: 439-41.

Rogers, Davip J. 1963. Studies on Manihot esculenta Crantz and
related species. Bull. Torrey Bot. Club 90: 43-54.

RoMINGER, JAMES M. 1962. Taxonomy of Setaria in North Amer-
ica. Illinois Biol. Monogr, 29: 1-132.

ROTHMALER, WERNER. 1944. Nomina generica neglecta 1753-1763.
Fedde, Repert. Sp. Nov. 53: 1-37.

Rupp, VELVA E. 1967. Oxyrhynchus and Monoplegma (Legumino-
sae). Phytologia 15: 289-94.

1968. Leguminosae of Mexico — Faboideae. T

Sophoreae and Podalyrieae. Rhodora 70: 492-532.

1969. A synopsis of the genus Piscidia (Legumino-
sae). Phytologia 18: 473-99.

St. JoHN, HanoLp. 1970. Classification and distribution of the
Ipomoea pes-caprae group (Convolvulaceae). Bot. Jahrb. 89:
563-83.

SAUER, JONATHAN. 1964. Revision of Canavalia. Brittonia 16:

106-81.

1972. Revision of Stenotaphrum (Gramineae:
Paniceae) with attention to its historical geography. Brittonia
24: 202-22.

SAVAGE, SPENCER. 1945. A catalogue of the Linnaean herbarium.
Linnaean Society of London. London. 225 pp.

ScHERY, ROBERT WALTER. 1942. Monograph of Malvaviscus. Ann.
Missouri Bot. Gard. 29: 183-244 + 17 pl.

136 Rhodora [Vol. 76

ScHINDLER, A. K. 1926. Die Desmodiinen in der botanischen Liter-
atur bis Linné. Fedde, Repert. Sp. Nov. 23: 71-127.

ScuuLz, Orro E. 1909. Citharexylum Linn. Sym. Antill. 6: 56-69.

ScuwaNiTZ G. 1967. Untersuchungen zur Postmeiotischen Mikro-
sporogenese. 1. Morphogenese des Ruppia-pollens. Pollen et
Spores 9: 9-48.

SEALY, J. ROBERT. 1954. Review of the genus Hymenocallis. Kew
Bull. 1954: 201-40.

SETCHELL, W. A. 1946. The genus Ruppia. Proc. Calif. Acad. Sci.
4a. S 25: 469-72.

SHERFF, EARL EDWARD. 1937. The genus Bidens. Field Mus. Nat.
Hist. Bot. ser. 16: 1-709.

1945. Some additions to the genus Dodon-

aea L. (Sapindaceae). Amer. J. Bot. 32: 202-14.

1947. Further studies in the genus Dodon-
aea L. (family Sapindaceae). Publ. Field Mus. Bot. 23: 269-317.

SHINNERS, L. H. 1951. The North Texas species of Heterotheca,
including Chrysopsis. Field and Lab. 19: 66-71.

1957. Synopsis of the genus Eustoma (Gentian-

aceae) Southwestern Nat. 2: 38-43.

1962. Micromeria brownei and its allies (Labi-
atae). Sida 1: 94-97.

SLEE, M. U. 1970. Crossability values within the slash-Caribbean
Pinus species complex. Euphytica 19: 184-89.

SPRAGUE, T. A. 1932. Laugeria “Vahl” = Terebraria Kuntze. Kew
Bull. 1926: 349.

STAFLEU, FRANS A. 1967. Taxonomic Literature. Regnum Vegeta-
bile 52. 556 pp. Utrecht.

STAFLEU, FRANS et al. 1972. International Code of Botanical No-
menclature. Reg. Veg. 82: 1-426.

STEARN, WILLIAM T. 1957. The boat-lily (Rhoeo spathacea). Bai-
leya 5: 195-98.

1958. A key to West Indian mangroves. Kew

Bull. 1958: 33-37.

1964. Catharanthus roseus, the correct name
of the Madagascar periwinkle. Lloydia 27: 196-200.

1966. Botanical Latin. Thomas Nelson and
Sons, Ltd. London. 566 pp.

1968. Jamaican and other species of Bumelia
(Sapotaceae). J. Arnold Arbor. 49: 280-89.

1969. A synopsis of Jamaican Myrsinaceae.
Bull. Brit. Mus. Nat. Hist. Bot. 4: 143-78 -- 8 pl.

1971. A survey of the tropical genera Oplonia
and Psilanthele (Acanthaceae). Bull. Brit. Mus. Nat. Hist. Bot.
4: 259-323.

1974] Bahama Flora — Gillis 137

1972. Typification of Evolvulus nummularius,
E. convolvuloides, and E. alsinoides (Convolvulaceae). Taxon 21:
647-50.

STUCKEY, RONALD L. 1972. Taxonomy and distribution of the genus
Rorippa (Cruciferae) in North America. Sida 4: 279-430.

TAYLOR, PETER. 1967. Lentibulariaceae. Mem. N.Y. Bot. Gard. 17:
201-28.

THARP, B. C. and MARSHALL C. JOHNSTON. 1961. Recharacteriza-
tion of Dichondra (Convolvulaceae) and a revision of the North
American species. Brittonia 13: 346-60.

TRELEASE, W. 1918. Agave in the West Indies. Mem. Natl. Acad.
Sci. 11: 1-55.

1916. The genus Phoradendron: a monographie re-
vision. Univ. Illinois Press. Urbana.

VERDCOURT, B. 1963. Convolvulaceae. in: Flora of Tropical East
Africa, C. E. Hubbard and E. Milne-Redhead. eds.; Crown Agents
for Overseas Governments and Administrations.

WAGENKNECHT, BURDETT L. 1960. Revision of Heterotheca, section
Heterotheca (Compositae). Rhodora 62: 61-76; 99-107.

WATERFALL, U. T. 1967. Physalis in Mexico, Central America, and
the West Indies. Rhodora 69: 82-102; 203-40; 319-29.

WEBSTER, GRADY L. 1956-58. A monographie study of the West
Indian species of Phyllanthus. J. Arnold Arbor. 37: 91-122; 217-
68; 340-59; pls. 1-14; 38: 51-80; 170-98; 295-373; pls. 15-23; 39:
49-100; 111-212; pls. 24-32.

1967. The genera of Euphorbiaceae in the

southeastern United States. J. Arnold Arbor. 48: 303-61; 363-430.

1970. A revision of Phyllanthus (Euphorbi-
aceae) in the continental United States. Brittonia 22: 44-76.

WHEELER, Louis C. 1938. The names of three species of Brassica.
Rhodora 40: 306-309.

WIJNANDS, D. O. 1973. Typification and nomenclature of two spe-
cies of Sansevieria (Agavaceae). Taxon 22: 109-114.

WILBUR, ROBERT L. 1955. A revision of the North American genus
Sabatia (Gentianaceae). Rhodora 57: 1-33; 43-71; 78-104.
WILSON, KENNETH A. 1960. The genera of Convolvulaceae in the
southeastern United States. J. Arnold Arbor. 41: 298-317.
WINDLER, D. R. 1966. A revision of the genus Neptunia (Legum-

inosae). Aust. J. Bot. 14: 379-420.

Woop, CARROLL E., JR. 1958. The genera of the woody Ranales in
the southeastern United States. J. Arnold Arbor. 39: 296-346.

1959. The genera of the Nymphaeaceae and

Ceratophyllaceae in the southeastern United States. J. Arnold

Arbor. 40: 94-112.

138 Rhodora [Vol. 76

and R. B. CHANNELL. 1960. The genera of
the Ebenales in the southeastern United States. J. Arnold Arbor.
41: 1-35.

Woopson, ROBERT E., JR. 1936. Studies in the Apocynaceae, IV.
the American genera of Echitoideae. Ann. Missouri Bot. Gard.
23: 169-438 + 7 pl.

1937. New or otherwise noteworthy Apo-
cynaceae of tropical America, V. Ann. Missouri Bot. Gard. 24:
11-16.

1938a. Studies in the Apocynaceae. VII.
An evaluation of the genera Plumeria L. and Himatanthus
Willd. Ann. Missouri Bot. Gard. 25: 189-224.

1938b. (Asclepiadales) Apocynaceae.
North Amer. Flora 29: 103-192.

YAKOVLEV, G. P. 1967a. Systematical and geographical studies of
the genus Sophora L. and allied genera. Prob. Pharmacog. 21:
42-62. (In Russian).

1967b. Some characteristics of structure of seeds
of Sophora L. and allied genera in connection with its systematics
and phylogeny. Repr. Proc. Leningrad Chem.-Pharm. Inst. 21:
90-98. (In Russian).

YATES, HARRIS O. 1966. Revision of grasses traditionally referred
to Uniola I. Uniola and Leptochloópsis. Southwestern Nat. 11:
372-394.

YUNCKER, TRUMAN GEORGE. 1932. The genus Cuscuta. Mem. Tor-
rey Bot. Club 18: 113-331.

ARNOLD ARBORETUM, HARVARD UNIVERSITY
CAMBRIDGE, MASS. 02138
AND
DEPARTMENT OF BIOLOGY
UNIVERSITY OF MIAMI
CORAL GABLES, FL. 33124

STONE'S PLANTS OF SOUTHERN NEW JERSEY:
A REPRINT'

To those who had always considered Witmer Stone as
an ornithologist — and one of the country's greatest — the
appearance in 1911 of his “Plants of Southern New Jersey"
came as a distinct surprise. Had they known more about
the man they would have realized that he was one of the
last of the all-around naturalists, interested in mammals,
insects and plants as well as birds.

The recent facsimile reprint of Stone's book, by the
Quarterman Publications, Inc., of Boston, makes available
once more a work which for more than sixty years has been
the standard reference for everyone who is interested in
the flora of the Pine Barrens, as well as the other plant
provinces of southern New Jersey.

Originally published as Part II of the Annual Report
of the New Jersey State Museum, this important volume
has for many years been out of print and has, indeed, be-
come something of a collector's item.

Although one can not be other than grateful that this
flora is once again in circulation, it is to be regretted that
the Foreword was not written by a professional botanist,
preferably one who was intimately acquainted with Witmer
Stone.

Since this reprint is a facsimile, every word, every punc-
tuation mark and every misprint appears exactly as it did
in the original. There are numerous discrepancies between
its nomenclature and that in use today. It would have been
helpful to the reader if Elizabeth M. Woodford, who wrote
the Foreword, had explained the reasons for this.

Partly because he was a zoologist and partly because of
his friendship with Dr. N. L. Britton of the New York
Botanical Garden, Stone's treatment of plant names follows
the now-abandoned American Code of Botanical] Nomen-

'Stone, Witmer. The Plants of Southern New Jersey. Reprint.
1973. Quarterman Publications, Inc., Boston, Massachusetts. 828 pp.
Illust. $25.00.

139

140 Rhodora [Vol. 76

clature. This code, like that of the zoologists, permits the
use of tautonyms or duplicate binomials. Hence we find
in Stone such combinations as Blephariglottis blephari-
glottis, Hepatica hepatica, Linaria linaria and others.

The American Code never embraced the principle of the
conservation of generic names and Stone therefore con-
sistently employed what he considered to be the earliest
generic epithets. As a consequence we have scores of
generic names which are unfamiliar to most present-day
taxonomists. A few examples will suffice: Spathyema for
Symplocarpus, Juncoides for Luzula, Abama for Narthe-
cium, Vagnera for Smilacina, Gyrostachys for Spiranthes,
Hicoria for Carya, Ammodenia for Arenaria (peploides),
Meibomia for Desmodium, etc.

Fortunately most of the presently accepted names are
cited in synonymy (although not always with the correct
authority), but it seems a pity that a republication of this
work should not have suggested some revision and up-
dating of its nomenclature. It might also have corrected
such misprints as “Sandy Hood” for “Sandy Hook" (page
432) and “dandelon” for “dandelion” (page 817).

Mrs. Woodford states that Stone’s collection of 12,000
specimens went to the herbarium of the Academy of Natu-
ral Sciences of Philadelphia. Such is not the case. Realizing
that many of his duplicates were already in the Academy
and that he had frequently been in the field with members
of the Philadelphia Botanical Club whose specimens had
likewise been deposited there, I suggested to Dr. Stone that
he leave his herbarium to the University of Pennsylvania.
He readily agreed, and in the fall of 1942 the University
received his collection of more than 14,000 sheets, rich not
only in plants from southern New Jersey, but also in ma-
terial from northern Pennsylvania. It was my pleasant
task to supervise the incorporation of this collection into
the herbarium of the University.

Witmer Stone once told me that his manuscript for the
New Jersey Flora had been prepared in a little more than
four months. Doubtless he had made some preliminary

1974] Book Review — Fogg 141

efforts to put together his observations accumulated over
many years of field work, but even so this was a prodigious
accomplishment. He added that the Director of the New
Jersey Museum had approached him in desperation to give
them something for their Annual Report and urged him
to prepare a paper on the Pine Barrens of New Jersey.
Stone's reply was that this would be like treating one
vegetational province in a vacuum and that he preferred
to deal with the entire flora of the lower three-fifths of
the state, at the same time emphasizing the geographic
origins of the various elements represented therein. This
he did superbly well, in a manner which certainly would
have elicited the approbation of Asa Gray.

Mrs. Woodford gives a picture of Witmer Stone as a
warm, friendly human being with a wide breadth of in-
terests and a well developed sense of humor. With this i
can heartily agree. It was my good fortune for many
years to spend my summers at Cape May, where Dr. Stone
also had a cottage, and in many walks with him I came
to appreciate his encyclopedic knowledge and great capacity
for friendship.

JOHN M. FOGG, JR.
ARBORETUM OF THE BARNES FOUNDATION
MERION, PA. 19066

THE ASIATIC SPECIES OF DESMODIUM:
A REVIEW

A new series of monographie studies, Ginkgoana,' was
inaugurated early in 1973 with the publication of Number 1,
“The Asiatic species of Desmodium and its allied genera
(Leguminosae)," by Hiroyoshi Ohashi of the Department
of Botany, University of Tokyo.

The first study in the series is truly a book, printed in a
clear and pleasing type face, and very well edited. The
text is written in English. This study is a portion of the
thesis presented by Mr. Ohashi for the degree of Doctor of
Science. It is an inclusive treatment with clearly stated
conclusions.

Since the genus Desmodium (in the broad sense) is one
in which I have long had a particular interest and since I
know the Asiatic species less well than the American and
African, I am pleased to see this detailed piece of work.
Of special importance are the pollen studies and the meticu-
lous investigations of floral parts, seeds, and seedlings.
Dr. Ohashi confirms the conclusions of Ambrose? for Des-
modium (i.e. the species of Desmodium ser. Americana have
hypogaeous germination, while those of other species ob-
served have an epigaeous type of germination), and pre-
sents new information for additional species.

The chapter considering observations on morphological
characters demonstrates well the care and scope of Dr.
Ohashi's observations on habit, leaves, inflorescences, bracts,
bracteoles, androecia or stamens, pollen grains, fruits
[*^pods"], seeds, seedlings, and chromosome numbers. He
has illustrated with clarity the basic structures and prob-
able evolutionary progression in the development of the

'Ginkgoana, Contributions to the Flora of Asia and the Pacific
Region. No. 1, The Asiatic Species of Desmodium and its Allied
Genera (Leguminosae), by Hiroyoshi Ohashi, 318pp., 86 figures, 76
plates [18.1 X 25.6 em.]. February 15, 1973. Academia Scientific
Book Ine., Tokyo, Japan.

"The Michigan Botanist 6: 97-99. 1967.

142

1974] Book Review — Schubert 143

inflorescence for the group (Desmodium sens. lat.) and also
of the bracts and bracteoles. In both cases the tendencies
indicated could be compared profitably with parallel or
similar progressions in the New World and African groups
of related genera. The resulting comparisons, particularly
with New World genera and species, should be of much
interest because of the numerous groups of closely related
species which radiate from the centers of diversity in Mex-
ico and Brazil.

The taxonomie treatment has well constructed keys, de-
tailed descriptions, ample citations of specimens, exceed-
ingly good illustrations of floral parts and of fruits, and
distribution maps. In general, I think Dr. Ohashi has rather
over-classified the group he has studied so carefully, follow-
ing Schindler and Hutchinson in large part. However, he
has documented his data and stated his reasons so carefully
that his occasionally debatable conclusions do not detract
from the usefulness of the work. The treatment of Des-
modium subgenus Sagotia is perhaps the one which troubles
me most. Dr. Ohashi says “The subgenus Sagotia is one of
the most polymorphic and widespread groups of the genus
Desmodium. It is characterized by having mostly the ter-
minal racemes composed of 2(-3)-flowered fascicles which
are subtended usually only by the primary bracts (i.e. lack-
ing the secondary bracts), ebracteolate flowers, diadelphous
androecia and rather small-sized articles of the fruits.” He
includes, however, species with secondary bracts, among
them Desmodium velutinum, D. gangeticum, and D. dichoto-
mum; some with bracteoles, such as D. caudatum and D.
benthamii; some with flowers borne singly; and some with
loment articles of considerable size.

It seems to me that the only reason the genus Sagotia
(sensu Ohashi) is so polymorphic is that Dr. Ohashi has
assigned to it very unlike elements which do not have a
close natural relationship. His arrangement may seem
feasible when studying only Asian species, but on a world
wide basis I think it would become wholly unworkable and
its defects would be much magnified. In some infrageneric

144 Rhodora [Vol. 76

groups in Asia as for most of the genus Desmodiwm in
Africa there are many unrelated, distinctive species in con-
trast to the situation in North and South America where
there are many small to large groups of closely related
species. This makes for a very different aspect of a pre-
sumed phylogeny, of proposed evolutionary series, and for
an understanding of interspecific relationships in Asia and
Africa on the one hand and in America on the other, even
though there are some species which bridge the gap by
natural or artificial means.

The only serious omissions in this careful study are in
the areas of ecology and phytogeography, discussion of the
former being almost totally lacking and of the latter only
minimal. This may indicate a lack of field experience which
can be remedied later by an active field program. Field
work might also give Dr. Ohashi a broader view of the taxa
with which he is working and a somewhat different ap-
proach to their interrelationships.

In concluding this review I would stress the meticulous
care of the observations Dr. Ohashi has made and the great
gap he has filled in our knowledge of one large complex in
the family Leguminosae.

BERNICE G. SCHUBERT
ARNOLD ARBORETUM
HARVARD UNIVERSITY

22 DIVINITY AVENUE
CAMBRIDGE, MASS. 02138

A MAGNIFICENT SWISS FLORA MANUAL'
A REVIEW

One of the most important tasks of botanists specializing
in taxonomy and geobotany is the compilation of flora
manuals for their colleagues in general and for the much
more numerous amateur botanists in particular. In this
way, the author himself gets a supplementary survey over
the plants of the region in which he works and gathers
experience that enables him to solve problems of unusual
importance. At the same time he gets a rare opportunity
to stimulate interest in his subject and to gather new prose-
lytes among the younger generation to carry on in his foot-
steps. Only a few botanists have the encyclopedic knowl-
edge and ability to write floras, and still fewer want to take
the risk of being unfairly criticized for this least gratifying
of scientific occupations, because in no other field is it easier
to be criticized for opinions and judgements. This is the
perennial trouble for compilers of floras and also the main
cause of the scarcity of good manuals. If the author also
ventures to make deviations from the accepted delimita-
tions, descriptions and names of the plants included, he
runs the additional risk of condemnation by his more con-
ventional colleagues, because scientists are no less con-
servative than are other citizens. Ordinary manuals are
always valuable, but unconventional floras ought to be
doubly welcome, because nothing else can stimulate future
research to a higher degree.

Although Switzerland is one of the smaller European
countries, it has fostered astonishingly many of the botani-
cal giants. Several of its botanists certainly have been con-
ventional and followed the trend of their times, whereas
others have dared to have new ideas and have looked at the

'Hans Ernst Hess, Elias Landolt, Rosemarie Hirzel: Flora der
Schweiz und angrenzender Gebiete. Three volumes, 29 X 23 cm.,
2690 pages, with 9 colored plates and about 3450 drawings. Birk-
háuser Verlag, Basel & Stuttgart, 1967, 1970, 1972. Price 450
Schweizer francs.

145

146 Rhodora [Vol. 76

plants of their country from new points of view. These
latter we have to thank for the unusually large number of
presently well-known species that were first described from
Switzerland. Although many of the great Swiss botanists
of the past are now always mentioned with reverence, there
were times when their contemporaries called them names
and were critical about their scientific abilities. Such is the
fate of those who are born too early or try to be prophets
in their own land, but time is their confederate.

It would be difficult to mention all the many flora man-
uals, which have been published in Switzerland in the past,
and the list of those which have reached fame far outside
the country because of scientific quality is far from being
short. One such flora was Albrecht von Haller’s great
*Historia stirpium indigenerorum Helvetiae inchoata",
which was published in 1768 in three folio volumes. That
work has stood in a class by itself for two centuries as the
most monumental botanical manual ever written in the field
of descriptive floristies. It still remains important for its
clear descriptions and other information, but the conserva-
tism in nomenclature and the reluctance to accept what
would be the future in that field has detracted from the
nomenclatural importance of the work though it always will
be a text of literary superiority rarely achieved in scientific
books.

Although many good Swiss floras have been published
since Haller's work, two centuries had to pass before a text
of similar magnitude and quality again was compiled for
this flora of more than 3500 species. This is the new “Flora
der Schweiz," by H. E. Hess, E. Landolt and R. Hirzel, the
three large volumes of which were published in 1967, 1970
and 1972. It is a work of extraordinary quality and quan-
tity, based on fifteen years of intensive work by the two
first authors and on equally intensive work of drawing the
excellent pictures of more than 3450 of the species by the
third author.

The virtues of this manual are too many to be even sim-
ply listed, and readers with different inclinations will al-

1974] Book Review — Lóve 147

ways find some matters of special interest. Considerable
information on the methods and philosophy adopted in the
compilation of this great work is given in an introductory
chapter, which then is followed by almost thirty pages on
the history and origin of the flora and its distribution, and
more than forty pages on its ecology. Specialists are apt
to find omissions in these chapters, and some may perhaps
join issue with the authors as to some of the explanations
given. That is to be expected in any work of this magnitude
and concentration. But this reviewer, who believes he has
gathered considerable knowledge about this flora through
visits and reading numerous good books and reports during
the past three decades, finds these chapters to be highly
educational and of great interest from more points of view
than he cares to mention. Numerous maps in these intro-
ductory chapters ought to make it easy for those unfamiliar
with the geography of the land to follow the detailed ac-
count.

The manual itself is arranged according to the Engler
system as are most modern European floras. A general key
to the higher groups leads to keys to the families of the
major groups where these commence, and keys to genera
and species are then given in appropriate places in the
text. Families and genera are described in detail, followed
under the latter by information about the size of the genus
and its distribution, and about cytological information avail-
able. After the generic key there are detailed reports of
possible hybridization observed, frequently with explana-
tions of the morphological, geographical, and cytological
characteristics of the plants.

Most species are represented by a good drawing, fre-
quently with additional details. Their Latin names are
followed by synonyms, if any, and German names, and a
very detailed description is given of the morphological
characteristics. For every species there is also information
on chromosome number and other cytological peculiarities,
and the authors have done an admirable work in looking up
the original papers in order to be able to exactly inform the

148 Rhodora [Vol. 76

reader about the geographical origin of the material. This
is the only place where such information has been compiled,
and it is admirably corect, although some few mistakes have
not been avoided. Then there are reports on the ecological
occurrence of each species in Switzerland, and on the gen-
eral distribution of the taxon elsewhere in the world. Fre-
quently, the authors then add taxonomical and geographical
remarks on the variation of the species within and outside
their country, especially when they feel that it is to be
regarded as a critical taxon in need of more close investiga-
tion, with references to the most recent literature in every
special case.

It is easy to find points of disagreement in a work of this
magnitude, although this reviewer is of the opinion that the
authors have sueceeded unusually well in avoiding harsh
judgements. They have made an effort to accept a modern
species concept although sometimes this may not have suc-
ceeded well enough, and the descriptions of the taxa are
clear and moderately concise. It is a pity that they have
found it impossible to mention subspecific categories, but
since Switzerland abounds in such variations which fre-
quently may be of doubtful value, it is understandable that
the authors avoided spending perhaps twice as much time in
evaluating these. The only serious remark to be made seems
to be that the authors adopted, for present-day European
botanists, an unusually conservative attitude towards the
nomenclature, so that sometimes their eccentricity may look
disturbing. Nevertheless, this is understandable, and the
explanation of this given in the last volume is not unrea-
sonable although I believe that a more liberal attitude would
have made the work better and also more useful for a longer
period of time.

After the description of the last species in the third vol-
ume, there are more than 90 pages of corrections and addi-
tions, half a page with an epilogue, where some criticism of
plant sociology and nomenclature by reviewers of the first
two volumes are met, and four pages on which the termi-
nology used is explained. There are good indexes of Latin

1974] Book Review — Lóve 149

and German names, and these are preceded by sixty pages
of a very important bibliography.

The three volumes are printed in the unusual size 29 X
23 em, with considerable empty space on every page that is
partially used for the large drawings. The heavy volumes
are strongly and tastefully bound in green linen, and they
are certainly reasonably priced for their size and quality.
Their weight would make it difficult to carry them into
the field, but the authors are compiling a smaller manual
with the keys only and new drawings of selected species
that will be a handy companion on excursions in the Alps
and elsewhere.

The “Flora der Schweiz” is an unusually thorough and
scholarly work which will long be the standard reference
for this part of Europe. It is also an invaluable source of
information for botanists from other parts of the continent,
and also for specialists in other fields, especially that of
cytotaxonomy. The text and pictures are so detailed and
exact that the book can be highly recommended to those
foreign botanists and institutions which need good and
reliable information on European plants which may have
been introduced overseas, because no other flora comple-
ments the comprehensive Flora Europaea more excellently
for southern and central European plant species.

ÁSKELL LÓVE
473 HARVARD LANE
BOULDER, COLORADO 80303

INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA

Manuscripts must be double-spaced or preferably triple-
spaced (not on corrasable bond), and a list of legends for
figures and maps provided on a separate page. Footnotes
should be used sparingly, as they are usually not necessary.
Do not indicate the style of type through the use of capitals
or underscoring, partieularly in the citations of specimens,
except that the names of species and genera may be under-
lined to indicate italies in discussions. Specimen citations
should be selected critically especially for common species
of broad distribution. Systematic revisions and similar
papers should be prepared in the format of “The System-
aties and Ecology of Poison-Ivy and the Poison-Oaks,"
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu-
larly with reference to the indentation of keys and syno-
nyms. Papers of a floristic nature should follow, as far as
possible, the format of ‘Contribution to the Fungus Flora
of Northeastern North America. V.," H. E. Bigelow & M. E.
Barr, Rhodora 71: 177-203. 1969. For bibliographic cita-
tions, a recommended list of standard journal abbreviations
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot.
Club 85: 277-300. 1958.

Volume 76, No. 805, including pages 1-150, was issued March 30, 1974.

150

CONTENTS: — continued

Studies on the Bryophytes of Southern Manitoba III.
Collections from Grand Beach Provincial Park

Paul W. Stringer and Muriel H. L. Stringer .................... 27
Some New or Noteworthy Vascular Plant Records from

Northwestern Saskatchewan

Vernon L. Harms and John H. Hudson ...............5.4....« 39
Parasitic Witchweed: Striga asiatica Versus S. lutea

(Scrophulariaceae)

F. N. Hepper eee eene ennt ent tna th tta anne tnntn seen tine th atas tna ta nan 45
Cacalia rugelia: A New Combination for a North American

Senecionoid

T. M. Barkley and Arthur Cronquist ............. eee 48
Typification and Application of the Names Scirpus ameri-

canus Pers. S. olneyi Gray, and S. pungens Vahl.

Alfred E. Schuyler ............ eee eee eee eee eene te nent tn nennen nnn 51
The Correct Authority for Cardamine clematitis

(Cruciferae)

y JU C209 207,177 NERS EE EATE AA IE E E T 53
A New Form of Digitaria sanguinalis

John E. Ebinger .rsccssosssssssessssssssecccserscsssnesensseterevssesacccssssssescescnns 57
The Growth of some New England Perennial Seaweeds

Narman EEO ER aai iori d diassec itaq E a 59
Chromosome Numbers in Kosteletzkya Presl. (Malvaceae)

Orland J. Blanchard, Jv. sscsssccersssescersrsssccccoveccessonconecssenseseceees 64
Name Changes for Seed Plants in the Bahama Flora

William T. Gillis eaea aa eaan A 67
Book Review

JORN EOI: IT: aeien oicseeri tis reei E 139
Book Review

Bornio G uSOhWDOT 5 un udis aszsessensoes iic iaa neem: 142

Book Review
Fer) ASE GS LEPC TYE) S EO E E ERE E TEM NE

Dodova

JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by

ALBION REED HODGDON, Editor-in-Chief
ROLLA MILTON TRYON )
RADCLIFFE BARNES PIKE
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
ALFRED LINN BOGLE
RICHARD EDWIN WEAVER

œ Associate Editors

L

Vol. 76 June, 1974 No. 806

CONTENTS:

A Systematic Treatment of the Native Unifoliolate Crotalarias
of North America (Leguminosae)

Donald R. Wiéndler ....cccccrccccscescsssssercosscessconessscssecceccssscessccosasences 151
The Flora of Mount Monadnock, New Hampshire
Er o e E A E T E E E T 205

Taxonomy of Perityle Section Perityle
(Compositae-Peritylinae)
NH MB AR eia A E A E ER A E A E 229

Carex exilis, Geocaulon lividum and Other Plants of Interest
in Pittsburg, New Hampshire
BEEN TONTO n A n a N eiii 307

(Continued on Inside Back Cover)

The Nem England Botanical Club, Ine.
Botanical Museum, Oxford St., Cambridge, Mass. 02138

RHODORA. — A quarterly journal of botany, devoted primarily to the
flora of North America and floristically related areas. Price $10.00
per year, net, postpaid, in funds payable at par in the United States
currency at Boston. Some back volumes, and single copies are
available. For information and prices write RHODORA at address
given below.

Scientific papers and notes, relating directly or indirectly to the
plants of North America, will be considered by the editorial com-
mittee for publication. Articles concerned with systematic botany
and cytotaxonomy in their broader implications are equally accept-
able. All manuscripts should be double-spaced throughout. Please
conform to the style of recent issues of the journal. Illustrations
can be used only if the cost of engraver's blocks is met through the
author or his institution. Forms may be closed five weeks in advance
of publication. Extracted reprints, if ordered in advance, will be
furnished at cost.

Address manuscripts and proofs to A. Linn Bogle,

Dept. of Botany and Plant Pathology,
University of New Hampshire,
Durham, New Hampshire 03824
Subscriptions and orders for back issues (making all remittances
payable to RHODORA) should be sent to RHODORA, Botanical

Museum, Oxford Street, Cambridge, Mass. 02138. In order to receive

the next number of RHODORA, changes of address must be received

prior to the first day of March, June, September or December.
Second Class Postage Paid at Boston, Mass.

MANUFACTURED BY
THE LEXINGTON PRESS, INC.
LEXINGTON, MASSACHUSETTS

QTRbooora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 76 June, 1974 No. 806

A SYSTEMATIC TREATMENT OF
THE NATIVE UNIFOLIOLATE CROTALARIAS
OF NORTH AMERICA (LEGUMINOSAE)"*

DONALD R. WINDLER

Crotalaria is a large genus of tropical, subtropical, and
temperate plants comprised of about 550 species, over 400
of which occur in Africa. In 1939 Senn reported 31 species
for North America with the majority growing in Mexico
and the West Indies; only 21 of these species were con-
sidered to be native. Eight of the 21 species, including the
Linnean C. sagittalis, form a group that is so distinctive
that Rafinesque (1837) separated it into the subgenus
Iocaulon. The Iocaulon? Crotalarias are annual or perennial
plants characterized by simple leaves; decurrent stipules,
when present; terminal or leaf-opposed inflorescences ; and
glabrous, inflated fruits that become black at maturity.
Senn (1939) indicated that this is the most difficult group
of North American Crotalarias in which to delimit species
and suggested the need for further work.

'A portion of a dissertation submitted as partial fulfillment of the
requirements for the Doctor of Philosophy Degree in the Department
of Botany, University of North Carolina at Chapel Hill.

"Contribution number 7 from Towson State College Herbarium. Cost
of Publication of illustrations assisted by the Towson State College
Faculty Research Fund.

"The name Iocaulon will be used as an adjective throughout this
paper to refer to the unifoliolate Crotalarias related to C. sagittalis.

151

152 Rhodora [Vol. 76

The history of the classification of Crotalaria has been
reviewed by Baker (1914), Senn (1939), and Polhill
(1968). While Baker treated the African species of Crota-
laria primarily, he also reviewed the history of the classi-
fication of the genus and discussed its separation from the
closely related genus Lotononis. Senn reviewed rather
briefly the nomenclatural history of Crotalaria in North
America, and discussed the placement of the North Amer-
ican species into sections described by Baker. Senn's treat-
ment includes a key to the species, nomenclature and
synonomy, Latin descriptions for newly described species,
lists of specimens examined, and a few comments on each
of the taxa. Most species lack descriptions and all lack
illustrations, which makes it difficult to identify a plant if
the key only narrows the choice to a group of species,

Polhill (1968), while dealing primarily with the African
species of Crotalaria, presented a discussion of the history
of generic subdivisions, discussed typification of the genus,
and proposed a new arrangement of species into sections
based on combinations of flower characters. The characters
used were: (1) type and position of standard appendages,
(2) twisting of the keel, (3) lobing of the calyx, (4) calyx-
keel length ratio, (5) pubescence of the style, (6) style
shape, and (7) receptacle size. Polhill placed C. sagittalis
and its relatives in the section Calycinae Wight & Arnott.
All the characters that Polhill used for this placement were
found to be consistently present in the Iocaulon Crotalarias
studied.

COLLECTIONS STUDIED

In addition to the personal collections made in the United
States and Mexico, specimens were borrowed from twenty-
one herbaria. The names and locations of the herbaria
from which specimens were borrowed appear below, ac-
companied by the abbreviation of each as listed in Index
Herbariorum, except in one case, marked by an asterisk,
where no listing could be found for the herbarium and an
arbitrary abbreviation is used.

1974] Crotalarias — Windler 153

A— Arnold Arboretum, Cambridge, Massachusetts.
ALU — University of Alabama, University, Alabama. AUA —
Auburn University, Auburn, Alabama. GA — University of
Georgia, Athens, Georgia. GH — Gray Herbarium, Cam-.
bridge, Massachusetts. LAF — University of Southwestern
Louisiana, Lafayette, Louisiana. MICH — University of
Michigan, Ann Arbor, Michigan. *MISS — University of
Mississippi, University, Mississippi. NCU — University of
North Carolina, Chapel Hill, North Carolina. NY — The
New York Botanical Garden, New York, New York. SIU —
Southern Illinois University, Carbondale, Illinois. TEX —
University of Texas, Austin, Texas. UC — University of
California, Berkeley, California. us — United States Na-
tional Museum, Washington, D.C. usF— University of
South Florida, Tampa, Florida. wis — University of Wis-
consin, Madison, Wisconsin,

BM — British Museum of Natural History, London, Eng-
land. G-— Conservatoire et Jardin Botanique, Geneva,
Switzerland. K-— Royal Botanic Gardens, Kew, Great
Britain. MEXU — Universidad Nacional de México, Mexico
City, Mexico. P — Museum Nacional d'Histoire Naturelle,
Paris, France.

Crotalaria plants from more than 60 locations in the
United States and Mexico were grown from seed at the
North Carolina Botanical Garden, Chapel Hill, during the
summer of 1968. Data from the garden plantings were
used in evaluating the status of taxa. Another paper de-
tailing the results of the field and garden studies is published
elsewhere (Windler, 1973).

SYSTEMATIC TREATMENT

CROTALARIA (Dillenius ex Linnaeus, Hort. Cliff. p. 218,
1731, pro parte) L., Sp. Pl. p. 714 pro maxima parte. Type
species: Crotalaria lotifolia L. (see Polhill, 1968 pp. 174-5).
GENERAL DESCRIPTION OF THE NATIVE IOCAULON CROTALARIAS

Erect, ascending, spreading, decumbent, or procumbent

154 Rhodora [Vol. 76

terrestrial herbs or shrubs. Root a taproot with slender
branches, nodules frequently present. Stems single in the
erect species to many in the decumbent and procumbent
species, the epidermis green, frequently becoming purple,
with an indument varying from nearly glabrous to serice-
ous, of spreading or appressed, simple trichomes, Stipules
absent or varying from a pair of subulate lobes projecting
from near the leaf base to inverted-sagittate, decurrent,
foliaceous appendages which wing the stem for a portion
of the subtending internode, a full internode, or for several
of the subtending internodes, the lobes spreading, incurved
toward the stem or parallel to the stem. Leaves alternate,
unifoliolate, undivided, entire, ciliate, membranaceous to
fleshy, broadly ovate to lanceolate or linear, 0.9-10.2 cm.
long, 0.3-4.6 cm. wide, the adaxial surface dark green, gla-
brous, or with spreading or appressed, simple trichomes,
the abaxial surface usually lighter green than the adaxial
and with spreading or appressed simple trichomes; petioles
0.5-3 mm. long.

Inflorescences loose racemes, peduncles borne terminally,
or opposite leaves and 0.0-3 cm. below the node. Bracts
lanceolate, elliptic, subulate, or linear, to 14 mm. long, one
at the base of each flower, frequently persistent after flower
or fruit abscission. Flowers papilionaceous. Calyces bilabi-
ate, deeply cleft, the upper two lobes free and broader than
the lower three, the lower usually remaining coherent, the
calyx bearing two lateral bracteoles, one on either side.
Standards (vexillum, banner) orbicular, obovate, or ellip-
tic-oblong, the apex rounded or retuse, 4 mm. shorter to 6
mm. longer than the upper calyx lobes, yellow, the adaxial
surface frequently becoming red-tinged, lined or unlined,
nearly glabrous with a few trichomes along the main vein
on the adaxial surface, with lamelliform appendages at the
base of the abaxial surface of the blade. Wings (alae) ob-
long, attached by a stalk continuous with the lower margin
(cleaver-shaped), the blades usually puckered between some
of the veins, yellow. Keel (carina) petals twisted at the
tip, white or greenish-white, lanate-ciliate. Stamens 10,

1974] Crotalarias — Windler 155

monadelphous, the staminal tube split on the upper side,
anthers dimorphie, with basifixed long anthers alternating
with medifixed short anthers, the short anthers positioned
above the elongate at anthesis. Styles geniculate at the
base, bearing a spiral line of trichomes transversing the
length and culminating in a tuft of trichomes on the stigma.

Fruit an inflated, sub-cylindric, short-stipitate legume,
glabrous inside and out, becoming black at maturity. Seeds
7-46 per pod, oblique-cordiform, smooth.

The systematic treatment presented here covers twelve
native North American unifoliolate species of Crotalaria
section Calycinae. The descriptions are based on materials
collected during the study and on numerous herbarium
specimens. Measurements of stipule width are taken at
the base of the sinus between the two lobes and include the
width of both at that point. Length measurements of the
calyx and standard are made from the base of the torus to
the tip of the respective structure in the undissected flower.
Illustrations were prepared by Mrs. Jerry Meyer.

Since data on cross-fertility between the numerous
populations sampled is not available, taxa have been dis-
tinguished primarily on patterns of morphological discon-
tinuity. It is assumed that these discontinuities also reflect
breeding discontinuities, especially in areas where species
are sympatric. In mountainous areas, however, many pop-
ulations are well isolated, frequently leading to morpholog-
ical discontinuity between populations. In these instances
greater emphasis has been placed on similarities in inter-
preting relationships.

Diagnostic characters that have been useful in the group
are plant habit; general size; position of inflorescence;
presence, absence, size, and shape of the stipules; length
and position of trichomes; and flower size. Most of the
diagnostic characters are present on the plants during the
entire growing season.

The synonomy includes only those names that were pos-
sible to check either against types or clear descriptions. A

156

Rhodora [Vol. 76

special effort is made to point out all places where the
present treatment differs from that of Senn's. l

A set of duplicate specimens from the author’s field col-
lections has been deposited in the herbarium of the Univer-
sity of North Carolina at Chapel Hill (NCU).

KEY TO THE SPECIES OF IOCAULON CROTALARIAS

WHICH OCCUR IN NORTH AMERICA

A. Inflorescence terminal, or terminal and also leaf-

opposed. ...........leeee RR heh B.
B. Plants with decurrent stipules. (West Indies, Cen-

tral and South America). ........... 1. C. pilosa.
B. Plants without decurrent stipules. ............ C.

C. Shrubs and subshrubs with diffuse habit. .. D.

D. Peduncles 9 cm. or more long; calyx 15-20
mm. long. (Mexico, Central and South
America). .......... esses 2. C, nitens.

D. Peduncles 6 cm. or less long; calyx 10-11
mm. long. (Northwestern Mexico). .....

3. C. brevipedunculata.

C. Herb with decumbent or ascending habit.

(Western Mexico). ........ 4. C. polyphylla.

A. Inflorescence not terminal, always leaf-opposed. .....
eee e hehe ehe E.
E. Plants erect or strongly ascending. ........... F.
F. Stem pubescence spreading; adaxial leaf sur-
faces regularly pubescent. ............... G.

G. Stipules with lobes recurved. (West Indies

and South America). .... 5. C. stipularia.

G. Stipules with lobes spreading, (Through-

out). co.cc eee eee ee eee 6. C. sagittalis.

F. Stem pubescence appressed, adaxial leaf sur-

faces glabrous or with appressed trichomes.

H. Upper leaf surface glabrous. ......... I.
I. Plants perennial; stipules usually con-
spicuous; peduncles 7.5-23 cm. long.

1974] Crotalarias — Windler 157

(Southern United States). ..........

NEP EL o M NC 7. C. purshii.

I. Plants annual; stipules absent or incon-
spicuous; peduncles 2.5-8 cm. long.

(Mexico and Central America). ......

"T c IEEE 8. C. quercetorum.

H. Upper leaf surface bearing trichomes. .. J.

J. Peduncles 0.5-2 cm. long, usually as

thick as the stem; stipules present;
trichomes dense, obscuring the stem.

HMETICO)S S.V VI 9. C. mexicana.

J. Peduncles 2.5 cm. long or longer, thinner

than the stem; stipules absent; tri-

chomes not obscuring the stem (except
occasionally near the top). (Southern

United States). .. 10. C. rotundifolia.

E. Plants decumbent, procumbent or spreading. .. K.
K. Mature plants decumbent or procumbent from
a thick taproot, stems usually less than 6 dm.
long; trichomes on stem 0.3-3.2 mm. long.
(Mexico and Southeastern United States). ..
eer er ere o o 10. C. rotundifolia.

K. Mature plants diffuse or spreading, stems usu-
ally more than 6 dm. long; trichomes on stem

usually less than 1 mm. long. (Mexico). .. L.
L. Calyx 13-20 mm. long; stipules 0.5-2.7 cm.
broad. (Mexico). .. 11. C. bupleurifolia.
L. Calyx 7.5-12 mm. long; stipules narrow or
absent. (Mexico). .. 12. C. nayaritensis.

1. Crotalaria pilosa Miller, Garden Dict. No. 2. 1768, non
C. pilosa Roxb. ex Mart. Denkschr. Konigl. Akad. Wiss.
München 6: 156, 1920, non C. pilosa Thunb. Prod. Pl.
Capensium 125.

Crotalaria pterocaula Desv. J. Bot. (Desvaux) 3: 76.
1814. Type: not located.

Crotalaria genistella HBK. Nov. Gen. et. Sp. Pl. 6: 398.
1824. Type: Humboldt 2050 (P, mc Microfiche observed).

158 Rhodora [Vol. 76

ac a

Figure 1. Crotalaria pilosa.
A. Habit and leaf variation
B. Stem

159

Crotalarias — Windler

1974]

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160 Rhodora [Vol. 76

Crotalaria pilosa Miller var. skutchii Senn, Rhodora 41:
331. 1939. Type: Skutch 3071 (GH).

Annual erect herb with a taproot to 0.4 cm. thick. Stem
single, 2.8-9.5 dm. long, 1.3-4 mm. thick, the longest inter-
node 2.8-8.2 cm. long; vestiture of appressed or loosely ap-
pressed trichomes 0.5-1.5 mm. long. Stipules present, de-
current more than the length of the internode, 0.3-0.8 cm.
wide, the width constant, the lobes 2.1-9.7 mm. long, usually
not spreading. Leaves elliptic, lanceolate, narrowly lanceo-
late, or linear, 3.3-8.6 cm. long, 3.5-25 mm. wide, base
rounded to cuneate, the apex acute; vestiture of both sur-
faces of appressed trichomes 0.4-2 mm. long; petioles 1-1.3
mm. long. Inflorescences terminal or leaf-opposed and ter-
minal, peduncles 4.5-12 cm. long. Bracts sessile, linear, or
subulate, 3.1-4 mm. long, 0.3-0.5 mm. wide; pedicels 3-4.8
mm. long. Calyces 9-11.5 mm. long, the tube 2.5-3.1 mm.
long, with appressed, loosely appressed, or somewhat
spreading trichomes 0.5-1 mm. long; bracteoles elliptic,
narrowly elliptic, or subulate, 2.5-4 mm. long, 0.4-0.5 mm.
wide. Corollas yellow, the standards 9-12.8 mm. long,
equaling or to 1.7 mm. longer than the upper calyx lobes;
elongate anthers 1.5-2 mm. long, short anthers 0.3-0.5 mm.
long ; style 5.5-5.8 mm. long. Fruits 2.5-3.6 cm. long, breadth
not known; seeds tan, brown, or red-brown, 15-23 per pod,
2.9.5 mm. long. Chromosome number: n—16. Flowering
time: July-March. Habitat: sandy and gravelly soils in
fields and open pine woods, elevation sea level to 2,400 feet.
Range: Costa Rica, Cuba, Jamaica, Panama, (Also widely
distributed in South America.) Figure 1, Map 1.

Type: “La Vera Cruz in New Spain" (Mexico, Cuba or
Jamaica?), Houstoun (BM; photo., NY!).

In spite of the fact that Miller, in his original descrip-
tion, cited Crotalaria pilosa as “growing naturally at La
Vera Cruz in New Spain", no specimens have been observed
from Veracruz, Mexico. The citation appears to be in
error. The probable true type locality was either in Cuba
or Jamaica. Senn, in his treatment of C. pilosa, recognized
the species as being composed of three varieties. Senn's

1974] Crotalarias — Windler 161

variety skutchii is here treated as being a part of variety
pilosa; his variety robusta is transferred to C. bupleurifolia
and treated as a variety of that species. No field collec-
tions were made of C. pilosa during this study and no seed
was obtained for garden plantings.

Crotalaria pilosa is characterized by its erect habit, ter-
minal or leaf-opposed and terminal inflorescences, and
stipules that wing the stem for more than one node below
the node of origin (Figure 1). Occasional specimens have
spreading lobes at the summit of the stipules. Such speci-
mens observed are Britton, Britton, & Shafer 734 (NY)
and Bro. Leon & Fr. M. Rosca 8837 (NY), both from Cuba.
Crotalaria pilosa is alone among the species covered in this
treatment in possessing both terminal inflorescences and
stipules.

REPRESENTATIVE SPECIMENS: COSTA RICA. Cina Grande, entre
San Ramon y Otevas, Brenes 11332 (NY). CUBA. Isle of Pines:
pinelands, Britton, Britton, and Wilson 14615 (NY, US); PINAR DEL
RIO: near Rio de las Vueltas, N of S. Diego de los Banos, Bro. Leon
4908 (GH). DOMINICAN REPUBLIC. TRUJILLO: between Baya-
guana and Guerra, Howard and Howard 9924 (cH). JAMAICA.
Upper Clarendon, James Hill Savannah, Harris 12844 (GH, NY, US).
PANAMA. Ancon Hill, Canal Zone, open hillside, Killip 12075 (us).
TRINIDAD. Without date or specific locality, de Suber 369 (P).

2. Crotalaria nitens HBK. Nov. Gen. et Sp. 6: 399. 1824.
Crotalaria bracteata Schlecht. et Cham. Linnaea 5: 575.
1830, non Roxburgh, Hort. Bengal 54. 1814. (Type:
Schiede 597, Hacienda de la Laguna, (B; photo., F!).
Crotalaria schiedeana Steudel, Nomen. Botanicus. 1: 445.
1840. Type: same as for C. bracteata for which this was
a nomen nov.

Shrub with unknown root type. Stems 5-15 dm. long, 2-4
mm. thick, the longest internode 1-5.5 em. long; vestiture
of appressed or loosely appressed trichomes, 0.5-1 mm. long.
Stipules absent. Leaves elliptic, lanceolate, oblong, or
oblanceolate, 2.5-7.5 (10) em. long, 3.5-18.5 mm. wide, base
rounded to cuneate, the apex rounded, acuminate, or acute;
vestiture of both surfaces of appresed or loosely appressed

162 Rhodora [Vol. 76

trichomes, 0.3-1 mm. long; petioles 1.3-3 mm. long. Inflor-
escenses terminal, peduncles 6-51 cm. long. Bracts petiolate,
lanceolate, the petioles 0.3-1 mm. long, the blades 4-14 mm.
long, 0.6-3 mm. wide; pedicels 4.1-7 mm. long. Calyces
15-18 (20) mm. long, the tubes 3.3-5 mm. long, bearing
appressed or loosely appressed trichomes 0.3-1 mm. long;
bracteoles lanceolate, elliptic-lanceolate or linear, 4.5-9 mm.
long, 0.5-1.4 mm. wide. Corollas yellow, the standards
13.5-19 (22) mm. long, equaling to 2.5 mm. longer than the
upper calyx lobes; elongate anthers 2.2-3.6 mm. long, short
anthers 0.6-1.2 mm. long; styles 8.5-11.5 mm. long. Fruits
2.5-3.2 em. long, breadth not known; seeds grey-brown,
brown, or red-brown, 26-40 per pod, 1.8-2.8 mm. long.
Chromosome number: not known. Flowering time: Sep-
tember-December. Habitat: bare soil, dry open mountain-
sides and pinelands, elevation 4,000-7,200 feet. Range:
Guatemala; Mexico: Chiapas, Jalisco, Oaxaca, Veracruz
(Also widely distributed in South America).

Type: Mariquita (Colombia), Humboldt (P, nc Micro-
fiche observed).

A photograph of the type of Crotalaria nitens was ob-
served in the Ipc microfiche photographs of the Humboldt,
Bonpland, and Kunth herbarium (P). It bears only the
name C. nitida and the location Mariquita which is also
mentioned in the location sites for C. nitens. It compares
favorably with the photograph examined by Senn (1939)
of the “co-type” in the Willdenow Herbarium. Senn’s appli-
cation of the name is followed in this treatment.

Crotalaria nitens is characterized by its shrubby habit,
lack of stipules, terminal inflorescences, and large flowers.
Of the North American species, it most resembles C. brevi-
pedunculata but differs from it in having a larger flower
size and longer peduncles.

Two varieties are recognized:
Peduncles stout, 1-2 mm. thick; bracts 7.5-14 mm. long,
1.2-3 mm. wide. .......... 2a. C. nitens var. nitens.
Peduncles slender, 0.5-0.6 mm. thick, bracts 4-5 mm. long,
0.6-0.8 mm. wide. ........ 2b. C. nitens var. gracilis.

1974] Crotalarias — Windler 163

mv,
WIN

(hi i |
y Wt

N
N

=—

a Z

Figure 2. Crotalaria nitens var. nitens.
A. Habit and leaf variation

B. Stem
Crotalaria nitens var. gracilis.
C. Habit

D. Stem

164 Rhodora [Vol. 76

2a. Crotalaria nitens HBK var. nitens Figure 2, Map 1.

REPRESENTATIVE SPECIMENS: BRITISH HONDURAS. EL CAYO:
near Hydram, Rio Frio Augustine, Mt. Pine Ridge, Hunt 214 (us).
GUATEMALA. VERAPAZ: Coban, 1,350 m., Türkheim 1948 (MICH,
NY, US). HONDURAS. Comayagua, near Siguatepeque, 1,080-1,400
m., Standley 56370 (Us). MEXICO. CHIAPAS: Muncipio of Tenejapa,
6,000 ft, Breedlove 10976 (US); OAXACA: Entre Puerto Eligio a
Comaltepec, Calderon 394 (MICH, US); VERACRUZ: Zacuapan, Purpus
8007 (NY, US).

2b. Crotalaria nitens HBK var. gracilis Windler, Phytologia
21: 263, 1971. Figure 2, Map 1.

Type: Mexico: Jalisco; Sierra de Halo, logging road
7 miles south southwest of Tecalitlan and extending south-
east toward San Isidaro, 13-16 miles from highway. Mc-
Vaugh & Koelz 1188 (MICH).

3. Crotalaria brevipedunculata Windler, Phytologia 21: 257.
1971.

Shrub or suffrutescent herb with root type unknown.
Stems more than 3 dm. long, 3.5 mm. thick, the longest
internode 1.2 em. long; vestiture of dense, short spreading
trichomes 0.6-0.7 mm. long. Stipules absent. Leaves elliptic
or elliptic-oblong, 2.1-4.4 em. long, 5-13.5 mm. wide, base
euneate to widely acute, the apex acuminate; vestiture of
loosely appressed trichomes 0.5-0.9 mm. long; petioles 1.6-
2.1 mm. long. Inflorescenses terminal and also leaf-opposed,
peduncles 1.2-6.1 cm. long. Bracts sessile, linear or nar-
rowly lanceolate, 4-4.4 mm. long, 0.3-0.6 mm, wide; pedicels
3.8-4.2 mm. long. Calyces 10.5-11 mm. long, the tubes 2.5
mm. long, bearing spreading trichomes 0.5-0.7 mm. long;
bracteoles linear, 3.5 mm. long, 0.2-0.3 mm. wide. Corollas
yellow, the standards 10.5-11 mm. long, equaling the upper
calyx lobes; elongate anthers 1.6-2 mm. long, short anthers
0.3-0.4 mm. long; style 6.4 mm. long. Fruits and seeds not
known. Chromosome number: not known. Flowering date:
December 20. Habitat: shady canyon slope with oaks and
palms, elevation 3,500 feet. Range: Mexico: Durango,
Sinaloa. Figure 3, Map 2.
= Type: Sierra Tres Picos, Durango, infrequent, scattered.
Gentry 5311 (GH).

1974] Crotalarias — Windler 165

Figure 3. Crotalaria brevipedunculata.
A. Habit
B. Stem

166 Rhodora [Vol. 76

X C.brevipedunculata |

€ C.mexicana M LL—— —
6 C. polyphylla

e C.quercetorum
——. | |
Map 2. Distribution of Crotalaria brevipedunculata, C. mexicana,
C. polyphylla, and C. quercetorum.

Crotalaria brevipedunculata is characterized by its lack
of stipules, its short terminal inflorescences and small
flowers. It most nearly resembles C. nitens, but differs
from it in its smaller flower size and shorter peduncles.

In addition to the holotype only one other collection of
this species has been observed: Mexico. SINALOA: Puerto
a Tamiapa, Gentry 5815 (MICH, NY).

4. Crotalaria polyphylla Riley, Kew Bulletin 43: 333. 1923.

Perennial decumbent or ascending herb with a taproot
to 1.4 em. thick. Stems 4-10, 1.2-3 dm. long, 1-2.5 mm.
thick, the longest internode 1-3.2 cm, long; vestiture of
short appressed or long spreading trichomes, the appressed
0.3-0.6 mm. long, the spreading 0.8-1.2 mm. long. Stipules

1974] Crotalarias — Windler 167

Figure 4. Crotalaria polyphylla.
A. Habit and leaf variation
B. Stem

168 Rhodora [Vol. 76

absent. Leaves elliptic, oval, or elliptic-oblong, 1.6-4.2 em.
long, 5-183 mm. wide, base rounded to cuneate, the apex
obtuse, rounded, or acuminate; vestiture of both surfaces
of loosely appressed trichomes 0.5-1.1 mm. long; petioles
1-2.5 mm. long. Inflorescences terminal or terminal and also
leaf-opposed, peduncles 4-16 em. long. Bracts sessile, nar-
rowly elliptic to narrowly lanceolate, 2.5-9.5 mm. long,
0.3-1.4 mm. wide; pedicels 4-6 mm. long, Calyces 9-13.5
mm. long, the tubes 2-3 mm. long, bearing appressed tri-
chomes 0.3-1 mm. long; bracteoles linear, narrowly lanceo-
late or narrowly elliptic, 3-7 mm. long, 0.4-1 mm. wide.
Corollas yellow, the standards 10-14 mm. long, 0.5-1.5 mm.
longer than the upper calyx lobes; elongate anthers 1.7-2.3
mm. long, short anthers 0.5-0.7 mm. long; styles 6-7.3 mm.
long. Fruits 1.4-2.4 em. long, 0.5-0.9 em. broad; seeds green,
grey-green, green-brown, brown, or tan, 7-18 per pod,
1.8-2.5 mm. long. Chromosome number n—16. Flowering
time: June-October. Habitat: slopes in open pine woods,
elevation 6,500-9,500 feet, Range: Mexico: Chihuahua,
Durango, Jalisco, Sinaloa. Figure 4, Map 2.

Type: Mexico: Sierra Madre, Seeman 2187 (K!).

Material here treated as Crotalaria polyphylla was in-
cluded by Senn (1939) in C, maritima var. maritima;
however, its terminal inflorescences and complete lack of
stipules clearly separate it from C. rotundifolia (for treat-
ment of Senn's C. maritima, see C. rotundifolia var. rotundi-
folia). The epithet polyphylla was used by Senn as the
varietal name for Mexican plants he considered to be C.
purshii (C. purshii var. polyphylla). Personal examination
of the type has led to the recognition of C. polyphylla at
the specific level. (For treatment of plants previously
placed in C. purshii var. polyphylla by Senn see C. querce-
torum.) Crotalaria polyphylla is characterized by its
decumbent or ascending habit, lack of stipules, and terminal
inflorescences.

REPRESENTATIVE SPECIMENS: MEXICO. CHIHUAHUA: Memelichi,
Rio Mayo, pine slopes, Standley 2701 (GH, MEXU, US). DURANGO:

1974] Crotalarias — Windler 169

32.7 mi. W of Durango in allano slope below pine forest, 8,500 ft.,
Johnston 2694 (MEXU, MICH). JALISCO: Sierra de la Campana,
along road to Mascota, 1,900-2,000 m., McVaugh 13777 (MEXU, MICH,
US). SINALOA: Ocurahui, Sierra Surotato, Gentry 6227 (MICH, NY).

5. Crotalaria stipularia Desvaux, J. Bot. (Desvaux) 3: 76.
1814.

Crotalaria espadilla HBK. Nov. Gen. et Sp. Plantarum
6: 399. 1824. Type: Humboldt 1025, (P; observed IDC
Microfiche).

Crotalaria stipularia var. serpyllifolia DC., Prodromus
2: 124. 1825. Type: (G; observed DC Microfiche in Pro-
dromus Herbarium).

Crotalaria stipularis, orthographic error by Grisebach,
Flora Brit. West Ind. Isl. 178. 1864.

Crotalaria stipularis var. oblongata Grisebach, Flora
Brit. West Ind. Isl. 178. 1864.

Crotalaria stipularis var. sericea Grisebach, Flora Brit.
West Ind. Isl. 178. 1864.

Crotalaria stipularis var. espadilla Kuntze, Revisio
Generum Plantarum 1: 175, 1891.

Crotalaria stipularia forma eliptica Chodat & Hassler,
4: 835. 1904.

Crotalaria stipularia var. grandifolia Senn, Rhodora 41:
333. 1939. Type: Hahn 239 (GH).

Annual, erect herb with a taproot to 0.6 cm. thick. Stem
solitary, 1-6.3 dm. long, 1.2-5 mm. thick, the longest inter-
node 3-8 em. long; vestiture of loose appressed and/or
spreading trichomes, 0.5-3 mm. long. Stipules present,
decurrent the length of the internode, 1.1-3.2 cm. wide at
the top, tapering to the subtending node, the stipule lobes
ineurved 1.2-1.6 em. long. Leaves oval, oblong, elliptie, or
lanceolate, 2.2-8.2 cm. long, 7.5-25 (-37) mm. wide, base
rounded to cuneate, the apex retuse, rounded, acuminate,
or acute; vestiture of both surfaces of appressed or loosely
appressed trichomes 0.5-2.1 mm. long; petioles 0.5-1.5 mm.
long. Inflorescences leaf-opposed, peduncles 3.1-10 cm. long.
Bracts petiolate, elliptic, lanceolate to linear, the petioles

170 Rhodora [Vol. 76

Figure 5. Crotalaria stipularia.
A. Habit and leaf variation
B. Stem

1974] Crotalarias — Windler 171

1-1.6 mm. long, the blades 3-5 mm, long, 0.3-1.2 mm. wide;
pedicels 2-4 mm. long. Calyces 8-10 mm, long, the tubes
2.2.3 mm. long, bearing appressed or loosely appressed
trichomes 0.7-1.2 mm. long; bracteoles elliptic, lanceolate,
or linear, 3-4.5 mm. long, 0.4-0.7 mm. wide, Corollas yel-
low, the standards 6-10 mm. long, equaling to 1.5 mm.
shorter than the upper calyx lobes; elongate anthers 1.1-1.3
mm. long, short anthers 0.3-0.5 mm. long; styles 4.5-5.2
mm. long. Fruits 2.5-3.3 cm. long, width not known; seeds
tan, bronze, red-brown, or dark brown, 17-41 per pod, 2-3
mm. long. Chromosome number: n—16. Flowering time:
February-December. Habitat: sandy soils, savannahs,
clearings, fields and roadsides, elevations to 2,300 feet.
Range: West Indies; Dominican Republic, Dominica,
Guadeloupe, Haiti, Martinique, Puerto Rico, Trinidad.
(Also widely distributed in South America.) Figure 5,
Map 1.
Type: Not located.

Crotalaria stipularia, as treated by Senn, was divided
into three varieties. Varieties "typica" and “orandifolia”
are not different except in overall plant size. Both varia-
tions are found in a single location mass collection from
Puerto Rico sent to me by Dr. R. A. Howard of the Arnold
Arboretum. This size variation is probably typical of plants
of this species at most locations. Senn’s application of De
Candolle’s C. stipularia var. serpyllifolia is to a plant in
which all of the original leaves were lost during drought
(or some other catastrophe) and new branches with small
leaves and shorter internodes had developed before the
plant was collected. The wide decurrent stipules were not
lost when the leaves were lost, which accounts for the
variety having large stipules. The IDC Microfiche photo-
graph of De Candolle’s specimen in the Prodromus Her-
barium shows a plant with smaller leaves, but not quite
so small as those in Senn’s cited specimens. No field col-
lections were made of C. stipularia during this study, but
seeds were obtained for garden studies (Irwin 1050 and
Stimson 4264).

172 Rhodora [Vol. 76

Crotalaria stipularia is characterized by its erect habit,
leaf-opposed inflorescences, small flowers, and large stipules
with incurved lobes. It most nearly resembles C. sagittalis,
but the two taxa may be separated by stipule characters.

REPRESENTATIVE SPECIMENS: DOMINICA. Grand Savannah, Lloyd
855 (NY). DOMINICAN REPUBLIC. Prov. de la Vega: near Gara-
bawa at Rio Yaquir, Fuertes 1669 (A, NY, P). GUADELOUPE.
Montebello, alt. 50 m., Questel 432 (P, us). HAITI. Dept. du Nord:
east of Marmelade, grassy summit of mountain, Leonard 7222 (us).
MARTINIQUE. Fort Vaillant, Hahn 239 (GH; Photo., Us). PUERTO
RICO. Along railroad N of Mayaguez, Heller 4574 (MICH, NY, US).
ST. VINCENT. near Barrovollic (Leeward), Smith & Smith 1053
(NY). TRINIDAD. south of Dabadie, Bulton & Hazen 702 (GH).

6. Crotalaria sagittalis L., Sp. Pl. p, 714. 1753.

Crotalaria fruticosa Miller, Gard. Dict. (pages not num-
bered) 1768. Type: Jamaica, Houstoun, (BM, photo., NCU).

Anonymos sagittalis (L.) Walt., Flora Carol. 181. 1788.
Type: same as for C. sagittalis L.

Crotalaria parviflora Roth, Catalecta Bot. 1: 83. 1797.
Type: not located.

Crotalaria sagittalis var. B oblonga Michx. Flora Boreali-
Am. 2: 55. 1803. Type locality: "Virginia et Carolina".

Crotalaria platycarpa Link, Enum. Pl. Hort. Reg. Bot.
Berol. Alt. part 2, 227. 1822. Type: destroyed in Berlin.

Crotalaria pringlei A. Gray, Proc. Am. Acad. 17: 200.
1881-2. Type: Arizona, Pringle, summer 1881 (GH).

Crotalaria sagittalis var. fruticosa (Miller) Fawcett and
Rendle, Fl. Jamaica 4: 10. 1920.

Crotalaria sagittalis var. blumeriana Senn, Rhodora 41:
339. 1939. Type: Blumer 1772 (GH).

Crotalaria tuerckheimii Senn, Rhodora 41: 339. 1939.
Type: von Tuerckheim II 1282 (GH).

Crotalaria belizensis Lundell, Amer. Midl, Naturalist 29:
473. 1943. Type: Gentle 4144 at (NY).

Annual or rarely perennial, erect herb with a taproot to
0.6 cm. thick. Stems 1 (-7), 0.25-4.1 dm. long, 1-2.5 mm.
thick, the longest internode 0.4-4.2 em. long; vestiture of
long spreading trichomes, 1.5-3 mm. long. Stipules present,

1974] Crotalarias — Windler his

Figure 6. Crotalaria sagittalis.
A. Habit and leaf variation
B. Stem

Rhodora [ Vol.

Map 3.

FER Oa DON A
Tet "ese
e! NA

ir

C. sagittalis

- YT) = on
- > J
252525» u o 1 ka
d
o i
3
“ee
*]
(e
^ —
} 297...
W " —7»

Distribution of Crotalaria sagittalis.

1974] Crotalarias — Windler 175

decurrent from 1% to the full length of the internode, 0.35-
1.7 cm. wide at the top, tapering to the subtending node, the
stipule lobes spreading 0.4-1.8 cm. long. Leaves elliptic,
oblong, lanceolate, or linear-lanceolate, 1.1-10 cm. long,
2.5-15 (22.5) mm. wide, base rounded to cuneate, the apex
obtuse, mucronate, acuminate, or acute; vestiture of both
surfaces of spreading or loosely appressed trichomes 1-2.9
mm. long; petioles 1.2-2.5 mm. long. Inflorescences leaf-
opposed, peduncles 2-8 (-12) cm. long. Bracts petiolate,
lanceolate to elliptie lanceolate, the petioles 0.5-1.5 mm.
long, the blades 4-8.1 mm. long, 0.5-1.3 mm. wide; pedicels
3.5-5.5 mm. long. Calyces 7-15 mm. long, the tubes 2.3-3.9
mm. long, bearing loosely appressed trichomes 0.8-2 mm.
long; bracteoles elliptic lanceolate, lanceolate, or linear,
2.5-7 mm. long, 0.7-1.2 mm. wide. Corollas yellow, the
standards 6-12.5 mm. long, equaling to 4 mm. shorter
than the upper calyx lobes; elongate anthers 0.9-2 mm.
long, short anthers 0.3-0.5 mm. long; styles 4-5.3 mm, long.
Fruits 1-2.7 (3.6) cm. long, 0.5-1.1 cm. broad; seeds tan,
tan-green, olive, green-brown, or coffee-brown, 7-32 per pod,
1.4-3 mm. long. Chromosome number: n=16. Flowering
time: May-October. Habitat: road and railroad rights-of-
way, disturbed ground, sand deposits, fields, prairies, lake
margins, and dry oak and pine forests, elevation sea level
— 8,500 feet. Range: Eastern and Central United States
through Mexico and Central America to Panama and on
the Greater Antilles. Figure 6, Map 3.

Lectotype: “2 Kalm sagittalis" (LINN) (see Taxon 21:
545.)

Type locality: “Brasilia, Virginia" as cited in Species
Plantarum.

Plants included here under Crotalaria sagittalis include
material cited by Senn as variety blumeriana and variety
fruticosa, as well as Senn’s variety "typica". Early leaves
of plants of C. sagittalis tend to be broader than the later
leaves, and all branches tend to have narrower leaves than
the main stem. In addition, the further south a plant occurs,
the longer the plant grows, and the more early leaves are

176 Rhodora [Vol. 76

lost before the end of the growing season. There is also
a tendency for southern plants to have narrower early
leaves than northern plants. Garden studies have shown
Crotalaria sagittalis var. blumeriana to be a dry habitat
ecophene of the typical species. Although a great deal of
variation exists in C. sagittalis, especially in the Mexican
representatives, little in the way of a clear pattern that
would allow a useful segregation of the plants (and popu-
lations) into varieties has been detected. Further study of
this wide-spread but poorly understood species, to gain a
more complete picture of the relationships between plant
variation, geographic, and ecological factors, may prove
helpful, especially in Mexico, Central and South America.

REPRESENTATIVE SPECIMENS: BRITISH HONDURAS. Honey Camp,
Lundell 665 (vus). COSTA RICA. Paturage a San Jose de Costa
Rica, Tonduz 443 (MicH, P). CUBA. Pinar del Rio: Herradura,
Britton, Britton, Earle, & Gager 6404 (NY). GUATEMALA. Alta
Verpaz: Coban, alt. 4,300 ft., Tuerckheim 239 (GH, Us). HAITI.
Massif de la Selle Nouvelle Touraine, alt. 1,400 m., Ekman 1501
(Ny, US). HONDURAS, Morazan, hills around Zamorano Valley,
alt. 800-900 m., Swallen 11252, (us). JAMAICA. Castleton, 490 ft.,
Harris 11855 (GH, NY, P, US). NICARAGUA. Comarca ElCabo:
LaTronquera, alt. 60 m., Molina 14881 (Ny). PANAMA. Panama:
Sabana de Dormisolo, near Chepo, Pittier 4686 (Us). EL SALVA-
DOR. Ahuachapan: Ahuachapan, alt. 800-1,000 m., Standley 19747
(GH, NY, US). MEXICO. Baja California: S side of La Carrerita,
Sierra El Taste, 1,500-1,600 m., Carter & Chisaka 8504 (MEXU).
Chiapas: Tenejapa, 7,800 ft., Breedlove 19718 (us). Chihuahua:
Madera, Pennell 19228 (vs). Colima: Colima, Palmer 130 (vs).
Durango: without locality, Garcia 942 (vs). Guanajuato: NE of
Guanajuato, Ripley & Barneby 13351 (NY). Jalisco: near Chapala,
Rose & Painter 7636 (NY). Mexico: Puerto del Salitre, 1,300 m.,
Hinton 1780 (GH, MEXU, NY). Michoacan: vicinity of Coalcoman,
1,000 m., Hinton 12862 (Ny, US). Morelos: Cuernavaca, Rose &
Painter 6850 (GH, Ny, US), Nayarit: Cerro de la Cruz, E of Tepic,
Mexia 669 (GH). Oaxaca: near city of Oaxaca, Rose & Hough 4623
(NY, US). Puebla: 72 km. SE of Mexico City, Weaver 995 (US).
San Luis Potosi: 7-8 mi. NE of Xilitla, King 4360 (us). Sinaloa:
near Mazatlan, Windler & Windler 2885 (NcU). Sonora: NE cf El
Tigre, 6,000 ft, White 4370 (GH, MICH). Tamaulipas: vicinity of
Gomez Farias, ca. 350 m., Palmer 341 (us). Veracruz: SE of Cate-
maco, meadow, Dressler & Jones 114 (MEXU, MICH, NY, US), UNITED

1974] Crotalarias — Windler 177

STATES. Alabama: MOBILE C0.: Dauphin Island, Jones 585 (ALA).
Arkansas: SALINE CO.: ridges, Benton, Demaree 37333 (stu). Con-
necticut: MIDDLESEX CO.: Middlefield, Kofoid (GH). Georgia: JACK-
SON CO.: 11 mi. N of Athens, Cronquist 4571 (GH). Illinois: JACKSON
co.: roadside S of Elkville, Bailey & Swayne 462 (stu, NCU). Indiana:
LAKE CO.: East Gary, Hull 18 July 1936 (GH). Iowa: JONES CO.:
railroad right-of way, Sect. 7, Oxford Twp., Isely 5913 (GA). Kansas:
RILEY Co.: ravines, Norton 80 (GH, P). Kentucky: GRAVES CO.: rail-
road embankment, Fancy Farm, O'Dell & Windler 936 (stv). Louis-
iana: CALCASIEU PARISH: 114 miles NNW of Starks, Sect. 26, Las-
seigne 1579 (LAF). Maryland: Montgomery?, Wildwood, Dowell 1626
(GH). Massachusetts: MIDDLESEX CO.: shore of Winter Pond, Bartlett
224 (GA, GH, TEX, WIS). Michigan: KALAMAZOO CO.: 1 mi. E of
Schoolcraft, Hanes 1796 (GH). Minnesota: CHISAGO CO.: Lindstrom,
Anderson 6458 (GH, WIS). Mississippi: LAYAYETTE CO.: near Tula
Cemetery, 18 mi. SE of Oxford, Pullen 64294 (GA, MISS, NCU).
Missouri: BARTON CO.: prairies, Milford, elev. 750 ft., Demaree 39377
(GA, NCU, USF). New Jersey: BERGEN CO.: Hackensack, route 4,
sandy field, Rush & Svenson 6769 (GH, wis). New York: SARATOGA
co.: N of Crescent, Burnham 8 Sept. 1906 (GH). North Carolina:
CALDWELL CO.: 1 mi. SE of Hudson, Randolph & Randolph 1096
(GH, NCU). Oklahoma: MUSKOGEE CO.: 2 mi. S of Braggs, Edmond
164 (TEx). Pennsylvania: DELAWARE CO.: Wayne, dry gravel bank,
Bartram 1162 (GH). Rhode Island: Cumberland (?) without collec-
tion name & number, Aug. 1840 (GH). South Carolina: GREENVILLE
CO.: 215 mi. SW of Berry's Pond, Freeman 56577A (NCU). Tennessee:
KNOX CO.: sandy soil near Knoxville, Ruth 661 (wis). Texas: JASPER
co.: 3 mi. N of Evadale, Turner & Tharp 3079 (TEX). Vermont:
WINDHAM CO.: Vernon, Blanchard 1901 (GH). Virginia: CULPEPER
co.: 0.3 mi. N of Lakota on Co. Rt. 621, Ahles & James 61692 (NCU).
West Virginia: HARDY co.: Lost River Bridge, wasteground, Core
18 Aug. 1931 (GH).

7. Crotalaria purshii DC., Prodromus 2: 124. 1825.
Crotalaria sagittalis var. B L. Sp. Pl. 714. 1753.
Crotalaria sagittalis var. a linearis Michx., Fl. Boreali-

Am. 2: 55. 1808. Type locality: “Virginia et Carolina".
Crotalaria laevigata Pursh, Fl. Am. Sept. 469, non Lam.,

Encyc., 2: 198. 1786. Type: Pursh specimen not located ;

Syntype was plate in Plukenet, Phytographia-Amagestum

Bot. t. 277, fig. 2. 1694. Specimen from which this figure

was drawn is in Sloane collection at BM.

Perennial, erect herb with a taproot to 0.7 cm. thick.

178 Rhodora [Vol. 76

Figure 7. Crotalaria purshii.
A. Habit and leaf variation
B. Stem

Crotalarias — Windler

C. purshii X

Map 4. Distribution of Crotalaria purshii.

HAA A as ae
Pe HO UI i Aro - C
: VETE CHI Lad Re

179

180 Rhodora [Vol. 76

Stems 1-4 (8), 1.5-5 dm. long, 1-1.5 mm. thick, the longest
internode 3.3-8.5 cm. long; vestiture of short appressed or
loosely appressed trichomes 0.4-0.8 mm. long. Stipules
present, decurrent one-half to the full length of the inter-
node, 0.3-0.9 cm. wide at the top tapering to the base, the
stipule lobes spreading 0.5-1.7 cm. long. Leaves linear,
oblong, linear-lanceolate or elliptic, 4.9-10 cm. long, 1.7-12
mm. wide, base rounded to cuneate, the apex obtuse or
acute; the lower surface bearing appressed trichomes 0.3-
0.7 mm. long, the upper surface glabrous; petioles 0.5-2
mm. long. Inflorescences leaf-opposed, peduncles 7.5-23 cm.
long. Bracts petiolate, elliptic-lanceolate to lanceolate, the
petioles 0.5-1.6 mm. long, the blades 4.5-7 (8.5) mm. long,
0.6-1.8 (2) mm. wide; pedicels 4.5-6 mm. long. Calyces
9-12.65 mm. long, the tubes 2.5-3.5 mm. long, bearing ap-
pressed trichomes 0.2-0.5 mm. long; bracteoles lanceolate
or ellipite-lanceolate, 3.5-5 mm. long, 0.8-1.2 mm. wide.
Corollas yellow, the standards 8-13.5 mm. long, 2 mm.
shorter to 0.5 mm. longer than the upper calyx lobes;
elongate anthers 1.2-1.6 mm. long, short anthers 0.3-0.6
mm. long; styles 2.7-4.6 mm. long. Fruits 1.4-2.8 cm. long,
0.6-1 cm. broad; seeds tan, olive, green-brown, or dark
brown, 11-37 per pod, 1.8-2.9 mm. long. Chromosome num-
ber: n=16. Flowering time: March-October. Habitat:
savannahs, sandhills, and open pine woods, coastal plain.
Range: United States: Alabama, Florida, Georgia, Louis-
iana, Mississippi, North Carolina, South Carolina, Virginia.
Figure 7, Map 4.
Type: Based on Crotalaria laevigata Pursh.
Type locality: Pine woods of Virginia and Carolina.

Crotalaria purshii, named by De Candolle, was based on
material from Virginia and Carolina. He cites C. laevigata
of Pursh as being the same, but proposes the name C.
purshii since C. laevigata had been previously used by
Lamarck. Crotalaria sagittalis var. B L. is based on
Plukenet's figure (Plukenet, 1694, t. 277, Fig. 2) which
is purshii.

Senn (1939) recognized a variety, polyphylla, under

1974] Crotalarias — Windler 181

Crotalaria purshii. Most specimens of this variety are here
treated as C. quercetorum. For further information con-
cerning the epithet polyphylla see the discussion under C.
polyphylla.

Crotalaria purshii is characterized by its erect habit;
sparse, short, appressed pubescence; spreading stipule
lobes; frequently narrow leaves, the upper surfaces of which
are glabrous; and long peduncles. Crotalaria purshii is
similar to C. sagittalis in habit and general appearance in
parts of its range, but it may be distinguished from C.
sagittalis by its glabrous upper leaf surfaces, appressed
pubescence, and long peduncles. The extremely variable
C. rotundifolia var. rotundifolia may occasionally bear a
resemblance to C. purshii, but these plants lack glabrous
upper leaf surfaces.

REPRESENTATIVE SPECIMENS: UNITED STATES. Alabama: BALD-
WIN co.: sandy field, Jones 496 (aua). Florida: CALHOUN CO.:
Cypress Park, Dead Lakes, Godfrey & Clewell 62916 (LAF). Georgia:
WAYNE CO.: open pinelands, 5 mi. S. of Jessup, Duncan 6941 (GA).
Louisiana: ST. TAMMANY PARISH: 7% mi. S of town of Pearl River,
Sect. 13, Lasseigne 1480 (LAF). Mississippi: PEARL RIVER CO.: 1 mile
N of Picayune along Hwy. 12, low pine lands, Bollwinkel & Wun-
derlin 9 (stu). North Carolina: CRAVEN CO.: pine savannah, 0.6 mi.
SW of Havelock, Radford 34504 (NcU). Virginia: ISLE OF WIGHT CO.:
dry sandy pine and oak woods ca. 1 mile SE of Zuni, Fernald &
Long 6233 (GH).

8. Crotalaria quercetorum Brandegee, U. Cal. Pub. Bot.
10: 407. 1924.

Crotalaria purshii var. polyphylla (Riley) Senn, Rhodora
41: 346. 1939. (see note below)

Annual, erect, or ascending herb with a slender taproot
to 0.25 em. thick. Stem solitary, 1.1-5.3 dm. long, 1-3.1 mm.
thick, the longest internode 1.3-5.2 cm. long; vestiture of
short appressed trichomes, 0.3-0.4 mm. long. Stiples usually
absent or very narrow, decurrent 1/5 the length of the
internode, 0.3-0.4 cm. wide at the top narrowing to the
base, the stipule lobes spreading 0.1-0.25 cm. long. Leaves
elliptic, narrowly oblong, narrowly elliptic, narrowly lanceo-

182 Rhodora [Vol. 76

Figure 8. Crotalaria quercetorum.
A. Habit and leaf variation
B. Stem

1974] Crotalarias — Windler 183

late, or linear, 3-6.8 cm. long, 2-5.5 (-8) mm. wide, base
rounded to cuneate, the apex acute or acuminate; vestiture
of appressed trichomes 0.2-0.5 mm. long, the upper surface
usually glabrous; petioles 1.5-3 mm. long. Inflorescences
leaf-opposed, peduncles 2.5-8 cm. long. Bracts sessile, nar-
rowly elliptic, or linear, 2.5-3.5 mm. long, 0.2-0.3 mm. wide;
pedicels 2.5-4.2 mm. long. Calyces 8.2-11 mm. long, the
tubes 1.5-2 mm. long, bearing appressed trichomes 0.2-0.8
mm. long; bracteoles linear or narrowly elliptic, 1.5-3.5 mm.
long, 0.2-0.3 mm. wide. Corollas yellow, the standards
7.5-10.5 mm. long, equaling to 2 mm. shorter than the
upper calyx lobes; elongate anthers 0.8-1.4 mm. long,
short anthers 0.2-0.3 mm. long; styles 3.5-3.6 mm. long.
Fruits 1-2.4 cm. long, 0.4-0.7 cm. broad; seeds olive, green-
brown, or brown, 24-30 per pod, 1.4-2 mm. long. Chromo-
some number: n—16. Flowering time: August-March.
Habitat: ridges and slopes in oak, oak-pine, and pine woods,
elevation 2,600-7,000 feet. Range: Mexico: Chiapas,
Jalisco, Michoacan, Nayarit, Oaxaca, Sinaloa. Figure 8.
Map 2.

Type: Mexico: Chiapas: Hacienda Monserrate, Purpus
9144 (uc!).

Crotalaria quercetorum was included under C. purshii
by Senn. Unfortunately, Senn chose to use the epithet
polyphylla for what he recognized as a variety of C. purshit.
Examination of the type of C. polyphylla Riley leads to the
conclusion that the name is erroneously applied to the ma-
terial here treated as C. quercetorum. The holotype of
C. quercetorum is a mixed sheet with one specimen present
which may be interpreted as intermediate between C.
quercetorum and C. sagittalis. The isotype (US) examined
is also composed of several elements, but all are referable
to C. quercetorum.

Crotalaria quercetorum is characterized by its short
appressed pubescence, lack or near lack or stipules, usually
narrow foliage, short peduncles and small flowers. Crota-
laria quercetorum is similar to C. purshii but may be dis-
tinguished by its much shorter peduncles and its usually

184 Rhodora [Vol. 76

smaller size. Because a number of specimens with inter-
mediate morphology have been collected, hybridization
between C. quercetorum and C. sagittalis is suspected in
parts of their range where they are sympatric. Windler
& Windler 2994 (NCU) is a collection in which some speci-
mens seem to reflect this putative hybridization.

REPRESENTATIVE. SPECIMENS: GUATEMALA. Exact locality not
given, Bernoulli 282 (G). MEXICO. cnurAPAS: Hacienda Monser-
rate, Purpus 9144 (uc, Us); Jalisco: 15-20 mi. SE of Autlan-Sierra
Manantlan ca. 1,700 ft., McVaugh 13973 (MICH); Michoacan: 32 mi.
W of Jiquilpan along road to Manzanillo, 7,000 ft, Fearing 1195
(TEX); Nayarit: 1 mi. from town of Santa Maria del Oro on way
to Laguna Santa Maria, 3,700 ft., Windler & Windler 2917 (NCU);
Oaxaca: 5-6 Km NE of Putla, 350 m., McVaugh 22248 (MICH);
Sinaloa: Puerto a Tamiapa, 4,500 ft., Gentry 5815 (MEXU).

9. Crotalaria mexicana Windler, Phytologia 21: 259. 1971.
Crotalaria sagittalis var. fruticosa (Miller) Fawcett and
Rendle, 4: 10, 1920, pro parte, sensu Senn, non sensu typus.

Annual erect herb with a slender taproot to 0.3 cm. thick.
Stems solitary, 1.2-2.3 dm. long, 1.6-2.4 mm. thick, the
longest internode 1-1.5 em. long; vestiture of dense, ap-
pressed trichomes, 1.2-2.5 mm. long. Stipules absent.
Leaves narrowly elliptic, linear-lanceolate or linear, 2.2-4.6
cm. long, 4-8 mm. wide, base rounded to cuneate, the apex
rounded, acute or acuminate; vestiture of both surfaces of
dense, loosly appressed trichomes 1.1-2.1 mm. long; petioles
0.5-0.6 mm. long. Inflorescences leaf-opposed, peduncles
0.8-2 cm. long. Bracts sessile, lanceolate, 3.3-3.6 mm. long,
0.7-0.8 mm. wide; pedicels 2.5-4 mm. long. Calyces 10.5-11.5
mm. long, the tubes 2-2.5 mm. long, bearing loosely ap-
pressed and spreading trichomes 0.8-2 mm. long ; bracteoles
linear-lanceolate, 4-4.5 mm. long, 0.5-0.6 mm. wide. Corollas
yellow, the standards 10-10.5 mm. long, equaling to 0.5 mm.
shorter than the upper calyx lobes; elongate anthers 1.5-1.7
mm. long, short anthers 0.5-0.6 mm. long; styles 4.8-5.3
mm. long. Fruits 2.1-2.5 em. long, 0.8-1 cm. wide; seeds
brown, number per pod not known, 1.8-2 mm. long.
Chromosome number: not known. Flowering time: Sep-

1974] Crotalarias — Windler 185

—

EREE eee,

= ee er te
€

JL
WEEE

[rJ

Figure 9. Crotalaria mexicana.
A. Habit and leaf variation
B. Stem
C. Leaf pubescence

186 Rhodora [Vol. 76

tember-October. Habitat: dry slopes of mountains, eleva-
tion ca. 6,000 ft. Range: Mexico: Jalisco. Figure 9,
Map 2.

Type: Mexico: Jalisco, mountainside above Etzatlan,
Pringle 8855 (holotype, GH; isotypes, TEX, US).

Crotalaria mexicana is a new species, the representatives
of which were referred by Senn (1939) to C. sagittalis
var. fruiticosa (here treated as C. sagittalis var. sagittalis).
Crotalaria mexicana is most similar to C. sagittalis and
C. quercetorum. It differs from C. sagittalis in its lack of
stipules and absence of spreading pubescence, and from
C. quercetorum in its short thick peduncles and in its
dense pubescence. Crotalaria mexicana is characterized by
its erect habit, dense, appressed pubescence, lack of stipules,
and extremely short, leaf-opposed peduncles.

REPRESENTATIVE SPECIMENS: MEXICO. JALISCO: near Etzatlan,
Pringle 8855 (—type), Pringle 11807 (GH, US), Rose & Painter 7571
(US); near Guadalajara, Rose & Painter 7469 (US).

10. Crotalaria rotundifolia (Walt.) Gmelin in Linne .
Systema Naturae 2: 1095. 1792.

Anonymos rotundifolia Walter, Flora Caroliniana, 181.
1788. Type: BM, photo., NCU!

Crotalaria sagittalis var. ovalis Michx. Flora Boreali-Am.
2: 55. 1803. Type locality: “Virginia et Carolina".

Crotalaria rotundifolia Poiret, Encycl. Meth. 2: 402.
1812, non Gmelin, 1792. (see Ward, 1962).

Crotalaria ovalis (Michx.) Pursh, 469. 1814, nomen
illegit.

Crotalaria hookeriana A. DC. in A.P. & A. DC., Mem.
Soc. Phys. Genéve 9: 97. 1841. Type: not observed.

Crotalaria leptoclona Schauer, Linnaea 20: 737. 1847.
Type: Mexico, Aschenborn 172, not located.

Crotalaria maritima Chapman, Flora South. United
States, Suppl. 614. 1883. Type: not located.

Crotalaria linaria Small, Flora SE United States 679.
1933. Type: Pine Key, Fla., Blodgett (NY!).

1974] Crotalarias — Windler 187

Crotalaria maritima var. linaria (Small) Senn, Rhodora
41: 347. 1939.

Crotalaria rotundifolia var. linaria (Small) Fernald &
Schubert, Rhodora 50: 208. 1948.

Perennial procumbent, decumbent, or ascending herb
with a taproot to 1.2 cm. thick. Stems 2-20, 0.3-6.5 dm.
long, 0.5-3.1 mm. thick, the longest internode 0.6-4.8 cm.
long; vestiture of appressed, loosely appressed, or spreading
trichomes, appressed 0.3-0.6 mm. long, spreading 0.8-3.2
mm. long. Stipules present, with only the stipule lobes
present or the stipules decurrent from 1, to the full length
of the internode, 0.13-2.3 cm. wide at the top tapering to
the base, the stipule lobes spreading 0.15-1.7 cm. long.
Leaves round, oval, oblong, ovate, lanceolate, elliptic, or
linear, 0.9-5.5 em. long, 1.0-33.1 mm. wide, base rounded to
widely cuneate, the apex rounded, obtuse, mucronate, acu-
minate, or acute; vestiture of both surfaces of erect, ap-
pressed, or loosely appressed trichomes 0.3-2.8 mm. long;
petioles 0.5-3 mm. long. Inflorescences leaf-opposed, pe-
duncles 1.3-33.9 cm. long. Bracts with or without petioles,
elliptic, lanceolate, or linear, the petioles to 1.5 mm. long,
the blade 2-7.1 mm. long, 0.3-2 mm. wide; pedicel 2-5 mm.
long. Calyces 7-14.9 mm. long; the tubes 1.5-4.5 mm. long,
bearing appressed, loosely appressed or spreading trichomes
0.2-1.5 mm. long; bracteoles broadly to narrowly elliptic,
lanceolate, or linear, 1.8-6.5 mm. long, 0.3-1.3 mm. wide.
Corollas yellow, the standards 6-15 mm. long, 2.5 mm.
shorter to 1.9 mm, longer than the upper calyx lobes;
elongate anthers 0.9-2.4 mm. long, short anthers 0.3-0.7
mm. long; styles 4-6.9 mm. long. Fruits 1.2-2.8 cm. long,
0.5-1.1 cm. broad; seeds tan, bronze, green-tan, olive, green-
brown, or red-brown, 12-46 per pod, 1.5-2.7 mm. long.
Chromosome number: n—16. Flowering time: United
States, January-October; Mexico, July-March. Habitat:
road and railroad rights-of-way, rocky slopes, sandridges,
sandhills, open pine, and pine-hardwood forests, elevation
sea level — 8,250 feet. Range: Costa Rica; Guatemala;
Mexico: Chihuahua, Guanajuato, Hidalgo, Jalisco, Mexico,

188 Rhodora [Vol. 76

Michoacan, Morelos, Nayarit, Oaxaca, Puebla, San Luis
Potosi, Sonora, Veracruz; El Salvador; United States:
Alabama, Florida, Georgia, Louisiana, Mississippi, North
Carolina, South Carolina, Virginia.

Type: In Walter Herbarium at BM, photo observed.

Crotalaria rotundifolia was first recognized as a distinct
species by Walter (1788) who also recognized it as being
congeneric with C. sagittalis L. Walter apparently thought
that C. rotundifolia and C. sagittalis represented a new
genus distinct from Crotalaria, but, instead of describing
a new genus, he placed them in a miscellaneous category
under the genus name Anonymos. A few years later, Gme-
lin placed Walter's Anonymos species under other genera
(Sprague, 1939). Anonymos rotundifolia was placed with
C. sagittalis L. under Crotalaria. In 1812, Poiret again
published the combination, and he, rather than Gmelin, is
frequently attributed the authorship of the combination
C. rotundifolia. In 1814, Pursh also placed Walter’s
Anonymos rotundifolia into Crotalaria. He, however, pro-
posed a new species name, C. ovalis, and cited Walter’s
name as a synonym, making C. ovalis a superfluous epithet
when published and therefore illegitimate.

Senn (1939), after examining a photograph of Miller's
type of Crotalaria angulata, thought that it was synony-
mous with C. rotundifolia, and the name has since been
used for the American plants. In 1897, Britten and Baker
examined the specimen and concluded it did not differ
from C. biflora L. A comparison of the photograph of
Miller’s specimen with an IDC microfiche of a C. biflora
specimen in the Linnaean Herbarium leads me to agree
with Britten and Baker. In addition, Miller indicated that
the flowers were produced singly on the sides of the
branches and that the leaves were sessile. Neither of these
characters agrees with the type from the Miller Herbarium.
It may also be possible that Miller made a mistake in indi-
cating the location of the place of origin of the seed. It
now seems better to recognize the epithet of Walter and to
reject the Miller name for our American plants.

1974] Crotalarias — Windler 189

Fernald and Schubert (1948), after studying photo-
graphs of Walter's Herbarium (taken by Schubert) at the
British Museum of Natural History, suggested that the
name Crotalaria rotundifolia actually refers to the plants
with short appressed pubescence previously referred to as
C. maritima. A more recent photograph of the specimen
figured by Fernald and Schubert has been supplied by Dr.
Norman K. B. Robson of the British Museum. Dr. Robson
(personal communication) indicates that the plant has
“appressed-sericeous” pubescence and would “run down
easily in Senn's key to C. maritima Chap.". In light of this
information, the name C. rotundifolia is recognized as ap-
plying to the material previously recognized as C. maritima,
as Fernald and Schubert suggested.

Crotalaria rotundifolia may be distinguished by its pro-
cumbent, decumbent, or occasionally ascending habit; leaf-
opposed inflorescences; and the presence of stipules. It is
an extremely variable species, especially in the Florida
peninsula.

Two varieties may be distinguished in the following
way:
Plants with appressed stem pubescence. ..............
RONDE n 10a. C. rotundifolia var. rotundifolia.
Plants with spreading stem pubescence, ..............
10b. C. rotundifolia var. vulgaris.

10a. Crotalaria rotundifolia var. rotundifolia Figure 10,
Map 5.

While plants of some populations in the higher Florida
Keys and along the Florida Gulf coast have only narrow
leaves, plants at most locations in the state are extremely
variable. Crotalaria linaria Small was based on narrow
leaved plants and was maintained by Senn as a variety of
C. maritima. Since a narrow leaved plant usually cannot
be differentiated as being from either the entirely narrow
leaved populations or the heterogeneous populations on the
peninsula, no purpose is served by maintaining the variety
or by reducing it to the status of forma.

190 Rhodora

Figure 10. Crotalaria rotundifolia var. rotundifolia.
A., B. Habit and leaf variation
C. Stem
Crotalaria rotundifolia var. vulgaris
D. Habit and leaf variation
E. Stem

1974] Crotalarias — Windler 191

C. rotundifolia

Map 5. Distribution of Crotalaria rotundifolia, var. rotundifolia.

[Vol. 76

Rhodora

192

C. rotundifolia var vulgaris

Map 6. Distribution of Crotalaria rotundifolia var. vulgaris.

1974] Crotalarias — Windler 193

REPRESENTATIVE SPECIMENS: PUERTO RICO. Vicinity of Dorado,
white sand, Britton, Britton, & Brown 6653 (US). UNITED STATES.
Florida: LEVY CO.: evergreen scrub oak sandridge, Cedar Key,
Godfrey 56571 (GA, GH, USF). MONROE CO.: open dry sandy swale
behind low coastal shell ridge, Middle Cape, Cape Sable, Everglades
National Park, Ward 3939 (GA, LAF, NCU). Georgia: MCINTOSH CO.:
sandhill thicket 1.3 mi. N of Ft. Barrington, Cox Road on ridge,
Boseman 458 (NCU). Mississippi: JACKSON C0.: Ocean Springs
and vicinity of Gulf Coast Research Laboratory, sandy soil under
pines, west end of Deer Island, Channell 414 (usF). South Carolina:
GEORGETOWN CO.: roadsides, 5.6 mi. S of Maryville, Radford 28670
(NCU).

10b. Crotalaria rotundifolia var. vulgaris Windler, Phyto-
logia 21: 264. 1971.

Crotalaria angulata Miller, 1768, sensu Senn, 1939. (See
discussion above.

Figure 10, Map 6.

Type: South Carolina: Hampton County, about three
miles northwest of Yemassee on South Carolina Highway
68. Sandhill. 23 July 1967, D. R. & B, K. Windler 2769
(NCU).

Crotalaria rotundifolia var. vulgaris is distinguished by
its spreading pubescence. Over most of its range var.
vulgaris is also characterized by round to oval leaves, but
in northern Florida and southern Georgia it intergrades
with the usually narrower leaved var. rotundifolia.

REPRESENTATIVE SPECIMENS: COSTA RICA. Cartago, Elev. 4,500
ft., Cooper 165 (us). GUATEMALA. Chimaltenango: alt. 2,100 m.,
Standley 61485 (NY). PANAMA. Panama: roadside, Rio Tapia,
Standley 28214 (us). EL SALVADOR. vicinity of San Salvador,
alt. 650-850 m., Standley 22444 (NY). MEXICO. Baja California:
San Antonio, Cape region, Purpus 475 (US). Guanajuato: 20 mi.
NW of Irapuato, Barkley, Rowel, & Parson 752 (TEx). Hidalgo:
Dist. Zimapan, on road from Hwy. at km. 242 to Encarnacion, Moore
& Wood 4319 (MEX). Jalisco: 5 mi. W of Guadalajara on Hwy. 15,
beneath cliff face in road cut, brown sandy soil, Irwin 1257 (TEX).
Mexico: Lecheria, Pringle 13411 (GH, US). Michoacan: growing in
oak, 2 mi. above Tancitaro, elev. 7,000 ft, Leavenworth 517 (GH).
Morelos: km. 56-57, road from D.F. to Cuernavaca, roadside em-
bankment, Moore 125 (GH). Nayarit: 13 mi. SE of Tepic, in red
clay, Irwin 1293 (TEX). Oaxaca: Valley of Oaxaca, alt. 5,500-7,500

194 Rhodora [Vol. 76

ft., Nelson 1481 (GH). Puebla: vicinity of Puebla, Arsene 1414 (Us).
San Luis Potosi: 15 mi. E of Ciudad del Marz on Rt. 80, ca. 1,000
m., Mickel 544 (MICH). Sinaloa: dry hill, vicinity of Mazatlan,
Rose, Standley, & Russell 13823 (US). Sonora: 3 mi. NE of Matape,
Wiggins & Rollins 400 (GH). Veracruz: Misantla, Purpus 5907
(GH, NY, US). UNITED STATES. Alabama: CLARKE CO.: near
Choctaw Lake, elev. 170 ft., Demaree 50694A (LAF, NCU). Florida:
HERNANDO CO.: Chinsegut Hill Bird Sanctury, open area along
margins of pine woods, Windler 1136 (NCU). Georgia: BEN HILL CO.:
open pine-oak on sandy upland, 7.1 mi. N of Fitzgerald, Duncan &
Hardin 14301 (GA). Louisiana: WASHINGTON PARISH: mixed woods,
edge, along Pushepetappa Creek, 2 mi. WNW of Varnado, Thieret
25910 (LAF). Mississippi: LAMAR C0.: 12 mi. WSW of Hattiesburg
along Black Creek, low area, Doughty 162 (MISS). North Carolina:
BLADEN CO.: wet ditch on 701 to White Lake, Radford & Radford
2065 (NcU). South Carolina: BERKELEY CO.: common on sandy bank
ea. 15 mi. S of Moncks Corner on 52, Wilbur 3547 (GA). Virginia:
NANSEMOND CO.: dry white sand of pineland SW of Marsh Hill
School S of South Quay, Fernald & Long 11050 (GH).

11. Crotalaria bupleurifolia Schlecht. & Cham., Linnaea
5: 575. 1830.

Crotalaria heldiana A. DC. in A. & A.P. DC. Mem. Soc.
Phys. Geneve 9: 97. 1841. Type: Grown from seed of un-
known source in the garden at Carlsruhe, (G!).

Annual or perennial spreading suffrutescent herb with
a taproot to 2 cm. thick. Stems few-many, 5-27 dm. long,
2.3.5 mm. thick, the longest internode 3.5-10 cm. long;
vestiture of spreading or loosely appressed trichomes 0.2-
1.3 mm. long. Stipules present, decurrent for 14, to the
full length of the internode, 0.5-2.7 cm. wide at the top
tapering slightly to or past the subtending node, the stipule
lobes absent or spreading 0.5-3 cm. long. Leaves oval, ellip-
tie, ovate, broadly lanceolate, or lanceolate, 3.2-10.2 cm.
long, 7-46 mm. wide, base obtuse to cuneate, the apex
obtuse, mucronate, acuminate, or acute; vestiture of both
sides of spreading or loosely appressed trichomes 0.2-1.6
mm. long or nearly glabrous; petioles 2-3 mm. long. Inflor-
escences leaf-opposed, peduncles 2-16 cm. long. Bracts
sessile, linear, subulate, or narrowly lanceolate, 4.5-7.5 mm.
long, 0.3-0.6 mm. wide; pedicels 4-5.5 mm. long. Calyces

1974] Crotalarias — Windler 195
13.5-19 mm. long, the tubes 2.5-5 mm. long, bearing
loosely appressed or spreading trichomes 0.1-1.1 mm. long;
bracteoles linear, subulate, or narrowly lanceolate, 3-6 mm.
long, 0.2-0.7 mm. wide. Corollas yellow, the standards
16-25 mm. long, equaling to 6 mm. longer than the upper
calyx lobes; elongate anthers 2-2.8 mm. long, short anthers
0.5-0.7 mm. long; styles 7-11 mm, long. Fruits 3-4.1 cm.
long, breadth not known; seeds tan, green-brown, or brown,
number per pod not known, 2.2-2.5 mm. long. Chromosome
number: »—16. Flowering time: August-March. Habitat:
North-facing slopes and bluffs and cloud forests with firs
and broad leaf trees, elevation 2,500-7,500 feet. Range:
Mexico: Chiapas, Guerrero, Jalisco, Mexico, Michoacan,
Sinaloa, Veracruz.

Type: Hacienda de la Laguna? Schiede et Deppe s.n.
(G!).

Crotalaria bupleurifolia was described by Schlechtendal
and Chamisso in an article describing the collections of
Schiede and Deppe in Mexico. A specimen collected by
Schiede and Deppe at G is marked “Typus” and may well
represent the type even though the location on the label
reals *Mexique. Cuerta grande de Chiconquiaco" rather
than “Hacienda de la Laguna" cited with the original
description of C. bupleurifolia. The number 596 cited with
Schiede's name in Senn (1939) refers to the species number
which accompanied Schlechtendal et Chamisso’s description.

Crotalaria bupleurifolia is characterized by its generally
large size, unusual stipules, and large flowers and habit.
It most closely resembles C. nayaritensis but may be dis-
tinguished from it by the large flower size, larger overall
size, and usually wider stipules.

Two varieties may be recognized:

Stipules present only at the base of peduncles, decurrent
for only a single mnternodes ys a es sete ee
ee es LAM 11a. C. bupleurifolia var. bupleurifolia.

Stipules present at the base of most leaves, frequently
decurrent for more than one internode. ............

11b. C. bupleurifolia var. robusta.

196 Rhodora [Vol. 76

Figure 11. Crotalaria bupleurifolia var. bupleurifolia.
A. Habit and leaf variation
B. Stem

Crotalarias — Windler

197

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10

Map. 7. Distribution of Crotalaria bupleurifolia var. bupleurifolia,

C. bupleurifolia var. robusta, and C. nayaritensis.

198 Rhodora [Vol. 76

lla. Crotalaria bupleurifolia var. bupleurifolia Figure 11.
Map 7.

REPRESENTATIVE SPECIMENS: MEXICO. Chiapas: NE of Bochil,
Miranda 5746 (MEXU). Guerrero: San Antonio Buenos Aires,
Hinton 11708 (GH, TEX, US). Jalisco: near Santa Monica, 1,950-2,050
m, McVaugh 14086 (MEXU, MICH, US). Mexico: Rincon del Carmen,
Dist. Tamascaltepec, 1,340 m, Hinton 1730 (MEXU, NY, US). Michoacan:
NW of Aguililla, 15 Km S of Aserradero Dos Aguas, 1,650-1,700 m,
McVaugh 22767 (MICH). San Luis Potosi: Las Canoas, Pringle 8
(MEXU). Sinaloa: Batel, Concordia, Dehesa 1619 (US). Tamaulipas:
10 Km NW of El Progresso, 1,450 m, Stanford, Retherford, & North-
craft 1009 (GH). Veracruz: Barranca de Tenampa, Zacuapan, Pur-
pus 3663 (GH, NY, US).

llb. Crotalaria bupleurifolia var. robusta (Senn) Windler,
Phytologia 21: 265. 1971.

Crotalaria pilosa var. robusta Senn, 1939, 41: 331. Figure
12, Map 7.

Type: Temascaltepec, Cumbre de Tejupilco, Hinton 2686
(Us!).

REPRESENTATIVE SPECIMENS: MEXICO. Jalisco: 10 Km al N de
La Cuesta, sobre el camino a Talpa, 1,100 m, Rzedowski 15134
(MEXU). Mexico: Plaza de Gallos, 1,200 m. Hinton 4595 (GH, NY).
Sinaloa: Km 1206 on Mexico Hwy. 40, ca. 30 mi, E of Mazatlan-
Guadalajara Junction, 2,700 ft., Windler & Windler 2869 (NCU).

12. Crotalaria nayaritensis Windler, Phytologia 21: 261.
1971.

Annual or perennial spreading herb with a taproot to
1.5 em. thick. Stems 1-many, 6-12 dm. long, 1.5-2.5 mm.
thick, the longest internode 3.8-10 cm. long; vestiture of
short appressed trichomes, 0.2-0.7 mm. long. Stipules
present, decurrent the length of the internode, 0.15-1.1 em.
wide at the top tapering to or past the subtending node,
the stipule lobes parallel with the stem or spreading, 0.1-
1.3 cm. long. Leaves oval, elliptic, ovate, narrowly ovate,
oblong or lanceolate, 3.5-7.8 cm. long, 7-26 mm. wide, base
obtuse to cuneate, the apex obtuse, mucronate, acuminate
or acute; vestiture of both surfaces of appressed or loosely
appressed trichomes 0.3-0.8 mm. long; petioles 1.2-2.5 mm.
long. Inflorescences leaf-opposed, peduncles 3.2-16 cm. long.

1974] Crotalarias — Windler 199

Figure 12. Crotalaria bupleurifolia var. robusta.
A. Habit and leaf variation
B. Stem

200 Rhodora [Vol. 76

Braets sessile, linear or elliptic lanceolate, 3-4.5 mm. long,
0.2-0.5 mm. wide; pedicels 2.8-3.8 mm. long. Calyces 7.5-
12 mm. long, the tubes 2-3.2 mm. long, bearing appressed
trichomes 0.1-0.5 mm. long; bracteoles linear or narrowly
lanceolate, 1.5-3 mm. long, 0.2-0.3 mm. wide, Corollas yel-
low, the standards 7-12.5 mm. long, 2 mm. shorter to 1 mm.
longer than the upper calyx lobes; elongate anthers 1.3-2.1
mm. long, short anthers 0.4-0.5 mm. long; styles 5-6 mm.
long. Fruits 1.3-2.3 em. long, 0.5-0.8 em. broad; seeds tan,
grey-tan, green-brown, brown or red-brown, 30-35 per pod,
1.6-2.4 mm. long. Chromosome number: n=16. Flowering
time: August-February. Habitat: steep moist slopes and
pine woods, elevation 2,500-6,600 feet. Range: Mexico:
Jalisco, Nayarit. Figure 13, Map 7.

Type: Mexico: Nayarit, North of Compostella (near
Km. 24), about 7 miles southwest of Tepic, along road
between Tepic and Compostella. Road-cut through moun-
tain on moist steep slope, Windler & Windler 2902 (NCU).

Crotalaria nayaritensis is a new species named for the
Mexican state from which the holotype was collected. It is
characterized by its spreading or diffuse habit, leaf-opposed
inflorescences, small flower size, and appressed pubescence.
Of the Mexican species it most nearly resembles C. bupleu-
rifolia, but differs from it in having a smaller flower,
appressed pubescence and usually narrower stipules.

REPRESENTATIVE SPECIMENS: MEXICO. Jalisco: 13 mi. SW of
Autlan, 1,000 m, MceVaugh 19886 (MICH); Llano Verde, municipio
de Tecalitlan, 1,600 m, Rzedowski 17417 (MiCH); 3 mi. S of Maza-
mitla, 2,100-2,200 m, MeVaugh 12997 (micH, US); San Sebastian,
W to Mascota, 1,425 m, Mexia 1408 (vs); Tepic, Palmer 1869 (NY,
US). Nayarit: 10 mi. SE of Ahuacatlan, 1,100-1,200 m, Fedema. 287
(MICH) ; N of Compostella, 3,000 ft., Windler & Windler 2902 (NCU);
Mina Esperanza Rosa Morada, Ortega 6682 (US); 2 mi. N of Tepic,
3,000 ft., Windler & Windler 2897 (NCU).

PUTATIVE HYBRIDS BETWEEN CROTALARIA SPECIES

During the course of the study a number of putative
hybrids were observed, The following presentation sets

1974] Crotalarias — Windler 201

Figure 18. Crotalaria nayaritensis.
A. Habit and leaf variation
B. Stem

202 Rhodora [Vol. 76

forth the probable parents and lists of representative speci-
mens which appear to be the results of crosses between
the parents.

Crotalaria sagittalis X C. stipularia
ST. KITTS. Pastures near Sandy Point, Britton & Cowell 128
(NY).

Crotalaria sagittalis X C. rotundifolia var. vulgaris
UNITED STATES. Alabama: Mobile, Mohr s.n. 1873 (SIU).
Florida: BAY CO., E of Panama City, Windler & Windler 2723 (NCU);
N of Betts, Windler & Windler 2721 (NCU); ESCAMBIA CO., W of
Century, Windler & Windler 2688 (NCU); JACKSON CO., Grand Ridge,
Windler & Windler 2712 (NCU); OKALOOSA CO. 4 mi. W of Black-
water River Bridge on Fla. Hwy. 4, Windler & Windler 2700 (NCU);
SANTA ROSA CO., E of Chumuckla, Windler & Windler 2693 (NCU);
S of Milton, Windler & Windler 2697 (NcU). Mississippi: FOREST
co., S of Hattiesburg, Windler & Windler 2659 (NCU); HARRISON CO.,
N of Biloxi, Windler & Windler 2666 (NCU); LAMAR CO., 8.5 mi. SE
of Hattiesburg, Cooley 62 (Miss); 7.5 mi. SW of Hattiesburg, Cooper
93 (MISS); Pearl River, 1 mi. N of Picayune, Pullen 641208 (MIss).

Crotalaria sagittalis X C. quercetorum
MEXICO. Chiapas: Bochil, Windler & Windler 2994 (NCU).

Crotalaria rotundifolia X C. purshii

UNITED STATES. Alabama: BALDWIN CO. Foley, Windler &
Windler 2681 (NCU). Florida: CALHOUN CO., 12 mi. S of Clarksville,
Windler & Windler 2425 (NCU); ESCAMBIA CO., 235 miles N of Jet.
Fla. Hwy. 97 — US Hwy. 29, Windler & Windler 2686 (NCU); SANTA
ROSA CO, S of Milton, Windler & Windler 2699 (NCU). Georgia:
GLYNN CO., Brythe Island, Hardin & Duncan 13785 (NCU); MCINTOSH
CO., 2.6 mi. W of Cox, Bozeman 1078 (NCU); 1 mi. SE of Ft. Bar-
rington, Boseman & Radford 2146 (NCU); THOMAS CO, SW of
Thomasville, Windler & Windler 2734 (NCU). Mississippi: HARRISON
CO.: Ship Island, Demaree 35028 (LAF); JACKSON CO., Round Island,
Demaree 33742 (LAF).

ACKNOWLEDGMENTS

I am especially grateful to my wife, Bonnie Kay, whose
moral support and technical and clerical assistance speeded
the completion of this study.

Dr. C. Ritchie Bell, who served as research adviser during
the study, deserves special thanks for his helpful sugges-

1974] Crotalarias — Windler 203

tions, criticisms, and editorial work on the manuscript.
Drs. Albert E. Radford and Clifford R. Parks are also
thanked for their helpful suggestions and cooperation
during the course of the study and the preparation of the
thesis. Dr. Velva Rudd, Smithsonian Institution, is thanked
for her critical review of the Systematic Treatment manu-
script.

I would like to acknowledge my fellow students at the
University of North Carolina at Chapel Hill and thank
them for their aid, counsel, and criticisms during my
years of study at the University.

During the extensive collecting trips, help was needed
on numerous occasions and was always cheerfully supplied
by colleagues at other institutions. Special mention should
be made of Dr. Arturo Gomez-Pampa, Universidad Nacional
de México; Dr. Sidney McDaniels, Florida State University ;
Dr. John Thieret, Southwestern Louisiana State University ;
and Dr. D. B. Ward, University of Florida, and their
respective staffs that provided plant dryers, and green-
house and storage space at various times during collection
trips. Dr. Andre Clewell and the staff of the Tall Timber
Research Station in Tallahassee, Florida, are also due thanks
for assistance and lodging that they provided while collec-
tions were being made in the vicinity.

Special thanks are extended to the curators of the
numerous herbaria (A, ALU, AUA, BM, G, GA, GH, K, LAF,
MEXU, MICH, MISS, NCU, NY, P, SIU, TEX, UC, US, USF, WIS)
from which specimens were borrowed and to Dr. Norman
K. B. Robson, in particular, whose efforts in my behalf at
the British Museum (Natural History) in checking some
of the early American collections proved invaluable.

LITERATURE CITED

Baker, E. 1914. The African species of Crotalaria. Journal Lin-
nean Society, Botany 42: 241-425.

BRITTEN, J. & E. BAKER. 1897. Houstoun’s Central American Legu-
minosae. Journal of Botany 35: 225-234.

FERNALD, M. L. & B. SCHUBERT. 1948. Studies of American types
in British Herbaria, IV. Some species of Thomas Walter.
Rhodora 50: 190-208.

204 Rhodora [Vol. 76

GRAUSTEIN, J. 1954. The date of Pursh’s “Flora Americae Septen-
trionalis." Rhodora 56: 275.

LANJOUW, J. & F. STAFLEU. 1964. Index Herbariorum, Regnum
Vegetabile, Vol. 31. 251 pp. Utrecht.

POLHILL, R. 1968. Miscellaneous notes on African species of Cro-
talaria L., II. Kew Bull. 22: 169-348.

SAVAGE, S. 1945. A Catalogue of the Linnaean Herbarium. 225
pages. London.

SENN, H. 1939. The North American species of Crotalaria. Rhodora
41: 317-366.

SPRAGUE, T. 1939. Analysis of binary combinations published under
Anonymos by Walter, Flora Caroliniana (1788). Kew Bull. 1939:
331-334.

Warp, D. 1962. The genus Anonymos and its nomenclatural sur-
vivors. Rhodora 64: 87-92.

WINDLER, D. R. 1970. Systematic studies in Crotalaria sagittalis
and related species. Unpub. diss, Univ. of North Carolina at
Chapel Hill. 258 pages.

1971. New North American unifoliolate Crotalaria

taxa. Phytologia 21: 257-266.

1973. Field and garden studies in Crotalaria

sagittalis and related species. Phytologia 26: 289-354.

TOWSON STATE COLLEGE
BALTIMORE, MARYLAND 21204

THE FLORA OF MOUNT MONADNOCK,
NEW HAMPSHIRE

HENRY I. BALDWIN

Mt. Monadnock (Elevation 3,165 feet) lies in Southern
New Hampshire at a distance from other hills of compara-
ble height. This mass of granite and gneiss rising from a
plain has given rise to the generic term “monadnock” for
similar isolated rock peaks. The Grand Monadnock, as it is
sometimes called, to distinguish it from other monadnocks,
is situated in the towns of Dublin and Jaffrey. Roads sur-
round the base of the mountain at elevations of 1,200 to
1,500 feet above sea level.

Botanically Mt. Monadnock is of interest because of the
presence of a few arctic-alpine plants on the bare rocky
summit area, far from other stations for them, and because
of the records made by Thoreau on his four visits from
1844 to 1860, and later by other botanists, such as Stone
(1885) and Deane (1890) and many later ones, some of
whom deposited specimens in New England herbaria. It
was my purpose to compare the present flora with that
found by Thoreau.

The student of land use can read much of its history from
the flora, since plant habitat is affected by what has hap-
pened to the plants. The history of Mt. Monadnock is of
interest because of three distinct periods: (1) the develop-
ment of complete forest cover since the time of glacial re-
treat, (2) its complete removal on the upper slopes by fire,
and partial removal on the lower by clearing for pasture
and fire, and (3) the slow renewal of vegetative cover dur-
ing the last 150 years.

It is generally believed on the basis of geological and
pollen evidence and observation of succession on rock and
soil recently exposed by the retreat of present day glaciers,
that the succession on Mt. Monadnock was probably as fol-
lows: the first pioneer plants, following lichens and mosses,
may have been shrubs and forbs capable of surviving under

205

206 Rhodora [Vol. 76

severe cold, wind and fluctuating moisture. These may have
included species that colonize mineral soil and rock crevices,
including some of the arctic-alpines. Later, as climate mod-
erated, spruces and pines (possibly Pinus Banksiana) oc-
cupied the accumulations of sand and organic matter in
depressions. Finally, as humus accumulated, Red Spruce
(Picea rubens) came to dominate the flora of the upper part
of the mountain where soil development was indefinitely
delayed.

Mt. Monadnock was completely forested at the coming of
the White Man, the summit area covered with Red Spruce
to the very summit, and the lower slopes with spruce and
hardwood, much as the case of lesser mountains much
farther north. All similar rocky peaks below the approxi-
mately 4,500 feet climatic timberline in New Hampshire
are or were completely forest-clad. Mt. Monadnock is about
1,000 feet lower than climatic timberline and lies almost
100 miles south of the nearest timberline. Tree line is
higher at lower latitudes. The bare rocky summits of New
Hampshire such as Chocorua, Cardigan, Firescrew, and
Percy Peaks all have documented history of fire.

Until recent years there were many old stumps and logs
near the summit. Thoreau wrote in his Journal (June
1858) : “The whole rocky part of the mountain, except the
extreme summit, is strewn with the relics of spruce trees a
dozen or fifteen feet long, and long since dead and bleached,
so that there is plenty of fuel at hand." The remains near-
est the summit were no doubt first consumed by visitors
campfires. Visitors were very numerous in Thoreau’s day
and before, and in several of his writings he reports having
seen fires at the top. Since the area at the extreme summit
is quite level, with depressions, it is probable that it was
originally as densely forested as the rest of the mountain.
Chamberlain (1936) shows a picture (p. 73) of stumps near
the top. Obviously, there could not have been such severe
fires at the summit had there not been timber to burn. The
1953 fire on the east slope stopped when it reached the
scrub zone where there was no fuel.

1974] Flora of Mount Monadnock — Baldwin 207

There are many published reports that confirm the orig-
inal forested nature of the summit. Later the practice of
clearing the slopes for pasture gave rise to numerous fires.

During the celebration of the centennial of the Town of
Dublin in 1852, Charles Mason made an address on June 17
(Leonard 1855) including the following remark:

“Formerly covered with trees almost to the summit.”

At the same celebration in Dublin (Leonard op. cit.) a
letter was read from Daniel Elliott, M.D. in which he re-
called: “I remember (Monadnock) when clothed with ver-
dant foliage to the very summit.” Similar records are
found in the Town of Jaffrey: “When the town was first
settled the mountain was covered to its summit with forest
trees, principally spruce, excepting a small peak southeast
of the top, which was called the “Bald Rocks” (Cutter
1881). “Belts of heavy spruce timber on the highest por-
tions” were reported by Child (1885). Similar references
may be found in Dana (1816) and Ellis (1880).

Ecologists recognized that forest had come to occupy the
mountain much as peaks farther north. Braun (1950)
refers to Monadnock (p. 423) as “covered with Red Spruce
from 2,300 feet upward, thus producing a small outlier of
the prevailing mountain vegetation in the north.” Spurr
(1964) states that “the summit of Mt. Monadnock ... was
clothed with a Red Spruce forest until destroyed by fire...”

Many fires have occurred during the recorded history of
Mt. Monadnock, and it is possible that lightning fires may
have burned in earlier times, but it is doubtful if Indians
caused any fires on its forbidding slopes. In the same ad-
dress referred to above (Leonard, loc. cit.) Charles Mason
said: “A great fire spread over a great part of the superior
portion of the mountain killing every tree and shrub. Dead
trees falling, furnished materials for another conflagration.
Some 30 years ago in the latter part of a dry summer, a fire
in a clearing made its way up to the higher regions." “Some
30 years ago” would have placed this fire about 1820, al-
though the cause of the fire differs from other accounts
given below. Also the statement of Dr. Elliott on the same

208 Rhodora [Vol. 76

occasion: “I saw, year after year, the devouring flames
climbing his lofty sides", suggests many fires.

Thoreau, on his second visit to Monadnock in 1852 (his
first was in 1844, the journal report of which has not been
preserved) related this incident during his passage through
Peterborough: *A man in Peterborough told me that his
father told him that Monadnock used to be covered with
forest; that fires ran through it and killed the turf; then
the trees were blown down, and their roots turned up and
formed a dense and impenetrable thicket in which wolves
abounded . . . till finally they set fire to this thicket and it
made the greatest fire they had ever had in the county.”

These and other records fail to give exact dates. As
Chamberlain (1936) says (p. 97): “No documentary evi-
dence has been found to show exactly when the fires oc-
curred.” He also quotes (p. 98) from the New Hampshire
Sentinel of September 30, 1825, reporting a great fire of
“some years ago”. “This fire consumed only the turf. The
trees turned over in every direction, and afforded about
three years later fuel for a more tremendous scene.” This
places the two “great” fires closer together. The best evi-
dence suggests a first fire about 1800, then the 1815 hurri-
cane, and the final great fire in 1820, There were later ones,
too. Thoreau reported (“A Walk to Wachusett”, July 19,
1842) “A fire blazing on Monadnock that night, which
lighted up the whole western horizon.”

The first inroads of man occurred on the lower slopes that
were cleared for pasture. Later, clearing progressed higher
where sheep were grazed. Fire was used both for opening
the areas and for keeping them so, and many of the fires
that reached the upper levels both before and after the
1820 fire were caused by fire escaping from pasture burn-
ing. The repeated burning promoted the growth of blue-
berries, which I believe may have spread from lower
pastures to the summit after the great fires. At present
these are rather scarce.

Following abandonment of the pastures, these seeded into
Red Spruce, and the process of reclothing the upper reaches

1974] Flora of Mount Monadnock — Baldwin 209

of the mountain progressed rapidly at first, and ever slower
on the more exposed parts of the mountain. Here denuda-
tion resulted in a worsening of the climate. Wind movement
increased over the bare rock, and extremes of temperature
and soil moisture retarded growth. Recently a cycle of
retarded spread of spruce seems to have set in, since almost
no spruce trees less than forty years old can now be found
on the upper part of the mountain and a high percentage
of the trees display dead tops and reduction in diameter
and height growth.

The large area laid bare by the fires apparently provided
a favorable habitat for a few arctic-alpine plants that may
have been present before the fires or had been transported
from other sources. In any case they were well-established
at the time of Thoreau's first visit, little over twenty years
after the great fire. Those recorded by Thoreau (Journal,
August 8, 1860, 25-28; 1670-1671) were: Arenaria
groenlandica, Vaccinium Vitis-Idaea var. minus and Juncus
trifidus. Others with northern or high altitude affinities,
not found generally in the lowland, especially in the vicinity
of Mt. Monadnock were Potentilla tridentata and Smilacina
trifolia. During the present survey Lycopodium Selago var.
Selago and var. appressum as well as L. annotinum var.
pungens have been added to the list.

Thoreau limited his observations to the "upper part of
the mountain," and essentially all the taxa reported by him
in 1858 “within 200 feet of the summit" were also collected
in 1968. Less complete were the lists of Stone (1885),
Deane (1890) and Emerson (1869), probably due to the
restricted area that they examined. While almost all the
taxa recorded earlier are present now (at least those easily
discoverable), and a substantial number of new ones have
been added, there are doubtless more to be found. There
have been changes in distribution and abundance of the
plants recorded by the early botanists. The encroachment
of Red Spruce and more especially shrubs such as Nemo-
panthus mucronata, Rhododendron canadense and Kalmia
angustifolia appears to have reduced the habitat available

210 Rhodora [Vol. 76

to the arctic-alpines. Vaccinium Vitis-Idaea can persist a
long time under these shrubs, but fruiting is diminished.
Some thirty years ago the proprietors of the Half Way
House used to collect enough mountain cranberries on the
mountain to supply hotel guests, while now only small
quantities can be picked. This plant has almost disappeared
from the southwest side where the hotel supply was largely
obtained. The alpine flora seems to be increasingly re-
stricted to the northern side of the peak where a cooler
climate prevails. Further, observations indicate that there
may be a successional change going on in the mats of Vac-
cinium Vitis-Idaea, that are invaded more and more by
Potentilla tridentata. Another major change from 100
years ago is the vast reduction in Cladonia lichens. Thoreau
remarks frequently how dense they were, crunching under
foot, and dry so that fire would spread through them.
Cladonia persists around the margins of shrubby hollows,
but today does not cover extensive areas. Other lichens
mentioned by Thoreau on the summit are now mostly con-
fined to moist ravines.

Originally it was planned to limit collections to the area
studied by Thoreau, “the upper part of the mountain". Be-
cause of the difficulty of determining just what this em-
braced, the area was extended to include land above 2,000
feet above sea level. (The Half Way House site is at 2,100
feet) Present collections have been supplemented by recent
herbarium records kindly supplied by Mr. and Mrs. David
E. Boufford of Keene State College. Where several collec-
tions of the same taxon were found, only the earliest record
is given, in order to save space.* All records that stated
Mt. Monadnock as the place of collection were included,
although often the specific location was not given. Genera
and species are listed in the order of Gray's Manual, 8th
Edition (Fernald 1950). The following symbols are used:

(*) asterisk: reported by Thoreau 1858-1860.

(—) dash: collected 1968-1972.

*Complete lists have been deposited in several New England her-
baria.

1974] Flora of Mount Monadnock — Baldwin 211

(+) dagger: collected 1968-1972, not previously reported
(as far as known)

Most of the specimens collected in 1968-1972 were de-
posited in the University of New Hampshire and Franklin
Pierce College herbaria. Abbreviations for all Herbaria
consulted in this study are as follows: HNH Dartmouth
College, FPC Franklin Pierce College, KSC Keene State Col-
lege, NEBC New England Botanical Club, MASS University
of Massachusetts, and NHA University of New Hampshire.

LIST OF THE VASCULAR FLORA OF MOUNT MONADNOCK

-Lycopodium Selago L. var. Selago L. W. Deane, Aug.,
1891 (NEBC).
+_L. Selago L. var. appressum Desv. H. I. Baldwin, July.
1968 (FPC).
* L.lucidulum Michx. H. I. Baldwin, Aug., 1968 (FPC,
NHA).
+—L. inundatum L. H. I. Baldwin, Oct., 1968 (FPC).
* L.annotinum L. H. D. Thoreau, Aug., 1869 (?) (NEBC).
+_L. annotinum L. var. pungens (La Pylaie) Desv. H. I.
Baldwin, June, 1971 (FPC, NHA).
*_L. clavatum L. T. L. Steiger, Oct., 1935 (NHA).
L. clavatum L. var. megastachyon Fern. & Bissell, C.
Jenks, Aug. 1883 (NEBC).
_L. obscurum L. var. obscurum. A. W. Driggs. Aug., 1916
(NEBC).
L. obscurum L. var. dendroideum (Michx.) D. C. Eat.
B. L. Robinson, Aug., 1869 (NEBC).
*_L. complanatum L. A. W. Driggs, Aug., 1916 (NEBC).
Botrychium matricariaefolium A. Br. E. L. Rand, May,
1897 (NEBC).
i-Osmunda cinnamomea L. H. I. Baldwin, Aug., 1968 (FPC).
Woodsia ilvensis (L.) R. Br. B. L. Robinson, Aug., 1898
(NEBC).
—Thelypteris Phegopteris (L.) Slosson. E. F. Williams,
June, 1894 (NEBC).
—Dryopteris spinulosa (O. F. Muell.) Watt. var. americana
(Fisch.) Fernald. E. F. Williams, June, 1894 (NEBC).

212 Rhodora [Vol. 76

—D. marginalis (L.) Gray. D. S. Boufford, June, 1971 (KSC).
+—Polystichum acrostichoides (Michx.) Schott. H. I. Bald-
win, Aug., 1971 (FPC, NHA).
*—Dennstaedtia punctilobula (Michx.) Moore. H. I. Baldwin,
Oct., 1968 (FPC).

-Pteridium aquilinum ( L.) Kuhn var. latiusculum (Desv.)
Underwood. H. I. Baldwin, Aug., 1968 (FPC).

*—Polypodium virginianum L. W. Deane, Aug., 1889 (HNH).

"—Abies balsamea (L.) Mill, J. R. Churchill, May, 1895
(NEBC).

*—Tsuga canadensis (L.) Carr. H. I. Baldwin, July, 1968
(FPC).

*—Picea rubens Sarg. E. F. Williams, May, 1898 (HNH).

T-Larix laricina (DuRoi) K. Koch. H. I. Baldwin, July,
1968 (FPC). (one tree at 2,900 feet on west side, two
in Thoreau Bog).

*-Pinus Strobus L. A. R. Hodgdon et al., Aug., 1956 (NHA).

Poa compressa L. W. Deane, July, 1884 (HNH).
P. pratensis L. E. F. Williams, June, 1894 (NEBC).
P. palustris L. J. R. Churchill, July, 1890 (MASS, NEBC).

*—_Festuca ovina L. E. L. Dickinson and F. C. Seymour,

July, 1930 (Mass).
T-F. rubra L. H. I. Baldwin, July, 1968 (FPC).
*_Deschampsia flexuosa (L.) Trin. H. D. Thoreau, Aug.,
1860 (NEBC).
*_Danthonia spicata (L.) Beauv. H. D. Thoreau, Aug., 1860
(NEBC).
—D. compressa Aust. A. S. Goodale, July, 1927 (MASS).
*Calamagrostis canadensis ( Michx.) Nutt. H. D. Thoreau,
Aug., 1860 (NEBC).

*—Agrostis scabra Willd. H. D. Thoreau, Aug., 1860 (NEBC).
-Scirpus atrovirens Willd. C. W. J enks, Aug., 1883 (NEBC).
—S. atrocinctus Fern. A. S. Goodale, July, 1927 (Mass).

*—Eriophorum spissum Fern., H. D. Thoreau, Aug., 1860

(?) (NEBC).
*E. gracile W. D. J. Koch. A. S. Goodale, July, 1927 (MASS).
-E. tenellum Nutt. R. M. Harper, July, 1899 (NEBC).
“3,150 feet”.

1974] Flora of Mount Monadnock — Baldwin 213

E. viridi-carinatum (Engelm.) Fern. C. F. Batchelder,
Sept., 1909 (NHA).
—E. virginicum L. C. W. Jenks and C. W. Swan, Aug.,
1883 (NEBC, NHA).
Rhynchospora capitellata (Michx.) Vahl. B. L. Robinson,
July, 1897 (NEBC).
+—R. alba (L.) Vahl. H. I. Baldwin, Oct., 1968 (FPC).
+—Carex stipata Muhl. H. I. Baldwin, June, 1969 (FPC).
*C. trisperma Dew. H. D. Thoreau, Aug., 1860 (NEBC).
“NE swamp”.
+-C. trisperma Dew. var. Billingsii Knight, H. I. Baldwin,
Oct., 1968 (FPC).
* C. canescens L. “E. F. Jr." (?), 1838 (MASS).
C. canescens L. var. subloliacea Laested. Sydney Harris,
June, 1894 (NEBC).
C. brunnescens (Pers.) Poir. W. Deane, July, 1889 (MASS).
C. brunnescens (Pers.) Poir. var. sphaerostachya (Tuk-
erm.) Kükenth. A. W. Driggs, Aug., 1916 (NEBC).
C. cumulata (Bailey) Mackenz. C. W. Jenks, Aug., 1885
(NEBC).
—C. communis Bailey. L. Russell, 1898 (NHA).
C. novae-angliae Schwein. W. Deane, July, 1891 (HNH,
MASS). “Mt. road below house".
C. deflexa Hornem. J. R. Churchill, May, 1895 (NEBC).
C. umbellata Schkuhr. J. R. Churchill, May, 1896 (NEBC).
C. gracillima Schwein. R. J. Eaton, July, 1957 (NHA).
C.aestivalis M. A. Curtis. J. R. Churchill, July, 1890
(MASS, NEBC).
C. leptonervia Fern. E. F. Williams, June, 1894 (NEBC).
C. intumescens Rudge. E. F. Williams, June, 1894 (NEBC).
+-C. nigra L. H. I. Baldwin. July, 1971 (FPC, NHA).
i-Arisaema triphyllum (L.) Schott. D. W. Allen, May, 1968
(FPC).
-Juncus bufonius L. A. W. Driggs, Aug., 1916 (NEBC).
*_J. trifidus L. C. W. Swan, Aug., 1883 (NEBC).
J.Greenei Oakes and Tuckerm. B. L. Robinson, Aug.,
1896 (NEBC).

214 Rhodora [Vol. 76

-J. brevicaudatus (Engelm.) Fern. A. W. Driggs, Aug.,

1916 (NEBC).
J. articulatus L. C. A. and A. W. Cheever, Sept., 1910

( NEBC).

T-Luzula, multiflora (Retz.) Lejeune. D. W. Allen, May,
1968 (FPC, NHA).

*—Uvularia sessilifolia L. D. W. Allen, May, 1968 (FPC).

*—Clintonia borealis (Ait.) Raf. T. L. Steiger, May, 1937
(NHA).

r-Smilacina racemosa (L.) Desf. H. I. Baldwin, Sept., 1970
(FPC).

*—S. trifolia (L.) Desf. Sydney Harris, June, 1894 (NEBC).

r-Maianthemwm canadense Desf. D. W. Allen, May, 1968
(FPC).

—Streptopus roseus Michx. var. perspectus Fassett. W.
Deane, July, 1889 (NEBC).

*-Polygonatum pubescens (Willd.) Pursh. H. I. Baldwin,
May, 1969 (FPC, NHA).

-Medeola virginiana L. D. W. Allen, May, 1968 (FPC).

-Trillium erectum L. J. R. Churchill, May, 1883 (Mass).

*—T. undulatum Willd. T. L. Steiger, May, 1937 (NHA).

r-Sisyrinchium montanum Greene. H. I. Baldwin, July,

1968 (NHA).
*-Cypripedium acaule Ait. D. W. Allen, May, 1968 (FPC,
NHA).
Habenaria Hookeri Torr. Henry A. Purdie, June, 1905
(NEBC).

Spiranthes lacera Raf. Hodgdon and Ken Welsh, July,
1957 (NHA). “2,500 feet”.
Malaxis uniflora Michx. F. Hunt, Aug., 1946 (NHA).
*Salix Bebbiana Sarg. A. R. Hodgdon, June, 1933 (NHA).
*—S. discolor Muhl. D. W. Allen, May, 1968 (FPC).
*—Populus tremuloides Michx. H. I. Baldwin, Aug., 1968
(FPC).
*_Betula alleghaniensis Britton. Hodgdon and Welsh, July,
1957 (NHA).
-B. populifolia Marsh. Hodgdon et al., Aug., 1956 (NHA).
*—B. papyrifera Marsh. Hattie Merrifield, July, 1879 (NEBC).

1974] Flora of Mount Monadnock — Baldwin 215

—B. papyrifera Marsh. var. cordifolia (Regel) Fern. Hodg-
don and Welsh, July, 1957 (NHA).
B. papyrifera Marsh. X B. populifolia Marsh. Hodgdon
et al., Aug., 1956 (NHA). “2,800 feet".
*_Ostrya virginiana (Mill. K. Koch. H. I. Baldwin, Aug.,
1971 (FPC).
—Fagus grandifolia Ehrh. Hodgdon et al., Aug., 1956 (NHA,
NEBC).
i-Quercus rubra L. D. W. Allen, May, 1968 (FPC).
Polygonum aviculare L. var. angustissimum Meisn. B. L.
Robinson, July, 1897 (NEBC).
* P. cilinode Michx. W. Deane, Aug., 1889 (HNH).
+_P. scandens L. H. I. Baldwin, Aug., 1972 (FPC, NHA).
“one plant, Red Spot Trail”.
+_Claytonia caroliniana Michx. H. I. Baldwin, June, 1971
(FPC).
*_Arenaria groenlandica Du Roi. Thomas Hope, Aug., 1879
(NEBC).
Ranunculus abortivus L. Rand and Robinson, June, 1898
(NEBC).
+—R. acris L. H. I. Baldwin, July, 1971 (NHA).
+_R. bulbosus L. D. W. Allen, June, 1968 (NHA).
i-Thalictrum polygamum Muhl. H. I. Baldwin, May, 1969
(NHA).
*_Coptis groenlandica (Oeder) Fern. D. W. Allen, May,
1968 (FPC).
3—Actaea pachypoda Ell. H. I. Baldwin, May, 1969 (FPC).
*_Aquilegia canadensis L. Rand and Robinson, May, 1897

(NEBC).

-Berberis Thunbergii DC. W. H. Manning, Oct., 1913
(NEBC).

_Drosera intermedia Hayne. Fred M. Hunt, Aug., 1946
(NHA).

+_D. rotundifolia L. H. I. Baldwin, July, 1968 (FPC, NHA).
+-Chrysosplenium americanum Schwein. D. E. Boufford,
1971 (KSC).
*Ribes cynosbati L. T. Otis Fuller, June, 1883 (NEBC).
* R. glandulosum Grauer. J. R. Churchill, May, 1883 (NHA).

216 Rhodora [Vol. 76

R. triste Pall. T. L. Steiger, May, 1937 (NHA).
*_Spiraea latifolia (Ait.) Borkh. A. W. Driggs, Aug., 1916
(NEBC). "above timberline".
1-8. tomentosa L. H. I. Baldwin, July, 1968 (FPC).
*—Pyrus melanocarpa (Michx.) Willd. C. F. Batchelder.
Sept., 1909 (NHA).
*—P. americana (Marsh.) DC. J. R. Churchill, July, 1890
(HNH).
P. decora, (Sarg.) Hyland. Hodgdon and Kenneth Welsh,
July, 1957 (NEBC). ‘2,500 feet".
P. americana L. X P. decora (Sarg.) Hyland. Hodgdon
and Kenneth Welsh, July, 1957 (NEBC). “2,500 feet".
*—Amelanchier laevis Wieg. Rand and Robinson, May, 1897
(NEBC).
*_A. Bartramiana (Tausch.) Roemer. J. R. Churchill, May,
1885 (NEBC).
A. Bartramiana (Tausch.) Roemer. X A. neglecta (?)
Rand and Robinson, May, 1897 (NEBC).
T-Crataegus macrosperma Ashe. D. W. Allen, May, 1968
(NHA).
T-Fragaria virginiana Duchesne. D. W. Allen, May, 1968
(FPC).
Potentilla fruticosa L. J. A. Bates, Aug., 1900 (NEBC).
*—P. tridentata Ait. E. S. Hoar, July, 1857 (NEBC).
P. norwegica L. A. W. Driggs, Aug., 1916 (NEBC). “sum-
mit".
T—P. canadensis L. H. I. Baldwin, June, 1969 (FPC, NHA).
T—P. simplex Michx. H. I. Baldwin, May, 1970 (FPC).
Geum aleppicum Jacq. var. strictum (Ait.) Fern. Henry
Purdie, July, 1905 (HNH).
Rubus hispidus L. J. P. Poole, July, 1939 (HNH, NHA).
R. setosus Bigel. A. W. Driggs, Aug., 1916 (NEBC).
*_R. vermontanus Blanch. B. L. Robinson, July, 1897 (NEBC).
R. allegheniensis Porter. A. W. Driggs, Aug., 1916 (NEBC).
T—R. frondosus Bigel. H. I. Baldwin, July, 1968 (FPC).
*—Prunus pensylvanica L. f. C. W. Jenks, Aug., 1883 (NEBC).
T—P. serotina Ehrh. H. I. Baldwin, July, 1968 (FPC).
t-Trifolium repens L. D. W. Allen, June, 1968 (FPC).

1974] Flora of Mount Monadnock — Baldwin 217

_Oxalis montana Raf. T. Otis Fuller, June, 1883 (NHA).
Polygala sanguinea L. R. H. Piper, July, 1927 (MASS).
* Nemopanthus mucronata (L.) Trel. Rand and. Robinson,
May, 1897 (NEBC).
* Acer spicatum Lam. Sydney Harris, June, 1894 (NEBC).
* A. pensylvanicum L. C. W. Jenks, Aug., 1883 (NEBC).
1—A. saccharum Marsh. H. I. Baldwin, June, 1968 (FPC).
* A. rubrum L. H. I. Baldwin, June, 1968 (FPC, NHA).
Hypericum perforatum L. J. P. Poole, July, 1939 (HNH).
_H. canadense L. Mary E. Gilbreth, Aug., 1889 (NEBC).
(one extremely small plant found in 1971, H. I. Bald-
win)
+—Viola papilionacea Pursh. H. I. Baldwin, May, 1969 (NHA).
V. sororia Willd. E. F. Williams, May, 1898 (NEBC).
V. fimbriatula Sm. E. F. Williams, May, 1898 (NEBC).
—V. pallens (Banks) Brainerd. A. W. Cheever, Sept., 1910
(HNH).
V. blanda Willd. J. R. Churchill, July, 1890 (HNH, NEBC).
V. lanceolata L. A. R. Hodgdon, June, 1933 (NHA).

1-V. rotundifolia Michx. H. I. Baldwin, April, 1969 (NHA).
V. adunca Sm. Rand and. Robinson, May, 1897 (NEBC).
1—V. conspersa, Reichenb. H. I. Baldwin, May, 1971 (FPC).

Circaea alpina L. Edward S. Hoar, 1879 (NEBC).
* Epilobium angustifolium L. H. I. Baldwin, Aug., 1971
(FPC).
E. leptophyllum Raf. C. W. Jenks, Aug., 1883 (NEBC).
Aralia racemosa L. Hattie Merrifield, July, 1879 (NEBC).
* A4. hispida Vent. J. P. Poole, July, 1939 (HNH, NHA).
_A. nudicaulis L. A. S. Goodale, May, 1926 (MASS).
1—Panaz trifolius L. D. W. Allen, May, 1968 (FPC, NHA).
* Cornus canadensis L. C. W. Jenks, Aug., 1883 (NEBC).
—C. alternifolia L. f. Hodgdon et al., Aug., 1956 (NHA).
Chimaphila umbellata (L.) Bart. var. cisatlantica Blake.
J. P. Poole, July, 1939, (HNH, NHA).
+-Monotropa uniflora L. H. I. Baldwin, Sept., 1968 (FPC).
(rare).
* Ledum groenlandicum Oeder. Sydney Harris, June, 1894
(NEBC).

218 Rhodora [Vol. 76

*—Rhododendron canadense (L.) Torr. Sydney Harris, June,
1883 (NEBC).
*—-Kalmia angustifolia L. J. P. Poole, July, 1939 (HNH).
*-Cassandra calyculata (L.) D. Don. D. W. Allen, May,
1968 (FPC).
*—Epigaea repens L. H. I. Baldwin, April, 1969 (FPC).
“(one plant only, on cliff) ."
*Gaultheria hispidula (L.) Bigel. C. G. Trow, 1871 (MASS).
1T-Calluna vulgaris (L.) Hull. H. I. Baldwin, May, 1965
(NHA). (one isolated colony (introd)).
-Gaylussacia baccata (Wand.) K. Koch. A. W. Driggs,
Aug., 1916 (NEBC).
Vaccinium myrtilloides Michx. Hodgdon et al., Aug., 1956
(NHA). “2,800 feet",
*_V. angustifolium Ait. W. Deane, Aug., 1889 (HNH).
-V. angustifolium Ait. var. laevifolium House. B. L. Rob-
inson, July, 1897 (NEBC).
T-V. augustifolium Ait, X V. corymbosum L. H. I. Baldwin,
Aug., 1969 (NHA).
V. angustifolium Ait. var. nigrum ( Wood) Dole. Fred M.
Hunt, Aug., 1946 (NHA).
V. corymbosum L. A. S. Goodale, May, 1926 (Mass).
*-V. Vitis-Idaea L. var. minus Lodd. C. G. Trow, Aug., 1864
(MASS).
T—Asclepias syriaca L. H. I. Baldwin, July, 1971 (FPC).
*—Trientalis borealis Raf. D. W. Allen, May, 1968 (FPC).
Lysimachia terrestris (L.) BSP. J. P. Poole, July, 1939
(HNH, NHA).
T-Fraxinus americana L. H. I. Baldwin, July, 1968 (FPC).
Hedeoma pulegioides (L.) Pers. B. L. Robinson, Aug.,
1896 (NEBC).
f-Veronica longifolia L. Hodgdon and Hehre, Aug., 1969
(NHA).
1—-V. officinalis L. H. I. Baldwin, June, 1969 (FPC, NHA).
Gerardia tenuifolia Vahl. B. L. Robinson, Aug., 1898
(NEBC).
*“_Melampyrum lineare Desr. var. americanum (Michx.)
Beauv. D. W. Allen, June, 1968 (FPC).

1974] Flora of Mount Monadnock — Baldwin 219

1-Mitchella repens L. H. I. Baldwin, Aug., 1968 (FPC, NHA).
rare.
tT-Houstonia caerulea L. D. W. Allen, May, 1968 (FPC, NHA).
*_Diervilla Lonicera Mill. C. W. Jenks, Aug., 1883 (NEBC).
*_Viburnum alnifolium Marsh. C. W. Jenks, Aug., 1883
(NEBC).
*_V. cassinoides L. C. L. Canfield, June, 1931 (NHA).
*_Sambucus pubens Michx. W. Deane, July, 1891 (NEBC).
T-Solidago caesia L. H. I. Baldwin, Sept., 1970 (FPC).
*_S. macrophylla Pursh. W. Deane, Aug. 1891 (HNH, NEHA,
MASS).
—S. puberula Nutt. Mary E. Gilbreth, Aug., 1889 (NEBC).
—S. Randii (Porter) Britt. C. W. Jenks, Aug., 1883 (NHA).
S. Randii (Porter) Britt. var. monticola Porter. C. F.
Batchelder, June, 1919 (NHA).
S. juncea Ait. Parsons and. Seymour, Aug., 1927 (MASS).
—S. nemoralis Ait. Fred M. Hunt, Aug., 1945 (NHA).
*—S. rugosa Ait. var. rugosa H. I. Baldwin, Oct., 1968 (NHA).
*_S. graminifolia (L.) Salisb. H. I. Baldwin, Sept., 1968
(FPC, NHA).
T—Aster divaricatus L. H. I. Baldwin, Sept., 1968 (FPC).
*—A. acuminatus Michx. B. L. Robinson, Aug., 1896 (NEBC).
T-Erigeron pulchellus Michx. H. I. Baldwin, July, 1968

(FPC).
*—Anaphalis margaritacea (L.) B.&H. D. W. Allen, May,
1968 (FPC).
Gnaphalium Macounii Greene. C. W. Jenks, Aug., 1883
(NHA).

G. obtusifolium L. C. A. Cheever, Sept., 1910 (HNH).

G. uliginosum L. W. Deane, Aug. 1891 (HNH).
*—Achillea Millefolium L. D. W. Allen, May, 1968 (FPC).
t-Chrysanthemum Leucanthemum L. var. pinnatifidum Le-

coq and Lamotte. H. I. Baldwin, July, 1971 (FPC, NHA).

Taraxacum erythrospermum Andrz. E. F. Williams, May,

1898 (HNH).
—T. officinale Weber. E. F. Williams, May, 1898 (NEBC).
T-Prenanthes trifoliata (Cass.) Fern. var. nana (Bigel.)
Fern. D. W. Allen, May, 1968 (FPC).
1—P. altissima L. H. I. Baldwin, Aug., 1968 (NHA).

220

Rhodora

[Vol. 76

Additional taxa reported by Thoreau in his journals of
1858 and 1860. The names, as given by Thoreau, including
his question marks are at the left in the following list:
Contemporary accepted binomials are listed at the right
opposite the name he used if that name is no longer used
or if only a common name were given by him.

Sensitive fern
Black spruce

Cinna arundinacea
Glyceria elongata

Eleocharis tenuis
Scirpus Eriophorum
Carex straminea

C. scoparia ?

C. debilis

C. Deweyana
Juncus acuminatus ?

J. parodoxus ?

Salix lucida
S. humilis
Tiarella cordifolia
Amelanchier canadensis
var Botryapium
A. canadensis
var oblongifolia
Geranium Robertianum
Epilobium palustre
Water andromeda

Checkerberry
Chelone glabra
Solidago virgaurea

Onoclea sensibilis L.

No Picea mariana (Mill) BSP.
has been found since Tho-
reau’s visits; this may be P.
rubens Sarg.

G. melicaria (Michx.) F. T.

Hubbard

S. cyperinus (L.) Kunth.
C. tenera Desv.

J. acuminatus Michx. or J. can-
adensis J. Gay

perhaps J. canadensis or J. acu-
minatus

A. arborea (Michx. F.) Fern.

A. canadensis (L.) Medic.

A. glaucophylla Link or Kalmia
polifolia Wang.

Gaultheria procumbens L.

?

1974] Flora of Mount Monadnock — Baldwin 221

Three-ribbed goldenrod

(not canadensis) S. altissima L. or S. gigantea Ait.
Aster thyrsoidea ?
Diplopappus umbellatus Aster umbellatus Mill.

DISCUSSION

The present flora of the upper part of Mt. Monadnock is
as noteworthy for the absence of certain plants as by the
presence of unusual ones such as the arctic-alpines. Thus
Alnus has not been found anywhere on the slopes of the
mountain, nor has Picea mariana been recorded from the
spruce-lined bogs. Acer saccharum is confined to a small
area on the east side. Further there are many instances of
lowland adventives that take root and may thrive for a
time and then disappear. This may explain why some spe-
cies recorded earlier have not now been found (aside from
less keen observation). A total of 220 taxa are represented
by specimens in herbaria, including 152 taxa collected from
1965 to 1972. Fifty-nine taxa collected during this period
were not previously found in herbaria, and may possibly
have been collected for the first time on the mountain. Of
the 94 taxa reported by Thoreau, 68 of which are in her-
baria, 62 were collected during the present survey.

Thoreau observed seeds of many lowland plants blowing
over the summit, and reflected on the origin of adventives
(and perhaps transitory taxa) :

“I saw what I took to be a thistle-down going low over
the summit, and might have caught it, though I saw no
thistle on the mountain top, nor any other plant from which
this could have come. (I have no doubt it was a thistle by
its appearance and its season.) It had evidently come up
from the country below. This shows that it may carry its
seeds to higher regions than it inhabits, and it suggests
how the seeds of some mountain plants, such as the Solidago
thyrsoidea may be conveyed from mountain to mountain,
also other solidagos, asters, epilobiums, willows, etc." Tho-
reau: Journal August 9, 1860, (p. 50) (page 1677 in the
1962 Dover edition).

222 Rhodora [Vol. 76

It seems obvious that most of the wind-disseminated spe-
cies that have restocked the mountain since the 1820 fire,
including Red Spruce, have come by the aerial route, mostly
probably from the adjacent lowland. Light-seeded arctic-
alpines may be included. It is less easy to explain how
heavy-seeded plants such as Vaccinium arrived. These may
have been present before the fires, surviving the shade of
spruce on some steep ledges, or they may have been carried
by birds migrating from timberline areas in the White
Mountains.

According to Pease (1964 p. 15) the following taxa col-
lected or reported from Mt. Monadnock “may be listed as
essentially alpine":

Lycopodium Selago (including var. appressum)

Lycopodium annotinum var. pungens

Juncus trifidus

Arenaria groenlandica

Solidago macrophylla var. thyrsoidea
Mt. Monadnock appears to be the southern limit for some
of these.

The relation of fire to the occurrence of arctic-alpine
plants needs more investigation. It is curious that they be-
came so abundant on Mt. Monadnock 29-30 years after the
great fire. Do the ashes from fire favor their establishment
(Fernald (1907) thought alkaline soil most unfavorable)
or is it open space, exposed rock crevices, or mineral soil
alone? Many years ago, searching for Vaccinium Vitis-
Idaea on Cape Breton, I could find it only on an area that
had been swept by fire about ten years before.

There are thus at least two possible explanations for the
presence on Mt. Monadnock of taxa usually found above
timberline:

(1) They have survived since post-glacial times in re-
fugia on the mountain itself possibly on lower slopes as pro-
posed by Fernald (1907, 1925), and then survived the
fires (which were very severe indeed), and then restocked
the bare rock and open soil.

(2) The summit flora may have arisen from disseminules

1974] Flora of Mount Monadnock — Baldwin 223

carried from timberline areas by wind, animals or birds,
or even men.

Steele and Hodgdon (1973) reached the same possible
explanations for the presence of Carex capitata and Carex
Bigelowii on Mt. Cardigan.

Regardless of the origin of the present flora, processes
of change are at work, reducing the abundance and area
occupied by some plants, while others are assuming dom-
inance. Many lowland forms may invade and thrive for a
time and then disappear. Neither the rate nor direction of
succession is uniform. Invasion of spruce on the upper
rocks has slowed to a temporary halt, even while the extent
of arctic-alpine plants declines.

The Problem of the Source and Persistence of
Arctic-Alpine Plants

How did arctic-alpine plants survive from post-glacial
times on lower mountains and escape competition with the
forest?

One explanation for the presence of arctic-alpine plants
below timberline is that they survived in refugia, rock
crevices or smal] exposed ledges since post-glacial times,
and that these crevices and ledges did not become over-
grown by other plants and that the microclimate remained
favorable. Then, when more open habitat was exposed by
fire, these plants re-invaded the bare summits and increased
greatly. Fernald (1907) postulated that alpine plants,
“forced to find temporary homes during the ice age on
potassic, calcareous or magnesium areas, found in the allu-
vial deposits a mixed soil in which they were able to spread.
As climate at low altitudes became warmer, these plants
found on cold cliffs and exposed mountain summits the only
habitats in which they have been able to persist within the
temperate regions” It is fairly easy to understand that they
were able to persist on alpine tundra above timberline, and
indeed on some “cold cliffs’ like those in Dixville Notch,
New Hampshire, and possibly elsewhere. Some plants
usually restricted to high altitudes and open tundra habitats

224 Rhodora [Vol. 76

are occasionally found below timberline, e.g. on talus and
rock piles in Carter Notch (a cool place) and even in some
open pastures (where they may be rather temporary resi-
dents). They cannot tolerate shade or root competition of
lowland plants. A small colony of Mountain Cranberry is
found along the Half Way House road on Monadnock (pos-
sibly introduced from the summit by berry pickers).

However, most steep rocky mountains that, like Monad-
nock, do not reach timberline and have not been burned
recently, are densely forested with spruce and at higher
levels, fir, with a heavy ground cover of moss, ferns and
herbs. There are probably hundreds of such mountains in
the Adirondacks and New England. Some of them have
exposed rocky cliffs, but records showing these to be havens
for alpine plants in undisturbed condition are few or want-
ing, Pease (op. cit.) reported finding some arctic-alpines
on open ledges on secondary mountains in the White Moun-
tains, *often caused by the destruction of the peaty humus
by fires," but he did not specify just how often these cases
were, nor did he give examples of such cases where there
was no trace or history of fire, He ventured no theory as
to how these plants reached the lowland habitat.

If arctic-alpine plants persisted in refugia in lower moun-
tains after the latter became forest-covered, it should be
possible to find these plants on unburned summits. They
have not been found, aside from the cases cited by Pease,
which were all close to high mountains supporting stable
colonies of these plants, whence invasion of even small
burned areas would be relatively easy compared to Mt.
Monadnock, which is far more remote.

More examples of bona fide refugia from ice-age times
need to be found, but even if these can be discovered and
dated, it may be impossible to demonstrate that no fires
occurred very long ago, or to prove how long the plants
have been on the site. Arctic flora of the Pleistocene became
adapted to the prevailing habitat conditions (Cf. Fernald,
1925) and it is still largely restricted to areas of open
ground and a cool, moist climate.

1974] Flora of Mount Monadnock — Baldwin 225

Did Arctic-alpine Plants Survive Fire and Spread
Over the Denuded Areas?

Assuming that these plants had succeeded in resisting
the competition of shade-tolerant trees and other plants,
and had remained in some secluded pocket since post-glacial
times, did they then survive the very intense fires that
burned these peaks, more destructive than any fires in
modern times? Fire destroyed all organic matter, and what
little mineral soil remained was then at the mercy of wind
and water. It must be remembered that two great fires on
Monadnock combined to make a tremendously hot and ex-
tensive burn. The 1800 fire consumed the humus and killed
the trees. The 1815 hurricane completed toppling them, so
that a formidible quantity of dry fuel was present. (It was
after a long hot, dry summer.) No fires in recent times
have approached it in heat and ferocity. Emerson (1896)
remarked that the heat was so great that it “blasted the
rock" into the ravines and caused slides.

How did Arctic-alpine Plants Come to Occupy Areas
Denuded by Fire?

Such plants must have arisen from disseminules brought
from nearby areas (refugia on the same mountain or in the
lowland) or from more distant sources, probably stretches
of alpine tundra on Mt. Moosilaukee or on the Presidential
Range. Since the lowland and lower slopes around Monad-
nock were largely cleared and cultivated or pastured, it is
possible that some of these plants had escaped from rocky
crevices on the mountain before the fire and had become
established on the lower open land. Here they might main-
tain themselves until forest succession once more eliminated
them. At the time of the fire, this land was still open. Even
if restocking of the burned mountain had proceeded from
this source, it is hard to see how the spread of heavy-seeded
species could have occurred uphill without the aid of birds
or animals. The latter carry seeds in burs that cling to
animal fur, but animals have not been found to be effective
carriers of berries and similar hard seeds.

226 Rhodora [Vol. 76

Were Birds Partly Responsible for the Spread of Plants
From Alpine Tundra to Monadnock?

Birds are well-known to be active and efficient carriers
of seeds. Ridley (1930) noted that birds are attracted by
seed color. Red seems to be the most attractive. Vaccinium
is a “bird-dispersed genus", he wrote. Forty-three species
of birds have been recorded to disperse plants of this genus,
and it seems likely that the red fruit of Mountain Cranberry
would be very much sought by birds. Taylor (1954) sug-
gested that birds are probably responsible for carrying
seeds to Macquaric Island, a small sub-antarctic island far
from any large land area that was severely glaciated during
the Pleistocene. He believed it unlikely that any of the 35
plant species could have survived the ice. Most of these
have relatively large seeds, but all have propagules suited
to bird transport.

Birds are important agents in the natural dissemination
of seeds of tree species having fleshy fruits, and presum-
ably shrubs as well. Red Cedar, Black Cherry and Black
Gum are examples. Seeds of Juniperus that have been
eaten by birds are said to germinate more promptly. (Anon.
1948) Indeed such hard-seeded species that display seed
coat dormancy may require the acid treatment received
during passage through a bird’s intestinal tract in order to
germinate. Phillips (1910) made an especially thorough
and wide-ranging study of bird dissemination of Juniperus
seed, and concluded that birds are responsible for most of
the spread of junipers, especially upwards on slopes and
up stream courses.

It is possible that the most likely candidate for being a
vector of seeds from alpine fell fields is the Slate-Colored
Junco. Not only did Thoreau remark about its being the
most common bird on Monadnock, but it is also abundant
there today. Forbush (1929) states that it is probably the
only bird that nests above tree limit in the White Moun-
tains. (I have found a nest with eggs in late June on Boott
Spur.) Forbush (op. cit. p. 90) reported that it eats many

1974] Flora of Mount Monadnock — Baldwin 227

dried berries and seeds. Objections to the theory of bird
transport include the questions concerning birds eating be-
fore migration, seeds passing undamaged, and limits set by
the time to pass through the digestive tract. Egler (1972)
questions the explanation “by having far-flying constipated
birds eject over distant lands.”

While there is thus evidence that some birds do carry
seeds, and birds are present on Mt. Monadnock, and that
seeds survive the treatment, or may be even benefited by
it, it is still not proven that heavy-seeded arctic-alpine
plants did actually become established on Monadnock by
this means.

Since this was written, Steele and Hodgdon (1973) have
examined these theories for the arrival on Mt. Cardigan of
two alpine species of Carex. They suggest a third possibil-
ity, that man may have consciously or unconsciously intro-
duced these plants to the bare rocky area. This appears
least likely in the case of the plants found on Monadnock,
because of the distance. It is also improbable that these
plants should appear on Mt. Chocorua and other sites. It
would be a very diligent Johnny Appleseed indeed. More
important, a prolonged period must be assumed for estab-
lishment. Success probably did not come with the first seed.

I am very grateful to Dr. Albion R. Hodgdon for checking
the identity of specimens collected, and for helpful sugges-
tions on the manuscript, also to Mr. David E. Boufford for
searching herbaria and listing Monadnock records.

LITERATURE CITED

ANON. 1948. Woody-Plant Seed Manual. Misc. Pub. 654, U.S. Dept.
Agri. 416 p.

BRAUN, E. Lucy. 1950. Deciduous Forests of Eastern North Amer-
ica. Blakiston, 596 p.

CHAMBERLAIN, ALLEN. 1936. Annals of the Grand Monadnock.
SPNHF, Concord, N.H. 195 p.

CHILD, HAMILTON. 1885. Gazetteer of Cheshire County, N.H. 1736-
1885. Pub. by the Daily Journal, Syracuse, N. Y. 560 p.

CUTTER, DANIEL B., M.D. 1881. History of the Town of Jaffrey,
N.H. Concord, N.H. 648 p.

228 Rhodora [Vol. 76

DANA, JAMES FREEMAN. 1816. Some Account of the Grand Monad-
nock. New England Jour. of Medicine and Surgery 5: 252-257.
Boston, Mass.

DEANE, WALTER. 1890. Flora of the Summit of Mt. Monadnock,
N.H. in July. Bull. Torr. Bot. Club 17: 316-318.

EGLER, FRANK E. 1972. (in a review) Ecology 53(5): 988.

ELLIS, GEORGE B. 1880. Memoir of Jacob Bigelow. Proc. Mass. Hist.
Soc. 17: 383-467.

EMERSON, EDWARD W. 1896. The Grand Monadnock. New England
Magazine 15: 33-51.

FERNALD, M. L. 1907. The Soil Preferences of Certain Alpine and
Subalpine Plants. Rhodora 9: 149-193.

1925. Persistence of Plants in Unglaciated Areas

of Boreal America. Memoirs Am. Acad Arts & Sciences 15: 235-

342.

. 1950. Gray's Manual of Botany. Eighth Edition.
Am. Book Co., N.Y. 1632 p.

ForBUSH, Epwarp Hower. 1929. Birds of Massachusetts and other
New England States. Part III. Mass. Dept. of Agri. Norwood,
Mass. 466 p.

LEONARD, LEVI W. 1855. History of Dublin, New Hampshire. Bos-
ton, Mass. 433 p.

PEASE, A. S. 1964. A Flora of Northern New Hampshire. New
England Botanical Club. Cambridge, Mass. 278 p.

PHILLIPS, FRANK J. 1910. The Dissemination of Junipers by Birds.
Forestry Quarterly 8: 60-73.

RIDLEY, HENRY N. 1930. The Dispersal of Plants Throughout the
World. L. Reeve & Co., Ashford, Kent, England. 744 p.

Spurr, S. H. 1964. Forest Ecology. Ronald Press, N.Y. 352 p.

STEELE, F. L. and A. R. Hopepon. 1973. Two Interesting Plants on
Mt. Cardigan. Rhodora 75: 155-157.

STONE, WILLIAM H. 1885. The Flora of Mt. Monadnock. Appalachia
4: 145-150.

TAYLOR, R. W. 1954. An Example of Long Distance Dispersal.
Ecology 35: 569-572.

THOREAU, HENRY D. (1906) 1962. The Journals of Henry D. Tho-
reau 1852-1860. Edited by Bradford Torrey and Francis H. Allen.
14 Vol. bound as two. Dover Pub. N.Y.

HILLSBORO, NEW HAMPSHIRE 03244.

TAXONOMY OF PERITYLE SECTION PERITYLE
(COMPOSITAE — PERITYLINAE) *

A. MICHAEL POWELL

Section PERITYLE, with 27 species and three varieties, is
the largest of three sections of the genus Perityle (Powell,
1968a). The distributional center of sect. PERITYLE is in
northwestern Mexico and the Baja Californian peninsula,
but the total natural range is extended by several taxa that
occur in the southwestern United States, Guadalupe Island,
the Revillagigedo Islands, and west-central Mexico (Jal-
isco). One weedy taxon, P. emoryi, has bicontinental dis-
tribution in North America and in Chile and Peru in South
America. Separate taxonomic works have been prepared
for the other two sections of the genus; Pappothrix (Powell,
1969) and Laphamia (Powell, 1973).

The species of sect. PERITYLE comprise a taxon that was
recognized as the genus Perityle by Everly (1947) who
compiled a taxonomic treatment of the group. In connection
with her morphologically oriented investigations of Perityle,
Everly also studied the closely related taxa Pappothrix and
Laphamia, which she regarded as genera. These taxa are
distinguished primarily by somewhat variable pappus and
achene characteristies, and several workers have questioned
their status as genera. Shinners (1959) merged Pappothrix
and Laphamia with Perityle, noting their close morpho-
logical relationship and contending that pappus differences
were not sufficient to warrant separate generic status. In
recognizing Pappothrix and Laphamia as sections of Peri-
tyle, I have followed Shinners’ congeneric disposition of
the taxa (Powell, 1968a). My systematic judgements were
based upon the evaluation of new information from cyto-
logical, chromatographic, and hybridization techniques, as
well as a re-evaluation of morphological] features. The re-

'Supported by National Science Foundation Grants GB-7740 and
GB-20361.

229

230 Rhodora [Vol. 76

sults have suggested that Pappothrix, Laphamia, and Peri-
tyle are closely related, essentially natural phylads, and
that they are best treated as sections of a single genus.
Niles (1970), however, has maintained Perityle and Lapha-
mia as distinct genera, while combining Pappothrix with
Laphamia. Niles did not conduct a comprehensive study of
the sect. PERITYLE species. A more complete review of the
taxonomic history concerning Perityle is available elsewhere
(Shinners, 1959; Powell, 1968a; Niles, 1970).

Perityle is closely allied to Pericome, Amauria, and
Eutetras of the subtribe Peritylinae (Helenieae). Rydberg
(1914) assigned Perityle and Pericome to the Peritylinae
and Amauria and Ewutetras to the subtribe Amaurinae.
Turner (1966) and Powell (1972a) have suggested that
Amaurinae (in part) should be combined with Peritylinae.
Presently the Peritylinae is being re-evaluated with refer-
ence to Rydberg's concepts, and the proper tribal position
of the subtribe is also being investigated (Powell & Turner,
unpublished).

CHROMOSOMAL CONSIDERATIONS

A discussion of the chromosome numbers of Perityle and
related genera is presented elsewhere (Powell, 1968b).
Additional counts for the species of sect. PERITYLE are
reported in Powell and Sikes (1970) and Powell (1972b).
Chromosome numbers have been obtained for 20 of the 27
species of sect. PERITYLE. The counts for specific taxa are
included with a list of the species under Phylogenetic
Considerations (Table 1), and they are also included with
species discussions in the taxonomic portion of this paper.

In Powell (1968b), a few changes must be noted with
regard to the counts for sect. PERITYLE. When the 1968
paper was compiled, the taxonomic studies of sect. PERITYLE
were in early stages. The reports of n — 16 for P. cali-
fornica should be attributed to P. cuneata var. marginata.
Perityle incompta is now considered as synonymous with
P. crassifolia var. robusta, and the counts listed for the
former (n = 19, 18) should be attributed to the latter

1974] Perityle — Powell 231

taxon. The diploid P. microglossa (Sikes and Babcock 185
a,b,c,f,) is P. microglossa var. saxosa, and the polyploid
P. microglossa (Sikes and Babcock 177 192 a-d, 152, 157 a,
206 a,b,) is var. microglossa. Sikes and Babcock 188 a,b,d,f,
and 190 a,b, listed as P. palmeri, are P. cordifolia (n = YT).
Perityle cf. spilanthoides is P. turneri (n — 1T) and P.
vaseyi (Sikes 116) is P. parryi (n — YT).

Most species of sect. PERITYLE are diploid, with counts of
n — 19, 18, 17, 16, 13, 12, and 11 having been reported for
the various taxa (Powell, 1968b; 1972b). Only P. micro-
glossa var. microglossa (n = 34, 51), P. emoryi (n = 32-36,
50-57), and P. incana (n = 50-57) are known to be poly-
ploid. The first two taxa are relatively well established as
exclusively polyploid, while only one count for the Guada-
lupe Island endemic P. incana (Powell, 1972b) has been
recorded. Turner and Flyr (1966) reported n = ca. 46 for
P. crassifolia var. robusta (Cowan 2265), but I have since
identified this collection as P. emoryi. A base number for
P. emoryi has not been determined in spite of numerous
attempts to obtain exact counts (Powell, 1968b).

The ancestral base number for sect. PERITYLE has not
been established. Subsequent to an earlier discussion of
the subject (Powell, 1968b), all attempts to resolve the
question have been unsuccessful, including those which in-
volved the analysis of experimenta] hybrids (Powell, un-
published). Most chromosomal] and distributional evidence
suggest a base of x — 17 for sect. PERITYLE, and the genus
as a whole (see Phylogenetic Considerations). Fourteen
species of sect PERITYLE are n = 17, and both sect. PAP-
POTHRIX (5 spp.) and sect. LAPHAMIA (21 spp.) are x = 17
(Powell, 1969; 1973). The taxa of sect. PERITYLE with
chromosome numbers below n = 17, P. californica (n = 13,
12, 11) and P. cuneata (n = 16, 12), are clearly aneuploid
derivatives, probably of the » — 17 line. However, the
origin of taxa with n = 19 (P. crassifolia and allies) is of
prime concern in evaluating the phylogeny of sect. PERI-
TYLE. Distributionally, the n = 19 “alliance” ( (southern
Baja California peninsula and neighboring islands) is pe-

232 Rhodora [Vol. 76

ripheral to the group where n — 17, which is centered on
the mainland of Mexico (see Phylogenetic Considerations).
The » — 19 group may be either ancestral to or derived
from the » — 17 group (Powell, 1968b), but I now believe
that the species with » — 19 and the species with » — 17
both evolved as separate lineages from a common “main-
land" ancestor that had a chromosome number of n = 19
(or 18). Consideration of the latter hypothesis in a broader
sense is supported by the base numbers of the related
genera Pericome (x = 18), Eutetras (x = 18), and Amau-
ria (x = 18).

ARTIFICIAL HYBRIDIZATIONS

Artificial hybridizations have been conducted with 16
species of sect. PERITYLE. Many of the crosses were of
intergeneric and intersectiona] nature, although several
interspecific combinations have been obtained. The results
of most of these crosses are discussed in another paper
(Powell, 1972c) where emphasis was placed upon evalua-
tion of the intergeneric and intersectional relationships of
Perityle. A hybridization program aimed at the evaluation
of interspecific relationships within sect. Perityle is still
in progress (Powell, unpublished). Where appropriate,
some of the preliminary information derived from artificial
crosses will be brought into species discussions in the taxo-
nomic portion of this paper.

Natural hybridization is rare in sect. PERITYLE, and only
one such occurrence has been documented (Powell, 1970).
Most of the species are geographically isolated, but a few
taxa do have overlapping ranges and may occasionally hy-
bridize.

CHROMATOGRAPHIC CONSIDERATIONS

Nineteen species of sect. PERITYLE were analyzed by 2-
dimensional paper chromatography of leaf extracts. In
general, the techniques outlined by Alston and Turner
(1963) were utilized. The procedures for sampling popu-

1974] Perityle — Powell 233

lations and analyzing chromatograms have followed those
discussed elsewhere (Powell, 1973). The major components
of chromatographic patterns were characterized as flavo-
noids (Powell and Tsang, 1966; Powell and Averett, un-
published), although the specific identity of only one com-
pound is known (Southwick et al., 1972).

The use of chromatographic techniques has been a valu-
able asset to previous taxonomic studies in the Peritylinae
(Powell and Tsang, 1966; Powell, 1968a; 1969; 1973).
Intergeneric and infrageneric relationships have been clari-
fied through the comparative analysis of pattern data. In
previous work I treated the crude chemical information
from a chromatographic pattern as a single taxonomic
character, and pattern data have not been given more
weight than any other chraacter.

The results of chromatographic studies in sect. PERITYLE
have revealed two strikingly different chemical patterns
based upon the presence or absence of yellow compounds
as seen under ultraviolet light. The white-flowered species
centered in the Sierra Madre Occidental (e.g., P. turneri),
and the yellow-flowered species of northwestern Mexico and
Baja California (e.g., P. cordifolia and P. californica) have
simple patterns consisting of a few dark purple spots. The
Baja Californian white-flowered species (e.g. P. crassi-
folia), and the Texas yellow-flowered species (e.g., P. par-
ryi) have relatively complex patterns that include promi-
nent yellow spots. Yellow compounds are considered im-
portant because they comprise the basic chromatographic
profile of sect. PAPPOTHRIX, sect. LAPHAMIA, and the genera
related to Perityle. The actual chromatographic data are
not presented here in the form of figures or tables. Instead,
where appropriate, the data are included in succeeding dis-
cussions concerning relationships of species and species
groups.

ECOLOGICAL AND REPRODUCTIVE CONSIDERATIONS

The species of sect. PERITYLE occur in rock and in soil
habitats, unlike the species of sect. PAPPOTHRIX and LAPHA-

234 Rhodora [Vol. 76

MIA, which are exclusively rock-dwelling (Powell, 1969;
1973). Certain species of sect. PERITYLE are obligate, rock-
dwelling perennials, while several perennial and annual
taxa are soil-obligates. In addition, a few taxa display
facultative existence in rock and soil habitats.

The rock-dwelling members of sect. PERITYLE occur under
a wide range of climatic conditions and at various eleva-
tions. About half of the petrophilic taxa are occupants of
protected canyons and exposures in relatively xeric desert
mountains at altitudes of 1000-4000 feet or less, and the
others thrive under relatively mesic conditions at altitudes
of 4000-8000 feet. Perityle turneri also occurs under mesic
conditions, at 7000-9500 feet in and near Durango, Mexico,
but may be rooted primarily in matted soils. The other soil-
dwelling taxa occur almost exclusively under desert condi-
tions.

Perityle is essentially a genus of self-incompatible spe-
cies (Powell, 1972c). Only two of the taxa, P. emoryi and
P. microglossa var. microglossa, are known to be self-com-
patible, and these are members of sect. PERITYLE. Ten spe-
cies have not been tested for the compatibility factor, two
of sect. LAPHAMIA and eight of sect. PERITYLE, but these
taxa are also probably self-incompatible. Both of the self-
compatible taxa are widespread polyploids (Powell, 1968b;
1972c) that are “weedy” and annual in habit. Perityle
emoryi is the most widespread species of the genus, having
bi-continental distribution in North and South America,
and P. microglossa var. microglossa probably ranks second
in distributional range. Several other species of the genus
are polyploid, and a few others are annual, thus suggesting
that the capacity for reproductive self-compatibility has
allowed for the “weedy” nature of two taxa.

PHYLOGENETIC CONSIDERATIONS

Speculation concerning phylogeny of the subtribe Perity-
linae and the genus Perityle has been discussed generally
elsewhere (Powell, 1973), and will not be repezted in detail
here. Essentially, it was proposed that Peritylinae origi-

1974] Perityle — Powell 235

nated in the southern Sierra Madre Occidental of Mexico.
Perityle is by far the most successful genus of the subtribe,
in terms of number of species, with only 2 or 3 species being
recognized for each of the other genera (Powell, 1972a;
1972d; and Turner, 1966). Section PERITYLE is assumed to
be ancestral to the other two sections of the genus. Based
upon distributional, morphological, chromosomal, and other
considerations, it is believed that sects. PAPPOTHRIX and
LAPHAMIA underwent rapid evolution in late Pliocene or
Pleistocene emanating from an ancestral sect. PERITYLE
complex (Powell, 1975).

Modes of speciation in the subtribe Peritylinae, and Peri-
tyle in particular, have been discussed elsewhere (Powell,
1969; 1972c; 1973). In sects. PAPPOTHRIX and LAPHAMIA,
speciation is believed to have occurred mostly, if not en-
tirely, as a result of geographic isolation. In sect. PERITYLE,
speciation probably has occurred as a result of geographic
isolation and other, more nearly reproductive phenomena
(Powell, 1972c). This subject wil be diseussed further
below.

In Table 1 the list of sect. PERITYLE species is presented
as an aid to discussing phylogenetic considerations. The
list has been organized to depict the hypothetical evolution-
ary groupings within the section. The arrangements are
based upon morphological, chromosomal, experimental,
chromatographic, and eco-geographical data. Additional
discussions of species relationships, including the evidence
upon which such judgements are based, are included in the
taxonomic portion of this paper.

The total data suggest that sect. PERITYLE comprises at
least two distinct evolutionary series. One series, including
the first 19 taxa, is marked by white ligules and yellow (or
cream-white) disc corollas, with a few exceptions, as indi-
cated in Table 1. Perityle incana and the P. jaliscana group
are relatively anomalous in the “white-flowered series" and
they may comprise separate and equal phyletic entities. A
second major series can be delimited by its exclusively
yellow flowers.

76

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1974] Perityle — Powell 237

Several related-species groups are further delimited
within both the ‘“white-flowered series" and “yellow-
flowered series” (Table 1). The groupings themselves com-
prise clearly related taxa, but the phyletic unity of the
related-species groups within monophyletic white- or yellow-
flowered series is indefinite. For example, origin of the
Baja Californian P. crassifolia group has not been con-
nected resolutely with the Sierra Madre Occidental P.
turneri group, even though both exhibit white ligules. And
the anamalous nature of P. incana and the P. jaliscana
group is indicated above. Also, there is no certainty that
the P. parryi group of Texas and the aneuploid P. cali-
fornica group of Baja California have monophyletic ties
within a yellow-flowered series.

White-Flowered Groups (Table 1). Three subgroups of
the white-flowered series are recognized. These are desig-
nated as the Perityle crassifolia, P. turneri, and P. jaliscana
related-species groups. Distributional information, habitat
preferences, and life form habits are given in the following
diseussions because of their supposed importance in under-
standing speciation in the genus.

The Perityle crassifolia group, also including P. socor-
rensis, P. emoryi, and P. aurea, is distributed in southern
Baja California and nearby islands, except for P. emoryi
which is a widespread weed. Also, P. socorrensis is found
only in the Revillagigedo Islands, which are over 200 miles
south of Baja California.

Perityle emoryi (n = 32-36, 50-57) is believed to be a
polyploid derivative of P. crassifolia var. robusta. Morpho-
logical and chromatographic data strongly suggest that
P. aurea belongs with the P. crassifolia (x — 19) group
even though this taxon has yellow-ligules and a chromo-
some number of n = 17.

Both Perityle emoryi and P. aurea are annual in habit,
while the other taxa of the P. crassifolia group are essen-
tially soil-dwelling perennials. All of the taxa, however,
with the exception of P. crassifolia var. crassifolia which
seems restricted to sand, might also occur in soil-filled

238 Rhodora [Vol. 76

crevices of rocks especially at sea side. The Guadalupe
Island endemic P. incana, is possibly allied with the P.
crassifolia group, but evidence as to its true relationship
has not been conclusive.

The base chromosome number of the Perityle crassifolia
group is presumed to be x — 19, a number that might be
ancestral in sect. Perityle. On morphological and distribu-
tional grounds, however, it is practically impossible to en-
vision the other related species groups as having been de-
rived from the P. crassifolia line. I believe that the P.
crassifolia line was an early offshoot from ancestral Peri-
tylinae.

The largest related-species group of the white-flowered
series is distributionally centered in the Sierra Madre
Occidental] of mainland Mexico. Two assemblages of four
species each are further distinguishable among this Perityle
turneri group, primarily on the basis of floral characters.
Perityle turneri, P. microcephala, and P. lineariloba are
clearly allied taxa which occur at relatively high altitudes
from Durango to central Chihuahua. The fourth member
of this assemblage is P. microglossa, which ranges from
northern Sonora along the west coast of Mexico south to
Colima, into the foothills of the Sierra Madre Occidental,
and east to San Luis Potosi and Texas. Habit and distribu-
tional considerations (see Taxonomy) strongly suggest that
P. microglossa is derived from P. microcephala. I believe
that P. turneri is primitive in this assemblage, on the basis
of features discussed elsewhere (Powell, 1972e).

Perityle coronopifolia, P. ciliata, P. hofmeisteria, and P.
canescens are also clearly allied members of the P. turneri
group. Perityle coronopifolia extends from northern Chi-
huahua into the mountains of southern Arizona and south-
ern New Mexico. Perityle ciliata is restricted to central
Arizona, while P. hofmeisteria and P. canescens apparently
are endemics respectively in Durango and in Sinaloa,
Mexico.

Most members of the Perityle turneri group are hardy,
suffrutescent perennials that live in crevices of rock bluffs.

1974] Perityle — Powell 239

Perityle turneri is a rhizomatous perennial, apparently
occurring typically in thin soil covering rocky outcrops, but
according to label data, the taxon might also be rock-dwell-
ing. Perityle microglossa is a soil-dwelling annual, also
found occasionally among rocks, and one collection of P.
hofmeisteria (Flyr 332) is said by the collector to be “seem-
ingly an annual.” Judging from known distributional data,
these species are strictly allopatric, suggesting that they
have undergone the type of geographic speciation that is
considered typical for sect. PAPPOTHRIX and sect. LAPHAMIA
(Powell, 1969; 1973).

The white-rayed (or discoid) Perityle jaliscana group
with cream-white discs comprises very closely related rock-
dwelling perennial taxa that are restricted to Jalisco, Mex-
ico. These taxa represent a third distinct assemblage within
the white-flowered species of sect. PERITYLE.

The three white-flowered subgroups are clearly delimited
morphologically, with perhaps the most useful distinguish-
ing features being those of the habits, achenes, and pappus
(see Taxonomy). The yellow-flowered species are quite
different superficially, but the generic unity of both white-
and yellow-flowered assemblages is evident.

Yellow-Flowered Groups (Table 1). Geographically, the
yellow-flowered species are divided into two major assem-
blages separated by the Sierra Madre Occidental. One as-
semblage, designated as the Perityle parryi group, has a
limited distribution in the Big Bend area of western Texas
and adjacent Mexico, and the other geographic assemblage,
including the P. cordifolia and P. californica groups, is
more widely distributed in Sonora, Sinaloa, and Baja
California, Mexico.

The three species of the Perityle parryi group are closely
related perennials, being distinguished on the basis of a
few morphological characters. One of the taxa, P. aglossa,
apparently is exclusively rock-dwelling in habit, while P.
vaseyi seemingly occurs only in desert soils. Perityle parryt,
however, is facultative in its existence in rocks and soils.

There are interesting biological parallels between the

240 Rhodora [Vol. 76

Perityle parryi group and the P. cordifolia group of west-
ern Mexico, and despite their geographic separation, most
evidence suggests that both groups had common origin.
Both alliances comprise closely related taxa, their vegeta-
tive and floral morphologies are strikingly similar, and
species of both groups have chromosome numbers of n = 17
with no polyploidy having been discovered in any of the
taxa. Additionally, and most remarkably, the evolutionary
pattern of the disjunct groups seems to be that soil forms
were established when woody perennials “came down out
of the rocks and adapted to life in the soil.” Perityle cordi-
folia, P. leptoglossa, and P. lloydii seem to be obligately (at
least typically) rock-dwelling, while P. palmeri and P.
lobata apparently are facultative in their tolerance for
existence in rocks or in soils.

The presumed “close” phylogenetic relationship between
the Perityle parryi and P. cordifolia groups is not supported
by their chemical profiles. Members of the P. parryi group
produce abundant yellow components, as do P. crassifolia
and allies, while chromatographic samples of the P. cordi-
folia group have revealed only simple patterns of purplish
spots. The chemical indication of affinity between the P.
parryt and P. crassifolia groups is complemented by a
tenuous morphological feature they have in common. Peri-
tyle vaseyi (rarely P. parryi, also) of the former alliance
and pappose members of the P. crassifolia group are the
only taxa of the genus which have awns with retrorsely-
barbed tips. In view of the morphological dissimilarity
and geographic distance between these groups, however, I
believe that the crude chemical data and pappus structure
merely reflect an ancestral connection between the P. crassi-
folia and yellow-flowered alliances. If this is true, then the
P. cordifolia group must have lost its ability to produce the
prominent yellow flavonoids.

Both species of the Perityle californica group are soil-
dwelling annuals. The taxa are of special systematic inter-
est in sect. PERITYLE because of their reduced chromosome
numbers. All other yellow-flowered taxa have m — 17.

1974] Perityle — Powell 241

Perityle californica (n = 13,12,11) is distributed in Sonora
and Sinaloa of mainland Mexico and along most of the Baja
California peninsula to just south of La Paz. Perityle
cuneata (n = 16,12) is restricted to the southern tip of
Baja California with northern limits to about the latitude
of La Paz.

Judging from overall morphological similarity and dis-
tributional considerations, it is assumed that Perityle cali-
fornica and P. cuneata are aneuploid derivatives of the P.
cordifolia line. With the exception of the P. parryi group,
which seemingly would be ruled out on distributional
grounds, only the P. cordifolia group bears any close re-
semblance to the aneuploids. Perityle californica occurs
sympatrically with P. cordifolia and P. palmeri in Sonora
and Sinaloa, apparently without hybridization. Since most
species of Perityle are allopatric, but artificially interfertile
(Powell, 1972c), it is reasonable to conclude that P. cali-
forniea achieved reproductive isolation and speciation
through reduction in chromosome number.

Summary of Phylogenetic Considerations. Species of the
genus Perityle are distributed primarily in the arid to semi-
mesic mountains of desert North America, but also in the
higher mountains, for example in the Sierra Madre Occi-
dental. The majority of the species are obligate rock-
dwellers. All five species of sect. PAPPOTHRIX and all 21
species of sect. LAPHAMIA occur as perennials in crevices
of rock bluffs, apparently never growing in soil at the base
of bluffs. That these species can survive only in such re-
strictive habitats surely imposes upon them evolutionary
limitations. The basic evolutionary trends of sect. PAPPO-
THRIX and LAPHAMIA have followed a pattern of speciation
by geographic isolation; thus evolved the groups of closely
related, mostly endenmic species, characteristic of the above
sections.

Greater evolutionary diversity is exhibited among the
species of sect. PERITYLE. Indeed, some species of this sec-
tion are obligate, rock-dwelling perennials that presumably
underwent the type of geographic speciation mentioned

242 Rhodora [Vol. 76

above. But several taxa have adapted to a facultative ex-
istence in rocks and in soils, and a few species are exclu-
sively soil-dwelling with perennial or annual habits. For
certain taxa, the adaptation to life in the soil apparently
has been facilitated by two evolutionary mechanisms, poly-
ploidy and aneuploidy. Emanating from the white-flowered
groups, the widespread weeds Perityle microglossa var.
microglossa (n = 34,51) and P. emoryi (n = 32-36, 50-57)
exemplify the adaptive success of polyploidy. Perityle cali-
fornica (n = 11, 12, 19) and P. cuneata (n = 12, 16) are
successful aneuploid species of yellow-flowered lineage.
Thus, along with the annual habit displayed by the above
species, the phenomena of polyploidy and aneuploidy have
independently advanced the adaptive capacity of species
belonging to distinct evolutionary units of sect. PERITYLE.

ACKNOWLEDGEMENTS

I am grateful to the curators of herbaria from which
specimens were borrowed. I am also grateful to B. L.
Turner who suggested the study of Perityle and provided
encouragement and advice throughout the study. I extend
thanks to John Averett for his assistance in gathering
literature; to Reid Moran for his help in providing vouch-
ers, buds, and ecological information for P. incana and
other species; to Rogers McVaugh for providing helpful
observations and material of P. microglossa and P. fedde-
mae; and to Annetta Carter for her assistance in providing
specimens and ecological information for several species.
Many other people have contributed to my studies of Peri-
tyle and related genera; they know of my appreciation.
One person who did not know fully of my deep gratitude
was the late Lloyd Shinners. His special character will
linger to inspire many taxonomists.

TAXONOMY

PERITYLE Benth. Bot. Sulph. 23. 1844.
Perityle section Perityle, see Powell, Sida 3: 277. 1968.

1974] Perityle — Powell 243

Galinsogeopsis Schultz-Bip. in Seem. Bot. Herald. 306. 1856.
Nesothamnus Rydb. N. Amer. Fl. 34: 12. 1914.
Leptopharynz Rydb. N. Amer. Fl. 34: 21. 1914. (in part).
Closia Remy in Gay, Fl. Chile 4: 120. 1849. (in part).

Plants shrubby, suffruticose, or herbaceous perennials,
or delicate to robust annuals, (2)10-80 cm high, growing
in rock crevices or various soils, variously pubescent, to-
mentulose to nearly glabrous, frequently glandular-pu-
bescent; leaves opposite or alternate, variable in size and
shape, 0.7-14 cm long including the petiole, 0.2-7.0 cm wide,
ovate, cordate, deltoid, subhastate, reniform, or suborbicu-
lar in outline, the margins subentire, serrate to doubly so,
or shallow to deeply lobed or divided, frequently 3-5 lobed,
pedate, subcruciform, or pinnatifed with narrow segments,
nearly glabrous, puberulent, to arachnoid-villous, often
glandular-pubescent, thick and semisucculent to thin in
texture, viscid, bitter tasting, usually aromatic; capitu-
lescence of solitary heads, or heads loosely to tightly aggre-
gated in clusters of 3 or more; peduncles short or long;
involucres subcylindric to hemispheric; heads 3-12 mm high,
4-15 mm wide; involucres of 2(3) subequal series, bracts
ovate or obovate to narrowly oblanceolate, obtuse to at-
tenuate at the apex, flattened, ribbed, or keeled, usually
spreading or reflexing at maturity; heads radiate or discoid
(in 4 spp.), ligules yellow or white, showy or rudimentary ;
disc flowers numerous, corollas 4-lobed, yellow or cream-
white (in 3-4 spp.), throats tubular to broadly funnelform
or narrowly campanulate; style branches, flattened, linear,
usually tapering to a fine, minutely pubescent tip; achenes
black when mature, 1.3-5.0 mm long, distinctly flattened
radially, outer achenes often 3-angled, slightly curved and
clasped by bracts, linear, oblong, oblong-elliptic, or oblance-
olate, the margins densely ciliate, rarely merely puberulent,
prominently calloused or the margins thin and not cal-
loused, the faces glabrous and shiny or short-pubescent;
pappus of a conspicuous, rarely inconspicuous, crown of
laciniate, hyaline, squamellae, and 0-2 (rarely 3-4) delicate
or rather stout bristles, 0.5-7.0 mm long, the bristles naked,

244 Rhodora [Vol. 76

barbellate, or subplumose throughout or only at the tips,
the barbs antrorse, lateral, or retrorse; base chromosome
number, x = 17 or 19.

Type: P. californica Benth.

Key to the Species

1. Heads discoid. 2

a somos otov t 9 9 n nm m m m rs n, n n n n n. n s s n. d

2. Leaves canescent; Guadalupe Lsland, Mexico. ....
TEM 5. P. incana.

2. Leaves not canescent; Texas or Jalisco, Mexico . 3.
3. Pappus bristle 1, 3.7-5.5 mm long; disc corollas

yellow; Texas. ................ 20. P. aglossa.

2 Pappus bristles 2(3), 0.5-1 mm long; dise corol-

las cream-white or pale yellow; Jalisco. ...... 4.

4. Leaves dissected. ........ 17. P. feddemae.

4. Leaves deltoid to subhalberd. 14. P. jaliscana.

1. Heads radiate. .... ""arzsKAazPe»saerszrresruss D
5. Ray and disc corollas yellow. ................. 6.

6. Plants distinctly annual; pappus bristles 1 or 2

(or rarely absent). ....................... T.

7. Achene margins thin, not calloused; single
pappus bristle delicate (rarely absent), and
retrorsely barbellate only at the tip. ........
MM 4. P. aurea.

7. Achene margins usually prominently cal-
loused; single pappus bristle rather stout
and antrorsely subplumose, or 2 delicate

bristles. .... 0.0.0.0... .0.0............ 8.
8. Pappus bristle 1, subplumose, 1.5-3.5 mm
long. ...... 2 ee. 26. P. californica.
8. Pappus bristles 2 (rarely absent), delicate,
0.5-2.0 mm long. .................... 9.

9. Achenes (2.5) 3-3.8 mm long, obcor-
date-cuneate, with broad, callous mar-
gins. ... 27a. P. cuneata var. cuneata.

9. Achenes 1.5-2 (2.5) mm long, obovate
to subcuneate with prominent callous

1974]

Perityle — Powell 245

TOTS suc Se a
27b. P. cuneata var. marginata.

6. Plants perennial, with woody bases, or herba-

ceous with fleshy taproots or rather thin branch

Toota 2 ee ec ele oe 10.
10. Plants of West Texas and adjacent Mexico.
ACTI NS DEUS or weer US sot FP

11. Leaves typically 3-lobed or subcruci-

form; taprooted perennials in soil. ..
DE AE cod DOT TON 10. P. vaseyt.
11. Leaves typically 3-lobed but not divided;
woody-based perennials in rock crevices,
or taprooted in soil ..... 18 P. parryi.
10. Plants of Sonora and Sinaloa, Mexico, and
Baja California Sur. ..... eoe 12.
19. Plants subshrubs or suffrutescent peren-
nidis. eur NEZX C S 13.
13. Leaves densely puberulent and sub-
canescent. ..... 22. P. leptoglossa.
13. Leaves pilose and green. ...... 14.
14. Leaves 3.5-11 em long; involu-

eral bracts 10-13 mm long.

EM _.. 21. P. cordifolia.
14. Leaves 2.5-4.5 cm long; involu-
eral bracts 5-7 mm long. .. .
ee NICE 23. P. lloydii.
12. Plants herbaceous perennials with
rather thin, fleshy roots. ........- 15.
15. Leaf blades 3-5 lobed, cleft, parted
or rarely divided, the margins den-
tate-lobed and acuminate; Baja
California Sur. ..... 24. P. lobata.
15. Leaf blades shallowly or strongly
3.lobed, the margins serrate-den-
tate; Sonora and Sinaloa, Mexico.
Rl) MN OCA Um 25. P. palmeri.

5. Ray corollas white, disc corollas yellow, except disc

corollas cream-white in P. rosea and P. trichodonta.
meds cp c a 16.

Rhodora [Vol. 76

16. Disc corollas eream-white. ... .. ....... 17.
17. Leaves 1.8-3 cm long, 1-1.5 (2) em wide;

ligules 2-2.5 mm long. ....... 15. P. rosei.

17. Leaves 0.7-0.9 em long, 0.25-0.5 cm wide;

ligules 1-1.5 mm long. .. 16. P. trichodonta.

16. Disc corollas yellow. .................... 18.

18.

18.

Achene margins thin, not calloused ; heads
0.6-1 cm high, 0.6-1.5 cm wide, but possibly
smaller in P. emoryi; Baja California Sur
and islands, except P. emoryi widespread.
DNUS 19.
19. Ligules 6-10 mm long. ......... 20.
20. Leaves usually thick and crisped,
arachnoid-villous; Baja California

Sur, coastal dune sand. [.

. la. P. crassifolia var. crassifolia.

20. Leaves usually rather thin and not
crisped, densely short-pubescent to
glabrous; Baja California Sur, in

various soils. ..................

. lb. P. crassifolia var. robusta.

19. Ligules 1.5-4 (6) mm long (rarely ab-
sent). ........ eee e eese. 21.

2]. Plants suffruticose perennials;
Revillagigedo Islands. ..........

ss 2. P. socorrensis.

21. Plants delicate or robust annuals;
widespread weed. .. 3. P. emoryi.

Achene margins thin or prominently cal-
loused; heads 3-7 mm high, 4-8 mm wide,
but may be wider in P. turneri; Sierra
Madre Occidental, Mexico, and foothills,
and Arizona, except P. microglossa wide-

spread. ........0.00.0.-0..0.000 004. 22.
22. Achenes 1.8-3.5 mm long; pappus bris-
tles 2 (3), 1.5-3 mm long. ....... 23.

23. Leaves deltoid-ovate to ovate-
rhombic, the margins serrate,

1974]

lS
N

Perityle — Powell 247

shallow-lobed, or serrate-crenate;
central Arizona. ... 11. P. ciliata.
Leaves ovate to subspathulate and
entire to shallow-lobed, or ovate-
cordate and serrate to serrate-
lobed, or 2-3-pinnatifid to pedately
divided; southern Arizona and
Mexico. . ... SUM DNE
24. Leaves entire, shallow-lobed
or serrate; Durango, Mexico.
ANES 13. P. hofmeisteria.

24. Leaves pinnatifid or pedately
divided with spathulate or
linear segments; southern
Arizona and adjacent Mexico.

ow Haani P re er 25.

25. Capitulescence of several
heads clustered on short
peduncles; achene mar-

gins typically long-ciliate.

_.. 12. P. coronopifolia.

25. Capitulescence essentially

of solitary heads; achene

margins merely puberu-

lent. 10. P. canescens.

Achenes 1.3-2 mm long; pappus bris-
tles 2 (or 0-2), 0.5-1.5 mm long. . 26.

26.

26.

Leaves pinnately 3-5 divided, the
divisions linear or nearly so.
— D ! . 7. P. lineariloba.

Leaves otherwise. ..... IDEE

27. Heads 0.7-1.4 cm wide. ..
MINNS 6. P. turneri.

27. Heads 4-7 mm wide. . 28.

28. Plants suffrutescent per-
ennials; leaves densely
grayish-puberulent.

. 8. P. microcephala.

248 Rhodora [Vol. 76

28. Plants delicate or robust
annuals; leaves puberu-
lent, glandular-puberu-
lent, or glabrous. 29.
29. Ligules 1.5-3.5 mm

long; upper pedun-
cles usually copi-
ously glandular-pu-
berulent. ....
.. 9a. P. microglossa
var. microglossa.

29. Ligules 3.5-4.5 mm
long; upper pedun-
cles usually sparsely
to densely puberu-
lent, rarely moder-
ately glandular-pu-
berulent, .
.. 9b. P. microglossa
var. saxosa.

1. Perityle crassifolia Brandeg. Proc. Calif. Acad. II, 3:
147. 1891. TYPE: Mexico: BAJA CALIFORNIA: San Jose del
Cabo, 6 Oct. 1890, T. S. Brandegee (Holotype, UC; isotypes,
GH !, US(2) !).

la. Perityle crassifolia var. crassifolia.

Plants perennial, 10-75 cm high, younger plants with
slender taproots, older plants with fleshy-woody taproots,
younger plants erect and branching toward the top, older
plants branching at the base, tending to be decumbent,
stems pilose-villous to hirsute and glandular-pubescent;
leaves mostly alternate, felty to touch, arachnoid-villous to
short pilose-hirsute and glandular-pubescent, 2-3 (5) cm
long, 1.5-2.0 (3.0) em wide, reniform to cordate in outline,
thick and crisped, crenate to 3-5 lobed or dissected ; subses-
sile to petiolate, the petioles 0.5-2.0 em long; capitulescence
of 1-3 heads borne on peduncles 1-4 (6) em long; heads
radiate, ca. 1.0 em high, 1.0-1.5 em wide, involucres hemi-

1974] Perityle — Powell 249

spherical; bracts numerous, ovate-lanceolate; ray flowers
12-16, ligules white, 6-8 mm long often pinkish-tinged, ob-
long to elliptic or subspathulate; dise flowers numerous,
corollas yellow, 3.0-3.5 (4.0) mm long, throats tubular-
funnelform; achenes 2.5-3.0 (4.0) mm long, lanceolate-
obovate and curved, callous margins absent or very thin, the
margins densely ciliate, ray achenes 3-angled and pubescent
on faces, disc achenes 2-angled and glabrous on faces;
pappus of a short crown of squamellae and 1 (rarely 2)
slender, barbellate bristle, 2-3 mm long, the distal barbs
retrorse; chromosome number, n = 19.

Apparently restricted to deep coastal sand from Punta
Arena to San Jose del Cabo, Cape Region of Baja California
Sur, and Isla Coronados. Flowering year around. (Fig. 1).

REPRESENTATIVE SPECIMENS:

Mexico: BAJA CALIFORNIA SUR: Los Frailes, S of Cabo Pulmo,
Arnaud (ps); San Jose del Cabo, Brandegee (DS, GH, NY); SW end
of Isla Coronados, Carter 4274 (DS, SD, UC, US); Punta Frailes, Daw-
son 1140 (us); 3 mi N of Los Frailes, Hastings and Turner 64-279
(ARIZ) ; sand near shore, Isla Coronados, Moran 9121 (sp, vs); 0.5
mi E of Eureka, Powell and Turner 1849 (SRSC, TEX) ; San Jose del
Cabo, Purpus 444 (us) 274, 446 (vc); 0.6 mi S of Buena Vista,
Wiggins 14747 (CAS, DS, GH, TEX).

This taxon is best distinguished from var. robusta by
its arachnoid-villous pubescence, distinct perennial habit
with fleshy to woody taproots in older plants, and habitat
in coastal sand.

Some considerations suggest that var. crassifolia and var.
robusta should be accorded specific rank. Plants from one
collection of var. crassifolia (Powell and Turner 1849) and
several collections of var. robusta have been grown from
seed under identical greenhouse conditions. The character-
istie morphologies of both taxa persisted under artificial
conditions, but var. crassifolia flowered rarely while var.
robusta flowered profusely. Vigorous intervarietal hybrids
were obtained, but only two flowering heads were produced
on only one of several plants. Meiosis was regular and
pollen stainability was ca. 3%. The reproductive data are

250 Rhodora [Vol. 76

too meager to allow speculation about relationships. Also,
I have seen specimens of var. robusta (among borrowed
material) which approach var. crassifolia morphologically.
I believe that it is best to retain varietal status for the taxa,
following Everly (1947), pending populational study of the
habit and habitat of var. crassifolia.

1b. Perityle crassifolia var. robusta (Rydb.) Everly, Con-
trib, Dudley Herb. 3: 382. 1947.

Perityle robusta Rydb. N. Amer. Fl. 34: 16. 1914. TvPE:
Mexico: BAJA CALIFORNIA: Cerralvo Island, 19 April 1911,
J. N. Rose 16880 (Holotype, NY!; isotype, US!; isotype
fragments UC (2) !).

Perityle incompta Brandeg. Univ. Calif. Pub. Bot. 6:
508. 1919. TYPE: Mexico: BAJA CALIFORNIA : Los Dolores,
W. E. Bryant (uc!).

Perityle macromeres Blake, Proc. Biol. Soc. Wash. 37:
59. 1924. TYPE: Mexico: BAJA CALIFORNIA: La Paz 3 Feb
1906, E. W. Nelson and E. A. Goldman 1483 (us!).

Plants perennial or annual, with fleshy taproots to slen-
der branching roots, erect with few branches to spreading
in large clumps with many branches, stems short-hirsute to
glabrous; leaves densely short-hirsute and glandular-pu-
bescent to glabrous, 2-10 cm long, 1-7 cm wide, ovate to
cordate in outline, rather thick and crisped to thin and not
crisped, typically deeply 3-5 lobed with the lobes also in-
dented or with somewhat irregularly dissected margins;
heads 0.6-1.0 cm high, 0.6-1.5 em wide, involucres hemi-
spherical to campanulate; ligules 6-10 mm long; disc corol-
las 2-8 (4) mm long; pappus bristle usually 1, rarely 0-3;
chromosome number, n = 19, 18.

Growing in various soils, including those which are sandy
or saline near the sea and occasionally among rocks, rather
common in southern Baja California Sur and neighboring
islands. Flowering year around. (Fig. 1).

REPRESENTATIVE SPECIMENS:

Mexico: BAJA CALIFORNIA SUR: Isla Espiritu Santo, Berry (CAS);
Isla Magdalena, Brandegee (NY, UC, US); Isla Santa Margarita,

1974]

Perityle — Powell

a

~

Ln

Fig. 1. Distribution of P. crassifolia var. crassifolia (closed cir-
cles); P. crassifolia var. robusta (open circles); P. aurea (closed

squares); not plotted are P. socorrensis of the Revillagigedo Islands,
and P. incana of Guadalupe Island.

251

252 Rhodora [Vol. 76

Brandegee (UC, US); San Jose del Cabo, Brandegee (POM); El
Mogote Peninsula, La Paz Bay, Carter 2729 (CAS, DS, GH, UC,
us); 45 km N of El Refugio, Carter, Alexander, and Kellogg
2152 (DS, UC, US); Puerto Escondido, Carter and Kellogg 2875 (ARIZ,
DS, GH, SD, UC, US); Isla Partida, Collins, Kearney, and Kempton 166
(us); Isla San Francisco, Collins, Kearney, and Kempton 198 (vs);
10 mi W of Comondu, Gentry 4083 (ARIZ, DS, GH, UC); dunes, San
Nicholas Bay, Johnston 3720 (CAS, GH, UC, us); Isla Coronados,
Johnston 3756 (CAS, DS, GH, NY, UC, US); dunes, Isla Monserrate,
Johnston 3865 (CAS, GH, UC, US); beach, Agua Verde Bay, Johnston
3893 (CAS, GH, NY, UC, US) ; dunes, Isla San Francisco, Johnston 3946
(CAS, DS, GH, NY, UC, US) ; Isla Cerralvo, Johnston 4046 (CAS); Isla
Espiritu Santo, Johnston 4081 (CAS, GH, NY, UC, US) ; Guadalupe Point,
Concepcion Bay, Johnston 4150 (CAS, DS, NY, uc); E base of San
Lazaro, Santa Maria Bay, Moram 3537 (ps, UC); Ensenada de los
Muertos, Moran 3560 (ps, sD); S end of Isla Cerralvo, Moran 9616
(ns, UC); Isla San Jose, Moran 3751 (ps, vC); Isla San Marcos,
Moran 3948 (ps, UC) ; Isla Danzante, Moran 9209 (ps); W side of Isla
Catalina, Moran 9329 (sp, UC); NE side of Isla San Jose, Moran
9387 (sp); Isla Las Animas Rock, Moran 9433 (sp, vC, US); Isla
San Diego, Moran 9593 (sp); Isla Carmen, Palmer 1 (GH, NY, US);
10 mi N of Loreto, Powell and Sikes 1662 (SRSC, TEX); 2 mi NE of
La Paz, Powell and Sikes 1681 (SRSC, TEX); 33 mi W of San Javier,
Powell and Turner 1847 (sRSC, TEX); Isla Magdalena, Rose 16319
(Ny, US); Isla Pichilinque, Rose 16529 (GH, NY, US); 8 mi W of
San Miguel, Shreve 7125 (ARIZ, DS, GH, us); 17 mi N of La Paz, Sikes
and Babcock 270 (SRSC, TEX); 2 mi S of Villa Constitucion, Sikes and
Babcock 276 (SRSC, TEX); ca. 29 mi N of Villa Constitucion, Thomas
8403 (CAS, DS, GH, US); 9 mi E of San Ignacio Wiggins 11360 (CAS,
DS, GH, UC, US); San Gregoria, 12 mi NW of La Purisima, Wiggins
11467 (ps, GH, UC); 0.6 mi S of Buena Vista, Wiggins 14748 (CAS,
DS, TEX, UC); Salino Flat, near S end of Isla Espiritu Santo, Wiggins
15595 (ps); Isla Partida, Wiggins 16160A (Ds).

Perityle crassifolia var. robusta is more widespread than
is var. crassifolia, and occupies a variety of edaphic condi-
tions in coastal, insular, and inland areas in the southern
portion of Baja California Sur. Two extreme forms of the
morphologically variable var. robusta can be recognized.
The plants of coastal habitats (e.g., near La Paz) usually
are robust with fleshy, perennial taproots, while plants of
the inland forms (e.g., near Commondu) are typically
smaller with annual or weak perennial habits. Also, the
inland form tends to be less pubescent and has thinner, less

1974] Perityle — Powell 253

crisped leaves than coastal plants. According to my inter-
pretation, the “inland form” corresponds to P. incompta
which Everly (1947) recognized as a distinct species. The
“coastal form” conforms with Everly’s P. crassifolia var.
robusta. Although the extremes of coastal and inland forms
are evident, the existence of separate taxonomic entities
has not been indicated. Instead, examination of exsiccata
material and field studies have shown a rather complete
morphological intergradation (e.g. Johnston 3946) between
the two extremes. Hence P. incompta is treated here as
synonymous with var. robusta.

Further study is needed to clarify the adaptable habit of
var. robusta, i.e., the perennial vs. annual conditions. Also,
particular attention should be given to the habitats in which
life forms occur. Those plants at La Paz grow in crusty
saline soils, while plants of other populations may be in
crevices of granite rocks, in rocky soil, or in fine sand.

Another aspect of variability in var. robusta is presence
or absence of pappus bristles, which seemingly is not taxo-
nomically significant. As a generality, however, awnless
forms occur on islands while awned forms are peninsular,
but there are exceptions.

2. Perityle socorrensis Rose, Bot. Gaz. 15: 118. 1890.
TYPE: Mexico: Socorro Island, Revillagigedo Islands, Mar.
1889, C. H. Townsend (Holotype, Us!; isotypes, GH !, NY !,
US!).

Plants suffruticose perennials, 10-40 cm high, many
branched and densely leafy; leaves typically alternate,
densely short-hirsute and glandular-pubescent, semisuccu-
lent, 2-6 cm long, 0.8-3.0 cm wide, ovate, deltoid-ovate, cor-
date to subhastate, 3-5 shallow-lobed and serrate to crenate ;
petioles 1.5-3.0 cm long; capitulescence of 1-2 (3-5) heads
borne on peduncles 1.0-3.5 (6.0) cm long; heads typically
radiate, rarely discoid, ca. 6 mm high, 7-10 mm wide, in-
volucres campanulate to narrowly so; bracts lanceolate to
oblanceolate or oblong-lanceolate; ray flowers ca. 10 or
fewer, ligules white, 2-4 mm long, oblong to oblong-elliptic,
rarely with a prominent inner lobelet; disc corollas yellow

254 Rhodora [Vol. 76

(?), 2.0-2.5 mm long, throats tubular-funnelform ; achenes
2.9.8.0 mm long, oblanceolate-obiconical, oblong-oblanceo-
late, to narrowly obconical typically curved, callous margins
absent or very thin, the margins ciliate, ray achenes
sparsely and minutely pubescent on faces, disc achenes
glabrous or nearly so; pappus of a short to vestigial crown
of squamellae and typically 2 (0-1) bristles, 1.5-2.0 mm
long, the bristles retrorse barbellate (rarely lateral or an-
trorse) distally; chromosome number, n = 19.

Endemic to the Revillagigedo Islands, growing on sea
cliffs and in soils near the shore. Flowering mostly in
winter and spring.

REPRESENTATIVE SPECIMENS:

Mexico: REVILLAGIGEDO ISLANDS: Isla San Benedicto: Anthony 372
(CAS, DS, GH, NY, UC, US); Barkelew 175 (ARIZ, DS, GH, NY, POM, UC,
us); Mason 1684 (cas, GH, UC, US); Isla Clarion: Anthony 415
(CAS, DS, POM, UC, US); Sulfur Bay, Howell 8347 (CAS, DS, NY, POM,
us); W end of island, Mason 1578 (CAS, DS, GH, NY, POM, UC, US);
Isla Socorro: Anthony 383 (ARIZ, CAS, DS, GH, POM, SD, UC, US);
Barkelew 192 (ARIZ, DS, GH, NY, POM, UC, US); Academy Bay, Carl-
quist 368 (CAS, RSA, UC); Brenner's Cove, Howell 8423 (CAS, DS, GH,
uc, US); Graysons Cove, Moran 5922 (ARIZ, CAS, DS, GH, NY, RSA, SD,
UC, US).

Perityle socorrensis is a distinct species closely related to
P. crassifolia from which it is distinguished by its typically
2 pappus bristles, reduced squamellae, generally smaller
floral features, short ligules (or absent), woody bases, leaf
morphology, and its distribution.

As discussed by Everly (1947), Perityle socorrensis is
notably variable in ligule and pappus features. The ligules,
typically short at 2-4 mm, are even shorter and aberrant-
looking in many collections, while other specimens are
without ligules. According to label data, discoid individuals
may occur in populations with radiate forms, and thus no
taxonomic significance is attributed to ligule variability.
Typically, the number of pappus bristles per achene in P.
socorrensis is two, but some individuals or populations may
have only one bristle or none. Bristle number is correlated

1974] Perityle — Powell 255

with the island to island distribution: plants with two bris-
tles on Socorro and San Benedicto Islands, and awnless
forms on Clarion Island. No taxonomic significance is at-
tached to bristle variability since exceptions are found on
all the islands, and because other features are not corre-
lated with the pappus differences.

3. Perityle emoryi Torr. in Emory, Notes Mil. Rec. 142.
1848. TYPE: California: mountains E of San Diego, 29 Nov.
1846, Emory (NY!).

Perityle nuda Torr. Pacif. R. Rep. 4: 100. 1857. TYPE:
California: Williams’ River, 7 Feb. 1853-4, J. M. Bigelow
(NY!).

Perityle emoryi var. nuda A. Gray, Bot. Calif. 1: 397.
1876.

Perityle emoryi S. Wats. Proc. Amer. Acad. 11: 116. 1876.
Not P. emoryi Torr. 1848.

Perityle californica A. Gray, Syn. Fl. N. Amer. 1: 321.
1884. Not P. californica Benth. 1844.

Perityle californica var. nuda A. Gray, Syn. Fl. N. Amer.
1: 321. 1884.

Perityle fitchii var. palmeri A. Gray, Syn. Fl. N. Amer. 1:
321. 1884.

Perityle fitehii Green, Bull. Calif. Acad. 2: 403. 1887.
Not P. fitchii Torr. 1857.

Perityle californica Vasey, Proc. U, S. Nat. Mus. 11: 368.
1889. Not P. californica Benth. 1844.

Perityle rothrockii Rose, Bot. Gaz. 15: 114. 1890. TYPE:
Nevada: 1872, Wheeler (us!).

Perityle greenei Rose, Bot. Gaz. 15: 117. 1890. TYPE:
California: Santa Cruz Isl, July-Aug. 1886, E. L. Green
(Holotype, ND?; isotypes, DS!, NY!, UC!, US!).

Perityle emoryi var. orcuttii Rose, Bot. Gaz. 15: 117.
1890. TYPE: Mexico: BAJA CALIFORNIA. Canyon Cambel-
los(?) July, 1884, C. R. Orcutt (Holotype, US!; isotype
GH!).

Perityle grayi Rose, Bot, Gaz. 15: 118. 1890. TYPE:
Mexico: Guadalupe Isl., 1875, E. Palmer 44 (Holotype, Us?;
isotypes, GH !, NY!).

~
"T

256 Rhodora [Vol. 7

IN

Laphamia nuda Benth & Hook. ex Jacks. Ind. Kew.
30. 1895.

Laphamia emoryi Benth & Hook, ex J acks. Ind. Kew. 2:
30. 1895. Partial Synonomy of South American P. emoryi
(= Closia).

Closia elata Phil Fl. Atac. 31 and Viage Des. Atac. 19,
205. 1860. Type not examined.

Closia discoidea Phil. Fl. Atac. 31 and Viage Des. Atac.
205. 1860. Type not examined.

Perityle emoryi Torr. var. elata. (Phil.) I. M. Johnston,
Contr. Gray Herb. 85: 127. 1929.

Perityle discoidea (Phil) I. M. Johnston, Contr. Gray
Herb. 85: 128. 1929.

Plants delicate or robust annuals, 2-60 cm high, usually
herbaceous or the lower stems woody, erect or spreading
with few to many stems, puberulent to hirsute and glandu-
lar-pubescent; leaves mostly alternate, hirsute to glandular-
pubescent, 2-10 cm long 1-5 cm wide, ovate, cordate, sub-
orbieular, to triangular in outline, the margins deeply
toothed, lobed, cleft, or divided, with the segments also
indented to irregularly dissected; petioles 0.3-4.0 cm long ;
capitulescence of 1 to many heads borne on peduncles 0.1-
7.0 em long ; heads radiate (rarely discoid), 0.4-1.0 cm high,
0.4-1.0 em wide; involucres hemispherical or campanulate;
bracts numerous, lanceolate, oblanceolate, to ovate-lanceo-
late; ray flowers 8-12 (14), ligules white, usually oblong,
1.5-4.0 (6.0) mm long, 1.5-3.0 mm wide, rarely rudiment-
ary; disc flowers numerous, corollas yellow, 2.0-2.5 (3.8)
mm long, throats tubular to tubular-funnelform; achenes
(1.5) 2-3 mm long, suboblong, oblanceolate, or subcuneate,
the outer ones often curved, margins thin, not calloused,
the margins long- or short-pubescent (ciliate), outer
achenes 2-3 (4) angled and often pubescent on the faces,
inner achenes 2-angled with glabrous (rarely puberulent)
faces; pappus of a vestigial or conspicuous crown of squa-
mellae and 1 slender bristle, (0.8) 1.0-2.5 (3.5) mm long,
antrorse-, lateral-, or retrorse-barbed especially at the tip.

1974] Perityle — Powell 257

Fig. 2. Generalized distribution of P. emoryt (stippled); South
American distribution in Chile and Peru not plotted.

258 Rhodora [Vol. 76

or the bristle absent; chromosome number, tetraploid (n =
32-36) or hexaploid (n = 50-58).

Widespread weed of desert southwestern United States,
Sonora, Mexico, Baja Californian peninsula, and neighbor-
ing islands; continental disjunct in Chile and Peru. Fower-
ing mostly in winter and spring, but also year around, de-
pending upon latitude. (Fig. 2).

REPRESENTATIVE SPECIMENS:

MEXICO: Baja California: Ensenada, Anthony 180 (CAS, Ds, UC,
us); 8.2 mi S of Socorro, Constance 3119 (DS, GH, LL, UC, US);
Bahia de los Angeles, Cowan 2265 (TEX); 14 mi S of Santa Rosa-
lillita, Powell and Sikes 1647 (SRSC, TEX); 41.6 mi S of Mexicali,
Powell and Turner 1710 (SRSC, TEX); Arroyo Calmalli, Purpus 23
(CAS, NY, POM); 14 km NW of Colonia Guerrero, Raven, Lewis, and
Thompson 12193 (GH); 8.5 km E of El Rosario, Raven, Mathias, and
Turner 12433 (ARIZ, GH); 15.9 mi N of San Felipe, Raven 14775
(ps, uc); N side of El Arco, Sikes and Babcock 295 (SRSC, TEX);
2 mi N of Miission de San Borja, Wiggins and Wiggins 14851 (DS);
9 mi S of Puertocito, Wiggins and Wiggins 15877 (Ds, US). Baja
California Sur: Sierra de la Trinidad, Cape Region, Brandegee (UC,
US); near Santa Rosalia, Ferris 8702 (Ds); Santo Domingo, Gander
9792 (sp); 10 mi W of Comondu, Gentry 4083 (ARIZ, UC); San
Francisquito Bay, Johnston 3566 (CAS, GH, US); Concepcion Bay,
Johnston 4150 (GH, US); E base of San Lazaro, Santa Maria Bay,
Moran 3537 (sp); 18.6 mi. S of Mulege, Powell and Turner 1836
(SRSC, TEX); ca. 20 mi N of Santa Rosalia, Sikes and Babcock 285
(SRSC, TEX); 8 mi NW of San Ignacio, Sikes and Babcock 288 (SRSC,
TEX); San Ignacio, Wiggins 16233 (ps); 3 mi N of El Barril, Wig-
gins 16864 (ns); Mulege, Wiggins and Wiggins 18070 (us); 4 mi
S of El Arco, Wiggins and Wiggins 18191 (CAS, DS). Islands: ISLA
ANGEL DE LA GAURDA: Wiggins 17010 (ps). ISLA CARMEN: Moran
9181 (sp). ISLA CEDROS: Anthony 283 (CAS, DS, GH, NY, POM, UC, US);
Haines and Hale (CAS, GH, LL, NY, SD, UC, US); Howell 10684 (CAS,
DS, GH, NY, POM, US). ISLA CORONADOS: Moran 6556 (sD); Palmer 16
(GH, NY, US). ISLA GUADALUPE: Anthony 233 (CAS, DS, GH, UC, US);
Carlquist 489 (RSA); Mason 1502 (CAs, GH, US); Moran 2900 (DS,
GH), 5628 (DS, RSA, SD, UC); Palmer 891 (Ny, UC, US); Rose 16003
(GH, NY, US). ISLA ILDEFONSO: Moran 9062 (sp); Wiggins 18254
(CAS, DS). ISLA NATIVIDAD: Brandegee (UC); Moran 10797 (sp).
ISLA PARTIDA: Johnston 3235 (CAS, US); Wiggins 17268 (DS). ISLA
PARTOS: Johnston 3240 (CAS, GH, NY, UC, US). ISLAS REVILLAGIGEDO:
Mason (CAS, US). ISLA SALSIPUEDES: Moran 8871 (SD, US). ISLA SAN
BENITO: Anthony 271 (DS, GH, POM, UC); Palmer 914 (ARIZ, CAS,

1974] Perityle — Powell 259

DS, NY, US). ISLA SAN ESTEBAN: Wiggins 17218 (DS). ISLA SAN
LORENZO: Moran 8895 (SD). ISLA SAN MARCOS: Johnston 3622 (CAS,
GH, UC, US). ISLA SAN PEDRO MARTIR: Johnston 3148 (CAS, GH, UC,
us); Moran 8810 (SD). ISLA TIBURON: Johnston 3268 (CAS, us).
ISLA TODOS SANTOS: Moran 16214 (sD). ISLA TORTUGA: Wiggins
17367 (ps). Sonora: Puerto Penasco, Breedlove 1397 (ns); 27.4 mi
S of Sonoyta, Breedlove 1389 (Ds, TEX); vicinity of Libertad, Graham
3803 (ps); 3 mi W of Kino, Heckard 1575 (JEPS); 1 mi E of San
Carlos Resort, Powell and Sikes 1685 (SRSC, TEX); Empalme, Rose,
Standley, and Russell 12636 (GH, NY, us); 5 mi NW of Caborea,
Shreve 1532 (ARIZ); 4 mi NW of Caborea, Wiggins 8233 (DS, GH,
UC, US).

U.S.A.: Arizona: COCONINO CO.: Havasu Canyon, Grand Canyon,
Howell 26537 (ARIZ, CAS, SMU); GILA CO.: Roosevelt Dam, Eastwood
6247A (CAS). MARICOPA CO.: Phoenix, Eastwood 6159 (CAS); near
Tempe, Gillespie 8808 (DS, GH, POM, UC, us); 2 mi NW of Scottsdale,
Russell 10911 (SMU, UC). MOHAVE CO.: 7 mi NW of Alamo, Benson
10074 (pom); 3.4 mi SE of Boulder Dam, Carter and Chisaki 3230
(ARIZ, DS, LL, NY, RSA, SMU, UC, US) ; Toroweap, Grand Canyon, Cottam
13001 (UT); 62 mi N of Wickenburg, Sikes 106 (SRSC). PIMA CO.:
10 mi W of Bates Well, Growler Mts., Benson 9926 (PoM); Gunsight
Peak, Fosberg 7865 (POM); ca. 5 mi N of Ajo, Gould and. Macbride
4130 (ARIZ, GH, NY, UC); Alamo Canyon, Ajo Mts, Huey 24351
(cH, sp); Dripping Spring, Organ Cactus Natl. Mon. McClintock
52-37 (CAS). PINAL CO.: Sacaton, Gilman 350 (ARIZ); 10 mi W of
Maricopa, Russell 11348 (smu). YUMA CO.: Gila Mts., near US 80,
Barr and Lange 64-174 (ARIZ); S end of Castle Dome Mts., Parker,
Wright and Lowe 7789 (ARIZ, DS, NY, RSA, us); 15.6 mi E of Yuma,
Powell and Turner 1704 (SESC, TEX); S end of Cunningham Pass,
Harcuvar Mts., Wiggins 8452 (ps, GH, UC, us). California: IMPERIAL
co.: 8 mi from Niland to Blythe, Balls 12921 (RSA); 7 mi N of
All American Canal spillway, Wiggins 8613 (DS, NY, POM, UC, US).
Inyo co.: Furnace Creek, Death Valley, Carpenter (JEPS) ; Funeral
Mts., Coville and Funston 324 (us); Hanaupah Canyon, Coville and
Gilman 392 (vs); Darwin Falls, Hitchcock 6219 (uc); Surprise
Canyon, Howell 3964 (CAS); Panamint Valley Smith 86 (JEPS).
KERN CO.: Last Chance Canyon, El Paso Range, Twisselman 11863
(JEPS). LOS ANGELES CO.: Santa Catalina Isl., Brandegee (UC);
Santa Monica Mts., Epling (DS, NY, RSA); Anacapa Isl., Howell 3816
(JEPS); San Clemente Isl., Raven 17345 (RSA, UC). RIVERSIDE CO.:
Palm Springs, Abrams 11012 (ps); San Jacinto Range, Benson 4167
(pom): 8 mi NE of Desert Center, Wiggins 9675 (DS, GH. RSA, UC).
SAN BERNARDINO CO.: 29 Palms, Alexander and Kellogg 870 (UC);
near Parker Dam, Brenckle 51140 (SMU, uc); 89 mi N of Needles,
Ferris 7224 (ps); between Kelso and Baker, Jepson 20590 (JEPS).
SAN DIEGO CO.: 23.1 mi NW of Coyote Wells, Breedlove 1856 (DS);

260 Rhodora [Vol. 76

Yaqui Wells, Eastwood 2644 (CAS, GH, UC, US); Jacumba, Nelson
11182A (DS, GH, NY, POM, UC); Borego Park, Wolf 8462 (ARIZ, au,
NY, RSA). SANTA BARBARA CO.: Santa Cruz Isl., Balls and Blakley
23727 (RSA, UC). VENTURA CO.: Point Mugu, Howell 3733 (cas).
Nevada: CLARK COo.: between Las Vegas and Boulder Beach, Cron-
quist 9844 (NY, UC); 1 mi below Boulder Dam, Grater 51 (vc); 8 mi
SW of Davis Dam, Gullion 258 (Uc).

CHILE: Atacama: Chanaral, Beetle 26164 (GH, UC); vicinity of
Caldera, Gigoux (GH); below Agua EI Huerto, Johnston 3678 (vs);
vicinity of Potrerillos, Johnston 4740 (GH, US); vicinity of Puerto
de Chanaral, Johnston 4794 (GH, US); vicinity of Copiapo, Johnston
5028 (GH); Caldera, Johnston 5056 (GH); vicinity of Caleta Pan de
Azucar, Johnston 5829 (Us); Vallenar, Werdermann 160 (GH);
Tierra Amarilla, Werdermann 406 (GH, NY); Quebrada Paipote,
Werdermann 448 (GH, NY). Antofagasta: Taltal, Jaffuel 984 (GH):
Tocopilla, Jaffuel 1014 (GH); Antofagasta, Jaffuel 1126 (GH); 6 km
N of Puerto Tocopilla, Johnston 3585 (Us).

ECUADOR: Galapagos Isles: Cerros Isl, Stewart 40 (cas).

PERU: Mts. near Chosica, Lima-Oroya Railroad, Weberbauer
5820 (GH, US).

The morphological variation of Perityle emoryi is at-
tested to by its considerable synonymy. This widespread
annual, polyploid weed exhibits variation in nearly all as-
pects of plant form. I have carefully examined the bulky
exsiccatae available to me, giving particular attention to
plant size, leaf morphology, head size, presence or absence
of ligules and pappus bristles, and geographic distributions.
None of the variable morphological aspects appear to have
populational significance, and thus, in my judgement, do
not require taxonomic recognition. I do note, however, that
several atypical collections of P. emoryi from Magdalena
Island resemble P. crassifolia and Amauria brandegeana,
and thus the desirability of further study of Perityle from
this locality is indicated.

Perityle emoryi is related to and probably derived from
the diploid P. crassifolia var. robusta, from which it is
delimited by habit, smaller heads, shorter ligules, usually
Shorter disc corollas, and chromosome number. With poorly
preserved specimens it is often difficult to distinguish these
taxa, but I have not had difficulty recognizing them in the
field. Perityle emoryi is also remarkably similar to Amauria

1974] Perityle — Powell 261

brandegeana in superficial morphology. A discussion of the
later similarity and distinguishing traits is to be found
elsewhere (Powell, 1972a).

It is assumed that P. emoryi achieved wide distribution
as a result of the increased adaptability provided through
a combination of polyploidy, reproductive self-compatibility
and annual habit. That polyploids often exceed their di-
ploids in distribution is well-known (Stebbins, 1950), and
the advantage of self-compatibility in the establishment of
disjunct colonies is clear. Indeed, Raven (1963) has sug-
gested that P. emoryi might have attained bicontinental
distribution as a result of long-distance dispersal from the
Sonoran Desert, and he indicated the advantage of self-
compatibility to such long-distance dispersal. More spe-
cifically, I believe that the South American P. emoryt orig-
inated from the vicinity of southern Baja California. It is
in southern Baja California that the ancestors of P. emoryt
are found. Furthermore, geographic variation in length of
pollen spines (the spines ornamenting pollen walls) sug-
gests that South American and Mexican P. emoryi had
common areal origin. In measuring the pollen spines of
some 20 populations of P. emoryi, it was found that the
spine length of South American populations averaged the
same or slightly longer than that of Mexican populations,
while the spine length of United States populations was
nearly twice as short as the others (Powell & Miller, un-
published). From this information it can be postulated
that P. emoryi originated and became established in Baja
California, migrated to South America by long-distance
dispersal in late Pliocene or Pleistocene (Raven, 1963), and
subsequently advanced to the north, occupying desert areas
in southwestern United States.

Further study of the South American Perityle emoryi is
needed, particularly chromosomal analyses and living plant
comparisons. At least two forms can be recognized among
the South American specimens I have seen, and this was
discussed to some extent by Johnston (1929). One form
(= P. discoidea (Phil.) I. M. Johnst.) deviates from North

262 Rhodora [Vol. 76

American P. emoryi in smaller habit, numerous, smaller
and tightly clustered heads with no conspicuous ligules, and
a pappus bristle on the achenes. Another form (= P.
emoryi var. elata (Phil.) I. M. J ohnst.) is variable in habit,
has large and small heads, longer and short ligules, and
awnless achenes. I have not been able to make taxonomic
distinctions between the South American and North Amer-
ican forms, pending further study, and thus have placed
Johnston’s combinations in Synonymy. It should be noted
that only a partial synonymy of South American P. emoryi
(Closia, in part?) is included since I have not seen speci-
mens or types upon which several other Closia names are
based. A few other collections from South America closely
resemble P. crassifolia (e.g., Johnston 5023) or Amauria
brandegeana (e.g., Werdermann 160; Morong 93). As
mentioned above, these similarities are also seen in a few
Mexican specimens. But it is also possible that both P.
crassifolia and Amauria brandegeana once made their way
to South America even though they have not become estab-
lished there.

4. Perityle aurea Rose, Contr. U. S. Nat. Herb. 1: 84. 1890.
TYPE: Mexico: BAJA CALIFORNIA: Santa Rosalia, 24 Feb.-
3 Mar. 1889, E. Palmer 185a (Holotype, Us!; isotypes, CAS !,
GH !, NY!).

Plants annual, 10-60 cm high, erect or spreading, stems
of larger plants succulent, reddish, nearly glabrous; leaves
alternate, puberulent to glabrous, slightly thick in texture,
1.7-9.0 em long, 1-4 cm wide, subreniform to ovate in out-
line, basically 3-lobed with dentate-serrate lobes and mar-
gins; petioles 0.6-5.0 em long; capitulescence of 1-3 heads
borne on rather short peduncles 1-2 (3) em long; heads
radiate, 5-8 mm high, 5-8 mm wide, involucres campanulate
to narrowly so; bracts broadly lanceolate; ray flowers ca.
10, ligules yellow, oblong-elliptie to obovate; dise corollas
yellow, 2-3 mm long, throats tubular to funnelform; achenes
2-3 mm long, narrowly oblanceolate to suboblong, with thin
callous margins, the margins densely white-ciliate, ray
achenes pubescent on faces, disc achenes glabrous or nearly

1974] Perityle — Powell 263

so on faces, pappus of a conspicuous crown of squamellae
and 1 bristle (rarely 0) 1.5-2.5 mm long, the bristle retrorse
barbelate with few barbs at tip; chromosome number,
n= M.

Restricted in soils and among rocks, near Santa Rosalia
of Baja California Sur and a few islands in the Sea of
Cortez. Flowering Jan-April. (Fig. i9»

SPECIMENS EXAMINED:

Mexico: BAJA CALIFORNIA: summit and N slope of high peak near
SE corner of Isla San Esteban, Moran 8845 (sD, US); summit of NE
peak, Isla San Esteban, Moran 13044 (sb); arroyo on S Isla San
Lorenzo, Moran 13060 (SD). BAJA CALIFORNIA SUR: Isla San Marcos,
Johnston 3614 (CAS, GH, UC, US); Isla San Marcos, Moran 3965
(ps); Santa Rosalia, Palmer 185a, 185 (CAS, GH, us); 8 mi W of
Santa Rosalia, Powell and Turner 1826, 1829 (SRSC, TEX); 9 mi N
of Santa Rosalia, Powell and Twrner 22219 (SRSC, TEX); 10 mi N
of Santa Rosalia, Reed 6225 (ps, POM) ; ca 12 mi N of Santa Rosalia,
Sikes and Babcock 281 (SRSC, TEX).

Two growth forms of Perityle aurea are evident in the
field, and both have been preserved in existing collections
of the species. One form is of small plants, superficially
resembling the Baja California populations of P. californ-
ica, and the other form is of taller, more robust and succu-
lent plants resembling P. crassifolia. The small form is
known to occur in roadside soils where dry conditions pre-
vail, and the large form occurs in places that receive and
retain more water. The growth forms likely are ecologically
controlled. I have seen both forms growing only a few
paces apart, and when seeds of both plant sizes were grown
under identical greenhouse conditions, only the larger, more
succulent forms developed. The island specimens of P.
aurea differ slightly from the mainland forms, most notably
in achene morphology and in being awnless, although awned
forms also occur on San Esteban Island.

Perityle aurea seemingly is related to P. crassifolia, from
which it is readily delimited by yellow ligules, chromosome
number (n = 17), and distribution. An affinity of P. aurea
with the P. crassifolia alliance is not indicated by its yellow

264 Rhodora [Vol. 76

ligules and chromosome number. All other members of the
P. crassifolia group have white ligules and have x — 19.
My first impression from superficial morphological exam-
inations was that the species was related to P. californica
(n = 13, 12, 11) and P. cuneata (n = 16, 12), both annual
yellow-rayed taxa with habits similar to the small growth
form of P. aurea. But P. aurea exhibits a combination of
vegetative and floral features which characterize only the
P. crassifolia alliance: robust and rather succulent habit,
achenes with callous margins absent or thin, other unique
achene morphology, and pappus bristles retrorsely barbed
at the tips. Also, chromatographic studies have shown that
P. aurea, like members of the P. crassifolia group, pro-
duces the abundant yellow flavonoid compounds that have
been detected elsewhere in sect. PERITYLE only in the P.
parryi group of Texas. I conclude that P. aurea evolved,
probably from P. crassifolia var. robusta, through specia-
tion that involved adaptation of the annual habit and aneu-
ploid reduction in chromosome number. Artifieial hybrid-
izations have strongly suggested that the above taxa are
reproductively isolated (Powell, 1972c).

5. Perityle incana Gray, Proc. Amer, Acad. 11: 78. 1876.
TYPE: Mexico: BAJA CALIFORNIA: Guadalupe Island, Feb.-
May, 1875, E. Palmer 43 (Holotype, GH!; isotypes, NY,
US!),

Nesothamnus incanus (Gray) Rydb. N. Amer. Fl. 34: 12.
1914.

Plants shrubby, 10-40 (80) em high, many branched and
densely leafy, stems tomentulose-canescent; leaves alter-
nate, thickish, tomentulose-canescent, 6-14 em long, 4-7 em
wide, deeply 3-divided or palmate, the divisions again
deeply lobed or cleft; petioles 1.5-7.0 em long; capitules-
cence of many tightly clustered heads (naked corymbs)
borne on short peduncles; heads discoid, 5-7 mm high, 4-8
mm wide, involucres campanulate; bracts broadly oblong-
lanceolate to broadly linear; disc corollas yellow, 2-3 mm
long, throats narrowly campanulate or broadly tubular;

1974] Perityle — Powell 265

achenes 2.0-3.8 mm long, narrowly obdeltoid to narrowly
oblanceolate, with prominent callous margins, the margins
coarse-ciliate, the faces coarse-pubescent ; pappus crown of
2 broad, laciniate squamellae, 0.5-0.9 mm long, and rarely a
narrow, fimbriate scale, 1.0-1.2 mm long; chromosome num-
ber, n = ca. 50-57.

Endemic to Guadalupe Island and islets, growing on cliffs
near the sea. Flowering winter, spring, and early summer.

REPRESENTATIVE SPECIMENS:

MEXICO: Baja California: ISLA GUADALUPE: Anthony 249 (DS);
Carlquist 440 (RSA, uc); Barracks Cove, Copp 157 (DS) ; Franceschi
7 (A, DS, POM, UC); Howell 8172 (CAS); Outer Islet, Lindsay 2621
(sp); Islote Negro, Mason 1519 (ARIZ, CAS, DS, us); Moran 2904
(ps, RSA); Outer Islet, Moran 5682 (CAS, DS, NY, SD, uc); North
Twin Canyon, Moran 6133 (sp); N end of island, Moran 6435 (DS,
sp); cliffs of the Lower Circus, Moran 12023 (sp); N side of Mt.
Augusta, Moran 12042 (DS, SD, uc); SW Oak Canyon, Moran 13792
(SD).

This cliff-dwelling Guadalupe Island endemic is perhaps
the most distinct species of sect. PERITYLE. Its identifying
features include: shrubby habit; tomentulose-canescent
pubescence ; clustered, naked capitulescence; coarse pubes-
cenct achenes; pappus of broad, laciniate squamellae and
rarely à narrow, fimbriate scale. Considering its relatively
large, shrubby habit, unique character, and isolated distri-
bution, Perityle incana would seem to be a primitive mem-
ber of the genus, although this possibility is not strength-
ened by its hexaploid chromosome number. According to
Reid Moran (personal communication), the taxon is not in
danger of extinction, largely because its habitat is out of
reach of the goats which have overrun the island for more
than a century.

It is clear that Perityle incana does not belong with any
of the related-species groups, and thus stands as an anoma-
lous species. In fact, the species could be recognized as a
montotypic genus with little change in the taxonomic nat-
uralness of the subtribe. I suspect, however, that the taxon
is distantly related to the P. crassifolia group and is best

266 Rhodora [Vol. 76

treated as a member of Sect. PERITYLE. Perityle incana is
similar to Pericome in habit and capitulescence characters,
and perhaps shares ancestral affinity with that genus.

6. Perityle turneri Powell, Madrofio 21: 456-457. 1972.
TYPE: Mexico: DURANGO: 3.4 mi E of Ey Palmito, 2 Apr.
1970, A. M. Powell 1858 (Holotype, TEX ! ; isotype, SRSC!),

Plants herbaceous to suffrutescent perennials, decumbent,
prostrate, or semierect, stems Spreading 20-45 em long,
often purplish, lower stems often rooting at the nodes,
upper stems puberulent to densely so, short-pilose, or sub-
tomentose; leaves mostly opposite, puberulent to densely so,
short-pilose to densely so, or subtomentose, often purplish,
3.0-4.5 (8.5) em long, 1.5-3.0 (4.0) em wide, ovate, deltoid,
or subcordate, the apexes acute or attenuate, the Margins
serrate, serrate-crenate, Serrate-lobed to doubly so, the ser-
rations or lobes acute or acuminate; petioles 1.0-1.5 (3.0)
cm long; capitulescence of 1-3 (4) heads on peduncles (1.0)
2-6 cm long; heads radiate, subglobose, 5-7 mm high, 0.7-1.4
cm wide, involucres hemispherical to broadly campanulate;
receptacles conical; ray flowers ca. 12-18, ligules white, (3)
6-9 mm long, oblong; disc corollas yellow, (1.8) 2.0-2.8
(3.0) mm long, throats short-campanulate, short-funnel-
form, rarely narrowly tubular-funnelform ; achenes 1.5-2.0
mm long, obovate to oblong-ovate, with thin or prominent
callous margins, the margins ciliate, the faces puberulent in
the center; pappus of a prominent crown of squamellae and
2 (rarely 1) unequal bristles, the longest 0.6-1.5 mm long;
chromosome numbers n — 17,n — 17 II 4 I.

Moist seeps and canyons, higher elevations to ca. 9500
feet, southern Chihuahua and Durango in the Sierra Madre
Occidental. Flowering spring and fall. (Fig. 3).

REPRESENTATIVE SPECIMENS:

Mexico: CHIHUAHUA, 4 mi SW of Villa Matamoros, Correll and
Gentry 22819 (LL); near La Rocha, NE slope of Sierra Mohinora,
Correll and Gentry 23109 (LL); Burro Canyon near Parral, Pringle
18650 (ARIZ, Cas, GH, SMU, TEX, UC, US). Durango. Quebrada San
Juan, ca. 50 mi W of Durango and 23 mi NW of Los Coyctes RR,

1974] Perityle — Powell 267

Fig. 3. Distribution of P. turneri (closed squares) ; P. lineariloba
(closed star); P. microcephala (closed triangles) ; P. microglossa var.
microglossa (closed circles); P. microglossa var. saxosa (open tri-
angles); P. ciliata (open squares); P. coronopifolia (open circles) ;
P. hofmeisteria (open star); not plotted is P. canescens.

268 Rhodora [Vol. 76

Cronquist 9568 (NY, SMU, TEX, US); 2 km S of El Espinozo del
Diablo, and 65 km SW of El Salto, Cronquist and Fay 10786 (NY);
6 mi W of La Ciudad, Flyr 273 (TEX); ca. 35 mi W of El Salto,
Gentry and Arguelles 18210 (Us) ; San Ramon, Palmer 60 (GH, NY,
UC, US); 2.2 mi E of El Palmito, Powell and Turner 1857 (SRSC,
TEX); 12 mi W of La Ciudad, Sikes and Babcock 380 (SRSC, TEX).
SINALOA. 0.9 mi W of El Palmita, ca. 47 mi E of Concordia, Breed-
love 1721 (Ds).

Plants of this remarkable species were rarely collected
until part of its high-altitude distributional range was made
accessible by the road cut across the Sierra Madre Occiden-
tal from Durango to Mazatlan. At the type locality near
Mex. 40, Perityle turneri is abundant in wet mats formed
by dense growths of Selaginella and small ferns. In overall
aspect, at least at the type locality, the taxon shows striking
resemblance to members of the tribe Astereae.

Perityle turneri is related to P. lineariloba, from which
it is easily delimited by leaf shape, and to P. microcephala,
from which it can be distinguished by its capitulescence and
head size. Further discussion concerning the taxonomy of
this species is available elsewhere (Powell, 1972e).

7. Perityle lineariloba Rydb. N. Amer. Fl. 34: 18. 1914.
TYPE: Mexico: DURANGO: San Ramon, 21 Apr.-18 May 1906,
E. Palmer 89 (Holotype, NY!; isotypes, GH!, UC!, US!).
Plants seemingly perennial, with semi-annual roots, stems
lignescent near the base, decumbent, ca. 30 cm long, grayish-
puberulent above; leaves mostly opposite, finely puberulent,
2.5-6.0 em long, 1.5-3.5 cm wide, pinnately 3-5 divided, the
divisions linear or nearly so and entire or cleft into linear
segments; petioles 0.7-2.0 em long; capitulescence of solitary
heads on long peduncles, 4-10 cm long; heads radiate, 5-6
mm high, 5-7 mm wide, involucres broadly campanulate;
ray flowers ca. 10-15, ligules white, 4-7 mm long, oblong ;
disc corollas yellow (?), 1.8-2.0 mm long, throats narrowly
campanulate-funnelform; achenes 1.3-1.5 mm long, nar-
rowly obovate to oblong-obovate, with thin or prominent
callous margins, the margins and faces short-pubescent ;
pappus of a conspicuous crown of squamellae and 1 (rarely

1974] Perityle — Powell 269

none) delicate bristle, ca. 0.5 mm long; chromosome num-
ber, unknown.

Known only from the type collection. (Fig. 3).

The distinguishing characteristics of Perityle lineariloba
include its pinnately divided leaves, long peduncles, and rel-
atively large heads with long ligules. Most of its floral
features are like those of P. turneri, to which the species is
related.

8. Perityle microcephala A, Gray, Proc. Amer. Acad. 21:
391. 1886, TYPE: Mexico: CHIHUAHUA: rocky hills near
Chihuahua, 8 Oct. 1885, C. G. Pringle 571 (Lectotype, GH ! ;
isolectotype, US!). LECTOPARATYPE: Chihuahua: mountains
above Batopilas, Oct, 1885, E. Palmer 268 (GH) ; isolecto-
paratypes, NY! US !).

Plants suffrutescent perennials, 20-40 cm high, many-
stemmed and densely leafy, erect or decumbent, densely
grayish-puberulent to pilosulous; leaves mostly opposite,
densely grayish-puberulent, 1.5-3.5 (5.0) em long, 0.8-2.0
(3.0) em wide, deltoid-ovate, ovate, to subcordate, the mar-
gins serrate-crenate to doubly so, or shallow-lobed; petioles
0.3-1.0 (1.8) cm long; capitulescence of severa] heads tight-
ly clustered on short peduncles; heads radiate, 3-4 mm high,
4-5 mm wide, involucres short-campanulate; ray flowers
ca. 8-12, ligules white, 2-5 mm long, oblong; disc corollas
yellow, often purple-tinged, 1.5-2.0 mm long, throats nar-
rowly campanulate-funnelform; achenes 1.5-1.8 mm long,
narrowly obovate to oblong-ovate, with callous margins, the
margins and faces short-pubescent; pappus of a prominent
crown of squamellae and 2 (rarely 1) unequal, delicate
bristles, the longest 0.5-1.0 mm long; chromosome number,
unknown.

Sierra Madre Occidental of Chihuahua and Sonora.
Flowering Aug-Nov, and probably in the spring also.
(Fig. 3).

REPRESENTATIVE SPECIMENS:
Mexico: CHIHUAHUA. Damon s.n. (UC); Guicorichi, Rio Mayo,

2'(0 Rhodora [Vol. 76

Gentry 1973 (ARIZ, UC, US); near Batopilas, Goldman 215 (GH, NY,
us); Madrono Mine, ca. 5 km NW of Pinos Altos, Hewitt 171 (GH);
Guayanopa Canyon, Sierra Madre, Jones s.n. (POM, US); Santa Clara
Mts., LeSueur 340 (ARIZ, CAS, GH, LL, TEX, UC); Mapula Mts., Pringle
764 (LL, NY, RSA, UC, US). SONORA. La Mesa Colorado, Gentry 541,
541m (ARIZ, DS); Sierra de Papas, Gentry 630 (Ds).

Everly (1947) treated Perityle microcephala as a syn-
onym of P. spilanthoides. I have found, however, that P.
microcephala represents a valid species, while P. spilan-
thoides is synonymous with the weedy variety of P. micro-
glossa.

Perityle microcephala belongs with the group of white-
rayed species which includes P. turneri, P. lineariloba, and
P. microglossa. Allof these taxa are quite similar in achene,
pappus, and most other floral characters, but P. micro-
cephala is clearly delimited from P. turneri and P. lineari-
loba by its capitulescence of small, tightly clustered heads,
grayish pubescence, and is easily separated from P. micro-
glossa by its pubescence and perennial habit. Overall
morphological similarity and distributional considerations
(Fig. 3) suggest that P. microcephala is most closely re-
lated to P. microglossa. Pringle 764 and Gentry 630, here
ascribed to P. microcephala, approach P. microglossa in
vegetative characteristics.

9. Perityle microglossa Benth. Bot. Sulph. 119. 1844. TYPE:
Nicaragua (?): Realejo, Nicaragua (?), 1837, R. B. Hinds
(K!).

9a. Perityle microglossa var. microglossa

Perityle acmella Harv. & Gray, Mem. Amer. Acad. II. 4:
77. 1849. TYPE: Mexico: “California”, Coulter 278 (Holo-
type, GH!; isotypes, NY!).

Galinsogeopsis spilanthoides Schultz-Bip. in Seem, Bot.
Herald 307. 1856. TYPE: Mexico: SINALOA: Sierra Madre,
1849, B. Seeman 1982 (Lectotype, K!). LECTOPARATYPE:
same data, B. Seeman 1981 (K).

Pericome spilanthoides Benth. & Hook. ex. Hemsl. Biol.
Centr. Amer. Bot. 2: 215. 1881.

Perityle microglossa var. effusa A. Gray, Syn. Fl. N.

1974] Perityle — Powell 271

Amer. 1: 332. 1884. TYPE: Arizona: Santa Catalina Mts.,
5 June 1882, C. G. Pringle (Holotype, GH!; fragment, us!;
isotype NY!).

Perityle effusa Rose, Contr. U.S. Nat. Herb. 1: 104. 1891.
TYPE: Mexico: SONORA: Alamos, 26 Mar.-8 Apr. 1890, E.
Palmer 350 (Lectotype, US!; isolectotypes GH !, NY !). LECTO-
PARATYPE: same data, E. Palmer 377 (US!) ; isolectopara-
types, (GH, NY !).

Perityle spilanthoides (Schultz-Bip.) Rydb. N. Amer. FI.
34: 17. 1914.

Plants weedy, herbaceous annuals, 20-60 cm high, erect
or decumbent, mostly branching above the base, stems few
or many, upper portions, especially peduncles, copiously
glandular-puberulent, rarely merely puberulent with few
glandular hairs; leaves opposite or alternate, mostly oppo-
site, puberulent, glandular-puberulent or glabrous, often
turning purplish, (2.0) 3.0-7.0 (10.0) em long, (1.5) 2.0-
5.0 (9.0) em wide, extremely variable in size and shape,
cordate, ovate, broadly ovate-cordate to subreniform, or
subdeltoid, the margins merely singly or doubly crenate-
even or irregular, to strongly 3-lobed or cleft, pedately
divided, or subhastate; petioles 0.5-3.0 (4.0) em long; capit-
ulescence of 1-3 or many heads clustered on relatively
short peduncles 0.6-3.5 (6.0) em long; heads radiate, 3.5-
4.5 (6.0) mm high, 4.0-6.5 (7.0) mm wide, involucres cam-
panulate; ray flowers 6-12, ligules white, 1.5-3.5 mm long,
oblong; disc corollas yellow, 1.2-2.2 mm long, throats tubu-
lar-funnelform; achenes 1.5-2.0 mm long, linear-oblong to
linear-elliptic, rarely very narrow and short obovate, with
thin to prominent callous margins, the margins ciliate, the
faces glabrous or puberulent; pappus of a crown of squa-
mellae and 2 unequal bristles, the longest 0.8-1.2 mm long;
chromosome numbers, n = 34, 51.

Rather widespread weed of northwestern Mexico, most
common in Sonora, Sinaloa, and Baja California Sur, rang-
ing south to Colima, seemingly introduced in northeastern
Mexico and extreme south Texas. Flowering probably year
around. (Fig. 3).

272 Rhodora [Vol. 76

REPRESENTATIVE SPECIMENS:

MEXICO: Baja California Sur. San Jose del Cabo, Anthony 326
(CAS, DS, GH, POM, UC, US) ; ca. 6 km NW of Mira Flores, Carter 2663
(ns, UC, US) ; Arroyo de los Pozos, Moran 6886 (sp); La Paz, Palmer
92 (CAS, GH, NY, US); Boca de la Sierra, Powell and Sikes 1672
(SRSC, TEX); 1 mi N of Pescadero, Powell and Sikes 1676 (SRSC,
TEX); vicinity of San Jose del Cabo, Wiggins 5689 (DS, NY, UC, US).
Chihuahua. Tres Hermanos, SW of Batopilas, Hewitt 36 (GH).
Colima. Colima, Oreutt 4551 (ps). Nayarit. Acaponeta, Jones s.n.
(PoM, UC); San Blas, Maltby 22 (us); Maria Magdalena Islands,
Maltby 175 (Ny, US); Tepic, Palmer s.n. (us). San Luis Potosi.
near waterfall at El Salto, King 3877 (NY, TEX, UC, us); from San
Luis Potosi to Tampico, Palmer 1093 (GH, NY, US). Sinaloa. Culiacan,
Brandegee s.n. (UC); ca. 64 mi S of Culiacan, Breedlove 1545 (bs);
Imala, Gentry 5455 (ARIZ, DS, NY, UC); San Blas, Jones, s.n. (CAS, NY,
POM, UC); Lodiego, Palmer 1614 (GH, NY); 56 mi NW of Mazatlan,
Powell and Sikes 1682 (sRsc, TEX); Rio del Fuerte, near El Fuerte,
Rose, Standley and Russell 13587 (NY, US); vicinity of Villa Union,
Rose, Standley and Russell 13932 (NY, US); ca. 5 mi E of Costa Rica,
Sikes and Babcock 192 (SRSC, TEX); 70 mi S of Mazatlan, Sikes and
Babcock 206 (SRSC, TEX). Sonora. 15 mi NE of Alamitos, Rio San
Miguel, Abrams 13357 (ps); Hermosillo, Brandegee s.n. (DS, GH);
12 mi W of Navojoa, Gentry 7950 (uc, US); Alamos, Palmer 673
(us); Rio Mayo, Navojoa, Sikes and Babcock 177 (SRSC, TEX);
2.4 mi N of Soyopa, Sikes and Babcock 157 (SRSC, TEX); 28 mi E
of Navojoa, Sikes and Babcock 185 (SRSC, TEX). Zacatecas. near
San Juan Capistrano, Rose 2427 (Ny, US). U.S.A.: TEXAS: Came-
ron Co. Rio Hondo, Chandler 7007 (us); E of Rio Hondo, Clover
1742 (ARIZ, TEX); Laguna Atacosa Refuge, Fleetwood 6069 (TEX).
Willacy Co. Sauz Ranch, Johnston 53, 280 (TEX).

The type locality originally given for Perityle m icroglossa,
Realejo, Nicaragua, is probably in error. The southern-most
eollection I have seen was from Colima, Mexico (Fig. 3),
and it seems likely that the species would have been col-
lected again in Nicaragua or in intermediate localities if it
were there. The type locality probably was San Blas, Naya-
rit, which also was visited during the Voyage of the Sulphur
and where P. microglossa is known to occur. According to
Rogers MeVaugh (personal communication), the localities
of many other Sulphur collections were mixed up in similar
fashion.

The variable nature of Perityle microglossa is evident

1974] Perityle — Powell 213

from the synonymy and from Everly's (1947) confused
treatment of the taxon. The current studies have led to the
recognition of two biological entities within P. microglossa.
One is var. microglossa (Fig. 3), a widespread, weedy poly-
ploid (n = 34, 51), and the other is var. saxosa (Fig. 3),
a more restricted "native" diploid (n — 17). Greenhouse
tests utilizing numerous collections of var. microglossa have
shown that it is self-compatible and that it reproduces
prolifically by seed. Only a few such tests have been
conducted with var. saxosa, but all have indicated self-in-
compatibility for the diploid taxon. The morphological char-
acters which distinguish the two varieties are not entirely
consistent, and several morphological intermediates, for
which chromosome numbers are not known, are found
among existing collections. Variety microglossa typically
exhibits shorter ligules, glandular-pubescent and shorter
peduncles, tighter capitulescences, and a weedy habit, as
compared to var. saxosa that usually displays longer ligules,
rather smooth and longer peduncles, more lax capitules-
cences, and a “native” habit.

Morphologically, Perityle microglossa (particularly var.
saxosa) is close to P. microcephala. Considering the evi-
dence presented for these two species, it can be postulated
that P. microglossa evolved from the montain-dwelling P.
microcephala through derivation of annual habit and adap-
tation to soil habitats. Variety microglossa must have
evolved in response to polyploidization and development of
reproductive self-compatibility which allowed for its
“weedy” proliferation.

Additional studies of Perityle microglossa var. saxosa, as
well as P. microcephala, are most desirable. If var. saxosa
proves to be a “native,” diploid, and self-incompatible pro-
genitor of var. microglossa, then perhaps specific status
should be accorded the latter taxon.

9b. Perityle microglossa var. saxosa (Brandeg.) Powell,
comb. nov.

Perityle saxosa Brandeg. Zoe 5: 225. 1905. TYPE: Mexico:
SINALOA: vicinity of Culiacan, Cerro Colorado, 5 Nov. 1904,

274 Rhodora [Vol. 76

Brandegee (Holotype, UC!; isotypes, GH!, POM!, US).
Perityle urticifolia Rydb. N. Amer. Fl. 34: 15. 1914.

TYPE: Mexico: SINALOA: Tepic, 5 Jan.-6 Feb. 1892, E.

Palmer 1960 (Holotype, NY !; isotypes, GH !, UC!, Us!).

Plants herbaceous annuals, possibly rarely weak peren-
nials, 15-50 cm high, erect, upper stems and peduncles
sparsely to densely puberulent, rarely moderately glandular-
puberulent; leaves opposite or alternate, mostly opposite,
glabrous to puberulent, rarely glandular-puberulent, 2-6
(8) cm long, (1.0) 1.5-3.5 (5.0) em wide; capitulescence
usually open, of 1-3 heads on relatively long peduncles, 2-7
em long; ligules 3.5-4.5 mm long; dise corollas 1.7-2.0 mm
long; achenes 1.0-1.6 mm long; pappus bristles 2, unequal,
the longest 1.0-2.0 mm long; chromosome numbers n = 17,
18.

Mostly in foothills and lower elevations, both sides of the
Sierra Madre Occidental, perhaps most common in Chihua-
hua, Sonora, and Sinaloa, Flowering probably year around.
(Fig. 3).

REPRESENTATIVE SPECIMENS:

MEXICO: Chihuahua. Temores, Flyr 60 (TEX); near Descanso,
Flyr 77 (TEX); Palmer 238 (GH, NY, Us). Durango. Tamazula,
Gentry 5234 (ARIZ, DS, GH, NY, UC). Nayarit, 4 mi E of Jalcocotan
on road to Tepic, MeVaugh 12147 (Us). Sinaloa. 68 mi N of Ma-
zatlan, Flyr 128 (TEX); 4 mi S of La Cruz junction, on MEX 15,
Flyr 122 (TEX); Cerro Colorado, Gentry 5468 (ARIZ, DS, NY, UC, US);
Canyon de Tarahumare, Sierra Surotato, Gentry 7315 (GH, NY, US);
road from Las Flechas to La Rastra, Goldman 323 (GH, NY, US);
30.4 mi E of Villa Union, Powell and Turner 1853 (SRSC, TEX).
Sonora. just S of Jecori, Drouet, Richards, and Lockhart 3692 (ps,
GH, NY); 18 mi NW of Campas, Drouet, Richards, and Lockhart 3704
(Ds, GH); San Bernardo, Rio Mayo, Gentry 1255 (ARIZ, GH, UC);
near Alamos, Gentry 4825 (ARIZ, DS, GH, NY); 12 mi SE of Colonia
Oaxaca, Hastings and Turner 65-43 (ARIZ, DS, SD); Petaquilla Canyon,
White 3320 (ARIZ, GH); La Vega Azul, SW of Cononia Morelos, White
4852 (ARIZ, GH, NY, US); 35 mi NE of Cajeme, on road to Tesopaco,
Wiggins 6408 (Ds, US).

The Hastings and Turner and the White collections from
NE Sonora are tentatively assigned to var. saxosa, but their

1974] Perityle — Powell 275

somewhat doubtful affinity with this taxon should be noted.
The specimens have elongated achenes, approaching the
fruit characteristics of Perityle ciliata and P. coronopofolia,
and they are perennial. In vegatative features they are
more like P. microglossa. Perhaps the unusual collections
are from a relict population which is transitional between
the shorter-achened (e.g., P. microglossa) and the longer-
achened (e.g., P. ciliata) members of the white-rayed al-
liance. Again, however, as is the case with most of the
poorly collected Sierra Madre taxa, further study is re-
quired.

10. Perityle canescens Everly, Contrib. Dudley Herb. 3: 393.
1949. TvPE: Mexico: SINALOA: Capadero, Sierra Tacuicha-
mona, 12 Feb. 1940, H. S. Gentry 5588 (Holotype, DS!; iso-
types ARIZ!, GH !, NY !, UC], Us!).

Plants low, suffruticose perennials, 6-15 cm high, densely
leafed upper stems pilose; leaves opposite or alternate,
pilose-hirsute to villous, 1.2-1.7 em long, 0.5-0.9 cm wide,
3-pinnatifid with linear segments, ovate-cordate in outline,
crisped; petioles 5-9 mm long; capitulescence essentially of
solitary heads on short peduncles; heads radiate, 5-7 mm
high, 6-8 mm wide, involueres broad campanulate; ray
flowers ca. 10, ligules white, 2.5-3.5 mm long, oblong to
broadly so; disc corollas yellow, becoming purple-tinged
2.5.3.0 ram long, throats broad tubular; achenes 2.0-3.5 mm
long, oblong-oblanceolate and nearly truncate on both ends,
with thick callous margins, the margins merely puberulent,
faces evenly puberulent; pappus of a prominent crown of
united, erose squamellae, and 2 very unequal, rather stout,
bristles, the longest 1.5-2.5 mm long; chromosome number,
unknown.

Known only from the type collection.

Perityle canescens is related to P. coronopifolia from
which it is distinguished by habit; dense indument; tend-
ency for solitary heads; leaf morphology ; thickly calloused
achenes with short-pubescent margins; strong, more or less

276 Rhodora [Vol. 76

united crown of pappus squamellae; stout pappus bristles:
and distribution.

Two other species, Perityle ciliata and P. hofmeisteria
are grouped with the above taxa in the white-rayed alliance,
largely on the basis of their similar floral characteristics
(particularly achene and pappus). In habit and achene
shape, P. canescens resembles P. lemmoni (sect. Laphamia:
Powell, 1973), but the former taxon obviously belongs with
sect. Perityle on the basis of its pappus crown.

11. Perityle ciliata (L. H. Dewey) Rydb. N. Amer. Fl. 34:
17. 1914.

Laphamia ciliata L. H. Dewey, Bot. Gaz. 20: 425. 1895.
TYPE: Arizona: rocks along Pine Creek near Pine, Ariz.,
26 Aug. 1891, D. T. MacDougal 676 (Holotype, US!; iso-
type, US!).

Plants suffruticose perennials, 15-30 cm high, many-
stemmed, erect to pendulous, densely short-pubescent above;
leaves opposite or alternate, densely gray-pubescent under-
neath, pubescent to puberulent on upper surfaces, 0.8-2.0
(3.0) em long, 0.6-1.3 (2.4) cm wide, deltoid-ovate to ovate-
rhombic, the marigns serrate, shallow-lobed, or serrate-
crenate; capitulescence of several heads clustered on short
peduncles; heads radiate, 5-7 mm high, 5.5-7.0 mm wide,
involucres campanulate; ray flowers ca. 6-10, ligules white,
3-6 mm long, broadly oblong to oblong-elliptic; dise corollas
yellow, often purple tinged, 2.0-2.5 (3.0) mm long, throats
tubular to tubular-funnelform; achenes 2.0-2.8 mm long,
linear-oblong, with prominent callous margins, the margins
long-ciliate, the faces slightly angled and pubescent; pappus
of a crown of squamellae and 2 (rarely 3) slender bristles,
1.5-2.0 (2.5) mm long; chromosome number, n — 17.

Restricted in distribution to the mountains of central
Arizona; rock-dwelling. Flowering spring, summer, and
fall. (Fig. 3).

REPRESENTATIVE SPECIMENS:
Arizona: APACHE C0.: Springerville, MeGinnies (ARIZ). COCONINO
CO.: Long Valley, Coconino Natl. Forest, Coconino Rec. Crew 492

1974] Perityle — Powell 277

(us); Oak Creek, W of Troutdale, Goodding 198-47 (ARIZ, SMU);
4 mi W of Strawberry, Sikes 99 (SRSC, TEX). GILA CO.: Barnhart
Pass, Matzatzal Mts., Collom 102 (GH, NY, US); Fossil Creek Hill,
Collom 594 (us); Parker Creek Canyon, Sierra Ancha Mts., Gould
8632 (ARIZ, CAS, UC); Workman Creek Falls, Sierra Ancha, Johnson
(ARIZ); Peterson Ranch, Sierra Ancha, Pase 1306 (ARIZ); 5.8 mi W
of Strawberry, Sikes and Patterson 427 (SRSC, TEX). MOHAVE CO.:
Hualapai Mts., Braem (ns); trail to Potato Patch, Hualapai Mts.,
Braem 875 (Ds). YAVAPAI CO.: near Senator Mine, along road from
Prescott, Eastwood 16721 (CAS); between Prescott and Ash Fork,
Eastwood 16770 (cas); Groom Creek, mts. near Prescott, Kearney and
Peebles 9750 (ARIZ, uc); near Granite Dells, vicinity of Prescott,
Kearney and Peebles 12780 (ARIZ, GH, NY, us); Prescott, Peebles,
Harrison, and Kearney 2645 (ARIZ, US).

This species is clearly related to P. coronopifolia from
which it is conveniently delimited by leaf shape.

12. Perityle coronopifolia A. Gray, Pl. Wright. 2: 82. 1853.
TYPE: New Mexico: GRANT CO.: copper mines of Santa Rita
del Cobre, Sept., 1851, C. Wright 1196 ( Holotype, GH!;
isotypes, NY!, US!).

Laphamia coronopifolia (A. Gray) Hemsl. Biol. Centr.
Amer. Bot. 2: 210. 1881.

Laphamia scopulorum M. E. Jones, Contr. West. Bot. 12:
48. 1908. TYPE: Mexico: CHIHUAHUA: Colonia Juarez, 6000
ft., 12 Sept. 1903, M. E. Jones (Holotype, POM!; fragment,
Us!).

Plants low, suffruticose perennials, 6-36 cm high, erect or
pendulous, many-stemmed, very leafy, grayish-pubescent ;
leaves opposite or alternate, puberulent to densely so, or
nearly glabrous, 0.7-2.5 (3.5) cm long, 0.5-1.2 (2.0) cm
wide, 3-palmate and lobed with slender-spathulate seg-
ments; pedately divided, or delicately 2,3-pinnatifid with
linear-filiform segments; petioles 2-8 (12) mm long; capi-
tulescence of several heads clustered on short peduncles;
heads radiate, 5.0-6.5 mm high, 5-6 mm wide, involucres
campanulate; ray flowers ca. 8-12, ligules white, 3-7 mm
long, broadly oblong, oblong-elliptic, to subspathulate; disc
corolas yellow, often purple tinged, 2.0-2.8 mm long,
throats tubular, tubular-funnelform, or tubular-campanu-

278 Rhodora [Vol. 76

late; achenes 1.8-2.5 mm long, linear-oblong to narrowly
oblanceolate, with thin or prominent callous margins, the
margins typically long-white ciliate, rarely short ciliate, the
faces slightly angled and glabrous or pubescent; pappus of
a crown of squamellae and 2 (rarely 3) slender bristles,
1.5-2.5 mm long; chromosome number, n = 17.

Rather widsepread and common in southern New Mexico-
Arizona, and rare in northern Chihuahua, Mexico ; rock
dwelling. Flowering spring, summer, and fall. (Fig. 3).

REPRESENTATIVE SPECIMENS:

MEXICO: Chihuahua. near Colonia Juarez, 6000 ft, Sierra Madres,
Jones (POM, US).

U.S.A.: Arizona: COCHISE co.: Miller Canyon, Hauchuca Mts.,
Barneby 5172 (CAS, NY); Ramsey Canyon, Huachuca Mts., Goodding
758 (GH, NY, RSA). GRAHAM co.: Fry Canyon, Pinaleno Mts., Shreve
1365 (ARIZ); Graham Mt., Thronber and Shreve 7783 (ARIZ). GREEN-
LEE CO.: Mts. back of Clifton, Greene (NY); San Francisco Mts.,
Greene (POM); 0.5 mi N of Metcalf, Maguire, Richards and Moeller
11822 (ARIZ, GH, NY, US). PIMA CO.: Spud Ranch, Rincon Mts.,
Blumer 3331 (ARIZ, DS, GH, UC) ; north slope Baboquivari Peak, Clark
12565 (GH); San Pedro Vista, Santa Catalina Mts, Parker 8096
(ARIZ, NY, RSA, US); below Goose-Head Rock, Mt. Lemmon, Sikes 95
(SRSC). SANTA CRUZ co.: Flux Canyon, Patagonia Mts., Kearney
and Peebles 10193 (ARIZ, US); Nogales to Ruby, Kearney and Peebles
14961 (ARIZ, GH, NY). New Mexico: DOÑA ANA CO0.: W side, Organ
Mts., Dunn 7274 (NMC, RSA); E peaks, Organ Mts., Powell 1393
(SRSC); Dripping Springs Canyon, Organ Mts., Sikes 108 (SRSC).
OTERO CO.: Dry Canyon, Rehn and Viereck (US). SOCORRO co.:
Mogollon Creek, Mogollon Mts., Metcalf 323 (NMC, NY, UC, US);
Gila Hot Springs, Mogollon Mts., Metcalf 828 (POM, UC, US).

Perityle coronopifolia is closely related to P. ciliata, from
which it is best distinguished by leaf Shape. The species
also has affinity with P. canescens. Both P. coronopifolia
and P. canescens have lobed leaves which are technically
distinguishable, and they are further delimited by the long
white-ciliate achene margins, clustered heads, pubescence,
and distribution.

The principal distribution of Perityle coronopifolia is in
southern Arizona and New Mexico. The one known Mexican
collection, originally described as Laphamia scopulorum, is

1974] Perityle — Powell 279

considered to be a morphological variant. The Jones speci-
mens from near Colonia Juarez are both of poor quality
but show delicate pinnatifid leaves, short-pubescent achene
margins, and inconspicuous pappus squamellae. These two
fruit characteristics indeed approach those of Laphamian
taxa (Powell, 1973), but the strong indication of its rela-
tionship with the Peritylean P. coronopifolia suggests its
correct placement with the latter species.

13. Perityle hofmeisteria Rydb. N. Amer. Fl. 34: 18. 1914.
TyPE: Mexico: DURANGO: vicinity of Durango, Apr.-Nov.,
1896, E. Palmer 28 (Holotype, NY !; isotypes, GH !, US!).

Plants suffrutescent to herbaceous perennials (possibly
also annuals), 12-25 cm high, puberulent; leaves opposite
or alternate, puberulent, 1.0-5.5 em long, 0.2-2.0 em wide,
ovate to subspathulate and entire to shallow-lobed, or ovate-
cordate and serrate to serrate-lobed ; petioles 0.5-2.0 (3.0)
em long; capitulescence of 1-3 heads borne on relatively
short peduncles, 1.5-2.8 cm long; heads radiate, 6-7 mm
high, 4-6 mm wide, involucres turbinate-campanulate; ray
flowers ca. 10-12, ligules white, 4-5 mm long, oblong; disc
corollas yellow, 2.5-3.0 mm long, throats tubular-funnel-
form; achenes 2.5-3.0 mm long, linear-oblong to narrowly
obovate, with thin or thick callous margins, the margins
prominently ciliate, the faces puberulent especially in the
centers; pappus of a crown of squamellae and 2 subequal
bristles, 2-3 mm long; chromosome number, n= 16 =+ 1.

Seemingly endemic in Durango, Mexico; probably rock-
dwelling. Flowering spring-fall. (Fig. 3).

SPECIMEN EXAMINED:
Mexico: DURANGO: 3 mi N of Rodeo, Flyr 332 (TEX).

The Flyr 332 specimen, said by the collector to be *seem-
ingly an annual," conforms to the types of Perityle hofmeis-
teria in floral but not in vegetative features. The main dif-
ferences are in habit appearance and leaf shape. The spe-
cies is poorly understood at present, but its affinity is with

280 Rhodora [Vol. 76

P. ciliata and P. coronopifolia from which it is readily
distinguished by leaf morphology and distribution (Fig. 3).

14. Perityle jaliscana A. Gray in S. Wats. Proc. Amer. Acad.
22: 431. 1887. TYPE: Mexico: JALISCO: Rio Blanco, ca. 10
mi NW of Guadalajara, 17-23 Sept. 1886, E. Palmer 554
(Holotype, GH ! ; isotypes, NY !, Us (2) !).

Plants low, suffruticose perennials, 5-17 cm high, stems
Spreading, densely puberulent; leaves opposite or alternate,
densely puberulent to nearly glabrous, 2.0-3.5 em long, 0.8-
1.8 (2.0) em wide, deltoid, deltoid-rhombic, deltoid-ovate,
or subhastate to subhalberd, usually 2-4 lobed proximally,
rarely serrate-lobed; petioles 0.7-1.8 (1.7) em long; capit-
ulescence of 1-several heads clustered on short, slender
peduncles; heads discoid, 5.0-6.5 mm high, 4-5 mm wide,
involucres funnelform-campanulate; disc corollas cream-
white and purple-tinged, 2.0-2.5 (3.0) mm long, throats
tubular-campanulate; achenes 1.8-2.2 (2.8) mm long, ob-
long or nearly so, slightly tapering to the base, truncate,
with prominent callous margins, rarely with 3-callous mar-
£ins, angled on both faces, the margins and faces short-
pubescent; pappus of a vestigial erown of squamellae, and
2 (3) slender bristles, 0.5-1.0 mm long, the bristles rarely
flattened ; chromosome number, n = 17.

Rock-dwelling endemic in Jalisco, Mexico. Flowering
spring-fall. (Fig. 4).

SPECIMENS EXAMINED:

Mexico: JALISCO: ca. 40 km N of Guadalajara, road to San Cristo-
bal de la Barranca, MeVaugh 22114 (NY, sD); Sierra San Esteban.
near Guadalajara, Pringle 2352 (GH, UC, US) ; Sierra de San Esteban,
Pringle 156834 ( ARIZ, CAS, LL, SMU, US); Rose and Painter 7476 (NY,
US); 5 mi SW, by road, from San Cristobal de la Barranca, Sikes
and Babcock 383 (SRSC, TEX).

Although relatively few collections of Perityle jaliscana
are known, it is the best-collected of four closely related
endemic species of Jalisco, Mexico. The others, P. rosei,
P. trichodonta, and the recently described P. feddemae, are
known only from type collections.

1974] Perityle — Powell 281

jaliscana (open circles); P. rosei
(open square); P. feddemae (closed

Fig. 4. Distribution of P.
(closed square); P. trichodonta

circle).

282 Rhodora [Vol. 76

Perityle jaliscana and P. feddemae are the most distinct
of the four species. I have followed Everly (1947) in recog-
nizing P. rosei and P. trichodonta, even though future col-
lections might indicate that they should be merged as a
single species. Furthermore, both P. rosei and P. tricho-
donta might simply be ecological variants of P. jaliscana.
I recognize the four Species at this time mainly to preserve
taxonomic consistency. In all three sections of the genus
Perityle there are groups of closely related species that are
no more distinctive than are the Jalisco endemics. In
Jalisco, P. feddemae, with its dissected leaves, seems to be
quite distinct, and it was recently discovered in a locality
not far from the other species (Fig. 4). Strict endemism
is common among the rock-dwelling taxa of Perityle, and
lacking evidence to the contrary, I think it possible that the
taxa in question are endemic species.

Perityle jaliscana is most closely related to P. rosei. Most
of its distinctive features, such as discoid heads, smaller,
oblong achenes, shorter pappus bristles and squamellae,
subhastate leaves, and short, dense pubescence, appear as
if they could have been derived from P. rosei.

15. Perityle rosei Greenm. Proc, Amer. Acad. 40: 45. 1905.
TYPE: Mexico: JALISCO: in the Sierra Madre W of Bolanos,
15-17 Sept. 1897, J. N. Rose 2947 (Holotype, Us!; isotypes,
GH !, NY!).

Plants suffruticose perennials, 10-20 em high, stems erect
or spreading, densely short-pilose; leaves opposite or alter-
nate, short-pilose to puberulent, 1.8-3.0 em long, 1.0-1.5
(2.0) em wide, deltoid to deltoid-ovate or deltoid-rhombic,
rarely subhastate, the margins entire or nearly so; petioles
5-9 mm long; capitulescence of 1.-several heads on short
peduncles; heads radiate, ca. 5 mm high, 4-5 mm wide,
involucres campanulate; ray flowers ca. 13, ligules white,
2.0-2.5 mm long; dise corollas cream-white and purple-
tinged, 2.0-2.3 mm long, throats tubular-campanulate ;
achenes 2.0-2.5 mm long, narrowly obdeltoid, with promi-
nent callous margins, slightly angled on both faces, the

1974] Perityle — Powell 283

margins and faces puberulent to subglabrous; pappus of a
short crown of squamellae, more or less united, and 2 (3-4)
slender bristles, 1.5-2.0 mm long, the 3-4 bristles, if present,
reduced in length; chromosome number, unknown.

Known only from the type collection. (Fig. 4).

Perityle rosei is best delimited from P. jaliscana by its
radiate heads, nearly obconical achenes, longer crown of
pappus squamellae and longer bristles, nearly deltoid, sub-
entire leaves, and longer, less dense pubescence.

16. Perityle trichodonta Blake, Proc. Biol. Soc. Wash. 37:
60. 1924. TYPE: Mexico: JALISCO: Sierra Madre W of Bo-
lanos, 15-17 Sept. 1897, J. N. Rose 2978 (Holotype, us!;
isotype, GH !).

Plants low, suffruticose perennials, ca. 10 cm high, many-
stemmed and spreading, densely gray-pubescent above;
leaves mostly opposite, densely hirtellous-pilose and gray-
ish, 7-9 mm long, 2.5-5.0 mm wide, ovate to ovate-rhombic
and subhastate; petioles 2-4 mm long; capitulescence of
1-several heads clustered on short peduncles; heads radiate,
ca. 4.5 mm high, ca. 4 mm wide, involucres funnelform-
campanulate; ray flowers ca. 8, ligules white, 1.0-1.5 mm
long; disc corollas cream-white and purple-tinged, 2.0-2.5
mm long, throats tubular-campanulate; achenes 1.5-2.2 mm
long, narrowly obdeltoid, with prominent callous margins,
slightly angled on both faces, the margins and faces puberu-
lent to subglabrous; pappus of a short crown of squamellae,
more or less united, and 2 slender bristles, 1.5-2.2 mm long,
chromosome number, unknown.

Known only from the type collection. (Fig. 4).

Perityle trichodonta looks as though it could be a small-
leafed, more densely pubescent, shorter liguled form of P.
rosei, but it does differ in the characters listed. Future col-
lections might reveal that P. trichodonta should be merged
with P. rosei.

17. Perityle feddemae McVaugh, Contr. Univ. Mich. Herb.

284 Rhodora [Vol. 76

9: 431-433. 1972. TvPE: Mexico: JALISCO: Rancho Viejo,
ca. 10 km ENE of Huejuquilla el Alto, rock-crevices, open
oak-woodland, elev. 1900 m, 31: Oct. 1963, Feddema 2278
(Holotype, MICH ; isotype, SRSC!).

Plants suffruticose perennials, 10-25 cm high, stems
spreading, puberulent; leaves mostly opposite, puberulent,
glandular-dotted, 1.7-3.5 em long including petioles, dis-
sected-pinnatifid with linear-spatulate segments; capitu-
lescence of 1-3 heads on short, slender peduncles; heads
discoid, 5-6 mm high, 3-5 mm wide, involucres funnelform-
campanulate; disc corollas pale yellow, 2.3-2.8 mm long,
throats tubular-campanulate; achenes 2-3 mm long, oblong-
oblanceolate, with prominent callous margins, angled on
both surfaces, the margins and faces short-pubescent; pap-
pus of a short crown of squamellae and 2 slender, unequal
bristles, ca. 1.0 mm long; chromosome number, unknown.

Known only from the type collection. (Fig. 4).

This taxon possibly is most closely related to Perityle
jaliscana that also has discoid heads, but its dissected leaves
mark it as the most distinctive of the Jalisco endemics.

18. Perityle parryi Gray, Pl. Wright. 2: 106. 1853. TYPE:
Texas: Canyon of the Rio Grande, Aug. 1852, C. C. Parry
521 (Isotype, NY!).

Laphamia parryi (Gray) Benth. & Hook. ex Hemsl. Biol.
Centr. Amer. Bot. 2: 210. 1881.

Leptopharyna trisecta Rydb. N. Amer. Fl. 34: 23. 1914.
TYPE: Mexico: CHIHUAHUA: dry cliffs, rocky hills near
Chihuahua, 20 Mar. 1885, C. G. Pringle 183 (Holotype, NY l;
isotypes, DS!, GH !, UC!, US!).

Plants low, woody-based perennials in crevices of rocks
or tall, spreading, herbaceous to suffruticose perennials in
soil, 10-75 em high; leaves opposite or alternate, glandular-
pubescent, 2-6 (9) cm long, 0.8-4.0 (6.0) cm wide, cordate
to subreniform in outline, margins irregularly lobed, den-
tate, or laciniate, but typically shallow to deeply 3-lobed,
and in some specimens even subcruciform; petioles 1.2-

1974] Perityle — Powell 285

Fig. 5. Distribution of P. parryi (open triangles); P. vaseyi
(closed triangles); P. aglossa (open squares); P. cordifolia (closed
circles); P. leptoglossa (closed squares); P. lloydii (open star);
P. lobata (closed stars) ; P. palmeri (open circles).

286 Rhodora [Vol. 76

4.0 em long; capitulescence of 1-3 heads borne on pe-
duncles 1-7 cm long; heads radiate (rarely discoid), 0.8-1.0
em high, 0.7-1.4 em wide, involucres hemispherical to campa-
nulate; bracts numerous, lanceolate to linear, rarely ob-
lanceolate, acute to attenuate at the apex; ray flowers ca.
12-16 (rarely 1-6), ligules yellow, oblong, showy; disc flow-
ers numerous, corollas yellow, (3.2) 4-6 mm long, throats
tubular; achenes (2) 3-4 mm long, linear-elliptic to ob-
conical-elliptic, with thin to prominent callous margins, the
margins short-pubescent to dense-pilose, faces pubescent ;
pappus of a crown of squamellae, and 1 (rarely 0 or 2)
barbellate bristle, (1.0-2.5) 3-6 mm long, typically tapering
to a point, the barbs typically antrorse throughout, rarely
becoming lateral or retrorse toward the tip; chromosome
number, n = 17.

Western Big Bend area of Texas and adjacent Mexico;
occurring in rocks and soils. Flowering essentially year
around. (Fig. 5).

REPRESENTATIVE SPECIMENS:

MEXICO: Chihuahua. 11 mi NE of Camargo, Johnston 7920
(GH); road to San Carlos Mines, Johnston and Muller 39 (GH);
12 mi S of Ojinaga, Johnston and Muller 1443 (GH); near new lake
on Rio Conchos, Powell, Turner, and Magill 2030 (SRSC, TEX); rocky
hills near Chihuahua, Pringle 183 (Ds, GH, NY, UC, US); 1 mi E of
km 100, Ojinaga-Camargo road, Sikes and Patterson 397 (SRSC, TEX) ;
20 km N of Chihuahua, Stewart and Johnston 2119 (GH). U.S.A.:
Texas: Brewster Co. Lost Mine Peak, Chisos Mts, Correll and
Johnston 24540 (LL); Green Valley Ranch, Correll and Wasshausen
27825 (LL); Agua Fria Spring, Cory 18638 (GH); Santa Helena
Canyon, Cory 26475 (us); N side Whirlwind Spring area, 02 Ranch,
Fletcher 1143 (srsc); Packsaddle Mt., McAfee 261 (SRSC); Chisos
Mts., Muller 8247 (GH, NY, US, TEX); lower Window Trail, Big Bend
Park, Powell and Sikes 1528 (sRSC, TEX); Basin, Chisos Mts., Sikes
32 (SRSC, TEX); Boot Springs Trail, Sikes 116 (SRSC, TEX); Nine-
Point Mesa, Warnock and Johnston 17670 (srsC); Rosillos Mts.,
Warnock 20603 (sRsc). Presidio Co. S edge of Presidio, Correll and
Johnston 21867 (LL); 10 mi SE of Redford, Correll and Johnston
21898 (LL); Casa Piedra road, 20 mi S of Marfa, Cox s.n. (SRSC) ;
Capote Creek, Havard s.n. (US); near Cottonwood Springs, Sierra
Tierra Vieja, Hinckley 1850 (GH, NY, US); ca. 9 mi N of Lajitas,
Powell 1243 (sRSC) ; 5 mi S of Redford, Powell, Sikes, and Watson

1974] Perityle — Powell 287

1396 (sRsc); 12 mi S of Redford, Powell, Sikes, and Watson 1398
(SRSC); 3 mi E of Ruidosa, Powell and Brey 1513 (sRsc); Capote
Creek, ca. 1 mi below Capote Falls, Powell 1518 (sRsC).

The three yellow-flowered species of west Texas and ad-
jacent Mexico (Fig. 5), Perityle parryi, P. vaseyi, and P.
aglossa, are morphologically similar. Perityle parryi and
P. vaseyi are particularly close and somewhat confusing
taxonomically, primarily because of intermediate leaf forms
and uncertain distributional integrity.

Perityle parryi occurs as a rock-dwelling subshrub and as
a soil-dwelling perennial, typically with leaves that are
merely 3-lobed. Perityle vaseyi is a soil-dwelling, taprooted
perennial, with deeply trisected leaves. However, some
Mexican populations of P. parryi (— L. trisecta) have
deeply trisected leaves, as do some soil-dwelling Texas
forms, and it is not always possible to distinguish the taxa
by leaf morphology. Perityle parryi is further delimited by
long-tapering pappus bristles (one per achene) which are
antrorsely barbellate, bracts usually lanceolate or linear and
somewhat attenuate, and disc corollas with usually narrowly
tubular throats, as opposed to P. vaseyi with shorter pappus
bristles that are retrorsely barbellate at the tips, bracts
usually oblanceolate and acute, and more broadly tubular
dise corollas.

It is suspected that the soil-dwelling forms of Perityle
parryi and P. vaseyi may occasionally hybridize. Any hy-
bridization, however, would be difficult to detect morpho-
logically. It is not known whether the leaf variability of
P. parryi is intrinsic or whether it might be the result of
hybridization with P. vaseyi. In spite of marginal sympatry
(Fig. 5), mixed populations of the two species have not
been found, and no clear-cut hybrids have been detected.
The subject is under experimental study.

The populations of Perityle parryi near Ruidosa, Texas,
in the Sierra Vieja and one collection 12 mi S of Ojinaga
(Johnston and Muller 1443) are notably anomalous. Their
pappus bristles are short and have retrorsely barbed tips,
resembling those of P. vaseyi. In addition, the Sierra Vieja

288 Rhodora [Vol. 76

populations may be nearly rayless or entirely discoid. In
other morphological characters the anomalous populations
conform with P. parryi. The discoid forms should not be
confused with P. aglossa, which is always rayless.

19. Perityle vaseyi Coult. Contr. U. S, Nat. Herb. 1: 42.
1890. TYPE: Texas: BREWSTER CO. Chisos Mts., 1889, G. C.
Nealley (245?), (Holotype, Us!).

Plants perennial with fleshy taproots, (10) 15-75 cm
high, herbaceous to suffruticose, stems erect or spreading;
leaves opposite or alternate, glandular-pubescent, 2.5-6.5
em long, 2-5 em wide, typically palmately divided into 3
lobes or subcruciform, with the lobes also dissected, cleft,
or parted, rarely the leaves are strongly 3-lobed, cleft, or
parted; petioles 1.4-3.5 cm long; capitulescence of 1-3 heads
borne on peduncles 1-8 cm long; heads radiate, 0.8-1.0 cm
high, 1.0-1.3 em wide, involucres hemispherical to campanu-
late; bracts numerous, oblanceolate to linear-lanceolate,
typically acute, less often alternate at the apex; ray flowers
ca. 14-16, ligules yellow, oblong, showy; disc flowers numer-
ous, corollas yellow, 3.2-5 (6) mm long, throats tubular to
broadly so; achenes (2.0) 3.0-4.0 mm long, linear-elliptic
to obconical-elliptic, with thin to prominent callous margins,
the margins short-pubescent to dense pilose, faces pubes-
cent; pappus of a crown of squamellae, and 1 (rarely 0 or
2) barbellate bristle, 1.5-4.0 mm long, typically truncate,
the barbs retrorse or lateral toward the tip; chromosome
number, n — 17.

Desert soils, especially gypsiferous clay, southwestern
Big Bend area of Texas and adjacent Mexico. Flowering
essentially year around. (Fig. 5).

REPRESENTATIVE SPECIMENS:

MEXICO: Chihuahua. 6.5 mi S of Ojinaga, Powell, Turner, and
Magill 2002 (sRsC, TEX); 5 mi S of Ojinaga, Powell, Turner, and
Magill 2079 (sRsc, TEX). Coahuila. Picachos Colorados, Johnston
and Muller 141 (GH). U.S.A.: Texas: BREWSTER CO. petrified forest,
Cory 2706 (GH); Hot Springs, Cory 6477 (POM); Glenn Springs,
Cory 44014 (TEX); Terlingua Creek on 02 Ranch, Fletcher 917

1974] Perityle — Powell 289

(SRSC); 2-8 mi E of Terlingua, Flyr 16 (SMU, SRSC, TEX); 8 mi E
of Lajitas, Flyr 19 (SMU, TEX); 1 mi SE of Terlingua, Flyr 180
(SMU, TEX) ; 2 mi SE of Hen Egg Mt., Johnston and Warnock 3656
(SRSC); top third of Packsaddle Mt., McAfee 184 (snsc); 70 mi S
of Alpine, Powell and Sikes 1530 (sRSC, TEX); 18 mi SE of Castolon,
Powell, Averett, and Watson 1539 (sRsC, TEX) ; 0.4 mi N of Castolon,
Powell, Averett, and Watson 1544 (SRSC, TEX); near Big Bend Tun-
nel, Sikes 39 (SRSC, TEX) ; near hwy. 118 entrance to Big Bend Park,
Sikes 40 (SRSC, TEX); 7 mi E of Castolon, Sikes and Averett 330
(SRSC, TEX); road to Fisk Canyon, Warnock 1148 (sRsc); Avary
Canyon, Big Bend Park, Warnock 9139 (sRsC); Big Bend Park,
Warnock and Wallmo 12211 (sRSC); San Vicente, near the Rio
Grande, Young s.n. (TEX).

As discused previously under Phylogenetic Considera-
tions, an ancestral connection is hypothesized for Perityle
vaseyi and its allies with the P. crassifolia group of Baja
California. The primary evidence for this possible relation-
ship are the similar habits and pappus bristles (with re-
trorsely barbed tips) of P. vaseyi and P. crassifolia and the
chromatographic profiles with yellow compounds, common
to both groups.

Morphologically, Perityle vaseyi and its relatives are
closer to the P. cordifolia group of western Mexico. The
distribution of the Texas species (Fig. 5) could have been
achieved by migration from a Sierra Madre Occidental
origin. The headwaters of the Rio Conchos are in the Sierra
Madre near Creel, just across the divide from P. cordifolia
country. The distribution of P. parryi near the Rio Conchos
in eastern Chihuahua and the location of P. vaseyi and P.
aglossa near the Rio Grande, suggest the possibility of
eastern migration along the river systems.

20. Perityle aglossa Gray, Pl. Wright 2: 107. 1853. TYPE:
Texas: Rio Grande below Presidio del Norte, Aug. 1852,
C. C. Parry (Lectotype, GH !).

Laphamia aglossa (Gray) Benth. & Hook. ex Hemsl. Biol.
Centr. Amer. Bot. 2: 210. 1881.

Leptopharynx aglossa (Gray) Rydb. N. Amer. Fl. 34:
24. 1914.

Plants woody-stemmed perennials in rock crevices, 15-45

290 Rhodora [Vol. 76

cm high; leaves opposite or alternate, glandular-pubescent,
2.4 em long, 0.6-2.3 cm wide, ovate, subeordate or subdel-
toid, irregularly lobed, laciniate, or serrate-dentate, often
shallow-3-lobed; petioles 0.8-2.3 em long; capitulescence of
1-3 heads borne on peduncles 1.0-2.5 em long; heads discoid,
8-11 mm high, 4-8 mm wide, involucres campanulate; bracts
linear to linear-lanceolate, attenuated at the apex; disc
flowers numerous, corollas yellow, typically pink- or purple-
tinged, 4.2-6.0 mm long, throats tubular; achenes (2.0)
2.5.3.2 mm long, oblong-obconical, with prominent callous
margins, the margins short-pubescent to dense-pilose, faces
pubescent; pappus of a crown of squamellae and 1 (rarely
2) barbellate bristle, 3.7-5.5 mm long, tapering to a point,
the barbs antrorse throughout; chromosome number,
n = 17.

Restricted in southern Brewster Co. and Terrell Co.,
Texas, along and near the Rio Grande; crevices of lime-
stone. Spring-fall. (Fig. 5).

REPRESENTATIVE SPECIMENS:

Texas: BREWSTER CO.: mouth of Regan Canyon on Rio Grande,
Hinckley and Warnock 3737 (US, SRSC); near Boquillas, Marsh. 306
(GH); near Big Bend Tunnel, Sikes 38 (sRsC); 4 mi W of Hot
Springs, Big Bend Park, Warnock and Turner 8291 (sRsC); San
Vicente Canyon, Big Bend Park, Warnock and Parks 864 (LL, SMU,
SRSC); limestone hills, Terlingua area, Warnock 13980 (SRSC) ;
Heath Canyon, Black Gap Refuge, Warnock 20615 (sRSC); mouth
of Regan Canyon, Warnock and Hinckley 461050 (SMU, SRSC, TEX).
TERRELL CO.: mouth of San Francisco Canyon, Surratt 133 (SRSC) ;
mouth of San Francisco Canyon, Warnock and Surratt 9817 (LL,
SMU, SRSC) ; 9 mi E of Sanderson, Warnock 13155 (sRsC) ; along Rio
Grande, between Regan Canyon and Sanderson Canyon, Warnock
15854 (sRsC).

Although only one type specimen is cited for Perityle
aglossa (Parry, Aug. 1852), I have seen another specimen
which probably came from Parry's original collection. At
NY a sheet labeled *P. aglossa, Arroyo San Carlos, April
1852," displays specimens of both P. aglossa and P. parryi.
Judging from present-day distributions of the species, I
believe that the specimens were probably mixed after

1974] Perityle — Powell 291

Parry's collections were made. Furthermore, the P. aglossa
specimen on the NY sheet probably was from the original
colleetion and perhaps should be recognized as an isolecto-
type. The P. parryi specimen is discoid and probably came
from the vicinity of Ruidosa above Presidio.

Perityle aglossa is a limestone-dwelling subshrub with
good distributional integrity. In addition to habit and dis-
coid heads, the species is distinguished from P. parryi, its
closest relative, by subtle characters of the leaves, involu-
eral bracts, and disc corollas.

21. Perityle cordifolia (Rydb.) Blake, Contr. U. S. Nat.
Herb. 23: 1602. 1962.

Leptopharynx cordifolia Rydb. N. Amer. Fl. 34: 22.
1914. TYPE: Mexico: SINALOA: vicinity of Topolobampo,
23 Mar. 1910, J. N. Rose, P. C. Standley, and P. G. Russell
13294 (Holotype, NY!?; fragment of type, UC!; isotype,
GH!).

Plants suffrutescent perennials, rather low and spreading
or semi-pendulent, 10-20 cm high, stems strong but herba-
ceous except at the bases, pilose; leaves mostly alternate,
bright green, pilose, 3.5-11.0 cm long, 1.5-9.0 cm wide, cor-
date to broadly so, veins prominent, the margins doubly
serrate-dentate, rarely crenate; petioles 1.5-4.5 mm long;
capitulescence of solitary heads borne on rather stout pe-
duncles, 2.5-6.0 em long; heads radiate, rarely cernuous,
10-12 mm high, 9-15 mm wide, involucres hemispherical] to
campanulate, typically truncate at the base; bracts 10-13
mm long, linear and attenuate; ray flowers numerous, lig-
ules yellow, 9-15 mm long, oblorg-elliptic; disc flowers
numerous, corollas yellow, 5-7 mm long, throats tubular ;
achenes 2.5-3.0 mm long, narrowly oblong, with thin or
prominent callous margins, the margins typically densely
ciliate, rarely puberulent, faces glabrous or puberulent in
the center; pappus of a crown of squamellae, ca. 0.5 mm or
less, and a single bristle, 4-7 mm long, the bristle tapering
from base, a fine tip and densely barbellate; chromosome
number, n = 17.

292 Rhodora [Vol. 76

A petrophilic, cliff-dwelling species, possibly also rooted
in soils of shaded bluffs, Sonora and Sinaloa, Mexico. Flow-
ering probably year around. (Fig. 5).

REPRESENTATIVE SPECIMENS:

Mexico: SINALOA: hill near Topolobampo, Flyr 63 (TEX); N of
Topolobampo, Flyr 76 (TEX); road to Los Animas beach from Los
Mochis, Flyr 105 (TEX); 23 mi N of Culiacan, Flyr 111B (TEX);
Cerro Tecomate, W of Pericos, Gentry 5743 (ARIZ, DS, GH, NY, UC);
ca. 1 mi E of Topolobampo, Sikes and Babcock 188 (SRSC, TEX);
bluffs W of Topolobampo, Sikes and Babcock 190 (SRSC, TEX).
SONORA: 32 mi N of Navojoa, Flyr 103 (TEx); Canyon Sapopa,
Rio Mayo, Gentry 1282 (ARIZ, GH, NY, UC); Aquibiquichi, near Bata-
cosa, Gentry 3003 (ARIZ, CAS, GH, UC, US); Cerro de Bayajori, 12 mi
W of Navojoa, Gentry 7949 (uc, US); 23 km E of Navojoa, Moran
4032 (DS, sD, UC); 1-2 mi N of San Carlos Resort, Powell and Sikes
1689 (SRSC, TEX); 10 mi N of Guaymas, Randle and Werner 1 (ARIZ).

Perityle cordifolia belongs with a group of four other
morphologically similar species; P. palmeri, P. leptoglossa,
and P. lloydii also of northwestern (mainland) Mexico, and
P. lobata of Baja California. In spite of their similarities,
the taxa are rather clearly delimited by vegetative features,
excepting P. lloydii, which has characters of the other three
mainland species. I have grown all of the taxa except P.
lloydii under identical conditions, and each did maintain its
characteristic morphology.

Everly (1947) treated Perityle cordifolia as synonymous
with P. palmeri. I recognize these taxa as related but cer-
tainly distinct species. In fact, the soil-dwelling P. palmeri
could have evolved from the rock-dwelling P. cordifolia, but
on morphological grounds, P. palmeri shares even closer
relationship with P. lobata. Perityle cordifolia is readily
delimited from P. palmeri by its woody habit; bright green,
pilose, cordate leaves with prominent veins and doubly ser-
rate margins; long ligules; long attenuate involucral bracts;
long pappus bristles; and long style branches.

22. Perityle leptoglossa Harv. & Gray in A. Gray, Mem.
Amer. Acad. II. 4: 77. 1849. TYPE: Mexico: SONORA: “vi-
cinity of Hermosillo,” 1829-1830, Coulter 277 (Holotype,
GH!; fragment, Us!).

1974] Perityle — Powell 293

Leptopharynx leptoglossa (Harv. & Gray) Rydb. N.
Amer. Fl]. 34: 22. 1914.

Plants shrubby, spreading or erect, 10-60 cm high, stems
densely grayish-puberulent; leaves mostly alternate, densely
puberulent, gray-green, 3.0-5.5 em long, 1.3-3.5 cm wide,
cordate to cordate-ovate, the margins shallow-lobed and
crenate to serrate, or merely crenate-serrate; petioles 1.3-
2.5 cm long; capitulescence of solitary heads borne on rela-
tively stout peduncles, 2-5 em long; heads radiate, rarely
cernuous, 8-10 mm high, 7-12 mm wide, involucres cam-
panulate-cylindroidal and truncate at the base; bracts 5-8
mm long, linear to linear-lanceolate and attenuate to sub-
attenuate; ray flowers numerous, ligules yellow, 6-10 mm
long, oblong to oblong-elliptic; disc flowers numerous, corol-
las yellow, 4-5 mm long, throats tubular; achenes (2.2)
2.5-3.0 mm long, narrowly oblong, with thin or prominent
callous margins, the margins long- or short-ciliate, the faces
glabrous or minutely pubescent at the center; pappus of a
crown of squamellae less than 0.5 mm long, and a single
bristle, 2.5-4.5 mm long, the bristle tapering from base to
apex and densely barbellate; chromosome number, n = 17.

Strictly rock-dwelling, in the vicinity of Hermosillo, So-
nora, Mexico. Flowering probably year around. (Fig. 5).

REPRESENTATIVE SPECIMENS:

Mexico: SONORA: 3 mi S of Magdalena, Aleramo 13216 (ps);
rocky sides of El Cerro de la Campana, Drouet, Richards and Alva-
rado 3387 (DS, GH, NY); 6 km N of Hermosillo, Drouet and Richards
3757 (ps); San Miguel de Horcasitas, Eisen (UC); 41 mi N of
Hermosillo, Flyr 96 (TEX) ; 26 mi S of Hermosillo, Gentry 45'12 (UC) ;
ca. 28 mi N of Hermosillo, Powell and Averett 1520 (SRSC, TEX) ;
ca. 28 mi N of Hermosillo, Powell and Sikes 1683 (SRSC, TEX); near
Hermosillo, Ripley 14309 (CAS) ; vicinity of Hermosillo, Rose, Standley,
and Russell 12346 (NY, US); near Magdalena, Shreve (NY); near
Estacion Torres, Shreve 6070 (ARIZ); 8 mi W of Hermosillo, Wiggins
and Rollins 96 (ns, GH, NY); ca. 1 mi N of Torres, Wiggins 6266
(DS, GH, POM, US); 12 mi from Ures on road to Babiacari, Wiggins
7359 (DS, TEX, US).

The most distinctive features of Perityle leptoglossa are:

294 Rhodora [Vol. 76

shrubby habit; leaf margins shallow-lobed or merely cre-
nate-serrate; densely puberulent upper stems and leaves;
small heads and short ligules; short bracts; achene margins
typically short-ciliate; pappus bristles typically slender at
bases. At one locality ca. 28 mi N of Hermosillo, I observed
a few plants that were 214 ft. high, the largest woody plants
in the genus. The species is related to P. palmeri and P.
lloydii.

23. Perityle lloydii Rob. & Fern. Proc. Amer. Acad. 30: 118.
1894. TvPE: Mexico: SONORA. Badehuachi, 2 Dec. 1890,
C. E. Lloyd 400 (Holotype, GH ! ; isotypes, NY !, UC!, US(3)!).

Leptopharynz lloydii (Rob. & Fern.) Rydb. N. Amer. Fl.
34: 22. 1914.

Plants semishrubby, erect, 10-40 cm high, stems woody
to herbaceous, pilose; leaves mostly opposite, pilose, 2.5-
4.5 em long, 1.5-2.5 em wide, cordate to cordate-ovate, the
margins doubly serrate; petioles 1.5-3.0 em long; capitu-
lescence of solitary heads borne on peduncles 2.0-3.5 cm
long; heads radiate, rarely cernuous, 8-9 mm high, 6-7 mm
wide, involucres campanulate-cylindroidal and truncate at
the base; bracts 5-7 mm long, linear and attenuate; ray
flowers numerous, ligules yellow, 5-8 mm long, oblong to
oblong-elliptic; disc flowers numerous, corollas yellow, ca.
4 mm long, throats tubular; achenes ca. 3 mm long, nar-
rowly oblong, the callous margins short-pubescent; pappus
of a minute crown of squamellae (ca. 0.2 mm long), and a
single bristle ca. 3 mm long, the bristle slightly tapering
from base to apex and densely barbellate; chromosome
number unknown.

Known only from the type collection. (Fig. 5).

Perityle lloydii is poorly delimited from the other main-
land taxa of the P. cordifolia species-group. In leaf shape,
pubescence, and achene morphology (short-pubescent mar-
gins and reduced pappus squamellae) P. lloydii resembles
P. cordifolia. In shrubby habit, small heads, short ligules,
and other floral characters, the taxon approaches P. lepto-

1974] Perityle — Powell 295

glossa. Its pubescence is like that of P. palmeri. Overall,
P. lloydii possibly is closer to P. leptoglossa, and further
collections might well provide evidence for merging the
taxa.

24. Perityle lobata (Rydb.) I. M, Johnston, Proc. Calif.
Acad. IV. 12: 1205. 1924.

Leptopharynz lobata Rydb. N. Amer. Fl 34: 23. 1914.
TYPE: Mexico: BAJA CALIFORNIA: Commondu, 19 Feb. 1889,
T. S. Brandegee (Holotype, GH!; isotype, uc!).

Plants fleshy-rooted perennials, possibly also annuals,
low and spreading, semierect, or semipendulent, 10-40 cm
high, stems weak, herbaceous, and rather brittle, usually
purplish, glabrous to sparse-pilose ; leaves mostly alternate,
bright green, turning purplish, glabrous to sparse-pilose,
3-8 (12) cm long, 1.8-6.0 (8) cm wide, cordate, cordate-
orbiculate, or subreniform, strongly 3-lobed, cleft, or parted,
rarely divided, the segments further shallow-lobed or the
blades even 5-cleft, the margins dentate-lobed and acumi-
nate, overall the blades appear regularly laciniate; capitu-
lescence of solitary heads borne on weak peduncles 3-8 (15)
em long; heads radiate, cernuous, 10-12 mm high, (5) 7-9
mm wide, involucres cylindroidal to campanulate, truncate
at the base; bracts 5.0-9.5 mm long, linear-lanceolate to
lanceolate-ovate, subattenuate, often purplish; ray flowers
8-12, ligules yellow, often turning purplish when dry, 2.5-
9.0 mm long, oblong-elliptic; disc flowers numerous, corollas
yellow, often purple-tinged, 3.8-5.0 mm long, throats tubu-
lar-funnelform; achenes (3) 3.5-5.0 mm long, narrowly
oblong to oblong-elliptic, with prominent callous margins,
the margins densely ciliate, faces glabrous to puberulent
at the center; pappus of a crown of squamellae, 0.7-1.0 mm
lonz, and a single bristle 3.5-6.0 mm long, the bristle very
stout at the base, tapering to a point, densely barbellate ;
chromosome number, n = 17.

Restricted to Baja California Sur in the area of Sierra
de la Giganta. Flowering year around. (Fig. 5).

296 Rhodora [Vol. 76

REPRESENTATIVE SPECIMENS:

MEXICO: BAJA CALIFORNIA SUR. Arroyo Carrizal, E of Rancho El
Horno, NE of San Javier, Carter and Ferris 3811 (UC, US) ; Arroyo
Santo Domingo, Rancho El Horno, Carter and Ferris 3867 (DS, SD,
UC); vicinity of Portezuelo E of La Victoria, Carter and Ferris
3911 (ns, UC); NW of Rancho El Horno, on steep escarpment of
Mesa de San Alejo, W of San Javier, Carter and Sharsmith 4199
(Uc); Purisima, Gentry 4213 (ARIZ, Ds, GH, UC); Puerto Escondido,
Johnston 4115 (CAS, GH, NY, UC, US); ca. 10 mi W of Loreto, Powell
and Sikes 1663 (SRSC, TEX); ca. 10 mi S of Mission Dolores, Wiggins,
Carter, Ernst 287 (ps, vs); arroyo Quisapol, E of La Presa, trail
to Laguna Caquihui, Sierra de la Giganta, Wiggins 15544 (ps).

Perityle lobata has perhaps been separated from main-
land Mexico and its relatives there for about four million
years. At least according to Moore and Buffington (1968),
this is how long the Baja California peninsula has been
drifting westward from the mainland. Perityle lobata is
clearly related to P. palmeri. Distributional considerations
suggest that the two present-day taxa once had a common
range (Fig. 5), or else proximal distributions before the
Sea of Cortez became a barrier of its present width.

It is possible that Perityle lobata has been introduced to
the mainland and has hybridized there, accounting for some
of the variability in mainland taxa such as P. palmeri, In
fact, it might be postulated that P. palmeri has evolved
from P. lobata instead of the other way around as previ-
ously assumed. However, if the latter conjecture were ac-
curate, one might expect that P. palmeri would be more
like P. lobata of Baja California. I have not found any
evidence that P. lobata has been introduced recently to the
mainland,

As opposed to Perityle palmeri, P. lobata can be recog-
nized by its leaf architecture, its purplish stems, pubescence,
ligule length and number, bract width, achene length, and
its stout bristles.

25. Perityle palmeri S. Wats. Proc, Amer. Acad. 24: 57.
1889. TYPE: Mexico: SONORA: Guaymas, Oct., 1887, E.
Palmer 308 (Holotype, aH !; isotypes, NY!, uc!, us (3) !).

1974] Perityle — Powell 297

Leptopharynx palmeri (S. Wats.) Rydb. N. Amer. Fl. 34:
23. 1914.

Plants fleshy-rooted perennials, low and spreading, semi-
erect, or semipendulent, 10-25 cm high, stems herbaceous
and rather brittle, often purplish, short or long pilose;
leaves mostly alternate, short or long pilose, grayish-green,
6-10 em long, 3.0-5.5 cm wide, cordate, cordate-ovate, to
cordate-orbiculate, the margins acutely shallow-lobed and
serrate-dentate, the lobes several, or the blades strongly
3-lobed, overall the blades may appear irregularly laciniate;
capitulescence of solitary heads borne on peduncles 2-6 (8)
cm long; heads radiate, often cernuous at maturity, 8-11
mm high, 6-13 mm wide, involucres hemispherical to cam-
panulate and truncate at the base; bracts 7-10 mm long,
linear to linear-lanceolate and attenuate; ray flowers nu-
merous, ligules yellow, 7-8 (10) mm long, oblong-elliptic ;
disc flowers numerous, corollas yellow, 4-6 mm long, throats
tubular; achenes 2.5-3.5 mm long, narrowly oblong, with
prominent callous margins, the margins densely long-ciliate,
faces glabrous or puberulent; pappus of a crown of squa-
mellae, 0.5-0.9 mm long, and a single bristle, 3-5 mm long,
the bristle stout, tapering from base to apex, and densely
barbellate; chromosome number, n = 17.

Primarily, if not entirely, soil-dwelling, most common in
the vicinity of Guaymas, Sonora, but also in northern Sina-
loa, Mexico. Flowering probably year around. (Fig. 5).

REPRESENTATIVE SPECIMENS:

Mexico: SINALOA: Topolobampo, Blakey (ARIZ.) SONORA: Guay-
mas, Brandegee (POM, UC); Plam Canyon, W side of Sierra Libre,
35 mi S of Hermosillo, Carter (ARIZ); San Pedro Bay, Craig 691
(POM); Guaymas, Dawson 1003 (ps, US); cliffs, E side of Guaymas,
Drouet and Richards 3846 (ps); 2.8 mi N of Guaymas, Lewis 5334
(sMU); 36 mi S of Hermosillo, Lindsay 1125 (ps); near Guaymas,
Palmer 1218 (us); Sahuaral Bay, Powell and Averett 1523 (SRSC,
TEX); 1 mi E of San Carlos Resort, Powell and Sikes 1687 (SRSC,
TEX); 6 km N of Empalme, Ripley 14307 (CAS); 1 mi E of San
Carlos Resort, Sikes and Babcock 169 (sRSC, TEX); 1 mi N of Naval
Base, Guaymas Bay, Sikes and Babcock 176 (SRSC, TEX).

298 Rhodora [Vol. 76

Perityle palmeri is best distinguished from P. lobata,
perhaps its closest relative, and from P. cordifolia by the
following characters: fleshy roots, weak perennial habit in
soils, lobing of the leaves, ligule length, bract length, and
pilose pubescence.

26. Perityle californica Benth, Bot. Sulph. 23. 1844.
TYPE: Mexico: BAJA CALIFORNIA: Magdalena Bay, 1837?,
R. B. Hinds (K!).

Perityle plumigera Harv. & Gray, Mem. Amer. Acad, II.
4: 77. 1849. TYPE: Mexico: SONORA: “California” (vicinity
of Hermosillo), Coulter 279 (Holotype, GH!; fragment,
uc!, us!).

Perityle deltoidea S. Wats. Proc. Amer. Acad. 24: 57.
1889. TYPE: Mexico: BAJA CALIFORNIA: Los Angeles Bay,
1887, E. Palmer 568 (Holotype, GH!; isotypes, NY!, UC!
US!),

Plants delicate or robust annuals, 6-35 cm high, erect or
decumbent, sparsely or densely leaved; leaves opposite or
alternate, glandular-puberulent to short pilose, 1-6 (10) em
long, 0.5-2.5 (6.0) cm wide, often purplish when mature,
deltoid, ovate, or cordate, 3-lobed to subcruciform with the
segments serrate-crenate or shallow-lobed, or the margins
merely shallow-lobed to serrate; petioles 0.3-1.5 (4.0) em
long; capitulescence of 1-5 heads borne on delicate pe-
duncles; heads radiate, 3.5-7.5 mm high, 3.0-5.0 (7.5) mm
wide, involucres narrowly to broadly campanulate; bracts
linear, oblanceolate, or obovate; ray flowers 10-15, ligules
yellow, oblong-elliptic; disc corollas yellow, 2.0-3.0 (3.5) mm
long, throats tubular to subfunnelform; achenes 1.5-2.5
(3.0) mm long, narrowly obovate, oblanceolate, to oblong-
oblanceolate, with thin or prominent callous margins, the
margins short or long ciliate, faces puberulent to densely
so; pappus of a crown of squamellae and a single bristle,
1.5-3.0 (3.5) mm long, the bristle barbellate to subplumose;
chromosome numbers, n — 13, 12, 11.

Common in desert soils, Sonora and Sinaloa of mainland

1974] Perityle — Powell 299

Mexico, and along most of the Baja California peninsula to
just S of La Paz. Flowering Dec.-June. (Fig. 6).

REPRESENTATIVE SPECIMENS:

MEXICO: BAJA CALIFORNIA. San Juan Mine, Moran 8117 (DS, Sb,
uc); Los Angeles Bay, Palmer 568 (GH, NY, UC, US); 4 mi S of
Rancho Rosarito, Powell and Turner 1808 (sRSC, TEX); El Arco, N
end of town, Sikes and Babcock 297 (SRSC, TEX); mesa S of Rancho
Mesquital, Wiggins 11326 (DS, GH, UC). BAJA CALIFORNIA SUR. Isla
Magdalena, Brandegee s.n. (GH, UC, US); 20 km E of San Ignacio,
Carter, Alexander, and Kellogg 1972 (ps, UC, US); 26 km NW of
San Ignacio, Carter, Alexander, and Kellogg 2521 (DS, GH, UC, US);
Arroyo del Cajon de Tecomaja, SW of Puerto Escondido, Carter and
Kellogg 2906 (uc); Rancho El Horno, NE of San Javier, Carter
and Ferris 3789 (uc); SE of La Soledad, Carter 5418 (uc); Arroyo,
4 mi above La Purisima, Constance 3146 (DS, GH, LL, US); Isla San
Jose, Moran 3785 (DS, SD, UC); Isla Santa Catalina, Moran 3867 (DS,
sp); Isla San Marcos, Moran 8983 (sp, US); Isla. Carmen, Moran
9133 (sp); Isla Danzante, Moran 9253 (sp); Santa Rosalia, Palmer
185 (CAS,GH, US); ca. 19 mi N of La Paz, Porter 414 (CAS, Ds, Uc);
20 mi S of El Arco, Powell and Sikes 1650 (SRSC, TEX); 0.5 mi S of
San Ignacio, Powell and Sikes 1653 (SRSC, TEX); ca. 8 mi S of Santa
Rosalia, Powell and Sikes 1657 (SRSC, TEX); 33 mi S of Santa Rosalia,
Powell and Sikes 1659 (SRSC, TEX); Bahia. de la Concepcion, Powell
and Sikes 1661 (sRSC, TEX); 10 mi W of Loreto, Powell and Sikes
1664 (SRSC, TEX); 13.6 mi S of Mulege, Powell and Turner 1835
(SRSC, TEX); 11.5 mi W of Loreto, Powell and Turner 1837 (SRSC,
TEX); 15 mi W of San Javier, Powell and Turner 1845 (SRSC, TEX) ;
3 mi E of La Paz, Sikes and Babcock 216 (SRSC, TEX); near km
40 on hwy between La Paz and Santo Domingo, Thomas 8455 (CAS) ;
Isla Partida, Wiggins, Carter, and Ernst 412 (DS); Commondu,
Wiggins 5502 (CAS, DS, GH, NY, UC, US); 22 mi N of San Ignacio,
Wiggins 7888 (DS, GH, TEX, UC, US); 20 mi S of Mulege, Wiggins
11408 (CAs, ps, GH, UC, US); 6 mi W of Canipole, Wiggins 11489
(CAS, DS, GH, Uc, US); Llano Caquihui, W of Los Dolores, Wiggins
15516 (ps, US); 16 mi E of La Paz, Wiggins 15625 (Ds, us); Isla
Carmen, Wiggins 17497 (ps). SINALOA. N of Topolobampo, Flyr 72
(TEX); vicinity of San Blas, Rose, Standley, and Russell 13244
(NY, US); vicinity of Fuerte, Rose, Standley and Russell 13472 (NY,
us); W side of Topolobampo Harbor, Sikes and Babcock 189 (SRSC,
TEX). SONORA. 8 mi N of Hermosillo, Breedlove 1758 (ns); San
Pedro Bay, Craig 662 (PoM); E side of Guaymas, Drouet and
Richards 3844 (CAS, DS, GH, US); 73 mi S of Hermosillo, Flyr 98A
(TEX); 47 mi N of Hermosillo, Frye and Frye 2305 (GH, us); 1 mi
E of San Carlos Resort, Powell and Sikes 1686 (SRSC, TEX); 1-2 mi

300 Rhodora [Vol. 76

Fig. 6. Distribution of P. californica (closed circles); the bars
indicate populations from which chromosome numbers were taken,
with (1) n — 11 II, (2) n = 12 II, (3) n — 13 II, and (4) n —10
II + 1 III; all mainland counts have been n= 13. P. cuneata var.
cuneata (open circles) ; P. cuneata. var. marginata, (closed triangles).

1974] Perityle — Powell 301

N of San Carlos Resort, Powell and Sikes 1691 (SRSC, TEX) ; vicinity
of Navojoa, Rose, Standley, and Russell 13140 (Ny, US); 110 mi
E of Hermosillo, Sikes and Babcock 151 (SRSC, TEX); 32 mi N of
Hermosillo, Shreve 10053 (LL); 10 mi N of Hermosillo, Wiggins 6250
(ns, POM, US); 5 mi SE of Torres, Wiggins 6273 (DS, GH, POM, US).

In her discussion of P. californica, Everly (1947) em-
phasized the variable habit of this taxon. The species oc-
curs in Baja California and in Sinaloa-Sonora of mainland
Mexico. Most of its morphological variability is expressed
in plant size, number of leaves, and leaf shape. Floral char-
acteristics are essentially the same throughout the distribu-
tional range.

Chromosomal variability also exists in Perityle californica
where numbers of n — 13, 12, 11, and 10 II + 1 III have
been reported (Powell, 1968; 1972b). Some distributional
and morphological correlation with chromosome number
has been observed, although no taxonomic status is war-
ranted for the cytotypes. Plants of the mainland popula-
tions are n = 13, as so far determined, and tend to be
rather tall and robust with deeply lobed (often trilobed)
leaves. In wet years P. californica is a common weed in
some parts of Sonora and Sinaloa. Chromosome numbers
of n = 13, 12, 11, and 10 II + 1 III have been found in
Baja California populations (Fig. 6), and the plants are
usually small with leaves not deeply lobed (mostly ser-
rately-lobed). I have found the Baja California populations
to be invariably small and scattered, but I have not seen
the peninsula in a wet year. Populations with » — 11 are
most common in Baja California and probably extend
throughout the range of the species (Fig. 6). One popula-
tion with n = 12 has been found, and those with n = 10
II + III presumably arose through hybridization of cyto-
types with n = 12 and n = 11 (Fig. 6). The one peninsula
collection with » — 13 (Fig. 6) could have been introduced
from the mainland, or seemingly less likely at present, could
represent the presence of native n — 13's on the peninsula.
It seem reasonable to conclude that P. californica in Baja
California evolved by aneuploid reduction in chromosome
number from mainland cytotypes.

302 Rhodora [Vol. 76

Perityle californica does not share close or obvious rela-
tionship with any species, and its origin as a low-numbered
aneuploid is not clear. The species is superficially similar
to P. cuneata (n = 16, 12), but these taxa differ consider-
ably in floral characters, particularly of the achenes and
pappus. Indeed, the floral] differences between these two
aneuploid species are significant enough to cast considerable
doubt upon their common origin. Perityle californica has
narrow oblanceolate achenes with a single, long, subplumose
pappus bristle, while P. cuneata has broad, obcordate or
obovate achenes with two delicate pappus bristles. The
morphology of P. californica (including flower color and
achene and pappus features) resembles that of the P. cordi-
folia group (x = 17), and possibly originated therefrom by
aneuploid reduction.

Certain aspects of the Perityle californica synonymy are
somewhat confusing, but a good explanation of the histori-
cal situations was given by Everly (1947).

27. Perityle cuneata Brandeg. Zoe 1: 54. 1890. TYPE:
Mexico: BAJA CALIFORNIA: Sierra de Laguna, near Todos
Santos, 21 Jan. 1890, T. S. Brandegee (Holotype, UC!; iso-
type, US!).

27a. Perityle cuneata var. cuneata.

Plants robust annuals, 25-50 (80) cm high, erect or de-
cumbent, sparsely to densely leaved; leaves opposite or
alternate, glandular-puberulent to nearly glabrous, 3-8 (15)
em long, 1.5-4.5 (11.0) cm wide, often purplish when ma-
ture, cordate to broadly so, ovate, or subhastate, the mar-
gins serrate-lobed to serrate-crenate; petioles 0.6-2.0 (5.0)
em long; capitulescence of 1-3 heads borne on rather stout
peduncles; heads radiate, 5-8 (10) mm high, 6-10 (12) mm
wide, involucres campanulate to hemispherical; bracts ob-
lanceolate to obovate and attenuate; ray flowers 10-15,
ligules yellow, oblong-elliptie; disc corollas yellow, 3.0-3.5
mm long, throats tubular; achenes (2.5) 3.0-3.8 mm long,
obcordate-cuneate to oblanceolate, with broad callous mar-

1974] Perityle — Powell 303

gins, often purplish, the margins puberulent, faces glabrous
to puberulent; pappus of a crown of squamellae and 2
(rarely 0-2) delicate bristles, 0.5-1.2 (2.0) mm long; chro-
mosome number, n = 12.

Infrequent in soils, Cape Region of Baja California Sur
to just S of La Paz, most common on eastern side. Fower-

ing Jan.-Sept. (Fig. 6).

REPRESENTATIVE SPECIMENS:

Mexico: BAJA CALIFORNIA SUR: San Jose del Cabo, Brandegee (DS,
GH, US) ; Sierra San Francisquito and La Chuparosa, E side of Sierra
de la Victoria, Carter and Ferris 3386 (DS, SD, UC); ca. 3 km above
Boca de la Sierra, Moran 7091 (CAS, DS, GH, SD, TEX); Potrero de
Almenta, S fork of Canyon San Pedro, Moran 7387 (CAS, DS, SD) ;
ca. 5 mi S of San Antonio, Powell and Sikes 1667 (SRSC, TEX) ; Boca de
la Sierra, Powell and Sikes 1671 (sRSC, TEX) ; 2 mi S of San Antonio,
Sikes and Babcock 233 (SRSC, TEX); ca. 2 mi N of San Antonio,
Sikes and Babcock 265 (sRSC, TEX); ca. 6 mi SW from Santiago,
Thomas 7715 (CAS, DS, SD, US); ca. 5 mi SW of La Palma and ca.
8 mi NW from Santa Anita, Thomas 7738 (DS, SD, UC).

The two varieties of Perityle cuneata are morphologi-
cally similar, although extreme forms of these taxa do ex-
hibit considerable differences. Everly (1947) recognized
P. cuneata and P. cuneata, var. marginata as conspecific.
In addition to the morphological differences, I have recog-
nized var. cuneata, (n = 12) and var. marginata (n = 16)
because of their chromosome numbers, Indeed, the chromo-
somal differences suggest that further study might demand
the elevation of var. marginata to specific rank.

Besides its chromosome number, Perityle cuneata var.
cuneata is characterized by robust habit, large cordate
leaves, large heads, tubular disc corollas, obcordate-cuneate
achenes, and attenuate involucral bracts. The plants of var.
marginata are usually smaller with smaller leaves, smaller
heads and other floral characters, campanulate-funnelform
dise corollas, obovate achenes, and acute involucral bracts.
In addition, var. cuneata usually occurs in higher hills and
canyons of the Cape Region while var. marginata occurs
mostly in sandy places not far above sea level.

304 Rhodora [Vol. 76

Perityle cuneata does not exhibit a close morphological
relationship with any other species. Superficially, P. cu-
neata most resembles P. californica, but it possibly evolved
independently of the latter taxon by aneuploid reduction
from the P. cordifolia line.

27b. Perityle cuneata var. marginata (Rydb.) I. M. Johns-
ton, Proc. Calif. Acad. IV, 12: 1204. 1924.

Perityle marginata Rydb. N. Amer. Fl. 34: 14. 1914.
TYPE: Mexico: BAJA CALIFORNIA: San Jose del Cabo, 25
Mar. 1911, J. N. Rose 16459 (Holotype, NY!; fragment,
UC!; isotypes, GH !, US!).

Plants delicate or robust annuals, 6-40 cm high, erect or
decumbent; leaves 1.5-5.0 (10) em long, 0.9-3.0 (5.0) cm
wide, cordate, ovate, or deltoid, the margins doubly serrate-
crenate to shallow-lobed; petioles 0.2-1.8 (4.0) em long;
heads radiate, 4-7 mm high, 4-7 mm wide; bracts subovate,
oblanceolate, or linear-lanceolate and acute; ligules yellow;
dise corollas yellow, 2 (2.5) mm long, throats narrowly
campanulate, campanulate-funnelform, rarely subtubular;
achenes 1.5-2.0 (2.5) mm long, obovate, subcuneate, rarely
almost oblong, with prominent callous margins, the margins
short or long-ciliate; pappus of a crown of squamellae and
2 (rarely none) delicate bristles, 0.5-1.0 (1.5) mm long;
chromosome number, n = 16.

Infrequent in desert soils, Cape Region of Baja California
Sur and N to near La Paz, most common on southern and
western sides. Flowering probably year around. (Fig. 6).

REPRESENTATIVE SPECIMENS:

Mexico: BAJA CALIFORNIA SUR. San Jose del Cabo, Jones s.n. (NY,
POM, SD, UC); Arroyo de los Pozos, Moran 6886 (CAS, DS, GH, TEX) ;
2.5 km NE of Cabo San Lucus, Moran 7049 (CAS, DS, GH, SD) ; 6.4 mi
N of Todos Santos, Porter 79 (CAS, DS, UC); 1.4 mi W of El Coyote,
Porter 118 (CAS, DS, UC); ca. 17 mi S of San Antonio, Powell and
Sikes 1668 (SRSC, TEX); ca. 9 mi S of San Bartolo, Powell and Sikes
1670 (SRSC, TEX); 26 mi NE of Todos Santos, Powell and Sikes 1680
(SRSC, TEX); 2 mi NE of Eureka, Powell and Turner 1848 (SRSC,
TEX); ca. 1 mi N of Pescadero, Sikes and Babcock 255 (SRSC, TEX) ;
near km 40 between La Paz and Santo Domingo, Thomas 8455 (Ds,

1974] Perityle — Powell 305

GH); 5 mi W of San Jose del Cabo, Wiggins 14370 (CAS, DS, TEX, UC) ;
8 mi E of Cabo San Lucas, Wiggins 14645 (CAS, DS, GH, TEX, UC).

The NY (holotype) and UC specimens of the type collec-
tion (Rose 16459) are typical of var. marginata, while the
US and GH specimens (isotypes) approach var. cuneata. It
is possible that the type collection may be mixed, containing
plants of both varieties.

Some of the small forms of var. marginata are so like P.
californica superficially that the two taxa can not be told
apart except by microscopic examination of the achenes
and pappus. The Thomas 8455 collection includes plants
of both var. marginata and P. californica, but I have not
otherwise noted sympatric occurrence of the taxa. No
interspecific hybridization has been detected.

LITERATURE CITED

ALSTON, R. E., & B. L. TURNER. 1963. Natural hybridization among
four species of Baptisia (Leguminosae). Am. Jour. Bot. 50:
159-173.

EvERLY, M. L. 1947. A taxonomic study of the genus Perityle and
related genera. Contrib. Dudley Herb. 3: 375-396.

JOHNSTON, I. M. 1929. In: Papers on the Flora of Northern Chile.
Contrib. Gray Herb. 85: 127-128.

Moore, D. G., & E. C. BUFFINGTON. 1968. Transform faulting and
growth of the Gulf of California since the late Pliocene. Science
161: 1238-1241.

NiLES, W. E. 1970. Taxonomic investigations in the genera Perityle
and Laphamia (Compositae). Mem. N. Y. Bot. Gard. 21: 1-82.

PowELL, A. M. 1968a. Additional discussions pertaining to the con-
generic status of Perityle and Laphamia (Compositae). Sida 3:
270-278.

1968b. Chromosome numbers in Perityle and related
genera (Peritylinae — Compositae). Am. Jour. Bot. 55: 820-828.
1969. Taxonomy of Perityle section Pappothrix
(Compositae — Peritylinae). Rhodora 71: 58-93.

. 1970. Natural intersectional hybridization in Peri-
tyle (Compositae). Brittonia 22: 3-10.

1972a. Taxonomy of Amauria (Compositae — Peri-
tylinae). Madroño 21: 516-525.

1972b. Peritylinae, In IOPB Chromosome Number
Reports. Taxon 21: 344.

306 Rhodora [Vol. 76

1972c. Artificial hybridizations in the subtribe Peri-

tylinae (Compositae — Helenieae). Am. Jour. Bot. 59: 760-768.

1972d. Taxonomy of Pericome (Compositae — Peri-

tylinae). Southwest. Nat. 18: 335-339.

1972e. New species of Perityle and Amauria (Com-

positae). Madroño 21: 456-458.

1973. Taxonomy of Perityle section Laphamia

(Compositae — Peritylinae). Sida 5: 61-128.

, & S. Sikes. 1970. Chromosome numbers of some

Chihuahuan Desert Compositae. Southwest. Nat. 15: 175-186.

,& H. S. Tsang. 1966. Preliminary chromatographic
studies of Laphamia and related genera of Compositae. South-
west. Nat. 11: 190-195.

RAVEN, P. H. 1963. Amphitropical relationships in the floras of
North and South America. Quart. Rev. Biol. 38: 151-177.

RYDBERG, P. A. 1914. Carduaceae — Helenieae — Peritylinae. N.
Am. Fl. 34: 11-27.

SHINNERS, L. H. 1959. Species of Laphamia transferred to Perityle
(Compositae — Helenieae). Southwest. Nat. 4: 204-209.

SOUTHWICK, L., T. J. MABRY, J. AVERETT, & A. M. POWELL. 1972.
Penduletin 4'-0-Methyl Ether, a new flavonoid from Perityle
vaseyi (Compositae). Phytochemistry 11: 2351.

STEBBINS, G. L. 1950. Variation and evolution in plants. Columbia
Univ. Press, New York.

TURNER, B. L. 1966. Taxonomy of Eutetras (Compositae — Heleni-
eae). Southwest. Nat. 11: 118-122.

TURNER, B. L. & D. FLYR. 1966. Chromosome numbers in the Com-
positae. X. North American species, Am. Jour. Bot. 53: 24-33.

DEPARTMENT OF BIOLOGY
SUL ROSS STATE UNIVERSITY
ALPINE, TEXAS 7983

CAREX EXILIS, GEOCAULON LIVIDUM AND
OTHER PLANTS OF INTEREST IN PITTSBURG, NEW
HAMPSHIRE. South Bay Bog at the southern end of First
Connecticut Lake in Pittsburg, Coos County, New Hamp-
shire apparently was not visited by A. S. Pease, S. K.
Harris and other botanists who collected extensively in
Coós County. The bog is very extensive, much of it open
but with scattered clumps of stunted Black Spruce and
Tamarack. Lying between 1600 and 1700 feet elevation,
it appears in every way to be boreal in character with
the following species of shrubs in abundance, Ledwm
groenlandicum, Rhododendron canadense, Andromeda glau-
cophylla, Kalmia polifolia, K. angustifolia, Cassandra. caly-
culata, Pyrus melanocarpa, Viburnum cassinoides and
Vaccinium angustifolium.

Bog Branch of Cedar Stream flows through or along the
northern edge of South Bay Bog and near it, in sites where
some drainage appears to run off the adjacent upland, there
are scattered trees of Thuja occidentalis L., some up to
30 inches d.b.h. These represent remnants of the primeval
forest since, along with dying trees of large size, there are
fallen ones in all stages of decay. Several species of orchids
grew in the immediate vicinity of trees of Arbor Vitae, the
rarest of which was Listera convallarioides (Liv.) Nutt.
A plant here of perhaps greater interest was Epilobium
palustre L. var. palustre of which Seymour (1969) states
“Often with Thuja,” but of which he cites only one locality
in New Hampshire, “Mt. Washington." Pease (1964) states
its habitat as “Boggy places, especially on the high moun-
tains,” but he gives two localities for it at lower elevations
both probably with some “Cedar” associated, “Stewarts-
town, Cedar bog” and “Columbia, near Lime Pond.”

In the more open part of South Bay Bog dominated by
sphagnum moss and the previously mentioned shrubs and
dwarf conifers there are plants of still greater interest to a
taxonomist, one being Carex exilis Dewey and the other
Geocaulon lividum (Richards.) Fern. Carex exilis is a sedge
of local but wide distribution in New England, it being

307

308 Rhodora [Vol. 76

found occasionally in all States except Connecticut. Fer-
nald (1950) omits mention of New Hampshire and Vermont
in his statement of range for the species though he does
mention Massachusetts, while Seymour (1969) gives but
one township for it in New Hampshire, “Tamworth” where
it was collected by F. L. Steele in a bog near Chocorua
Lake and where he has shown it to me. Specimens of it are
in the herbaria of the New England Botanical Club (NEBC)
and the University of New Hampshire (NHA). However
also at NHA are two other collections of fairly long stand-
ing, the earlier being from Windham in Rockingham County
collected on June 14, 1897 by W. S. Harris. This record
was included in the list published by Hodgdon and Fried-
lander (1949). The second collection of Carex exilis at
NHA is from a bog in Pittsburg known as “Moose Pasture”
lying along East Inlet about two miles northeast of Second
Connecticut Lake. This was collected by me in company with
Peter Allen on July 23, 1960 and bears my collection num-
ber 11636. Pease (1964) did not include C. exilis in his
"Flora" and it is quite possible that this specimen of it
was not available for inspection at the time that Pease
was gleaning additional records for Coós County and Harris
was carefully checking the manuscript before publication.
The specimens of C. exilis observed at South Bay Bog on
our visit of June 8, 1973 were completely dioecious, the
clones or colonies of female individuals being widely
separated from the males.

The other species of outstanding interest at South Bay
Bog was Geocaulon lividum. I have been quite familiar
with the Northern Comandra from the frequency of its
occurrence in bogs on Campobello Island, New Brunswick
and its presence on the Wolf Islands in the Bay of Fundy
where R. B. Pike and I have observed it. It seems strange
that a species inhabiting bogs at low elevations in coastal
eastern Maine and the Maritime Provinces should be con-
fined to montane areas inland though the same relationship
seems to hold for Rubus chamaemorus L. and Lycopodium
annotinum L. var. pungens (LaPyliae) Desv. Harris (1965)

1974] Pittsburg, N. H. Plants — Hodgdon 309

discussed the distribution of Geocaulon lividum in a critical
way, pointing out that its habitat in stations that he visited
was mostly damp but not boggy sites and that the lowest
elevation for it was at 2500 ft. on the Basin Rim in Coos
County. The station at South Bay Bog is in no way mon-
tane and the elevation is between 1600 and 1700 feet. It is
entirely possible that botanists have concentrated their
collecting more in the mountains than in the more prosaic
terrain of bogs that often lie between the mountains, thus
passing by an interesting intermediate type of habitat for
certain species that are supposed to occur only in coastal
areas or near the tops of mountains. In any event it is
evident that Geocaulon lividum should be searched for
widely as Stuart Harris pointed out, not only as he sug-
gested, in montane situations but also in bogs at inter-
mediate elevations in Northern New England.

Specimens of Carex exilis and Geocaulon lividum have
been deposited in the herbaria of the New England Bo-
tanical Club (NEBC) and the University of New Hampshire
(NHA).

These discoveries would not have been possible except
for the expert guidance of Dr. Peter Allen. In addition
I have borrowed freely in preparing this article from notes
that he made or that we recorded jointly on the occasion
of our visit to South Bay Bog on June 8, 1973.

LITERATURE CITED

FERNALD, M. L. 1950. Gray’s Manual of Botany, eighth edition,
American Book Company, N. Y. 1682 p.

Harris, S. K. 1965. Geocaulon lividum in New England. Rhodora
67: 407-409.

HopGDoN, A. R. and HERBERT FRIEDLANDER. 1949. Additions to the
Harris “Flora of Windham, New Hampshire.” Rhodora 51: 110.

PEASE, A. S. 1964. A Flora of Northern New Hampshire, New
England Botanical Club Inc. Cambridge, Mass. 278 p.

SEYMOUR, F. C. 1969. The Flora of New England. The Charles
Tuttle Co., Rutland, Vermont. 596 p.

A. R. HODGDON
DEPARTMENT OF BOTANY
UNIVERSITY OF NEW HAMPSHIRE, DURHAM 03824

NEW STATION FOR NUPHAR MICROPHYLLUM IN
DURHAM, NEW HAMPSHIRE. Pittsfield, N. H. (Mer-
rimack County) is no longer the southernmost collection
point in New Hampshire for Nuphar microphyllum (Pers.)
Fern. as reported in my article *Some Morphological Aids
in Distinguishing Nuphar microphyllum from Similar
Aquatics” (Rhodora 75: 65-74, March 1973). On October 8,
1973 while A. R. Hodgdon and I were scouting the Lamprey
River in Durham as a possible field trip site for his course
in aquatic plants at the University of New Hampshire,
I discovered a colony of N. microphyllum growing with
N. variegatum and Nymphoides cordata in about two feet
of water in a small cove near the upper end of Moat Island.
Perhaps closer observation may reveal some plants of the
hybrid, Nuphar X rubrodiscum. Undoubtedly other sta-
tions for N. microphyllum exist but its relative rarity in
southern New Hampshire has made it unnoticeable to the
casual observer. The two specimens collected late in the
season are a bit ragged and will be kept as vouchers in
the NHA Herbarium. An attempt will be made to secure
more presentable material for the New England Botanical
Club Herbarium.

JOHONET C. WICKS
DEPARTMENT OF BOTANY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, N. H. 03824

310

TYPIFICATION OF CHAMAESARACHA CORONOPUS

The type of Chamaesaracha coronopus Was erroneously
listed as Berlandier 1494 (= 234) (Rhodora 75: 339).
The section referring to the type material should read:

Chamaesaracha coronopus (Dunal) Gray, Bot. Calif. 1:
540. 1876. Holotype (G!), [Texas, Bexar Co.], Rio
Medina, between Laredo and Bejar [San Antonio]. 1828.
Berlandier 1513 (= 253). Isotypes: GH! & K!.

The locality indicated on the holotype and original de-
scription is only between Laredo and San Antonio, then
Bejar, Mexico. An isotype at Kew, however, has Rio
Medina on the label. The Medina River flows southeast
from Bandera County into Medina County, then east into
Bexar County. The route taken by Berlandier likely
erossed the Medina River in Bexar County, about 12 miles
south of San Antonio.

JOHN E. AVERETT
DEPARTMENT OF BIOLOGY
UNIVERSITY OF MISSOURI-ST. LOUIS

RANGE EXTENSION OF WOODWARDIA AREO-
LATA. While I was walking through a swampy area close
to Spruce Hole Bog in Durham, N.H. on October 20, 1973,
a fern frond came to my attention that from afar resembled
an aberrant form of sensitive fern, Onoclea sensibilis. Upon
further investigation, however, what was found turned out
to be the fertile frond of Woodwardia areolata (L.) Moore,
the netted-chain fern, so named for its long sori arranged
in a row on each side of the secondary midrib.

The discovery of this fern is noteworthy when one real-
izes that only two stations for it have ever been recorded
from New Hampshire, both in Rockingham County: Rye
Beach by Walter Deane in 1886 (N.E.B.C.) and Seabrook
by A. A. Eaton in 1896 (N.E.B.C.) In the seventy-seven
years since those recorded findings, Woodwardia areolata
has not been found in New Hampshire. In addition, this
new station extends the range of this coastal plain species
by approximately twenty miles.

Shaded by two bushes of Vaccinium corymbosum, a total
of six plants have been located within an area of three feet.
The plants are growing on rotting stump matter and are
surrounded most of the year by a foot of water or more.

Such a find always leads one to speculate as to how
this fern came to occur at this location, how long it has
been there, and why it was not discovered long before this.

A voucher specimen of the Woodwardia areolata is de-
posited in the University of New Hampshire herbarium.

AMINTA KITFIELD

DEPT. OF BOTANY

UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824

EDITORIAL ANNOUNCEMENT

With this number the present editor-in-chief with twelve
and a half years of service in that capacity, is completing
his tenure. Also Dr. Radcliffe B. Pike, as a valued associate
editor during this entire period, is completing his editorial
connection.

We trust that the younger group of editors who are now
in charge will find the journal to be prospering. Many
changes in Rhodora have been made during the past twelve
years; we express the hope that the journal can continue
to be molded to fit changing needs.

The new editor-in-chief is Dr. Alfred Linn Bogle,
Associate Professor of Botany at the University of New
Hampshire. Please send all manuscripts and communica-
tions to Dr. A. Linn Bogle, Department of Botany and
Plant Pathology, University of New Hampshire, Durham,
New Hampshire 03824.

A. R. HODGDON

DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
UNIVERSITY OF NEW HAMPSHIRE, DURHAM, N. H. 03824

313

INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA

Manuscripts must be double-spaced or preferably triple-
spaced (not on corrasable bond), and a list of legends for
figures and maps provided on a separate page. Footnotes
should be used sparingly, as they are usually not necessary.
Do not indicate the style of type through the use of capitals
or underscoring, partieularly in the citations of specimens,
except that the names of species and genera may be under-
lined to indicate italics in discussions. Specimen citations
should be selected critically especially for common species
of broad distribution. Systematic revisions and similar
papers should be prepared in the format of “The System-
aties and Ecology of Poison-Ivy and the Poison-Oaks,"
W. T. Gillis, Rhodora 73: 161-237, 370-443. 1971, particu-
larly with reference to the indentation of keys and syno-
nyms. Papers of a floristic nature should follow, as far as
possible, the format of “Contribution to the Fungus Flora
of Northeastern North America. V.," H. E. Bigelow & M. E.
Barr, Rhodora 71: 177-203. 1969. For bibliographie cita-
tions, a recommended list of standard journal abbreviations
is given by L. Schwarten & H. W. Rickett, Bull. Torrey Bot.
Club 85: 277-300. 1958.

Volume 76, No. s06, including pages 151-314, was issued July 3, 1924.

CONTENTS: — continued

New Station for Nuphar microphyllum in Durham, New
Hampshire

Johonet C. Wicks eese eee eene eene erano peonia ei sinit 310
Typification of Chamaesaracha coronopus

John B: Averell .cictcocesdacsncsisccanscnectsiescscavencecusscesepsatstgseesteduccsese 811
Range Extension of Woodwardia areolata

Aminta Kit field ............. 4. eee eere eese eese eee esee estan nost tatto a ah taa a 312
Editorial Announcement ................ eere eene eene nnne ennas 813

Instructions for Contributors .............. eene 314

ena

Hovova

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by
ALFRED LINN BOGLE, Editor-in-Chief

ROLLA MILTON TRYON
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
RICHARD EDWIN WEAVER

Associate Editors

Vol. 76 September, 1974 No. 807
CONTENTS:
A Systematic Study of the Genus Kalmia (Ericaceae)

TORN. E DIIP aoieanna aia E e 315
Marine Algae of Rhode Island

R. D. Wood and Martine Villalard-Bohnsack .................. 399

A Biosystematic Revision of Lupinus lyallii
20) 08 OT es Ts canioniusnndctsvsensereaseesanssise A E 422

The Role of Some Haloragaceae in Algal Ecology
L, C. Colt, Jr., and C. Barre Hellquist ................... 446

Illinois Field and Herbarium Studies
Robert H. Mohlenbrock and Dan K. Evans ......... 460

(Continued on Inside Back Cover)

Che Nef England Botanical Club, Ine.
Botanical Museum. Oxford St.. Cambridge, Mass. 02138

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Rhodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 76 September, 1974 No. 807

A SYSTEMATIC STUDY OF THE GENUS KALMIA
(ERICACEAE)

JoHN E. EBINGER

A study of the cytology and genetics of the genus Kalmia
was started in 1961 by Dr. Richard A. Jaynes of the Con-
necticut Agricultural Experiment Station. At that time
little was known about the genetics of this group although
a few of the species are important ornamental shrubs. Dr.
Jaynes has now accumulated many living representatives
of most of the species due to the efforts of numerous bota-
nists and friends. As a result of his studies (Jaynes 1968a,
1968b, 1969, 1971b) information is now available concern-
ing cross compatibility, speciation, and possible evolution-
ary relationships within the genus.

As treated here, the genus is composed of seven species,
six varieties and numerous forms, both natural and culti-
vated. The genus is restricted to North America and Cuba,
being particularly abundant along the east and west coasts
and at higher elevations and latitudes. No monographic
study of the genus has been undertaken, but the species
have been adequately described in many floristic works, and
Small (1914) treated all of the species that are presently
recognized. Since one of the species is commonly cultivated
and all are probably poisonous, this taxon has also been
reviewed in horticultural and agricultural literature. This
present study is an attempt to bring together most of this

315

316 Rhodora [Vol. 76

information in the light of what is now know about the
geneties and cytology of the genus.

ACKNOWLEDGEMENTS

The author is grateful to the many people who helped in
the completion of this study. Dr. Richard A. Jaynes pro-
vided access to his herbarium specimens and living material
as well as his notes and extensive knowledge concerning
the cytology and genetics of the genus. The Connecticut
Agricultural Experiment Station provided space and equip-
ment for 4 months to examine and collect material. Also,
many individuals at Eastern Illinois University provided
helpful criticism and advice in the preparation of the manu-
script. The author is also indebted to the many curators
who sent specimens for examination. Over 8,000 specimens
were examined from 75 herbaria from the United States,
Canada, and Europe. In the lists of representative speci-
mens these herbaria are indicated by their abbreviations
following Lanjouw and Stafleu (1964).

HISTORY OF THE GENUS

References to mountain laurel, Kalmia latifolia, appear
in the early literature of the American colonies. The Cape
Cod “Rose-trees” of Henry Hudson's log (Purchas, 1625)
may have been this species (Holmes, 1956), and Smith
(1624) mentions that laurel occurs in Virginia. This spe-
cies was first described and figured by Plukenet (1696)
under the name Cistus sempervirens, described by Grono-
vius (1739) who called it “The Common Laurel/vulgarly
called Ivy," while the first color print was. figured by
Catesby (1743) under the name Chamaedaphne.

Because of their potential as ornamentals, some of the
species were introduced into Europe. According to Aiton
(1789) Kalmia latifolia and K. angustifolia were intro-
duced into England in 1734 and 1736 by Peter Collinson,
an English merchant and fellow of the English and Swedish
Royal Societies. A third species, K. polifolia, was intro-

1974] Genus Kalmia — Ebinger 317

duced in 1767. Catesby (1743) reported that K. latifolia
from Pennsylvania flowered in his garden at Fulham in
1740. Numerous horticultural forms have been developed
as a result of breeding and selection in Europe for the past
200 years.

Probably the first detailed account of the toxic character-
istics of the genus occur in Peter Kalm's journal (Benson,
1937) in which he described in detail the poisonous proper-
ties of the “Laurel Trees" (Kalmia latifolia). He also
described both K. angustifolia and K. latifolia as to their
characteristics, economic importance, habitat requirements
and general distribution.

The generic name Kalmia was first proposed by Chenon
(1751) in a doctoral thesis, and was based on specimens
that Peter Kalm had collected during his trip to the New
World. Kalmia latifolia and K. angustifolia are described
in this thesis, and Linnaeus (1753) included both in his
Species Plantarum. Later, Linnaeus (1754) described the
genus and placed it in his first order (Monogynia) of his
tenth class (Decandria).

The genus is now regarded as a relatively primitive mem-
ber of the Ericaceae. Drude ( 1889) placed it in the Tribe
Phyllodoceae Drude of the subfamily Rhododendroideae
Drude, and its position has not been changed by most
subsequent authors. The members of this tribe are gen-
erally evergreen plants, usually with buxoid or ericoid
leaves, with regular flowers, and with seeds that usually
lack appendages. Two microscopic characteristics of the
anthers have also been used to characterize the tribe: the
presence of resorption tissue involving the epidermis of
the anther and the absence of filaments among the pollen
tetrads. Copeland (1943) discussed in detail the morphol-
ogy and anatomy of the tribe Phyllodoceae and considered
the genera Rhodothamnus Reichenb., Kalmia, and Phyllo-
doce Salisb. to be relatively primitive, an opinion also ex-
pressed by Wood (1961). Studies of wood anatomy by Cox
(1948) indicate that the genus Kalmia is closely related to

318 Rhodora [Vol. 76

Rhodothamnus and that these two genera represent one of
the three lines of evolution in the Phyllodoceae.

There have been few nomenclatorial complications in the
genus, and only two other generic names have been pro-
posed. Kuntze (1891) used Catesby's pre-Linnean name of
Chamaedaphne and made wholesale transfers to it, while
Small (1914) divided Kalmia into two genera. He proposed
the genus Kalmiella for Kalmia hirsuta and its related
species that differ by having a deciduous calyx and flowers
borne singly in the axils of ordinary or somewhat reduced
leaves. In all other morphological and anatomical character-
istics these species do not differ from the other members
of the genus, and there is no real justifieation for their
removal. Except for Alain (1957), all subsequent authors
have treated these species as members of the genus Kalmia.

ECONOMIC IMPORTANCE OF KALMIA

The genus Kalmia is economically important mainly be-
cause a few of the species are commonly grown as orna-
mentals. By far the most important species is the moun-
tain laurel, Kalmia latifolia, which is commonly planted as
an ornamental throughout its natural range as well as in
other parts of North America and Europe where climate
and soil are satisfactory. This species varies widely in
flower color, foliage, and size, and many cultivars have been
named and described (Holmes, 1956; Dudley, 1967). Culti-
vars of mountain laurel had been named by 1840, and at
the present time at least 90 names have been applied to
different forms of the species. This species has been sug-
gested as a possible choice for the national flower of the
United States (Van Rensselaer, 1889), and presently it is
the state flower of Connecticut and Pennsylvania.

The sheep laurel, Kalmia angustifolia, is also becoming
an important cultivated plant. Unlike the mountain laurel,
this species can be propagated by cuttings. A number of
cultivated forms are available, and selections should become
available as soon as nurserymen become familiar with hand-

1974] Genus Kalmia — Ebinger 319

ling them. This species is not important for its wood as
few specimens exceed 3 feet in height, but it is economically
important as a weed. Smith, Hodgdon and Eggert (1947)
considered it an important weed in low-bush blueberry fields
and tried fire, continued cutting, and lime to control it.
Later, Trevett (1956) reported on the effects of 2,4-D and
24,5-T in controlling sheep laurel in blueberry fields.
Porcher (1869) found that the leaves and twigs of this
species will dye cotton a fine drab color if a copper mordant
is used.

The foliage of Kalmia latifolia is commonly gathered in
eastern United States for floral displays as well as for
Christmas decorations and other festivities, continuing a
tradition started in colonial days. As early as 1913, Britton
suggested that this species should probably be protected
since it was becoming rare in some areas due to indiscrimi-
nate collecting. Buttrick (1924), in an extensive article
concerning growth and cultivation of mountain laurel in
Connecticut, estimated that the amount of foliage of this
species used in New York City probably exceeded 1,000
tons per year and that the maximum amount used in the
United States for decorative purposes probably exceeded
10,000 tons per year. It is still commonly found in decora-
tions, but no figures are presently available concerning the
amount used. The foliage of mountain laure] can also be
used to dye cloth. Nicholson and Clovis (1967) found that
the boiled leaves of this species produced a yellow tan color
on wool if a chrome mordant was used.

In the past, the wood of the mountain laurel was occa-
sionally used to make small items. Peter Kalm in his journal
(Benson, 1937) mentions that the strong wood is carved
into weaver's shuttles, pulleys, spoons and trowels. Browne
(1857) considered the wood to be similar to that of the
European Box and that it was sometimes used for the
handles of light tools, for screws, boxes, and that it was
supposed to make good clarinets. He also mentioned that
the American Indians used the wood for making small
dishes, spoons, and other domestic utensils, and according

320 Rhodora [Vol. 76

to Benson (1937), this was the source of the common name
"spoonwood." Other uses include the making of rustic
furniture and as a substitute for brier pipes (Britton,
1913). Due to the small size of most mountain laurel
shrubs, the wood is rarely used at the present time except
locally for tool handles and novelties.

Kalmia latifolia, as a common understory shrub, is im-
portant in preventing water runoff and soil erosion. Studies
by Johnson and Kovner (1956) in the southern Appalachian
Mountains show that the cutting of dense stands of laurel
and rhododendron in hardwood stands decreased the evapo-
transpiration rate and correspondingly increased the annual
streamflow on the average of 2 inches per year.

Extracts from the leaves of most species of Kalmia were
at one time considered important as medicine, Barton
(1802, 1804) mentions that a saturated tincture prepared
from the leaves of Kalmia latifolia in a proof spirit could
be used for certain fevers. According to Rafinesque (1830)
preparations of this species were used as a narcotic, errhine,
antisyphilitic and antiherpetic, while Griffith (1847) con-
sidered it useful for fever, itching and in cases of hemor-
rhaging and dysentery. Wood and Bache (1851) considered
it useful mainly for cutaneous affections when applied as
an ointment or decoction. During the latter part of the 19th
century the leaves of K. latifolia were still commonly found
in drug stores. According to Paschkis (1881), they were
dried, rolled into a pipe-form and used principally as a
remedy for diarrhea. The leaves of K. hirsuta were used
in the south as a cure for itching and for mange in dogs
(Porcher, 1869), while Holmes (1884) reports that K.
angustifolia was commonly used by the Cree Indians of the
Hudson Bay region for bowel complaints and as a tonic.

PROPAGATION OF KALMIA

Numerous cultivars of Kalmia latifolia have been named;
no large scale nursery production of any of these forms
exists, however, because they have been considered difficult

1974] Genus Kalmia — Ebinger 321

to root, hard to graft, and are untrue from seed. As a
result, one common source of this species for ornamental
purposes has been native woodland stands. Recent studies,
however, indicate that it is possible to root cuttings of this
species and that selected types do breed true.

According to Jaynes (1971d) most Kalmia species can be
started from cuttings. He found that greenwood cuttings
of Kalmia angustifolia, K. hirsuta, K. polifolia, and K.
microphylla root readily under mist or a plastic tent. By
contrast, K. cuneata cuttings were difficult to root, while
K. latifolia was highly variable in its rooting ability. In this
latter species the ability to root is related to the age of the
stock plant, since 89% of the cuttings from 1-year-old
plants rooted, while only 21% of those from 4 or more
year-old plants rooted. Jaynes also reported that most of
the sterile hybrids can be propagated readily from cuttings,
even when K. latifolia is one of the parents. Jaynes (1972)
found that clones of K. latifolia vary in their ability to root
from cuttings and that cuttings of cuttings or young grafts
root more readily than cuttings from the original stock
plant.

Few attempts have been made to graft any of the species
of Kalmia. Trumpy (1893) had some success using side
grafts with current year’s wood of Kalmia latifolia, while
Jaynes (1971d, 1972) grafted current year’s growth of
K. latifolia onto stock of the previous year using cleft or
side-veneer grafts. The grafts were made in mid-June,
kept in plastic tents for 6 weeks, hardened for 2 weeks, and
overwintered in a lath house. The first report on the suc-
cess of layering K. angustifolia was probably by Curtis
(1795).

The propagation of Kalmia from seeds has been exten-
sively studied. Nichols (1934) found that Kalmia polifolia
seeds germinated only after stratification, while K. angusti-
folia, and K. latifolia seeds had a germination rate 5 times
higher after stratification for 12 weeks. In contrast, Jaynes
(1971c) reported that of these 3 species, only K. latifolia
required stratification, the others germinating without any

322 Rhodora [Vol. 76

pregermination treatment. Jaynes also found that K. cu-
neata, as well as K. latifolia, required stratification, but that
soaking the seeds in gibberellin for 24 hours created the
same response as stratification. The most difficult species
to propagate appears to be K. hirsuta, which germinated
best after a heat-humidity treatment (heating the seeds in
a moist chamber at 80°C for 30-60 minutes). Jaynes
(1968c) suggested that this response to high temperature
may have adaptive value and could have evolved as a result
of recurring ground fires. He also suggests that K. ericoides,
a closely related species that is native to sandy pine areas
of Cuba, may require a similar pregermination treatment.
Kalmia microphylla is similar to K. polifolia in that it needs
no pregermination treatment. The studies of Flemer (1949)
and Jaynes (1971c) indicate that all species of Kalmia
germinate best at 22°C under light in a mixture of sand,
ground sphagnum, and peat moss that has a pH of 4.2.

A mycorrhizal association, without which growth will
not occur, has been demonstrated for Kalmia latifolia, and
it is probably necessary for all species of the genus. Flemer
(1949) found that mountain laurel seedlings grew very
poorly in sterile culture without the associated endophyte,
but when the seedlings were combined with the fungal
associate they rapidly outgrew the controls. Microscopic
examination revealed hyphae sheathing the younger por-
tions of the roots and frequent hyphal connections with
mycorrhizal coils in the cortical cells. No attempt was made
to identify the fungus involved.

GENETICS OF THE GENUS KALMIA

Many species of Kalmia are sympatric, and most have
similar habitat requirements. Even so, no natural hybrids
have been reported, and recent studies suggest that such
hybrids are unlikely to occur. Jaynes (1968a, 1971d) made
all possible crosses, including reciprocals, between 6 species
of the genus (omitting only Kalmia ericoides) and found
that the barriers to gene flow are well developed. Most of

1974] Genus Kalmia — Ebinger 323

the interspecific crosses were difficult to complete since no
seed was set, the resulting seed failed to germinate, or the
seed produced albino or yellow-green seedlings. Stylar inhi-
bition of pollen tube growth appeared to be the most com-
mon reason for the lack of seed set. Only 9 combinations
yielded viable seedlings and all were sterile. By contrast,
intra-specific crosses between plants differing in flower color
and growth habit within and between geographie sources
showed no evidence of genetic barriers.

Little is known about the inheritance of traits in the
genus; the only detailed study involves flower color in
Kalmia angustifolia. In this species, Jaynes (1971b) found
that the presence of anthocyanin pigment is determined by
a single dominant gene, with the homozygous recessive
lacking red pigment in the leaves and stem and having
white flowers. This trait is carried in the population as a
heterozygote, but is rarely expressed. These white forms
are at a selective disadvantage in the wild as the seedlings
are weak, especially when small.

Inbreeding depression and self incompatibility have been
observed in some species of Kalmia. Jaynes (1968b) found
that self pollinated Kalmia latifolia and K. angustifolia set
85-90% less seed than cross pollinated plants, while K. poli-
folia, in contrast, showed no self incompatibility. The seed-
lings of all three species, however, showed inbreeding de-
pression. As measured by height growth, the vigor of the
outcrossed seedlings was generally twice that of the selfs.
Jaynes (1971a) also noticed inbreeding depression when
sibling red-budded mountain laurel was crossed. It is possi-
ble that compact seedlings of Kalmia species of potential
ornamental value can be obtained by selfing.

Few cytological studies of the genus have been under-
taken. Hagerup (1928) in an extensive cytological study
of the bicornes reported a chromosome number of n = 12
for Kalmia latifolia and n = 24 for K. glauca (K. poli-
folia). He found that the chromosomes of K. polifolia were
about half as large as those of K. latifolia, so that the
amount of chromatin in the two species seemed to be about

324 Rhodora [Vol. 76

the same. Later, Callan (1941) reported a chromosome
number of 2n — 44 for K. polifolia, which in light of recent
studies is an error. Jaynes (1969) studied cytologically all
of the taxa except for the Cuban species. He found that
with the exception of K. polifolia (n — 24), all had a chro-
mosome number of » — 12. No differences were found in
the size of the chromosomes, and their smallness prevented
a karyotype analysis.

POLLINATION IN KALMIA

The most distinctive feature of the genus is its pollen-
discharge mechanism. Near the middle of the corolla are
10 sacs that form small lobes on each keel of the flower bud.
Just before anthesis the anthers are pushed upward into
these sacs by the elongating filaments. As the corolla opens
the anthers are held in the sacs and are carried down and
outward, the elastic filaments being bent backward under
tension. When the flower is disturbed by a large insect, one
or more of the anthers are released from their sacs. The
filament, which is under tension, snaps the anther upward,
showering the insect with pollen. This mechanism was first
described by Sprengel (1793) and later by Hasskar] (1863)
for Kalmia latifolia, while Rothrock (1867) observed the
same mechanism in K. angustifolia, Studies by Penhallow
(1882) showed that the pollen of K. latifolia could be
thrown up to 14 cm. from the flower while in K. angusti-
folia it was thrown up to 8 cm. away. These authors sug-
gested that this was a method of insuring self-pollination
since the pollen was thrown toward the stigma of the flower.
Beal (1867) was the first to describe the way in which the
flowers were cross-pollinated by insects. He observed that
while a bumble bee was searching around the base of the
flower, the proboscis liberated the stamens which projected
their pollen onto the underside of the insect’s body. This
pollen was then rubbed onto the stigma of other flowers
visited. This process was also discussed by Gray (1876,
1879).

1974] Genus Kalmia — Ebinger 325

When an anther is released, the pollen tetrads are dis-
charged as a fine powder in some species, and in others as
a sticky net caused by the presence of viscid threads that
hold the tetrads together. According to Matthews and Knox
(1926) these extremely fine, non-cellular threads are prob-
ably derived from small quantities of microspore mother
cell protoplasm which are excluded from the pollen tetrads.
They also suggest that this feature is an adaptation that
facilitates pollination in relatively large, upright flowers
like those of the genus Rhododendron. These threads were
found in Kalmia latifolia by Ikuse (1954) and have been
observed by the present author in K. hirsuta and K. eri-
coides. Their presence or absence was used by Copeland
(1943) as a major characteristic in separating the tribes of
the subfamily Rhododendroideae, with the tribe Phyllodo-
ceae being the only one that consistently lacked these
threads. According to Wood (1961) it is now apparent that
viscid threads do occur in some species of Kalmia, Phyllo-
doce and Loiseleuria, all members of the Phyllodoceae.
Because of this, the usefulness of this characteristic to
distinguish tribes is doubtful.

Only a few species of insects have been observed polli-
nating Kalmia species. Beal (1867) listed bumble bees and
other Hymenoptera, but mentions no species. The only de-
tailed account was by Lovell and Lovell (1934) who col-
lected and identified the pollinators found on a population
of Kalmia angustifolia near Waldoboro, Maine. Their ob-
servations show that K. angustifolia is effectively cross-
pollinated by bees and that a total of 14 species of insects
is capable of springing the stamens. Of these, Bombus
ternarius and Andrena vicina were the most common visi-
tors to the flowers. During the present study, populations
of K. angustifolia and K. latifolia were examined in south-
ern Connecticut, and all insects found to release the stamens
were collected and identified. These studies were made for
2 hour periods, at various times each day, from June 15 to
June 25, 1970. In most instances the insect did not release
the stamen by just landing on the flower. Usually the sta-

326 Rhodora [Vol. 76

mens were sprung by the proboscis while searching for
nectar, but sometimes the legs set the stamens free. No
honey bees were found on the flowers, and none were re-
ported by Lovell and Lovell (1934).

Recently bees have been used by Jaynes (1971a) to polli-
nate caged plants of Kalmia latifolia and K. angustifolia for
the large scale production of seed. In these experiments
selected plants were caged before the flower buds opened.
Honey bees (Apis spp.) or bumble bees (Bumbus spp.)
were introduced into the cages when the flowers started to
open. The plants that were in cages with bumble bees set
large quantities of seeds (58 to 95% of the flowers forming
capsules), while plants in cages with honey bees set fewer
seeds (2 to 31% of the flowers forming capsules). Plants
that were in cages with no bees rarely set seed (less than
1% of the flowers forming capsules).

CHEMISTRY OF KALMIA

Catesby (1743) was one of the first to report on the
poisonous properties of members of this taxon. He found
that when cattle and sheep were deprived of better food
during severe winters, many died from eating the leaves of
Kalmia latifolia (as Chamaedaphne). An extensive report
by Kalm (Benson, 1937) mentions that young sheep die
immediately or become very sick after eating leaves of
Kalmia latifolia, but calves “were cured by giving them
gunpowder and other medicines.” According to Kalm,
larger domestic animals are also affected by the leaves.

Most of the literature on livestock poisoning by Kalmia
latifolia and K. angustifolia has been reviewed by Crawford
(1908), Muenscher (1957) and Kingsbury (1964). Some
experimental work has been done to determine the symp-
toms, dosage, and treatment of the poisoning by these spe-
cies. One of the first studies was made by Wood (1883),
who fed a decoction of K. angustifolia leaves to a sheep.
The symptoms were similar to those observed by Pritchard
(1956) who studied K. angustifolia poisoning in a flock of

1974] Genus Kalmia — Ebinger 327

Minnesota sheep. Experiments with K. latifolia by Cary
and Matthews (1903), Crawford (1908) and Marsh and
Clawson (1930) showed that all livestock tested were poi-
soned by this species. The western species, K. microphylla,
is also poisonous but, because of its habitat, is seldom
encountered by livestock. Fleming (1920) fed this species
to some calves and sheep, most of which became sick, while
Clawson (1933) found that the symptoms exhibited by
sheep and goats were similar to those observed in animals
fed K. latifolia and K. angustifolia. No information is
available concerning the poisonous properties of the other
members of the genus.

Catesby (1743), Kalm (Benson, 1937), Bigelow (1818),
and other early authors mentioned that deer can eat the
leaves of mountain laurel without suffering ill effects.
Forbes and Bechdel (1930, 1931) indicate that laurel is
eaten in considerable quantity by deer, especially in time of
food shortage. When restricted to a laurel diet, however,
deer become thin and weak and usually develop a mild con-
dition of rickets. Death results when the animals are fed
in excess of 1.29% of their live weight of laurel leaves.

All species of Kalmia are probably poisonous to humans
(Hardin and Arena, 1969), although no deaths have been
attributed directly to this genus. Bigelow (1818) and Grif-
fith (1847) have reported instances of poisoning in humans.
In these cases, a decoction made from the leaves of Kalmia
latifolia produced toxic symptoms. The tough leaves and
their bitter taste make it unlikely that anyone would eat
them. Also, the quantity required to produce illness is
greater than what a person would probably consume. Ac-
cording to Barton (1802) the Delaware Indians used the
leaves of K. latifolia for suicide.

It has been suggested by various authors that the honey
produced from Kalmia flowers is poisonous, but no definite
proof is available. Barton (1802) reported that the honey
from Kalmia angustifolia, K. latifolia, and K. hirsuta is
poisonous to humans and listed a few instances of suspected
poisoning by honey of these species. Miller (1895) and

328 Rhodora [Vol. 76

Pellett (1920), in contrast, doubted that mountain laurel
was responsible for poisonous honey. They felt that due to
the abundance of this species it would seem that the cases
of poisoning would be reported much more frequently if
there was good reason to suspect the honey from this source.
Some doubt is also expressed by Lovell (1926, 1951) since
bees are rarely found on mountain laurel. Howes (1949),
as well as Hardin and Arena (1969), indicate that Kalmia
latifolia honey is an extremely distressing narcotic, varying
in its effect in proportion to the quantity eaten. In none of
the reports is the source of the poisonous honey known for
certain, the genus Kalmia being suspected because other
parts of these plants are known to be poisonous.

Andromedotoxin, the poisonous compound found in the
genus Kalmia, is known to occur also in other genera of the
Ericaceae. Numerous attempts have been made to isolate
and determine the chemical structure of this compound.
Lasché (1889) reported the occurrence of andromedotoxin
in the leaves and twigs of Kalmia angustifolia and also in
the fruits of K. latifolia, while Matusow (1897) found what
he considered andromedotoxin in the roots of K. latifolia.
More recently Jacobs and Lloyd (1939) and Waud (1940)
isolated andromedotoxin from K. angustifolia. The chemical
structure of andromedotoxin has been studied by Wood,
Stromberg, Keresztesy and Horning (1954) and more re-
cently by Tallent, Riethof and Horning (1957). The latter
authors renamed the substance acetylandromedol and de-
veloped a procedure for detecting its presence in plant
extracts through the use of paper electrophoresis. Though
the chemical structure of andromedotoxin is still in doubt,
its empirical formula is probably C..H,,0..

FOSSILS OF KALMIA

Only a few fossil species of Kalmia have been named.
Most are described from leaf impressions, and except for
size, shape, and probable coriaceous texture, there is little
reason to consider them members of the genus Kalmia, or

1974] Genus Kalmia — Ebinger 329

members of the family Ericaceae. One living species is also
known from fossil remains.

Kalmia brittoniana Hollick (1892).

This fossil was first described from clay beds of the
Raritan formation, near Kerischerville, Staten Island, New
York and is Upper Cretaceous in age (Museum, Staten
Island Association of Arts and Science). The material il-
lustrated consists of parts of three oblong, entire, probably
coriaceous leaves, 0.7 X 2.7 cm., with prominent midribs
and no secondary veins. Hollick (1906) mentions that the
absence of secondary nervation, indicating a leaf of thick,
coriaceous texture, was what largely influenced his refer-
ring the leaf to the genus Kalmia. This species has also
been reported from the Upper Cretaceous deposits of North
Carolina (Berry, 1907, 1920) and Alabama (Berry, 1913,
1919). When he described Kalmia brittoniana from the
Tuscaloosa formation of Alabama, Berry (1919) suggested
that there is no reason for its placement in the genus
Kalmia, or even in the Ericaceae. He further sugested that
the fossil probably represents some Upper Cretaceous spe-
cies of the Proteaceae or the Myrtaceae.

Kalmia ? elongata Ashlee (1932).

This species was found in the Latah formation, Elk River,
Idaho, and is Upper Miocene in age (University of Idaho
collection No. 303). The specimen consists of two narrowly
elliptical, subcoriaceous leaves, 0.9 X 1.7 cm., with entire
margins and a petiole 2-4 mm. long. The leaves are super-
ficially similar to Kalmia microphylla, but since the midrib
is grooved and the secondary and tertiary veins are present,
this fossil must be considered a doubtful member of the
genus.

Kalmia elliptica Brown (1962).

This species was found in the Fort Union formation (Ilo
Post Office, Wyoming) and the Denver formation (Golden,
Colorado) and is Paleocene in age. The specimen illustrated
by Brown (1962) consists of a single elliptic leaf with
entire margins and secondary and tertiary veins that ir-

330 Rhodora [Vol. 76

regularly fork and loop near the margins. The specimen is
similar in outline to a typical leaf of Kalmia latifolia but
is about twice as large (7 X 17 cm.). The presence of the
well developed and large number of secondary and tertiary
veins, as well as their angle to the midrib, is not typical of
any present day species of Kalmia. It must therefore be
considered a doubtful member of the genus.

Kalmia saxonica Litke (1968).

This species was found at Brandis, northwest Saxony,
Germany, and is Lower Miocene in age. It consists of pieces
of cuticle with the upper epidermis occasionally attached.
'These fragments indicate that the leaf was entire margined,
1-1.3 em. wide, and had an obvious midvein. The cells are
polygonal with wavy margins, the stomata are round, 18u
wide, surrounded by a double cuticular ring and occasionally
by radiating cuticular striations. The basal portion of glan-
dular hairs are also present. The structure and arrange-
ment of the cells is similar to that found in Kalmia latifolia,
and it is possible that this fossil represents a member of
this genus or a closely related genus of the Ericaceae.
Similar cuticle remains were placed in the form genus
Kalmiophyllum by Kráusel and Weyland (1959) when they
described the species Kalmiophyllum marcodurense.

Kalmia polifolia, the Bog Laurel, has also been found in
fossi] form. It was first reported by Berry and Johnston
(1922, as K. glauca Ait.) from interglacial deposits at
Point Grey near Vancouver, British Columbia. The fossil
specimen illustrated is of the same shape and size as those
of living specimens of K. polifolia. Since the other fossil
plants found clearly indicate a bog habitat, it is very pos-
sible that this fossil is a species of Kalmia. Kalmia polifolia
has also been reported from Pleistocene lake deposits of the
upper Connecticut River Valley in northern New Hamp-
shire (Hollick, 1931). These fossils are post-glacial in age
and according to Hollick are representative of the flora that
migrated northward in the wake of the retreating Wiscon-
sin ice sheet. The other fossil species found in the same
deposits indicate a habitat and climate similar to that pres-

1974] Genus Kalmia — Ebinger 331

ently prevailing in the area. The plates in this article show
excellent examples of leaves very similar in shape and size
to those of living K. polifolia. As mentioned by Hollick,
however, positive identification would “merely represent a
matter of personal opinion" since the leaves of K. polifolia,
Andromeda polifolia, and Ledum groenlandicum appear al-
most identical in outline.

TAXONOMIC TREATMENT

KALMIA Linnaeus, Sp. Pl. 1: 391. 1753.
Kalmiella Small, Fl. Southeast. U. S. 886. 1908.
Chamaedaphne Catesby ex Kuntze, Rev. Gen. Pl. 2: 388.
1891, non Moench.

Evergreen or rarely deciduous shrubs or small trees with
pubescent to occasionally glabrous, terete or 2-edged twigs.
Lateral buds very small, flattened, with 2 exposed valvate
to overlapping scales. Leaves alternate, opposite, or in
whorls of 3, exstipulate, usually coriaceous, petioled to sub-
sessile, margins entire. Flowers solitary in the axils of
leaves, or in terminal or axillary corymbiform racemes, or
in terminal compound corymbs. Pedicels slender, terete,
usually pubescent and with stalked glandular hairs, sub-
tended by a pair of bractlets in the axil of a small leaf-like
bract. Flowers 5-merous, actinomorphic, with hypogynous
insertion. Calyx synsepalous, usually green and persistent
in fruit; lobes 5, longer than the tube. Corolla sympetalous,
white to pink, red or purple, with a short cylindrical tube
extending into a rotate limb that is shallowly 5-lobed and
commonly 10-keeled, the limb with 10 saccate depressions
in which the anthers are held under tension. Stamens 10,
shorter than the corolla; filaments usually filiform, glabrous
or pubescent, dorsifixed; anthers 2-celled, unappendaged,
short, ovoid, opening by large terminal slits; pollen dis-
charged in tetrads, with or without viscid strands. Ovary
superior, 5-celled, placentation axile, placentae massive with
numerous ovules; style slender, terete, nearly straight;
stigma unexpanded, slightly grooved. Fruit a subglobose

332 Rhodora [Vol. 76

to globose, 5-valved, septicidal capsule usually subtended by
the persistent calyx. Seeds numerous, small and light-
weight, testa tight or loose and extending well past the
ends of the remainder of the seed. A North American
genus extending from Alaska south in the mountains to
California and Utah, eastward through Canada to the At-
lantic Ocean and southward through eastern United States
to Florida and Cuba.

Lectotype Species: Kalmia latifolia L. (cf. Britton 1908).

KEY TO THE SPECIES OF KALMIA

A. Leaves opposite; inflorescence a terminal corymbiform

raceme.
B. Midrib of leaves lacking stalked glands; seeds less
than 1.5 mm long . .. ..... 1. K. microphylla.

B. Midrib of leaves with stalked glands on both sur-
faces; seeds more than 1.5 mm long. . 2. K. polifolia.
A. Leaves alternate or in whorls; inflorescence various.

C. Inflorescence a terminal compound corymb; leaves

usually more than 2 em broad. ..... 3. K. latifolia.

C. Inflorescence a simple corymbiform raceme or the

flowers solitary in the axils of the leaves; leaves
usually less than 2 cm broad.

D. Leaves more than 1.5 em long; inflorescence a

corymbiform raceme from the axils of the leaves.

E. Leaves in whorls of 3, evergreen. ..........
IDEM ... 4. K. angustifolia.

E. Leaves alternate, deciduous. .. 5. K. cuneata.

D. Leaves less than 1.5 cm long; flowers mostly soli-
tary in the axils of the leaves.

F. Leaves broad, flat, the margins only slightly
revolute; flowers scattered along the stem in
the leaf axils. .............. 6. K. hirsuta.

F. Leaves narrow, strongly revolute; flowers
clustered toward the ends of the twig giving
the appearance of a terminal corymbiform
raceme. ........... ls 7. K. ericoides.

1974] Genus Kalmia — Ebinger 333

1. Kalmia microphylla (Hooker) Heller, Bull. Torrey Bot.
Club 25: 581. 1898.

Low growing alpine plant to sparingly branched shrub
to 6 dm tall; branchlets terete to slightly 2-edged below the
leaves, glabrous to lightly pubescent; leaves opposite, sub-
sessile, the broad petiole to 2 mm long, the blades coriaceous,
ovate to oval to narrowly lanceolate, to 4 em long, 0.4-1.1 cm
broad, whitened beneath with small hairs, a band of white
hairs usually present on the upper surface on either side
of the midrib, midrib lacking stipitate glands, margins flat
to slightly revolute; inflorescence a few-flowered, terminal,
racemose cluster, each flower on a glabrous pedicel 1-3 cm
long, each pedicel subtended by a pair of bractlets in the
axis of a leaf-like bract; calyx pink to light green, 5-10 mm
broad, the lobes obtuse, margins ciliate; corolla rose-purple
(rarely white), 0.8-2.0 cm broad; anthers purple, 0.6-1.6
mm long; fruit subglobose, 4-7 mm broad, 3.5-6 mm long;
seeds 0.8-1.4 mm long, the testa extending past the ends of
the seed. Chromosome number n — 12.

Kalmia microphylla has rarely been considered specifically
distinct from K. polifolia, Hall (1912), when he transferred
the taxon to a variety of K. polifolia mentioned that “No
characters upon which the variety might be raised to spe-
cific rank have as yet been pointed out, except the variable
one of size in plant and leaf." Most present floristic works
maintain this taxon as a variety or subspecies of K. poli-
folia, though it has been considered a separate species by
Small (1914), Rydberg (1954), Hultén (1948) and Abrams
(1951). Hultén (1948) mentioned that it was with some
hesitation that he accepted K. microphylla as being specifi-
cally different from K. polifolia, but that in Alaska, there
does not seem to be any intermediate types. However, in
1968 he considered K. microphylla a subspecies of K. poli-
folia.

When these taxa are considered distinct the characters
of plant and leaf size are the most commonly used for
separation. Kalmia polifolia is a large shrub, 3 or more dm

384 Rhodora [Vol. 76

tall, the leaves are 2-4 cm long, 214 to 3 times longer
than broad, and the margins are usually strongly revolute.
Kalmia microphylla, in contrast, is a much smaller plant,
usually less than 1.5 dm tall, the leaves are usually less than
1.5 em long, less than 215 times as long as broad, and the
margins are little if at all revolute. In most instances, these
characters are sufficient to separate the two taxa; however,
variation and overlapping of characters do occur. Other
characters that have previously been overlooked are much
more reliable in separating the two species. In K. polifolia
clavate glands are present on the upper and lower surface
of the leaf midrib, and the seeds are 1.5 to 2.2 mm long,
while in K. microphylla these glands are absent, and the
seeds are usually less than 1.3 mm long.

Kalmia microphylla is the only species of the genus found
west of the Rocky Mountains. It extends from central
California north to Alaska, and east through the Yukon
and Northwest Territories to the extreme northwest corner
of Manitoba. It is primarily a low-growing alpine shrub
rarely exceeding 1 dm in height, but in bogs at lower eleva-
tions it may reach a height of 6 dm. The species consists
of two varieties, one restricted to alpine and tundra areas,
and one found in coastal lowland regions.

KEY TO VARIETIES

1. Leaves oval to broadly elliptic, usually less than 2 cm
long and less than 214 times as long as broad; calyx
less than 7 mm broad. .......... la. var. microphylla.

1. Leaves lanceolate, usually 2.5 cm long or longer, 21% to
4 times as long as broad; calyx over 7 mm broad. ....

gaa new wea nEy ecaey eenerhu names 1b. var. occidentalis.

1a. Kalmia microphylla (Hooker) Heller var. microphylla.

Kalmia glauca Ait. y microphylla Hook. Fl. Bor.-Amer. 2:
41. 1834. TYPE: swamps in the Rocky Mountains, Drum-
mond (K). Kalmia polifolia var. microphylla Rehder in
Bailey, Cycl. Am. Hort. 2: 854. 1900. (without basionym).
Kalmia polifolia var. microphylla (Hooker) Hall, Univ.

1974] Genus Kalmia — Ebinger 335

California Publ. Bot. 4: 201. 1912. Kalmia polifolia ssp.
microphylla (Hooker) Calder & Taylor, Canad. Jour. Bot.
43: 1398. 1965.

Sparingly branched alpine shrub usually less than 1 dm
tall (to 3 dm tall at lower elevations) ; nodes usually less
than 1 em apart; leaves ovate to oval to broadly elliptic, to
1.5 (rarely 2.5) cm long, less than 214 times as long as
broad; calyx pink to light green, 5-7 mm broad; corolla
rose-purple 0.8-1.2 (rarely 1.5) cm broad; fruit 4-5 mm
broad, 3.5-4.5 mm long; seeds 0.8-1.3 mm long.

Distribution: Mountainous regions of western North
America from California, Nevada, Utah and Colorado north
through the Rocky Mountains to the Yukon and Northwest
Territories.

Common name: Alpine Laurel, Small-leaved Kalmia.

REPRESENTATIVE SPECIMENS

UNITED STATES: Alaska: Skagway, Anderson 1608 (1sc),
Eastwood (GH). California: EL DORADO CO.: Echo Lake, Beatle &
Beatle 3959 (ARIZ, DUKE, NY). FRESNO CO.: Marie Lake, Kruckeberg
3487 (RM, WS). HUMBOLDT CO.: Trinity Summit, Davy & Blasdale
5823 (Us). INYO CO.: Pine Creek, Duran 3338 (GH, MICH, US).
MADERA Co.: Shadow Creek, Raven (NY). MARIPOSA CO.: Crescent
Lake, Congdon (MIN, WS). MODOC CO.: Dismal Swamp, Weatherby
1598 (NY, RM). MONO CO.: Treble Lake, Munz 19998 (NY). NEVADA
co.: Frog Lake, Sonne 438 (MU, PH, US). PLACER CO.: Squaw Valley,
Eggleston 21607 (GH, NY, US, WS). PLUMAS C0.: Long Lake, Hall
9325 (GH, RM, US). SHASTA C0.: Lassen Nat Park, Ferris & Lorraine
10459 (GH, RM, US). SISKIYOU CO.: Mt. Shasta, Cooke 11296 (GH,
osc), 17654 (WS, WIU). TULARE CO.: Sky Blue Lake, Howell 25555
(US). TUOLUMNE CO.: Unicorn Creek, Mason 698 (GH). Colorado:
BOULDER C0.: Brainard Lake, Green 279 (PENN). CLEAR CREEK CO.:
Loveland Pass, Robbins 732 (ARIZ). GILPIN C0.: Forest Lake, Roberts
(CM). JACKSON CO.: Cameron Pass, Beaman & Erbisch 1362 (MSC,
RM). JEFFERSON CO.: Geneva Creek, Livingston 298 (DUKE). LARIMER
co.: Estes Park, Cooper 116 (RM), 157 (ARIZ, MIN, NMC, RM).
ROUTT CO.: Routt Natl Forest, Weber 6991 (ws). Idaho: BLAINE
co.: Sawtooth Peaks, Macbride & Payson 3707 (GH, RM, US).
BOISE CO.: Middle Spangle Lake, Hitchcock & Muhlick 10130 (GH,
NY, RM). BONNER CO.: Priest Lake, MacDougal 292 (NY). BOUNDARY
co.: Mt. Roothaan, Daubenmire 44289 (WTU). CLEARWATER CO.:
Lolo Trail Rd., Joslyn 953 (1D), 954 (ID). FREMONT CO.: Henry Lake,

336 Rhodora [Vol. 76

Payson & Payson 2047 (cH, NY, RM). IDAHO CO.: Big Fog Mt.
Baker 14762 (1D). SHOSHONE co.: Little Lost Lake, Wilson 318
(GH, IDS, WS). VALLEY CO.: Boulder Lake, Boone 177 (1D). Montana:
BEAVERHEAD CO.: Odell Lake, Hitchcock & Muhlick 14915 (NY, WS,
WTU). CARBON CO.: Beartooth Mts., Cronquist 8068 (GH, NY, ws,
WTU). DEER LODGE CO.: Wate (MONT). FLATHEAD CO.: Johns Lake,
Standley 18513 (US). GALLATIN CO.: Spanish Peaks, Vogel (cu,
MONT, US). GLACIER CO.: Hidden Lake, Hitchcock 1905 (MONT, RM).
GRANITE CO.: Helena, Kelsey (MU, NY). LAKE CO.: McDonald Lake,
Hitchcock 18343 (RM, ws, WTU). MADISON C0.: Old Hollowtop,
Rydberg & Bessy 4654 (MONT). MISSOULA CO.: Lace Lake, Rossbach
& Rossbach 184 (WTU). PARK CO.: Lady of the Lake, Witt 1233
(MINN, NY, WS, WTU). POWELL CO.: Upper Tineup Creek, Trask 117
(MONT). RAVALLI CO.: St. Mary's Peak, Hitchcock & Muhlick 15332
(NY, WS, WTU). STILLWATER CO.: Stillwater, Hawkins 35502 (MONT).
SWEET GRASS CO.: Rainbow Lake, Hitchcock 16501 (CAN, IDS, NY, WS,
WTU). TETON C0.: Henry Mts., Williams (GH). Nevada: ELKO Co.:
Lamoille Lake, Holmgren 1404 (BRY, NY). WASHOE Co.: Mt. Rose,
Archer 6425 (ARIZ, GH, NY, RENO). Oregon: BENTON Co.: Three
Sisters, Vechten 212 (osc). CLACKAMAS CO.: Clackamas Lake, Bart-
lett & Grayson 791 (NY). CROOK COo.: Des Chutes Nat'l. Forest,
Ferris & Duthie 488 (RM). DESCHUTES CO.: Sparks Lake, Bellinger
(WILLU). DOUGLAS CO.: Umpqua, Ingram 1515 (ORE). JACKSON CO.:
Mt. Pitt, Peck 5184 (WILLU). KLAMATH CO.: Crater Lake Applegate
319 (US, wILLU), 345 (Us). LANE Co.: McKenzie Pass, Henderson
16315 (ORE). LINN C0.: Hensley Meadows, Coville & Applegate 1152
(US). MARION CO.: Russell Lake, Lee 44 (osc). UNION co.: Head
1298 (osc). WALLOWA CO.: Mirror Lake, Peck 1777 (NY, WILLU).
Utah: DUCHESNE CO.: Mt. Agassiz, Cottam & Biddulph 3703 (BRY).
SUMIT CO.: Motte Peak, Payson & Payson 5139 (CHRB, GH, PH, RM,
US, WS). UINTAH CO.: Leidy Peak, Maguire 17679 (RM). UTAH CO.:
Grandaddy Lakes Basin, Stutz 224 (BRY). Washington: CHELAN CO.:
Ice Lake, Morrill 332 ( MONT, ND, WTU). KITTITAS CO.: Mt. Stuart,
Elmer 1103 (GH, K, MIN, US, WS). KLICKITAT CO.: Falcon Valley,
Suksdorf (PH, ws, WTU). LEWIS CO.: Reflection Lake, Townsend
(WS). OKANOGAN CO.: Slate Peak, Ownbey & Meyer 2302 (an, IDs,
ISC, MIN, NY, OSC, US). PEND OREILLE CO.: Sullivan Lake Dist.,
Layser 866 (WS). PIERCE CO.: Mt. Rainier, Allen 96 (CAN, GH, K,
MIN, MSC, NY, US, WS). SKAMANIA CO.: Chiquash Mts., Suksdorf
(WS). WHATCOM CO.: Mt. Baker, Muenscher 8077 (ws). YAKIMA
co.: Bird Creek Meadows, Zuberbuhler (ws). Wyoming: ALBANY
co.: Brooklyn Lake, Nelson 9226 (GH, MIN, RM). BIG HORN CO.:
Upper Crater Lake, Lofgren 13 (MIN, NY, RM). CARBON CO.: Silver
Lake, Henry (CM). FREMONT CO.: Gannett Creek, Jozwik 386 (Gu,
RM). JOHNSON CO.: Powder R. Pass, Stephens 40827 (VDB). LARIMER

1974] Genus Kalmia — Ebinger 337

co.: North Park Range, Goodding 1839 (PENN). PARK CO.: Island
Lake, Truman 54298 (WTU). SHERIDAN CO.: Dome Lake, Pammel &
Stanton 62 (MIN). TETON CO.: Taggart Lake, Williams 1119 (ID,
MIN, OSC, RM). YELLOWSTONE NAT'L PARK: Artists Paint Pots,
Nelson & Nelson 6170 (CM, ISC, MIN, GH, NY, RM, US).

CANADA: Alberta: Banff Natl. Park: Bow Pass, Saunders 11
(rRT); Lake Louise, Brown 578 (GH, PH, US), McCalla 2162 (ALTA,
NY, US); Waterfowl Lake, Walker 2443 (PENN). Jasper Nat. Park:
Athabasca Falls, Moss 2752C (ALTA, GH); Maligne Lake, Scamman
2664 (GH); Amethyst Lake, MacFadden 3246 (No). Waterton Lakes
Nat! Park: Summit Lake, Moss 975 (GH, NY, US); Upper Twin
Lake, Blais & Nagy 1793 (CAN). Fairfax Lake, Damais & Anderson
2335 (ALTA); Mercoal, Woollven 6 (ALTA); Viewpoint, Damais &
Kempinsky 5683 (ALTA); Virginia Hills, Moss 12469 (ALTA), 12505
(ALTA, CAN). British Columbia: Ernest C. Manning Prov. Park:
Timberline Valley, Hardy 18501 (usc, v); Three Brother Mts.,
Underhill 782 (v). Garibaldi Prov. Park: Castle Towers, Davidson
(usc). Glacier Nat'l. Park: Bald Mt., Fogg, Jr. 1273 (PENN, UBC);
Eagle Mt., Rosendahl 944 (MIN). Kokanee Glacier Prov. Park:
Thompson 14447 (CAN, GH, MICH, NY, PH, US, WTU). Kootenay Natl.
Park: Ross Lake, McCalla 9603 (uBC, v); Emerald Lake, Brown
(TRT). Mt. Assiniboine Prov. Park: Magog Lake, Green 551 (PENN).
Mt. Revelstroke Nat. Park: Calder & Savile 10861 (Ny, US),
8884 (Nv, US). Tweedsmuir Prov. Park: Trew 17177 (v). Wells
Gray Prov. Park: Battle Mt., Ahti & Ahti 6975 (v, wTU). Yoho
Nat'l Park: Emerald Lake, Beer (cM); Selkirk, Shaw 41 (GH,
MIN, NY, PENN, US); Yoho Lake, Pease 22371 (MASS). Vancouver
Island: Mt. Arrowsmith, Taylor 164 (TRT); Mt. Albert Edward,
Hett 352 (v); Moat Lake, Underhill 299 (wTU). Atlin, Szezawinski
(UBC, V). Caribou Pass, Henry 608 (GH, ISC); Ft. Steele, Johnstone
(UBC, v); Lillooet, Beamish & Vrugtman 610710 (vBC); Mt. Hazel,
Slocomb (v); Penticton, Calder & Savile 10751 (NY, RM, UBC, V, WS) ;
Prince George, Holland (v); Sikanni Chief R., Henry 743 (PH);
Summit Lake Landing, Weber 2589 (GH, Ws); Skagit Valley, Macoun
72406 (ND, NY); Tenquille Lake, Beamish & Vrugtman 61010 (1D,
UBC, WTU). Manitoba: Churchill, Beckett M-45-5 (MIN), Ellis 1287
(Us), Gillett 1832 (Isc, us), Gillis 3244 (msc), Voss 8999 (MICH),
Walker 5090 (WIN); Tidal, Brown 178 (CAN, SASK). Northwest
Territories: Aberdeen Lake, Macpherson 327 (CAN); Beverly Lake
cabin, Kuyt 107 (CAN); Contwoyto Lake, Thieret 9415 (GH, US);
Ennadai Lake, Brown 1320 (CAN): Gordon Lake, Henderson 19
(CAN); Great Bear Lake, Porsild 17117 (cAN), Porsild & Porsild
3429 (CAN), 3509 (CAN, US); Great Slave Lake, Pike (K); Mac-
Millan Pass, Porsild & Breitung 11258 (CAN, GH, US) ; Maguse Lake,
Gussow 33 (CAN); Nueltin Lake, Harper & Porsild 2211 (CAN, PH),

338 Rhodora [Vol. 76

2246 (CAN), 2271 (MICH, MIN, PH), 2554 (MICH, MIN, NY); Small
Tree Lake, Maini & Swan 506 (RM, SASK); Thelon Game Sanctuary,
Carroll 65 (CAN); Yellowknife Highway, Thieret & Reich 7742
(CAN, MIN), 8121 (Us). Yukon: Granite Mt., Porsild & Breitung
10576 (CAN, ISC, GH, US, WTU); Mayo Dist., Bostock 102 (CAN);
McQueston Area, Campbell 493 (CAN); Mt. Sheldon, Porsild & Brei-
tung 11094 (CAN, GH, US, WTU), 11485 (CAN, GH); pass between
Teslin & Nisutlin R., Porsild & Breitung 11045 (CAN, Ny), 11148
(CAN); St. Elias Mt., Pearson & Porsild 67-166A (CAN); Yukon-
Mackenzie border, Calder & Kukkonen 27664 (Isc, NY); White Pass
Summit, Bolton (us).

Plant and leaf size in Kalmia microphylla var. micro-
phylla is controlled to some extent by the environment. In
the typical alpine form the leaves are extremely small,
usually less than 1.2 em long. The type specimen, which
represents this extreme, has leaves less than 1 cm long and
is extremely short, less than 1 dm tall. At lower elevations
the leaves are not uncommonly 2 em long and the plant 2 dm
tall. Rarely taller plants are found with leaves to 2.6 cm
long. These larger leaved forms are superficially similar to
K. polifolia and probably account for the variation in tax-
onomic treatment of K. microphylla in the past.

Kalmia microphylla and K. polifolia probably never come
into contact with each other. In the Northwest Territories,
however, the northern part of both their ranges, it is possi-
ble that the two taxa may occur together. The author has
seen very few specimens of K. polifolia from the Northwest
Territories, but this species is common in northern Alberta,
Saskatchewan, and Manitoba. A few specimens of K. micro-
phylla have been examined from the Northwest Territories,
and this species extends all the way to the northwestern
corner of Manitoba. No specimens, however, have been
observed from Alberta or Saskatchewan. Sterile hybrids
have been produced by crossing these two taxa (Jaynes,
1968a), but no specimens from northern Canada have been
found that are morphologically similar to these hybrids.

The only other taxon with which Kalmia microphylla var.
microphylla might be confused is K. microphylla var. occi-
dentalis. Both taxa occur in western North America and

1974] Genus Kalmia — Ebinger 339

their ranges overlap. The two are easily separated, how-
ever, since var. occidentalis has lanceolate leaves that are
usually more than 2.5 cm long and 214-4 times as long as
broad, while var. microphylla has oval to eliptical leaves
that are smaller and less than 215 times as long as broad.
Also, var. occidentalis has larger flowers, with the corolla
more than 1.5 em broad and the calyx 7-10 mm wide. Vari-
ety microphylla, by contrast, has smaller flowers, with the
corolla less than 1.5 cm broad (usually less than 1.2 cm
broad) and the calyx less than 7 mm wide.

Although both varieties are found in marshes, bogs, and
other open wet areas, it appears that they rarely come into
contact with one another. Of the var. microphylla speci-
mens examined, most were collected at relatively high
elevations aecording to the information available on the
herbarium labels. The elevation at which this taxon was
collected varied from an average altitude of 8,500 feet above
sea level in California (5,200 to 11,500), to about 10,000 in
Colorado, Nevada, Utah and Wyoming (7,000 to 12,500),
to about 6,500 in Oregon (5,800 to 7,700), and to about
6,000 in Washington, Alberta and British Columbia (3,000
to 7,300). The altitudes in parentheses indicate the mini-
mum and maximum elevations recorded from the labels.
The information on the labels of specimens of var. occi-
dentalis indicates that this species is found at lower eleva-
tions. Most of the specimens were from at or near sea level,
but a few had been collected as high as 3,000 feet.

Kalmia microphylla var. microphylla and K. microphylla
var. occidentalis apparently hybridize when they come into
contact. A few specimens from Vancouver Island and the
Cascade Mountains of Washington appear to be intermedi-
ate morphologically. These intermediate individuals are
robust plants, 2-4 dm tall, similar in habit to the large-
leaved var. microphylla. The leaves are 1.8-2.5 cm long,
usually more ovate than lanceolate, and about twice as long
as broad. The flowers are large, more typical of var. occi-
dentalis. A few hybrids have been produced by crossing
these varieties and they are highly fertile and set large

340 Rhodora [Vol. 76

quantities of seed (Jaynes, personal communication). Of the

few suspected hybrids found, the pollen stainability varied

from 10-90% normal pollen, indicating that some sterility

occasionally results.

1b. Kalmia microphylla var. occidentalis (Small) Ebinger,
coinb. et stat. nov.

Kalmia occidentalis Small, N. Am. Fl. 29: 53. 1914.
TvPE: Washington: foothills of Mt. Rainier, 1885, Mrs.
Bailey Willis (NY). Kalmia polifolia subsp. occidentalis
(Small) Abrams, Ill. Fl. Pacific St. 3: 302. 1951.

Sparingly branched shrub to 6 dm tall; leaves lanceolate,
flat to slightly revolute margins, 2-4 cm long, some of the
leaves always over 2 cm long and 215-4 times as long as
broad; calyx light pink, 7-10 mm broad; corolla usually
rose-purple (rarely white), 1.5-2.0 cm broad; fruit 5-7 mm
broad, 4.5-6 mm long; seeds 1-1.4 mm long.

Distribution: Along the western coast of North America
from southern Alaska south to Oregon.

Common name: Western Swamp Kalmia; Western Bog

Laurel.

REPRESENTATIVE SPECIMENS

UNITED STATES: Alaska: Admiralty Isl., Hibber 236 (UNM);
Annette Isl, Coville & Kearney 381 (vs); Douglas Isl, Thorne
10280 (UWM); Gravina Isl., Anderson 5352 (1sc), 24599 (CAN, ISC);
Kuiu Isl, Rigg 17 (wTU); Metlakatlan Isl, Emgstrom (WTU);
Mitkof Isl, Brown (ALA); Pleasant Isl, Reiners (HNH); Prince of
Wales Isl., Flett 1976 (vs); Revillagigedo Isl., McCabe 8590 (WTU);
Woronkofski Isl., Klein 486 (ALA); Yakobi Isl., Anderson 1389 (US);
Juneau, Anderson 6338 (ALA, BRY, CAN, GH, ISC, PH, RM, V) ; Menden-
hall, Anderson 88 (CHRB); Petersburg, Scamman 515 (ALA, GH);
Sitka, Anderson 160 (1sc, US); Wrangell, Walker & Walker 638
(CM, GH, NY, RM, US, ws); Yes Bay, Gorman 21 (K, NO, NY, US).
Oregon: CLATSOP CO.: Gearhart, Adair (osc); Knappa, Bouch
(WTU); Delmoor, Cooke 10828 (osc). Washington: GRAY HARBOUR
co.: Lake Quinault, Thompson 11429 (ALTA, GH, MONT, NY, PH,
US, WS, WTU). ISLAND CO.: Whidbey Isl, Smith 480 (ws, WTU).
KING CO.: Green Lake, Benson 1474 (WTU). MASON CO.: Lost Lake,
Jones 6527 (PH, V, WTU). PACIFIC CO.: Ilwaco, Foster 865 (ORE, US).
PIERCE C0.: Tacoma, Sloan (WTU). SAN JUAN CO.: Mt. Constitution,
Zeller & Zeller 1193 (CM, GH, MIN, NY, US). SKAGIT CO.: Mt.

1974] Genus Kalmia — Ebinger 341

Higgins, Thompson 9639 (ND, US, WTU). SNOHOMISH CO.: Marys-
ville, Grant (ARIZ, OAC, MU, RM, US). THURSTON CO.: Upper Valley
of the Nisqually, Allen 4 (CAN, K, MIN, MSC, NY, US, WS). WHATCOM
co.: Custer, Muencher & Muencher 6007 (PH, Us, WS).

CANADA: British Columbia: Aristazabal Isl, McCabe 3424
(wTU); Dewdney Isl, Schofield 70-27 (UBC); Goose Isl, Martin &
Guiquet (v); Hope Isl., Calder & Mackay 31240 (cM, MIN); Lulu Isl.,
Beamish & Vrugtman 60519 (Ny, osc, UBC); Graham Isl., Calder &
Savile 21349 (osc, TRT, ws); Kunghit Isl., Foster & Bigg (UBC);
Langara Isl., Beebe (v); Moresby Isl., Calder & Taylor 35771 (UBC);
Prince Royal Isl, McCabe 3358 (wTU); Saltspring Isl, Ashlee
(UBC, V); Vancouver Isl.: Comox, Anderson 483 (v); McCoy Lake,
Carter 473 (us); Lost Lake, Macoun 618 (CAN, US); Port Hardy,
Racey (UBC); Trout Lake, Davidson 6192 (usc); Alice Arm,
Szczawinski (V); Alouette R., Garman (v); Garibaldi Prov. Park,
Dow D-2A-C (v); Haney, Taylor 1002 (UBC); Hope-Prince Hwy.,
Brayshaw 49470 (uBC); Kelsey Bay, Calder & MacKay 29204 (WTU);
Kitsault, Fisher (UBC); New Westminster, McCalla 2944 (ALTA);
Prince Rupert, Mendel 78 (v); Prudhomme Lake, Taylor & Lewis
618 (UBC); Terrace, Szczawinski (UBC, V).

Kalmia microphylla var. occidentalis has been considered
specifically distinct only by Small (1914). In all floristic
works of the west coast of North America this taxon has
been considered a part of the eastern K. polifolia, although
it nas been rarely considered as a separate subspecies
(Abrams, 1951). Hultén (1948) mentions that he was not
able to separate the Alaskan specimens from those of East-
ern America. He indicates, however, that a slight racial
difference might exist, but that a large body of material
would be necessary for a definite statement.

Kalmia microphylla var. occidentalis and K. polifolia are
strikingly similar. Both have the same general habit and
size, and except for a slight difference in flower size these
structures are also the same. Under close morphological
examination, however, these two species are easily sepa-
rated. In K. polifolia, the leaf margins are usually revolute,
and well developed stalked glands are present on both sides
of the midrib. In K. microphylla var. occidentalis, by con-
trast, the leaves are not revolute, or only slightly so, and
the stalked glands are absent. The two taxa also differ in
seed size and calyx width. In K. polifolia, the seeds are long

342 Rhodora [Vol. 76

and narrow, 1.5-2.1 mm long, while the calyx varies in
width from 5-7 mm. In K. microphylla var. occidentalis,
the seeds are shorter and thicker, 1-1.4 mm long, and the
calyx varies from 7-10 mm broad.

The geographic distribution of the two taxa also differs,
and they never come into contact with one another. Kalmia
polifolia is mainly an eastern species, occurring as far west
as Saskatchewan and northeastern Alberta, while K. micro-
phylla var. occidentalis occurs along the west coast from
southern Alaska, south through British Columbia and from
Washington to Oregon. Most of the specimens of var. occi-
dentalis examined were from marshes, bogs and wet open
areas at lower elevations from coastal areas and the islands
off the coast of Alaska and British Columbia. Hultén
(1968), who did not distinguish between these two species,
shows this difference in distribution and also has an excel-
lent drawing of K. microphylla var. ? occidentalis (as K.
polifolia subsp. polifolia) .

Cytologically, Kalmia microphylla var. occidentalis (n =
12) is more closely related to K. microphylla var. micro-
phylla (n = 12) than it is to K. polifolia (n = 24). Crosses
between var. occidentalis and K. polifolia result in triploid
F, hybrids that produce less than 10% stainable pollen and
no seeds (Jaynes, 1968a). In contrast, crosses between K.
microphylla var. microphylla and var. occidentalis were
highly fertile with the F,’s being intermediate between the
two parents, producing viable seed and pollen that looked
normal and fertile (Jaynes, persona] communication).

Kalmia microphylla var. occidentalis f. alba Ebinger,
forma nov.

Forma typicae habita statura etc. similis, floribus omni-
bus albis.

This white flowering form, which is similar to that found
in some other species of Kalmia, has been collected at two
different localities. Alaska: near Wrangell, Mrs. J. C. Dart,
1922. (us #1,118,961, Holotype). British Columbia: Lulu
Island, peat bog between N. Westminster Highway and
No. 4 road, H. Rhodes, et al., 29 May 1954 (UBC).

1974] Genus Kalmia — Ebinger 343

29. Kalmia polifolia Wang., Schr. Berlin Ges. Naturf.
Freunde 8: 130. 1788. TYPE: unknown.

Kalmia polifolia Ludwig, Neu. Wilde Baumz. 25. 1783.
(nom. nud).

Kalmia glauca Loddiges ex Ludwig, Neu. Wilde Baumz.
25. 1783. (nom. nud.), Ait. Hort. Kew. 2: 64. 1789. TYPE:
Plant growing in the Kew Gardens, (K).

Chamaedaphne glauca | ( Ait.) Kuntze, Rev. Gen. Pl. 2:
388. 1891.

Kalmia glauca var. praecox Lavallée Arbor. Segrez. 159.
1877. (nom. nud.).

Kalmia glauca var. stricta Jager, Ziergeholze 279. 1865.

Kalmia glauca var. superba Makoy ex Bosse, Vollst.
Handb. Blumengart. 2: 348. 1841. (nom. nud.), Jager &
Beissner, Ziergeholze 194. 1884. Kalmia superba K. Koch,
Dendrologie 2(1): 154. 1872. (nom. nud.).

Kalmia rosmarinifolia Dum.-Cours. Bot. Cult. 2: 250.
1802. TYPE: unknown. Kalmia glauca p rosmarinifolia
(Dum.-Cours.) Pursh, Fl. Am. Sept. 296. 1814. Kalmia
polifolia var. rosmarinifolia (Dum.-Cours.) . Rehd. in Bailey,
Cycl. Amer. Hort. 2: 854. 1900.

Kalmia oleaefolia Dum.-Cours. Bot. Cult. ed. 2. 3: 322.
1811. (as synonym).

Sparingly branched shrub to 1 m tall; branchlets slightly
flattened, 2-edged, glabrous or puberulent, commonly with
decurrent ciliate lines from the petiole base; leaves oppo-
site, subsessile with a broad petiole to 2 mm long, coriaceous,
linear to oblong, the margins usually revolute, 1.5-4 cm
long, 0.3-1.3 em broad, 214 to 4 times as long as broad,
whitened beneath with small hairs, the midrib covered with
purple clavate hairs on both surfaces; inflorescence a few-
flowered, terminal corymbiform raceme, each flower on a
glabrous pedicel 1.5-3 cm long, each pedicel subtended by
a pair of bractlets in the axis of a leaf-like bract; calyx
deeply 5-lobed, colorless to light pink, 5-7 mm broad, the
lobes obtuse, margins ciliate ; corolla rose-purple, 1.2-1.8 cm
broad; anthers purple, 1-1.5 mm long; fruit subglobose,

344 Rhodora [Vol. 76

4-7 mm broad, 4-6 mm long; seeds 1.5-2.2 mm long, the
testa extending past the ends of the seed. Chromosome
number n — 24,

Distribution: Bogs, swamps, and other wet, open areas
from northeastern Alberta east to the Atlantic Ocean and

south to the Great Lakes, Pennsylvania, New York and
New Jersey.

Common name: Bog Laurel, Swamp Laurel, Pale Laurel,
Gold Withy.

REPRESENTATIVE SPECIMENS

UNITED STATES: Connecticut: HARTFORD CO.: Suffield, Harger
6524 (ARIZ), Blewitt 3490 (PH). LITCHFIELD CO.: Bingham Pond,
Weber 1489 (Isc). NEW HAVEN CO.: Middlebury, Shepard (conn).
Maine: ANDROSCOGGIN CO.: Livermore, Gordon 657 (MAINE). AROOS-
TOOK CO.: Knoles Corner, Neal 1908 (MAINE), True 6068 (PENN).
CUMBERLAND CO.: Standish, Rossbach 5599 (ACAD, MAINE). FRANKLIN
co.: Mt. Abraham, Rossbach 5389 (MAINE). HANCOCK CO.: Mt.
Desert Isl, Redfield 4266 (US). KENNEBEC CO.: Monmouth, Hyland
90 (MAINE). KNOX CO.: Rockland, Rossbach 1429 (ACAD). OXFORD
co.: Old Spec, Richards 6032 (MAINE). PENOBSCOT CO.: Orono,
Fernald 349 (GH, Mass, MIN, MSC, ND, NY, PH, US). PISCATAQUIS CO.:
Greenville, True 4701 (PAC), 4814 (PENN). SOMERSET CO.: Jackman,
True 4668 (PENN). WALDO CO.: Northport, Rossbach 2499 (ACAD),
5349 (MAINE). WASHINGTON CO.: Jonesport, Steinmetz 844 (MAINE).
YORK CO.: Sunken Pond, True 214 (GH, PENN). Massachusetts:
BERKSHIRE C0.: Wolf Swamp, Walters (NHA). ESSEX CO.: Andover,
Steele (MU). FRANKLIN CO.: Hawley, Forbes 24 (MASS). HAMPDEN
co.: Ludlow, Seymour 557 (DUKE, GH, MASS, NY). HAMPSHIRE CO.:
Belchertown, Torrey (MASS). MIDDLESEX CO.: Concord, Eaton
(MAINE). WORCESTER C0.: Worcester, Dowling 818 (NY). Michigan:
ALCONA CO.: Hall & Hagenah 447 (BLH). ALPENA co.: Hatt (BLH).
BARAGA CO.: Baraga, Richards 1783 (MICH, NY), 2956 (MAINE, MICH).
CHARLEVOIX CO.: Beaver Isl, Reis 906 (BLH, MSC). CHEBOYGAN CO.:
Mud Lake, Gleason, Jr. 665 (DUKE, GH). CHIPPEWA CO.: Sugar Isl.,
McVaugh 8745 (BLH, CAN, MICH). CLARE CO.: Lake Station, Dreis-
bach 5650 (MICH). CRAWFORD Co.: Hartwick, Bingham (BLH).
DELTA CO.: Bull Run Creek, Hagenah 6723 (BLH). EMMET CO.: Carp
Lake, Gates & Gates 10334 (RM). GOGEBIC CO.: Watersmeet, Voss
7846 (MICH), 12472 (MICH). GRAND TRAVERSE CO.: "Traverse City,
Dieterle 1299 (cM, MICH). GRATIOT C0.: Alma, Davis (MONT).
HOUGHTON CO.: Laurium, Herman 7591 (NY, US). IRON CO.: Golden
Lake, Voss 4122 (MICH), KALKASKA CO.: Carlisle, Hagenah 4602
(BLH). KEWEENAW CO.: Mott Isl, Bailey & Bailey 4131 (MICH).

1974] Genus Kalmia — Ebinger 345

LEELANAU CO.: Maple City, Thompson L-1693 (BLH). LENAWEE CO.:
Clinton, Wood & Wood (us). LUCE CO.: Spring Lake, Hagenah 1913
(BLH). MACKINAC CO.: Hagenah 4876 (BLH). MANISTEE CO.: Manis-
tee, Daniels 4267 (MONT). MARQUETTE CO.: Turin, Barlow (NY).
MIDLAND CO.: Sanford Game Preserve, Dreisbach 7941 (MICH).
MONTMORENCY CO.: Barding Swamp, Case (MICH). OAKLAND CO.:
Lakeville, Farwell 4905 (BLH). OGEMAW CO.: Clear Lake, Case, Jr.
(MICH). ONTONAGON CO.: Berglund, Beaman 1848 (MS). OSCEOLA
co.: Evart, Monaghan (MSC). OSCODA CO.: Zimmerman 416 (MICH).
OTSEGO COo.: Otsego Lake Village, Voss 10847 (MICH). ROSCOMMON
co.: Higgins Lake, Denton 1038 (MSC). ST. CLAIR CO.: Capae, Dodge
(MU, TENN). SANILAC CO.: Minden City, Voss 8341 (MICH). SCHOOL-
CRAFT CO.: Long Lake, Voss 2657 (MICH). WEXFORD CO.: Cadillae,
House 4660 (Us). Minnesota: BELTRAMI CO.: Star Isl, Butters &
Rosendahl 6584 (MIN). CARLTON CO.: Atkinson, Lakela 3550 (DUL,
MIN). CASS CO.: Lake Kilpatrick, Ballard (ORE, RM, US, WS). CLEAR-
WATER CO.: Long Lake, Morley 1106 (MIN). COOK CO.: Sea Gull
Lake, Lakela 3669 (DUL, MIN). CROW WING CO.: Bay Lake, Sheldon
2172 (MIN). ITASCA CO.: Sand Lake, Johnson 2042 (PH), 2207 (GH,
US). KANABEC C0.: Mora, Moore & Tryon, Jr. 17504 (MIN). KOOCHI-
CHING Co.: Rainy Lake, Moore & Moore 11793 (ISC, MIN). LAKE
co.: Basswood Lake, Lakela 8898 (DUL). PINE CO.: Bruno, Jerabek
66 (MIN). ST. LOUIS CO.: Palo, Nickelson 2432 (ISC, NY, ws). New
Hampshire: BELKNAP CO.: Crystal Lake, Hodgdon & Steele 9125
(NHA). CARROLL CO.: Church Pond, Gale (NHA). CHESHIRE CO.:
Keene, Gilbert (NY, PH). coos Co.: Mt. Washington, Hartmann
207 (NY); Moore 260 (Gv), 1081 (GY). GRAFTON CO.: Mt. Lafayette,
Fernald & Smiley 11839 (GH). HILLSBORO CO.: Peterborough, Batch-
elder 1891 (KY). ROCKINGHAM CO.: Derry, Batchelder (NHA).
STRAFFORD CO.: Barrington, Hodgdon & White 11935 (NHA). SULLI-
VAN Co.: Springfield, Hodgdon & Steele 9128 (NHA). New Jersey:
MORRIS CO.: Mt. Hope, Edwards & Rusling (CHRB). SUSSEX CO.:
Kittatinney Ridge, Griscom 9562 (GH). New York: CHAUTAUQUA CO.:
Petit, Thurber (GH). COLUMBIA CO.: Knickerbocker Lake, McVaugh
1013 (PENN). ERIE CO.: Protection, Miller 3764 (MSC). ESSEX CO.:
Whiteface Mt., Rudd 824 (US). FRANKLIN C0.: Upper Savanac Lake,
Guest 182 (PAC, PENN). HAMILTON CO.: Bern Bog, Henry (CM).
HERKIMER CO.: Twin Lake, VanPelt & Long (PH). MADISON CO.:
Recksport, House 6930 (US). ONEIDA CO.: White Lake, True 38
(PENN). ONONDAGA CO.: Labrador Pond, Clausen 2129 ANY, PAC,
PENN). OSWEGO CO.: Mud Pond, Wiegand 13764 (GH). ST. LAWRENCE
co.: Norfolk, Phelps 757 (GH, NY, US). SARATOGA CO.: Saratoga
Lake, House 27663 (PENN). SCHENECTADY CO.: Princetown, Svenson
(GH). SULLIVAN CO.: Neversink, Jones, Phillips & Stebbins 21 (GH).
TIOGA CO.: Owego, Mapes 4767 (GH). ULSTER CO.: Rockland, Camp
3036 (NY). WASHINGTON CO.: Salem, Stewart 63 (NY). Pennsylvania:

346 Rhodora [Vol. 76

CARBON CO.: Albrightsville, Pretz 13321 (PH). LACKAWANNA CO.:
Elmhurst, Glowenke 4820 (PENN). MONROE CO.: Tannerville, Leeds
2237 (MIN, PH). PIKE CO.: Shohola Falls, Byhouwer & Kobuski 226
(GH). SUSQUEHANNA Co.: Ararat, Glowenke 184 (PENN). WAYNE
co.: Lehigh Pond, Adams 3699 (GH). Rhode Island: PROVIDENCE CO.:
Glocester, Collins (CONN, GH, MASS, NHA, NY, PENN, PH). Vermont:
ADDISON CO.: Ripton, Eggleston 280 (GH). CALEDONIA CO.: Peacham,
Blanchard (PH, US). CHITTENDEN CO.: Colchester, Pringle (BRY).
LAMOILLE CO.: Mt. Mansfield, Moseley (MASS). ORLEANS CO.: Lake
Willoughby, Stevens (US). RUTLAND CO.: Mt. Tabor, Seymour 24407
(BRY). WINDHAM CO.: Tyson, Moldenke & Moldenke 10263 (Ny).
Wisconsin: BAYFIELD C0.: Sand Bay, Cheney 6371 (UWM). DOOR CO.:
Bailey’s Harbor Ridge, Shinners (UWM). FLORENCE CO.: Spruce
Lake, Mau (UWM). FOREST CO.: Crandon, Palmer 27776 (GY). IRON
co.: Mercer, Salamun 228 (UWM). JACKSON CO.: Ketchum, Hartley
& Morrissey 9158 (ISC). KEWAUNEE CO.: Gregor (OS). LINCOLN CO.:
Tomahawk, Engel (UWM). MARINETTE CO.: Walsh, Throne (UWM).
MONROE CO.: La Grange Twp., Hartley 7884 (MIN). ONEIDA CO.:
Three Lakes, Wadmond 2758 (MIN). RUSK CO0.: Ladysmith, Goessl
2162 (UWM). SHAWANO CO.: Camp Tivol, Keefe (UWM). TAYLOR
co.: Buch Lake Area, Piehl (UWM). viLAS co.: Eagle River,
Hollister 334 (US). WASHBURN CO.: Spooner, Cheney 6135 (UWM).

CANADA: Alberta: Lake Athabaska: Smith Landing, Preble &
Preble 99a (US); Alta, Raupp & Abbe 4529 (ALTA, CAN, MIN, PH).
Gregoire Lake, Dumais & Anderson 2693 (ALTA); Richardson Lake,
Whitehorn & Barber 185 (SASK). Labrador: Anatolak, Sewall 469
(GH, US), 471 (GH, US); Ashuanipi Lake, Harper 3877 (CAN, MIN,
us); Attikamagen Lake, Harper 3501 (CAN, ISC, MIN, PENN); Battle
Harbour, Koelz 36 (MICH, US); Cape St. Charles, Potter & Brierly
3410 (GH, NMC); Cartwright, Hitchcock 23818 (vs); Gannet Isl,
Potter & Brierly 3411 (GH); Goose Bay, Gillett & Findley 4931
(ACAD, NY); Gready Isl., Bishop 474 (GH), 475 (GH); Hopedale,
Walker 370 (PENN); Mealy Mts., Gillett & Findley 5273 (ACAD,
NY, TRT, US); Mills Harbour, Cook 88 (GH, US); Petty Harbour,
Bishop 476 (CAN, GH); Spotted Isl, Wallace 42 (CONN); Sandwich
Bay, Woodworth 329 (GH), 330 (GH); Windy Tickle, Potter & Brierly
3407 (MASS). Manitoba: Baralzon Lake, Scoggan & Baldwin 8216
(GH, MIN, WIN); Cranberry Portage, Freedmann (WIN); Churchill
Dist., Schofield 1001 (DUL, GH, MIN, WS); Duck Lake, Scoggan &
Baldwin 8245 (ALTA, CAN, GH); Gillam, Punter 97 (WIN); Goldesand
Lake, Keleher 107 (WIN); Heming Lake, Keleher 159 (WIN); Herch-
nier, Core 215 (WVA); Lake Winnipeg, Lowe (WIN); Lake Winni-
pegosis, Scoggan & Baldwin 8152 (CAN); Nelson River, Scoggan
3016 (ALTA, CAN, MIN, WIN); Nueltin Lake, Baldwin 2180 (CAN, TRT,
WIN, WS); Oxford Lake, Scoggan 5390 (CAN); Reindeer Lake, Bald-

1974] Genus Kalmia — Ebinger 347

win 2028 (CAN, TRT, WIN); Seal River, Ritchie 1794 (CAN, MIN);
Third Cranberry Lake, Freedman (WIN); Wabowden, Gillett 2790
(us); Whiteshell Forest Reserve, Scoggan 8781 (ALTA, WIN); York
Factory, Punter 134 (WIN). New Brunswick: ALBERT CO.: Grindstone
Isl, Christie 1576 (NBM). CHARLOTTE CO.: Campobello Isl., Malte
955/29 (CAN). GLOUCESTER CO.: Bathurst, Malte 13 (CAN). KENT
co.: Little Forks Brook, Squires & Squires (NBM). MADAWASKA CO.:
St. Leonard, Maite 605 (CAN). QUEENS CO.: Castaway, Gorham &
Squires (NBM). SAINT JOHN CO.: Portland, Matthew (NBM). WEST-
MORELAND CO.: Sackville, Fowler (QUK). YORK CO.: Fredericton,
Dore & Gorham 4569 (ACAD). Newfoundland: BONAVISTA SOUTH DIST.:
Lethbridge, Rouleau 8363 (NFLD). BURGEO-LA POILE DIST.: Grandy
Brook, Fernald, Long & Fogg 359 (GH, PENN, PH). FERRYLAND DIST.:
Witless Bay, Rouleau 5391 (CAN, NFLD, US). FOGO DIST.: Fogo Isl.,
Sornborger (GH). FORTUNE BAY-HERMITAGE DIST.: Hermitage Bay,
Palmer 1353 (GH, US). GRAND FALLS DIST.: Bishop Falls, Fernald &
Wiegand 6023 (CAN, GH, K, NY, PH). HUMBER DIST.: Eagle's Nest
Brook, Rouleau 3069 (NFLD). ST. BARBE DIST.: Trout R., Rouleau
3515 (NFLD). ST. MARY'S DIST.: Colinet, Rouleau 4175 (NFLD). ST.
JOHN’S DIST.: Doctor Hill, Fernald, Long & Fogg 1916 (GH, PENN,
PH, US). TRINITY NORTH DIST.: George’s Brook, Rouleau 6078 (NFLD).
WHITE BAY DIST.: Twin Lakes, Rouleau 6525 (NFLD, US). Northwest
Territories: Baralzo Lake, Scoggan & Baldwin 8335 (CAN), 8379
(CAN); Gagnon Lake, Scotter 3162 (CAN); Ingalls Lake, S.C.Z.
(CAFB). Nova Scotia: ANNAPOLIS CO.: Karsdale, Johnson (ACAD).
CAPE BRETON CO.: Scatari Isl, Ershine 51-1167 (QUK). COLCHESTER
co.: Folleigh Lake, Perry (ACAD). CUMBERLAND CO.: Sand R.,
Schofield & Webster 5592 (ACAD). DIGBY CO.: Central Grove, Fernald
& Long 24285 (GH). GUYSBOROUGH CO.: Tor Bay, Eshbaugh 522
(IND). HALIFAX CO.: Fernald, et al. 24284 (GH, PH). HANTS CO.:
Mt. Uniacke, Jack 3230 (GH), 3231 (GH). INVERNESS CO.: Mackenzie
Mt., Rossbach 4235 (ACAD). KINGS CO.: Caribou Bog, Erskine
(ACAD). LUNENBURG CO.: Deep Cove, Smith, et al. 19604 (ACAD,
QUK). PICTOU co.: New Glasgow, Dore & Gorham 45-389 (ACAD).
QUEENS C0.: Eagle Head, Donly 1009 (ACAD). RICHMOND CO.: Grand
R., Smith, et. al. 10110 (ACAD, TRT). SHELBURNE CO.: Barrington,
Fernald, Long & Linder 22154 (CAN, GH). VICTORIA CO.: Cape Breton
Nat’l. Park, Armstrong 452 (QUK). YARMOUTH CO.: Yarmouth,
Pease & Long 22152 (GH, PENN). Ontario: ALGOMA DIST.: Michi-
picoten Harbour, Hosie, Harrison & Hughes 1223 (CAN, TRT).
CARLETON CO.: Mer Blue, Porsild 7743 (CAN). COCHRANE DIST.:
Fraserdale, Kirk 75 (TRT): Timmins, Baldwin 4761 (CAN, TRT).
DURHAM C0.: Newtonville, Krug & Purchase 473 (TRT). FRONTENAC
co.: Kingston, Beschel 10128 (QUK). GREY CO.: Euphrasia Twp.,
Jackson (TRT). HALIBURTON CO.: Dorset, Walshe 12 (CAN, TRT).
HURON C0.: Wingham, Anderson 724 (TRT). KENORA DIST.: Rich-

348 Rhodora [Vol. 76

mond Gulf, Abbe & Abbe 3383 (CAN, MIN); Las Seul, Baldwin 8739
(CAN, MIN, QUK, TRT); Big Trout Lake, Moir 13 (MINN), 21 (CAN,
MIN), 1744 (MIN); Swan Lake, Moir 156 (MIN); Fort Severn,
Moir 1337 (CAN, MIN); Black Duck R., Moir 1831 (MIN), 2037 (CAN,
MIN, TRT) ; Windigo R., Cringan P-26 (ACAD, TRT). MANITOULIN DIST.:
Great Duck Isl, Grassel 6028 (NY). MIDDLESEX CO.: London, Baker
1638 (OAC). MUSKOKA DIST.: Huntsville, Soper & Shields 4927
(TRT), 4967 (TRT); Margaret Lake, Chow 6415 (TRT). NIPISSING
DIST.: Algonquin Park, Watson 3967 (TRT). PARRY SOUND DIST.:
Axe Lake, Shields & Shields 1640 (TRT). PEEL CO.: Heart Lake,
Catling & Sparling (TRT). RUSSELL CO.: Bourget, Heimburger 2612
(TRT). SUDBURY DIST.: Skead, Winterhalder 66114 (CAN). THUNDER
BAY DIST.: Bear Trap Lake, Voss 10361 (MICH, TRT). TIMISKAMING
DIST.: Hilliard Twp., Baldwin & Breitung 2564 (CAN, SASK, TRT).
WATERLOO CO.: Baden, Montgomery 997 (au, OAC). WELLAND CO.:
Humberstone, Soper & Dale 3776 (GH, MIN, TRT, US). WELLINGTON
Co.: Puslinch, Stroud (TRT). YORK co.: Vandorf, Robb (TRT).
Prince Edward Island: KING Co.: Bothwell, Fernald & St. John 11148
(CAN, GH, US, WS). PRINCE CO.: North Point, Erskine & Dore 1080
(ACAD, NY). QUEENS CO.: Brackley Point Rd., Fernald, Long & St.
John 7901 (CAN, GH, PH, US, WS). Quebec: ABITIBI CO.: Watson
Lake Area, Bentley 58145 (TRT), 5837 (CAN, TRT). BERTHIER CO.:
Lanoraie, Scoggan 1538 (CAN). BONAVENTURE CO.: Nouvelle, Collins
& Fernald (GY). CHAMBLY CO.: Saint-Bruno, M.-Victoria & R.-
Germain 43884 (cM, GH, ND, PH). CHICOUTIMI CO.: Lac Merlac,
Brisson 63124 (CAN). GASPE WEST CO.: Mt. Albert, Scoggan 1086
(CAN). GATINEAU CO.: Eardley, Cody & van Rens 11167 (DUKE, GH,
MSC, NY, TENN). HULL CO.: King Mt., Zavitz 1107 (TRT). HUNT-
INGTON CO.: Covey Hill, M.-Victoria & R.-Germain 43300 (GH).
LABELLE C0.: Nominingue, Roy 1167 (wvA), 3650 (ACAD, CONN, NY,
RM, US). LAKE ST. JOHN WEST CO.: Peribonka, M.-Victorin 15849
(GH). LEVIS CO.: St. Nicolas, Cayouette & Cayouette 54-89 (TRT).
MAGDALEN ISLANDS CO.: Grindstone Isl, Fernald, et. al. 7900 (CAN,
PH, WS). MATANE CO.: Montagne Lake, Boivin 787 (PENN). MATA-
PEDIA CO.: Sayabee, Scoggan 13600 (CAN). MEGANTIC CO.: Black
Lake, Fernald & Jackson 12137 (GH). MONTCALM CO.: Lac Tapani,
Pennell 16686 (PH). MONTMAGNY CO.: Lake Monroe, R.-Germain &
Courtemanche 454 (US). PAPINEAU CO.: Notre Dame du Laus,
C.-Joseph 8508 (ACAD). PONTIAC COo.: Danford Lake, Dore 19994
(TRT). QUEBEC CO0.: Montmorency Fall, Macoun 68560 (am).
RIMOUSKI CO.: Father Point, Bartram & Long 581 (PH). SAGUENAY
co.: Fort Chimo, Porsild 21873 (CAN); Knob Lake, Harper 3350
(HNH, QUK); Cairn Isl., Abbe, et. al. 3067 (RM): Seven Isl., Robinson
868 (CAN, GH, NY); Mutton Bay, Lewis 582 (CAN). ST. JEAN CO.:
Villeneuve, M.-Victorin, et. al. 43047 (MU). SHERBROOKE CO.: Orford,
Forest 16058 (CAN, CM, QUK, SASK). TERREBONNE CO.: St. Janvier,

1974] Genus Kalmia — Ebinger 349

Scoggan 314 (CAN). TWO MOUNTAINS CO.: Oka, M.-Victorin 8228
(NY, US). WOLFE C0.: Lac Aylmer, Hamel 13217 (CAN). Saskatche-
wan: Athabaska Lake, Lang 188 (Us); Raupp 6175 (CAN, GH, NY),
6657 (GH), 6731 (GH, NY); Bittern Cr., Wheaton (CAFB); Candle
Lake, Boivin & Breitung 6157 (NY); Cree Lake, Maini 21 (RM,
SASK), 235 (SASK), 236 (CAN, RM, SASK); Crooked R., Breitung 70
(SASK); Cumberland House, Argus 4083 (SASK); Hatchet Lake,
Maini 91 (OAC, RM, SASK); Nipawin Prov. Park, Argus 4122 (SASK);
Orley, Breitung 268 (ALTA, CAN, NY, US); Prince Albert Nat’l. Park,
Fraser (PAC, SASK); Stoney Rapids, Maini 205 (RM, SASK), 275
(SASK), 462 (RM, SASK); White Fox, Fraser (SASK) ; Windrum Lake,
Bryenton 166 (CAN).

The name Kalmia polifolia was first used by Ludwig
(1783), who gave it the common name Canada Laurel or
Poley Leaved Laurel, but did not describe it. This name
first appears with a valid description a few years later
(Wangenheim, 1788). He gives its distribution as extend-
ing northward from 45 degrees latitude in America, Nova
Scotia and the colder parts of Canada. His description,
illustration, and the distribution leave no doubt as to the
taxon referred to. Recently, Boivin (1967) suggested that
the name be spelled poliifolia to conform to article 73 of the
International Code of Botanical Nomenclature (Lanjouw,
1966). The name K. glauca was first used as a synonym
of K. polifolia by Ludwig (1783) and later described by
Aiton (1789). The type was based on a plant from New-
foundland growing at Kew. A specimen in the Kew Her-
barium has written on the label “Kalmia glauca Hort.
Kews. sp. nov. 1784” and probably represents the type of
this name. Kalmia rosmarinifolia was used by Dumont de
Courset (1802) to distinguish individuals that have narrow
leaves with strongly revolute margins. Later, he listed the
name as a synonym of K. glauca (Dumont de Courset,
1811).

Kalmia polifolia is found in bogs, swamps, and other wet
places, commonly forming a border around ponds and
lakes. In bogs, its roots usually form dense mats that
extend out over the water. It is also found at higher ele-
vations in the mountains of northeastern United States and
Canada. This species is the most widely distributed member

350 Rhodora [Vol. 76

of the genus, extending from northeastern Alberta, across
Canada to the east coast and south to New Jersey. Its
entire range, however, is hard to determine with certainty.
There are two reports of its occurrence as far north as the
Arctic Circle in the Northwest Territories (Gussow, 1933,
and Thieret, 1963). Both of these (Maguse Lake, Gussow
33; Contwoyto Lake, Thieret 9415) are specimens of K.
microphylla var. microphylla. The only specimens of K.
polifolia seen from the Northwest Territories are from the
extreme southern part (Scotter, 1966). In the Great Lakes
region this taxon extends as far south as northern Illinois
(Fuller, 1944), and along the east coast to New Jersey
(Griscom, 1931). This species has also become naturalized
in Europe since Scott (1962) reported that it has been col-
lected from Flanders Moss, Perthshire, Scotland.

Besides leaf size and shape, the character most commonly
used by previous authors to separate Kalmia polifolia from
K. microphylla is revolute leaf margins. In K. microphylla,
the leaf margins are not revolute, or only slightly so, while
in K. polifolia the margins are usually strongly revolute.
This characteristic is somewhat variable. In about 80% of
the specimens examined, the leaf margins were strongly
revolute, resulting in a narrow, linear leaf. In the remain-
ing specimens, some lacked revolute margins entirely, while
in others the younger leaves were revolute while older
leaves were not. The variability of this characteristic is
responsible, in part, for the confused taxonomic status of
K. microphylla.

The major distinguishing vegetative character that sep-
arates Kalmia polifolia from the related species, K. micro-
phylla, is the presence of purple clavate hairs on the leaf
midrib. Boivin (1968) appears to be the first to have made
use of this character when he separated variety microphylla
from variety polifolia. He mentions, however, that west of
Alberta the distinction of the two varieties becomes mean-
ingless, as most of the British Columbia material is inter-
mediate, the leaves being long and narrow like variety poli-
folia, but glandless on the midrib. It appears that he was

1974] Genus Kalmia — Ebinger 851

confusing K. polifolia with the western K. microphylla var.
occidentalis, which is similar vegetatively, except for the
glandless midribs. Clavate glands were found on all speci-
mens of K. polifolia examined. These glands vary in size,
shape and color, and are more plentiful on some specimens
than on others. The glands average about 0.3 mm long, are
colorless near the base and become deep purple to black
toward the swollen apex. In some specimens, however, they
are nearly colorless throughout, and in others barely ex-
panded above. Also, their distribution varies, being more
common on some leaves than on others. On older leaves, it
is not uncommon to find that many of the glands have
broken off. The glands on the upper side of the midrib are
commonly shorter, darker colored and not as obvious as
those on the under side of the leaf. For determining the
presence of these glands it is best to look at the developing
leaves.

Morphologically and genetically Kalmia polifolia (n =
24) is most closely related to K. microphylla (n = 12). The
F, of a cross between these two species is a highly viable
triploid which has a pollen stainability of 0-15% (Jaynes,
1968a). These similarities suggest that these two taxa have
probably become specifically distinct as a result of the last
ice age and the resulting development of a tetraploid taxon
from a wide ranging ancestral diploid species. Jaynes
(1969) suggested that these taxa should be considered sep-
arate species based on his cytological and genetic studies.
Kalmia polifolia can also be crossed with other members of
the genus, but few viable offspring are produced.

Kalmia polifolia forma leucantha Schofield & Smith,
Canad. Field Nat. 67: 94. 1953. TYPE: Newfoundland:
near Hodgewater Line, Trinity South, E. E. Smith and
A. C. Smith 1055 (ACAD). i

This white flowering form, which is similar to that found
in some of the other species of Kalmia, is known from one
collection. This single specimen was found in a bog where
the typical form was very common.

352 Rhodora [Vol. 76

3. Kalmia latifolia Linnaeus, Sp. Pl. 1: 391. 1753.
TYPE: “Habitat in Marilandia, Virginia, Pennsylvania"
(LINN 560.1, lectotype, specimen at the top right).

Chamaedaphne latifolia (L.) Kuntze, Rev. Gen. Pl. 2:
388. 1891.

Kalmia latifolia var. acuminata Raf. Medical Fl. 2: 17.
1830. (nom. nud.).

Kalmia latifolia var. alba Raf. Medical Fl. 2: 17. 1830.
Kalmia latifolia f. alba (Mouillefert) Rehd. Rhodora 12: 2.
1910.

Kalmia latifolia var. arborea Raf. Medical Fl. 2: 17. 1830
(nom. nud.).

Kalmia latifolia var. pavarti André, Revue Hort. 60: 541.
1888.

Kalmia latifolia f. rubra K. Koch, Dendrologie 2(1) : 152.
1872. Kalmia latifolia var. rubra (K. Koch) Rehd. in
Bailey, Cycl. Amer. Hort. 2: 854. 1900.

Kalmia latifolia var. ternata Raf. Medical Fl. 2: 17. 1830.

Kalmia latifolia var. laevipes Fern. Rhodora 42: 53. 1940.
TYPE: Virginia: NORFOLK CO.: dry woods of a *hammock",
Great Dismal Swamp, west of Yadkin, Fernald & Long
11101 (GH).

Kalmia lucida K. Koch, Dendrologie 2(1): 152. 1872.
(as synonym).

Kalmia nitida Forbes, Hort, Woburn. 93. 1833. TYPE:
unknown, probably a living plant.

Much branched shrub to 4 m tall (rarely a small tree) ;
branchlets terete, reddish brown, becoming gray, stipitate-
glandular to glabrous; leaves alternate (appearing nearly
whorled on slow growing twigs), flat, coriaceous, elliptic to
elliptic-lanceolate, 4-12 cm long, 1.5-4 (rarely 5) em wide,
dark green and glabrous above, light green to reddish brown
and with short glandular hairs beneath, apex acute, base
narrowly cuneate, midrib puberulent above; petiole 1-3 em
long, glabrous to puberulent, sometimes glandular-stipitate;
inflorescence a terminal, compound corymb; pedicels 2-4 cm
long, puberulent to stipitate-glandular, often floccose, viscid

1974] Genus Kalmia — Ebinger S00

(rarely glabrous), subtended by 2 bracteoles in the axil of
a puberulent, lanceolate bract, 1-5 mm long; major inflor-
escence branches puberulent, usually stipitate-glandular, to
4 cm long; calyx glabrous to stipitate glandular, green to
reddish, 5-8 mm broad, the lobes usually oblong, acute;
corolla usually light pink (white to deep red) with purple
spots around each anther pocket, usually lightly stipitate-
glandular outside, puberulent inside, 1.5-3 cm broad; style
1-1.8 cm long; anthers dark purple to brown, 1.2-1.7 mm
long; filaments puberulent; capsule depressed globose, stipi-
tate-glandular, 4-7 mm broad, 3-5 mm long; seeds .7-1 mm
long, the testa loose and extending slightly past the ends of
the seed.

Distribution: Mostly rocky or sandy hardwood forests in
eastern United States from southern Maine, west through
southern New York to central Ohio, south to southern Mis-
sissippi, Alabama, and Georgia and northwestern Florida.

Common Name: Mountain Laurel, Broad-leaved Laurel,
Calico-bush, Spoonwood, Ivy, Mt. Ivy, Big-leaved Ivy,
Laurel-leaves, Calmoun.

REPRESENTATIVE SPECIMENS

Alabama: CALHOUN CO.: Pendergrass (TENN). CHEROKEE CO.:
Centre, Clonts 95 (1sC). CHOCTAW CO.: Silas, McDaniel 4218 (VDB).
CLARKE CO.: Tallahatta Creek, Kral 19749 (VDB). CLEBURNE CO.:
Kral 31843 (VDB). CONECUH CO.: Castleberry, Howell 760 (us).
CULLMAN CO.: Cullman, Kral 28911 (VDB). DALE CO.: Newton, God-
frey 55477 (TENN). DE KALB CO.: Mentone, Demaree 50521 (TENN).
ESCAMBIA CO.: Little Escambia River, Ahles 7229 (MISSA). FRANK-
LIN CO.: Russellville, Kral 26321 (VDB). HENRY co.: Headland,
Wiegand & Manning 2370 (CU). JACKSON co.: Flat Rock, Kral
34948 (VDB). LEE CO.: Chewacla, Barnes 119 (VDB). MARION CO.:
Hackelberg, Kral 20024 (VDB). MARSHALL CO.: Albertville, Harbison
15887 (TENN). MOBILE CO.: Mt. Vernon, Burnham 84 (VDB). PIKE
co.: Troy, McDaniel 7830 (vpB). TUSCALOOSA Co.: Lake Nichols,
Williams 104 (VDB). WASHINGTON CO.: Bates Creek, Segars 191
(WVA). WINSTON cCOo.: Haleyville, Kral 29476 (vpB). Connecticut:
FAIRFIELD CO.: Wilton, Eames 10426 (CONN). HARTFORD CO.: Wind-
sor, Ahles 65351 (MASS). LITCHFIELD CO.: Preston, Lumsden (MU).
MIDDLESEX CO.: Millers Pond, North 51 (MICH, US). NEW HAVEN CO.:
Milford, Fames 2813 (CONN). NEW LONDON CO.: Lantern Hill, Glea-

354 Rhodora [Vol. 76

son (DUKE). TOLLAND CO.: Andover, Travis 2408 (CONN). WINDHAM
co.: Crystal Lake, Peterson (cU). Delaware: NEW CASTLE CO.:
Arden, Travis 1086 (PENN). SUSSEX CO.: Gumboro, Beaven 241
(DUKE). Florida: ESCAMBIA CO.: McDavid, Goodale (MASS). GADSDEN
co.: Sycamore, Godfrey 61925 (Msc). HOLMES CO.: Darlington,
Godfrey 55498 (TENN). LIBERTY CO.: Greensboro, Karl 1916 (TENN).
OKALOOSA CO.: Laurel Hill, Godfrey 61284 (US). WALTON CO.: De
Funiak Springs, Perkins & Hall 2510 (cv). Georgia: CLARK CO.:
Middle Oconee River, Duncan & Duncan 4358 (ID, IND, ISC, PENN,
RENO, TENN, US). CLAY CO.: Cemochechahee Creek, Thorne 3453
(CU). DADE CO.: Lookout Mt., MeVaugh 9023 (MICH), EARLY CO.:
Sawhatchee Creek, Duncan & Thorne 1778 (CU, TENN). EFFINGHAM
co.: Savannah River, Harper 1814 (US). GWINNETT CO.: McGuires
Mill, Small (US). MORGAN CO.: Hard Labor Creek St. Park, Jones
15456 (VDB). PICKENS CO.: Jasper, Cronquist 4483 (NO, US). RABUN
cc.: Darling Spring, J.H.M. E1525 (WVA). RANDOLPH CO.: Cuthbert,
‘Harbison 13729 (TENN). SCREVEN CO.: Newington, Cronquist 5023
(MICH, US, WS). SUMTER CO.: Americus, Harper 523 (US). UNION
co.: Vogel St. Park, McVaugh & Pyron 841 (ORE). UPSON co.: Flint
River, Browne & Browne, Jr. 6587 (KY). WALKER CO.: Pigeon Mt.,
Wilson 176 (US). WALTON Co.: Athens, Cronquist 4468 (MICH, NO,
US). WHITE CO.: Cleveland, Lovett (DUKE). WHITFIELD CO.: Dalton,
Cronquist 5045 (MICH, US, WS). Indiana: CLARK CO.: Borden, Deam
24495 (IND, MIN). CRAWFORD CO.: Riddle, Deam 16497 (IND). PERRY
co.: Derby, Ramsey 10876 (IND). Kentucky: BELL co.: Pike Mt.
St. Park, Collins 180 (KY). BREATHITT CO.: Robinson Forest, Smith
1726 (KY). CARTER CO.: Cascade Cavern, Gilbert & McCoy 93513
(us). CASEY CO.: Liberty, Wharton 4366 (KY). EDMONSON CO.:
Mammoth Cave Nat’l. Park, Pittillo 2452 (Kv). ESTILL CO.: South
Irvine, Wharton 2298b (Kv, MICH). FLEMING CO.: Plummer's Mill,
Wharton 3873 (KY). FLOYD CO.: Hueysville, Centers & Blair TT
(KY). GRAYSON CO.: Clifty Creek, Browne & Browne, Jr. 6494 (KY).
HARLAN CO.: Poor Fork, Kearney 182 (ISC, MSC, OS, US). JESSAMINE
co.: Camp Daniel Boone, Webb 104 (Ky). KNOTT CO.: Littcarr,
Smith 59 (KY). LETCHER CO.: Browne & Browne, Jr. 5400 (KY).
LEWIS CO.: Vanceburg, Wharton 5012 (KY, MICH). LINCOLN CO.:
Ottenheim, Wharton 4866 (KY, MICH). MCCREARY CO.: Cumberland
Falls, Rogers 55 (DUKE, IND, MIN, PENN, TENN, US, WVA). MADISON
co.: Berea, Nelson (MICH, MIN, MU). MENIFEE CO.: Frenchburg,
MelInteer 2302 (KY). MORGAN CO.: Ezel, Oldfield 57 (KY). POWELL
co.: Stamping Ground, Singer (BRY). PULASKI CO.: Mt. Victory,
Warden 129 (Kv). TODD CO.: Clifty, McCoy (WVA). TRIMBLE CO.:
Bedford, Born (KY). WOLFE CO.: Sky Bridge, McInteer 1055 (KY).
Louisiana: WASHINGTON PARISH: Bogalusa, Cocks (NO); Varnodo,
Thieret 23481 (DUKE).

Maine: CUMBERLAND CO.: Richville, Hyland 444 (MAINE). OXFORD

1974] Genus Kalmia — Ebinger 355

co.: Fryeburg, Hyland 446 (MAINE). PENOBSCOT CO.: Carmel,
Harvey 622 (US). SAGADAHOC CO.: Phippsburg, Gordon 592 (MAINE).
WASHINGTON CO.: Cherryfield, Hyland 760a (MAINE). YORK CO.:
Kennebunk, Hyland 252 (MAINE). Maryland: ALLEGANY CO.: Cum-
berland, Richards 128 (MAINE). ANNE ARUNDEL CO.: Camp Whip-
poorwil, Grimes (WVA). BALTIMORE C0.: Abingdon, Nieuwland
(ND). CAROLINE CO.: Newton, Quidas 37 (MAINE, PENN). CECIL CO.:
Elkton, Randolph & Randolph 124 (CU). FREDERICK Co.: Hunting
Creek, Earle 1872 (PENN). GARRETT CO.: Grantsville, Stone 14111
(PENN). KENT CO.: Chestertown, Vanatta (CHRB). PRINCE GEORGE
co.: Suitland, Meyer 9178 (ISC, WVA). QUEEN ANNES CO.: Chester
River, Thatcher 5 (MIN). ST. MARY'S CO.: Patuxent River, Grether
8276 (MICH). TALBOT CO.: Easton, Earle 810 (PENN). WASHINGTON
co.: Boonsboro, Tidestrom 724a (US). WICOMICO CO.: Sharptown,
Tidestrom 12228 (MICH). WORCESTER CO.: Ferry Creek, Redmond
225 (MICH). Massachusetts: BERKSHIRE CO.: Williamstown, Ames
(MICH). BRISTOL CO.: New Bedford, Bartlett 319 (IND, MICH).
ESSEX CO.: Newburyport, Mock (NHA). FRANKLIN CO.: Mt. Toby,
Elwell (MASS). HAMPDEN CO.: Granville, Seymour 197 (MASS).
HAMPSHIRE CO.: Amherst, Elwell (MASS). MIDDLESEX CO.: Concord,
Mann (HNH). NORFOLK CO.: Dedham, Greenman 2371 (MIN).
PLYMOUTH CO.: Plymouth, Morong 1184 (US). WORCESTER CO.:
Brookfield, Markert (MASS). Mississippi: CLARKE CO.: Burstone
Cuesta, Jones 12050 (VDB). FORREST CO.: McCallum, Ray, Jr. 6045
(MISSA). GEORGE CO.: Cedar Creek, Demaree 35222 (VDB). HARRISON
co.: Gulfport, Bomhard 33 (NO). HINDS CO.: Natchez Trace Park-
way, Price (MISSA). JACKSON CO.: Van Cleave, Demaree 82454
(DUKE). JASPER CO.: Rose Hill, Reynolds 96 (MISSA). LAUDERDALE
co.: Meridian, Hayes 20 (MISSA). LINCOLN CO.: Boque Chitto, Ray,
Jr. 7830a (MISSA). PEARL RIVER Co.: Picayune, Ray, Jr. 6519
(MISSA). STONE CO.: Perkinson, Ray, Jr. 7713 (MISSA). TISHOMINGO
co.: Fisherman Ford, Ray, Jr. 7507 (MISSA, VDB). WAYNE CO.: Clara,
Graham 118 (MISSA). New Hampshire: BELKNAP CO.: Center Harbor,
Hodgdon, et. al. 9507 (NHA). CARROLL CO.: Albany, Pease 25358
(MAINE). CHESHIRE CO.: Fitzwilliam, Hodgdon 7069 (HNA). GRAFTON
co.: Hanover, Drew (HNH). HILLSBORO Co.: Manchester, Hodgdon
& Kent 7070 (NHA). MERRIMACK CO.: Hooksett, Hall (NHA). ROCK-
INGHAM CO.: Windham Center, Harris 430 (NHA). STRAFFORD CO.:
Farmington, Lord (NHA). New Jersey: ATLANTIC CO.: Alsion, Crow
126 (MSC). BERGEN CO.: Palisades, Gershoy (CU). BURLINGTON CO.:
Pemberton, Palmer 89752 (MICH, MIN). CAMDEN CO.: Gibbsboro,
Hynes (PENN). CAPE MAY CO.: Dennisville, Fender 5049 (PENN).
CUMBERLAND CO.: Gouldtown, Long 575 (ARIZ, CAN, CU, IND, ISC, KY,
NHA, NO, TENN, UARK, WVA). ESSEX CO.: Millburn, Rusby 1710
(MICH). GLOUCESTER CO.: Mantua, Adams 258 (PENN). HUNTERDON
co.: Fairmount, Good (CHRB). MIDDLESEX CO.: Spotswood, Laugh-

356 Rhodora [Vol. 76

ridge & Chrysler (CHRB). MONMOUTH CO.: Clarksburg, Fogg, Jr.
14129 (PENN). MORRIS CO.: Longwood, Davidson 764 (CHRB). OCEAN
C0.: South Lakewood, MacKenzie 3116 (US). PASSAIC co.: Utter-
town Bog, Montgomery (CHRB). SALEM CO.: Auburn, Fogg, Jr. 6606
(CHRB, PENN). SOMERSET CO.: Willowood estate, Blackburn (CHRB).
SUSSEX CO.: Kittatinny Ridge, Davidson 1509 (CHRB). WARREN CO.:
Sand Pond, Large & Clausen 1280 (CU). New York: BRONX CO.:
Bronxdale, Burnham 428 (CU). CATTARAUGUS CO.: Rock City Park,
Muenscher 15841 (CU). CHEMUNG CO.: Susquehana River, Lucy 6616
(MU). COLUMBIA CO.: Ghent, McVaugh 881 (PENN). DELAWARE CO.:
Deposit, Muenscher 16244 (cu). DUTCHESS CO.: Clove, Standley &
Bollman 12138 (US). GREENE CO.: Winter Clove Mt., Gershoy (CU).
NASSAU CO.: Woodbury, Stewart (CU). ONEIDA CO.: New London,
House 27859 (CU). RICHMOND CoO.:. Haverstraw, Fosberg & Fosberg
14897 (PENN). ROCKLAND CO.: Ramapo, Gershoy (CU). SCHUYLER CO.:
Cayuta Lake, Muenscher & Brown 21703 (cU). STEUBEN CO.: Painted
Post, Muenscher 15842 (CU). SUFFOLK CO.: Southampton, St. John
2852 (CU). SULLIVAN CO0.: Camp Ranachqua, Weber 272 (isc).
TIOGA CO.: Spencer, Hames & McDaniels 4765 (CU). TOMPKINS
co.: Enfield, Palmer 928 (IND). ULSTER C0.: Lake Minnewaska,
Gillis 3090 (MSC). WESTCHESTER CO.: Ossining, Gershoy (CU, MU).

North Carolina: ALEXANDER CO.: Keever 21 (DUKE). BUNCOMBE
co.: Asheville, McCarthy (US). BURKE CO.: Shortoff Mt., Wilbur
7004 (VDB). CALDWELL CO.: Granite Falls, Randolph & Randolph
1085 (CU). CHATHAM CO.: Costello 14 (DUKE). CLAY CO.: Perry
Gap Road, Wilbur 1275 (MIN, DUKE). CUMBERLAND CO.: Rockfish
Creek, Correll & Rankin 9053 (DUKE). DURHAM CO.: Catsburg,
Wilbur 12230 (DUKE). EDGECOMBE CO.: Tar River, Fox & Whitford
1746 (DUKE). FORSYTH CO.: Schallert 1092 (DUKE). GASTON CO.:
Kings Mt., Oosting 1646 (DUKE). GRAHAM CO.: Thunderhind Mt.,
Beardslee & Kofoid (MU). GRANVILLE CO.: Tar River, Correll 516
(DUKE, MASS). HARNETT CO.: Bunnlevel, Laing 1104 (MIN). HAY-
woop co.: Waynesville, Standley 5401 (US). HENDERSON CO.: Hen-
dersonville, Cottam 11140 (UT). JACKSON CO.: Wesner Bald, Ramseur
171 (MAINE). JOHNSTON CO.: Neuse River, Fox & Whitford 1293
(DUKE, WS). LEE CO.: Deep River, Beard 405 (KY). MCDOWELL CO.:
Rogers 8358 (DUKE, US). MACON CO.: Mt. Satula, Harbison 1848
(TENN). MADISON C0.: Hot Springs, Wehmeyer 533 (MICH). MIT-
CHELL CO.: Roan Mt., Cannon 143 (CU). MOORE CO.: Carthage,
Harriot (NO). ORANGE Co.: Hillsboro, Correll 325 (DUKE). POLK
co.: Columbus, Townsend 236 (CU). ROCKINGHAM. CO.: Smith River,
Correll & McDowell 10695 (DUKE). SCOTLAND CO.: Laurel Hill,
Wiegand & Manning 2369 (cu). STANLY CO.: Badin, Fox, Godfrey
& Boyce 1427 (DUKE, TENN, US, WS). STOKES CO.: Sauratown Mt.,
Radford 34665 (1sc). SURRY co.: Pilot Mt., Williams 296 (DUKE).
SWAIN CO.: Andrews Bald, Bruhn 4 (TENN). TRANSYLVANIA CO.:

1974] Genus Kalmia — Ebinger 357

Rosman, Morley 891 (18C). UNION CO.: Rocky River, Ahles & Haesloop
31521 (UARK). WAKE CO0.: Raleigh, Godfrey 3765 (US). WATAUGA
co.: Boone, Ahles & Duke 43728 (PAC). WAYNE CO.: Mt. Olive,
Rose 36 (MASS). WILKES CO.: Wyatt 832 (ARIZ, BRY, NHA, VDB).
Ohio: ADAMS CO.: Mineral Springs, Herrick (0S). ATHENS CO.:
Athens, Kellerman (0S). COLUMBIANA CO.: Sanders (OS). COSHOCTON
co.: New Castle Twp., Selby (os). CUYAHOGA Co.: Bayville, Biebel
(OS). FAIRFIELD C0.: Sugar Grove, Potzger 2969 (ND). GALLIA CO.:
Jones (0S). GUERNSEY CO.: Liberty Twp., Thomas (0S). HOCKING
co.: Clear Creek, Stuckey 2912 (0s). HOLMES CO.: Killbuck Twp.,
Herrick (08S). JACKSON CO.: Liberty Twp., Bartley & Pontius (os).
JEFFERSON C0.: Amsterdam, Cusick (08). KNOX co.: Pipesville,
Moldenke 12186 (0S). LAWRENCE CO.: Hanging Rock, Werner (os).
LICKING C0.: Blackland Gorge, Stuckey 6603 (os). MEIGS CO.:
Columbia Twp., Jones (0S). MONROE CO.: Adams Twp., Thomas
(OS). MORGAN CO.: Bur Oak St. Park, ‘Herrich (0S). MUSKINGUM
co.: Laurel Hill, Reed (0s). PERRY CO.: Jonathan Creek, Hicks &
Dambach (os). PIKE CO.: Chimney Rocks, Crowl (0S). ROSS CO.:
Tar Hollow, Crowl (os). SCIOTA CO.: Cooper Head Fire Tower,
Demaree 10664 (os). TUSCARAWAS CO.: Wayne Twp., Brown (os).
VINTON C0.: Lake Hope St. Park, Stuckey 6407 (0S). WASHINGTON
co.: Vincent, Walpole 1129 (BLH). Pennsylvania: ADAMS CO: Birch
River, Tanger 4434 (PENN, WS). ALLEGHENY C0.: Tom's Run Rd.,
Henry & Buker (PENN). ARMSTRONG CO.: Sherrett, Henry (PENN).
BEDFORD CO.: Tatesville, Berkheimer 982 (PENN). BERKS CO.: Bech-
telsville, Fender 1022 (PENN). BLAIR CO.: Blue Knob, Westerfeld
16177 (PAC). BRADFORD Co.: Canton, Westerfeld 907 (PAC, PENN).
BUCKS CO.: Janney, Long 76067 (PENN). BUTLER CO.: Clintonville,
Brubaker 2537 (PENN). CAMBRIA CO.: Johnstown, Tinan (os).
CAMERON CO.: Sinnemahoning Creek. Fogg, Jr. 11555 (PENN).
CARBON CO.: Little Gap, Fogg, Jr. 11900 (MIN, PENN). CENTRE CO.:
Yarnell, Keener 2581 (PAC). CHESTER CO.: Harmonvville, Fender
1401 (PENN). CLARION CO.: Callensbure, Wood. Jr. 2170 (PENN).
CLEARFIELD CO.: Troutville, Ehrle 2475 (PAC). CLINTON CO.: Boone-
ville, Wahl 3198 (Pac). COLUMBIA co.: Mill Grove, Pohl 2554
(PENN). CUMBERLAND CO.: Shipnensbere, Saxe, Jr. 36 (PENN).
DAUPHIN Co.: Pillow. Berkheimer 12883 (PAC, PENN). DELAWARE
co: Darby, Baker (Pac). ELK CO.: Medix Run, McClelland (PENN).
FAYETTE CO.: Wvymps Gan, Bwinger (PENN). FOREST CO.: Brooks-
ton. Shields D-2157 (PAC). FRANKLIN CO.: Mercersburg, Kriner 104
(PAC). FULTON CO.: Deneen Gap, Westerfeld 7798 (Pac. PENN).
GREENE CO.: Jefferson. Wherry & Bell (PENN). HUNTINGDON CO.:
Maddensville, Westerfeld 9823 (PAC). JEFFERSON CO.: Sieel, Wahl
10944 (PAC). JUNIATA CO.: Cross Keys, Westerfeld 14535 (PAC).
LACKAWANNA CO.: Montdale, Adams 3542 (PENN). LANCASTER CO.:
Collins, Brubaker 1521 (PENN). LAWRENCE CO.: McConnells Mills,

358 Rhodora [Vol. 76

Russell 1582 (PENN), LEBANON CO.: Cold Spring, Travis 1170
(PENN). LUZERNE CO.: Mountaintop, Adams 3484 (PAC), LYCOMING
co.: Cedar Run, Wahl 19261 (PAC). MIFFLIN CO.: Standing Stone
Creek, Westerfeld 3961 (PAC). MONROE CO.: Scotrun, Niering 1214
(PAC). MONTGOMERY CO.: Audubon, Wherry (PENN). NORTHAMPTON
co.: Wind Gap, Schaeffer, Jr. 18252 (PENN). NORTHUMBERLAND C0.:
Sunbury, Wagner 8987 (PENN). PERRY CO.: Bloomfield, Adams &
Thebes 1232 (PENN). PIKE CO.: Porter’s Lake, DePue 136 (PAC).
SCHUYLKILL C0.: Delano, Wagner 3227 (PENN). SNYDER CO.: Paxton-
ville, Wade 783 (PENN). SOMERSET CO.: Jennerstown, Fogg, Jr. 15258
(PENN). SULLIVAN CO.: Laporte, Westerfeld 381 (PAC, PENN).
SUSQUEHANNA CO.: North Knob, Adams 3612 (PENN). TIOGA CO.:
Mainesburg, Fender 1537 (PENN). UNION CO.: Laurelton, Ludwig
1521 (PAC). VENANGO CO.: Franklin, Wahl 12633 (PENN). WARREN
co.: Irvington, Moldenke 17479 (ND). WAYNE CO.: White Mills,
Adams 4624 (PENN). WESTMORELAND CO.: Ligonier, Matthews 2399
(PAC). WYOMING CO.: Crane Swamp, Stone (PENN). YORK CO.:
Dillsburg, Hoover 3078 (PAC).

Rhode Island: NEWPORT CO.: Tiverton, Mearns 211 (US). PROVI-
DENCE CO.: Pascoag, Myers 350 (WVA). WASHINGTON CO.: Kingston
Hill, Reynolds 649 (HNH). South Carolina: ANDERSON CO.: Carpen-
ter’s Mill, Davis (MICH, MIN, UT). GREENVILLE CO.: Saluda River,
Freckman 1239 (ISC). LEXINGTON CO.: Batesburg, McGregor 297
(US). MARLBORO CO.: Little Pee Dee River, Radford 12494 (TENN).
OCONEE CO.: Whitewater, Davis 392 (MASS). PICKENS CO.: Table
Rock, Rodgers 540 (DUKE). SPARTANBURG CO.: Campobello, Erdman
973 (BRY). SUMTER CO.: Poinsette St. Park, Holdaway (DUKE).
Tennessee: ANDERSON CO.: Coal Creek, Kearney (MIN, OS, US).
BLOUT CO.: Cades Cove, Wallace & Jennison 1621 (TENN). CAMP-
BELL CO.: Jellico, Lamb & Williams (TENN). CHEATHAM CO.: Mar-
rowbone Creek, De Selm 1258 (TENN). CLAIBORNE CO.: Lone Mt.
Ridge, Kelley 2459 (TENN). COFFEE CO.: Old Stone Fort, Blum 3732
(VDB). CUMBERLAND CO.: Ozone, Demaree 47496 (KY, NO, VDB).
DAVIDSON CO.: Nashville, Hubbard 9112 (IND). DE KALB CO.: Smith-
ville, De Selme 173 (TENN). DICKSON CO.: Sycamore School, Jobe
(TENN). FENTRESS CO.: Clear Fork, Sharp 25648 (TENN). FRANKLIN
co.: Elk River, Sharp & Adams 10785 (TENN). GRAINGER CO.: Blaine,
Hodges 20 (TENN). GREENE CO.: Horse Creek Rd., Sherman & Sharp
22280 (TENN). GRUNDY CO.: Mont Eagle, Stifler (CU). HAMILTON
co.: Lookout Mt., Scribner (US). HARDIN CO.: Kral 36161 (VDB).
HAWKINS CO.: Clinch Mt., Wolfe 19180 (TENN). HOUSTON CO.: Erin,
Palmer 17610 (vs). JOHNSON CO.: Barclay 12B (TENN). KNOX CO.:
New Hopewell, Sharp 40454 (TENN). LAUDERDALE CO.: Summertown,
Postal (IND). LAWRENCE C0.: Shoal Creek, Sharp, Adams & Felix
11058 (TENN). LEWIS CO.: Trace Creek, Sharp, Adams & Felix
10065 (TENN). MARION CO.: Fosters Falls, Webb 368 (TENN).

1974] Genus Kalmia — Ebinger 359

MONROE CO.: Skona River, Galyon & Sharp 21603 (TENN). MORGAN
CO.: University Forest, Shanks 3141 (TENN). POLK CO.: Ocoee River,
Wilson 3726 (TENN). PUTNAM CO.: Monterey, Norris & Shanks 2475
(TENN). RHEA CO.: Spring City, James (08). ROANE C0.: Caney Creek,
Shanks 22078 (TENN). ROBERTSON CO0.: Pleasant View, Shanks,
Woods, & Cooley 14776 (TENN). SEVIER CO.: Pittman Center, Jen-
nison 3350 (TENN). UNICOI CO.: Rich Mt., Price 658 (DUKE). UNION
co.: Norris Lake, Kelley (TENN). VAN BUREN CO.: Spencer, Shanks
1288 (TENN). WASHINGTON CO.: Cherokee Mt., Sharp 3819 (TENN).
WAYNE CO.: Butler Creek, Shanks, Woods & Cooley 9156 (TENN).
WHITE CO0.: Caney Fork, Channell 8956 (VDB). WILLIAMSON CO.:
Ferndale, Major 6 (vpB). Vermont: BENNINGTON C0.: Pownal,
Eggleston (HNH, US). Virginia: ALBEMARLE CO.: Miller School,
Tinsley (NMC). AUGUSTA CO.: Stuarts Draft, Churchill (Msc).
BLANO CO.: Suiter, Kral 10453 (VDB). CRAIG CO.: Newcastle, Fogg,
Jr. 13211 (PENN). FAIRFAX CO.: Great Falls, Bartlet 1783 (IND).
GILES CO.: Salt Pond Mt., Fogg, Jr. 12688 (PENN). GREENE CO.:
Bear Fence Mt., Fosberg 17205 (PENN). ISLE OF WIGHT CO.: Smith-
field. Wise (KY). JAMES CITY C0.: Gordon Creek, Fernald & Long
11393 (DUKE). KING GEORGE CO.: King George, Muller 3787 (ND).
LOUDOUN CO.: Ashburn, Pratt (MU). MADISON CO.: Old Ray Mt.,
Fosberg 16909 (PENN). MONTGOMERY CO.: Blacksburg, Musselman
645 (UWM). NANSEMOND CO.: Nansemond River, Fernald & Long
13719 (US). NELSON CO.: Hall 3755 (VDB). NORFOLK CO.: Great
Dismal Swamn. Fernald & Long 12147 (TENN). PITTSYLVANLA CO.:
Smith Mt., Zobel (DUKE). ROANOKE CO.: Surarloaf Mt., Ebinger 4041
(MU). ROCKBRIDGE CO.: G. Washington Nat’l. Forest, Hall 3716
(VDB). ROCKINGHAM CO.: Shenandow Forest, Eggleston 18624 (us).
RUSSELL CO.: Clinch Mt., Kral 23850 (VDB). SMYTH CO.: Pond Mt.,
Small (msc, US). WISE CO.: Norton, Pennell 11813 (Isc). West
Virginia: BARBOUR CO.: Nestorville, Core 5922 (WVA). BERKELEY
co.: Back Creek. Core 5847 (WVA). BRAXTON CO.: Little Birch,
Harris (WVA). CALHOUN CO.: Pink, Harris (WVA). CLAY CO.: Clay,
Core 6372 (WVA). DODDRIDGE CO.: West Union, Bartholomew (WVA).
GRANT CO.: Dolly Sods Mt., Davis & Davis 7129 (WS). GREENBRIER
co.: Muddy Creek Mt., Fox (WVA). HAMPSHIRE CO.: Ice Mt., Frye
1269 (WVA). HARDY CO.: North Mt. Core & Ludwia 399 (WVA).
HARRISON CO.: Duck Creek, Martin 327 (WVA). JEFFERSON CO.:
Charlestown, Harper (CU). KANAWHA CO.: Nitro. Greenlee (WVA).
MCDOWELL CO.: Panther, Music (WVA). MARION CO.: Pinesville,
Bodola (WVA). MARSHALL CO.: Calis, Bartholomew 1941-217 (WVA).
MERCER CO.: Pinnacle Rock, Fox (WVA). MINERAL CO.: Patterson
Creek, Brown (WVA). MINGO CO.: Chattaroy, Bowen (WVA).
MONONGALIA CO.: Cassville, Watkins (WVA). MONROE CO.: Wayside,
Fox (WVA). MORGAN CO.: Langent, Core 5739 (WVA). NICHOLAS CO.:
Calvin, Creasy (WVA). PENDLETON CO: Seneca Rocks, Clarkson 2001

360 Rhodora [Vol. 76

(WVA). POCAHONTAS CO.: Droop Mt., Ludwig 77 (WVA). PRESTON
co.: Pisqah Rd., Meyer 247 (HNH, PENN, TENN, WVA). RANDOLPH
co.: Roaring Plains, Core 3171 (WVA). RITCHIE CO.: Cairo, Good-
win 251 (WVA). SUMMERS C0.: Barger Springs, Boone (WVA).
TAYLOR CO.: Wickwire Run, Core 5475 (WVA). TUCKER CO.: Black-
bird Knob, Gibson 1039 (WVA). UPSHUR C0.: Bucklin, Pollock (MIN,
US). WAYNE CO.: Hurricane Creek, Lycan 59 (WVA). WEBSTER CO.:
Camp Caesar, Fox (WVA). WETZEL CO.: Smithfield, Wagner (wva).
WIRT CO.: Standingstone Creek, Bartholomew W-5007 (WvA).

Mountain laurel commonly forms dense thickets in rocky
and sandy forests throughout most of its range, particularly
where there are openings in the canopy. It is also found in
pastures and open fields and commonly forms thickets at
the edge of roads. A detailed map of the known range of
Kalmia latifolia is given by Kurmes (1967) who considers
the report of this species from Ontario and New Brunswick
to be an error, as is the report from the valley of the Red
River in Arkansas (Britton, 1908). The report of this spe-
cies from Canada probably originates from Pursh (1814),
and it has been perpetuated by Brunet (1867), Sargent
(1893), and more recently by Munns (1938). There is no
evidence that this species is native to any part of Canada.
It is possible that these early reports were based on culti-
vated plants or large-leaved specimens of K. augustifolia
(Macoun, 1884). Presently the species is known from south-
ern and southeastern Maine (Fernald, 1911; Lamson-
Scribner, 1891; Norton, 1913, 1935; and Perkins, 1935),
southern New Hampshire (Hodgdon, 1955) and southern
Vermont (Deane, 1899).

The ecological requirements of Kalmia latifolia in the
deciduous forests of eastern United States have been exam-
ined by a few authors. Kurmes (1961) studied the ecology
of mountain laurel in southern New England and deter-
mined its exact range from herbarium material and field
observations. The relationship of this species to water
run-off and the water-holding capacity of the soil was
studied by Colvin and Eisenmenger (1943) and Johnson
and Kovner (1956). Studies have also been made of the
rate of water-vapor loss from its leaves (Meyer, 1927), the

1974] Genus Kalmia — Ebinger 361

seasonal variation in the moisture content of its leaves
(Reifsnyder, 1961) and seasonal changes in cold resistance
(Parker, 1961). The competition of mountain laurel with
tree seedlings and sprouts has also been studied. Wahlen-
berg and Doolittle (1950) found that under normal condi-
tions in the southern Appalachian mountains K. latifolia
forms dense thickets that preclude any natural regeneration
of timber trees. Their studies indicate that partial clearing
of these thickets followed by planting of desirable tree spe-
cies is an effective method of forest regeneration. They also
found that mountain laurel has a mean annual height
growth of 0.4 feet and a lateral growth of 0.3 feet.

Kalmia latifolia is usually a tall, spreading shrub that
throughout most of its range rarely exceeds a height of
12 feet. In the fertile valleys between the Blue Ridge and
the Alleghany Mountains, however, this species may attain
the size of a small tree. Engelmann, Canby, and Gray
(1877) observed a number of large individuals growing
in the bottom of a dell, back of Caesar’s Head, on the ex-
treme western border of South Carolina. One of the trunks
measured 4 feet 11/4, inches in circumference about 1 foot
above the ground while another in the same area was 3 feet
4 inches above the first fork, and at 1 foot above the ground
was 4 feet 4 inches in circumference. Later, Sargent (1893)
records its maximum size as 30-40 feet tall with a diameter
of 18-20 inches. The largest specimen on record, as listed
by the American Forestry Association (1955), is a plant
from the Great Smoky Mountains National Park with a
circumference of 3 feet 6 inches, a height of 25 feet, and a
spread of 45 feet. This specimen is probably located at
Forney Ridge, Swain County, Tennessee (Barksdale & Jen-
nison 1285).

Besides the numerous studies concerned with the poison-
ous properties of mountain laurel, some early attempts were
undertaken to determine what other substances were pres-
ent. These early studies reported the presence of arbutin,
gums, and tannins (Kennedy, 1875), bark tannins (De-
Graffe, 1896) and fat deposits (Stanley, 1931). More re-

962 Rhodora [Vol. 76

cently the effects of herbicides on Kalmia latifolia have
been studied. Egler (1947, 1948) found that 2,4-D has no
permanent effect, causing only temporary stunting and cur-
vature of the foliage. Later, Egler (1949, 1950) found that
a weak solution (.25%) of a 2,4-D and 2,4,5-T combination
causes temporary cessation of growth, but a strong solution
(596) is necessary to ensure death.

The synonomy of mountain laurel is not very extensive.
Most of the names proposed are for forms and varieties
which are the result of the extreme variation that exists in
flower color, leaf shape and size, plant size and pubescence.
In 1940, Fernald described a variety (laevipes) of Kalmia
latifolia from the coastal plain on southeastern United
States. He considered this variety in its extreme develop-
ment as having pedicels that are “quite smooth or with the
merest suggestion of sessile glandular spots". In typical K.
latifolia, in contrast, the pedicels are densely floccose and
viscid with stalked glandular hairs. Many individuals with
few or no glands on the pedicels are found in Indiana and
Ohio, as well as on the coastal plain from Virginia to
Louisiana. In some populations specimens with both gland-
less and glandular pedicels are found. Also, Radford, Ahles,
and Bell (1968) mention that this condition is very variable,
with a tendency for pedicels to be floccose on plants in the
mountains and less glandular on the coastal plain. Because
of this variation it appears best to consider these individuals
as an extreme that occurs in the southern and western parts
of the range of K. latifolia.

A total of 7 forms of Kalmia latifolia have been recog-
nized by Rehder (1910, 1945, 1949). Many of these were
originally described as varieties, but that classification is
not appropriate for these taxa in either the horticultural or
botanical sense. It is possible that these taxa could be
treated as cultivars as done by Holmes (1956) and Dudley
(1967), but most are better treated as forms as defined by
Davis and Heywood (1963). The genetic evidence indicates
that most of these taxa are not unique clones, but rather
are sporadic variants distinguished by a single or a few

1974] Genus Kalmia — Ebinger 363

linked characters, without a distinct distribution. Also,
they do not represent the normal extremes of most popula-
tions, but are the result of the repeated association of re-
cessive genes that are of a limited occurrence in some popu-
lations. Of the 7 forms recognized by Rehder, 5 fit the
criteria of a form as described above. The remainder, both
variations in flower color (alba and rubra), unlike the
forms, are not aberrant or disjunct, but are connected to
the typical light pink flower by intermediate color types.

Kalmia latifolia f. angustata Rehd. Jour. Arnold Arb. 26:
481. 1945. TYPE: New Jersey: CAPE MAY CO.: Dennis town-
ship, H. A. Scribner (GH).

Kalmia latifolia B. salicifolia Forbes, Hort. Woburn. 93.
1833. (nom. nud.).

This very rare foliage form has narrowly oblanceolate to
linear leaves that are 4-8 cm long and 4-10 mm wide. Liv-
ing material of the type is cultivated at the Arnold Arbore-
tum (Rehder, 1945). The only specimens of f. angustata
seen by the author are cultivated plants.

Common name: Willow-leaved Mountain Laurel.

Kalmia latifolia f. fuscata (Rehd.) Rehd. Rhodora 12: 2.
1910.

Kalmia latifolia var. fuscata Rehd. Moller's Deutsch.
Gart. Zeit. 18: 578. 1903. TvPE: New York: Canaan Four
Corners, Dr. Childs (GH).

Kalmia latifolia var. coronata Gray, Amer. Nat. 2: 324.
1868. TYPE: Massachusetts: woods near Framingham,
James Parker (GH).

Kalmia latifolia var. maculata Raf. Medical Flora 2: 17.
1830. TvPE: unknown.

This form is distinguished by white to pink flowers that
have a heavily pigmented, usually continuous, brownish-
purple or cinnamon band on the inside of the corolla at the
level of the anther pockets. This band breaks up into
brownish dots toward the base and the margin of the
corolla. It also shows through to the outside of the bud
giving it a muddy appearance.

364 Rhodora [Vol. 76

Some variation does exist in the size, shape and color of
the band with an interrupted band existing in some of the
specimens examined. According to Jaynes (personal com-
munication) controlled crosses with specimens showing this
banded characteristic strongly suggests that the character
is determined by a single dominant gene.

Common name: Banded Mountain Laurel, Crowned Moun-
tain Laurel.

SPECIMENS EXAMINED

Alabama: near Citronella, Milligan (US). Connecticut: near
Chaplin, Jaynes (EIU). Massachusetts: near Framingham, Parker
(GH); Wales, Clark (MASS); Guterman (MASS). New York: Brew-
ster, Jaynes 240 (EIU). North Carolina: Highlands, Anderson (US).
Ohio: Holden Arboretum, Mentor, Jaynes 94 (EIU). Pennsylvania:
nursery in Stroudsburg, Jaynes 168 (EIU).

Kalmia latifolia f. myrtifolia (Bosse) K. Koch, Dendrol-
ogie 2(1) : 153. 1872.

Kalmia latifolia var. myrtifolia Bosse, Vollst. Handb.
Blumengart. 2: 348. 1841. Kalmia myrtifolia Andre, Revue
Hort. 55: 10. 1883. TYPE: unknown, probably a living plant.

Kalmia latifolia f. minor K. Koch, Dendrologie 2(1) : 153.
1872. TYPE: unknown, probably a living plant.

Kalmia latifolia var. nana Lavallée, Arbor. Segrez. 159.
1877. (nom, nud.); Mouillefert, Traité Arb. Arbrisse 2:
1027. 1897. TYPE: unknown, probably a living plant.

This form is in all respects a miniature mountain laurel,
being compact and slow-growing, rarely exceeding a height
of 3 feet. In all specimens examined the leaves are much
smaller than in typical Kalmia latifolia, averaging 2-4 cm
long and about 1 cm wide. Also, flower size and length of
the internodes are approximately 1/3 to 1/2 normal. Recent
evidence (Jaynes, personal communication) suggests that
this dwarf condition is controlled by a single recessive gene.

According to Dudley (1967) this form has been grown
since 1840 and is occasionally found in gardens where space
is limited. Though occasionally mentioned in the early
horticulture literature of Europe, the first reference to
this form in North American literature appears to be by

1974] Genus Kalmia — Ebinger 365

Sprague (1871). The only report of wild plants of f. myrti-

folia appeared in 1895 (Anonymus) which, according to the

author, were identical to the cultivated dwarf form.
Common name: Miniature Mountain Laurel, Dwarf

Mountain Laurel.

SPECIMENS EXAMINED

Connecticut: Portland, Jaynes 25 (EIU). Massachusetts: Arnold
Arboretum, Hornby (MASS).

Kalmia latifolia f. obtusata (Rehd.) Rehd. Rhodora 12: 2.
1910.

Kalmia latifolia var. obtusata Rehd. Móller's Deutsch.
Gart. Zeit. 18: 577. 1903. TYPE: Connecticut: near Pom-
fret, Bowditch.

Kalmia latifolia var. ovatifolia Raf. Medical Flora 2: 17.
1830. (nom. nud.).

This foliage form has leaves that are oval to oblong-
obovate and rounded at both ends. In the specimens exam-
ined the leaves average 4-7 cm long and 2.5-4 cm broad, are
apiculate, and have short petioles that rarely exceed 1 cm
in length. According to Rehder (1910) this form is slow
growing, forms a compact bush, and has been cultivated at
the Arnold Arboretum since 1886 when it was propagated
from the type plant. Dudley (1967) suggested that the
cultivar ‘Ovata’ which is available from some European
sources is a synonym of this variant.

Common name: Hedge Mountain Laurel.

SPECIMENS EXAMINED
Connecticut: Riverton, Jaynes 291 (EIU). Massachusetts: 1 mi.

east of Ashby, Ahles 65185 (MASS); Arnold Arboretum, Wyman
2854 (MSC), Jaynes 238 (EIU).

Kalmia latifolia f. polypetala (Nickolson) Beissner,
Schelle, & Zabel, Handb. Laubholz-Benennung 386. 1908.

Kalmia latifolia var. polypetala Nicholson, Hand-list
Trees Shrubs Arb. 2: 49. 1896. Kalmia latifolia f. polype-
tala (Nicholson) Rehd. Rhodora 12: 1. 1910. TYPE: Mas-
sachusetts: near South Deerfield, Miss M. Bryant (GH).

Kalmia latifolia var. monstruosa Mouillefert, Traité Arb.

366 Rhodora [Vol. 76

Arbrisse 2: 1027. 1897. TYPE: Unknown, probably a living
plant.

This form has the corolla deeply divided into 5 narrow
to fairly broad, strap-shaped petals. In some specimens
these petals are extremely narrow and thread-like due to
the revolution of the margins. Gray (1871) reports that
these narrow petals resemble filaments and are sometimes
tipped with imperfect anthers. More commonly the petals
are broader, and a few specimens have been found in which
the individual flowers appear much like apple blossoms.
Other variations include an apetalous type which has been
observed in some nursery stock from Massachusetts, and a
cultivar named ‘Bettina’ (Dudley, 1967) in which the corolla
is deeply lobed. According to Jaynes (personal communica-
tion) the inheritance data fits the hypothesis that the genetic
control of the polypetalous condition is by a single recessive
gene.

This form was first described by Gray (1871) and later
by Sargent (1890), while Bean (1897) mentioned that this
unusual form flowers each year at the Kew Gardens. It was
given the name var. polypetala by Nicholson (1896) and
the name var. monstruosa by Mouillefert (1897), both
based on Sargent's description of 1890. This taxon was
first considered as a form by Beissner, Schelle, and Zabel
(1903) who use the name polypetala and cite Sargent as
the author.

This form appears to occur most commonly in Massachu-
setts. Stone (1909) reported it growing at the edge of a
road in Leverett, not far from Mt. Toby, while Britton
(1925) reports that Dr. Stone has found it from three dif-
ferent stations, all on or near Mt. Toby. Mt. Toby is close
to South Deerfield, the type locality of this form.

Common name: Feather Petal Mountain Laurel.
SPECIMENS EXAMINED

Massachusetts: Arnold Arboretum, Wyman 2458 (Msc); Leverett,
Mt. Toby, Henry (MASS), Osmun (MASS), Stone (MASS), 67570
(Mass). North Carolina: Danbury, Hanging Rock State Park,
Rogers (EIU).

1974] Genus Kalmia — Ebinger 367

4. Kalmia angustifolia Linnaeus, Sp, Pl. 1: 391. 1753.
Type: “Habitat in Pensylvania, Nova Caesarea, Nove-
boraco" (LINN 560.2, lectotype, specimen on the right).

Chamaedaphne angustifolia (L.) Kuntze, Rev. Gen. PLA
388. 1891.

Much branched shrub to 1.5 m tall; branchlets terete,
reddish-brown, glabrous to pubescent and stipitate-glandu-
lar, usually ascending; leaves in whorls of 3 (rarely a few
alternate or opposite), slightly coriaceous, evergreen, flat,
oblong to elliptic-lanceolate, 2-5 (rarely 7) cm long, 0.5-2 cm
wide, usually lightly puberulent above with short white
hairs, sometimes glabrate with age, midrib puberulent
above, glabrous to puberulent beneath, sometimes stipitate-
glandular; petioles to 1 (rarely 1.5) cm long, usually pu-
berulent; inflorescence of numerous corymbiform racemes
from the axils of last year’s leaves; pedicels 0.5-2 cm long,
puberulent and stipitate-glandular, subtended by 2 bracte-
oles in the axil of a lanceolate bract, 1-6 mm long; main
axis of the inflorescence less than 1.5 cm long; calyx pu-
berulent outside, glabrous within, usually green, sometimes
the tip reddish, or reddish throughout, 3-6 mm broad, the
lobes usually acuminate; corolla reddish-purple to pink
(rarely white), puberulent outside, puberulent toward the
base within, 6-13 mm broad; anthers purplish-brown, .5-1
mm long; filaments puberulent on the lower half; capsule
depressed globose, 3-5 mm broad, 2-3.5 mm long, puberu-
lent and stipitate-glandular; seed stramineous, 0.6-1 mm
long, and testa loose and extending past the ends of the
seed. Chromosome number n — 12.

Kalmia angustifolia is a common species in eastern North
America. It is found from northern Ontario, Quebec and
Labrador, through the Great Lakes region, and south along
the east coast of the United States to the Carolinas. It is
primarily a plant of bogs and other wet, open areas, but is
also found in open woods and as a weed in open areas, some-
times forming dense thickets.

Genetically this species is most closely related to Kalmia
latifolia. When K. angustifolia is the male parent the cross

368 Rhodora [Vol. 76

is not successful because the sheep laurel pollen will not
grow down the style of mountain laurel (Jaynes, 19682).
The reciprocal cross, however, resulted in about 18% of
the flowers producing capsules which contained some viable
seed. These resulting interspecific hybrids were intermedi-
ate to the parental types, although leaf arrangement was
primarily alternate, as in K. latifolia. Jaynes (1968a) also
reported that K. angustifolia could be crossed with both
K. hirsuta and K. polifolia. Both crosses produced mostly
weak, yellow-green individuals that died within the year.
A few individuals lived longer, and some produced flowers
with sterile pollen.

Small (1903, 1914) considered this taxon as two separate
species, a procedure not followed by subsequent authors.
Both Fernald (1950) and Gleason (1952) considered this
complex as being one species with two fairly distinct vari-
eties, a view that has recently been followed by Radford,
Ahles, and Bell (1968). Considering the genetic and mor-
phological similarities of these taxa it is more realistic to
follow the latter view and consider it as a single species
with two fairly distinct varieties.

KEY TO VARIETIES

l. Calyx densely stipitate-glandular; mature leaves not
puberulent beneath. ............ 4a. var. angustifolia.
l. Calyx lacking stipitate-glands, or nearly so; mature
leaves densely puberulent beneath. . 4b. var. caroliniana.

4a. Kalmia angustifolia L. var. angustifolia.
Kalmia angustifolia B carnea Ait. Hort. Kew. 2: 64. 1789.
Kalmia angustifolia var. hirsuta Jager, Ziergeholze 278.

1865. Kalmia angustifolia f. hirsuta Voss, Blumengart. 1:
591. 1894. (without basonym).

Kalmia angustifolia var. lucida Lavalée, Arbor. Segrez.
159. 1877. (nom. nud.). Kalmia angustifolia f. lucida
Beissner, Schelle, & Zabel, Handb. Laubholz-Benennung 386.
1903. (nom. nud.).

1974] Genus Kalmia — Ebinger 369

Kalmia angustifolia var. minima Bosse, Vollst. Handb.
Blumengart. 2: 347. 1841.

Kalmia angustifolia var. nana Bosse, Vollst. Handb.
Blumengart. 2: 347. 1841. Kalmia angustifolia f. nana
Voss, Blumengart. 1: 591. 1894. (as snyonym).

Kalmia angustifolia B ovata Pursh, Fl. Amer. Sept. 296.
1814.

Kalmia angustifolia f. procumbens Beissner, Schelle, &
Zabel, Handb. Laubholz-Benennung 386. 1903. (nom.
nud.).

Kalmia angustifolia, var. rosea Forbes, Hort. Woburn. 93.
1833. Kalmia angustifolia f. rosea Beissner, Schelle, &
Zabel. Handb. Laubholz-Benennung 386. 1903. (nom.
nud.).

Kalmia angustifolia var. variegata Forbes, Hort. Woburn.
93. 1833.

Kalmia elliptica Raf. Autikon Bot. 87. 1840. (as syno-
nym).

Kalmia ferruginea Raf. Autikon Bot. 86. 1840. TYPE:
unknown.

Kalmia media K. Koch, Dendrologie II; 1: 153. 1872.
Kalmia angustifolia f. media Beissner, Schelle, & Zabel,
Handb. Laudholz-Benennung 386. 1903. (nom. nud.) TYPE:
unknown.

Kalmia intermedia Lange Ind. Sem. Hort. Hauniensi
Nom. emendata 3. 1873. Kalmia glauca B intermedia Lange
Ind. Sem. Hort. Hauniensi Nom. emendata 3. 1873. (as
synonym). TYPE: a herbarium specimen in the Botanical
Museum of Copenhagen (C) labeled Kalmia intermedia
Lge ad int. Hort. bot. Haun. 16. Juni 1873, sem. ex hort.
Paris s.n.

Kalmia angustifolia B oleifolia Pers. Syn. Pl. 1: 4T".
1805. Kalmia oleifolia (Pers.) Bosse, Vollst. Handb. Blu-
mengart. 2: 347. 1841. (as synonym). TYPE: unknown.

Kalmia angustifolia var. pumila Forbes, Hort. Woburn.
93. 1833. Kalmia pumila Bosse, Vollst. Handb. Blumengart.
2:347. 1841. (as synonym).

370 Rhodora [Vol. 76

Kalmia pumila rubra Bosse, Vollst. Handb. Blumengart.
2:347. 1841. (as synonym).

Kalmia pumila serotina Bosse, Vollst. Handb. Blumengart.
2: 347. 1841. (as synonym). Kalmia angustifolia f. pumila
Voss, Blumengart. 1: 591, 1894. (without basonym).
TYPE: unknown.

Kalmia angustifolia « rubra Ait. Hort. Kew 2: 64. 1789.
Kalmia angustifolia var. rubra (Ait.) Loddiges, Bot. Cab.
6: 502. 1821. Kalmia rubra (Loddiges) Bosse, Vollst.
Handb. Blumengart. 2: 347. 1841. (as synonym). Kalmia
angustifolia f. rubra Voss, Blumengart. 1: 591. 1894.
(without basonym). TYPE: unknown.

Kalmia angustifolia var. serotina Bosse, Vollst. Handb.
Blumengart. 2: 347. 1841. Kalmia serotina Bosse, Vollst.
Handb, Blumengart. 2: 347. 1841. (as synonym). TYPE:
unknown.

Mature leaves glabrous or with scattered stalked glandu-
lar hairs beneath, usually not puberulent; calyx puberulent
and strongly stipitate-glandular; corolla glabrous or lightly
puberulent on the outside.

Distribution: Bogs, Swamps, and other wet open areas
from Ontario east to the Atlantic Ocean and southeast
through the Great Lakes to Pennsylvania and south along
the coast to Virginia.

Common name: Sheep Laurel, Lambkill, Sheepkill,
Wicky, Narrow-leaved Kalmia, Dwarf Laurel, Pig Laurel.

REPRESENTATIVE SPECIMENS
UNITED STATES: Connecticut: FAIRFIELD CO.: Trumbull, Eames
2809 (CONN). HARTFORD CO.: Kensington, Cowles (MU). LITCHFIELD
CO.: Salisbury, Ebinger 697 (MU). NEW HAVEN co.: New Haven,
Safford 163 (US). NEW LONDON CO.: Groton, Jausson (CONN, UT).
TOLLAND CO.: Donovans Bog, Travis 1959 (PENN). WINDHAM CO.:
Hasting’s Woods, Petersen (VDB). Delaware: NEW CASTLE CO.: Hog-
swamp Road, Tatnall 4858 (GH). SUSSEX CO.: Canby (Us). Maine:
ANDROSCOGGIN CO.: Lewiston, Bean 14793 (VDB). AROOSTOOK CO.:
Knoles Corner, Neal 1907 (MAINE). CUMBERLAND CO.: Cape Eliza-
beth, Gayle 766 (Us). HANCOCK CO.: Bar Harbor, Rossbach 2757
(ACAD, NCU). KENNEBEC CO.: Clinton, Bean (MAINE). KNOX CO.:

1974] Genus Kalmia — Ebinger 371

Glencove, F'riesner 7721 (MU, RENO, UT, WS, WVA). LINCOLN CO.:
Ocean Point, Fassett 3946 (DUKE). OXFORD CO.: White Mt. Nat.
Forest, Richards 6066 (MAINE). PENOBSCOT CO.: Old Town, Davis
(MAINE). PISCATAQUIS CO.: Sangerville, Fernald 127 (MIN). SAGA-
DAHOC CO.: Five Islands, Rehder 1018 (GH). SOMERSET CO.: Baker
Brook, St. John & Nichols 2431 (CAN, NY). WALDO CO.: Searsmont,
Rossbach 1932 (ACAD, MAINE). WASHINGTON CO.: Joe Dyer's Point,
Walker 3039 (MAINE). YORK CO.: Sand Pond, Richards 5951 (MAINE).
Maryland: ANNE ARUNDEL C0.: Severn Run, Adams & Wherry 3200
(GH). cECIL co.: North East, Long 54422 (GH). CHARLES CO.:
Waldorf, Niewwland (ND). PRINCE GEORGES CO.: Airport Bog, Her-
man 11546 (NY). Wicomico CO.: Sharptown, Tidestrom 12229 (GH).
WORCESTER CO.: Furnace, Redmond 237 (MICH). Massachusetts:
BARNSTABLE CO.: Falmouth, Greenman 3043 (MIN). BERKSHIRE CO.:
Great Barrington, Eggleston 21413 (MIN, NY). ESSEX CO.: Marble-
head, Nelson 341 (CAN). FRANKLIN CO.: Mt. Toby, Seymour 2566
(DUKE, MASS). HAMPDEN CO.: Granville, Seymowr 137 (MASS, MIN,
NY). HAMPSHIRE CO.: Amherst, Torrey (MASS). MIDDLESEX CO.:
Townsend, Ahles 65188 (MASS). NANTUCKET CO.: Polpis, Mackeever
504 (PENN). NORFOLK CO.: Medfield, Batchelder 234 (NCU).
PLYMOUTH CO.: Ellisville, Erdman 4223 (BRY). WORCESTER CO.:
Mount Wachusett, Palmer 43196 (MIN). Michigan: ALCONA CO.:
Lincoln, Voss 4583 (MICH). ALPENA CO.: Alpena, McVaugh
11405 (MICH). | ARENAC CO.: Omer, Voss 6794 (MICH).
CHARLEVOIX CO.: Garden Isl, Cain 149 (TRT). CHEBOYGAN CO.:
Wolverine, Gleason & Gleason, Jr. 248 (GH, IND, ISC, NY, WVA).
CRAWFORD CO.: Grayling, Voss 3098 (MICH). 10sCO CO.: Sand Lake,
Dreisbach 7243 (CONN). KALKASKA CO.: Zavitz 1168 (QUK). MIS-
SAUKEE CO.: Moorestown, Voss 3087 (MICH, OS). MONTMORENCY CO.:
Hagenah 4936 (BLH). oscopa Co.: Rhodes Lake, Voss 3578 (MICH).
OTSEGO CO.: Hardwood Lake, MeVaugh 10888 (BLH, CAN, MICH, MO).
PRESQUE ISLE CO.: Emma, Clover 26193 (MICH). ROSCOMMON CO.:
Roscommon, Voss 4258 (MICH). SAGINAW CO.: Sage Lake, Case, Jr.
(MICH). New Hampshire: BELKNAP CO.: Crystal Lake, Hodgdon
& Steele 9124 (NHA). CARROLL CO.: Albany, Gale (NHA). CHESHIRE
co.: Winchester Manning (BRY). coos Co.: Randolph, Pease 4696
(GH, MICH). GRAFTON CO.: Pike, Mathias 163 (MIN). HILLSBORO
co.: Mt. Monadnock, Andrews 895 (ORE). MERRIMACK CO.: Wilmot,
Hodgdon & Steele 9949 (NHA). ROCKINGHAM CO.: Harris 224 (NHA).
STRAFFORD CO.: Rochester, Barrett & Hodgdon 15522 (NHA). SULLI-
VAN CO.: Springfield, Hodgdon & Steele 9129 (NHA). New Jersey:
ATLANTIC CO.: Hammonton, Keefe 123 (UWM). BERGEN CO.: Carl-
stadt, Niederer (NY). BURLINGTON CO.: Rancocas River, Leonard
6314 (US). CAMDEN CO.: Atco, Painter 684 (MIN, US). CAPE MAY
co.: Belleplain, Fender 4916 (PAC, PENN), CUMBERLAND CO.: Mill-
ville, Adams 324 (GH, PENN). ESSEX CO.: Franklin, Rusby (CHRB).

372 Rhodora [Vol. 76

GLOUCESTER CO.: Repaupo, Long 16065 (GH). MIDDLESEX CO.: Mill-
town, House (US). MONMOUTH CO.: Farmingdale, Long & Brown
9557 (PENN). MORRIS CO.: Mt. Hope, Britton (CHRB). OCEAN CO.:
Lakewood, Mackenzie 3112 (Us). PASSAIC co.: Uttertown Bog,
Montgomery (CHRB). SALEM CO.: Auburn, Fogg, Jr. 6616 (PENN).
SOMERSET CO.: Sourland Mts., Fogg, Jr. 12406 (PENN). SUSSEX CO.:
Cranberry Lake, Mackenzie 717 (ARIZ, IND).

New York: ALBANY CO.: Karmer, House 7850 (CAN). CHENANGO
co.: Plymouth Pond, Wiegand 6976 (GH). CLINTON CO.: Rudd 816
(US). COLUMBIA CO.: Ghent, McVaugh 362 (PENN). DUTCHESS CO.:
Susan Court, Ahles 64464 (MASS). ESSEX CO.: Lake Harris, House
8072 (CAN). FRANKLIN CO.: Upper Saranac Lake, Muenscher, et. al.
476 (NY). FULTON CO.: Gloversville, Haught (WVA). GREENE CO.:
Coville (GH). HAMILTON CO.: Little Tupper Lake, Fosberg 16101
(PENN). JEFFERSON CO.: Perch Lake, Beschel 17144 (QUK). LEWIS
co.: Martinsburg, ‘Hotchkiss 2654 (GH). NASSAU CO.: Massapequa
St. Park, Cain 93 (MO, PENN, TENN). ONEIDA CO.: Sylvan Beach,
Maxon (US). ORANGE CO.: Sutherland Pond, Mearns 125 (US).
RENSSELAER CO.: West Sand Lake, Brown 652 (DUKE). RICHMOND
co.: Staten Isl, Dowell & Davis 2224 (GH). ST. LAWRENCE CO.:
Norfolk, Phelps 756 (GH, NY, US). SARATOGA CO.: Corinth, Whitney
4124 (PENN). SUFFOLK CO.: South Haven, Moldenke 18685 (NY).
SULLIVAN CO.: Wurtshore, Hess 1267 (ARIZ). ULSTER CO.: Highland,
Chrysler (CHRB). WARREN CO.: Luzerne, Fogg, Jr. 20309 (PENN).
WASHINGTON CO.: Lake Nebo, Burnham 2756 (MIN). Pennsylvania:
BERKS C0.: Hamburg, Berkheimer 1734 (PENN). BRADFORD CO.: Troy,
Beardslee (MU). BUCKS CO.: Bristol, Adams 4514 (MAINE, PENN).
CARBON CO.: Little Gap, Adams 3331 (GH, ISC, PENN). CHESTER CO.:
Warwick, Wherry (GH). COLUMBIA CO.: Centralia, Fogg, Jr. 14256
(NY, PENN). CUMBERLAND CO.: Sand Springs, Ward (PAC), DELA-
WARE CO.: Tinicum, Baker (PENN). FRANKLIN CO.: Caledonia St.
Park, Overhots (PAC). LACKAWANNA CO.: West Mt., Glowenke 6528
(MIN, NY). LANCASTER CO.: Smithville, Small (US). LEHIGH CO.:
Allentown, Dowell 1115 (Us). LUZERNE CO.: Freeland, Orton (PAC).
LYCOMING CO.: Bald Eagle Mt., Keener 2561 (PAC). MONROE CO.:
Tannersville, Swartley 791 (MAINE). NORTHAMPTON CO.: Daniels-
ville, Schaeffer, Jr. 6184 (PAC). NORTHUMBERLAND CO.: Montandon,
Reed 235 (CHRB). PIKE CO.: Twin Lakes, De Pue 878 (PENN).
SCHUYLKILL CO.: Tamaqua, Kriebel 8586 (ND). SOMERSET CO.:
Salisbury, Caffrey (PAC). SULLIVAN CO.: Double Run, Fogg, Jr.
14437 (PENN). WAYNE CO.: Big Spring Pond, Glowenke 2434
(PENN). WYOMING CO.: Falls, Glowenke 2023 (PENN). YORK CO.:
McCall’s Ferry, Heller & Halbach (MIN). Rhode Island: KENT CO.:
Warwick, Congdon (08). NEWPORT CO.: Newport, Mearns 254 (GH).
PROVIDENCE CO.: Slaterville, Palmer 43348 (ISC, MIN). WASHINGTON
co.: South Kingstown, Collins (CONN). Vermont: ADDISON CO.:

1974] Genus Kalmia — Ebinger 373

Bristol Pond, Eggleston 21453 (US). BENNINGTON CO.: Manchester,
Day 267 (GH). CALEDONIA CO.: Barnet, Blanchard (HNH, MO).
CHITTENDEN CO.: S. Burlington Twp., Charette 2190 (BRY, MIN).
FRANKLIN CO.: East Berkshire, Clausen (MO, US). ORANGE CO.:
Fairlee, Grover (MU). RUTLAND CO.: Middletown Springs, Carpenter
(NCU). WASHINGTON CO.: Roxbury, Countryman 820 (HNH, NHA).
WINDHAM CO.: Westminster, Robinson 30 (GH). WINDSOR CO.:
Reading, Kittredge (NY). Virginia: ISLE OF WIGHT Co.: Franklin,
Heller 1124 (GH, MIN, MO, PENN, US, WS); Lee’s Mill, Fernald &
Long 12146 (GH, MO, NY, US), Harvill 15946 (NCU); Zuni, Fernald,
Griscom & Long 6665 (GH, NY), 6666 (GH, PENN). NANSEMOND CO.:
Cherry Grove, Fernald & Long 11392 (DUKE, GH, US); County Rd.
665, Duke 1067 (NCU); South Quay, Fernald & Long 10366 (GH,
NY).

CANADA: Labrador: Assizes Isl., Potter & Brierly 3424 (GH);
Cape St. Charles, Potter & Brierly 3427 (GH); Cartwright, Brown
(GH); Forteau, Long 28832 (GH); Goose Bay, Gillett & Findley 5352
(MIN, MO, TRT, WS), 5584 (ACAD, NY, US); Kenamu River, Gillett
& McKay 5831 (ACAD, GH); St. Clary’s River, Wallace 23 (CONN).
New Brunswick: ALBERTA CO.: Grindstone Isl. Christie 1575 (NBM).
CARLETON CO.: East Wolf Isl, Hodgdon & Pike (NCU). CHARLOTTE
co.: Deer Isl., Squires & Squires (NBM), GLOUCESTER CO.: Four
Roads, Roberts & Pugh 65-4358 (CAN). KENT CO.: Richibucto,
Scoggan 12135 (CAN). KINGS CO.: Havelock, Perry (ACAD). MADA-
WASKA CO.: St. Leonard, Malte 294 (CAN). NORTHUMBERLAND CO.:
Mt. Carleton, Smith & Clattenburg 20066 (ACAD, QUK). QUEENS CO.:
Mill Brook, Squires & Christie (NBM). RESTIGOUCHE CO.: Dalhousie,
Scoggan 11721 (ACAD, CAN). ST. JOHN CO.; Mispec, Crawford &
Patterson (ACAD). SUNBURY CO.: Mill Settlement, Christie 245 (NBM).
WESTMORLAND CO.: Johnson Lake, Beschel 5508 (QUK). Newfound-
land: BONAVISTA NORTH DIST.: Glovertown, Smith, Squires, & Smith
116 (ACAD), 135 (ACAD). BONAVISTA SOUTH DIST.: Lethbridge,
Rouleau 8370 (NFLD). BURGEO-LA POILE DIST.: Osmond, Rouleau 7375
(NFLD, US). CARBONEAR DIST.: Old Perlican, Torrey 25 (CAN, GH).
FERRYLAND DIST.: Witless Bay, Rouleau 5364 (NFLD). FOGO DIST.:
Stag Harbour, Ludlow 125 (ACAD). FORTUNE BAY-HERMITAGE DIST.:
Wallace Cove, Rouleau 8089 (NFLD). GANDER DIST.: Gander, Rouleau
6133 (CAN, NFLD, US). GRAND FALL DIST.: Overflow Pond, Rouleaw
7961 (NFLD). HUMBER DIST.: Corner Brook, Rouleau 3960 (ACAD,
CAN, GH, NFLD, NY, US). ST. BARBE DIST.: Bonne Bay, Bishop 471
(GH, CAN). ST. JOHN’s DIST.: St. John's, Robinson & Schrenck 41
(GH, CAN, MIN, MO, NY, P). TRINITY SOUTH DIST.: Whitbourne,
Rouleaw 6069 (NFLD). TWILLINGATE DIST.: Loon Bay, Rouleau 4866
(NFLD). WHITE BAY DIST.: St. Anthony, Savile & Vaillancourt 2561
(NCU, NFLD, NY). Nova Scotia: ANNAPOLIS CO.: Belleisle, Bent
12723 (ACAD). ANTIGONISH co.: Linwood, Smith, et. al. 10765

374 Rhodora [Vol. 76

(MAINE). CAPE BRETON CO.: Frenchvale, Smith, et. al. 15094 (ACAD).
COLCHESTER CO.: Truro, Malte, (CAN, QUK). CUMBERLAND CO.:
Wharton, Schofield 3197 (ACAD). DIGBY CO.: Corberrie, Jack 3121
(MAINE, NY, US). GUYSBOROUGH CO.: Canso, Rousseau 35450 (CAN).
HALIFAX C0.: Halifax, Jack 3235 (NY, TENN). HANTS CO.: Bishop-
ville Rd., Wellwood 12662 (ACAD). INVERNESS CO.: North Mt., Smith,
et. al. 11275 (ACAD). KING CO.: Wolfville, Anderson (ACAD).
LUNENBURG C0.: Beck Lake, Smith, et. al. 18551 (ACAD, NCU). PICTOU
co.: Westville, Sampson 28 (ACAD). QUEENS CO.: Tobeatic Lake,
Martin 5210 (CAN). RICHMOND CO.: Grand Lake, Ershine 51993
(QUK). SHELBURN C0.: Barrington Passage, Macoun 81371 (CAN).
VICTORIA CO.: Cape Breton, Senn 1553 (MIN, US). YARMOUTH CO.:
Seal Isl, Erskine 54.977 (ACAD). Ontario: BRUCE CO.: Big Bay,
Krotkov 9295 (NY). CARLETON CO.: Wright’s Cove, Minshall 236
(MIN, NY). COCHRANE CO.: Abitibi, Pritchard 208A (MASS). DUNDAS
co.: Elma, House 23357 (PENN). FRONTENAC CO.: Kingston, Fowler
(US). HALIBURTON CO.: Minden Twp., Roberts (TRT). KENORA DIST.:
Attawapiskat River, Porsild, et. al. 19986 (CAN). LENNOX CO.: Skoota-
matta, Jackson (TRT). MANITOULIN DIST.: Honora, Soper & Bell 9819
(CAN, QUK, TRT). MUSKOKA DIST.: Sparrow Lake, Walden 110 (TRT).
NIPISSING DIST.: Stuarts, Ross 2 (TRT). ONTARIO CO.: Cannington,
Thompson 516 (TRT). PARRY SOUND DIST.: Ahmic Lake, Soper 5225
(CAN, TRT). PRESCOTT CO.: Alfred Twp., Calder & Cody 922 (PENN).
RENFREW CO.: Renfrew, Stroud (TRT). STORMONT CO.: Newington,
Jenkins 8297 (NCU). SUDBURY DIST.: Sudbury, Ostenfeld 613 (CAN).
THUNDER BAY DIST.: Longlac, Baldwin & Breitung 3693 (CAN, TRT).
TIMISKAMING DIST.: Cattle Isl., Moss 77 (ALTA). WELLAND C0.:
Niagara, Zenkert (OAC). Prince Edward Island: KING CO.: Dundee,
Fernald, Long, & St. John 7899 (GH). QUEENS C0.: Brackley Beach,
Grandtner 18518 (CAN). Quebec: ABITIBI CO.: Amos, Morton 11035
(US). ARGENTEUIL CO.: Lachutem, Porsild 6337 (CAN). BERTHIER
co.: Lanoraie, Scoggan 370 (CAN). BONAVENTURE CO.: New Rich-
mond, Chrysler 1228 (CHRB). BROME CO.: Bolton, Knowlton (GH).
CHICOUTIMI CO.: Lac Kenogami, Lavoie, Doyon, & Lord 600706-01
(ACAD, TRT). GASPE WEST CO.: Mt. Albert, Cottam 11852 (UT).
GATINEAU CO.: Wilson Corners, Mitchell 1750 (TENN). KAMOURASKA
co.: Ouelle, Gauthier 65-207 (QUK). LABELLE CO.: Nomininque, Roy
1053 (ACAD). LEVIS CO.: Chaudiere, Forbes (CAN). MAGDALEN ISLAND
co.: Amherst Isl, Potter & Brierly 3425 (NMC, UT). MASKINONGE
co.: Metis Beach, DuBoulay & DuBoulay 2589 (QUK). MATANE CO.:
Lake Montagne, Boivin 786 (PENN). MATAPEDIA CO.: Saint Zenon,
Blain 321 (CAN). MEGANTIC CO.: Black Lake, Blais, Deshaies, &
Forest 10379 (CAN, QUK, SASK). MISSISQUOI CO.: Farnham, Maycock
1863 (QUK). MONTCALM CO.: Lac Tapani, Pennell 16678 (NY).
MONTMORENCY CO.: Boischatel, Cauchon, et. al. 69-111 (MU, TRT).
PORTNEUF CO.: Marie-Victorin 9437 (WS). RICHELIEU CO.: Saint-ours,

1974] Genus Kalmia — Ebinger 975

Rolland-Germain 48518 (GH). RIMOUSKI CO.: Cap Enrage, Rousseau
26877 (GH). RIVIERE DU LOUP CO.: Soper 3670 (TRT). SAGUENAY CO.:
Mingan, Townsend (GH); Natashquan, Abbe 1188 (GH); Romaine,
St. John 90652 (GH); Round Lake, Gillett 5208 (GH); Seven Isl.,
Harper 3309 (CAN, HNH, PENN, US), Robinson 769 (CAN, GH, NY).
SHERBROOKE CO.: Orford, Martineau 35 (CAN). TEMISCOUATA CO.:
Trois Pistoles, Fox (TRT). TERREBONNE CO.: St. Sophe, Wahl 19419
(PAC). TIMISKAMING CO.: Ottawa River, Baldwin 5823 (CAN, SASK).
TWO MOUNTAINS CO.: La Trappe, Louis-Marie 10 (GH). WOLFE CO.:
Lake Nicolet, Brisson & Hamel] 11295 (ACAD, OAC, SASK, TRT).

The synonomy of this taxon is fairly extensive. Numerous
varietal and forma names have been proposed for the vari-
ation that exists, and some of these names have been ele-
vated to specifie rank. Many of these lack descriptions or
are listed as synonyms under a variety or form name. Ex-
cept for these subspecific categories only a few other names
have been applied to this taxon. One of these was when
Kuntze (1891) transferred the specific epithet angustifolia
to another genus while the others were proposed by Rafin-
esque (1840).

Most of the subspecific categories that have been pro-
posed for Kalmia angustifolia var. angustifolia are for
variation and extremes in flower color (carnea, rosea,
rubra), variation in size and habit of the plant (media,
minima, nana, procumbens, pumila) and variation in shape
and color of the leaves (lucida, oleifolia, ovata, variegata) .
In general, the differences in flower color appears to repre-
sent natural variation within populations and the above are
better treated as cultivars than as forms. This is also true
of the variation in leaf shape and color, and, in part, for
variation in plant height. Holmes (1956) discusses some
of these cultivars.

The dwarf condition reported for Kalmia angustifolia
var. angustifolia is either the result of selection of more
northern or high altitude sources or because of inbreeding
depression. Jaynes (19682) found that seedlings from
selfed plants average about half the height of seedlings
from outcrossed plants. He also found that seed set from
self-pollinated plants is 85-90% less than that from cross

376 Rhodora [Vol. 76

pollination and that there is a marked reduction in survival
of the seedlings from selfed plants. This indicates that this
species is only weakly self compatible and that outcrossing
usually occurs under natural conditions.

Kalmia angustifolia var. angustifolia commonly occurs
in bogs, swamps, and other wet places, usually forming
dense thickets around ponds and lakes. It is also found in
open woods, as a weed in fields, and in moist openings of
pine savannas. It is distributed from the northeastern part
of the Michigan peninsula and the eastern half of Ontario,
east through Quebec, the Maritime Provinces, and New-
foundland; north to at least the Attawapiskat River (Ken-
ora District) in Ontario and to Goose Bay and Cartwright
in Labrador; and south in eastern United States through
New England and eastern New York, eastern Pennsylvania
and Maryland to the coast in New Jersey and Delaware and
the southeastern tip of Virginia. One specimen (Davis &
Davis, 4691) has been seen which is reputed to have come
from Randolph Co., West Virginia, but this collection should
be considered doubtful. This species has also been reported
from Indiana, but according to Deam (1940) this is an
error.

It is possible that extracts from sheep laurel may inhibit
the growth of other plants, particularly coniferous trees,
thereby maintaining open areas for extended periods of
time. Peterson (1965) found that the leaves of this species
contain a substance that hinders the primary root develop-
ment of black spruce by destruction of the epidermal and
cortical cells. He further suggests that this may be the
reason for the abnormally poor growth of trees associated
with Kalmia angustifolia on upland sites in the boreal
region of eastern Canada.

Studies by Jaynes (personal communication) reveal that
some incompatibility exists between the two varieties of
Kalmia angustifolia. This incompatibility is highly variable,
depending upon the parents used, with the resulting off-
spring varying from completely sterile to highly fertile.
Many of these hybrids produced a large quantity of viable

1974] Genus Kalmia — Ebinger 377

seed while in others the capsules remained small and lacked
seeds. From some crosses, both sterile and fertile individu-
als were produced. Also, among the offspring that pro-
duced no seeds, stainable pollen varied from 64% to 2%.

All artificially produced hybrids between these two taxa
are similar to Kalmia angustifolia var. angustifolia, and
except for the lack of seed set in some, could not be dis-
tinguished from it. All individuals have glandular calyces
and most have leaves that are not puberulent beneath, both
characteristics of typical var. angustifolia. Only 2 hybrids
were found that had leaves which were puberulent beneath.
More studies are needed to clarify the status of these taxa,
but until more information is available, it seems best to
maintain them as varieties.

Kalmia angustifolia L. var. angustifolia f. candida Fern.
Rhodora 15: 151. 1913.

Kalmia angustifolia var. candida (Fern.) Rehd. In:
Bailey, Stand. Cycl. Hort. 3: 1734. 1915. TYPE: Newfound-
land: damp mossy or turfy hollows on hill south of St.
John’s, Fernald & Wiegand 6019 (GH).

According to Jaynes (1971a, 1971b) the presence of pig-
ment in Kalmia angustifolia is determined by a single
dominant gene, and the true-breeding recessive is white
flowered. These white flowered individuals are anthocyan-
inless with green stems, as contrasted to the normal wild
types with reddish stems. This characteristic is particularly
obvious in seedlings.

It is possible that the recessive genes for the anthocyan-
inless condition are farily common in populations of this
species, but are rarely expressed. In a population of Kalmia
angustifolia from Madison, Connecticut, Jaynes (1971b)
found that about 24% of the population was heterozygous
for the anthocyaninless condition, but no homozygous re-
cessive plants were found. It appears that the anthocyanin-
less plants are at a selective disadvantage in the wild.

SPECIMENS EXAMINED
UNITED STATES: Maine: ANDROSCOGGIN CO.: Gordon 407

378 Rhodora [Vol. 76

(MAINE). LINCOLN CO.: Hyland 69 (MAINE). Massachusetts: Sher-

born, Loomis (GH).
CANADA: Newfoundland: St. Johns, Fernald & Wiegand 6019
(GH).

4b. Kalmia angustifolia L. var. caroliniana (Small) Fern.
Rhodora 39: 476, 498. 1937.
Kalmia caroliniana Small, Fl. Southeast. U.S. 1336. 1903.
TYPE: North Carolina: Flat Rock, L. R. Gibbs (NY).
Kalmia carolina Small, Fl. Southeast. U. S. 886. 1903.
Kalmia angustifolia var. carolina (Small) Fern. Rhodora
39: 437. 1937.

Mature leaves permanently puberulent beneath with a
dense covering of whitish, simple hairs, scattered stalked
glandular hairs also present; calyx strongly puberulent,
lacking stalked glandular hairs; corolla usually strongly
puberulent on the outside.

Distribution: Open woods and shrubby bogs in the moun-
tains and sandy woods, pocosins, savannas and bogs on the
coastal plains. Extending from extreme southern Virginia
through North Carolina to eastern South Carolina and west
to eastern Tennessee and northeastern Georgia.

Common name: Sheep Laurel, Lambkill.

REPRESENTATIVE SPECIMENS
Georgia: RABUN C0.: Oakey Mt., Duncan 3281 (GH, ID, ISC, MO,
NY, TENN, US), 4312 (Isc); Tom's Swamp, Bookout & McDowell 65
(GH). North Carolina: ALLEGHANY CO.: Edmonds, Radford 38191
(NCU). AVERY CO.: Pineola Bog, Cain & Duncan 251-3 (IND, ND).
BEAUFORT CO.: Chocowinity, Radford 33396 (NCU). BLADEN CO.:
White Lake, Kral 27210 (VDB). BUNCOMBE CO.: Asheville, McGarthy
(US). CARTERET CO.: Beaufort, Bentley (VDB). CHOWAN CO.: Tyner,
Ahles 39907 (NCU). COLUMBUS C0.: Chadbourn, Wilbur 5315 (DUKE).
CRAVEN C0.: New Bern, Harbison 14080 (NcU), 14932 (NCU).
CUMBERLAND CO.: Stedman, Wilbur 5169 (DUKE). DARE CO.: Manns
Harbor, Radford 8197 (NCU). DUPLIN Co.: Charity, Fogg, Jr. 5481
(GH, PENN). GATES CO.: Gatesville, Godfrey 5214 (DUKE, GH).
GREENE CO.: Walstonburg, Radford 36508 (NCU). HARNETT CO.:
Blomquist 4486 (DUKE). HENDERSON CO.: Muddy Creek, Smith 221
(180, US). HOKE CO.: Timberland, Ahles 29514 (IND). HYDE CO.:
Radford & Stewart 803 (NCU). JACKSON CO.: Cullowhee, Biltmore

1974] Genus Kalmia — Ebinger 379

Herb. 1344e (ALTA, GH, US). JOHNSTON CO.: Benson, Smith (IND,
NCU). JONES CO.: Hargetts Store, Radford 36956 (NCU). LENOIR
co.: Pine Hill, Radford 31651 (NCU). MACON co.: Highlands,
Wilson 1854 (TENN). MARTIN CO.: Robersonville, Radford 35364
(Isc). MONTGOMERY CO.: Pekin, Radford 14595 (NCU). NEW HAN-
OVER CO.: Wilmington, Newber 165 (UARK). ONSLOW co.: Dixon,
Moldenke 1242 (DUKE, MIN, MO, NY, PENN, US). PASQUOTANK CO.:
Elizabeth City, Ashe (NCU). PENDER CO.: Wards Corner, Ahles &
Bell 7856 (MISSA, MSC). PERQUIMANS CO.: Beach Spring, Radford
5470 (NCU). PITT CO.: Staton, Radford 34940 (NCU). RICHMOND
co.: Rockingham, Coker (NCU). ROBESON CO.: Allenton, Ahles &
Ramseur 23770 (ND, PAC). SAMPSON CO.: Tomahawk, Fox & Boyce
3792 (KY). SCOTLAND CO.: Wagram, Correll 1181 (DUKE). TRAN-
SYLVANIA CO.: Cedar Mt., Freeman 58130 (NCU). TYRRELL CO.:
Cross Landing, Radford 33820 (NCU). WASHINGTON CO.: Hoke,
Radford 35264 (NCU). WATAUGA CO.: Deep Gap, Radford 6239 (NCU).
WAYNE CO.: Goldsboro, Radford 22006 (NCU). South Carolina:
DARLINGTON CO.: between Darlington & Society Hill, Smith 1068
(NCU). HORRY CO.: Radford & Stewart 979 (NCU, NY). ORANGEBURG
co.: Edisto Swamp, Ravenel (GH). SUMTER CO.: Poinsette St. Park.
Holdaway (DUKE). Tennessee: JOHNSON CO.: Cole’s Bog, Shanks
& Sharp 1115 (PENN, TENN); Shady Valley, Barclay (TENN), Sharp
& Underwood 1252 (vps), 1552 (NCU, NY), 3161 (NY). KNOX CO.:
Kingston Pike, Jennison 717 (TENN). Virginia: CARROL CO.: Wood-
lawn, Massey 5058 (GH). NANSEMOND CO.: Whaleyville, Fernald &
Long 7564 (GH, NY, US). NORFOLK CO.: Dismal Swamp, Coville &
Kearney, Jr. 91 (08, US), Portsmouth Ditch, Grimes 4517 (MIN, NY).
SOUTHAMPTON CO.: Franklin, Fernald & Long 10002 (GH).

The name first applied to this taxon was Kalmia carolina
when Small (1903) separated the plants with puberulent
leaves from typical K. angustifolia. Later in a list of genera
and species in the same publication (page 1336) the name
was changed to K. caroliniana. It is very possible that this
later name is a printing error, particularly since the name
K. carolina was used by the author in a later publication
(Small, 1914). Fernald (1937) transferred the epithet
carolina to a variety of K. angustifolia (page 437), but later
in the same publication (page 476, 498) changed the name
to variety caroliniana, stating that this spelling, “since it
accompanied the citation of the type, should prevail.”

Kalmia angustifolia var. caroliniana has a fairly re-
stricted range. Most of the specimens examined are from

380 Rhodora [Vol. 76

the coastal plain and the mountains of North Carolina with
the species being extremely common on the pocosins and in
bogs. This taxon also has a sporadic occurrence in the
coastal plain of South Carolina (Radford, Ahles, and Bell,
1968) and in southern Virginia (Fernald, 1938, 1939),
while it has been reported from a few localities in the moun-
tains of extreme eastern Tennessee by Jennison (1935) and
Shanks (1952) and from two mountain bogs in the Blue
Ridge of northeastern Georgia by Duncan (1948).

The characteristics used to separate Kalmia angustifolia
var. caroiimiana are very reliable, but occasionally some
overlap does occur. Usually, var. angustifolia has glandular
calyces and leaves that are not puberulent beneath while in
var. caroliniana the calyces are eglandular and the leaves
are strongly puberulent beneath. All but two specimens of
var. caroliniana examined had strongly puberulent leaves
and even those were lightly puberulent on the undersurface
of the leaf. This was also noticed by Radford, Ahles, and
Bell (1968) who mentioned that a few specimens of var.
caroliniana with glabrate leaves were found in the Caro-
linas. A few specimens of var. angustifolia from Nantucket
Island and the coast of Massachusetts and Connecticut have
leaves that are strongly puberulent beneath. All of these,
however, have strongly glandular calyces, typical of var.
angustifolia, and are referred to that taxon. The best char-
acter to separate the two varieties is the presence or absence
of stalked glandular hairs on the calyx. Close examination,
however, reveals that on some specimens of var. caroliniana
a few glandular hairs are present on the calyces. The glands
are not very numerous, however, and mostly restricted to
the margins of the lobes or to the base of the calyx. Even
on the holotype a few of these stalked glandular hairs are
found. Because of this variability it is probably best to
maintain these taxa as varieties of one species rather than
as separate species.

A white flowered, anthocyaninless form of var. carolin-
iana also exists. It is similar genetically to Kalmia angusti-
folia var. angustifolia f. candida in that it is a homozygous

1974] Genus Kalmia — Ebinger 381

recessive plant in which flower color is determined by a
single pair of genes (Jaynes, 1971b). A specimen of this
form was obtained by Dr. Jaynes from the Garden-in-the-
Woods, in Sudbury, Massachusetts. No wild individuals
have been found and its origin is not known. The best
crossing results obtained by Jaynes (1968a) between K.
angustifolia and other species of the genus were when this
white flowered form was used as the female parent (Jaynes,
1971b).

5. Kalmia cuneata Michaux, Fl. Bor.-Amer. 1: 257. 1803.
Chamaedaphne cuneata (Michx.) Kuntze, Rev. Gen. Pl.
2: 388. 1891. TYPE: In Carolina, Michaux s.n., (GY).

Much branched rhizomatous shrub to 1.5 m tall; branch-
lets terete, reddish brown, usually adscending, stipitate-
glandular and puberulent, becoming glabrous and light
gray; leaves alternate, deciduous, flat, oblanceolate to ellip-
tic-lanceolate, 2-6 cm long, 0.5-1.5 cm wide, lightly stipitate
glandular beneath, glabrous above, apex obtuse to acute,
apiculate, base narrowly cuneate, midrib puberulent above,
stipitate-glandular beneath; petiole to 4 mm long; inflor-
escence a reduced corymbiform raceme or fascicle of 3-10
flowers in the upper axils of the last season’s growth, pedi-
cels 1-3 mm long, glabrous to lightly stipitate-glandular,
recurved in fruit, subtended by 2 bracteoles in the axil of a
small lanceolate bract about 1 mm long, main axis of the
inflorescence less than 3 mm long; calyx glabrous to lightly
stipitate-glandular, 6-7.5 mm broad, the lobes oblong, apex
acute; corolla creamy white with a red band within, lightly
stipitate-glandular outside, puberulent near the base within,
1.3-1.8 mm broad; anthers purplish brown, 1-1.3 mm long;
filaments puberulent toward the base; style 7-10 mm long;
ovary stipitate-glandular; capsule depressed globose, 4-6
mm broad, 2.5-3.5 mm long, stipitate-glandular; seeds light
brown, 0.6-0.9 mm long, the testa not loose or extended past
the ends of the seed. Chromosome number n — 12.

Distribution: Wet thickets and shrub bogs in southeastern

382 Rhodora [Vol. 76

North Carolina and adjacent South Carolina.
Common name: White Wicky.

REPRESENTATIVE SPECIMENS

North Carolina: BLADEN CO.: Colly Creek, Ashe (PH); 10 miles
north of Elizabethtown, Ebinger 11271 (EIU); Jerome Pocosin, Buell
& West 1740 (MIN). CUMBERLAND C0.: Hope Mill, Alexander (NY).
MOORE CO.: Pinehurst, Wicker (NY). PENDER CO.: Squalling Bluff
Rd., Coville (MICH, NY, US); Burgaw, Chrysler 2262 (CHRB). RICH-
MOND CO.: Hamlet, Wiegand & Manning 2365 (GH). South Carolina:
DARLINGTON CO.: Hartsville, Smith (NY), Coker (NY).

A small fragment of the type of Kalmia cuneata is at the
Gray Herbarium. It consists of one fascicle of 5 flowers and
a small stem with a few leaves attached. On the label is
written “Ex Herb. Michaux Kalmia cuneata." At the pres-
ent time the rest of the type has not been located.

Kalmia cuneata is very distinct. The only species that it
could possibly be confused with is K. angustifolia since
their general habit and leaf size are similar. Kalmia cune-
ata, however, is the only deciduous member of the genus
and the only one in which the flowers are borne in fascicles
from the axils of last season’s growth. Other distinguishing
characteristics are the creamy-white flowers with a red
band within, the oblanceolate leaves, and the recurved fruit-
ing pedicels.

Genetically, this species appears to be most closely re-
lated to Kalmia hirsuta, another alternate leaved species of
the southeastern United States. Jaynes (1968a) reported
that K. cuneata set no seed when used as the female parent,
but as the male parent crossed with both K. angustifolia
and K. hirsuta. The cross with K. angustifolia produced
only inviable hybrids. Most of the hybrids with K. hirsuta
were also inviable, producing sectors of albino or yellow-
green tissue, but one hybrid flowered and was pollen sterile.

This species is one of the rarest shrubs in North America
and few early reports exist concerning its distribution.
Pursh (1814) reports that it occurs on the mountains of
Carolina, while Nuttall collected it between Camden, South
Carolina and Statesville, North Carolina (Wood, 1961).

1974] Genus Kalmia — Ebinger 383

Curtis (1835) reported its occurrence about 30 miles NE
of Wilmington, North Carolina. Present studies indicate
that it grows only in the sandhills and bay margins of the
pocosin ecotones of the Carolinas. It now is known from
8 counties in this region (Radford, Ahles & Bell, 1968).

6. Kalmia hirsuta Walt. Fl. Carol. 138. 1788.
Chamaedaphne hirsuta (Walt.) Kuntze, Rev. Gen. Pl. 2:
388. 1891. Kalmiella hirsuta (Walt.) Small, Fl. Southeast.
U.S. 886. 1903. TYPE: unknown.
Kalmia ciliata Bartram, Trav. N. & S. Carol. v, 7, 94, 171,
242,1791. (nom. nud.).

Low, sparingly branched shrub, to 6 dm tall with a thick
basal caudex; branchlets terete, reddish brown, puberulent
and densely hispid; leaves alternate, sessile or with a petiole
to 1 mm long, elliptic to ovate, 5-14 mm long, 1.5-8 mm
broad, the margins usually slightly revolute, usually pu-
berulent and hispid on both surfaces, small stipitate glands
usually present beneath, apex acute, base broadly cuneate
to rounded; flowers solitary (rarely in fascicles of 2-5 or
in compact racemes) in the axils of the leaves of new
growth; pedicels to 1.5 cm long, stipitate-glandular and
hispid, subtended by a pair of bracts; calyx green, the lobes
foliaceous, lanceolate, 3-8 mm long, hispid, tardily deciduous
in fruit; corolla pink (rarely white), marked with red
around the anther pockets and a red ring near the base,
1-1.5 em broad, usually lightly hirsute and stipitate-glandu-
lar on the keels outside, puberulent at the base within;
anthers purplish brown, 0.7-1.3 mm long; filaments puberu-
lent toward the base; ovary usually stipitate glandular;
style 5-7 mm long; fruit 2-4 mm broad, 2-3.5 mm long,
subglobose to ovoid, thinly stipitate-glandular; seeds light
brown, 0.3-0.5 mm long, the testa not loose or extended past
the ends of the seed. Chromosome number n = 12.

Distribution: Low, sandy pine savanna, sand hills, dunes
and flat pine woods on the coastal plains of southern Ala-

384 Rhodora [Vol. 76

bama and Georgia, northern Florida and the southern tip
of South Carolina.
Common name: Sandhill Laurel, Calico-bush.

REPRESENTATIVE SPECIMENS
Alabama: BALDWIN CO.: Gulf State Park, Webster & Wilbur 3539
(GH, NY, US). MOBILE C0.: Theodore, Pennell 4513 (MICH, NY, PENN,
PH). Florida: ALACHUA CO.: Cross Creek, Ford 2250 (GH, TENN).
BAY CO.: Lynn Haven, Billington (MicH, us); Lazor 5006 (Pac).
CLAY CO.: Kingsley Lake, Henry 3828 (PH). COLUMBIA CO.: Lake
City, Nash 2193 (GH, MASS, MICH, MIN, MSC, ND, NY, OS, PH, US).
DIXIE C0.: Old Town, West & Arnold (PH). DUVAL CO.: Jacksonville,
Curtis 4359 (ARIZ, ND, NY, US), 4921 (GH, ISC, MIN, MSC, NY, US).
ESCAMBIA CO.: Pensacola, Webster & Wilbur 3572 (GH, NY, US).
FLAGLER CO.: Bunnell, Ames (GH). FRANKLIN CO.: Carrabelle, Mosier
884 (MIN, NY, PENN, US); Moldenke 1134 (MIN, NY, PENN, US).
GADSDEN CO.: Quincy, Sargent (MIN). GILCHRIST CO.: Trenton, Ward,
West & Will 2651 (GH, US). GULF CO.: Port St. Joe, Ward & Ward
1460 (GH). HAMILTON CO.: Bowers & Wofford 71-550 (TENN).
HOLMES CO.: Bonifay, Blanton 6582 (US). JACKSON CO.: Compass
Lake, Knight & Arnold (PAC). JEFFERSON Co.: Wacissa, Godfrey
53552 (GH). LAFAYETTE CO.: Smith & Myint 189 (GH). LEON CO.:
Silver Lake, Godfrey 53660 (GH, MISSA, NY). LEVY CO.: Bronson,
Godfrey & Lindsey 56989 (GH). LIBERTY CO.: Bristol, Small, Small
& DeWinkeler 11448 (Gn, NY, PENN, TENN). MADISON CO.: Green-
ville, Ramsey 106 (MSC). MARION CO.: Ocala, Beardslee, Jr. (MU).
OKALOOSA CO.: Crestview, McArthur (GH). PUTNAM CO.: Interlachen,
Fox 5703 (GH, MICH, MIN, NY, TENN). ST. JOHNS CO.: Anastasia
Island, Crawford (NY, PH). ST. LUCIE CO.: Fort Walton, Henry
1087 (PH). SANTA ROSA CO.: Munson, Ford & Arnold 4190 (us).
WAKULLA CO.: St. Marks Wildlife Refuge, Godfrey 57860 (GH).
WALTON CO.: Santa Rosa, Ripley & Barneby 10128 (NY). Georgia:
APPLING CO.: Baxley, Cottam 11060 (UT); Ebinger 11204 (E10).
BRANTLEY CO.: Waynesville, Ebinger 11232 (EIU). BULLOCH CO.:
Portal, Henry 7311 (PH). CAMDEN CO.: Waverly, Ebinger 11228
(EIU). CANDLER CO.: Metter, Henry 7277 (PH); Hermann 10122
(GH, US). CHARLTON CO.: Folkston, Harbison 5816 (GH). COFFEE
co.: Harper 687 (GH, US). EFFINGHAM CO.: Harper 961 (GH).
GLYNN C0.: Emanuel Church Rd., Ebinger 11231 (EIU). JEFF DAVIS
cc.: Hazlehurst, Ebinger 11195 (EIU). LANIER CO0.: Adams 50 (Ky).
LIBERTY CO.: Altamaha River, Small (GH, MIN, NY, PH). LOWNDES
co.: Valdosta, Beardslee, Jr. (MU). MCINTOSH CO.: Darien, Cron-
quist 5366 (GH, MICH, NY, PH). TATTNALL CO.: Reidsville. Cronquist
5840 (GH, IND, MICH, NY, PH, US). THOMAS CO.: Thomasville, Taylor
(GH, MIN): TIFT CO.: Tifton, Hardin & Duncan 13649 (US). WARE

1974] Genus Kalmia — Ebinger 385

co.: Suwannee Lake, Harper 48 (PH), 298 (GH, NY, PH, US). WAYNE
co:. Grangerville, Ebinger 11233 (EIU). WORTH CO.: Poulan, Pollard
& Maxon 563 (NY, Us). South Carolina: BEAUFORT CO.: Ferry
Landing, Bell 3805 (NY). HAMPTON CO.: Hampton, Leonard & Rad-
ford 1662 (ARIZ, BRY, CHRB, NHA). JASPER CO.: Ridgeland, Ebinger
11242 (EIU).

Except for transfers of the specific epithet, hirsuta, to
other genera, only one other name has been proposed for
this taxon. Bartram (1791) used Kalmia ciliata for a plant
he observed on his travels through the southeastern United
States. He used this name a number of times but did not
include a description (Merrill, 1945). He did, however,
describe “a diminutive but elegant Kalmia" (page 18)
which undoubtedly refers to K. ciliata. This description
leaves no doubt that his K. ciliata is the same as K. hirsuta.
The first time the name K. ciliata appears with a descrip-
tion is in the work of Dumont de Courset (1811) who
attributes the name to Bartram.

Kalmia hirsuta has a relatively limited distribution, oc-
curring along the coastal plain in southeastern United
States. Small (1903, 1914) indicated that it extended from
Virginia to Florida, perpetuating an error started by Gray
(1856). Its occurrence in Virginia is extremely doubtful
(Fernald, 1947) and is probably due to a few herbarium
specimens with incomplete or erroneous collecting data.
Two such specimens have been found during the present
study. Both were collected during the middle of the last
century and beside the collector’s name and the year col-
lected include the name Virginia.

Most of the specimens of Kalmia hirsuta examined are
hispid on the stems, leaves, and pedicels. A few specimens
were found, however, that were completely glabrous, lack-
ing the long coarse hairs as well as the stipitate-glands and
smaller hairs commonly found on most parts of the plant.
Variation also occurs in the arrangement of the flowers on
the plant. Usually the flowers are solitary in the axils of
the leaves, but occasionally the flowers occur in fascicles of
2-5, while a few specimens were found in which short

386 Rhodora [Vol. 76

racemes occur in the axils of the leaves. In other specimens
the flowers appeared as a terminal raceme due to a shorten-
ing of the internodes and a reduction in the size of the sub-
tending leaves. The pedicels also vary extensively in length.
The flowers of some plants are nearly sessile, while in others
the pedicels are 1.5 cm long with the two basal bracts
scattered along the stalk.

Genetically Kalmia hirsuta appears to be closely related
to K. latifolia with which it can be crossed to produce viable
hybrids with 0-10% pollen stainability. Jaynes (1968a) also
reported that it can be crossed with both K. angustifolia
and K. cuneata producing viable but sterile hybrids. Based
on habit and morphology K. hirsuta is most closely related
to K. ericoides. These species were placed in the genus
Kalmiella by Small (1914) since they have a deciduous
calyx and their flowers are borne singly in the axils of the
leaves. Unfortunately the Cuban member of the genus is
not available for genetic studies.

7. Kalmia ericoides Wright ex Griseb. Cat. Pl. Cubensium
51. 1866.

Chamaedaphne ericoides (Wright ex Griseb.) Kuntze,
Rev. Gen. Pl. 2: 388. 1891. Kalmiella ericoides (Wright ex
Griseb.) Small, North Amer. Fl. 29: 54. 1914. TYPE: Cuba
occ. in distr. Guane, in pinetis arenosis pr. La Grifa, C.
Wright 2199 (GH, NY, US - Isotypes).

A sparingly branched, erect to spreading shrub, to 10 dm
tall with a thick basal caudex; branchlets light gray to
reddish-gray, puberulent and commonly densely hispid and
stipitate-glandular (rarely glabrous) ; leaves alternate, per-
sistant, thick, leathery, subsessile or with a puberulent
petiole to 1 mm long, linear, 4-13 mm long, 0.6-2 mm broad,
the margins strongly revolute, usually lightly hispid and
stipitate-glandular on both surfaces, sometimes strongly
puberulent (rarely glabrous); flowers solitary (rarely in
fascicles or in compact racemes) in the axils of the leaves
at the end of the branches, forming a terminal cluster;
pedicels to 13 mm long, commonly hispid and stipitate-

1974] Genus Kalmia — Ebinger 387

glandular, subtended by a pair of puberulent bracts 1-3 mm
long; calyx green, the lobes foliaceous, lanceolate, 3-6 mm
long, usually hispid and stipitate-glandular, tardily decidu-
ous in fruit; corolla light pink to white, 10-17 mm broad,
usually lightly hirsute and stipitate-glandular on the keels
outside, lightly puberulent at the base within; anthers pur-
plish brown, 0.8-1.2 mm long; filaments puberulent toward
the base; ovary usually stipitate-glandular; style 4-7 mm
long; fruit 2-4 mm broad, 2-3 mm long, subglobose to ovoid,
lightly stipitate-glandular ; seeds reddish-brown, 0.4-0.7 mm
long, the testa not loose or extending past the ends of the
seed.

Kalmia ericoides is the only member of the genus that
has not been studied cytologically. Morphologically, how-
ever, it is most closely related to K. hirsuta, a species re-
stricted to southeastern United States. Both have relatively
small leaves (less than 1.5 em long), that are usually hispid,
their flowers are borne singly in the axils of the leaves, and
the calyx is tardily deciduous. The two are easily separated,
however, since the leaves of K. ericoides are thick and
leathery with revolute margins while in K. hirsuta the
leaves are thin and lack a strong revolute margin. Also,
the flowers are borne scattered along the stem in the axils
of the leaves in K. hirsuta while in K. ericoides, they are
solitary in the leaf axils, but clustered toward the ends of
the stems giving the appearance of a terminal corymbiform
raceme.

Kalmia ericoides is endemic to savannas and pine barrens
of western Cuba. Although it has a very limited distribu-
tion, the variation that exists in pubescence and compact-
ness of the inflorescence has resulted in a division of this
complex into three species by Alain (1957), while others
considered it a single highly variable species (Wood, 1961).
Present studies indicate that the compactness of the inflor-
escence is not completely reliable in separating the taxa.
The variation in leaf pubescence, however, is relatively
stable. As a result, the following two varieties are recog-
nized.

888 Rhodora [Vol. 76

KEY TO VARIETIES

1. Leaves glabrous or lightly hispid and with a few scat-
tered stalked glandular hairs on the upper surface. ....
rr rr Ta. var. ericoides.

1. Leaves densely puberulent and stipitate-glandular on
the upper surface. .............. 7b. var. aggregata.

Ta. Kalmia ericoides Wright ex. Griseb. var. ericoides.

Kalmiella simulata Britt. & Wils. Mem. Torrey Bot. Club
16: 93. 1920. TYPE: Cuba: vicinity of Los Indios, Isle of
Pines, Britton, Britton, & Wilson 14205 (NY).

Shrub with ascending, hispid, stipitate-glandular and
sometimes puberulent branchlets; leaves scattered, usually
only 5-12 per cm of branchlet, glabrous to lightly hispid and
stipitate-glandular above, not puberulent, 4-9 mm long,
0.6-1.5 mm wide; pedicels usually longer than the subtend-
ing leaves, giving the inflorescence an open appearance;
calyx lobes 3-4.5 mm long, hispid and lightly puberulent on
the outside, usually glabrous within.

Distribution: Low, white-sand savanna and pine barrens
on the Isle of Pines and in the Province of Pinar del Rio,
Cuba.

REPRESENTATIVE SPECIMENS

Cuba: PINAR DEL RIO: La Grifa, Ekman 18165 (NY, Us), Wright
2199 (GH, NY, US); Laguna de Alcatraz Grande, León & Victorin
18706 (GH, US), 20474 (GH, Us); El Sabalo, Alain 1326 (GH, US).
ISLE OF PINES: Los Indios, Britton Britton, & Wilson 14205 (GH,
NY), Killip 42882 (GH, NY, US), León & Victorin 17853 (GH), Vic-
torin & Alain TTa (US).

Kalmia ericoides var. ericoides has a very restricted range
and few herbarium specimens are available. It appears to
be a fairly stable variety that varies mostly in the extent of
leaf pubescence. All of the specimens seen from the prov-
ince of Pinar del Rio are typical of the type specimen which
has a few scattered glandular hairs and is lightly hispid on
the upper leaf surface. In contrast, most of the specimens
of this variety from the Isle of Pines are glabrous on the

1974] Genus Kalmia — Ebinger 389

upper leaf surface. These glabrous specimens were sepa-
rated as Kalmiella simulata by Britton and Wilson (Britton,
1920). These glabrous specimens all have small, narrow
leaves (less than 9 mm long and 0.6-1.2 mm wide), pedicels
that are much longer than the subtending leaves, and calyx
lobes that are fairly short (less than 4.5 mm long) all char-
acteristics of variety ericoides.

7b. Kalmia ericoides var. aggregata (Small) Ebinger,
comb. nov.
Kalmiella aggregata Small, North Amer. FI. 29: 54. 1914.
Kalmia aggregata (Small) Copeland, Amer. Midl. Nat. 30:
571. 1943. TYPE: Cuba: pinelands, Los Indios, Isle of Pines,

Jennings 324 (NY).

Shrub with adscending, lightly hispid and strongly pu-
berulent branchlets; leaves crowded, usually 15-25 per cm
of branchlet, densely puberulent and stipitate-glandular
above, 8-13 mm long, 1-2 mm wide; pedicels usually as short
as the subtending leaves, giving the inflorescence a compact
appearance; calyx lobes 3.5-5.5 mm long, hispid and densely
puberulent on the outside, densely puberulent within.

Distribution: Low, white-sand savanna and pine barrens
on the Isle of Pines, Cuba.

REPRESENTATIVE SPECIMENS

Cuba: ISLE OF PINES: Beach at Siquanea, Morton 10098 (us);
between Mina de Oro and Playa del Soldado, Killip 43905 (us),
45385 (GH); Lama Daquille, Ekman 72492 (NY); Los Indios, Jen-
nings 324 (GH, NY, US), León & Victorin 18857 (GH, US), Victorin
& Alain 77 (GH, NY, US), Playa Roja, Killip 48001 (GH, NY, US);
San Pedro, Britton, Britton, & Wilson 14146 (GH, NY, US), Killip
43698 (US), 45566 (vs), Morton 10018 (Us), 10053 (US); Siquanea
region, Killip 44043 (us), 44044 (us), 44753 (us), 44755 (US).

This variety appears to be more robust than the preced-
ing. Specimens to 13 dm have been reported, and the leaves
and flowers are, in general, slightly larger than in variety
ericoides. Besides this difference in size, all of the speci-
mens of variety aggregata examined are strongly puberu-
lent throughout. In particular the leaves are strongly

390 Rhodora [Vol. 76

puberulent on both surfaces, but this is also characteristic
of the branches, pedicels and sepals.

UNCERTAIN OR EXCLUDED NAMES

Kalmia lanceolata Raf. Autikon Bot. 86. 1840. This
name should probably be referred to Kalmia polifolia but
the description could apply to other taxa. No specimen to
represent this name has been found.

Kalmia serotina Hoffmgg. Verz. Pflanzenkulturen 70.
1824. Nomen nudum.

Kalmia spuria Bartr. Trav. N. & S. Carol. 308. 1791.
Nomen nudum.

LITERATURE CITED

ABRAMS, L. 1951. Illustrated Flora of the Pacific States: Wash-
ington, Oregon, and California, Vol. 3. Stanford.

AITON, W. 1789. Hortus Kewensis. Vol. 2. London.

ALAIN, BR. H. 1957. Ericaceae. In: LEON, BR. H. & BR. H. ALAIN,
Flora de Cuba. Vol. 4. Havana.

AMERICAN FORESTRY ASSOCIATION. 1955. These are the Champs.
Am. Forests 61(9): 31-40.

ANONYMUs. 1895. New or little-known plants. Garden Forest 8:
315.

ASHLEE, T. R. 1932. A contribution to the Latah flora of Idaho.

Northw. Sci. 6: 69-82.

BARTON, B. S. 1802. Some account of the poisonous and injurious

honey of North America. Trans. Am. Phil. Soc. 5: 51-70.
1804. Collections for an essay toward a materia medica

of the United States. Philadelphia.

BaRTRAM, W. 1791. Travels through North and South Carolina,
Georgia, East and West Florida. Philadelphia.

BEAL, W. J. 1867. Agency of insects in fertilizing plants. Am.
Nat. 1: 254-260.

BEAN, W. J. 1897. Kalmias. Garden 52: 77-78.

BEISSNER, L., E. SCHELLE, & H. ZABEL. 1903. Handbuch der
Laubholz-Benennung. Berlin.

Benson, A. B. 1937. Peter Kalm’s travels in North America. New
York.

Berry, E. W. 1907. Contributions to the Mesozoic flora of the
Atlantie coastal plain. II. North Carolina. Bull. Torrey Bot.
Club 34: 185-206.

1974] Genus Kalmia — Ebinger 391

1913. Contributions to the Mesozoic flora of the At-
lantie coastal plain. IX. Alabama. Bull. Torrey Bot. Club
40: 567-574.

1919. Upper Cretaceous floras of the eastern gulf
region in Tennessee, Mississippi, Alabama, and Georgia. U. S.
Geol. Surv. Prof. Pap. 112: 1-177.

1920. Contributions to the Mesozoic flora of the At-
lantic coastal plain. XII. North Carolina. Bull. Torrey Bot.
Club 47: 397-406. |

, & W. A. JouNSTON. 1922. Pleistocene interglacial
deposits in the Vancouver region, British Columbia. Proc. Trans.
Roy. Soc, Canada 16(4): 133-140.

BrGELOW, J. 1818. American medical botany. 1(II): 133-141;
3(II): 185-186. Boston.

BorviN, B. 1967. Connecting vowels in epithets of Latin origin.
Rhodora 69: 451-455.

1968. Flora of the prairie provinces. A handbook to
the flora of the provinces of Manitoba, Saskatchewan and
Alberta. Phytologia 16: 1-47.

BRITTON, E. G. 1913. Wild plants needing protection. Jour. N. Y.
Bot. Gard. 14: 121-123.

1925. A freak of the mountain laurel. Ibid. 26: 187-
190.

Britton, N. L. 1908. North American trees. New York.

1920. Description of Cuban plants new to science.
Mem. Torrey Bot. Club 26: 57-118.

BRowN, R. W. 1962. Paleocene flora of the Rocky Mountains and
Great Plains. U. S. Geol. Surv. Prof. Pap. 375: iv + 119 + 69.

BROWNE, D. J. 1857. The trees of America. New York.

BRUNET, L. O. 1867. Catalogue des végétaux ligneux du Canada.
Quebec.
BUTTRICK, P. L. 1924. Connecticut's state flower, the mountain
laurel, a forest plant. Marsh Bot. Gard. Publ. 1: 1-28.
CALLAN, H. G. 1941. The cytology of Gaulthettya wisleyensis
(Marchant) Rehder. A new mode of species formation. Ann.
Bot. II. 5: 579-585.

Cary, C. A, & F. G. MATTHEWS. 1903. Some diseases of cattle.
Alabama Agr. Exp. Sta. Bull. 125: 105-168.

CaTEsBY, M. 1743. The natural history of Carolina, Florida and
the Bahama Islands. London.

CHENON, L. J. 1751. Dissertation Botanica, qua Nova Plantarum
Genera. Upsaliae.

CLAWSON, A. B. 1933. Alpine Kalmia (Kalmia microphylla) as a
stock-poisoning plant. U. S. Dept. Agr. Tech. Bull. 391: 1-9.

392 Rhodora [Vol. 76

COLVIN, W. S., & W. S. EISENMENGER. 1943. Relationships of natu-
ral vegetation to the water holding capacity of the soils in New
England. Soil Sci. 55: 433-446.

CoPELAND, H. F. 1943. A study, anatomical and taxonomic, of the
genera of Rhododendroideae. Am. Midl. Nat. 30: 533-625.

Cox, H. T. 1948. Studies in the comparative anatomy of the
Ericales. I. Ericaceae — Subfamily Rhododendroideae. Am.
Midl. Nat. 39: 220-245.

CRAWFORD, A. C. 1908. Mountain laurel, a poisonous plant. U. S.
Dept. Agr. Bur. Pl. Ind. Bull. 121: 21-35.

Curtis, M. A. 1835. Enumeraticn of plants growing spontaneously
around Wilmington, North Carolina, with remarks on some new
and obscure species. Boston Jour. Nat. Hist. 1: 82-141.

CurTis, W. 1795. Kalmia angustifolia. Narrow-leaved Kalmia. Bot.
Mag. 9: 331.

Davis, P. H., & V. H. HEvwoop. 1963. Principles of Angiosperm
taxonomy. Princeton.

DEAM, C. C. 1940. Flora of Indiana. Indianapolis.

DEANE, W. 1899. Kalmia latifolia in Vermont. Rhodora 1: 136.

DEGRAFFE, B. L. 1896. The tannins of some Ericaceae. Am. Jour.
Pharm. 68: 313-321.

DRUDE, O. 1889. Ericaceae. In: ENGLER, A., & K. PRANTL, Die
naturlichen Pflanzenfamilien. IV(1): 15-65.

DuprEY, T. R. 1967. Ornamental mountain-laurel and a new cul-
tivar: Kalmia latifolia ‘Bettina’. Am. Hort. Mag. 46: 245-248.

DUMONT DE CoursET, G. L. M. 1802. Le botaniste cultivateur.
Vol. 2. Paris; 1811. Ed. 2, Vol. 3. Paris.

Duncan. W. H. 1948. Preliminary reports on the flora of Georgia.
I. The distribution in Georgia of Spermatophytes new to or
rare in the state. Castanea 13: 70-83.

EGLER, F. E. 1947. Effects of 2,4-D on woody plants in Connecti-
cut. Jour. Forest. 45: 449-452.

—. 1948. 2.4-D effects in Connecticut vegetation, 1947.

Ecology 29: 382-386.
1949. Herbicide effects in Connecticut vegetation, 1948.

Ecology 30: 248-256.
. 1950. Herbicide effects in Connecticut vegetation, 1949.

Bot. Gaz. 112: 76-85.

ENGELMANN. G.. W. M. CANBY. & A. GRAY. 1877. Large trunks of
Kalmia latifol'a. Amer. Nat. 11: 175.

FERN^'Dp. M. L. 1911. Notes from the Phaenogamic Herbarium of
the New England Botanical Club. II. Rhodora 13: 177-183.

1937. Local plants of the inner coastal plain of south-

eastern Virginia. Rhodora 39: 321-366, 379-415, 433-459, 465-491.

. 1938. Noteworthy plants of southeastern Virginia.
Rhodora 40: 434-459.

1974] Genus Kalmia — Ebinger 393

1939. Last survivors in the flora of tidewater Vir-
ginia. Rhodora 41: 529-559.
1940. A new southern Kalmia. Rhodora 42: 53-54.

— — — —— 1947. Additions to and subtractions from the flora of
Virginia. Rhodora 49: 145-159.

1950. Gray's Manual of Botany. 8th ed. New York.

FLEMING, C. E. 1920. Project 22 — Poisonous range plants. Nevada
Agr. Exp. Sta. Ann. Rep. 1920: 39-43.

FLEMER, W. 1949. The propagation of Kalmia latifolia from seed.
Bull. Torrey Bot. Club 76: 12-16.

Forges, E. B. & S. I. BECHDEL. 1930. The chemical composition
and feeding value of Laurel and Rhododendron, and the condi-
tions of browsing in forests overstocked with deer, in relation
to the death of young deer. Penn. Agr. Exp. Sta. Bull. 258:
39-41.

& .1931. Mountain Laurel and Rhododendron
as food for the white tailed deer. Ecology 12: 323-333.

FULLER, G. D. 1944. Notes on some plants that are rare in Illinois.
Trans. Ill. St. Acad. Sci. 37: 57-58.

GLEASON, H. A. 1952. The new Britton and Brown Illustrated
Flora of the Northeastern United States and adjacent Canada.
Vol. 3. Lancaster.

GRAY, A. 1856. Manual of the Botany of the Northeastern United
States. 2nd ed. New York.

1871. Dialysis with staminody in Kalmia latifolia.
Amer. Nat. 1: 373-374.

1876. How flowers are fertilized. Amer. Agr. 35: 262.

1879. Gray’s Botanical Text-book. 6th ed. Vol. 1.
Structural Botany. New York.

GRIFFITH, R. E. 1847. Medical Botany. Philadelphia.

Griscom, L. 1931. Dwarf mistletoe and other plants new to New
Jersey. Rhodora 33: 101.

Gronovius. J. F. 1739. Flora Virginica. Leiden.

Gussow. W. C. 1933. Contributions to the knowledge of the flora
of Northern Manitoba and the North-western Territories, Do-
minion of Canada (Lat. 58°-62°; Long. 95°-100°. Hudson Bay).
Canad. Field-Nat. 47: 116-119.

HAGERUP, O. 1928. Morphological and cytological studies of Bi-
cornes. Dansk Bot. Ark. 6: 1-26.

HALL, H. M. 1912. New and noteworthy California plants, I.
Univ. Calif. Publ. Bot. 4: 195-208.

HARDIN. J. W., & J. M. ARENA. 1969. Human poisoning from native
and cultivated plants. Durham.

HASSKARL, J. C. 1863. Ueber Kalmia latifolia L. Bot. Zeit. 31:
237-239.

394 Rhodora [Vol. 76

Hopepon, A. R. 1955. Ilex glabra and a new station for Kalmia
latifolia in New Hampshire. Rhodora 57: 34-36.

HOLLICK, A. 1892. Additions to the Paleobotany of the Cretaceous
formation of Staten Island. Trans. N. Y. Acad. Sci. 12: 28-39.

1906. The Cretaceous flora of southern New York and
New England. U. S. Geol. Surv., Monogr. 50: 1-219.

1931. Plant remains from a Pleistocene lake deposit
in the Upper Connecticut River Valley. Brittonia 1: 35-56.

HorLMEs, E. M. 1884. Medical plants used by the Cree Indians.
Hudson's Bay territory. Am. Jour. Pharm. 56: 617-621.

HornMES, M. L. 1956. Kalmia, the American laurels. Baileya 4:
89-94,

Howes, F. N. 1949. Sources of poisonous honey. Kew Bull. 1949:
167-171.

HULTÉN, E. 1948. Flora of Alaska and Yukon. Vol. 8. Lund.

1968. Flora of Alaska and neighboring Territories.
Stanford.

IkUusE, M. 1954. The presence of the viscid threads among pollen
grains in Phyllodoceae, etc. of Ericaceae. Jour. Jap. Bot. 29:
146-148.

JacoBS, M. L, & W. R. Lrovp. 1939. A phytochemical study of
Kalmia angustifolia. Jour. Am. Pharm. Assoc. 28: 408-412.
JAYNES, R. A. 1968a. Interspecific crosses in Kalmia. Am. Jour.

Bot. 55: 1120-1125.
1968b. Self incompatibility and inbreeding depression
in three laurel (Kalmia) species. Proc. Amer. Soc. Hort. Sci.
93: 618-622.
1968c. Breaking seed dormancy of Kalmia hirsuta
with high temperatures. Ecology 49: 1196-1198.
1969. Chromosome counts of Kalmia species and re-
valuation of K. polifolia var. microphylla. Rhodora 71: 280-284.
1971a. Laurel selections from seed. True breeding red-
budded mountain laurel. Cire. Conn. Agr. Exp. Sta. 240: 1-10.
1971b. A gene controlling pigmentation in sheep laurel.
Jour. Hered. 62: 201-203.
1971c. Seed germination of six Kalmia species. Jour.
Am. Soc. Hort. Sci. 96: 668-672.
. 1971d. The Kalmias and their hybrids, Quart. Bull.
Am. Rhododendron Soc. 25: 160-164.
. 1972. Selection and propagation of improved Kalmia
latifolia cultivars. Int. Pl. Prop. Soc. 21: 366-373.

JENNISON, H. M. 1935. Notes on some plants of Tennessee. Rhodora
37: 309-323.

JOHNSON, E. A., & J. L. KovNER. 1956. Effects on stream flow of
cutting a forest understory. Forest Sci. 2: 82-91.

1974] Genus Kalmia — Ebinger 395

KENNEDY, G. W. 1875. Arbutin in Kalmia latifolia, Lin. Am.
Jour. Pharm. 47: 5-6.

KiNGSBURY, J. M. 1964. Poisonous plants of the United States and
Canada. Englewood Cliffs.

KRAUSEL, R., & H. WEYLAND. 1959. Kritische Untersuchungen zur
Kutikularanalyse Tertiàrer blátter. IV. Dikotyledonen aus der
braunkohle. Palaeontographica Abt. B. 105: 101-124.

KUNTZE, O. 1891. Revisio Generum Plantarum. Part II. Leipzig.

KunMES, E. A. 1961. The ecology of mountain laurel in southern
New England. Ph.D. Thesis, Yale Univ.

1967. The distribution of Kalmia latifolia L. Am.
Midl. Nat. 77: 525-526.

LAMSON-SCRIBNER, F. 1891. A sketch of the flora of Orono, Me.
Bot. Gaz. 16: 228-234.

LANJOUW, J. 1966. International code of botanical nomenclature.
Regnum Veg. 46. Utrecht.

, & F. A. STAFLEU. 1964. Index Herbariorum. Part. 1.
The herbaria of the world. 5th ed. Regnum Veg. 31. Utrecht.

Lascut, A. J. M. 1889. Examination of some of the poisonous
Ericaceae of North America. Pharm. Rund. 7: 208-213.

LINNAEUS C. 1753. Species Plantarum. Stockholm.

—— —————. 1754. Genera Plantarum. 5th ed. Stockholm.

LirkE, R. 1968. Pflanzenreste aus dem untermiozán in Nordwest-
sachsen. Palaeontographica Abt. B. 123: 173-183.

LovELL, J. H. 1926. Honey plants of North America. Medina.

, & H. B. LovELL. 1934. The pollination of Kalmia
angustifolia. Rhodora 36: 25-28.

Lovett, H. B. 1951. The mechanics of pollination. Bees 5: 18-23.

Lupwic, C. F. 1783. Die neuere wilde baumzucht in einem alpha-
betischen und systematischen verzeichisse. Leipzig.

MACOUN, J. 1884. Catalogue of Canadian Plants. Part II. Gamo-
petalae. Montreal.

MansH, C. D., & A. B. CLAWSON. 1930. Mountain-laurel ( Kalmia
latifolia) and sheep laurel (Kalmia angustifolia) as stock-
poisoning plants. U. S. Dept. Agr. Tech. Bull. 219: 1-22.

MATTHEWS, J. R., & E. M. KNox. 1926. The comparative morphology
of the stamens in the Ericaceae. Trans. Proc. Bot. Soc. Edinb.
29: 243-281.

Matusow, H. 1897. Analysis of the root of Kalmia latifolia. Am.
Jour. Pharm. 69: 341-343.

MERRILL, E. D. 1945. In defense of the validity of William Bar-
tram's binomials. Bartonia 23: 10-35.

MEYER, B. S. 1927. The measurement of the rate of water-vapor
loss from leaves under standard conditions. Am. Jour. Bot. 14:
582-591.

396 Rhodora [Vol. 76

MILLER, C. C. 1895. Honey from mountain laurel. Am. Bee Jour.
35: 825-826.

MOUILLEFERT, P. 1897. Traite des Arbres & Arbrisse. Vol. II.
Paris.

MUENSCHER, W. C. 1957. Poisonous plants of the United States.
Revised ed. New York.

Munns, E. N. 1938. The distribution of important forest trees of
the United States. U. S. Dept. Agr. Misc. Publ. 287: 1-176.

NICHOLS, G. E. 1934. The influence of exposure to winter tempera-
ture upon seed germination in various native American plants.
Ecology 15: 364-373.

NICHOLSON, G. 1896. Hand-list of trees and shrubs grown in Arbo-
retum. Part II. Gamopetalae to Monocotyledons. Royal Gardens,
Kew.

NICHOLSON, S. M, & J. F. CrLovis. 1967. Dye plants and dye
methods in West Virginia. Castanea 32: 111-116.

NORTON, A. H. 1913. Some noteworthy plants from the islands and
coast of Maine. Rhodora 15: 137-143.

1935. Dr. Aaron Young, Jr. and the botanical survey
of Maine. Rhodora 37: 1-16.

PARKER, J. 1961. Seasonal changes in cold resistance of some north-
eastern woody evergreens. Jour. Forest. 59: 108-111.

PASCHKIS, H. 1881. Contributions to a closer knowledge of some
little known leaves. Pharm. Jour. Trans. 12: 85-87.

PELLETT, F. C. 1920. American honey plants. Hamilton.

PENHALLOW, D. P. 1882. Fertilization of Kalmia. Gard. Month.
24: 54.

PERKINS, A. E. 1935. Notes on some rare plants of York County,
Maine. Rhodora 37: 415-416.

PETERSON, E. B. 1965. Inhibition of Black Spruce primary roots
by a water-soluble substance in Kalmia angustifolia. Forest Sci.
11: 473-479.

PLUKENET, L. 1696. Opera omnia botanica in sex tomos divisa.
IV. Almagestum botanicum. London.

PORCHER, F. P. 1869. Resources of the Southern field and forests.
Charleston.

PRITCHARD, W. R. 1956. Laurel (Kalmia angustifolia) poisoning
of sheep. N. Am. Veterinarian 37: 461-462.

PuRCHAS, S. 1625. Purchas His Pilgrimes. 3rd. Part. 16: 581-595.

PunsH, F. 1814. Flora Americae Septentrionalis. London.

RADFORD, A. E., H. E. AHLES, & C. R. BELL. 1968. Manual of the
vascular flora of the Carolinas. Chapel Hill.

RAFINESQUE, C. S. 1830. Medical Flora; or manual of the medical
botany of the United States of North America. Vol. 2. Phila-
delphia.

1974] Genus Kalmia — Ebinger 397

1840. Autikon Botanikum, Philadelphia.

REHDER, A. 1910. Notes on the forms of Kalmia latifolia. Rhodora
12: 1-3.

. 1945. Notes on some cultivated trees and shrubs, II.
Jour. Arnold Arb. 26: 472-481.

1949. Bibliography of cultivated trees and shrubs
hardy in the cooler temperate regions of the Northern Hemi-
sphere. Jamaica Plain.

REIFSNYDER, W. E. 1961. Seasonal variation in the moisture con-
tent of the green leaves of Mountain Laurel. Forest Sci. 7:
16-23.

RoTHROCK, J. T. 1867. The fertilization of flowering plants. Am.
Nat. 1: 64-72.

RYDBERG, P. A. 1954. Flora of the Rocky Mountains and adjacent
plains. New York.

SARGENT, C. S. 1890. New or little known plants. A curious form
of Kalmia. Garden Forest 3: 452.

1893. Silva of North America, Vol. 3. Boston.

Scott, G. D. 1962. Kalmia polifolia in Scotland. Glasgow Nat.
18: 196-197.

ScoTTER, G. W. 1966. A contributfon to the flora of the eastern
arm of Great Slave Lake, Northwest Territories. Canad. Field-
Nat. 80: 1-18.

SHANKS, R. E. 1952. Notes on woody plant distribution in Ten-
nessee. Castanea 17: 90-96.

SMALL, J. K. 1903. Flora of the Southeastern United States. New
York.

1914. Ericaceae. North Amer. Flora 29: 33-102.

SMITH, J. 1624. The general history of Virginia, New England, and
the Summer Isles. Book 5.

SMITH, W. W., A. R. HopGDoN, & R. EGGERT. 1947. Progress re-
port on chemical weed control in blueberry fields. Proc. Am.
Soc. Hort. Sci. 50: 233-238.

SPRAGUE, E. 1871. The Rhododendron and “American Plants.”
Boston.

SPRENGEL. C. K. 1793. Das entdeckte Geheimnifs der Natur im
Bau und in der Befruchrung der Blumen. Berlin.

STANTEY. O. B. 1931. Fat deposits in certain Ericaceae. Butler
Univ. Bot. Stud. 2: 33-41.

STONE, G. E. 1909. A remarkable form of Kalmia latifolia. Rho-
dora 11: 199-200.

TALLENT. W. H., M. L. RIETHOF, & E. C. HORNING. 1957. Studies
on the occurrence and structure of acetylandromedol (androme-
dotoxin). Jour. Am. Chem. Soc. 79: 4548-4554.

THIERET. J. W. 1963. Kalmia polifol’a: Second record from the
Arctic. Canad. Field-Nat. 77: 173.

398 Rhodora [Vol. 76

TREVETT, M. F. 1956. Control of lambkill (Kalmia angustifolia) in
lowbush blueberry fields. Northeastern Weed Control Conference
Proc. 10: 193-198.

TRUMPY, J. R. 1893. Propagating Kalmias. Gardening 1: 122.

VAN RENSSELAER, M. G. 1889. The choice of a national flower.
Garden Forest 2: 333-335.

WAHLENBERG, W. G., & W. T. DOOLITTLE. 1950. Reclaiming Appa-
lachian brush lands for economic forest production. Jour. Forest.
48: 170-174.

WANGENHEIM, F. U. J. 1788. Beschreibung der Polenblattrigen
Kalmia und der gelbbluhenden Rosskastanie. Schr. Berlin Ges.
Naturf. Freunde 8: 129-133.

Waup, R. A. 1940. The action of Kalmia angustifolia (Lambkill).
Jour. Pharm. Exp. Therap. 69: 103-111.

Woop, C. E. JR. 1961. The genera of Ericaceae in the South-
eastern United States. Jour. Arnold Arb. 42: 10-80.

Woop, G. B., & F. BACHE. 1851. The Dispensatory of the United
States of America. Philadelphia.

Woop, H. B., V. L. STROMBERG, J. C. KERESZTESY, & E. C. HORNING.
1954. Andromedotoxin. A potent hypotensive agent from Rhodo-
dendron maximum. Jour.“Am. Chem. Soc. 76: 5689-5692.

Woop, T. F. 1883. Is sheep laurel poisonous to sheep? Am. Agr.
42: 66.

EASTERN ILLINOIS UNIVERSITY
CHARLESTON, ILLINOIS 61920

MARINE ALGAE OF RHODE ISLAND
R. D. Woop AND MARTINE VILLALARD-BOHNSACK

With increasing marine research activity in the Narra-
gansett Bay area, the need for information about local
marine algae has become urgent. Therefore, the available
information has been compiled (Wood, in press) and a list
of species known to occur in the State is presented.

The earliest serious collecting of algae in Rhode Island
seems to have been by S. T. Olney and his associates, and
considerable effort was made in the years of 1846-1849.
But, also, during the summer of 1846, J. W. Bailey visited
Rhode Island and collected with Olney. Bailey, working
independently, obtained specimens from Europe and worked
up the collections systematically and included the Rhode
Island records in his publication of 1847. Olney, on the
other hand, sent his material to W. H. Harvey of Dublin,
who identified them. The records were published by Olney
in 1847, the same year that Bailey's appeared. Whose work
appeared first is not clear, but Olney was the first Rhode
Islander to publish on the marine algae of the State, while
Bailey was probably the first worker to both study and
publish on algae of Rhode Island. Subsequent records are
those by Harvey (1852-1858), Olney (1871), Farlow (1873,
1875, 1876, and 1881), and Bennett (1888), and the several
reports by Collins between 1882 and 1896 which were
largely summarized in his works of 1900 and 1909-1918.
More recent reports are those by Wood & Palmatier (1954),
the dissertation by Villalard (1971), and thesis by Vaughan
(1971). In addition, three collections of concern are the
*Algae Rhodiaceae" by Olney (1871) and the two large
exsiccata sets in which Rhode Island algae occur — Farlow,
Anderson & Eaton’s (1877-1889) “Algae Exsiccatae Ameri-
cae-Borealis” and Collins, Holden & Setchell’s (1895-1919)
“Phycotheca Boreali-Americana.”

The senior author (rdw) prepared a list of species re-
corded from Rhode Island by Taylor (1957), and then

399

400 Rhodora [Vol. 76

searched the literature for other early records. The sup-
porting specimens were sought and examined, especially
those at Brown University (courtesy of G. L. Church and
W. H. Snell) and at the Farlow Herbarium at Harvard
University (courtesy of I. M. Lamb). The complex of iden-
tifications and synonyms required careful tracing, but veri-
fication of most early names by examination of the actual
collections was possible. The background information is to
appear elsewhere (Wood, in press). The final list was
checked by the junior author (mvb), and the latter also
compiled the currently suggested synonyms that appear
parenthetically in the list. Lastly, an evaluation of the
frequency of occurrence of each taxon, based upon the
authors’ personal observations, was included. Where identi-
fications were still uncertain, Dr. Wm, Randolph Taylor
graciously aided. The final manuscript was examined and
edited by Urda K. Wood.

It is hoped that this list will provide a background for
future work on the algae of Rhode Island waters.

LIST OF MARINE ALGAE KNOWN FOR RHODE ISLAND

The nomenclature below follows Taylor (1957), but other
suggested names and references are indicated parentheti-
cally. However, author citations have been checked, and
those involving “in” and “ex” entries include both authors.
Species reported for the first time in Rhode Island are desig-
nated by an asterisk (*). The earliest known published rec-
ord for each species in Rhode Island is cited in the second
column, the abbreviations being: B — Bailey (1847), BENN.
= Bennett (1888), B,o = Bailey (1847) and Olney (1847),
c = Collins (1900), F = Farlow (1873), o = Olney (1847),
OAR [for Olney’s “Algae Rhodiaceae"] = Olney (1871), PBA
= in Collins, Holden & Setchell (1895-1919), T = Taylor
(1937), (VA) = thesis by Vaughan (1971), (VI) = disserta-
tion by Villalard (1971), W = Wood (1962), w&P = Wood
& Palmatier (1954).

Species which should be expected in the state, based on

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 401

Taylors (1957) range records, but for which no adequate
report is yet available, are summarized at the end of the list.

The frequency of occurrence of each species based on our
combined personal observations is indicated in the third
column as: t = trace, rare, + = local and uncommon, ++
= locally common, +++ = general and quite common.
Where a rare taxon has been seen by only one of us, it
is indicated as by “rdw” or “mvb” for R. D. Wood or M.
Villalard-Bohnsack, respectively.

CHLOROPHYTA

Ulotrichaceae
1. Stichococcus marinus
(Wille) Hazen T +
2. Ulothrix flacca
(Dillw.) Thuret in Le Jol. C dar

Chaetophoraceae

3. Pseudendoclonium submarinum
Wille T

Gomontiaceae

4. Gomontia polyrhiza
(Lagerh.) Born. et Flah. T

Ulvaceae
* 5. Capsosiphon fulvescens
(C.Ag.) Setch. et Gard. (VI) *
* 6. [Enteromorpha ahlneriana
Blid.] (VI) t
7. E. clathrata (Roth) J. Ag. OAR +
(E. clathrata (Roth) Grev. —
Parke & Dixon, 1964)
8. E. compressa (L.) Grev. B,O +
(ecotype of E. intestinalis (L.)
Link — DaSilva & Burrows in
Norton, 1972, p. 385)
9. E. cruciata Collins OAR

402

10

Rhodora

. E. erecta, (Lyngb.) J. Ag.
(E. flexuosa ssp. paradoxa
(Dillw.) Blid. — Bliding, 1963)

* 11. E. groenlandica (J.Ag.) Setch.

12.
13.
14.

15.

16.
17.

18.

20.

21.

23.

24.

et Gard.

E. intestinalis (L.) Link

E. linza (L.) J. Ag.

E. marginata J. Ag.
(Blidingia marginata (J. Ag.)
Dang. — Dangeard, 1958)

E. minima Nag.

(Blidingia minima (Nag. ex
Kitz.) Kylin — Kylin, 1947)
E. prolifera (Müll.) J. Ag.
Monostroma fuscum (Post. et
Rupr.) Witt.

(Ulvaria obscura (Kütz.) Gayr.
— Gayral, 1965)

M. grevillei (Thuret) Witt.
(Ulvopsis grevillei (Thuret)
Gayr. — Gayral, 1965)

. M. leptodermum Kjellm.

(Kornmannia leptoderma

(Kjellm.) Blid. — Bliding,

1968)

M. oxyspermum (Kütz.) Doty

(Ulvaria oxyspermum (Kiitz.)

Blid. — Bliding, 1968)

M. pulchrum Farlow

. Percursaria percursa (C. Ag.)
Rosenv.

Ulva lactuca L.
var. latissima (L.) DC.

Prasiolaceae
Prasiola stipitata Suhr. in Jess.

B,O

(VI)

B,O

B

[Vol. 76

+++

++

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 403

Cladophoraceae

25. Chaetomorpha aerea (Dillw.)
Kütz. B +
(Ch. linum (Müll. Kütz. —
Christensen, 1957)
26. Ch. linum (Miill.) Kütz. B ++
27. Ch. melagonium (Web. et
Mohr) Kiitz. t(rdw)
28. Cladophora albida (Huds.)
Kiitz. F tT
29. Cl. flexuosa (Dillw.) Harv. C +
(Cl. serica, (Huds.) Kütz. —
Hoek, 1963)
f. densa Collins PBA
30. Cl. glaucescens (Harv.) Harv. F
(Cl. albida (Huds.) Kutz. —
Hoek, 1963)
31. Cl. gracilis (Griff. ex Harv.)
Kütz. [o rain
(Cl. serica (Huds.) Kütz. —
Hoek, 1963)
32. Cl. hutchinsiae (Dillw.) Kütz.
var. distans (C. Ag.) Kütz. T
(Cl. hutchinsiae (Dillw.) Kütz.
— Hoek, 1963)
* 83. Cl. laetevirens (Dillw.) Kütz. (VI) m
34. Cl. magdalenae Harv. F
(Cl. aibida (Huds.) Kütz. —
Hoek, 1963)
35. Cl. refracta (Roth) Kütz. O
(Cl. albida (Huds.) Kütz. —
Hoek, 1963)
36. Cl. rudolphiana (C. Ag.) Harv. C
(Cl. serica (Huds.) Kütz. —
Hoek, 1963)
37. Cl. rupestris (L.) Kütz. OAR zu
38. Rhizoclonium riparium (Roth)
Harv. C t

404

* 89.

40.

* A1.

* 42.

43.

44.

45.

46.

Rhodora

var. implexum (Dillw.)
Rosenv.

var. polyrhizum (Lyngb.)
Rosenv.

R. tortuosum Kütz.

(R. riparium (Roth) Harv. —
Scagel, 1966)

Spongomorpha arcta (Dillw.)
Kütz.

(Acrosiphonia arcta (Dillw.)
J. Ag. — Kornmann, 1962)
S. lanosa (Roth) Kütz.

(S. aeruginosa (L.) Hoek —
Hoek, 1963)

S. spinescens Kütz.
(Acrosiphonia arcta (Dillw.)
J. Ag. — Kornmann, 1962)
Urospora penicilliformis
(Roth) Aresch.

Bryopsidaceae

Bryopsis plumosa (Huds.)
C. Ag.

Codiaceae
Codium fragile (Sur.) Hariot
ssp. tomentosoides (van Goor)
Silva

PHAEOPHYTA

Ectocarpaceae

Ectocarpus confervoides
(Roth) Le Jol.

(E. siliculosus (Dillw.)
Lyngb. — South & Cardinal,
1970)

var. pygmaeus (Aresch.)
Kjellm.

B,O

(VI)

0?,c

[Vol. 76

++

t(mvb)

+++

++

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 405

4T.
48.
49.
50.
51.

52.

53.

54.

55.
56.

57.

58.

E. dasycarpus Kuck.

E. fasciculatus Harv.

E. penicillatus (C. Ag.) Kjellm.
E. siliculosus (Dillw.) Lyngb.
E. tomentosus (Huds.) Lyngb.
(Spongonema tomentosum
(Huds.) Kütz. — Kuckuck,
1960)

Giffordia granulosa

(Smith) Hamel

G. mitchellae (Harv.) Hamel
(G. mitchelliae (Harv.) Hamel
— Earle, 1969)

G. ovata (Kjellm.) Kylin

G. secunda (Kiitz.) Batt.
Pylaiella littoralis (L.) Kjellm.
(Pilayella littoralis (L.)
Kjellm. — Dixon & Russell,
1964)

Sorocarpus micromorus
(Bory) Silva

Streblonema aecidioides
(Rosenv.) Fosl.

(Entonema aecidioides (Rosenv.)
Kjellm. — Edelstein &
McLachlan, 1968)

. S. chordariae (Farl.) De Toni

Sphacelariaceae

. Cladostephus verticillatus

(Lightf.) Lyngb.

. Sphacelaria cirrosa

(Roth) C. Ag.

. S. radicans (Dillw.) C. Ag.

Myrionemataceae

. Ascocyclus orbicularis Magnus

(Myrionema magnusii (Sauv.)
Lois. — Loiseaux, 1967)

B,O

++

+ (rdw)

406

64.

65.
66.

67.

68.

69.
70.

71.
72.

73.

14.

75.

76.

77.

Rhodora

Hecatonema terminalis
(Kiitz.) Kylin

Myrionema corunnae Sauv.
M. foecundum (Stromf.) Fosl.
(Hecatonema foecundum
(Strómf.) Lois. — Parke &
Dixon, 1968)

M. globosum (Reinke) Fosl.
(Microsporangium globosum
Reinke — Parke & Dixon, 1964)
M. strangulans Grev.

Ralfsiaceae
Ralfsia borneti Kuck.
R. clavata (Harv. in Hook.)
Crouan frat.
R. pusilla (Strómf.) Batt.
R. verrucosa (Aresch.) J. Ag.

Elachisteaceae

(Elachistaceae — Parke & Dixon (1964) )
PBA

Elachistea chondri Aresch.
(Elachista chondri Aresch. —
Fritsch, 1945; Parke & Dixon,
1964)

E. fucicola (Vell.) Aresch.

( Elaschista fucicola Aresch. —
Fritsch, 1945; Parke & Dixon,
1964)

Halothrix lumbricalis

(Kütz.) Reinke

Chordariaceae
Chordaria flagelliformis
(Müll.) C. Ag.
var. densa Farl.
Eudesme virescens (Carm. ex
Harv. in Hook.) J. Ag.

Q

[Vol. 76

++

++

++

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 407

78. Leathesia difformis (L.)

Aresch. OAR ++
79. Sphaerotrichia divaricata
(C. Ag.) Kylin OAR +
Stilophoraceae
80. Stilophora rhizodes
(Turn.) J. Ag. OAR
Desmarestiaceae
81. Arthrocladia villosa
(Huds.) Duby C t
82. Desmarestia aculeata (L.)
Lamour. B +
83. D. viridis (Müll. Lamour. B "n
Striariaceae
84. Striaria attenuata
(Grev.) Grev. C
Punctariaceae
* 85. Asperococcus echinatus
(Mert.) Grev. (VI) Tom

(A. fistulosus (Huds.) Hook. —
Edelstein, Wynne & McLachlan,

1970)
86. Desmotrichum undulatum

(J. Ag.) Reinke B,O
87. Litosiphon filiformis

(Reinke) Batt. PBA
88. Myriotrichia clavaeformis

Harv. C
89. M. filiformis Harv. F
90. Petalonia fascia

(Müll. Kuntze BENN. ++
91. Punctaria latifolia Grev. BENN. +
92. P. plantaginea (Roth) Grev. O ule ce
93. Rhadinocladia cylindrica

Schuh T

94. R. farlowii Schuh T

408

95.

96.

97.

98.
99.
100.
101.

102.
103.

104.
105.

*106.

107.
108.

109.

Rhodora

Scytosiphon lomentaria
(Lyngb.) Link

(S. lomentarius (Lyngb.) Link
— Farlow, 1881)

Dictyosiphonaceae
Dictyosiphon chordaria
Aresch. (as Gobia)
D. foeniculaceus
(Huds.) Grev.

Laminariaceae

Alaria esculenta (L.) Grev.
Chorda filum (L.) Lamour.
C. tomentosa Lyngb.
Laminaria agardhii Kjellm.
(L. saccharina (L.) Lamour. —
Wilce, 1965)

L. digitata (Huds.) Lamour.
L. intermedia Fosl.

(L. digitata (Huds.) Lamour.
— Sundene, 1958)

L. longicruris De la Pyl.

L. saccharina (L.) Lamour.

Fucaceae

Ascophyllum mackaii
(Turn.) Holmes et Batt.

(A. nodosum (L.) Le Jol. —
Gibb, 1957)

A. nodosum (L.) Le Jol.
Fucus edentatus De la Pyl.
(F. distichus L. ssp. edentatus
(De la Pyl.) Pow. — Powell,
1963)

F. evanescens C. Ag.

(F. distichus L. ssp.
evanescens (C. Ag.) Pow. —
Powell, 1963)

B,O

B,O

[Vol. 76

++ +++

+++

++

+ (rdw)

++, +++
TG

+?

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 409

110. F. spiralis L. B res
var. limitaneus Montagn. T

111. F. vesiculosus L. B,O ro
Sargassaceae

112. Sargassum filipendula C. Ag. B,O +

*113. S. natans (L.) J. Meyen (VA) t(drift)

RHODOPHYTA
Bangiaceae

114. Bangia ciliaris Carm. ex Harv. C t

(Erythrotrichia ciliaris (Carm.
ex Harv.) Thur. in Le Jol. —
Parke & Dixon, 1964)

115. B. fuscopurpurea

(Dillw.) Lyngb. B,O FT
116. Erythrotrichia carnea
(Dillw.) J. Ag. C
117. Goniotrichum alsidit
(Zanard.) Howe C t
118. Porphyra leucosticta
Thuret in Le Jol. t
*119. P. miniata (C. Ag.) C. Ag. (VI) x
120. P. umbilicalis (L.) J. Ag. B,O dT
Acrochaetiaceae
121. Acrochaetium daviesti
(Dillw.) Nag. C
122. A. flexuosum Vickers T
123. Kylinia secundatum
(Lyngb.) Papenf. C

(Acrochaetium secundatum
(Lyngb.) Nig. — Parke &
Dixon, 1968)
124. K. virgatula (Harv.) Papenf. C
(Acrochaetium virgatula
(Harv.) J. Ag. — Parke &
Dixon, 1968)

410

125.

126.

127.

128.

*129.

131.

132.

133.

Rhodora

Rhodochorton penicilliforme
(Kjellm.) Rosenv.

R. purpureum

(Lightf.) Rosenv.

Helminthocladiaceae

Nemalion multifidum
(Web. et Mohr) J. Ag.

(N. helminthoides (Vell. in
With.) Batt. — Parke &
Dixon, 1964)

Chaetangiaceae

Scinaia furcellata
(Turn.) Bivona

Bonnemaisoniaceae
Asparagopsis hamifera

(Hariot) Okam. (VI), (VA)

(Bonnemaisonia hamifera Har.
— Feldmann & Feldmann,

1942)
Gelidiaceae
. Gelidium crinale
(Turn.) Lamour.
Dumontiaceae

Dumontia incrassata
(Miill.) Lamour.

Rhizophyllidaceae
Polyides caprinus
(Gunn.) Papenf.
(P. rotundus (Huds.) Grev.
— Parke, 1953)
Squamariaceae

Hildenbrandia prototypus
Nardo

[Vol. 76

++

+ (mvb)

++

++

+++

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 411

134. Rhododermis georgii
(Batt.) Collins T
(Rhodophysema georgii Batt.
—Dixon, 1964)

Corallinaceae
135. Corallina officinalis L. BENN. mtm
136. Fosliella farinosa
(Lamour.) Howe e

(Melobesia farinosa Lamour.
— Hamel & Lemoine, 1952)
137. F.lejolisii (Rosan.) Howe C +
(Melobesia lejolisit Rosan. —
Hamel & Lemoine, 1952)
138. Lithophyllum corallinae

(Crouan frat.) Heydr. PBA
139. L. macrocarpum
(Rosan.) Fosl. T

(Dermatolithon pustulatum
(Lamour.) Fos]. — Edelstein
& McLachlan, 1968)

140. L. pustulatum

(Lamour.) Fosl. OAR
*141. L. lenormandi
(Aresch. in J. Ag.) Fosl. (VI) oe

(Phymatolithon lenormandii
(Aresch. in J. Ag.) Adey —
Adey, 1966)
142. Melobesia membranacea
(Esper) Lamour. C
(Epilithon membranaceum
(Esper) Heydr. — Parke &
Dixon, 1964)
143. Phymatolithon laevigatum
(Fosl.) Fosl. T ++
(Lithothamnion laevigatum
Fos]. — Parke & Dixon, 1964)

412

144

145

146.

147.

148.

149.

150.

151.

Rhodora

P. polymorphum (L.) Fosl. C
(Lithothamnion polymorphum

(L.) Aresch. in J. Ag. — Parke

& Dixon, 1964)

Gloiosiphoniaceae
Gloiosiphonia capillaris
(Huds.) Carm. ex Berk. C
Kallymeniaceae
Euthora cristata
(C. Ag.) J. Ag. F
Choreocolacaceae
Choreocolax polysiphoniae
Reinsch C
Harveyella mirabilis
(Reinsch) Reinke T
Solieriaceae
Agardhiella tenera
(J. Ag.) Schmitz B

(as used by Taylor (1957), but
designated A. baileyi (Harv. ex
Kütz.) Taylor in Taylor &
Rhyne (1970: 13), and since
changed to Neoagardhiella
baileyi (Harv. ex Kütz.) Wynne
& Taylor, Wynne & Taylor,

1973, p. 101)
Rhodophyllidaceae
Cystoclonium purpureum
(Huds.) Batt. B,O
var. cirrhosum Harv. C
Hypneaceae

Hypnea musciformis

(Wulf.) Lamour. BENN.

[Vol. 76

t—+
(periodic)

t(drift?)

ab

++ +++

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 413

Gracilariaceae
152. Gracilaria foliifera
(Forssk.) Borg. (0) t
153. G. verrucosa (Huds.) Papenf. W&P +++
Phyllophoraceae
154. Ahnfeltia plicata
(Huds.) Fries B,O T
155. Gymnogongrus griffithsiae
(Turn.) Mart. C t(mvb)
156. Phyllophora brodiaei
(Turn.) Endl. F +

(P. truncata (Pallas) Newr. et
Taylor — Newroth & Taylor,

1971)
var. catenata (C. Ag.) Collins T
157. P. membranifolia
(Good. et Woodw.) J. Ag. 0 mo

(P. pseudoceranoides (Gmel.)
Newr. et Taylor — Newroth &
Taylor, 1971)

158. P. traillii Holmes ex Batt. C
Gigartinaceae
159. Chondrus crispus Stackh. B,O ttt
160. Gigartina stellata
(Stackh.) Batt. G M

(Palmaria palmata (L.)
Stackhouse — Guiry, 1974,

p. 219.)
Rhodymeniaceae
161. Rhodymenia palmata
(L.) Grev. B,O TARERE
Champiaceae
162. Champia parvula
(C. Ag.) Harv. B,O bad a

163. Lomentaria baileyana
(Harv.) Farl. OAR TRT

414

*164.

165

166.

167.
168.

169.
170.

171.

172.

173.
174.
175.

176.
177.
178.
179.

180.
181.
182.
183.

184.

185.

Rhodora

[L. clavellosa (Turn.) Gail.]
L. orcadensis
(Harv.) Coll. ex Taylor

Ceramiaceae

Antithamnion americanum
(Harv.) Farl. in Kjellm.

A. cruciatum (C. Ag.) Nag.
A. plumula

(Ellis) Thuret in Le Jol.

A. pylaisaei (Mont.) Kjellm.
Callithamnion baileyi Harv.
var. laxum Farlow

C. byssoides

Arnott ex Harv. in Hook.

C. corymbosum

(Smith) Lyngb.

C. roseum (Roth) Lyngb.

C. tetragonum (With.) C. Ag.
Ceramium deslongchampii
Chauv. in Duby

C. diaphanum (Lightf.) Roth
C. fastigiatum Harv.

C. rubriforme Kylin (prox.)
C. rubrum (Huds.) C. Ag.
var. pedicellatum Duby

(C. pedicellatum DC. — Parke
& Dixon, 1964)

var. squarrosum Harv.

C. strictum Harv.

Griffithsia globulifera Harv.
G. tenuis C. Ag.
Pleonosporium borreri
(Smith) Näg. ex Hauck
Plumaria elegans

(Bonnem.) Schmitz

Ptilota serrata Kütz.

[Vol. 76

+ (mvb)
OAR
C
BENN. +
F
C
OAR t
T
OAR
B =t-
C +
O
O
B,O +,+ +
OAR ++
(VA) t?
B,O +++
C
OAR
C +
C +
O0 t
OAR
F Tte
B?,F t

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 415

186.

187.

188.

*189.

190.

191.

193.
194.
195.
196.

197.

198
199

Seirospora griffithsiana Harv.
(S. seirosperma (Harv.)
Dixon — Dixon, 1964)
Spermothamnion turneri
(Mert.) Aresch.

(S. repens (Dillw.) Rosenv. —
Rosenvinge, 1924)

Spyridia filamentosa

(Wulf.) Harv. in Hook.

T railliella, intricata

(J. Ag.) Batt.

(sporophyte of Asparagopsis
hamifera — Taylor, 1957,

p. 291)

Delesseriaceae
Grinnellia americana
(C. Ag.) Harv.
Phycodrys rubens (L.) Batt.

Dasyaceae

. Dasya pedicellata
(C. Ag.) C. Ag.
(D. baillouviana (Gmel.) Mont.
— Dixon & Irvine, 1970)

Rhodomelaceae
Chondria baileyana
(Mont.) Harv.
C. dasyphylla (Woodw.) C. Ag.
C. sedifolia Harv.
C. tenuissima
(Good. et Wooaw.) C. Ag.
Polysiphonia denudata
(Dillw.) Grev. ex Harv. in
Hook.
. P. elongata Harv. in Hook.
. P. fibrillosa (Dillw.) Spreng.

OAR

(VI)

B,O

++
++

++

ob
++

416 Rhodora [Vol. 76

200. P. harveyi Bail. 0 tt
var. olneyi (Harv.) Collins O
201. P. lanosa (L.) Tandy B,O ++, +++
202. P. nigra (Huds.) Batt. F +
203. P. nigrescens (Huds.) Grev. B +
204. P. novae-angliae Taylor B
205. P. subtilissima Mont. F +
206. P. urceolata
(Lightf. ex Dillw.) Grev. C ++
207. Rhodomela confervoides
(Huds.) Silva B,O +++

208. R. virgata Kjellm. C

SPECIES TO BE EXPECTED IN RHODE ISLAND,
AND REPORTED FROM ADJACENT STATES BY TAYLOR (1957)

CHLOROPHYTA
Chlorococcaceae
Codiolum gregarium A. Br.
Ulotrichaceae
Ulothrix implexa Kütz.
Chaetophoraceae

Bolbocoleon piliferum Pringsh.
Entocladia flustrae (Reinke) Batt.
(Epicladia flustrae Reinke — Parke & Dixon, 1968)
E. wittrockii Wille
(Ectochaete wittrockii (Wille) Kylin — Parke & Dixon, 1968)
Pringsheimiella scutata (Reinke) Marchew.
Protoderma marinum Reinke
(Pseudendoclonium marinum (Reinke) Aleem et Schulz — Loiseaux,
1967)

Ulvaceae

Enteromorpha plumosa Kütz.
(E. flexuosa ssp. paradoxa (Dillw.) Blid. — Bliding, 1963)

Cladophoraceae
Chaetomorpha atrovirens Taylor
Cladophora crystallina (Roth) Kütz.
(Cl. vagabunda (L.) Hoek; Cl. glomerata (L.) Kütz. — Hoek, 1963)
Cl. expansa (Mert.) Kütz.
(Cl. vagabunda (L.) Hoek — Hoek, 1963)

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack
Cl. flavescens (Roth) Kütz.
(a doubtful name — Hoek, 1963)

Derbesiaceae
Derbesia vaucheriaeformis (Harv.) J. Ag.

Bryopsidaceae
Bryopsis hypnoides Lamour.

Phyllosiphonaceae
Ostreobium quekettii Born. et Flah.
XANTHOPHYTA
Vaucheriaceae
Vaucheria litorea C. Ag.
V. thuretii Woronin
PHAEOPHYTA
Ectocarpaceae

Ectocarpus subcorymbosus Farl.
(Farlowiella onusta (Kütz.) Kuck. — Kuckuck, 1956)

Elachisteaceae
(Elachistaceae — Parke & Dixon, 1964)

Myriactula minor (Farl. Taylor

Chordariaceae
Acrothrix novae-angliae Taylor
Eudesme zosterae (Lyngb.) Kylin
(Cladosiphon zosterae (J. Ag.) Kylin — Parke, 1953)
Striariaceae
Stictyosiphon subsimplex Holden

Punctariaceae
Desmotrichum balticum Kiitz.
(D. undulatum (J. Ag.) Reinke — Kylin, 1947a)
Sargassaceae
Sargassum fluitans Borg.

RHODOPHYTA
Bangiaceae
Asterocytis ramosa (Thwaites in Harv.) Gobi ex Schmitz

Acrochaetiaceae
Audouinella membranacea (Magnus) Papenf.

417

418 Rhodora [Vol. 76

Corallinaceae

Lithothamnium laeve (Strómf.) Fosl.
(Leptophytum laeve (Strómf.) Adey — Adey, 1966)

Ceramiaceae
Callithamnion circinatum (Kütz.) J. Ag.

Delesseriaceae
Membranoptera denticulata (Mont.) Kylin

Rhodomelaceae

Bostrychia rivularis Harv.
Polysiphonia flexicaulis (Harv.) Collins

LITERATURE CITED

ADEY, W. H. 1966. The genera Lithothamnion, Leptophytum (nov.
gen.) and Phymatolithon in the Gulf of Maine. Hydrobiologia
28(3-4): 321-370.

BAILEY, J. W. 1847. Notes on the algae of the United States, Am.
J. Sci. & Arts 2: 80-85, 399-403.

BENNETT, J. L. 1888. Plants of Rhode Island, being an enumeration
of plants growing without cultivation in the State of Rhode
Island. Published by the author, Providence Press Co., Provi-
dence. 128 pp.

BLIDING, C. 1963. A critical survey of European taxa in Ulvales,
I, Capsosiphon, Percursaria, Blidingia, Enteromorpha. Opera
Bot. 8(3): 1-160.

1968. A critical survey of European taxa in Ulvales,
II, Ulva, Ulvaria, Monostroma, Kornmannia, Bot. Not. 121:
535-629.

CHRISTENSEN, T. 1957. Chaetomorpha linum in the attached state.
Bot. Tidsskr. 53: 311-316.

CoLLINS, F. S. 1900. Preliminary lists of New England plants,
V. Marine algae. Rhodora 2: 41-52.

1909-1918. The green algae of North America. Tufts
Coll. St. 2(3): 79-480; First Suppl, ibid. 3(2): 69-109, 1912;
Second Suppl., ibid. 4(7): 1-106, 1918.

CoLLINS, F. S., I. HOLDEN, & W. A. SETCHELL. 1895-1919. Phyco-
theca Boreali-Americana. (Exsiccata) Fasc. 1-46 + A-E.
Malden.

DANGEARD, P. 1958. La reproduction et le développement de
l'Enteromorpha marginata J. Ag. et le rattachement de cette
espéce au genre Blidingia. C. R. Acad. Sci., Paris 246: 347-351.

DixoN, P. S. 1964. Taxonomic and nomenclatural notes on the
Florideae IV. Bot. Not. 117: 56-78.

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 419

Drxon, P. S., & L. M. IRVINE. 1970. Miscellaneous notes on algal
taxonomy and nomenclature, III. Bot. Not. 123(4): 474-487.
DixoN, P. S, & G. RUSSELL. 1964. Miscellaneous notes on algal
taxonomy and nomenclature, I. Bot. Not. 117(3): 279-284.
EARLE, S. A. 1969. Phaeophyta of the eastern Gulf of Mexico.

Phycologia 7(2): 71-254.

EDELSTEIN, T., & J. MCLACHLAN. 1968. Investigations of the marine
algae of Nova Scotia, V. Additional species new or rare to
Nova Scotia. Can. J. Bot. 46: 993-1003.

EDELSTEIN, T., M. J. WYNNE, & J. MCLACHLAN. 1970. Melanosiphon
intestinalis (Saund.) Wynne, a new record for the Atlantic.
Phycologia 9(1): 5-9.

Fartow, W. G. 1873. List of seaweeds or marine algae of the
south coast of New England. Rep. U. S. Comm. Fish & Fisheries
1: 281-294,

1875. List of the marine algae of the United States
with notes of new or imperfectly known species. Proc. Am.
Acad. Arts & Sci. 10: 351-380.

. 1876. List of the marine algae of the United States.
Rep. U. S. Comm. Fish & Fisheries 5: 691-718.

1881. The marine algae of New England. Rep. U. S.
Comm. Fish & Fisheries for 1879, Append. A-1: 1-210.

FARLOW, W. G., C. L. ANDERSON, & D. C. EATON. 1877-1889. Algae
Exsiccatae Americae-Borealis. (Exsiccata) Fasc. 1-5. Boston.

FELDMANN, J., & G. FELDMANN. 1942. Recherches sur les Bonne-
maisoniacées et leur alternance de générations. Ann. Sci. Nat.
Bot., Ser. II, 3: 75-175.

FRITSCH, F. E. 1945. The structure and reproduction of the algae.
Vol. II. The Univ. Press, Cambridge. 939 pp.

GAYRAL, P. 1965. Monostroma Thuret, Ulvaria Rupr., emend.
Gayral, Ulvopsis Gayral. (Chlorophycées, Ulotrichales): struc-
ture, reproduction, cycles, position systematique. Rev. Gén. Bot.
72 (857): 627-638.

GIBB, D. C. 1957. The free-living forms of Ascophyllum nodosum
(L.) Le Jol. J. Ecol. 45(1): 49-83.

Guiry, M. D. 1974. A reappraisal of the genus Palmaria Stack-
house. Brit. Phyc. J. 9(2): 219. :
HAMEL, G., & P. LEMOINE. 1952. Corallinacées de France et
d'Afrique du Nord. Arch. Mus. Natn. Hist. Nat., Paris 7(1):

17-136.

Harvey, W. H. 1852-1858. Nereis boreali-americana. Part I.
Smithsonian Contrib. Knowledge 3(4): 1-150; Part II, ibid.,
5(5): 1-258, 1853; Part III, tbid., 10: 1-140, 1858 (including
supplements).

HokK, C. VAN DEN. 1963. Revision of the European species of
Cladophora. E. J. Brill, Leiden. 248 pp.

420 Rhodora [Vol. 76

KORNMANN, P. 1962. Eine Revision der Gattung Acrosiphonia.
Helgolànd. Wiss. Meeres. 8: 219-242.

Kuckuck, P. 1956. Ectocarpaceen-Studien IV. Herponema, Kutz-
ingiella nov. gen., Farlowiella nov. gen. herausgegeben von Peter
Kornmann. Helgolànd. Wiss. Meeres. 5: 292-325.

1960. Eectocarpaceen-Studien VI. Spongonema. Helgo-
lànd. Wiss. Meeres. 7: 93-113.

KvLIN, H. 1947. Ueber die Fortpflanzungsverhaltnisse in der Ord-

nung Ulvales. K. Fys. Sallsk. Lund Fórh. 17: 174-182.
1947a. Die Phaeophyceen der Schwedischen Westküste.
Lunds Univ. Arsskr. N. F., Avd. 2, 43(4) : 1-99.

LorsEAUX, S. 1967. Morphologie et cytologie des Myrionémacées.
Critères taxonomiques. Rev. Gén. Bot. 74(878): 329-947.

Lunp, S. 1959. The marine algae of East Greenland, I. Taxo-
nomical part. Medd. Gronland 156(1): 1-248.

NEwRoTH, P. R., & A. R. A. TAYLOR. 1971. The nomenclature of
the North American species of Phyllophora Greville. Phycologia
10(1): 93-97.

Norton, T. A. 1972. The marine algae of Lewis and Harris in the
Outer Hebrides. British Phycol. J. 7(3): 375-385,

OLNEY, S. T. 1847. Rhode Island plants, 1846, or additions to the
published lists of the Providence Franklin Society. Proc. Provi-
dence Franklin Soc. 1(2): 25-42.

1871. Algae Rhodiaceae. A list of Rhode Island algae,
collected and prepared by Stephen T. Olney, in the years 1846-
1849, now distributed from his own herbarium. Published by
the author, printed by Hammond, Angell & Co., Providence.
13 pp. (Exsiccata, few sets).

PARKE, M. 1953. A preliminary check-list of British marine algae.
J. Mar. Biol. Ass. U. K. 32: 497-520.

PARKE, M., & P. S. Dixon. 1964. A revised check-list of British
marine algae. J. Mar. Biol. Ass. U. K. 44: 499-542.

1968. Check-list of British marine algae — second re-
vision. J. Mar. Biol. Ass. U. K. 48: 783-832.

PowELL, H. T. 1963. Speciation in the genus Fucus L. and related
genera. In: J. P. Harding & N. Tebble (ed.), Speciation in
the sea. Syst. Ass. Publ. 5: 63-77.

RosENvINGE, K. L. 1924. The marine algae of Denmark. Vol. III.
Rhodophyceae. III. (Ceramiales). K. Danske Vidensk. Selsk.,
Ser. Nat. 7: 287-486.

ScaGeL, R. F. 1966. Marine algae of British Columbia and northern
Washington. Part I: Chlorophyceae (green algae). Nat. Mus.
Can., Biol. Ser. No. 74, Bull. 207: 1-257.

SOUTH, R., & A. CARDINAL. 1970. A check list of marine algae of
eastern Canada. Can. J. Bot. 48: 2077-2095.

1974] R. I. Marine Algae — Wood & Villalard-Bohnsack 421

SUNDENE, O. 1958. Interfertility between forms of Laminaria digi-
tata. Nytt. Mag. Bot. 6: 121-128.

TAYLOR, W. R. 1937. Marine algae of the northeastern coast of
North America. Univ. Michigan Press, Ann Arbor. 427 pp.

1957. Ibid., 2nd ed. 509 pp.

TAYLOR, W. R., & C. F. RHYNE. 1970. Marine algae of Dominica.
Smithsonian Contrib. Botany 3: 1-16.

VAUGHAN, C. F. 1971. An ecological survey of the macroscopic
algae of Point Judith Pond (Salt Pond), Rhode Island. M. S.
Thesis, University of Rhode Island. 179 pp.

VILLALARD, M. 1971. Phytosociology of intertidal algae of Conani-
cut Island, Rhode Island. Ph.D. Thesis. University of Rhode
Island. 227 pp.

WiLCE, R. T. 1965. Studies in the genus Laminaria. III. A re-
vision of the North Atlantic species of the Simplices section of
Laminaria. Bot. Goteborg. 3: 247-256.

Woop, R. D. 1962. Codium is carried to Cape Cod. Bull. Torrey
Bot. Club 89: 178-180.

————— ————. (in press). Information for a list of marine algae of
Rhode Island. New England Marine Resources Information Pro-
gram, University of Rhode Island, Kingston.

Woop, R. D, & E. A. PALMATIER. 1954. Macroscopic algae of the
coastal ponds of Rhode Island. Am. J. Bot. 41(2): 135-142.

WYNNE, M. J., & W. R. TAYLOR. 1973. The status of Agardhiella
tenera and Agardhiella baileyi (Rhodophyta, Gigartinales).
Hydrobiologia 43(15): 93-107.

R. D. WOOD
DEPT. OF BOTANY

UNIVERSITY OF RHODE ISLAND
KINGSTON, RHODE ISLAND 02881

M. VILLALARD-BOHNSACK
DEPT. OF BIOLOGY

ROGER WILLIAMS COLLEGE
BRISTOL, RHODE ISLAND 02809

A BIOSYSTEMATIC REVISION OF
LUPINUS LYALLII

B. J. Cox!

Systematics of the genus Lupinus is often viewed as one
of the most perplexing studies ever attempted. This is
attested by the diverse taxonomic treatments of various
floras and special publications. While several students of
the lupines have recognized many entities, others have
found these plants in such confusing mosaics that enormous
synonymic reductions were made; neither approach has
solved the taxonomic problems formed by lupine diversity.

In a biosystematic revision of the Lupinus lepidus-L.
caespitosus complex of the Sierra Nevada, Cascade and
Rocky Mountains, various techniques of study were em-
ployed in an effort to show the degree of relationship be-
tween the taxa (Cox 1972a). Over 75 epithets have been
applied to various members of this complex since 1828
when L. lepidus was described, and these epithets have
been considered by some as referable to a single species.

In order to present the concept of the L. lepidus-L. caes-
pitosus complex in a digestible and economically feasible
form, a taxonomic treatment of only Lupinus lyallii, its five
subspecies and six varieties will be offered here. More
complete distribution maps, cytological and chemotaxonomic
data, photographs of typical specimens, and illustrations of
typical leaflet and floral parts are given elsewhere (Cox
1972a, 1972b, 1973a, 1973b).

Chromosome numbers: Taxa within Lupinus lyallii have
chromosome number reports of 2n — 48 (subsp. lyallii, var.
lyallii and var. villosus, and subsp. minutifolius) by Cox
(1972b). The chromosomes are extremely small and counts
are difficult to obtain, undoubtedly explaining the lack of
previous reports. Although Phillips (1957) reported a

'Studies conducted at the Division of Biological Sciences, University
of Missouri, Columbia, Missouri.

422

1974] Genus Lupinus — Cox 423

chromosome count for L. lyallii of n = 24, his study plant
was L. minimus. Chromosome counts have offered little in
determining the genetic affinities within the L. lepidus-L.
caespitosus complex since all reported numbers are 2n — 48
(Cox 1972b).

Chemosystematic data. Seed proteins: Comparisons were
made of seed proteins of four of the L. lyallii taxa by poly-
acrylamide gel electrophoresis (Cox 1972a, 1973b). Con-
siderable variation was found in several polymorphic popu-
lations; largely, the electrophoresis disc protein data closely
resemble those data from the morphological study in the
degree of variation found. Although no protein band oc-
curred in common with all taxa studied, likewise no pro-
tein band was found unique to any one taxon. Each popu-
lation was found to have a distinct combination of proteins,
resulting in low percent similarity values between the taxa.

Seed Alkaloids: The seed alkaloids of all of the sub-
species and varieties of Lupinus lyallii were compared
by thin-layer chromatography (TLC) separation by Cox
(1973a). Although some interpopulational variation was
found, the alkaloids were generally quite consistent. Corre-
lations were made between alkaloid shapes, R: values, posi-
tive color reactions with two different visualizing agents,
and comparisons with standards. Percent similarity values
were high among the interspecific taxa, and identical alka-
loid combinations were shared among a few, these differing
only in relative amounts of individual alkaloids.

Morphological and. distributional considerations: Field
and herbarium-laboratory studies were conducted over a
three year period on the Lupinus lepidus-L. caespitosus
complex. Techniques and preparations of specimens for
measurements in the descriptions are given by Cox (1972a).

The following herbaria are acknowledged for the use of
over 1000 specimens of the taxa of Lupinus lyallii: AEC,
ARIZ, BRY, CAN, DS, GH, ISC, JEPS, MIN, MO, NMC, NY, POM,
ORE, OSC, RENO, RM, RSA, SMU, UBC, UC, UMO, US, UTC, V,
WILLU, WS.

424 Rhodora [Vol. 76

Lupinus lyallii A, Gray

Plants perennial, herbaceous, caespitose or mat-like,
less than 18 cm. tall; leaflets few (5-9), oblanceolate, the
largest less than 2 cm. long; banners glabrous, usually re-
flexed below the midpoint.

Distribution: The taxa are primarily found in the high
Sierra Nevada and Cascade Mountains with a few popula-
tions radiating out into isolated localities (Fig. 1).

KEY TO THE SUBSPECIES AND VARIETIES OF LUPINUS LYALLII

A. Plants with spreading villous-hirsute to shaggy pubes-
CENCE. aooaa lee. B.

B. Plants lees than 10 cm. tall; flowers 6-7 mm. long;
largest leaflets 4-6 mm. long; Sierra Nevada, alpine,

dwarf. ...... 1b. L. lyallii subsp. lyallii var. danaus.
B. Plants usually taller; flowers over 7 mm. long; larg-
est leaflets over 6 mm. long. ............ . C.

C. Plants decumbent, mat-like; leaflets 6- T, largest
10-13 mm. long; Sierra Nevada, with a few sta-
tions in Klamath Co., Ore.

. lf. L. lyallii subsp. lyallii var. villosus.

C. Plants erect to semi-decumbent, branching, 10-18
cm. tall; leaflets 5-6, largest 9-21 mm. long;
Washoe Co., Nevada. ...... ,

ZEE . 9. L. lyallii subsp. washoensis.
A. Plants with appressed to subappressed pubescence. .. D.
D. Plants erect in densely branching clumps; 12-18 cm.

tall; Rogue River-Crater Lake region, Oregon. .. E.

E. Branching stems mostly woody, less than 1.5 mm.
in diameter; leaflets 5-6; racemes 1.3-1.4 cm. in
diameter ; flowers 6-7.5 mm. long.

. 1c. L. lyallii subsp. lyallii var. fructiculosus.

E. Branching stems mostly herbaceous, more robust;
leaflets 7-9; racemes 1.9-2.5 cm. in diameter;
flowers 7-9.5 mm. long. ................. s.
e le. L. lyallii subsp. lyallii var. roguensis.

D. Plants decumbent or erect and densely caespitose.

1974] Genus Lupinus — Cox 425

F. Plants mat-like, usually less than 10 cm. tall;
flowers 6-7 mm. long; largest leaflets 4-6 mm.
long; alpine, Sierra Nevada Range. .........-
Mero Rd 1b. L. lyallii subsp. lyallii var. danaus.

F. Plants usually larger or flowers longer. ..... G.
G. Plants 4-6 cm. tall; flowers 7.5-10 mm. long;

largest leaflets 6-9 mm. long, upper surface
glabrous; Steens Mts, Harney Co., Oregon.
TM IO ay e «= 3. L. lyallii subsp. minutifolius.
G. Plants mostly taller; flowers various; largest
leaflets various lengths, mostly pubescent
(becoming somewhat glabrate in subsp. alcis-
temporis). occ ccc ecw cece gien p hh n H.
H. Plants mostly decumbent; Clearwater Co.,
Idaho. .. 2. L. lyallii subsp. alcis-temporis.

H. Plants erect or mat-like, not decumbent.
I Leaves all basal, erect, in small tufts;
petioles 3-9 em. long; largest leaflets

9-22 mm. long; SW Idaho & NE Ne-
vada. .. 4. L. lyallii subsp. subpandens.

I. Leaves cauline, basal, or both, generally
ascending or more or less spreading,
general not in small tufts; petioles
usually 2-5.5 em. long; largest leaflets
CEPEOGOEG o ni PTT J.

J. Flowers 10-12.5 mm. long; racemes
2.5-4.5 (5.5) cm. long, 2.5-3.0 cm.

in diameter ; leaflets 6-9; high alpine

areas of western Cascade Range,

Öre & Wash UU b e d nih s 1d. L.

lyallii subsp. lyallii var. macroflorus.

J. Flowers 7.5-10.5 mm. long; racemes

1.5-3 em. long, 2.0-2.3 cm. in di-
ameter; leaflets 5-7; alpine and sub-

alpine, wide distribution, Sierra Ne-

vada Range, Calif. & Cascade Range,

426 Rhodora [Vol. 76

Ore., Wash. & B. C. .......... la.
L. lyallii subsp. lyallii var. lyallii.

la. Lupinus lyallii subsp. lyallii var. lyallii

Lupinus lyallii A. Gray, Proc. Amer. Acad. 7: 334.
1868. Type: “Summit of the Cascade Mountains, lat 49°,
elev. 7500 ft.”, Lyall, Jul 1860 (Holotype: K; Isotypes:
BM, GH, POM). Photos: (RSA, UC, UMO, US).

L. lepidus subsp. lyallii (Gray) Detl., Amer. Mid. Nat.
45: 490-491. 1951.

L. alpinus Heller, Muhlenbergia 6: 22. 1910. Type:
NEVADA: WASHOE CO.: Mt Rose, Heller 9914a (Holo-
type: Nev. Agr.; Isotypes: GH, ISC, MIN, NMC, NY, POM,
RSA, UC, US, WS). Photos: (RSA, UMO).

L. paulimus Greene, Leafl. Bot. Obs. & Crit. 2: 234.
1912. Type: OREGON: DESCHUTES CO.: Paulina Lake,
W end, Leiberg 550 (Holotype: US; Isotypes: GH, UC).

L. rubro-soli Eastw., Leafl. West. Bot. 3: 203. 1941.
Type: CALIFORNIA: MENDOCINO CO.: Red Mt., Eastwood
& Howell 4674 (Holotype: CAS; Isotype: US). Photos:
(RSA, UMO).

Plants perennial, caespitose, forming mat-like radicle
clumps, 15-25 cm. in diameter, from a thick, woody caudex
which often branches in older plants, 6-17 cm. tall; foliage
2-7 cm. tall; stems poorly developed, less than 1.5 mm. in
diameter, semi-prostrate, some sprawling-decumbent, some
erect, subappressed silky sericeous throughout, hairs usually
less than 1 mm. long; cauline nodes 0-2; petioles slender,
2-5(7.5) cm. long; stipules generally inconspicuous; leaf-
lets 5-7, oblanceolate-linear to elliptic-obovate, apex acute to
short mucronate, largest 5-12(18) mm. long, av. 9.95 mm.,
1.5-4.5 mm. wide, length to width ratio 2.17-4.0(6.0), av.
3.18, appressed silky-sericeous on both surfaces; peduncles
usually much exceeding the foliage, 3-8(10) cm. long;
racemes capitate to subcapitate, 1.5-3 em. long, occasionally
to 4.0 cm. at anthesis, 20-23 mm. in diameter, rarely 2X
as long as wide; flowers dense, blue, 7.5-10.5 mm. long, av.
9.44; bracts persistent to caducous, 3-6.5 mm. long, subu-

1974] Genus Lupinus — Cox 427

late to lance-linear; pedicels 1.5-2.5 mm. long; calyx bi-
labiate, the upper-lip 3.5-6.0 mm. long, bilobed, the notch
1-2.5 mm. deep, the lower-lip 3.8-6.8 mm. long, apex serrate
to tridentate, the center tooth to 0.5 mm. long, lips connate
1-2 mm., bracteoles usually present but inconspicuous, to
0.5 mm. long, attached at the sinus, sericeous; banner ellip-
tic-oblong to suborbicular, glabrous, 7-10 mm. long, 5.1-8.0
mm. wide, length to width ratio 1.10-1.57, av. 1.29, reflexed
below or near the midpoint, the reflexed to appressed ratio
1-1.3, av. 1.14, angle 120-150°; wings 6.5-9.8 mm. long, 3.5-
5.5 mm. wide, length to width ratio 1.56-2.25 av. 1.90, claws
1.0-2.2 mm. long, glabrous; keel 2.5-3.0 mm. wide at the
widest point, angle 105-111°, ciliated along the upper mar-
gin near the acumen; ovules 3-5; pods 10-15 mm. long, 4-5
mm. wide, short sericeous; seeds 2-4, 1.8-3.0 mm. X 2.6-3.5
mm., 1-1.5 mm. thick, tan, beige, brown and green mottled.

This variety is primarily alpine to subalpine, ranging
latitudinally in its distribution from the high Sierra Nevada
Mountains of California through the Cascade Range of
Oregon and Washington into British Columbia. These
plants may be found in several soil types, such as volcanic
ash, disintegrated granite, sandy, gravelly rock of several
compositions to shale. They usually occur on slopes associ-
ated most frequently with Pinus contorta, P. murrayana,
P. albicaulis and Tsuga mertensiana. Altitudinally, the
plants are found above timberline at 3660 m. down to about
2439 m., or in the more northerly latitudes to about 1830 m.
Flowering occurs between June and September; the plants
flowering during the latter part of the season usually are
found in sites of lingering snowbanks.

Representative specimens: CANADA: British Columbia: Bomford
Mt, Ashnola Dist, Barr 9351 (UBC); Chileotin, Copley 472 (v);
Lillooet, Anderson, Jul 1916 (v); Manning Park, Hardy, Aug 1945
(v); Mt McLean, Anderson 7987 (v); Mt Moor, Bird 3436 (UBC) ;
Nicoamen River head, Dawson 51833 (CAN); Noaxe Lake, Brink, Aug
1957 (UBC); Yalakom River, Sugden 118 (v). USA: California:
ALPINE CO.: Carson Pass, Yates 3980 (UC). EL DORADO CO.: Desolation
Valley, Abrams 12740 (GH, NY, POM, UC). FRESNO CO.: Humphreys

428 Rhodora [Vol. 76

Basin, headwaters Piute Creek, Quibell 5118 (UC). LASSEN CO.:
Lassen Volcanic Natl. Park, Rickett, Aug 1968 (UMO). MADERA CO.:
Mt Lyell, at Moraine Meadows, Akey 318 (UC). MARIPOSA CO.: Mt
Connes, Tioga Pass, Ramsey & Ramsey 2496 (POM). MODOC CO.:
Antelope Valley, Applegate 847 (US). MONO CO.: Bodie, 2.2 mi
N-NW, Graham 274 (UC). NEVADA CO.: Mt Stanford, Sonne 54
(MO). PLACER CO.: Snow Mt, Bolt 103 (UC). SHASTA CO.: Bolam
Road, near the S.P. RR, Cooke 15003 (uc). SISKIYOU CO.: Bolam
Creek Canyon, opposite Coquette Falls, Mt Shasta. Cooke 25691
(UC). TEHAMA CO0.: King Creek Meadow, Lassen Volcanic Natl.
Park, Heller 15426 (Mo, NY). TRINITY CO.: Fisher Ridge, E of
Black Rock, North Yolla Bolly Mts, Munz 16743 (RSA). TULARE CO.:
Golden Trout Creek, Hall & Hall 8417 (uc). TUOLUMNE CO.: Bond
Pass, Winkelman 276 (Uc). Nevada: WASHOE CO.: Mt Rose, Ken-
nedy 1148 (NY, UC, US). Oregon: CURRY CO.: Bear Camp Ranger
Station, Leach 1692 (ORE). DESCHUTES CO.: Bend, Farnham & Beach
97 (OSC). DOUGLAS CO.: Bailey Mt, Applegate 4121 (uc). JACKSON
co.: Ashland Peak. Thompson 12383 (MO, NY, UC). JFEFFFPSON CO.:
Hoodoo, Big Lake Turnoff, US Hwy 20, Isely & Isely 11299 (ISC,
UMO). JOSEPHINE CO.: Free and Easy Trail, head of Babyfoot,
Leach 2661 (ORE). KLAMATH CO.: btn Fort Klamath & Summit,
Furlong et al, 1901 (UC). CRATER LAKE NATL. PARK: Summit Lake,
Henderson 9634 (ws). LAKE CO.: Drews Valley. Austin 1649 (Us).
LANE CO.: McKinzie Pass, Peck 5467 (WILLU). LINN CO.: Bie Lake,
Hawley & Anderson, Jul 1927 (osc). WALLOWA CO.: Wallowa Mts,
Aneroid, 2 mi S, Peck 17985 (uc). Washington: CHELAN CO.: Blewett
Pass region, Grant, Aug 1929 (UC). CLALLAM co: Hurricane Ridge,
Thompson 14226 (GH, NY, US). JEFFERSON CO.: Tron Mt, Thompson
11049 (MO. NY, POM. US). OKANOGAN CO.: Bunker Hill summit,
Edwards 32 (ws). PIERCE co.: Mt Rainier Natl. Park: Flett’s
Glacier, Flett 3124 (ws). wHATCOM co.: Mt Baker. Heliotrope
Ridge, Muenscher 8201 (GH, UC. US. WS). YAKIMA CO.: Hell Rearing
Canyon rim, Langdon 116 (osc).

The populations from the Wallowa Mountains (Wallowa
Co., Oregon) are isolated from other Lupinus lyallii subsp.
lyallii populations by 322 km.; the subsp. alcis-temporis
occupies areas along the Clearwater and Selway Rivers in
Idaho, but it is quite distinctive. The growth habit of the
Wallowa populations appears to be typically that of var.
lyallii specimens in the normal distribution, with short
vegetation to 7 cm. tall, capitate inflorescences 2-2.5 cm.
long, small flowers 6-8 mm. long and short leaves 7-10 mm.
long. But, these plants hold in common with L. minimus a

1974] Genus Lupinus — Cox 429

strong woody branching caudex at above ground level, many
stems in a clump with leaves appearing basal, not more
than 2-3 cauline leaves, and silky pubescence.

The specimens of Lupinus lyallii subsp. lyallii var. lyallii
in Deschutes Co., Oregon (Leiberg 550 — GH, UC, US), de-
scribed as L. paulinus by Greene, are somewhat larger than
other L. lyallii plants of the immediate region but closely
resemble several populations found in California. The
racemes of some plants range from 3-4 cm. to 7.5 cm. long
and from 16 mm. to 22 mm, in diameter, with the leaflets
11-15 mm. long, the upper surface of the leaflets sparsely
pubescent, flowers approaching 9 mm. long and peduncles
6-7 cm. long. Several plants resemble L. sellulus var. lobbii
with the flowers scattered in verticils somewhat like L.
sellulus var. medius (Cox 1973c). However, the population
sample indicates that these variants are more closely
related to L. lyallii subsp. lyallii var. lyallii than to any
other taxon and not distinctive enough to warrant a sepa-
rate epithet.

The material found on Mt. Rose in Nevada, which Heller
called Lupinus alpinus, flowers from late July to late Sep-
tember and occurs at elevations of 2927-3110 m.

The plant habit is low and spreading mats, much like
that on Mt. Rainier in Washington. Colonies occur in
grassy open meadows, in the alpine belt. Although the
plants may be somewhat atypical of the California L. lyallii
subsp. lyallii var. lyallii specimens, they are much like those
of Oregon and Washington.

1b. Lupinus lyallii subsp. lyallii var. danaus (A. Gray) S.

Wats., Proc. Amer. Acad. 8: 534. 1873.

L. danaus Gray, Proc. Amer. Acad. Arts & Sci. 7: 335.
1868. Type: CALIFORNIA: TUOLUMNE CO.: Mt Dana,
Bolander 5087 (Holotype: GH; Isotype: UC, US).

L. danaus var. bicolor Eastw., Leafl. West. Bot. 2: 201.
1939. Holotype: CALIFORNIA: TUOLUMNE C0.: Mt Dana,
Howell 14539 (CAS). Photos: (RSA, UMO).

L. oreocharis Eastw., Leafl. West. Bot. 3: 171. 1942.

430 Rhodora [Vol. 76

Holotype: CALIFORNIA: TULARE CO.: Little Five Lakes
Basin, Howell 17392 (CAS). Photos: (RSA, UMO).

Plants differing from L. lyallit subsp. lyallii var. lyallii
by matted vegetative growth, 1-4 cm. tall; plant overall
height usually less than 10 cm. tall; pubescence frequently
sparse, spreading; largest leaflets mostly 4-6 mm. long,
rarely to 10 mm. long; peduncles 1.5-3 em. long, rarely to
8 em. long; racemes capitate, 1.5-2.0 cm. long; flowers fre-
quently more loosely arranged than in var. lyallii, smaller,
6-7 mm. long.

Representative specimens: California: FRESNO C0.: Humphreys
Basin, headwaters Piute Creek, W of Mt Humphreys, Sharsmith 3136
(uc). INYO co.: Piute Pass, E slope, Peirson 2552 (RSA). MADERA
co.: Isberg Pk, 1 mi SW, Akey 332 (UC). MARIPOSA CO.: Mt Dana,
Congdon, Aug 1898 (Uc). MODOC CO.: Mt Warren, Payne 307
(JEPS). MONO C0.: San Juaquin Peak, Wright K3 (POM). TULARE
co.: Mt Whitney, Clemens, Jul 1910 (POM, UMO). TUOLUMNE CO.:
Gaylor Lakes, Howell 20401 (ws).

The depauperate forms of L. lyallii subsp. lyallii are
interpreted here as being ecological variants. Under a par-
ticular set of environmental conditions populations of L.
lyallii dwarfs to the matted form of var. danaus. However,
frequently within the same population typical var. lyallii
may occur in the larger form. Both varieties are found
in high alpine rocky, sunny areas. Although this depau-
perate or ecological variant of L. lyallii appears to be
morphologically blending with the larger variety, it is
considered here as a distinct taxon at the varietal level.

A variant of similar vegetative morphology was observed
among the Lupinus lyallii specimens of Canada and the
higher alpine areas of Washington and Oregon, but the
reproductive structures differ from that of var. danaus in
California.

Lupinus oreocharis is reduced to synonymy under var.
danaus. Although some of these plants range to 14 cm. tall
and resemble var. villosus in pubescence, and some may
differ from var. danaus by having slightly elongated stems

1974] Genus Lupinus — Cox 431

which may have 1-2 cauline leaves, the available evidence
does not indicate the warrant of a special epithet.

1c. Lupinus lyallii subsp. lyallii var. fruticulosus (Greene)

C. P. Sm., Bull. Torrey Bot. Club 51: 303. 1924.

L. fruticulosus Greene, Muhlenbergia 8: 117. 1912.
Type: OREGON: KLAMATH CO.: Anna Creek Valley,
Austin 1650 (Holotype: US; Isotypes: NY, POM). Photos:
(RSA, UMO, US).

L. perditorum Greene, Muhlenbergia 8: 117. 1912.
Holotype: OREGON: JACKSON CO.: Upper Rogue River,
Applegate 2636 (US). Photo: (US).

Plants in dense semi-shrubby clumps, branching pro-
fusely, erect to semi-decumbent; stems 19-17 em: tall,
usually less than 1.5 mm. in diameter; leaves cauline;
petioles slender, 2-5 cm. long; leaflets 5-6, elliptic-oblanceo-
late, largest 5-12 mm. long, av. 8.4, 1.8-3.5 mm. wide, length
to width ratio 2.78-4.0, av. 3.42, subappressed, strigose-
pilose on both surfaces; peduncles 2.5-5.5 em. long; racemes
numerous, 1.5-4.3 em. long, 13-14 mm. in diameter, capitate
and dense to somewhat scattered; flowers blue, 6.0-7.5 (8.0)
mm. long, av. 6.9 mm.; bracts 1.5-3.0 mm. long; pedicels
0.5-1.5 mm. long; banner elliptic to suborbicular, reflexed
below the midpoint; pods 7.5-10 mm. long, 3.3-4 mm. wide;
seeds 2-3.

The greatest development of var. fruticulosus is along
the flood plain of the Rogue River in Klamath Co., Oregon.
Plants are found at elevations of 1067-1372 m., flowering
in July.

Representative specimens: Oregon: JACKSON CO.: Ashland, 9.7 km
S, Sherwood 842 (WILLU). KLAMATH CO.: Crater Lake Natl. Park,
Evinger 608 (osc). LINN CO.: Hunts Cove, Mt Jefferson area,
Leach 4485 (ORE).

Variety fruticulosus differs morphologically from other
variants of Lupinus lyallii by its semi-shrubby base, pro-
fusely branching clump habit and smaller flowers. A few
L. lyallii specimens from Washington (mainly var. macro-

432 Rhodora [Vol. 76

florus) have thick, woody branching bases, but these are
part of the underground caudex system and differ by having
distinctively larger inflorescence structures. The Washing-
ton material is more robust than fruticulosus, Variety
fruticulosus is one of the most distinctive variations of
L. lyallii subsp. lyallii.

Variety fruticulosus is morphologically similar to var.
roguensis, of the same general geographical region, and
to subsp. alcis-temporis of the Clearwater and Selway
Rivers in Idaho. It may be separated from these two taxa
on the basis of its flower morphology, more scattered flower
arrangement on the racemes and smaller size, as well as
differences in vegetative traits. These plants exhibit a
more erect woody branching habit with more filiform
petioles and stems than any closely related entities. They
are generally not as leafy, and the leaflets are fewer (5-6
per leaf), smaller, and arranged in the upper portion of
the branching clump.

1d. Lupinus lyallii subsp. lyallii var. macroflorus B. J. Cox,
var. nov. Type: WASHINGTON: PIERCE CO.: Mt Rainier,
Yakima Park, Thompson 11086 (Holotype: MO; Isotypes:
NY, POM-in part, US). Photo of holotype: (Cox 1972 a).

Plantae perennes, humiles; caules ex caudice profuse
ramificanti lignoso, 5-17 cm. alti, 1-15 mm. diametro, nodis
0-2(8); petioli 3-5.5 em. longi; foliola 6-9, oblanceolata,
maxima 5.5-11.3 mm. longa, 2.5-5.0 mm. lata; pedunculi
3.0-5.5 em. longi, supra folia 1-2 em, extensi; racemi 2.5-5.5
cm. longi, 2.5-3.0 cm. diametro, capitati usque subcapitati,
floribus 5-17 in 1-5 verticillis; flores caerulei 10-12.5 mm.
longi; calyx gibbosus; vexillum suborbiculatum usque obo-
vatum ; ovula 3-5; legumina 13-15 mm. longa, 4-5.5 mm. lata.

Plants perennial, in dense low mats; stems from a pro-
fusely branching woody base, 5-17 cm. tall, 1-1.5 mm. in
diameter, pubescence appressed to ascending, some spread-
ing, cauline nodes 0-2(8); petioles 3-5.5 em. long; leaflets
6-9, oblanceolate, largest 5.5-11.3 mm. long, av. 8.29 mm.,
2.5-5.0 mm, wide, av. 2.56 mm. ; peduncles 3.0-5.5 cm. long,

1974] Genus Lupinus — Cox 433

exceeding the foliage by 1-2 cm. or more; racemes 2.5-5.5
em. long, 2.5-3.0 cm. in diameter, capitate to subcapitate,
flowers 5-17 in 1-5 verticils; flowers blue, 10-12.5 mm. long,
av. 11.2 mm. long; calyx gibbous at the base; banner sub-
orbiculate to obovate, glabrous; keel densely ciliated near
the acumen to glabrous in some; ovules 3-5; legumes 13-15
mm. long, 4-5.5 mm. wide.

Populations of var. macroflorus are found on the western
side of the Cascade Mountain Range on both sides of the
Columbia River in Washington and Oregon. They are found
on Mary’s Peak in Benton Co., on Mt. Hood of Clackamus
and Hood River Counties, Santiam Pass in Linn Co., Ore-
gon, and Mt. St. Hellens of Skamania Co., and Mt. Rainier
of Pierce Co., Washington. Like var. lyallii, var. macro-
florus occurs at the upper vegetational limit between tim-
berline and snowline at elevations of 1524-2378 m. Plant
communities commonly associated include the species:
Pinus contorta, P. albicaulis, Abies lasiocarpa, Tsuga. mer-
tensiana and less frequently Pseudotsuga menziesii.

Representative specimens: Oregon: BENTON CO.: Corvallis, W at
Mary's Peak, Gilbert 108 (osc, US). CLACKAMUS CO.: Mt Hood, Jct
of Hwys 26 & 35, Downey 4B (UMO, UTC). HOOD RIVER CO.: Mt Hood,
Abrams 9445 (MO, POM, RM, WILLU). LINN CO.: Santiam Pass,
Cronquist 7853 (GH, NY, RSA, UC, WS). Washington: PIERCE CO.:
Mt Rainier Natl. Park, Burrough’s Mt & Frozen Lake, Flett 3121
(WS). SKAMANIA CO.: Mt St Hellens, Buzzettii, Aug 1848 (osc).

Variety macroflorus is distinct primarily on the basis of
its larger flower size (10-12.5 mm. long) from var. lyallii
flower size (7.5-10.5 mm. long). There are several inter-
mediates between the variations of L. lyallii, but in general
var. macroflorus is the extreme in flower morphological
deviation.

le. Lupinus lyallii subsp. lyallii var, roguensis B. J. Cox,
var. nov. Type: OREGON: JACKSON co.: Farewell Bend
Forest Camp, 1.6 km N of Union Cr, on Rogue River flood
plain, Hitchcock & Martin 4997 (Holotype: RSA; Isotypes:
MO, NY, UC, WS). Photo of isotype: (Cox 1972a).

434 Rhodora [Vol. 76

Plantae perennes, herbaceae, caespites densos ramificantes
formantes, 2.5-3.5 dm. diametro, 12-18 cm. altae; caules
humifusi usque erecti, 1-3 mm. diametro, ex caudice dense
ramificanti lignoso, nodis 6-10, ad nodos ramificantes ; foliola
7-9, maxima 8-13 mm. longa, 2.5-3.0 mm. lata; racemi capi-
tati usque subcapitati 1.5-3.5 mm. longi, 4.5 mm. longi
fructiferi, 1.9-2.5 mm. diametro; flores 7-9.5 mm. longi;
vexillum et alae glabrae.

Plants perennial, herbaceous, in dense branching clumps,
2.5-3.5 dm. in diameter, 12-18 cm. tall; stems procumbent
to erect, 1-3 mm. in diameter, from densely branching
woody base, nodes 6-10, branching at the nodes; leaflets
7-9, largest 8-13 mm. long, 2.5-3.0 mm. wide; racemes capi-
tate to subeapitate, 1.5-3.5 mm. long, 4.5 mm. long in fruit,
1.9-2.5 mm. in diameter; flowers 7-9.5 mm. long; banner
and wings glabrous; keel ciliated along the upper margin
near the acumen.

Variety roguensis is found at Diamond Lake, Huckleberry
Mt., Davis Lake, and on the flood plain of the Rogue River,
in Jackson and Klamath Counties, Oregon, These plants
occur in sandy soil in pine forests at elevations of 915-1067
m. Flowering is from mid-June to mid-July.

Representative specimens: Oregon: JACKSON CO.: Huckleberry
Mt, Rogue River Natl. Forest, Thompson 130835 (MO, RSA, UC, WILLU,
WS). KLAMATH CO.: Davis Lake, W shore, Hitchcock & Martin 4935
(RSA).

Morphologically, these plants are most closely related to
var. fructiculosus, but they differ in several significant
characteristics. They are more robust, have larger branch-
ing procumbent to erect stems, have larger flowers that
are more densely arranged in racemes, and have more
leafy foliage. The leaves are more numerous and are scat-
tered throughout the clump, and the leaflets are larger
with more leaflets per leaf. The manner of branching re-
sembles that of L. aridus, especially subspecies lenorensis
(Cox 1972a) found along the Clearwater and Selway Rivers
in Idaho, Although the general branching pattern is often

1974] Genus Lupinus — Cox 435

difficult to detect, careful scrutiny finds verticillate spread-
ing branching from an erect shoot; these branches become
somewhat decumbent, but secondary and tertiary branches
arise from them either adaxially or twisting around the
decumbent stems and growing upward forming erect
clumps.

Several color variations are found in this segment of
Lupinus lyallit. Many plants of var. roguensis have albino
flowers (white to cream color — Maguire & Holmgren 26493
— GH, NY, UC, US) ; however, others within the same popu-
lation are intermediate in flower color to the albino and
typical blue form (white to cream colored petals except for
the purple banner and keel tip — Maguire & Holmgren
26494 — GH, MO, NY, UC, US, WS).

1f. Lupinus lyallii subsp. lyallii var. villosus Jeps., Jepson,
Fl. Calif. 2: 267. 1936. Holotype: CALIFORNIA: ALPINE
CO.: Carson Pass, Jepson 8100 (JEPS).

L. hypoleucus Greene, Leafl. Bot. Obs. & Crit. 1: 74.
1904. Type: CALIFORNIA: White Chief Peak, Culbert-
son 16, Jul 1904 (distributed by Baker 4416 — specimen
not located.) Paratype: CALIFORNIA: Mt, Goddard,
Hall & Chandler 707 (US).

Plants densely caespitose; stems decumbent to semi-erect,
from a thick branching woody caudex, sericeous with as-
cending to long spreading white hairs turning reddish-
brown with age; petioles 4-5 cm. long; leaflets 6-7, elliptic-
oblanceolate, apex acute-mucronate, largest 10-13 mm. long,
3-5 mm. wide, villous on both surfaces; peduncles 3-7 cm.
long; racemes densely verticilate 2.5-5 cm. long; bracts
persistent, 4-6 mm. long; pedicels 1-1.5 mm. long; banner
and wings glabrous; keel ciliate near the acumen; pods
10-11 mm. long, 4 mm. wide.

This variety occurs scattered throughout much of the
distributional range of L. lyallii in several counties of
California in the Sierra Nevada northward to several popu-
lations in Klamath Co., Oregon.

436 Rhodora [Vol. 76

Representative specimens: California: ALPINE CO.: Carson Pass,
Higgins 1752 (BRY). FRESNO CO.: Bubbs Creek, Peirson 531 (RSA).
HUMBOLDT CO.: Grouse Mt, back of Clear Lake, Tracy 16672 (UC).
LASSEN C0.: Grass Valley, Fisher & Johnson F203 (UC). MADERA
co.: Garnet Lake, Howell 16421 (RSA, UC, US). MARIPOSA CO.:
Donohue Pass, Schreiber 1767 (UC). MENDOCINO CO.: South Yolla
Bolly, Jepson 57t (JEPS). MONO CO.: San Juaquin Mt, Howell
16875 (Ws). PLACER CO.: Mt Lincoln, Howell 18670 (US). SHASTA
Yolla Bolly, Jepson 57t (JEPS). MONO C0.: San Juaquin Mt, Howell
16875 (ws). PLACIER CO.: Mt Lincoln, Howell 18670 (US). SHASTA
co.: Squaw Valley Creek Meadows, Cooke 13651 (NY). TRINITY CO.:
Trinity Center, 3 mi S, E of Scott Ranch, Cantelow, Aug 1936 (RSA).
TUOLUMNE C0.: Gaylor Lakes, Howell 20418 (GH). Oregon: KLA-
MATH CO.: Llao Rock, Baker 6245 (NY, OSC, UC, WS).

Variety villosus is recognized primarily on the basis of
its dense pubescence. In L. lyallii, pubescence ranges from
puberulent-sericeous to woolly-villous approaching spread-
ing hirsute. Several populations exhibit different forms of
pubescence within a single colony; this is especially true
of those specimens viewed from the Carson Pass vicinity
(the type locality of var. villosus), Alpine Co., California.

2. Lupinus lyallii subsp. alcis-temporis (C. P. Sm.) B. J.

Cox, comb. nov.

Lupinus alcis-temporis C. P. Sm., Sp. Lup. 558. 1946.
Type: IDAHO: CLEARWATER CO.: Pierce, 17.7 km. E,
Crystal Creek Grade, Christ 14634 (Holotype: DS; Iso-
types: NY, US). Photos: (RSA, UMO).

Plants sprawling caespitose, branching profusely from
a slender woody base, in clumps 11-30 cm. in diameter;
stems subdecumbent, 8-15 em. long, strigose-sericeous with
dense coarse ascending hairs, nodes 4-6; cauline petioles
2-4 em. long; stipules 7-9 mm. long, connate to the petioles
3-4.5 mm., filiform; leaflets 5-7, elliptic-oblanceolate, apex
acute to short mucronate, longest leaflets 2-18 mm., av.
10 mm. long, 0.8-5 mm. wide, length to width ratio 2.33-
4.33, av. 3.18, strigose-pilose to villous on both surfaces,
some more sparsely so on the upper surface to glabrous;
peduncles 1.5-3.5 cm. long; racemes capitate, 2-3.5(6) cm.
long, densely verticillate; flowers blue with yellow sulcus,

1974] Genus Lupinus — Cox 437

7-8.2 mm. long; bracts 5-9 mm. long, subulate-attenuate,
sericeous to villous; pedicels 1.2-1.8 mm. long; calyx upper-
lip 4.0-5.0 mm. long, bilobed, notch 2-3.5 mm. deep, lower-
lip 5.0-6.0 mm. long, apex tridentate, center tooth 0.2-0.5
mm. long, lips connate 1.0-1.5 mm. ; bracteoles 0.8-1.5 mm.
long, attached at the sinus, villous ; banner elliptic-oblanceo-
late, 7.0-8.0 mm. long, 4-4.5 mm. wide, length to width 1.56-
1.88, av. 1.74, reflexed to appressed ratio 1.25-1.50, av. 1.37,
angle 148-158°; wings 6.0-7.3 mm. long, 3.0-3.4 mm. wide,
length to width ratio 1.97-2.85, av. 2.16; keel 2 mm. wide
at the widest point, angle 111-122°, ciliated sparsely near
the acumen ; ovules 2-3; pods 8.5-9.5 mm. long, 3.4-4.0 mm.
wide, densely short woolly; seeds 2-3, 1.8-2.0 mm. X 2.0-
2.2 mm., 0.9-1.1 mm. thick, tan color.

Subspecies alcis-temporis is found in sandy and gravelly
slopes on road cuts, generally in otherwise barren areas.
One population occurs in a Pinus monticola forest; the
Christ & Smith collection (15385 — NY) was found on
“forest soil bordering beds of Boykinia major; the Dauben-
mire collection (47150 — WS) was made in a Spruce-Fir
Zone. Plants oecur at 915-1220 m. elevation.

Representative specimens: Idaho: CLEARWATER CO.: Pierce, 17.7
km E at the confluence of Big Breakfast Creek and Crystal Creek,
Christ & Smith 15385 (NY); Washington and Lodge Creeks, S31,
T39N, R7E, Lingenfelter 466 (NY, RSA, UC, WS).

This taxon was first described by C. P. Smith from a
Christ “little number picked up in Clearwater County
(Idaho) during elk season in October.” Hence the name
*Christ's Elk-Season Lupine”, Lupinus alcis-temporis in
Smith’s Latinum utilizes the genitive for both elements of
the compound name (alces — elk and tempus — season).
Both Christ and Smith were deeply interested in the fact
that a lupine in anthesis could be found in Clearwater
Co. in the middle of October “without a trace of fruit or
seed, and at only 4,000 feet elevation."

Several stages of development are found in more ex-
tensive study of other collections of these lupines, all of

438 Rhodora [Vol. 76

which have been collected on July 10th or July 11th. Small
clusters of young herbaceous-stemmed, procumbent plants
may be found flowering the same time as older plants with
semi-woody branching stems, sprawling 30-45 cm. in di-
ameter. The isotype material viewed from NY and US
appears merely to be atypically late in reaching anthesis.
It is in anthesis with some fruit in the initial stages of
enlargement, and all branches are woody including second-
ary and tertiary branches; only the leaves and inflorescences
remain herbaceous at this late seasonal date.

3. Lupinus lyallii subsp. minutifolius (Eastw.) B. J. Cox,
comb. & stat. nov.
Lupinus minutifolius Eastw. Leafl. West. Bot. 2: 267.
1940. Holotype: OREGON: HARNEY CO0.: Fish Lake,
Steens Mts., Henderson 8132 (CAS). Photos: (RSA, UMO).

Plants acaulescent; foliage to 4.5 cm. tall; stems 4-6 cm.
tall, appressed hispidulose with more coarse scaberose
hairs; petioles filiform, 2.5-4 cm. long; leaflets 5-7, elliptical,
glabrous above to sparse pubescence on the margins, short
coarse appressed hairs below, largest 6-9 mm., av. 6.8 mm.,
2.9.8 mm. wide, av. 2.45 mm., length to width ratio 2.4-3.0,
av. 2.75; peduncles 6-8 cm. long or only 3 cm. long in im-
mature plants; racemes 1.5-3.0 em. long, 3-4 compact verti-
cils, ascending or erect from nearly prostrate stems; flowers
7.5-10 mm. long, mostly blue with nearly white banners
and purple tipped keels; bracts caducous, 4.5-6 mm. long,
subulate; pedicels 1.0-2.0 mm. long; calyx upper lip 4-5 mm.
long, bilobed, notched 2.5-2.7 mm. deep, lower-lip 4.5-5.0
mm. long, apex entire-serrate to tridentate with the center
tooth approaching 0.7 mm. long, lips connate 1.2-1.3 mm.,
bracteoles 0.2 mm. long, attached at the sinus, strigose to
short ascending pilose; banner elliptic-oblong, 8-8.5 mm.
long, 4.2-5.5 mm. wide, length to width ratio 1.45-1.90, av.
1.65, reflexed to appressed ratio 1.08-1.57, av. 1.34, angle
150-160°; wings 8.8-9.0 mm. long, 3.8-4.2 mm. wide, length
to width ratio 2.10-2.57, av. 2.34, claws 1.0-1.8 mm. long,
glabrous; keel 2.0-2.2 mm, wide at the widest point, angle

1974] Genus Lupinus — Cox 439

118-1235, ciliated along the outer half of the upper margin,
densest toward the acumen; ovules 3; pods 9.0-10.5 mm.
long, 3.5-4.2 mm. wide, short appressed; seeds 2-4.

Subspecies minutifolius is found in the Steens Mountains
of Harney Co. Oregon, geographically replacing subsp.
lyallii. These plants are found on rocky, gravelly, gentle
basaltic slopes but frequently occur in alpine meadows on
hillside areas with grasses and sedges in sloping rocky
habitats. Several specimens have been collected in beds of
lingering snowbanks where snow has accumulated and
remained during the winter and spring. Anthesis generally
occurs during the latter part of July. Plants are found at
elevations of 2439-2896 m.

Representative specimens: Oregon: HARNEY CO.: Big Fish Creek
Canyon, Maguire & Holmgren 26789 (GH, NY, UC, US); Frenchglen,
31.4 km due E-SE, Hansen 490 (osc).

Although some populations of subsp. minutifolius re-
semble subsp. lyallii var. danaus, they have several distinct
morphological characteristics. The flowers of subsp. minu-
tifolius generally have more oblong shaped banners, more
linear wings, and therefore a greater length to width petal
ratio than subsp. lyallii. Although the vegetative mor-
phology of subsp. minutifolius resembles the alpine var.
danaus and other northern forms of subsp. lyallii, it shows
distinction in the filiform petioles from the basal clumps
of less than 4.5 cm. tall, and frequently by the fewer (4-7,
av. 5) leaflets that are glabrous on the upper surface. The
plants of the Cronquist collection. (8694 — NY, OSC, RM,
RSA, UC, WS) are morphologically larger than normal for
the taxon. The foliage of these specimens approach 5 cm.
tall, overall plant height approaches 9.5 cm.; flowers are
an intermediate 8.5 mm. long ; the leaflets are larger 10 mm.
x 8.5 mm.; peduncles are larger 6.5-7 mm. long, and
racemes approach 4 em. long.

4. Lupinus lyallii subsp. subpandens C. P. Sm. ex Dunn,
El Aliso 3: 171. 1955. Type: NEVADA: ELKO CO.: Elko,

440 Rhodora [Vol. 76

113 km. NW, Holmgren, Jun 1938 (Holotype: UTC; Iso-
types: NA, NEV, AGR.). Photos: (RSA, UMO).

Plants with many small tufts from a profusely branch-
ing woody caudex, erect; foliage 4.5-9.0 cm. in height,
mostly less than 6 cm. tall; stems 7-18 cm. tall, usually less
than 1 mm. in diameter, appressed strigose-pilose pubes-
cence less than 2 mm. long ; leaves in basal clusters ; petioles
slender, 3-9 cm. long; stipules inconspicuous, 6-10 mm. long,
connate to the petioles 4-5 mm, subulate; leaflets 5-6, linear-
oblanceolate, apex acute, largest 9-22 mm. long, 2-5 mm.
wide, length to width ratio 3.5-6.3, av. 4.8, sericeous-strigose
below, more sparsely so above with glabrous apex and mid-
rib; peduncle 5-9 cm. long, usually exceeding the foliage by
2-5 cm.; racemes capitate to subcapitate, 1.5-4 cm. long;
flowers dense, blue, 9.5-11.5 mm. long; bracts 4-8.5 mm.
long, appressed to ascending pilose; pedicels 1-2 mm. long;
calyx upper-lip 4.2-5.5 mm, long, notch 1.3-3.2 mm. deep,
lower-lip 4.5-5.5 mm. long, apex deeply serrate to tridentate,
center tooth to 0.5 mm. long, lips connate 1.0-1.5 mm.,
bracteoles slender, inconspicuous, 0.5-1.0 mm, long, attached
to the sinus, strigose-sericeous, hairs to 2 mm. long;
banner elliptic, 8-9.5 mm. long, 5-6.5 mm. wide, length to
width ratio 1.45-1.90, av. 1.56, reflexed to appressed ratio
1.13-1.71, av. 1.44, angle 146-150°; wings 8.2-9.8 mm. long,
3.4-4.8 mm. wide, length to width ratio 2.04-2.65, av. 2.36,
claws 1.5-2.0 mm. long, glabrous; keel 2.2-3.0 mm. wide at
the widest point, angle 108-120°, ciliated along the outer
half of the upper margin, most densely near the acumen,
cilia to 0.7 mm. long; ovules 3-5; pods 11-14 mm. long,
5-5.5 mm. wide, dense strigose to short villous; seeds 2-3,
2.0-2.2 mm. X 2.8-3.0 mm. 1.3 + 0.1 mm. in thickness,
beige to tan with darker brown mottling.

These plants are found in dry, rocky creek beds and
open well-drained draws in northeastern Nevada, in Elko
Co., and southwestern Idaho, in Camas and Owyhee Coun-
ties. The most common associate is Artemisia sp. Plants
flower from mid-July to the first part of August. Altitudi-

1974] Genus Lupinus — Cox 441

nally, they are found at 1677-1982 m., with one population
found on Alleghany Creek at 2439 m.

Representative specimens: Idaho: CAMAS CO.: Fairfield, 16 km.
SE, Ripley & Barneby 10671 (NY). OWYHEE Co.: Mud Flat, 4.8 km.
SW, on road to Juniper Mt, Maguire & Holmgren 26305 (GH, POM,
UC, US, UTC, WS). Nevada: ELKO Co.: O’Neil, 16 km. S, Holmgren,
Cooper & Cain 181 (GH, osc, UTC) ; Wells, 48 km. N, Jones, Jul 1901
(POM).

Dunn (1955) interpreted Lupinus lyallii subsp. sub-
pandens as an acquisition of a combination of the char-
acters of L. aridus with those of L. lyallii. The pubescence
type and system of branching from the caudex, as well
as certain flower morphology, is characteristic of L. aridus,
whereas the small caespitose habit with elongated peduncles
and capitate inflorescences with small flowers is more in-
dicative of L. lyallii. Dunn further suggests that the plant’s
occupancy of the flatland Artemisia habitat of 1677-1982 m.
elevation indicates a considerable amount of the L. aridus
physiological adaptations, contrasting the higher altitudinal
L. lyallii habitat.

This study concurs with Dunn's interpretation of the
affinity of subsp. subpandens with L. lyallii. These plants
inhabit a geographically distinct region, also having both
morphological and physiological distinction. Although each
subsp. subpandens population has aequired a unique com-
bination of characteristics, the plants can usually be recog-
nized by the erect habit of both flowering culms and petioles
as well as by the glabrous apex and midrib of the upper
leaflet surfaces.

The specimens from Camas Co., Idaho (Ripley & Barneby
10671 — NY) have larger flowers than the taxon mean, with
banners only slightly reflexed and frequently with one node
on the flowering culm above 2 cm. from the basal cluster
of leaves, and racemes of 3-5 cm. long, being about 2 cm.
in diameter.

The specimens collected south of O'Neil, Nevada (Holm-
gren, Cooper & Cain 181 — GH, OSC, UTC) are more slight
in habit, having shorter petioles with smaller leaflets that

442 Rhodora [Vol. 76

are more glabrous on the upper surface, and shorter, more
capitate racemes with the banners less reflexed.

The Mud Flat, Owyhee Co., Idaho specimens (Maguire
& Holmgren 26305 — GH, POM, UC, US, UTC, WS) have fewer
leaflets than other populations, and the petioles are more
elongated and slender.

5. Lupinus lyallii subsp, washoensis (Heller) B. J. Cox,

comb & stat. nov.

L. washoensis Heller, Muhlenbergia 6: 72. 1910. based
on L. pinetorum Heller, Muhlenbergia 6: 25. 1910, not L.
pinetorum Jones, Contr. West. Bot. 8: 25. 1898. Type:
NEVADA: WASHOE CO.: Ridge S of Alum Creek, Heller
9748 (Holotype: Nev. Agr.; Isotypes: DS, GH, ISC, MO,
NY, ORE, POM, UC, US, WS). Photos: (RSA, UMO).

L. aridus var. washoensis (Heller) C. P. Sm,, Bull.
Torrey Bot. Club 51: 3083. 1924.

Plants 10-18 cm. tall, in dense caespitose clumps, 15-30
cm. in diameter, from woody vertical caudex branching at
ground level, with 10-18 racemes flowering simultaneously;
stems procumbent to erect, 1-2.5 mm. in diameter, villous
to hirsute-shaggy white pubescence 2-4 mm. long; 2-3
nodes; cauline petioles 4.5-8 cm. long, pubescence as on the
stems; basal petioles on immature plants 4.5-7 cm. long;
stipules 6-10 mm. long, connate to petioles 4-6 mm., densely
white bearded pubescence; leaflets 5-6, oblanceolate to ellip-
tic-oblong, apex short mucronate, largest 9-21 mm. long,
av. 13.7 mm., 3-7 mm. wide, length to width ration 1.86-
3.27, av. 2.71, silvery appressed to ascending pubescence
below, more sparsely so above, upper surface appearing
more greenish; peduncles 3-5 cm. long, generally exceeding
the leaves; racemes 3-6 cm. long, 1.6-1.9 cm. in diameter,
densely subverticillate; flowers blue to violet, banner with
a white sulcus, 8-9 mm. long; bracts 3-5.5 mm. long, subu-
late-attenuate, pilose-sericeous; pedicels 1.5-2.0 mm, long;
calyx upper-lip 3.5-4.5 mm. long, bilobed, the notch 0.8-1.2
mm. deep, lower-lip 4-5.5 mm, long, apex serrate, lips con-
nate 1-2 mm., bracteoles 0.5-1.0 mm. long, attached at the

1974] Genus Lupinus — Cox 443

sinus, villous; banner elliptic-oblong, 7.8-8.0 mm. long, 5.0-
6.0 mm. wide, length to width ratio 1.33-1.67, av. 1.52,
reflexed to appressed ratio 1.25-1.35, angle 140-150? ; wings
7.5-8.0 mm. long, 4-4.5 mm. wide, length to width ratio
1.74-2.00, av. 1.87, claws 1.5-2.0, glabrous; keel 2.5-3.0 mm.
wide at the widest point, angle 118-125°, ciliated along the
upper margin near the acumen; ovules 2-3; pods 9-13 mm.
long, 5-6 mm. wide, villous to ascending woolly; seeds 1-3,
1.8-2.2 mm. X 2.4-2.7 mm., 0.9-1.2 mm. thick, tan with
brown mottling.

Subspecies washoensis has a restricted habitat on the
eastern slopes of the Sierra Nevada in the southern part
of Washoe Co., SW of Reno, Nevada. Plants occur on
gravelly montane open pine slopes at elevations of 1524-
2134 m., flowering in June.

Representative specimens: Nevada: WASHOE CO.: Dinsmore Camp,
Hunter Creek Canyon, Kennedy 1614 (Mo, Ny, UC); Reno, Hillman
(POM); 10-13 km. SW, Hunter’s Creek Road, Archer 6090 (NY).

The general appearance of some of these plants indicates
relationship with Lupinus aridus, a portion of the racemes
being exceeded in height by the foliage. Affinity is also
indicated with L. sellulus var. lobbii in general morpho-
logical structure (Cox 1972a, 1973c). Considering the
total conformation — narrow, short subcapitate or longer
racemes, and the procumbent, branching, sprawling vege-
tative habit — this study concludes that the washoensis
affinity lies most closely with L. lyallii. These plants re-
place L. lyallii subsp. lyallii geographically and ecologically,
aso being recognizably distinct by somewhat larger stature,
having longer racemes, larger leaves, more yellow-green
color and spreading silvery stem and petiole pubescence.

The Dinsmore population (Kennedy 1614 — MO, NY, UC),
which is only a few miles north of the type locality in
Hunter Creek Canyon, has been used in sheep feeding ex-
periment (Nev. Agr. Exp. Sta. Bull. 71: 30. 1909). The
plants are said to be relished by the sheep, proving to be a
good forage plant.

Rhodora [Vol. 76

` oo N
» 0O Q BRITISH COLUMBIA N
———— \
o% E
a |!
WASHINGTON i Y
N
|o M LUPINUS LYALLII
is we RN i (
* & i
/ 1
, \
OREGON C Lees
/ IDAHO |
^ |
PO
q—. de. |
: Ap ———
| AA!
^|
NEVADA '
b | subspecies: variety:
A |
odes. | o-lyallii ——— lyallii
33 N j +- lyallii ————— fruticulosus
3 x
EAT X | *- | yallii villosus
(]
Et i i e - | yallii roguensis
oo N M
go NOM x—lyallii danaus
ii .. CALIFORNIA N v - lyallii macroflorus

A - washoensis
a- minutifolius
x - alcis- temporis

A-subpandens

Figure 1. Distribution of Lupinus lyallii.

1974] Genus Lupinus — Cox 445

LITERATURE CITED

Cox, B. J. 1972a. Biosystematies of Lupinus lepidus-L. caespitosus
complex. Ph.D. dissertation. Univ. of Missouri, Columbia. 343 p.
1972b. IOPB Chromosome number reports XXXVIII.
Taxon 21: 680-681.
1973a. A chemosystematic comparison of the Lupinus
lepidus-L. caespitosus complex. Bull. Torrey Bot. Club 100: 12-17.
1973b. Protein relationships among the perennial cae-
spitose lupines. Bull. Torrey Bot. Club (in press).
1973c. Taxonomy of Lupinus sellulus (Papilionaceae).
Amer. Mid. Natur. (in press).
Dunn, D. B. 1955. A new subspecies of Lupinus lyallii. El Aliso

$2 2171-172.
PHILLIPS, L. L. 1957. Chromosome numbers in Lupinus. Madroño
14: 30-36.

INDUSTRIAL BIO-TEST LABORATORIES
NORTHBROOK, ILLINOIS 60062

THE ROLE OF SOME HALORAGACEAE
IN ALGAL ECOLOGY

L. C. COLT, JR., AND C. BARRE HELLQUIST

While collecting vascular aquatie plants in several ponds
in east-central Massachusetts we found that there was a
considerable accumulation of algae upon some of the vascu-
lar species, but very little on others. The present paper is
the result of our futher observations on and examination of
the relationship between the algae and the vascular plants
upon whose surfaces they were found.

There is little published data elaborating the substrate
preferences of fresh-water algae. Water chemistry, light,
temperature, water density, pH, and a wide variety of
metabolic processes have been treated frequently (see Jack-
son, 1964, and Smith, 1950) as environmental factors in
algal growth. Reference texts (e.g., Prescott, 1962) indi-
cate that some species of algae are epilithic, endophytic, or
epizooic, or that many are epiphytic upon vascular plants or
larger algae. Other than Chapman (1962) there seems to
be no treatment of situations in which the algae have the
opportunity to grow on or in association with a variety of
substrates and in which clearly some means of preference
for one another is established. Tiffany (1951), for exam-
ple, suggested that the amount of substrate available to the
spores for fixation is of major importance to algal growth,
but he fails to include any discussion of the character of
the substrate itself, or of the various environmental factors
that might be operative in the selection and utilization of a
particular substrate. Chapman suggests that, (1) the age
and nature of the substrate flora are important, (2) diatoms
are more likely to be on older leaves, (3) epiphytes are most
numerous on well lighted surfaces, (4) epiphytes are most
numerous on submerged material, (5) depressions between
cells of the host surface seem to provide better substrate for
attachment, and (6) ponds with muddy bottoms have a

446

1974] Algae on Haloragaceae — Colt & Hellquist 447

reduced number of epiphytes. With few epiphytes occur-
ring in our collections we tend to agree with the last prem-
ise, but we have insufficient evidence to either support or
refute the other ideas. We might generally suggest, how-
ever, that rather similar principles may operate for both
tychoplankters and epiphytes.

Our examination of material from different ponds clearly
showed that algae grew profusely upon some vascular plants
but sparsely or rarely on others. We also found that both
the number of species and the types of tychoplankter algae
varied with similar plants in different environments. In
order to identify, if possible, the operative environmental
factors, we selected two sites in which the algal habitats
might be contrasted. In both instances, (1) a small leaved
aquatic plant was the primary algal substrate, (2) the sub-
strate plant grew in the shallower portions of the pond,
(3) both ponds and collecting sites were in virtually iden-
tical locations beside highways, and (4) there was no shad-
ing of the collecting stations by macrovegetation on or near
the banks of the pond.

THE POND STATIONS

1. Pratt Pond. This pond is located at the junction of
Grove Street, Hopkinton Road, and Westboro Road in the
town of Upton, Worcester County, Massachusetts.

All collections were made in the western sector of Pratt,
from water varying between 1.5 and 2.0 meters in depth.
Most of the remainder of the pond is sufficiently deep so as
to permit swimming and diving. The bottom of the pond is
sandy-mud throughout. The pH averaged 6.5 and the alka-
linity was 3 ppm.

The dominant vascular aquatic in the collection area of
Pratt was Myriophyllum heterophyllum Michx., occurring
over more than one-third of the center of the bay. In so far
as we were able to determine during the several months of
field work, no other vascular plant grew in or near the stand
of Myriophyllum. This resulted in a clear zone around the
stand extending to within several meters of the pond’s edge.

448 Rhodora [Vol. 76

In this shallower water a few other rooted aquatics grew
sparsely. Of these Nymphaea odorata Ait., Pontederia cor-
data L., and Potamogeton spirillus Tuckerm., were the most
common.

2. Icehouse Pond. This pond is beside West Main Street,
0.9 kilometers east of Interstate Route 495, in the town of
Hopkinton, Middlesex County, Massachusetts.

Samples were collected from water up to 0.5 meters in
depth along the southern edge of the pond. The deepest
portion of this pond was approximately 1.5 meters, the
bottom being mud and gravel. Water loss from this pond
during the late summer months left some of the westerly
edges as exposed mud and rocks. The pH averaged 6.1, and
the alkalinity was 3.5 ppm.

The shallow waters of this pond were densely populated
with a variety of rooted aquatic plants. Algae were asso-
ciated most profusely with Proserpinaca palustris L., which
grew in scattered stands of a few plants each. Other com-
mon vascular plants in the collection area were Potamogeton
pulcher Tuckerm., P. capillaceus Poir., Eleocharis acicularis
(L.) R. & S., Pontederia cordata f. taenia Fassett, Nym-
phaea odorata Ait., Nuphar variegatum Engelm., Brasenia
sehreberi Gmel., Ludwigia palustris var. americana (DC.)
Fern. & Grisc., Callitriche verna L., Myriophyllum humile
(Raf.) Morong, and Utricularia purpurea Walt.

PROCEDURE

samples of the vascular plants were collected from the
two pond stations and the species determination made using
Fassett (1966) and Fernald (1950). Representative speci-
mens were pressed and are in the herbarium at Boston
State College.

samples of the primary substrate plants were collected,
some portions retained fresh for immediate scrutiny, and
the remainder preserved in a solution of 3% formalin. De-
termination of the associated algal species was made from
Smith (1924), Irenee-Marie (1939), Prescott (1964), and
Patrick & Reimer (1966). Microscope slides were made of

1974] Algae on Haloragaceae — Colt & Hellquist 449

some algae using the Kaiser Glycerin-Gelatin method in
order to facilitate drawing by camera lucida. All other
samples were placed in Transeau's Fluid for permanent
storage.

DISCUSSION

Twenty-one species of algae were found on or in asso-
ciation with Myriophyllum heterophyllum in Pratt Pond.
Sixty-one algal taxa were discovered in various degrees of
association with Proserpinaca palustris in Icehouse Pond.
Seven species were common to each pond. The wide dis-
parity in numbers despite virtually simultaneous collection
times strongly suggests that Proserpinaca offers a more
favorable habitat to the algae than does Myriophyllum. In-
asmuch as a difference in surface characteristies might
lead to the difference in habitats, it was the first possibility
examined. We were not able to substantiate such a premise
on any physical basis. One of the problems, however, is that
only five genera of epiphytes were recorded during the
study. Stigeoclonium and Characium each had one species
present infrequently in Pratt Pond, while Bulbochaete and
Gloeotrichia each had one commonly found species in Ice-
house Pond, and Oedogonium with one common species and
one infrequently encountered, both in Icehouse Pond. The
pattern of occurrence of these was generally the same as
for the tychoplankters and appeared to be due to the overall
differences in the habitats rather than individual differences
between the substrate plants.

Thus it seemed likely that there was a combination of
factors operating in each situation, with sufficient variance
between the two habitats such as to account for the differ-
ence in algal populations. This view is strengthened by the
difference in tychoplankter populations, Icehouse Pond hav-
ing approximately twice as many taxa. Inasmuch as the
tychoplankters do not adhere to a particular substrate, but
live in association with the substrate, they would not neces-
sarily be affected by physical characteristics.

Some previous work gave direction to our scrutiny. An-

450 Rhodora [Vol. 76

derson & Walker (1920) concluded from an investigation
of some shallow Nebraska lakes that light and mechanical
support are of primary influence to algal growth. They also
suggested that shallow water and wind protection give rise
to as nearly uniform conditions as possible. Needham &
Lloyd (1916) showed that in shallow waters with abundant
vegetation currents are reduced or eliminated and there is
little possibility of wind disturbance. The authors' own
experience has been that tychoplankter populations are par-
ticularly rich among the wet sphagnum of New England
bogs.

Comparison of the two sites on these bases yielded some
rather interesting results. It has already been stated that
sunlight was readily and equally available to the water in
which both sets of substrate plants grew. The collecting
area in Icehouse Pond, however, was much shallower than
that of Pratt Pond. This difference was negated by the
position of the tychoplankter algae in Pratt Pond; they
were all within 14 meter of the surface of the water. The
only living algae below that level in Pratt Pond were dia-
toms, and they were usually obscured by the detritus on the
lower branches of plants of Myriophyllum.

Physically, Icehouse Pond is protected from surface winds
by the mixed deciduous-coniferous forest around it. Al-
though its total surface area approximates that of the sec-
tion of Pratt Pond from which we collected, we were unable
to observe more than mere surface ripples on Icehouse Pond
during any visit to the site. Part of the reason for this is
the dense vegetation within the water itself. At Pratt Pond,
however, we were able to note considerable surface disturb-
ance by even relatively light wind currents.

We were unable to observe any current within Icehouse
Pond. This is largely due to the dense aquatic vegetation.
Water movement could easily be discerned in Pratt Pond.
As we collected samples in Pratt Pond, broken pieces of
Myriophyllum would rise within the stand of plants then
drift “downwind.” Our observation of this phenomenon
showed that the drag of the wind on the surface of the

1974] Algae on Haloragaceae — Colt & Hellquist 451

water caused the upper portion of the water to move, cre-
ating an effective pull on the lower water body. Water then
rose through the plants and downwind. Davis (1955) sug-
gested that tychoplankters would be unable to adapt evolu-
tionarily to such current conditions, On the basis of our
limited work we concur, finding no evidence of specialized
attachment structures or polymorphism which would give
any species an advantage in maintaining its position. It
appears to us that the lack of water movement within Ice-
house Pond is conducive to an optimum interchange of
chemical nutrients. On the other hand, the flow within
Pratt Pond apparently reduces a similar interchange, thus
inhibiting population growth to some degree.

Our final consideration was the growth habit of the small
leaved algal substrates and the associated macroflora. In
Pratt Pond Myriophyllum produced vertical axes greater
than the depth of the water. Near the surface of the water
the distal portions of the plants would bend with the drag
of the surface water. This appeared to produce a matting
effect at the surface which would effectively reduce the light
available to lower leveis. The Proserpinaca in Icehouse
Pond grew only to the surface of the water and distal flex-
ing was rare and then of a minor nature, Light could pass
easily through the finely dissected leaves to lower levels.
Moreover, the densely crowded vegetation of Icehouse Pond
effectively reduced lateral “waving” by the plant. Pro-
serpinaca occurred in larger clumps than any of the other
small leaved plants in Icehouse Pond (e.g., Myriophyllum,
Utricularia).

Examination of the broader-leaved aquatics in each pond
did not reveal any substantial amount of algae growing on
or associated with them. The most frequent species were
diatoms, Tabellaria, for example, being the most common.

SUMMARY

From our observations we conclude that for the optimum
growth of tychoplankter algae there are at least three basic
requirements. Not necessarily in order of importance they

452 Rhodora [Vol. 76

are: (1) a substrate which provides both maximum surface
area and exposure to the available light; (2) an absence of
strong or prolonged water currents; and (3) the presence
of a varied flora which provides a broad spectrum of nutri-
ents within the system.

Based on our observations at both ponds we suggest that
small leaved plants will most often provide the optimum
habitat for tychoplankter algae. We are not convinced,
however, that monospecific stands provide the optimum en-
vironment for tychoplankters, although such stands may be
the best of the available substrates. We have found com-
parable situations in bogs, as it has been our experience
that the sphagnum bogs with the greatest variety of macro-
flora usually yield the largest number of algal species.

We further suggest that pH and water temperatures are
primarily useful only in contrasting regions with geograph-
ical and/or geological differences. It can be reasonably
inferred that evolution has already adapted local algal
species to pH and water temperature as they occur in the
algal habitat.

Finally, we concur with previous workers who have sug-
gested that quiet, stable and uniform water conditions are
best for the growth of tychoplankter algal populations.

The list of algae below follows the sequence of Smith
(1950). Nomenclature of the Bacillariophyceae is according
to Patrick & Reimer (1966). Location and frequency are
indicated by the following: IP — Icehouse Pond, PP —
Pratt Pond, R — rare (less than 5 collected in all samples),
F — frequent (averaged up to 10 in all samples), C — com-
mon (averaged more than 10 in all samples), E — epiphyte,
T — tychoplankter.

CHLOROPHYTA
Stigeoclonium subsecundum Kuetzing PP-R-E
Bulbochaete Furberae Collins IP-C-E
Oedogonium michiganense Tiffany IP-C-E
O. pseudoplenum Tiffany IP-C-E

Characium ornithocephalum A. Braun PP-R-E

1974] Algae on Haloragaceae — Colt & Hellquist 453

Pediastrum Boryanwm (Turp.) Meneghini IP-R-T

P. duplex var. cohaerens Bohlin

P. tetras (Ehrenberg) Ralfs

P. tetras var. tetraodon (Corda)
Rabenhorst

Nephrocytium lunatum W. West

Kirchneriella contorta (Schmidle)
G. M. Smith

Scenedesmus abundans var. brevicauda

G. M. Smith
S. bijuga (Turp.) Lagerheim
S. dimorphus (Turp.) Lagerheim

S. quadricauda var. maxima West & West
- Actinastrum Hantzschii Lagerheim
two easily separable species

Spirogyra spp.

IP-R-T
IP, PP-R-T

IP-C-T
PP-R-T

PP-R-T

IP-R-T
IP, PP-C-T
IP-R-T
IP, PP-C-T
IP, PP-C-T

were common in the vegeta-
tive state in Icehouse Pond.

Gonatozygon pilosum Wolle
Triploceras gracile Bailey
Closterium Jenneri Ralfs
Cl. parvulum Naegeli

Cl. Ralfsii var. hybridum Rabenhorst

Cl. setaceum Ehrenberg

Pleurotaenium coronatum (Breb.)
Rabenhorst

P. subcoronulatum var. detum
West & West

P. Trabecula (Ehrenb.) Naegeli

P. Trabecula var. rectum (Delp.)
W. West

P. Trochiscum var. tuberculatum
G. M. Smith

Euastrum abruptum f. minor
West & West

E. binale (Turp.) Ehrenberg

E. ciastonii Raciborski

E. gemmatum Brebisson

Micrasterias Nordstedtiana, Wolle

IP-C-T, PP-R-T
IP-R-T
IP-R-T
IP-R-T
IP-F-T
IP-F-T

IP-F-T

IP-R-T
IP-C-T

IP-R-T
IP-R-T

pr

IP-R-T
IP-R-T
IP-R-T
IP-R-T

454 Rhodora
Cosmarium Blytii Wille

C. difficile var. dilatatwm Borge

C. furcatospermum West & West
C. granatum Brebisson

C. isthmium W. West

C. ornatum Ralfs

C. portianum Archer

C. pseudoconnatum Nordstedt

C. pseudopyramidatum Lundell

C. pyramidatum var. transitorium Heimer]

C. tumidum Lundell

Staurastrum dejectum Brebisson

S. gracile var. nanum Wille

S. gracile var. tenuissima Ralfs

S. Leptocladum Nordstedt

S. Ravenelii var. spinulosum Irenee-Marie

Spondylosium planum (Wolle)
West & West

Desmidium Aptogonum Brebisson

D. Baileyi (Ralfs) Nordstedt

Xanthidium cristatum Brebisson

CHRYSOPHYTA

Tabellaria fenestrata (Lyngb.) Kuetzing
T. flocculosa (Roth) Kuetzing

T. quadrisepta Knudson

Meridion circulare (Grev.) Agardh
Eunotia elegans Ost.

E. formica Ehrenberg

E. pectinalis (O.F.M.) Rabenhorst

E. pectinalis var. ventricosa Grunow

E. praerupta var. bidens (Ehr.) Grunow
E. tenella (Grun.) Cleve

E. serra var. diadema (Ehrenb.) Patrick
Hannaea arcus Patrick

Pinnularia flexuosa Cleve

P. formica (Ehrenb.) Patrick

Frustulia rhomboides (Ehrenb.) de Toni

IP-R-T
IP-R-T
IP-R-T
IP-R-T
IP-F-T
IP-F-T
IP-F-T
IP-F-T
PP-R-T
PP-R-T
IP-R-T
IP-F-T
IP-R-T
IP-F-T
IP-R-T
IP-R-T

IP-C-T
IP-R-T
IP-F-T
IP-F-T

IP-C-T
IP-C-T
PP-C-T

[Vol. 76

IP, PP-C-T

IP-C-T
IP-F-T
PP-F-T
PP-F-T
IP-F-T
PP-F-T
IP-F-T
PP-R-T
IP-F-T
IP-R-T
IP-C-T

1974] Algae on Haloragaceae — Colt & Hellquist 455

E. vulgaris Thwaites IP-F-T
Gomphonema acuminatum var. coronatum

(Ehrenb.) W. Smith IP, PP-C-T
G. augur Ehrenberg PP-C-T

CYANOPHYTA

Microchaete tenera (Thuret) de Toni PP-F-T
Hapalosiphon intricatus West & West IP-C-T
Gloeotrichia Pisum (C. A. Agardh)

Thuret IP-F-E

LITERATURE CITED

ANDERSON, E. N., & E. R. WALKER. 1920. An ecological study of
the algae of some sandhill lakes. Trans. Amer. Micros. Soc.
39: 51-85.

CHAPMAN, V. J. 1962. The Algae. MacMillan & Co., N. Y.

Davis, C. C. 1955. The marine and fresh-water plankton. Mich.
State Univ. Press, East Lansing, Mich.

Fassett, NoRMAN C. 1966. A manual of aquatic plants. Univ.
Wisc. Press, Madison, Wisc.

FERNALD, M. L. 1950. Gray's Manual of Botany. American Book
Co., New York, N. Y.

IRENEE-MARIE, FRERE. 1939. Flore Desmidiale de la Region de
Montreal. Laprairie, P. Q.

JACKSON, DANIEL F. 1964. Algae and Man. D. F. Jackson, Ed.
Plenum Press, New York, N. Y.

NEEDHAM, JAMES G., & J. T. LLOYD. 1916. The life of inland waters.
Comstock Publishing Co., Ithaca, N. Y.

PATRICK, R., & C. W. REIMER. 1966. The Diatoms of the United
States exclusive of Alaska and Hawaii. Monograph No. 13,
Acad. Nat. Sci. Phila.

Prescott, G. W. 1962. Algae of the Western Great Lakes Area.
Wm. C. Brown Co., Dubuque, Iowa.

1964. How to know the fresh-water Algae. Wm. C.
Brown Co., Dubuque, Iowa.

SMITH, G. M. 1924. Phytoplankton of the Inland Lakes of Wiscon-
sin. Bull Univ. Wisc. Press, Serial 1048.

1950. Fresh-water Algae of the United States. Mc-
Graw-Hill Book Co., N. Y., N. Y.

TIFFANY, L. H. 1951. Ecology of fresh water Algae. In: Manual

of Phycology, Chronica Botanica, Waltham, Mass.

BOSTON STATE COLLEGE
BOSTON, MASS. 02115

[Vol. 76

Rhodora

456

Plate 1

1974] Algae on Haloragaceae — Colt & Hellquist 457

Plate 1:

a) Eunotia formica Ehrenberg

b) E. elegans Ost.

c) Staurastrum leptocladum Nordstedt (semicell)

d) Gomphonema acuminatum var. coronata (Ehr.) W. Smith
e) G. auger Ehrenberg

f) Spondylosium planum (Wolle) West & West

g) Pinnularia formica, (Ehr.) Patrick

h) Pediastrum duplex var. cohaerens Bohlin

i) Desmidium Baileyi (Ralfs) Nordstedt

j) Gloeotrichia Pisum (C. A. Agardh) Thuret

k) Seytonema crispum (C. A. Agardh) Bornet

1) Cosmarium isthmium W. West

m) Tabellaria flocculosa (Roth) Kuetzing

n) Eunotia praerupta var. bidens (Ehr.) Grunow

o) Scenedesmus bijuga (Turp.) Lagerheim

p) Characium ornithocephalum A. Braun

q) Euastrum gemmatum Brebisson (semicell)

r) Scenedesmus abundans var. brevicauda G. M. Smith

458 Rhodora [Vol. 76

"75w A, 50w J, "Sue bl, H24 C-i, km-p

Plate 2

1974]

Algae on Haloragaceae — Colt & Hellquist 459

Plate 2:

a)
b)
c)
d)
e)
f)
g)
h)

i)
j)
k)

1)
m)
n)
o)
p)

Oedogonium pseudoplenum Tiffany

Oe. michiganense Tiffany

Scenedesmus dimorphus (Turp.) Kuetzing
Staurastrum dejectum Brebisson

Eunotia serra var. diadema (Ehr.) Patrick
Scenedesmus quadricauda var. maxima West & West
Desmidium aptogomum Brebisson

Staurastrum curvatum var. elongatum G. M. Smith
Eunotia pectinalis (O.F.M.) Rabenhorst
Pleurotaenium subcoronulatum var. detum West & West
Euastrum abruptum fa. minor West & West
Bulbochaete furberae Collins

Pediastrum tetras var. tetraodon (Corda) Rabenhorst
Euastrum ciastonii Rabenhorst

Xanthidium cristatum Brebisson (semicell)
Nephrocytium lunatum W. West

ILLINOIS FIELD AND HERBARIUM STUDIES
RoBERT H. MOHLENBROCK AND DAN K. EVANS

Further field and herbarium studies of Illinois plants have
added several new taxa to the state flora. Several are ad-
ventive species from the Chicago area, while others repre-
sent native species of taxonomically difficult groups and are
treated in some detail. Appreciation is accorded to Mr.
Floyd Swink and his colleagues at the Morton Arboretum
who kindly made available some of their recent collections.
All records cited are in the herbarium of Southern Illinois
University (siU), the Morton Arboretum (MOR), or the
Missouri Botanical Garden (MO).

Poa bulbosa L. The bulbous blue grass, native to Eurasia,
is previously unreported from Illinois. Collections have
been made of this from cultivated beds at the Morton Ar-
boretum. DU PAGE CO.: Morton Arboretum, within the shrub
collection in Lacy Land, May 20, 1971, F. Swink (MOR).

Phleum pratense L. f. viviparum (S. F. Gray) Louis-
Marie. Vivipary, whereby plantlets or vegetative buds re-
place the flowers in part or all of the inflorescence, may be
encountered in several genera. The phenomonon seems most
frequent in the Gramineae, especially such cool season
grasses as Poa and Festuca. Vivipary in Scirpus atrovirens
(Cyperaceae) is also commonly encountered. Because of
the apparent rare occurrence of this type of apomictic re-
production in Phleum pratense, this report seems worth-
while. DU PAGE CO.: In the Morton Arboretum, July 27,
1971, F. Swink (MOR).

Fuirena scirpoidea Michx. The previously known range
of this species was along the coastal plain of Georgia,
Florida, Alabama, and Louisiana. Its discovery in Illinois
at the Dolan Lake Conservation area marks a range exten-
tion of more than 500 miles. One colony was found in
shallow water growing beside Eleocharis obtusa. Svenson

460

1974] Illinois Studies — Mohlenbrock & Evans 461

(1957), the most recent monographer of the genus, com-
bines F. pumila Torrey and F. breviseta Coville with F.
squarrosa Michx. The three species now attributed to the
Gray's Manual region may be identified in the following key.

1. Leaves reduced to sheaths without blades. F. scirpoidea.
1. Leaf blades well developed. ...................... 2.
2. Achene dark brown with the beak pubescent; scale
awn terminal. ............ ET . F. squarrosa.

2. Achene yellow-brown with the beak 2 scale
awn arising below the apex. ......... F. simplex.

Although F. squarrosa (F. pumila) has been attributed
to Illinois (Fernald, 1950; Jones 1950, 1963; Patterson,
1876), a thorough search by the senior author has failed
to reveal any such specimens from this state. HAMILTON
CO. : Shallow water, Dolan Lake shore in Dolan Conservation
area, June 13, 1970, N. Tracy (SIU).

Salix X myricoides Muhl. This taxon is a reputed hybrid
between S. rigida Muhl. and S. sericea Marsh., although it
appears to have more of the characteristics of S. sericea.
It differs from S. sericea by its canescent twigs and thinly
sericeous capsules, while S. sericea has glabrous or glabrate
twigs and densely sericeous capsules. We have recently
collected it from Jackson County for its first known locality
in Illinois, but it undoubtedly occurs elsewhere in the state.
JACKSON CO.: Along rocky stream, Giant City State Park,
August 30, 1971, R. H. Mohlenbrock (SIU).

Populus X smithii Boivin. This is the hybrid between
P. grandidentata Michx. and P. tremuloides Michx., first
described from Michigan by Wagner in 1970. It is inter-
mediate in several characters between the parent species,
the most obvious being the number of dentations along the
leaf margins. The following key will serve to distinguish
this hybrid from its parents.

1. Margin of leaf dentate, with 5-25 teeth (averaging 10-
20)? budà pubescent. ............... ce wee ces 2.

462 Rhodora [Vol. 76

2. Margin of leaf with 5-15 teeth (averaging 10);
petiole 5-10 em long (averaging 7 em). ..........
SNNT? P. grandidentata.

2. Margin of leaf with 12-25 teeth (averaging 20);
petiole 3-6 cm long (averaging 5.5 em). ..........
EEDUVUOOÓO nul P. X smithü.

1. Margin of leaf finely crenate, with 20 or more teeth

(averaging 31) ; buds glabrous or nearly so. ...... ...

VN P. tremuloides.

LA SALLE CO.: Starved Rock State Park, June 16, 1943,
G. N. Jones 15791 (MO). PEORIA CO.: Horse Shoe Bottom,
July 27, 1919, V. H. Chase 3217 (Mo).

Carya pallida (Ashe) Engl. & Graebn. Correspondence
with Dr. Wayne Manning, noted authority on the genus
Carya, suggested the possibility of C. pallida in Illinois,
since it occurs in western Kentucky and southern Indiana.
Consequently, the senior author has kept a close vigil for
this species in southern Illinois and was rewarded with the
discovery of a specimen from Union County referable to
C. pallida. The distinction between C. pallida and C. texana
Buckl. is difficult. Both have yellow-lepidote buds, and the
fruits are not different, although the fruits in C. pallida are
usually smaller. Carya pallida usually has at least a few
leaves with tomentose rachises, while C. texana usually does
not except in var. villosa (Sarg.) Little. In the spring, C.
texana has distinctive red hairs, while C. pallida does not.
UNION CO.: Dry, acidic woods, along Grapevine Trail, May
21, 1972, R. H. Mohlenbrock (SIU).

Quercus veiutina Lam. f. missouriensis (Sarg.) Trel.
This form of Black Oak has the lower surface of the leaves
densely and permanently pubescent. In this respect, f. mis-
souriensis resembles Q. falcata Michx., but differs from this
species in leaf shape and bud and acorn characteristics.
SALINE CO.: Edge of dry woods, Old Stone Face, July 3,
1971, R. H. Mohlenbrock (SIU).

Chenopodium. While preparing the treatment of Cheno-

1974] Illinois Studies — Mohlenbrock & Evans 463

podium for The Illustrated Flora of Ilinois series, the
senior author has studied many collections of the genus
and has re-evaluated the species previously known from the
state. He has followed closely the treatment by Steyermark
(1963) in Flora of Missouri in order to make more uniform
the treatment of this genus in the central midwest. Three
additional recognizable species from Illinois emerged from
the study.

Chenopodium pallescens Standl. In general, all Illinois
Chenopodium with white-mealy, narrow, entire leaves have
been designated either C. leptophyllum Nutt. (Fernald,
1950; Gleason, 1952) or C. pratericola Rydb. (Jones, 1963).
Illinois material can be separated reliably into two taxa,
one to be called C. pallescens Standl, the other C. desic-
catum A. Nels. var. leptophylloides (Murr.) Wahl. The
following key serves to separate these taxa.

1. Leaves 1-nerved, linear; pericarp firmly attached to

readily removed from seed. ©... 1 eee eee nn nn
C. desiccatum var. leptophylloides.

Chenopodium pallescens is found in rocky ground and in
waste areas, primarily in the north-central counties of the
state. Chenopodium dissectum var. leptophylloides is almost
always in waste ground in the northern half of Illinois.

Chenopodium bushianum Aellen. Although Jones (1963)
combined this species with C. album L., the conspicuous
honeycombed surface of the seeds of C. bushianum makes
it a readily recognizable species. It is found occasionally
throughout the state in fields, woods, and waste places.

Chenopodium strictum Roth var. glaucophyllum (Aellen)
Wahl. This taxon is related to C. album L. and C. mis-
souriense Aellen. It differs from C. album by its smaller
seeds (0.8-1.2 mm broad), and from C. missouriense by its
coarsely toothed lower leaves and its calyx lobes which do

464 Rhodora [Vol. 76

not cover the fruit. JACKSON CO.: Waste ground, one mile
north of Carbondale, September 3, 1971, R. H. Mohlenbrock
(SIU).

Draba verna L. var. boerhaavii Van Hall. This variety
differs from typical var. verna by its fruits that are never
twice as long as broad and by the seeds fewer than 40 per
fruit. It is naturalized from Europe and adventive in a
mowed field in Illinois. JACKSON CO.: Field, Southern Illi-
nois University campus, Carbondale, April 4, 1972, R. H.
Mohlenbrock (SIU).

Draba reptans (Lam.) Fern. var. micrantha (Nutt.) Fern.
This variety, unreported previously from Illinois, occurs
occasionally in northern Illinois where it sometimes is found
growing with var. reptans. Variety micrantha differs by
its hispidulous fruits.

Suaeda depressa (Pursh) S. Wats. This species is nor-
mally found in the naturally occurring saline soils of south-
ern Canada, through the plains states, and south to Texas.
In an apparent response to salting by the highway depart-
ment, the plant grew along the highway median in the
Chicago area. COOK CO.: Median strip, one mile north of
Illinois 22, one mile south of Everett Pond, October 7, 1972,
R. Schulenberg (MOR).

Spergu'aria media (L.) C. Presl. Another adventive ele-
ment new to the Illinois flora is this member of the pink
family. The previous range of this species appears to be
the natural saline soils of central and coastal New York as
well as coastal California and Oregon. Illinois collections
were made near Elgin along a tollway where salt is fre-
quently used by the highway department. KANE CO.: 314
miles west of the Elgin toll booth on the Northwest Tollway
right-of-way bank, in sterile clay, September 3, 1972, R.
Read. (MOR).

Pyrus pyrifolia (Burm. f.) Nakai. The Chinese Pear is an
occasional cultivated plant in Illinois. A specimen about

1974] Illinois Studies — Mohlenbrock & Evans 465

thirty-five feet tall has been found growing along a rocky
stream in Giant City State Park. This species differs from
the more common P. communis L. by its rounder fruits,
slightly larger flowers, and sharply toothed leaves. JACKSON
CO.: Along Stonefort Creek, Giant City State Park, Septem-
ber 11, 1970, R. H. Mohlenbrock (SIU).

Crataegus marshallii Egglest. This is one of the few spe-
cies of Crataegus in Illinois which has the veins of the leaf
running to the sinuses as well as to the points of the lobes.
It differs from C. phaenopyrum (L. f.) Medic. by its petioles
much longer in relation to the length of the blade, its fewer
stamens with reddish anthers, and by its usually two nut-
lets per fruit. Its habitat in swampy woods is also dis-
tinctive. JACKSON CO.: Swampy woods, Greentree Reser-
voir, 2144 miles southwest of Gorham, R. Anderson (SIU).

Triadenum virginicum (L.) Raf. The discovery of this
species in Lake County brings to four the number of spe-
cies of Triadenum known from Illinois. The following key
separates these four species.

1. Leaves without punctations. . ....... T. tubulosum.
1. Leaves punctate, at least on the lower surface. .... . 2.
2. Leaves petiolate. ................ ... T. walteri.

2. Leaves sessile. ....... DEMNM o re 9.

3. Sepals obtuse, up to 5 mm long; styles up to 1.5

mm Jong, ........... aa REA T. fraseri.

3. Sepals acute, 5-8 mm long ; styles 2-3 mm long.
E. laeua gy ax v7 12 JE NEN T. virginicum.

LAKE CO.: Boggy ground, Illinois Beach State Park, Au-
gust 3, 1972, R. H. Mohlenbrock (SIU).

Mentzelia decapetala (Pursh) Urban & Gilg. This species
is native to dry prairies and plains west of the Mississippi
River. In Grundy county, apparently its easternmost site,
it was collected along a railway where it occurred with
other such adventives as Kochia scoparia, Setaria viridis,
Helianthus annuus, and Grindelia lanceolata f. latifolia.

466 Rhodora [Vol. 76

GRUNDY CO.: Cinder and gravel ballast, at Eileen, along
right-of-way of Santa Fe RR, August 13, 1972, R. Schulen-
berg, D. Kropp, & D. Ladd (MOR).

Polypremum procumbens L. This species was expected to
occur in southern Illinois since it is known from Kentucky
to the south and five counties in nearby southeastern Mis-
souri. Collections were made from a sandy field in the
Horseshoe Lake Conservation area. The inconspicuous
corolla, procumbent habit, and slender leaves probably have
resulted in this species being overlooked in other localities.
ALEXANDER CO.: Low, sandy field, Horseshoe Lake, August
11, 1971, J. Huston 920 (SIU).

Jacquemontia tamnifolia (L.) Griseb. This species is na-
tive to the southeastern United States and represents a
genus new to the Illinois flora. This probable adventive was
collected from a disturbed habitat where it grew in associa-
tion with Agropyron repens, Ipomoea purpurea, and Eu-
phorbia dentata, The following key serves to distinguish
Jacquemontia from Ipomoea, Convolvulus, and Calystegia in
Illinois.

1. Stigma 1, capitate or with 2 to 3 lobes. ...... Ipomoea.
1. Stigmas 2, noncapitate, without lobes. ............ 2.
2. Stigmas elliptic, oblong or flattened. . Jacquemontia.

2. Stigmas filiform or subulate. ... ........... . 9.

3. Calyx not concealed by large bracts; fruit 2-locu-

lar. ....... see Convolvulus.

3. Calyx concealed by 2 large bracts; fruit 1-locular.
BENNETT UU Calystegia.

GRUNDY CO.: Railroad tracks at Gardner, October 15, 1972,
R. Schulenberg & E. Lace (MOR).

Vinca major L. The large-flowered periwinkle, an intro-
duced European cultivar, was previously known in the wild
in the coastal states from North Carolina south to Missis-
sippi. In Illinois the species was collected around an old
homestead in Pope County where it grew without cultiva-
tion. POPE CO.: Randolph Farm, V. Randolph (SIU).

1974] Illinois Studies — Mohlenbrock & Evans 467

Matelea decipiens (Alex.) Woodson. This climbing mem-
ber of the milkweed family is widely distributed in central
and southern Missouri, but the record cited below is ap-
parently the first from Illinois. All three Illinois repre-
sentatives of this genus are restricted to the southernmost
counties. They may be distinguished by the following key.

1. Flowers greenish-yellow; pedicels glabrous; follicles
smooth, angular. ......... 0 .........., M. gonocarpa.
1. Flowers rose, maroon, or rarely cream; pedicels pubes-

2. Petals 1.5-2.5 mm wide, rose or rarely cream. .....
CETERAE id en vue 39 oa A RE yo G's UR tw deen M. obliqua.

WILLIAMSON CO.: Low floodplain woods, natural levee of
Big Muddy River, north of Colp, May 28, 1972, M. Swayne
& J. Swayne (SIU).

Physalis macrophysa Rydb. This plant, treated variously
as a species, variety, or form, differs from P. subglabrata
Mack. & Bush by its translucent leaves and its large, pyra-
midal calyces when in fruit. The calyx becomes 3-6 cm
long during fruiting. Illinois specimens, previously labelled
P. subglabrata, have been seen from Champaign and Peoria
counties.

Antirrhinum orontium L, The often cultivated lesser
snapdragon has been collected from a cultivated oat field.
Although it has probably escaped elsewhere in Illinois, this
is the first report of such an occurrence. DU PAGE CO.: Oat
field, National Accelerator Laboratory near Warrenville,
September 28, 1972, F. Swink (MOR).

Penstemon cobaea Nutt, This sometimes cultivated
beards-tongue was previously known from southern Mis-
souri northwest to Nebraska, south to Texas, and east to
Arkansas. Within this range the typical habitats are prai-
ries, glades, and bluffs. In Illinois this species was collected
from a dry meadow where it grew in association with such

468 Rhodora [Vol. 76

prairie elements as Asclepias verticillata, Poa compressa,
and Physalis subglabrata. Although this species is not
known to be native to northeastern Illinois, the plant asso-
ciates as well as the prairie habitat might suggest an indig-
enous condition. KANE CO.: Flat, dry meadow, west of
Montgomery, near Kendall County line, June 25, 1972, J.
Phillips, D. Young, & R. Schulenberg (MOR).

Lagenaria siceraria Standl. This highly variable gourd is
frequently planted in Illinois and occasionally escapes from
cultivation. An early collection from Hancock County and
a recent one from Jackson County verify its existence in
Illinois.

Cucurbita pepo L. var. ovifera (L.) Alef. This variety of
the common Field Pumpkin has extremely variable fruits,
many of them the source of interesting ornamental gourds
such as the Pear, the Bell, the Apple, the Egg, and the
Orange. A single collection has been made for this plant in
Illinois. UNION CO. : in disturbed soil along a stream, 2 miles
north of McClure, July 31, 1971, R. H. Mohlenbrock (SIU).

Sherardia arvensis L. This new element of the Illinois
flora is adventive but well established in areas of the SIU
campus at Carbondale. It may be confused with Galiwm
but is distinguished by its long, funnel-form, blue corolla
and narrow leaf-like involucres that subtend the flowers. In
addition to our location, the species is local in Nova Scotia,
southeastern Canada, Missouri, Tennessee, and North Caro-
lina. JACKSON CO.: Lawn, SIU campus by the Forest Ser-
vice Laboratory, April 30, 1971, D. Evans 11408 (SIU).

Campanula rotundifolia L. var. velutina A. DC. The oc-
currence of typical C. rotundifolia from the rocky environs
of northeastern Illinois is not uncommon. However, variety
velutina is heretofore reported only from a single collection
by E. J. Hill from Cheboygan County in northern Michigan.
We have this taxon from near Apple River Canyon State
Park. Here the plant grows in the crevices of an east-facing
limestone wall which borders Apple River. Associated spe-

1974] Illinois Studies — Mohlenbrock & Evans 469

cies include the typical form, Aquilegia canadensis, Carex
granularis, and the cliff fern Cryptogramma stelleri, Con-
sidering the close habitat association with the markedly
different typical form, it seems unlikely that this pubescent
variety is a mere ecological variant as claimed by some.
JO DAVIESS CO.: East-facing limestone wall bordering Apple
River, 14 mile northeast of Apple River Canyon State Park,
June 13, 1972, D. Evans 1156 (SIU).

Grindelia lanceolata Nutt, f. latifolia Steyerm. Prior to
this report from Grundy County, f. latifolia was unknown
in Illinois. The typical form remains unreported from the
state. The closest occurrence of this new taxon to the Il-
linois site is the type locality in Stone County, Missouri.
Collections are also known from Oklahoma. GRUNDY CO.:
At Eileen, in N 14 of Sect. 36, T33N, R6E, August 13, 1972,
R. Schulenberg & D. Ladd (MOR).

Senecio jacobaea L. Collections from railways and road-
sides often produce rare or unusual adventive species. This
waif recently has been collected from a railway in the Chi-
cago area. DU PAGE CO.: Great Western RR at Euclid Ave-
nue, Villa Park, August 17, 1972, F. Swink (MOR).

Thelesperma gracile (Torr.) Gray. The natura] habitat
and range of this unusual composite are the prairies and
plains from Nebraska to Wyoming, south to Arizona, and
east to Texas and Oklahoma; also Mexico and South Amer-
ica. In addition, Steyermark (1963) reports two adventive
sites in Missouri. This roadside collection from Kane county
represents the easternmost extension of its range in North
America. KANE CO.: Along Ancutt Road, west of Montgom-
ery, July 4, 1972, F. Swink (MOR).

LITERATURE CITED

FERNALD, M. L. 1950. Gray's Manual of Botany, ed. 8. The Ameri-
can Book Company, New York. 1632 pp.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated
Flora of Northeastern United States and Adjacent Canada. New
York Botanical Garden, New York. 3 Vol. 1762 pp.

470 Rhodora [Vol. 76

Jones, G. N. 1950. Flora of Illinois, ed. 2. Am. Midl. Nat. Monogr.

5: 1-368. University of Notre Dame Press, Notre Dame, Indiana.
1963. Flora of Illinois, ed. 3. Am. Midl. Nat. Monogr.

7: 1-401. University of Notre Dame Press, Notre Dame, Indiana.

PATTERSON, H. N. 1867. Catalogue of phaenogamous and vascular
eryptogamous plants of Illinois. Oquawka, Illinois. 54 pp.

STEYERMARK, J. A. 1963. Flora of Missouri. The Iowa State Uni-
versity Press, Ames, Iowa. 1725 pp.

SvENSON, H. K. 1957. Fuirena. In: North American Flora 18(9):
505-507.

SOUTHERN ILLINOIS UNIVERSITY
CARBONDALE, ILLINOIS 62901

TWO NEW COLOR-FORMS FROM SOUTHERN MAINE

While collecting in the Ocean Park section of Old Orchard
Beach, York County, Maine, in recent summers, I have
noticed certain plants of Spiraea latifolia, Meadowsweet,
with deep-pink flowers. Some of these plants have pubes-
cent panicles; others are glabrous. I have found this form
growing both in open wet spots and in shaded woodland
stations.

Spiraea latifolia (Ait.) Borkh. forma rosea T. W. Wells,
forma nova. TYPE: Maine: YORK co.: Old Orchard Beach,
11 October, 1969, T. W. WELLS (NEBC). Differt a forma
latifolia floribus valde roseis.

While collecting in salt-marshes along Goose Fare Creek,
which is the southern boundary of Ocean Park, I found
plants of Polygonum ramosissimum which differ from the
typical in that the sepals have bright-white margins.

Polygonum ramosissimum Michx. forma alba T. W. Wells,
forma nova. TYPE: Maine: YORK co.: Old Orchard Beach,
5 September, 1970 T. W. WELLS (NEBC). Differt a forma
ramosissimum sepalis albomarginatis.

THEODORE W. WELLS
MILTON ACADEMY
MILTON, MASSACHUSETTS 02186

RANGE EXTENSIONS AND NEW RECORDS
FOR THE BAHAMA FLORA

STEVEN R. HILL

In recent years the flora of the Bahama Islands has again
aroused the interest of several taxonomists, who have found
The Bahama Flora of N. L. Britton and C. F. Millspaugh
(1920) to be inadequate in many respects. Not only have
numerous nomenclatural changes been proposed, but also
since that time collections of species new to the islands have
been made (Howard, 1950; Howard & Dunbar, 1964; Lewis,
1971). At the present time The Fairchild Tropical Garden
is undertaking the much-needed revision of the outdated
work.

This writer has made two collecting trips to the Bahamas;
the first from April 28 to May 16, 1970, on New Providence
Island, Long Island, and Inagua Island, and the second from
May 18 to June 3, 1972, on Long Island alone. Both trips
were as an undergraduate at Bates College, Lewiston,
Maine, and were under the guidance and direction of Dr.
Harold E. Hackett of the Department of Biology at that
institution. Four collections were from New Providence,
147 from Long Island, and 51 from Inagua. It was noted
that the flora of Long Island, a flora which seems to re-
semble that of the eastern coast of Cuba, was particularly
poorly known. Long Island is an “out island" nearly one-
hundred miles long and four miles wide, trending north-
south, and bisected by the Tropic of Cancer. Since the flora
was poorly known, Walter Cerbin, SSJ, of Clarence Town,
Long Island agreed to collect in his area, with the process-
ing of specimens to be done at the New York Botanical
Garden. To date Father Cerbin's collections number 76 (nos.
102-167 from Long Island, and 1001-1010 from Inagua).
These are deposited in this writer's personal herbarium
(designated as SRH) housed at the New York Botanical
Garden, with duplicates at New York and the Arnold Ar-
boretum.

471

472 Rhodora [Vol. 76

According to Britton and Millspaugh (1920), five col-
lectors had gathered specimens on Long Island up to the
time of their work, and of their collections only 173 num-
bers were cited (nos. 495-520, 524 of William C. Coker,
1903; and nos. 6225-6307 of Britton and Millspaugh, 1907).
The two others considered to have collected on the island
were Swainson (1830-42), whose collections were not known
to the authors of The Bahama Flora, and H. F. A. von
Eggers (1888), whose collections were apparently not suf-
ficiently well-known to be cited. Among those who have
touched upon Long Island since that time are Dr. Harold
E. Hackett of Bates College (53 numbers in the Duke Uni-
versity Herbarium), Father Walter Cerbin, and this writer.

The results of the recent collections include several ncw
records: 49 species newly reported for Long Island, 4 new
to Inagua, and 8 new to the Bahama Islands. The collections
also include the rediscovery of a species formerly known
only from the type, Zamia lucayana Britton. It is interest-
ing to note that many of these new records are of rather
common species in the West Indies, reflecting the paucity
of the collections from the area to date. Cultivated plants
have been included since they commonly escape on the
islands and become a part of the local flora.

The list of taxa is divided into four sections: 1) species
newly reported for Long Island, 2) species newly reported
for Inagua, 3) species newly reported for the Bahamas, and
4) a rediscovery. The citations after the collection number
refer to those institutions in which the specimen is housed,
and the abbreviations are as in the Index Herbariorum
(Lanjouw and Stafleu, 1964).

The author wishes to thank Dr. William T. Gillis for his
help in reviewing the manuscript, Dr. Harold E. Hackett
for his guidance and encouragement while in the Bahamas
and at Bates College, Drs. Arthur and Noel Holmgren, Dr.
Gerrit Davidse, and Mr. Kenneth Becker for their help with
certain determinations, and Dr. John T. Mickel, his advisor
at the New York Botanical Garden.

1974] Bahama Flora — Hill 473

I. Species Newly Reported for Long Island.
A. Native or long established species.
POLYPODIACEAE: Acrostichum danaeifolium Langsd.
& Fisch.: Hill 840 (sRH) sinkhole, Village Road,
S. Clarence Town.

Asplenium dentatum L.: Hill 834 (A, SRH) sink-
hole, Deadman’s Cay.

Pteris longifolia L.: Hill 819b (SRH) moist de-
pression, Mangrove Bush.

Thelypteris normalis (C. Chr.) Moxley: Hill 125,
835 (SRH) sinkholes, Village Road, S. Clarence
Town.

GRAMINEAE: Chloris petraea Sw.: Hill 467 (SRH)
Clarence Town. Det. G. Davidse.

Distichlis spicata (L.) Greene: Hill 123 (A, SRH)
Harbor Point, Clarence Town. Det. A. Holmgren.

Uniola paniculata L.: Hill 826 (A, SRH) Harbor
Point, Clarence Town.

AMARYLLIDACEAE: Agave sisalama (Engelm.) Per-
rine: Hill 824 (sRH) Clarence Town.

Hymenocallis declinata (Jacq.) M. Roem.: Hill
829 (A, SRH) Turtle Cove.

BROMELIACEAE: Tillandsia usneoides L.: Hill 670b
(US, A, SRH) Galloway's Cave, South Clarence Town.

Tillandsia utriculata L.: Hill 529 (sRH) Village
Rd., S. Clarence Town.

COMMELINACEAE: Rhoeo spathacea (Sw.) Stearn:
Hill 244 (sRH) Clarence Town.

ORCHIDACEAE: Encyclia bahamensis (Griseb.) Brit-
ton: Hill 532 (US, A, SRH) 4 mi. W. of Clarence
Town.

POLYGONACEAE: Antigonum leptopus H. & A.: Cer-
bin 120 (NY, A, SRH) Clarence Town.

Coccoloba swartzii Meisner: Hill 471, 841 (SRH)
Clarence Town.

NYCTAGINACEAE: Commicarpus scandens — (L.)
Standley: Cerbin 104 (NY, A, SRH) Turtle Cove.

474

Rhodora [Vol. 76

Guapira bracei (Britt.) Little: Hill 848 (SRH)
Turtle Cove.
BATIDACEAE: Batis maritima L.: Hill 531 (SRH)
Clarence Town.
MIMOSACEAE: Acacia farnesiana (L.) Willd. : Cerbin
144 (NY, A, SRH) Village Road, South Clarence
Town.

Albizia lebbeck (L.) Benth.: Hill 540 (SRH)
Clarence Town.
CAESALPINIACEAE: Cassia biflora L.: Cerbin 116
(NY, A, SRH).
ZYGOPHYLLACEAE: Kallstroemia maxima (L.) T. &
G.: Hill 952 (SRH) Clarence Town.
EUPHORBIACEAE: Pedilanthus tithymaloides (L.)
Poit.: Hill 139 (SRH); Cerbin 139 (NY, A, SRH)
Clarence Town.

Ricinus communis L.: Hill 542 (SRH) common
escape in Clarence Town.
MALVACEAE: Abutilon trisulcatum (Jacq.) Urban:
Cerbin 160 (NY, A, SRH) Clarence Town.

Cienfuegosia yucatanensis Millspaugh: Hill 1015
(SRH) moist depression between Hamilton's and
Buckley's. Identified by Dr. Paul Fryxell.

Thespesia populnea (L.) Soland.: Hill 129 (a,
SRH) Clarence Town.
TURNERACEAE: Turnera diffusa Willd.: Hill 845
(SRH) Harbor Point, Clarence Town.
PASSIFLORACEAE: Passiflora bahamensis Britton:
Hill 515b (A, SRH); Cerbin 166 (NY, A, SRH) Clar-
ence Town.
CARICACEAE: Carica papaya L.: Hill 142 (A, SRH)
occasional in Clarence Town.
SAPOTACEAE: Mastichodendron foetidissimum
(Jaeq.) Cronq.: Hill 839 (A, sRH) Turtle Cove.
APOCYNACEAE: Plumeria obtusa L.: Hill 831 (SRH)
Turtle Cove.
ASCLEPIADACEAE: Cryptostegia grandiflora (Roxb.)

1974]

Bahama Flora — Hill 475

R. Br.: Hill 128 (SRH); Cerbin 153 (NY, SRH)
Clarence Town outskirts.
CONVOLVULACEAE: Evolvulus squamosus Britton:
Cerbin 113 (NY, A, SRH) Mangrove Bush.

Merremia dissecta (Jacq.) Hall: Cerbin 117 (Ny,
A, SRH) Morrisville.
SCROPHULARIACEAE: Stemodia maritima L.: Hill 947
(SRH); Cerbin 119 (NY, A, SRH) 4 miles west of
Clarence Town. Determined by Dr. Noel Holmgren.
ACANTHACEAE: Blechum brownei Juss.: Cerbin 126
(A, NY, SRH) Clements.

Ruellia tuberosa L.: Hill 846 (SRH) Clarence
Town.
RUBIACEAE: Casasia clustifolia (Jacq.) Hitch.: Cer-
bin 141, 152 (NY, A, SRH) Salt Pond.
ASTERACEAE: Bidens pilosa L.: Cerbin 111 (NY, A,
SRH) Mangrove Bush.

Wedelia trilobata (L.) Hitche.: Cerbin 136 (Ny,
A, SRH) Salt Pond.

Verbesina encelioides (Cav.) Benth. & Hook.:
Cerbin 159 (NY, A, SRH) Clarence Town. Deter-
mined by K. Becker.

B. Cultivated or escaped.

AMARYLLIDACEAE: Hippeastrum puniceum (Lam.)
Urban: Hill 1017 (sRH) Hamiltons.

FABACEAE: Sesbania grandiflora (L.) Pers.: Cerbin
102 (NY, A, SRH) Turtle Cove.

EUPHORBIACEAE: Codiaeum variegatum (L.) Blume:
Cerbin 122 (NY, A, SRH) Clarence Town.
ARALIACEAE: Polyscias guilfoylei (Bull. Bailey:
Cerbin 114 (NY, A, SRH) Morrisville.
APOCYNACEAE: Thevetia peruviana (Pers.) K.
Schum.: Hill (A, SRH) Clarence Town.

OLEACEAE: Jasminum fluminense Vell.: Cerbin 140
(NY, A, SRH) Salt Pond.

SCROPHULARIACEAE: Russellia equisetiformis Schl.
& Cham.: Cerbin 133 (NY, A, SRH) Deadman’s Cay.

476

II.

III.

Rhodora [Vol. 76

CUCURBITACEAE: Momordica charantia L.: Cerbin
125 (NY, A, SRH) Clements.

Species Newly Reported for Inagua.

LILIACEAE : Sansevieria hyacinthoides (L.) Druce: Cer-
bin 1001 (NY, A, SRH) Along road to Northwest Point.
PASSIFLORACEAE: Passiflora bahamensis Britton: Hill
515 (sRH) Union Creek.

APOCYNACEAE: Nerium oleander L.: Hill 488a & b.
(SRH) Matthew Town.

CONVOLVULACEAE: Merremia dissecta (Jacq.) Hall: Hill
489 (SRH) Matthew Town.

Species Newly Reported for the Bahama Islands.
AMARYLLIDACEAE: Crinum asiaticum L.: Cerbin 124
(NY, A, SRH) Clarence Town, Long Island. Cultivated
and possibly escaped in moist places.

Zephyranthes citrina Baker: Hill 827 (A, SRH) Clar-
ence Town, Long Island.

Zephyranthes puertoricensis Traub: Hill 836 (SRH)
Hamilton’s, Long Island. Also in cultivation from this
population at NY.

AMARANTHACEAE: Alternanthera brasiliana (L.)
Kuntze: Cerbin 127 (NY, A, SRH) Clements, Long Is-
land. Determined by Mr. Kenneth Becker, NYBG.
CAESALPINIACEAE: Bauhinia variegata L.: Hill 566
(SRH) Nassau, New Providence. Commonly cultivated
and escaped.

MALPIGHIACEAE: Malpighia glabra L.: Cerbin 1005
(NY, A, SRH) Matthew Town, Inagua. Cult. Cultivated
and possibly escaped.

CACTACEAE: Opuntia rubescens (Salm-Dyck) Lemains:
Hill 476 (sRH) Matthew Town, Inagua. Dooryard.
SOLANACEAE: Datura inoxia Mill.: Hill 850 (SRH) Clar-
ence Town, Long Island. Roadside. Also Nash and
Taylor 1357, formerly determined as D. metel L. but
re-identified by H. N. Moldenke. Inagua.

IV. Rediscovery.

CYCADACEAE: Zamia lucayana Britton: Hill 828 (Mo,

1974] Bahama Flora — Hill 477

K, US, A, F, BM, TRIN, SRH, NY). Found in limestone and
sand 100 yards from the sea, about five miles north of
the type locality; Turtle Cove, Long Island. Formerly
known only from the type collection “In a sandy coastal
thicket Clarence Town, Long Island, only one plant
found after a long search (Britton and Millspaugh
6271)” (Britton, 1907). This writer discovered an
extensive colony with many old plants as well as a
thriving understory of seedlings and hermit crabs
which were feeding on the fleshy seed coats. Plants
in the shade generally exhibited much longer leaves
than those in sun, those specimens in shade agreeing
more closely with the type. Male and female cones
were collected, and living material is in cultivation at
NYBG, Harvard, and the Fairchild Tropical Garden.

LITERATURE CITED

BRITTON, N. L. 1907. Contributions to the Flora of the Bahama
Islands. IV. Bull. N. Y. Bot. Gard. 5: 311.

BRITTON, N. L. & C. F. MiLLsPAUGH. 1920. The Bahama Flora.
Hafner Publishing Co., Inc. New York,

Howanp, R A. 1950. Vegetation of the Bimini Island Group. Ecol.
Monog. 20: 317-349.

, & H. F. DUNBAR. 1964. Additions to the Flora of

Inagua, the Bahamas. Rhodora 66: 6-15.

LANJOUW, J. & F. A. STAFLEU. 1964. Index Herbariorum. 5th.
ed. Part I. The Herbaria of the World. Reg. Veg. 31. Utrecht.

LEWIS, W. H. 1971. Additions to the Flora of the Bahama Islands.
Rhodora 73: 46-50.

NEW YORK BOTANICAL GARDEN
BRONX, NEW YORK 10458

A NEW SPECIES OF LECHEA (CISTACEAE)
FROM PENINSULAR FLORIDA

ROBERT L. WILBUR!

A dozen years ago a strikingly different Lechea was
noted (Rhodora 63: 117. 1961.) among collections from
southern Florida. It then seemed unwise, however, to de-
scribe the population that they represent as a distinct new
taxon since the specimens available were not as complete
or mature as would be desired in proposing a new entity in
such a perplexing and technical genus. However, additional
collections from southern Florida and the low-keyed but
persistent hints from Dr. Olga Lakela that something
wasn’t quite satisfactory with available treatments finally
prodded me into taking another look. Her own collections
have largely eliminated one of my reservations about for-
mally proposing a new taxon, although additional collections
from throughout the growing season are still very much to
be desired. Practically nothing is known of the plants ex-
cept in the fruiting stage and although that is the most
important stage for the accurate identification of members
of this genus, it still would be most desirable to know
something more about the vegetative features of this spe-
cies. Leaves in this species are almost completely lacking
by the time fruits are ripening. Still it seems probable that
more of a stimulus for additional collections of this taxon
and of the other highly distinctive representatives of the
genus in peninsular Florida will be forthcoming if the taxon
is formally proposed now than if we wait for more col-
lections.

KEY TO THE PENINSULAR FLORIDIAN SPECIES OF LECHEA

1. Pubescence of aerial stems mostly strongly divergent
and spreading. ......ssse e nn 2.

"This study was supported by NSF GB-13815, for which grateful
acknowledgment is made.

478

1974]

Genus Lechea — Wilbur 479

2. Internal sepals conspicuously V- or U-shaped in

cross-section with thin scarious margins and a
roughened, often sparsely pilose keel (the sepal
otherwise glabrous) ; leaves often over 1.5 cm. long;
capsule thin-walled, readily splitting into 3 valves
at maturity, subglobose, about equaling the calyx in
length. .... 1. L. mucronata Raf. (L. villosa Ell.).

Internal sepals but slightly bowed in cross-section
with texture appearing uniform and pubescence scat-
tered across surface; leaves all less than 1 cm. long;
capsule thick-walled, indehiscent, ellipsoid or some-
what rounded- barrell-shaped, exserted from the
calyx for at least 1/3-1/2 its length. ..............
ems Lis 2. L. divaricata Shuttlew. ex Britt.

1. Pubescence of aerial stems mostly closely appressed or
THOME ea iio ae on an a e oo owe ees ale ee 3.

3.

External sepals equaling or exceeding the internal
sepals in length. ........... 00. ce eee n n n 4.

4. External sepals at least one-fifth longer than the
inner sepals and usually equaling or exceeding
the capsule in length; capsule equaling or exceed-
ing the inner sepals by not more than one-fifth its
length; cauline leaves usually elliptic to elliptic-
oblong, usually less than 5 times as long as wide,
those below the inflorescence commonly appear-
ing whorled and often more than 2 mm. wide.
ck ct acs eo tie eet zu. De dA minor L.

4. External sepals about equaling the inner sepals
and never equalling the capsule in length; capsule
usually exceeding the inner sepals by about 1/3-
1/2 its length; cauline leaves narrowly oblong to
linear, usually 6 times (or more) as long as wide,
those below the inflorescent alternate and less
than 2 mm. wide. ..........-..- rnm n
HERR 4. L. sessiliflora Raf. (L. patula Legg.).

3. External sepals shorter than the internal sepals. . 5.

5. Leaves pubescent on both upper and lower sur-

480

Rhodora [Vol. 76

faces (at least those of the basal shoots conspicu-
ously pilose above and below while the cauline
and rameal leaves are usually inconspicuously
pubescent over entire surface); flowers or fruits
mostly clustered in 2's or 3's; capsule wall thick-
ened and indurate. ........ 5. L. cernua Small.

Leaves variously pubescent below but glabrous on
upper surface; flowers or fruits not appearing
fascicled (but obviously attached separately) ;
capsule wall thin or at least not conspicuously
indurate. 2.0.0.0... e 6.
6. Aerial stems perennial, suffruticose, clearly
woody at base, with wiry woody branches;
capsule exserted from the often spreading
calyx by 1/3-1/2 its length; calyx sparingly
short-pubescent to glabrous. ... ...... ...
Dee eee eee eee ees 6. L. deckertii Small.

6. Aerial stems annual, herbaceous, dying to the
base each year; capsule equaling the calyx or
exserted not more than 1/5 its length from
the closely enveloping sepals; calyx moder-
ately to densely pilose. ............. sus. 7.
7. Leaves abruptly tapering at apex into a

hardened, shiny, conical callosity about
0.25 mm. long; inner sepal clearly 3-nerved
(often best demonstrated by moistening) ;
pedicels averaging over 1.5 mm. long;
capsule exceeding the sepals by about 1/5
its length; seeds mostly 2. ... 7. L. pul-
chella Raf. (= L. leggettii Britt. & Holl.).
7. Leaves pointed but not differentiated into
a callosity; inner sepal 1-nerved; pedicels
averaging less than 1.5 mm. long; capsule
almost completely enveloped by the sepals;
seeds mostly 3-6. ...............00055 8.
8. Calyces densely appressed pilose;
stems and under surface of the leaves

1974] Genus Lechea — Wilbur 481

or at least the midvein and often the
margins appressed pilose; capsules
equaling or slightly shorter than the
closely enveloping inner sepals. ......
PW ga EE 8. L. torreyi Legg. ex Britt.

8. Calyces glabrous; stems and leaves
completely glabrous; capsules very
slightly exceeding the inner sepals at
Maturity l.c vs vs 9. L. lakelae.

Lechea lakelae Wilbur, sp, nov.

Caudex simplex vel paucis aliquotve ramis; folia ramealia
peranguste linearia-elliptica, acuta, 0.6-1.2 cm. longa et
0.3-1.0 mm. lata, subtus costa et margine glabra; pedicelli
glabri, (0.8) 1.0(1.2) mm. longi; calyx fructifer obovoideus,
1.5-1.9 mm. longus et 1.2-1.6 mm. latus; sepala interiora
glabra et leviter carinata, 1.5-1.9 mm. longa et 1.2-1.6 mm.
lata; sepala exteriora glabra linearia vel anguste lanceolata
1/2 vel 2/3 plo breviora quam sepala interiora; capsula
c. 1.2-1.6 mm. longa; semina 3, c. 0.8 mm. longa.

Caudex simple or with few to several branches; basal
resting shoots unknown; aerial (i.e. flowering and fruiting)
stems 1 to several, 2-3 (4) dm. tall, branching mostly above
the middle and forming a rather compact, wiry-stemmed,
bushy top with completely glabrous axes; cauline leaves
lacking on all specimens seen; rameal leaves very narrowly
linear-elliptic, tapering to both the apex and base, 0.6-1.2
em. long and 0.3-1 mm. wide, completely glabrous above and
below with the midvein somewhat elevated beneath; fruit-
ing branches and pedicels glabrous, the pedicels (0.8)1.0
(1.2) mm. long; fruiting calyx obovoid, broadest above the
middle and tapering into the broadly obpyramidal, coria-
ceous base, drying dark reddish brown, completely glabrous;
inner sepals about 1.5-1.9 mm. long, including the indurate,
obpyramidal base, and 1.2-1.6 mm. broad, slightly exceeded
by the mature capsule, very broadly elliptic to almost or-
bicular, apically broadly rounded, only the strongly elevated

482 Rhodora [Vol. 76

or slightly keeled midvein clearly visible; outer sepals linear
to narrowly lanceolate, about 1/2-2/3 as long as the inner;
capsules broadly barrel-shaped, cylindrical, about 1.2-1.6
mm. long and 1-1.2 mm. in diameter, the valves firm, in-
durate, tardily dehiscing about 1/2-2/3 the distance from
apex to base; seeds 3, equilateral, dorsiventrally compressed,
about 0.8 mm. long and 0.4 mm. wide, blackish.

TYPE: Collier Co., Florida: Marco Island, moist level
grassy area, higher beach of recently made lake. 7 August
1967. Lakela 30953 (Holotype, DUKE; Isotype, USF).

OTHER SPECIMENS EXAMINED: Collier Co.: coastal strand, Serenoa-
Ceratiola scrub, growing with Cyperus in white sand, Lakela 31879
(DUKE); Marco Island, coastal strand — Pinus elliottii association,
Lakela & Almeda 31567 (DUKE); Marco Island with Indigofera, Poly-
gonella, in Ceratiola-Quercus scrub, on U.S. 92, Lakela 27852A
(usF); lagoon head, upper beach, Lakela 31673 (DUKE).

My recollection and rather brief notes made more than a
dozen years ago indicate that the species also occurs or
occurred on Florida’s eastern coast in Broward Co.: sandy
scrub above Fort Lauderdale, Buswell 24 July 1936 (Bus!).

The apparent nearest relative of the new species, Lechea
lakelae, is L. torreyi. The most obviously distinctive fea-
tures are indicated in the key, but the two taxa may be
readily distinguished at a glance. Obviously I feel that both
merit recognition at specific rank. In the mind of some
there still may remain a nagging suspicion that the striking-
ly distinctive plants of the newly described taxon are merely
a glabrous form of L. torreyi. I feel, however, that the ap-
parently consistently longer capsules when compared to the
length of the inner sepals suggest that L. lakelae is more
than that.

This species is named in recognition of Dr. Olga Lakela
upon whose keen eye and careful collections our present
knowledge of this taxon is largely based. It is humbling to
recall that her recent significant contributions to south-
eastern botany were made in her “retirement” years after
an active career as a teacher and researcher in Minnesota.

1974] Genus Lechea — Wilbur 483

Not many botanists are destined to write floras of as distant
regions as one of Minnesota's northeastern counties and
Florida's southern tip.

DEPARTMENT OF BOTANY
DUKE UNIVERSITY
DURHAM, N.C. 27706

PIMPINELLA SAXIFRAGA L. (UMBELLIFERAE) IN
WISCONSIN: This species, adventive from Eurasia, has
heretofore been reported to extend from Newfoundland and
New Brunswick to Delaware and the District of Columbia,
west to Pennsylvania, Ohio, and Indiana (Mathias & Con-
stance, 1944. Umbelliferae, in North American Flora 28B
(1): p. 133; Fernald, Gray’s Manual, 1950; Gleason, Illus-
trated Flora, 1952; Gleason & Cronquist, Manual of Vascu-
lar Plants, 1963). Jones (Flora of Illinois, 1950) and
Steyermark (Flora of Missouri, 1963) do not mention the
plant, so it appears not to have spread westward. I report
here its discovery in Wisconsin: WINNEBAGO CO.: roadside
at junc. Country Club Road and Black Wolf Point Road,
sect. 24, TI7N, R16E, 24 September 1967, Below 146; in
roadside ditch ca. 144 mile from the junc. of county trunks
YY and N, on N, sect. 20, T17N, R16E, 26 September 1972,
Rhyner 044.

The two specimens are deposited in the herbarium here
at Oshkosh. The curators of the herbaria at MIL, UWM,
and WIS have no records of the plant from Wisconsin; Iltis
at WIS has kindly confirmed the identifications. The plant's
occurrence at two sites four miles apart and spanning a
five-year period warrants its inclusion as an element in the
Wisconsin flora.

NEIL A. HARRIMAN

BIOLOGY DEPARTMENT

U. OF WISCONSIN — OSHKOSH
54901

A SECOND REPORT OF THE PROTHALLIA OF
LYCOPODIUM INUNDATUM IN NORTH AMERICA'

Davip M. LANE AND A. LINN BOGLE

Gametophytes of Lycopodium inundatum L., first re-
corded for North America in 1972 (Bruce, 1972) from
two localities in Michigan, were found growing at the edges
of shallow wet depressions along the margins of a sandy
access road on the shore of an inactive and flooded sandpit
(Figs. 1, 2) in Somersworth, Strafford Co., New Hampshire.
The mature sporophytes of L. inundatum var. inundatum
(compare Gillespie, 1962) are abundant along the shoreline,
often forming dense mats (Fig. 3).

Samples of sandy soil containing the smallest visible
sporophytes were collected in October and examined accord-
ing to Bruce’s technique. Some of the young sporophytes
were still attached to living gametophytes, as adjudged by
the green color, turgidity, and characteristic lobing of the
latter. Gametophytes were also found which had not yet
produced sporophytes.

Gametophytes ranged in size from less than one mm. in
diameter for one which bore no sporophyte, up to about
three mm. in diameter among those bearing sporophytes.
Included in the latter category were one gametophyte bear-
ing a sporophyte which had produced about ten microphylls,
and two gametophytes each bearing two young sporophytes
(Fig. 4). In contrast, however, some very young sporo-
phytes were found to have no attached gametophytes, the
latter apparently having rotted away, even though the
sporophytes bore only one or two microphylls. This condi-
tion appeared particularly prevalent where the soil particles
were overgrown with moss protonemata and algae.

In addition to variation in size discussed by Bruce, vari-
ations in occurrence and condition, as well as in form (Bold,
1973), of the gametophyte in relation to differences in the

‘Published with the approval of the Director of the University of
New Hampshire Agricultural Experiment Station as Scientific Con-
tribution No. 756.

484

1974] Lycopodium inundatum — Lane & Bogle 485

microhabitat suggest the need for microecological studies
in the field and laboratory. These would complement lab-
oratory study of the culture and development of gameto-
phytes of other species (Freeberg, 1962; Freeberg and Wet-
more, 1957) and of the embryogeny of the sporophyte of
Lycopodium inundatum (Bruce, 1972).

The fact that spores of Lycopodium inundatum germinate
readily and produce green photosynthetic gametophytes
which live on the surface of the soil is of potential value to
biology teachers (Bierhorst, 1964) in areas where the spe-
cies occurs in New England (Fig. 5; for cireumpolar range
map see Hultén, 1968, p. 26), for the gametophytes might
be grown on artificially denuded patches of soil next to the
sporophytes. Although gametophytes of mosses and ferns
are readily available and commonly used for study in biology
classes, few students ever see the live, fleshy gametophytes
of Lycopodium. It is possible that careful searching in suit-
able locations near colonies of mature sporophytes would
produce numerous live gametophytes for class use.

LITERATURE CITED
BrERHORST, D. W. 1964. Suggestions and Comments on Teaching
Materials of the Non-Seed Bearing Plants. In: Vestal, P. A.
et al. 1964. The use of living material in the teaching of botany.
Amer. Biol. Teacher. 26: 89-113.
BorLp, H. C. 1973. Morphology of Plants, Ed. 3. Harper & Row,
New York. 688 p.
BRUCE, J. G. 1972. Observations on the Occurrence of the Pro-
thallia of Lycopodium inundatum. Amer. Fern Jour. 62: 82-87.
FREEBERG, J. A. 1962. Lycopodium prothalli and their endophytic
fungi as studied in vitro. Amer. Jour. Bot. 49: 530-535.
, and R. H. WETMORE. 1958 (dated 1957). Gametophytes
of Lycopodium as grown in vitro. Phytomorphology 7: 204-217.
GILLESPIE, J. P. 1962. A Theory of Relationships in the Lyco-
podium inundatum Complex. Amer. Fern Jour. 52: 19-26.
HuLTÉN, E. 1968. Flora of Alaska and Neighboring Territories.
Stanford Univ. Press. 1008 p.
Seymour, F. C. 1969. Flora of New England. Charles E. Tuttle
Co., Rutland, Vermont. 596 p.

DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824

486 Rhodora [Vol. 76

Figure 1. Typical macrohabitat with abundant mature sporo-
phytes along brook (to left) and pond (to right) of access road
(center). Note area of Fig. 2 (box).

Figure 2. Microhabitat of gametophytes at edges of depression
(left of center) darkened by growth of moss protonemata and algae.

1974] Lycopodium inundatum — Lane & Bogle 487

—

Figure 3. Vertical view of dense mat of mature sporophytes (each
branch less than one cm. thick).

Figure 4. Gametophyte bearing two young sporophytes (upper
right dark area was green and minutely lobed, the remainder of the
gametophyte was colorless).

488 Rhodora [Vol. 76

Figure 5. County distribution of Lycopodium inundatum varieties
in New England (plotted from Seymour, 1969): var. inundatum
(dots), var. Bigelovii Tuckerm. (circles), and var. robustum R. J.
Eat. (triangles). Note site of gametophytes (X) discussed here.

NEW AND INTERESTING PLANTS FROM
THE CENTRAL PLAINS STATES

LAWRENCE K. MAGRATH AND RONALD R. WEEDON

As a result of field and herbarium studies, a number of
new or interesting records of plants have been found for
Texas, Oklahoma, Kansas, and Nebraska. All specimens
cited, unless indicated otherwise, are located at KANU.

Corallorhiza wisteriana Conrad. Nebraska: DAWES CO.:
Chadron State Park, open ponderosa pine wooded hillside,
pine needle litter, clay loam soil, two stems growing in
ravine, in fruit, 14 July 1972, Magrath & Weedon 7620
(KANU). This collection represents an extension of the
range of the coralroot southward from the Black Hills
of South Dakota into the Pine Ridge of northwestern
Nebraska.

Holosteum umbellatum L. Texas: COLLINGSWORTH CO.: 0.3
mi E of the Salt Fork of the Red River on T 203, growing
in sandy soil in roadside ditch, scattered and uncommon,
10 April 1973, Magrath & Richardson 7936 (KANU). KEN-
EDY CO.: 18.3 mi S of Armstrong on US 77, growing in
railroad right-of-way, 20 March 1972, Magrath, Rice &
Johnston 7225 (KANU, OKLA). This collection represents
an extension of the range of the species westward from
Oklahoma into the Texas Panhandle and southward from
Oklahoma to the Gulf Coastal area of southern Texas.

Scleranthus annuus L. Oklahoma: GRADY CO.: Oklahoma
College of Liberal Arts Campus (17th and Grand Avenue)
in Chickasha; growing in flower bed on E side of Austin
Hall; sandy soil; rare; associated with Arenaria, Bromus
and Holosteum, flowers greenish-white, apetalous; 24 May
1973; Magrath 8032 (KANU). This collection represents an
extension of the range of this species southwestward from
Missouri and eastern Kansas.

489

490 Rhodora [Vol. 76

Ranunculus testiculatus Crantz. Nebraska: DAWES CO.:
Chadron State Park, 8 mi S of Chadron on Hwy #385, in
campground in waste areas in shade of deciduous trees,
weedy areas dominated by Taraxacum officinale, sandy soil,
abundant, 6 June 1972, Weedon & Gates 7011 (KANU);
Doris Gates, 29 April 1970. The range of this species is
now known to be extended eastward into northwestern
Nebraska. The above collection indicates the probable
establishment of the species in abundance in this area.

Thlaspi perfoliatum L. Oklahoma: KAY CO.: rest area on
I-35, 2.8 mi N of Blackwell-Medford Exit; clay loam soil,
somewhat sandy; small area near moist drainage ditch;
only one colony seen, apparently introduced; associated
with Bromus, Holosteum and Rumex; 6 May 1973; Magrath
7969 (OKL). This collection represents an extension of the
range of the species southwestward from Missouri and
Kansas.

Eryngium prostratum Nutt. var. prostratum. Kansas:
CHEROKEE Co.: 1.5 mi S of Galena on K 26, small cut-over
and disturbed marshy area in Ozarkian oak-hickory woods,
gravelly soil, associated with Juncus and sedges, apparently
rare — only one plant found, prostrate vine rooting at the
nodes, flowers bluish-purple, 22 October 1972, Magrath 7855
(KANU). This collection represents an extension of the
range of the taxon westward from the eastern Ozarks of
Missouri, and northward from southeastern Oklahoma.

Lactuca saligna L. f. ruppiana (Wallr.) G. Beck. Oklahoma:
OKLAHOMA C0.: ca. 3 mi N of the Jct of I-44 and I-35 on
I-35, a few scattered plants growing in road-cut, sandy
loam soil, 5 August 1972, Magrath 7700 (KANU, OKL, OKLA).
This collection represents an extension of the range of this
taxon southwestward from Missouri and Kansas.

LAWRENCE K. MAGRATH RONALD R. WEEDON
DEPARTMENT OF BIOLOGY DEPARTMENT OF BIOLOGY
OK. COLLEGE OF LIBERAL ARTS CHADRON STATE COLLEGE
CHICKASHA, OK. 73018 CHADRON, NEB. 69337

A NEW DROSERA FROM VENEZUELA
J. A. STEYERMARK AND L. B. SMITH

Shortly after entering the Grand Savanna of Venezuela
on the northern side we left the road to examine a sandy
area with scattered small shrubs. On the patches of white
sand between was a Drosera so small that it could easily
be covered by a quarter and with so short a peduncle that
the one or two flowers appeared practically sessile. This
new species was such a happy augury for the Smiths' intro-
duction to the Guayana Highland that we have christened
it as follows:

Drosera felix Steyermark & Smith, sp. nov. Fig. 1.

Plantae solitariae; caulibus curtissimis; foliis dense rosu-
latis, patentibus; stipulis ad petiolum 1/3 adnatis, ad me-
dium 3-laciniatis, membranaceis, 2.5-3 mm longis, basi
vaginantibus, solidis, 1 mm latis, segmentis laciniatis an-
guste triangulari-lanceolatis, longe attenuatis, 1.7-2.2 mm
longis, 0.2-0.3 mm latis, glabris; petiolis 5-5.5 mm longis,
0.7 mm crassis, supra minute papilloso-verruculosis et pilis
patentibus paucicellularibus acicularibus laxissime vestitis,
subtus pilis multis adpressis simplicibus acicularibus onus-
tis; laminis suborbicularibus, apice late rotundatis, 3 mm
longis, 2.5 mm latis, castaneo-purpureis, supra margine
glandulis 2-4 stipitatis et centro glandulis subsessilibus
vestitis, subtus pilis eis petioli similibus; pedunculo nullo
vel haud ultra 2.5 mm longo, eglanduloso; bractea lineari-
lanceolata, acuminata, 1.5 mm longa, 0.1 mm lata, ciliata;
floribus 1-2; pedicello 1-5 mm longo, pilis adpresso-adscen-
dentibus simplicibus et glandulis remotis sessilibus atro-
purpureis vestito; hypanthio cupulato-subcampanulato, 0.8
mm longo, 1.2 mm lato, pilis patentibus remotis vestito;
sepalis 5, lanceolatis, 1.5-1.8 mm longis, sub medium 0.8 mm
latis, e pilis laxis ascendentibus simplicibus paucicellularibus
hirsutulis; petalis 5, patentibus, albis, obovatis, apice ro-
tundatis, basi cuneatis, 2.5 mm longis, 1.5 mm latis; stami-

491

492 Rhodora [Vol. 76

Fig. 1, a-h, Drosera felix: a, habit; b, single leaf (upper side);
c, single leaf (lower side); d, hair detached from petiole; e, calyx
and pistil with pedicel; f, stamen; g, petal; h, seed, ventral and
dorsal views from left to right.

1974] Venezuela Drosera — Steyermark & Smith 493

nibus 5, quam petalis brevioribus; filamentis 1 mm longis,
glabris; antheris subhemisphaericis, 0.5 mm longis, 0.4-0.5
mm latis; ovario subgloboso, 1 mm longo, 1 mm lato, glabro;
stylis 3, ad basim bipartitis, ramis 1.7 mm longis, glabris.
Venezuela: BOLÍVAR: Carretera El Dorado to Santa Elena
de Uairen south of El Dorado, alt 1200-1400 m, 19 Feb 1972,
Julian A. Steyermark, Lyman & Ruth Smith, G. C. K. & E.
Dunsterville 105468 (holotype VEN, isotypes NY, US).

INSTITUTO BOTANICO
CARACAS, VENEZUELA

AND
DEPARTMENT OF BOTANY
U. S. NATIONAL MUSEUM
WASHINGTON, D. C. 20560

INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA

Manuscripts should be submitted in duplicate and should
be double-spaced or preferably triple-spaced (not on cor-
rasable bond), and a list of legends for figures and maps
provided on a separate page. Footnotes should be used
sparingly, as they are usually not necessary. Do not indi-
cate the style of type through the use of capitals or under-
scoring, particularly in the citations of specimens, except
that the names of species and genera may be underlined to
indicate italics in discussions. Specimen citations should
be selected critically especially for common species of broad
distribution. Systematic revisions and similar papers should
be prepared in the format of “The Systematics and Ecology
of Poison-Ivy and the Poison-Oaks,” W. T. Gillis, Rhodora
73: 161-237, 370-443. 1971, particularly with reference to
the indentation of keys and synonyms. Papers of a floristic
nature should follow, as far as possible, the format of
“Contribution to the Fungus Flora of Northeastern North
America. V.,” H. E. Bigelow & M. E. Barr, Rhodora 71: 177-
203. 1969. For bibliographic citations, a recommended list
of standard journal abbreviations is given by L. Schwarten
& H. W. Rickett, Bull. Torrey Bot. Club 85: 277-300, 1958.

Volume 76, No. 807, including pages 315-495, was issued Nov. 15, 1974.

495

CONTENTS: — continued
Two New Color-forms from Southern Maine

Theodore W. Wells .......... eren nnne ener
Range Extensions and New Records for the Bahama Flora

Steven R. Hill 1... eese eene tornato tne eo osten atteso ases e tenet otto

A New Species of Lechea (Cistaceae) from Peninsular
Florida
Robert L, Wilbur ccccccccccccsscccccsccssecssecesseesnseceeeseeseessseeenseneeeeaeees
Pimpinella Saxifraga L. (Umbelliferae) in Wisconsin
Neil A. Harriman ....... eere eene enne eet tnnt neenon seno eno

A Second Report of the Prothallia of Lycopodium
inundatum in North America

David M. Lane and A. Linn Bogle ............ ee

New and Interesting Plants from the Central Plains States
Lawrence K. Magrath and Ronald R. Weedon ..................

A New Drosera from Venezuela
J. A. Steyermark and L. B. Smith ........ eee

Instructions to Contributors ............ eene

470

471

478

483

484

————

Dodota

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by
ALFRED LINN BOGLE, Editor-in-Chief

ROLLA MILTON TRYON
STEPHEN ALAN SPONGBERG
GERALD JOSEPH GASTONY
RICHARD EDWIN WEAVER

Associate Editors

Vol. 76 December, 1974 No. 808

CONTENTS:

Phenology and Physiognomy of the Hydrophyte Community
in Otsego Lake, N. Y.

Willard N. Hammam eene nnne nennt nter

Panicum ensifolium Baldw. — New for Mexico
Monn EONIA a iedera a REED

Significant New Distributional Records for the Genus
Sphagnum in the Northeastern United States
Richard E. Andrus

ersessostastootestlüesesastssecesrepesosevveseveeevecectéasesoaoseneven

Environmental Control of Needle Characteristics in Sub-
alpine Black Spruce

Brian F. Chabot

eeseshssesssthsósssstsosshseshpseesseosttsesereeeveeceooccessausnsecnasne

(Continued on Inside Back Cover)

497

509

511

The Nem England Botanical Club, Jne.
Botanical Museum. Oxford St.. Cambridge. Mass. 02138

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Rhodora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 76 December, 1974 No. 808

PHENOLOGY AND PHYSIOGNOMY OF THE
HYDROPHYTE COMMUNITY IN OTSEGO LAKE, N.Y.

WILLARD N. HARMAN

As part of a biological survey conducted in 1969, the
species and distribution of aquatic macrophytes in Otsego
Lake, Otsego Co., N.Y., were determined (Harman 1970).
The 1969 information was compared with data collected by
Muencher (1936) in 1935 (Harman & Doane 1970). Emer-
gent plants were more abundant in 1969 than in 1935. The
number of species present in the lake had decreased between
1935 and 1969 and an introduced species, Potamogeton
erispus L., that was not present in 1935, was the most
obvious plant in the lake in early summer of 1969. The
reduction in species in this lake is similar, although not as
pronounced, as changes observed over 50 years in Oneida
Lake, N.Y. (Harman & Forney 1970) and over 70 years in
Put-in-bay Harbor, Lake Erie, Ohio (Stuckey 1971).

During the summer of 1969 it was noted that the zooben-
thos and zoo-meroplankton populations underwent drastic
fluctuations that correlated with the phenology of the aqua-
tic macrophytes and the seasonal changes in the physiog-
nomy of the plant community. A study was initiated to
more accurately determine the composition of the flora at
various times during the year so that we could quantita-
tively ascertain its effects on the associated fauna. The
purpose of this report is to describe the seasonal changes in

497

498 Rhodora [Vol. 76

the appearance of the littoral zone in Otsego Lake at Rat
Cove, at the SUNY Oneonta Biological Field Station at
Cooperstown, New York.

METHODS

In 1970, a transect about 500 m long was plotted from
the end of our docks through the littoral zone, to the deeper
water beyond. Plants growing along this line were meas-
ured by divers each week, from 1 June through 2 Septem-
ber, to determine the emergence time of the spring cohorts,
greatest maximum heights attained, times of flowering, and
times of death and decomposition of each species (Harman
1971). Because all growth represented even-aged stands,
sufficient data were obtained by measuring the tallest in-
dividual plants of each species each week. It was impossible
to measure the same individuals throughout their existence
because the act of measurement disrupted, if not destroyed
these fragile organisms. Also, movements along the tran-
sect disturbed the sediments so that by the end of Septem-
ber we had to move farther and farther away from the
original line in order to measure plants in an undisturbed
environment. Secchi transparency and surface water tem-
peratures in Rat Cove were recorded from 15 June through
25 August.

In 1971 the same procedures were followed with these
changes and additions. Samples were taken through the
ice on 5 February. Weekly observations were initiated
Shortly after the ice breakup (22 April) and continued
until 16 December after all plants had completed their
seasonal growth. 'The transect was marked with an an-
chored hand line for divers to follow in order to alleviate
the problems of getting lost in the often turbid water as
had happened a few times in 1970. Secchi transparency and
surface water temperatures in Rat Cove were recorded
weekly from 10 May 71 until 3 January 72.

Samples of benthic organisms were taken near the tran-
sect at depths of 1-2 m, 3-5 m, and 6-8 m each week, to

1974] Otsego Lake — Harman 499

determine and correlate seasonal changes of the macro-
benthos with those of the aquatic plants. Likewise, num-
ber 20 plankton nets were carried by divers through the
macrophytes occurring at 1-2 m, 3-5 m, and 6-8 m depths
each week, to correlate the successional changes in the zoo-
plankton community associated with them. The data result-
ing from these observations will be reported in a manuscript
now in preparation.

Surface temperatures were essentially the same for both
1970 and 1971 (high = 24°C; mean for June, July, and
August = 21.0 and 21.2°C, respectively). Mean Secchi
transparency for June, July, and August was 4.4 m in 1970
and 4.3 in 1971. One appreciable difference in transparency
was noted in June, the time that many of the plants were
growing most actively. In 1970 the mean Secchi transpar-
ency was 5.3 m, in 1971, 4.2 m (Harman 1972).

PHENOLOGY AND PHYSIOGNOMY

The zonation of hydrophytes according to water depth,
ie. emergents, floating leaved varieties, and submergents
(e.g., Sculthorpe 1967), is common knowledge. The com-
ponents of the submergent macrophyte flora also occur in
similar patterns, presumably because of the varying com-
pensation points of each species relative to decreasing light
intensity in deeper water, or to their growth in the shade
of other species, and to the severe competition for space on
eutrophic littoral substrates.

In any one area along the Rat Cove transect, between
50 em and 550 cm in depth, several species of hydrophytes
usually occur. This often results in a complex physiognomy
exhibiting an overstory (usually of low density), a moder-
ately dense intermediate stratum, and a low, very dense
understory, although up to 7 distinct strata may be present.
Vertical stratification of the community is continually
changing during the growing season as the various species
of macrophytes emerge from the substrate, attain maximum
height and density, and then decompose.

500 Rhodora [Vol. 76

The shallowest waters along the transect are barren be-
cause of the action of the waves on the shore. Nuphar
variegatum Engelm., the yellow water lily, grows in scat-
tered clones in water from 20 to 60 cm in depth in associa-
tion with Elodea canadensis Michx. and Megalodonta Beckii
(Torr.) Greene. Water from 50 em to 500 cm in depth
maintains all the remaining macrophytes studied except for
Nitella flexilis (L.) C. A. Agardh and Potamogeton crispus
which occur in water from 500 to 700 em in depth.

The growth of macrophytes in 1971 was similar to that
observed in 1970 with the following exceptions: Potamoge-
ton crispus, P. Richardsonii (Ar. Benn.) Rydb., and P.
illinoensis Morong reached heights of approximately 300 cm
in 1970, but attained less than 250 cm in 1971 before de-
composition began. In 1969 and 1970 Potamogeton crispus
reached the surface in some areas of the lake forming dense
beds that hindered navigation, but did not reach the surface
anywhere in 1971. After the spring cohort of the pond
weeds mentioned died back in 1970, comparatively few
scattered plants were observed during the rest of the sum-
mer. In 1971, after the first cohort decomposed, many
individuals resumed growth and died back sporadically until
October.

The following are descriptions of the phenology and phy-
siognomy of the plants at 4 sites (0.5 m, 2.0 m, 4.0 m, and
5.5 m in depth) along the transect (Fig. 1).

WATER DEPTH 0.5 M: Nuphar variegatum emerged from
the substrate the first week in May; by 15 June, 1971 the
leaves had reached the surface. They remained there until
decomposed in early September. Elodea canadensis and
Megalodonta Beckii started their seasonal growth in late
April and the middle of May respectively. They maintained
a maximum height of about 30 cm beneath the Nuphar
during August, By 15 September M. Beckii had collapsed
to the bottom and was decomposing. At the same time,
E. canadensis started to decompose from the base and the
entire plants became very brittle, By 15 November the

501

Otsego Lake — Harman

1974]

"Ádouso əsuəp = :Ádouvo əsnyıp = - - — -'sejep pejose[es ye sejAudoupAu
juepunqe jo sjudreQ Jurwous ‘YLE o3osj() 940) yey UI yydep ur ur G'Q uumn[OO I93€A gZ eun3rj

LZ 19quie^oN GL LL 19quieides GL LZ 15nBny GL LL Aine GL LZ eunr SL

ejuopoyebay

eapo|3 e3uopojeBa|A "9 eapoj[3

?epo|3

?apo|3 9 e3uopojebejA|

1eudnwN seydny seydny

ejensqng woi wo

'LL6T ‘GT Á[nf uo uorjejeZeA gueurwop Jo syry ejvorpur seul[
[fjuozHOH 'gG-G sein3g jo Suoreoo[ Burmoys j?esut1j 940) yey JO e[goid orjeunue?dSer[ ‘T ANB

L4 ie:0niiqng
—] =

9e Asqne

—____

aoeyins

L- -

31vHisg8ns WOHud SHu313W
+
L

a10ys

502 Rhodora [Vol. 76

winter condition was reached. Megalodonta Beckii and E.
canadensis were prone on the bottom with winter buds
1 to 2 em long developing at the nodes.

During July and August, two obvious layers of vegetation
were present, Nuphar variegatum at the water's surface and
Elodea canadensis and Megalodonta Beckii below. From
autumn into the winter there were no actively growing
macrophytes at this site. Figure 2 illustrates these changes
at five selected dates during the year.

WATER DEPTH 2.0 M: This site was always dominated by
dense stands of Chara vulgaris L. Only in June was Megalo-
donta Beckii able to emerge above this species. Elodea
canadensis occurred only slightly higher than the Chara in
June, July, and August before dying in the fall. In late
August and early September Najas flexilis (Willd.) Rostk.
& Schmidt was present, but never grew higher than the
Chara. During August and September there were two
definite vegetational strata present. A diffuse overstory of
Vallisneria americana Michx. and Myriophyllum exalbes-
cens Fern. occurred above C. vulgaris. With the decompo-
sition of V. americana and M. exalbescens in November,
only Chara remained.

The first week in May dense stands of Chara vulgaris
were at their lowest height of the entire year. Chara grew
steadily until the middle of November, when the plants
started to decrease in size, presumably from basal decom-
position, since the apices remained healthy all winter.
Megalodonta Beckii and Elodea canadensis started growth
at the same time as in the shallower water but attained
somewhat greater heights. Megalodonta Beckii reached ap-
proximately 35 cm in height in August, decomposing in Sep-
tember. Elodea canadensis attained about 75 cm in height
in early September and then began decomposing. Myrio-
phyllum exalbescens appeared in early May, grew steadily
until September and then fell to the bottom with new shoots
appearing at the nodes. Vallisneria americana emerged
from the substrate the middle of June; grew more rapidly

503

Otsego Lake — Harman

1974]

ayeaysqng wos) wo

00L

SCL

osL

SLL

ooz

"'Ádougo esuep =
quepunge jo sjugreu Sutmoys 'exe'[ o3esj() ‘ea0p Fey ur ujdep ur ur (rg uum[oo 1976M '$ əng

:Kdoueo esngrip = - — — 'sə}ep papəps ye sejAudoipáu

LZ 1equia^oN GL LL 19quiejdeg GL LL 15nBny GL LL Ainp GL LZ eunf SL
p ET e euopoebew| | | [seeN Marco eloh RESCUE NEMENES
f | winyjAydouAy dips seleN eueusieA
eapo|3 ç eapo|3 y eyg
Errr anar E E NU gae
sefeN L— — euopogBew| [t | esuopojebayl|
DUPPEPTTPETTTRETIS ese
8 supose eu) winyAydouAW
eeyo — se ee
ee mee |)
eapo[3 enaus je A
me s — winyjAydouAW
eey
x S umpAydouAw |
Ruin) AqdouAw — —À IFETTHTIETN
E ——— o o o o -—
m eIJ8USI] [EA

504 Rhodora [Vol. 76

than its competitors, attaining its maximum height of about
125 cm in September, and then rapidly decomposed. Najas
flexilis emerged from the substrate in the middle of July,
grew rapidly to about 60 cm high in early September and
disappeared. Figure 3 shows the strata in 2.0 m of water
at various times during 1971.

WATER DEPTH 4.0 M: As at the 2 m site Chara vulgaris
grew in dense beds at a consistent rate throughout the
summer. Elodea canadensis was present until August as
before. Early in May Potamogeton pusillus L., P. zosteri-
formis Fern., P. Richardsonii, and P. illinoensis emerged
from the substrate. By 15 July P. Richardsonii and P.
illinoensis attained about 200 cm in height while P. pusillus
and P. zosteriformis reached only about 100 cm heights.
All of the species of Potamogeton were decomposing by
August, although a second cohort of P. Richardsonii re-
mained until early September. In late May, Heteranthera
dubia (Jacq. MacM. emerged from the substrate and
maintained steady growth until late September when it
collapsed to the substrate.

In June the morphology of the plant community was very
complex with 7 species of actively growing hydrophytes all
at different heights in the water column. By J uly 3 obvious
Strata were present: a diffuse overstory composed of Pota-
mogeton Richardsonii and P. illinoensis, a discontinuous
stratum of intermediate height composed of P. pusillus and
P. zosteriformis, and a dense understory of Chara, Elodea
and Heteranthera. In September this same area was en-
tirely different in appearance with 2 major strata present.
There was a rather dense overstory of Heteranthera dubia.
underlain by Chara vulgaris. By November the Chara was
all that remained, Figure 4 illustrates the physiognomy at
this depth at several selected dates.

WATER DEPTH 5.5M: In these deeper waters Nitella
flexilis replaced Chara vulgaris. Dense beds attaining
95 em occurred in this area. The only macrophyte asso-
ciated with this alga was Potamogeton crispus, which ap-

505

Otsego Lake — Harman

1974]

ajeusqng uio4j w3

"'Ádoueo əsuəp =

:Adouvd osngtrp = - — — ‘sazep pəpəjəs e sayAydorpAy
juepunge jo sju3reu 3urMous IYE 03ƏS7O ‘ea0g yey ut ujdep ur ur (yp uum[0o I939AA `$ 9In3lLq

: LZ 18quie^O0N GL LZ 1equieideg GL LL 1snBny GL LZ Aine GL LZ eunf GL
E: vi ADD & iuosp1euatH 'd eee "di PAIJETUJSU
?8po|3 snjjisnd ^q s d:
S6 4 sisuaout ^d T ix ay
x^ eyg
ejaqyuejejeH| JfePOIl27o 0 0
os - eeu
eueyd
BS ae v^ S1WI404}143}S0Z "d
J e9pol3 —y PAPPI3| fuuospueuoni “dw
GZ - -— [epo 7 I1uospjeuotH "d,
eyg
00L eJou1ue1839H snyisnd d ^ cy ——— |
| "X "siünojaaisoz “g]
GZL 4
Sisuaout| `d
pdec ci qM oa eee | ccc cc cud dpt M
IIUOSpJEU2IH ‘d
em £J9Q1UPJ4919H
SISUBOUIII! ^d. ~N
ooz 4 [^ R^ muospieuol "d
l IR |
eus ed

CN NN WoW

506 Rhodora [Vol. 76

peared from the substrate the first week in May. The latter
species grew extremely rapidly, attaining about 250 cm in
height by the middle of July. It then collapsed to the sub-
strate and decomposed.

In June and July two definite layers of vegetation were
present at these depths. Potamogeton crispus formed a
diffuse overstory while Nitella flexilis composed a dense
understory. From August on into the winter N. flexilis was
the only species present at the site (Fig. 5).

The macrophytes studied can be separated into four
groups according to their morphology during the winter
season. These characterize the appearance of the substrate
during this period and provide food and cover for the local
benthos. Group 1. Chara vulgaris, Nitella flexilis. Once
maturity is reached the oldest parts of these algae are con-
tinually decomposing. During the winter period entire
stands decrease in height as the rate of decomposition
greatly exceeds the rate of growth. The apical meristems
remain in a healthy condition throughout the year. In early
spring much new growth is added from germinating
oospores that cannot be distinguished from the older plants
remaining during the same time periods. Group 2. Myrio-
phyllum exalbescens, Elodea canadensis. When these or-
ganisms decompose (late September and late August, re-
spectively), the old growth lays on the substrate and winter
buds, 2-3 cm in height, appear at once from the nodes.
These grow extremely slowly over the winter reaching 4-6
cm by early May. At that time the roots have become estab-
lished, internodal tissues from the parent plants have rotted
away and the new plants grow rapidly until the fall when
the process is repeated. Group 3. Heteranthera dubia. Like
the organisms in Group 2, this plant collapses to the bottom
in a living condition (in mid-October) but the new shoots
do not appear at the nodes until late May or early June of
the next year. They grow rapidly unti] fall and the cycle
begins again. The parent stems remain intact throughout
much of the growing season. Group 4. Potamogeton cris-
pus, P. Richardsonii, P. illinoensis, P. zosteriformis, P.

507

Otsego Lake — Harman

1974]

jugpunqe jo SJT Surmoys faye] o3esj() ‘eao0g FEY UI u3dep ur ur g'g uumn[oo 137e M

ejensqns wos uio

‘Adouvs esuep =

LL saquianon GL

LZ 1equieydes GL

:Ádoueo esnyip = - - — ‘sazep pejoe[es ye sazAydorpAy

LL 1snBny GL

LZ Aine GL

'G ONS

LZ eunr SL

06

SL 71

ela}! N

ella}! N

ool 4

Sil =

pem

sndsi42 `d

elə} N

S S

ella} N

PION

sndsio 'd

sndsiJ2 ‘d

NS

= i 35 S Y

508 Rhodora [Vol. 76

pusillus, Vallisneria americana, Najas flexilis, Megalodonta
Becki, Nuphar variegatum. This group contains all the
remaining plants studied. These macrophytes grow rapidly
from buried winter buds or underground stems or root-
stocks and then decompose, shoots not appearing above the
substrate until the next growing season.

LITERATURE CITED

HARMAN, W. N. 1970. Aquatic biology studies, pp. 15-16 + Ap-
pendix pp. 1-142. In: Second annual report (1969-70) SUNY
Oneonta Biological Field Station, SUNY, Oneonta (mimeo).

1971. Biological studies — Otsego Lake, pp. 1-28. Im:
Third annual report (1971) SUNY Oneonta Biological Field
Station, SUNY, Oneonta (mimeo).

1972. Aquatic biology studies, pp. 4-15. Im: Fourth
annual report (1971) SUNY Oneonta Biological Field Station,
SUNY, Oneonta (mimeo).

, & T. R. DOANE. 1970. Changes in the aquatic flora of
Otsego Lake, New York between 1935 and 1969. N. Y. Fish
Game Jour. 17: 121-123.

, & J. L. Forney. 1970. Changes in the molluscan com-
munity in Oneida Lake, N.Y. between 1917 and 1967. Limn.
Ocean. 15: 454-460.

MUENSCHER, W. C. 1936. Aquatic vegetation of the Susquehanna
and Delaware areas. pp. 205-221. In: A biological survey of the
Delaware and Susquehanna watersheds. N. Y. S. Cons. Dept.,
Albany.

SCHULTHORPE, C. D. 1967. The biology of aquatic vascular plants.
610 pp. St. Martin's Press, New York.

Stuckey, R. L. 1971. Changes of vascular aquatic flowering plants
during 70 years in Put-in-bay Harbor, Lake Erie, Ohio. Ohio
Jour. Sci. 71: 321-342.

BIOLOGY DEPARTMENT
STATE UNIVERSITY COLLEGE
ONEONTA, NEW YORK 13820

PANICUM ENSIFOLIUM BALDW. —
NEW FOR MEXICO

MELVIN L. CONRAD

During the summer of 1972 a survey of grasses ( Conrad,
et al., 1972) was conducted for the Food Agriculture Or-
ganization of the United Nations at the Instituto Tecno-
lógico y de Estudios Superiores de Monterrey which is
developing a range management experiment station near
Tempoal, Veracruz. On July 19th the group of investiga-
tors accompanied the coordinators of the program, Dr.
Donald Huss and Ing. Edmundo L. Aguirre, on an excur-
sion south of Tempoal toward Molango, Hidalgo, to observe
the cloud forest vegetation (Leopold, 1950) near the crest
of the Sierra Madre Oriental. At a point 95.2 km. from
Tempoal, before reaching Molango, the caravan of two
automobiles stopped at a small parking area on the steep
mountain slope to turn around due to heavy fog, rain, and
an increasing number of fallen boulders on the highway.
In the parking area I was observing a tree fern (Cyathea
sp.) that grew beside the bluff from nearly 10 meters below;
and after a moment, I realized that I was standing in a
shallow wet depression about 5 meters in diameter that was
filled with a Panicum different from several species that
had been collected at lower elevations. Collections were
taken.

In the laboratory the grass from the parking area on the
mountain was identified as Panicum ensifolium Baldw.
Intermixed with it was P. cordovense Fourn., (Hitchcock
and Chase, 1910). Other species associated with these were
Juncus dudleyi Wieg., Carex festucacea Schkuhr, Agrostis
perennans (Walt.) Tuckerm., and Sporobolus poirettii
(Roem. & Schult.) Hitche.

It was first hypothesized that this population of P. ensi-
folium had been introduced, due to the considerable distance
separating this station from its previously reported range

509

510 Rhodora [Vol. 76

in the southeastern United States, (Hitchcock, 1950). How-
ever, it is likely to be native, since reports by Sharp (1946)
of disjunction in other taxa indicate similar affinities be-
tween the floras of these two regions.

The assistance of Dr. A. A. Beetle at the University of
Wyoming for verification of the identification, and the con-
firmation by both Dr. Beetle and Dr. Arturo Gómez-Pompa
at the Instituto de Biologia in Mexico City that P. ensi-
folium is new for Mexico is hereby acknowledged with deep
appreciation. Thanks are also due Dr. David B. Dunn for
the use of the herbarium at the University of Missouri.

Voucher specimens of P. ensifolium Baldw. (No. FAO
Z-101) are deposited in herbaria at the Departamento
Zootécnia, I.T.E.S.M., in Monterrey, México, Northeast
Missouri State University, University of Missouri at Colum-
bia (UMO), and The University of Wyoming (RM).

LITERATURE CITED

CoNRAD, M. L., W. K. MEDLER, S. NEFF, & C. PEPKING. 1972. 'Some
Grasses (Gramineae), Sedges (Cyperaceae), and Rushes (Junca-
ceae) of the Huasteca Zone in Veracruz and Hidalgo, Mexico.
Instituto Technológico y de Estudios Superiores de Monterrey.
Monterrey, México. Unpublished report.

Hitcucock, A. S. 1950. Manual of the Grasses of the United States,
U. S. Dept. Agric. Misc. Pub. No. 200. Revised by Agnes Chase.

, & A. CHASE. 1910. The North American Species of
Panicum. Contrib. U. S. Natl. Herb. 15: 1-396.

LEOPOLD, A. S. 1950. Vegetation Zones in Mexico. Ecol. 31: 507-
518.

SHARP, A. J. 1946. Informe preliminár sobre algunos estudios
fitogeográficos efectuados en México y Guatemala. Rev. Soc. Mex.
Hist. Nat. 7: 35-40.

DIVISION OF SCIENCE
NORTHEAST MISSOURI STATE UNIVERSITY
KIRKSVILLE, MISSOURI 63501

SIGNIFICANT NEW DISTRIBUTIONAL RECORDS
FOR THE GENUS SPHAGNUM IN THE
NORTHEASTERN UNITED STATES

RICHARD E. ANDRUS

Recent collecting and herbarium studies reveal interest-
ing new distributional records for several species of Sphag-
num in the northeastern United States. Most of the records
are from New York, and these are presented on the accom-
panying maps. Collection numbers, unless otherwise indi-
cated, are those of the author. Duplicates for most stations
collected by the author are at NYS, and many are also at NY
and SU-CF (=S.U.N.Y. College of Environmental Science
and Forestry).

Sphagnum henryense Warnst. A more difficult taxon than
the literature would indicate, S. henryense has frequently
been mistaken for S. palustre L., S. papillosum Lindb., or
S. imbricatum Russow. Forms of S. henryense lacking the
characteristic worm-like ridges on the hyaline cell walls
where they overlie chlorophyll cells are especially difficult
to identify. Such forms can usually be separated from the
very similar S. palustre by the pore characteristics of
the convex surface of the branch leaf. In S. henryense, the
hyaline cells have numerous small round pores along the
commissures, often 10 to 20 per cell, and usually lack large
membrane gaps near the leaf apex. S. palustre, on the
other hand, has hyaline cells with fewer and more elliptic
pores as well as membrane gaps near the leaf apex. Macro-
scopically, well developed forms of S. henryense may be
recognized by their reddish-brown pigmentation (often
visible in the capitulum) and a flattened capitulum with
small pointed head branches. In contrast, S. palustre is
browner and typically has a rounded capitulum with the
head branches blunter and more obese.

Sphagnum henryense is fairly widespread near the coast
from Long Island, N.Y. (Ketchledge 1957) south to Florida

511

[Vol. 76

Rhodora

512

pulchrum

S.

A = S, henryense

subfulvus

O=s.

torreyanua1

O-s.

[0] = S, tenellum

*
D

jensenii

= 8.

A-s.

strictun

1974] Sphagnum — Andrus 513

and westward to Texas. The Massachusetts station reported
here represents a northward extension of this coastal range.

The inland stations extending into western New York
and Pennsylvania are more surprising, however, but can
be considered, perhaps, as analogous to similar vascular
plant distributions noted by Fernald (1937). On this basis,
one might expect to find Sphagnum henryense in other
Great Lakes states such as Ohio and Michigan as well as
southern Ontario.

In comparing Sphagnum henryense and S. palustre
from inland stations, one particularly interesting pattern
emerges. Where it is found, S. henryense is normally an
obvious and abundant member of its plant community.
S. palustre, on the other hand, although found at many
more inland locations, typically occurs as a minor vege-
tational element, usually as a few scattered hummocks.
Limited observations on Long Island indicate the same
pattern may also hold in coastal areas. The factors re-
sponsible for this difference are unknown.

New York: ERIE C0.: Collins, Glowny 10850, 10848 (NYS). GENESEE
co.: Bergen Swamp, Durand 3642, 3624 (cu), Winne 1590 (cu),
S. J. Smith 47742, 47982 (NYS). CHEMUNG CO.: Winter Bog, 1594,
2168, 2170, Winne 2094, 2083 (NYS). ONONDAGA CO.: Cicero Swamp,
2852. SCHENECTADY CO.: Featherstonhaugh Lake, 2142, 2146. GREENE
co.: Grapeville, S. J. Smith 48750 (NYS). ORANGE CO.: Sterling
Forest, G. L. Smith 2666 (NY). CATTARAUGUS CO.: St. Bonaventure
Campus, Boehner, Aug. 1, 1940 (St. Bonaventure Coll.. Massachu-
setts: SUFFOLK CO.: Milton, Eaton & Faxon, Sphagna Boreali-Ameri-
cana #159. Pennsylvania: MCKEAN CO.: Smethport, Glowny 12007.

Sphagnum portoricense Sull. The previously known
northern limit for S. portoricense was on Long Island,
N.Y. (Mapes 1962). The northern range extreme is now
extended to Mt. Desert Island, Maine, where a collection
was made at a site strikingly different from those at the
stations formerly thought to be the most northerly. On
Long Island and in New Jersey (Andrews 1912) S. por-
toricense is aquatic in nature, being confined to shallow
sandy-bottomed ponds. On Mt. Desert Island it was clearly

514 Rhodora [Vol. 76

non-aquatic, with the plants being found at the margin
of Big Heath, a large open bog, where they formed a carpet
well above the water table along a small water course in the
shade of black spruce. One possible explanation may lie
in the high humidity characteristic of Big Heath, which is
situated very near the coast and often shrouded in fog.
S. portoricense also forms non-aquatic carpets in the very
humid environment of the high-elevation cloud forests of-
Puerto Rico.

Maine: Mt. Desert Island, Big Heath, 2030.

Sphagnum strictum Sull. The discovery of S. strictum in
New York was not unexpected because of its occurrence
northward and southward along the Atlantic coast (Maass
1966). The record from the Shawangunk Mountains of
upstate New York is, however, unusual in view of the close
affinity for the coastline that S. strictum exhibits in the
northern part of its range (Maass 1966).

Sphagnum strictum, like the closely related S. compactum
DC., is a pioneer species. It is found on more organic and
shaded substrates than the latter species.

New York: ULSTER CO.: Lake Minnewaska, S. J. Smith 48953
(NYS). SUFFOLK CO.: Riverhead, Third Pond, Latham 34419 (cu,
NYS); Sandy Pond, 2251, 2205, 2193, Mapes 1435 (Nvs) ; Swan Pond,
S. J. Smith 48158 (Nvs); Manorville, Latham 33936 (NYS, CU).

Sphagnum torreyanum Sull. (S. cuspidatum var. torreyi
(Sull.) Braithw.). Sphagnum torreyanum is reported here
for the first time from upstate New York. Like S. henry-
ense, it was previously known only from coastal locations,
in this case from Newfoundland to North Carolina (An-
drews 1938).

In upstate New York, as on Long Island, it occurs as an
aquatic in poor fens, often intermixed with S. cuspidatum
Hoffm.

New York: osWEGO CO.: Kibbe Lake, 1521, 1610. SARATOGA CO.:
Mulleyville Pond, S. J. Smith 46881, 46882 (Nvs); Luther's Woods,
2163.

1974] Sphagnum — Andrus 515

Sphagnum riparium Angstr. The range of S. riparium,
previously reported southward to New Hampshire, Michi-
gan, Wisconsin, and Washington (Andrus & Layser 1971),
is now further extended into New York. A robust northern
species, it typically grows in moderately minerotrophic
sites near the margins of poor fens associated with such
Sphagnum species as S. angustifolium (Russow) C. Jens.,
S. fallax (Klinggr.) Klinggr. and S. fimbriatum Wils.
Present collecting data indicates that S. riparium is a very
minor element in New York mire vegetation.

New York: ESSEX co.: Raybrook Bog, 1661; Peninsula Nature
Trails, 2152. FRANKLIN CO., Upper St. Regis Lake, 1707; 0.6 mi. E.
of Corey’s, 2816. HAMILTON CO.: 4 mi. S. of Long Lake, 1969; Johnny
Mack Bk. fen, 2846. oSWEGO co.: Kibbe Lake, 1528, 1529. CORTLAND
co.: Little York Lake, 2795. MADISON CO.: Nelson Swamp, 2778.
SCHENECTADY CO.: Featherstonhaugh Lake 2134. RENSSELAER CO.:
Round Pond outlet, 1884; !4 mi. NE. cf Round Lake, 1914.

Sphagnum pulchrum (Braithw.) Warnst. A subocean-
ically distributed species, in eastern North America S. pul-
chrum is known from the arctic (Persson & Sjórs 1960)
south along the coast to New Jersey (Andrews 1938) and
inland around the Great Lakes to Michigan (Crum & Miller
1969) and Wisconsin (Andrews 1938). The only previous
New York report (Ketchledge 1957) is from Big Moose
Station, Herkimer Co. (Peck £45, NYS). The numerous
New York records are not unexpected and fill in the known
distributional pattern.

Sphagnum pulchrum is a characteristic poor fen species
and occurs in both mire wide and mire edge vegetation,
often in abundance. Typical associated Sphagnum species
include S. papillosum, S. majus, S. fallax, S. angustifolium
and S. rubellum.

New York: CLINTON CO.: Cannon Flatrock, 1997a. FRANKLIN CO.:
Upper St. Regis Lake, 1808, 1811, 1346, 1725. ST. LAWRENCE CO.:
Sevey Pond, 1978; Wanakena, 1481; Grass River Flow, 1990; 1 mi.
E. of Sevey Corners, 1967, 1968. ESSEX CO.: Algonquin Peak, 1245,
1246, 1247, 1248, 1249; Raybrook Bog, 1283, 1288, 1671, 1282, 1821;
Lower Cherry Patch Pond, 1389; Whiteface Mt., 1215; Lake Tear,

516 Rhodora [Vol. 76

1333; between Scott and Wallface Ponds, 1351; near Scott Pond,
2425; Elk Lake, Phelps 100 (NYS). HERKIMER CO.: Big Moose Sta.,
Peck 45 (NYS). RENSSELAER CO.: Sand Lake, Peck 43 (CU, NYS);
1.5 mi. E. of Taborton, 2175, 2176, 2177, 2178; Bucks Corners, 1922.
SUFFOLK CO.: Napeague Beach, 2357; 1 mi. E. of Manorville, 2867;
Cedar Pond, N. G. Miller 6865 (NCH).

Sphagnum jensenii H. Lindb. (S. annulatum var. poro-
sum (Schlieph. & Warnst.) Maass & Isov.). A far northern
species, S. jensenii is noted for the first time for the U.S.
east of Michigan (Maass 1967b). The collection site was
at the margin of a large poor fen where it was growing in
a shallow pool in the bog mat associated with S. pulchrum,
S. papillosum, S. subsecundum Nees, S. flexuosum Dozy &
Molk and S. rubellum Wils. Maass (1967b) gives an excel-
lent account of the separation of S. jensenii from closely
allied species.

New York: ST. LAWRENCE CO.: Sevey Pond, 2829, 2832, 2833.

Sphagnum tenellum (Brid.) Brid. Sphagnum tenellum
was collected among sand dunes in a small bog pocket where
S. pulchrum was also found. As in the case of S. strictum,
this report fills a coastal distribution gap between Maine
and New Jersey.

Wet hollows in ombrotrophic mires are the most frequent
habitat of Sphagnum tenellum. Although such sites are
found elsewhere on Long Island, they are devoid of the
species. Lack of suitable habitat, therefore, is an unlikely
explanation for its rarity on Long Island. Climatic factors
are probably responsible, since on Long Island S. tenellum
is very near the southern extreme of its eastern North
American coastal distribution — this extreme being in New
Jersey.

New York: SUFFOLK CO.: Napeague Beach, 2238.
Sphagnum subfulvum Sjórs. The single previous con-

terminous U.S. report for S. subfulvum is from Michigan
(Crum & Miller 1969). It undoubtedly reaches its southern

1974] Sphagnum — Andrus 517

range limit in New York. The only other published records
for North America (Maass 1967a) give a range of “Green-
land to Bay of Fundy and N. Ontario and from Alaska into
western Yukon."

The collection sites were medium to rich fens, uncommon
mire types in New York, and this may account for its ex-
treme rarity in the state. A plant that typically forms low
to medium hummocks, Sphagnum subfulvum was found
associated with S. warnstorfii Russow, S. contortum K. F.
Schultz, S. centrale C. Jens., Scorpidium scorpioides (Hedw.)
Limpr., Calliergonella cuspidata (Hedw.) Loeske and Cam-
pylium stellatum (Hedw.) C. Jens. — all common rich mire
species.

Sjórs' (1944) original species description contains an
excellent discussion on the distinction of Sphagnum sub-
fulvum from the similar S. subnitens Russow & Warnst.
and S. flavicomans (Card.) Warnst.

New York: ESSEX CO.: Lost Pond, 1475. WARREN CO0.: Rush Pond,
1860, 1865; Glen Lake, 1854, 1885, 1889, 1886, 1890; Jenck's Swamp,
2815.

BIBLIOGRAPH Y

ANDREWS, A. L. 1912. Notes on North American Sphagnum, II.
The Subgenus Inophloea Russow. Bryologist 15: 1-9.
1913. Sphagnaceae. N. Am. Fl. 15: 1-31.
1938. The North American Atlantic species of Sphag-
mum. Ann. Bryol. 11: 15-20.
ANDRUS, R. E. and E. F. LAYSER. 1971. Sphagnum riparium
Angstr., a new record for the western United States. Bryologist
74: 211.
Crum, H. and N. G. MILLER. 1969. Bryophytes new to Michigan.
II. Michigan Bot. 8: 28-29.
FERNALD, M. L. 1937. Local plants of the inner Coastal Plain of
southeastern Virginia. Rhodora 34: 465-491.

IsovirTA, P. 1966. Studies on Sphagnum L. I. Nomenclatural re-
vision of the European taxa. Ann. Bot. Fenn. 3: 199-264.
KETCHLEDGE, E. H. 1957. Checklist of the mosses of New York

State. N. Y. S. Mus. Bull. 363: 55 pp. Albany, N. Y.
Maass, W. S. G. 1966. Untersuchungen über die Taxonomie und

518 Rhodora [Vol. 76

Verbreitung von Sphagnum VI. Sphagnum pylaesii Brid. und
das boreo-atlantische Florenelement unter den Torfmoosen in
Südamerika. Nova Hedw. 12: 81-105 + 5 pls.

1967a. Studies on the taxonomy and distribution of
Sphagnum II. Sphagnum angermanicum Melin in North Ameri-
ca and its relation to allied species. Nova Hedw. 13: 449-465
+ pls. 116-128.

1967b. Studies on the taxonomy and distribution of
Sphagnum IV. Sphagnum majus, Sphagnum annulatum, Sphag-
num mendocimum and Sphagnum obtusum in North America.
Nova Hedw. 14: 187-214 + pls. 69-81.

Mares, H. M. 1962. Sphagnum portoricense and Discelium nudum

in New York. Bryologist. 65: 68.

PERSSON, H., and H. SJörs. 1960. Some bryophytes from the Hud-
son Bay Lowland of Ontario. Sv. Bot. Tidskr. 54: 247-268.
SJORs, H. 1944. Sphagnum subfulvum Sjórs and its relations to
S. flavicomans (Card.) Warnst. and S. plumulosum Warnst.

P.P. Sv. Bot. Tidskr. 38: 403-427 + pls. 7, 8.

DEPARTMENT OF FOREST BOTANY AND PATHOLOGY
STATE UNIVERSITY OF NEW YORK

COLLEGE OF ENVIRONMENTAL SCIENCE AND FORESTRY
SYRACUSE, NEW YORK 13210

ENVIRONMENTAL CONTROL OF
NEEDLE CHARACTERISTICS IN
SUBALPINE BLACK SPRUCE

BRIAN F. CHABOT'”

Two distinct forms of Picea mariana, one upright and
the other more or less prostrate, exist in the vicinity of
treeline on several New England mountains. Characters
distinguishing the two types were discussed by Teeri
(1969) in establishing the varietal status of Picea mariana
(Mill) BSP. var. semiprostrata (Peck) Teeri. Individuals
of this taxon are recognized by their prostrate growth
form, lack of a dominant upright axis, and short, glaucous
needles that are slightly adaxially curved. The most prom-
inent character other than the growth form is the needle
length. Needles average 4 mm. (2.2 to 5.5 mm.) in length
in the prostrate form as compared with upright individuals
that have needles averaging 7.5 to 8.5 mm. The study
reported here indicates that needle characteristics and,
perhaps, other features of the prostrate variety are subject
to eonsiderable modification by the environment.

Plants were collected during late August and September
1972 from Lion's Head (1450 m.) and Cape Horn (1280 m.)
on Mt. Washington, N.H., and in Bear Swamp, Wolcott, Vt.
(340 m.). Both upright and prostrate forms were collected
at the treeline sites on Mt. Washington. Only the typical
form was obtained from Wolcott. Individual plants from
all populations were transferred to a commercial potting
mixture (Jiffy Mix) and grown in a cool greenhouse (10
to 20°C). All individuals were watered daily and fertilized
weekly with a 20-20-20 nutrient solution (Universal Chem-

1Present address: Section of Ecology and Systematics, Langmuir
Laboratory, Cornell University, Ithaca, New York 14850.

*Published with the approval of the Director of the New Hampshire
Agricultural Experiment Station as Scientific Contribution Number
685.

519

520 Rhodora [Vol. 76

ical Co.). Buds expanded during April, and both the new
needles and the previous year's needles produced under
natural conditions were analyzed in early July. Fifty
needles were selected from each indicated age class on
three separate individuals from each of the populations
studied. The samples were pooled according to age class
and source population. A final sample of 100 needles was
taken from each pooled sample for actual measurement.
Length measurements were taken to the nearest 0.1 mm.
Photographs of sample individuals were also taken.

New growth needles of the short needle var. semipro-
strata contrasted dramatically in length compared with
growth which had occurred under natural conditions (Fig.
1). The new growth needles were not only longer by about
5 mm. (Table 1), but they were darker green in color and
had no pronounced curvature. New needles of the other
populations of Picea mariana also had a healthier appear-
ance than the naturally produced needles, but, otherwise,
differences in needle length were not at all pronounced
(Fig. 1, Table 1).

Average needle lengths along with standard deviations
are presented in Table 1. Tests for significance were ap-
plied using the analysis of variance technique. The natural
needle length (short form) of var. semiprostrata was sig-
nificantly different (5% level) from the other sample
lengths including the needles produced in this variety
under cultivation. The needle lengths resulting from culti-
vation of all three populations were not significantly dif-
ferent.

Table 1. Average needle length and standard deviation for three
populations of Picea mariana.

Source Elevation (m.) Needle Length (mm.)
Natural Cultivated
A) 1450 (var. semiprostrata) 4.5 + 0.54 9.4 + 1.2
B) 1450 and 1280 (var. mariana) 8.5 + 0.97 8.1 + 1.2

C) 340 (var. mariana) 6.4 + 0.73 8.5 + 1.8

1974] Black Spruce — Chabot 521

Figure 1. Branches of Picea mariana with needles produced
naturally (below arrow) and under cultivation (above arrow).
Sources of the original populations are: A) 1450 m., Mt. Wash-
ington, var. semiprostrata; B) 1450 m., Mt. Washington, long-needle
form; and C) 340 m., Bear Swamp.

The results indicate that needle length in var. semipro-
strata is under some degree of environmental control.
Needle curvature is also capable of being modified. Curva-
ture appears to be related to needle length in that the
Shorter needles (less than 5 mm.) of all populations had
varying degrees of adaxial bending, while the longest
needles were most frequently straight. Modified needle
growth in black spruce under cultivation has previously
been observed, but not fully reported (Kozlowski, 1971;

522 Rhodora [Vol. 76

R. Pike, pers. comm.). The precise environmental factors
which operate here have not been isolated, but they cer-
tainly include both the improved nutrient and thermal
regimes of the greenhouse. The nutrient regime may be
particularly important in that the bog population, from an
environment which has been characterized as having low
nutrient availability (Small, 1972), also tended to have
shorter needles.

Additional information regarding the genetic structure
of var. semiprostrata can be obtained from observations
concerning the prostrate form itself and the fact that both
short- and long-needle types exist in close proximity to each
other at treeline (Teeri, 1969). The latter observation may
be taken as presumptive evidence of genetic distinctiveness
assuming that the forms are not actually distributed in
different microenvironments. The environmental situation
must be studied carefully as personal observation and the
studies of Tiffney (1972) indicate that slight differences in
microtopography, slope, and degree of exposure can make
considerable difference in the success of individual plants
at treeline.

The genetic nature of the prostrate growth habit is dif-
fieult to demonstrate experimentally because of the long
period of time necessary to develop this form. There is a
tendency for new twigs on the uppermost branches to occur
vertically. Frequently the preceding year's needles on these
protruding branches have been removed on the windward
side. Thus we may be witnessing an example of wind trim-
ming as has been observed in other climates (Thomas,
1973). Additionally, the work of Jaffe (1973) suggests that
the stunting of individuals could be an aspect of thigmo-
morphogenesis resulting from the mechanical action of
wind.

The contiguous natural distribution of characteristic
forms of both varieties as well as character variation which
is essentially clinal in nature (Teeri, 1968) perhaps has
resulted from disruptive selection similar to processes
described by Antonovics and Bradshaw (1970) and Snay-

1974] Black Spruce — Chabot 523

don (1970). Phenotypic variation which has little im-
portance in other parts of the range of Picea mariana may
assume considerable significance at alpine treeline. Several
morphological features of var. semiprostrata make it par-
ticularly well adapted to the alpine environment (Teeri,
1969). Additionally, it has frequently been observed that
growth rates, such as needle elongation, and cold-hardiness
are inversely related (Alden and Hermann, 1971). Thus,
at the distributional limit for a species, a uniquely adapted
genotype could exist sympatrically with the more common
genotype even in a homogeneous environment. Some degree
of reproductive isolation exists between the varieties due
to the infrequency of cone production at treeline (Teeri,
1968).

It is not the intention of this study to reorganize the
taxonomy of black spruce. The variety semiprostrata as
defined from natural specimens is sufficiently unique to
merit distinction. However, it should be recognized that
the genetic basis for this taxon lies not in a narrowly
specified phenotype, but in its plasticity and the sensitivity
of the genotype to interaction with the environment.

ACKNOWLEDGMENT

Appreciation is expressed to the Superintendent, White
Mountain National Forest, and Peter Marchand for their
assistance in certain phases of this study.

LITERATURE CITED

ALDEN, J., & R. K. HERMANN. 1971. Aspects of the cold-hardiness
mechanism in plants. Bot. Rev. 37: 37-142.

ANTONOVICS, J., & A. D. BRADSHAW. 1970. Evolution in closely
adjacent plant populations. VIII. Clinal patterns at a mine
boundary. Heredity 25: 349-362.

JAFFE, M. J. 1973. Thigmomorphogenesis: the response of plant
growth and development to mechanical stimulation. Planta 114:
143-157.

KozLowsKr, T. T. 1971. Growth and development of trees. Vol. 2.
Academic Press, N.Y. 333 pp.

524 Rhodora [Vol. 76

SMALL, E. 1972. Ecological significance of four critical elements
in plants of raised sphagnum peat bogs. Ecology 53: 498-503.

SNAYDON, R. W. 1970. Rapid population differentiation in a mosaic
environment, I. Response of Amthoxanthum odoratum popula-
tions to soils, Evolution 24: 257-269.

TEERI, J. A. 1968. The ecology of subalpine black spruce in New
England. M.S. Thesis, Univ. of New Hampshire. 62 pp.

1969. The phytogeography of subalpine black spruce in

New England. Rhodora 71: 1-6.

Tuomas, T. M. 1973. Tree deformation by wind in Wales. Weather
28: 46-58.

TIFFNEY, W. N., JR. 1972. Snow cover relationships of White
Mountain alpine plants. Ph.D. Thesis, Univ. of New Hampshire.

75 pp.
DEPARTMENT OF BOTANY

UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824

LINDERA MELISSAEFOLIUM IN ARKANSAS. An
excellent description of Lindera melissaefolium (Walter)
Blume (Lauraceae) and a detailed account of its known
distribution in 1949 were given by Steyermark (Rhodora
51:153-162). He suggested that the shrub might some day
be found in Arkansas because of its occurrence in adjacent
Ripley County, Missouri. The discovery of an Arkansas
locality in 1972 is not unexpected, therefore, but is here
recorded in view of the extreme rarity of the species.

Several colonies representing hundreds of plants are now
known from the northern part of Clay County, Arkansas.
The area is a part of the Mississippi Embayment region
and is characterized locally by a series of low woods in
which swampy depressions are surrounded by sandy knolls.
The shrub occurs in the water of these depressions, where
it forms dense stands which average 2-4 feet in height.
Some of the colonies occurring at the woods margins grow
in close association with the rare and interesting Corkwood,
Leitneria floridana Chapman.

Specific collection data for specimens from the Arkansas
locality are as follows: CLAY COUNTY: Low woods having
depressions with standing water surrounded by sandy
knolls, Sect. 2, T21N, R4E, 10 June 1972 (with R. Davis),
Tucker 9642 (UARK, NCU, APC).

G. E. TUCKER

DEPARTMENT OF BIOLOGICAL SCIENCES
ARKANSAS POLYTECHNIC COLLEGE
RUSSELLVILLE, ARKANSAS 72801

SPIRANTHES CASEI, A NEW SPECIES
FROM NORTHEASTERN NORTH AMERICA

PAUL M. CATLING AND JAMES E. CRUISE

In 1923 Henry Mousley made a collection of Spiranthes
orchids from the vicinity of Hatley (45° 11' N., 71° 56’ W.),
Stanstead Co., Quebec, He reported these plants in 1924,
under the name of Spiranthes cernua (L.) Rich. var. ochro-
leuca (Rydberg) Ames (Can. Field-Nat. 38: 61-63, 86-88,
p. 88). About 17 years later he sent these collections to
Dr. D. S. Correll at Harvard University, who was then
engaged in a study of the genus. Correll reported back to
Mousley that the plants from Hatley represented a mixed
collection of S. cernua (L.) Rich. var. odorata (Nutt.)
Correll and S. vernalis Engelm. & Gray. This was reported
by Mousley in 1941 (Can. Field-Nat. 55: 19-80, pl. 1-2),
and by Correll in 1950 (p. 227). Another station of S.
vernalis was discovered by Mousley during 1941 near Ste.
Dorothée (45° 32’ N., 73° 39' W.), Quebec, and described
in some detail by him in 1942 (Can. Field-Nat. 56: 1-2,
pl. 1-2). Gleason and Cronquist (1963) reported south-
eastern Quebec to be within the range of S. vernalis, a
decision probably based on the report of Correll (1950). In
recent floristic work concerning eastern Canada (e.g. Marie-
Victorin, 1964; Roland & Smith, 1963-64; Boivin, 1967) no
further mention has been made of S. vernalis, and surpris-
ingly the reports of Mousley and Correll have not been
repeated. However, Case ( 1964) discussed the occurrence
of plants referable to S. vernalis in the western Great Lakes
region, and Voss (1972, p. 461) has mapped the distribution
of S. vernalis in Michigan, although both of these authors
assigned their plants to S. vernalis with some hesitation.

Over the past three years a species of Spiranthes that
compared favorably with Mousley's specimens from Quebec
was found to be frequent and locally abundant in some parts
of the Canadian Shield region of southern Ontario, and it

526

1974] Spiranthes — Catling & Cruise 521

was listed for the province as S. vernalis by Whiting and
Catling (1971). A survey of herbarium collections in east-
ern North America revealed many new localities for this
«northern S. vernalis" in Ontario, several new stations in
Quebec, Michigan, and New England, and two localities in
Nova Scotia where it was previously unknown.

As specimens were examined over this wide range, it be-
came clear that the northern plants were quite distinct and
easily separated from Spiranthes vernalis as it occurs in the
southeastern states. A series of specimens from the Coastal
Plain and the gulf states, including the type of S. vernalis
Engelm. and Gray (AMES 82967), differed consistently from
the northern plants in exhibiting a light-colored and fre-
quently more dense, non-glandular pubescence on the rachis,
and a denser, less robust spike with a larger number of
narrower and slightly longer flowers. Plants collected from
Massachusetts, including the types of Spiranthes X inter-
media Ames (AMES 2246) and Spiranthes neglecta Ames
(AMES 2518) and the justifying specimens for Spiranthes
cernua X gracilis Ames (AMES 17 391), are comparable with
the “southern S. vernalis.” In his enumeration of North
American orchids, Ames (1924) included S. X intermedia,
S. neglecta and S. cernua X gracilis in synonomy with his
x S. vernalis. A general northern limit for the *southern
vernalis" is approximated by a line joining Massachusetts,
Kentucky, and northern Missouri. The northern plants re-
ferred to vernalis are distributed in a broad band from
Michigan to northern New England and the Canadian
maritime provinces. As these northern plants are quite
distinct and different from the southern plants (including
the type of S. vernalis), and as they appear to have a dis-
crete distribution pattern we are herewith describing them

as a new species.
Spiranthes casei Catling et Cruise sp. nov.

Differt a S. vernalis pubescentia septata et rubri-glandu-
losa in inflorescentia, spica robustiore et minus densa, flori-
bus paucioribus, latioribus, brevioribus.

528 Rhodora [Vol.

&
Splranthes vernalis Engeln, and Gray
rooms Ont = = MN a
fes

* Simcoe
"ese Moist adge cf aspen woods,

a Lot 4, eone, X, l k
Innlafil tpe

ma 30 Aug. 1370 FX" T, Razniosk 1470
Soul, bel 410 sm T. Reantook

- s Thera pee" a ‘come ot s 40 of thes^
overlockei orchids in this area.

Figure 1. A typical herbarium specimen of Spiranthes casei. The

elongate spike with flowers loosely arranged in a single spiral, the
erect oblanceolate leaves, and the robust nature of the plant are
characteristic. (T. Reznicek 470, Simcoe County Museum).

i

Figure 2. Part cf a spike of Spiranthes casei from Dorset,
Ontario. (Photograph by Dr. Erich Haber).

530 Rhodora [Vol. 76

Plantae (13-) 27-33 (-43) cm. altae sub anthesi. Folia
omnia laevia, saepe glauca, usque ad tertiam caulis partem
assurgentia; infima ad anthesin non permanentia, ovato-
lanceolata, 1-2 cm. lata, 7 cm. longa; superiora ad et non-
numquam per anthesin permanentia, oblanceolata vel lin-
eari-lanceolata, usque ad 15 vel 20 cm. longa, plerumque
minus quam 1 cm. lata, brevia, sub inflorescentia ad vaginas
sine laminis reducta. Caulis basaliter glaber, in rachidi pilis
septatis rubri-glandulosis 0.10-0.30 mm. longis pubescens.
Spica plerumque 6-10 cm. interdum ad 15 cm. longa, floribus
laxe et saepe in spiram unicam dispositis. Bracteae florales
ovatae vel ovato-lanceolatae, longi-acuminatae, 7-12 mm.
longae, basaliter plus minusve pubescentes. Flores cremei.
Ovarium sub anthesi 4-6 mm. longum, supra oblique tumi-
dum. Pubescentia in ovarii, sepali dorsalis, sepalorum lat-
eralium superficie pubescentiae rachidis similis. Sepala
lateralia 5-7 mm. longa; sepalum petalaque dorsalia sursum
vix curvata, 5-7 mm. longa; petala dorsalia pagina externa
papillosa. Labellum florum inferiorum 6-6.5 mm., superi-
orum 5-6 mm. longum; calli basales crassi, 0.8-1.0 mm.
longi, incurvi; labelli inferior superiorque centralis pagina
et callorum regio brevi-pubescens vel papillosa.

Different from S. vernalis in its septate, reddish-glandu-
lar pubescence in the inflorescence, and its less dense, more
robust spike with fewer, wider and shorter flowers.

Plants (13-) 27-33 (-43) cm. tall at anthesis. Leaves glab-
rous, often glaucous, ascending up to 1/3 the length of the
stem; lowest leaves, not persisting unti] anthesis, ovate-
lanceolate, 1-2 cm. wide, 7 cm. long; upper leaves, persisting
until and sometimes throughout flowering, oblanceolate or
linear-lanceolate, to 15 or 20 cm. long, and usually less than
1 cm. wide, reduced to bladeless sheaths below the inflores-
cence. Stem glabrous basally, becoming pubescent in the
rachis with septate, reddish-glandular hairs 0.10-0.80 mm.
long. Spike usually 6-10 cm. long, occasionally to 15 cm.
long, with the cream-colored flowers arranged loosely and
often in a single spiral. Floral bracts ovate or ovate-lanceo-
late, long-acuminate, 7-12 mm. long, more or less pubescent

1974] Spiranthes — Catling & Cruise 531

basally. Ovary 4-6 mm. long at anthesis, obliquely swollen
on the upper side. Pubescence on the surface of the ovary,
dorsal sepal, and lateral sepals, similar to that of the rachis.
Lateral sepals 5-7 mm. long. Dorsal sepal and petals barely
upcurved, 5-7 mm. long, the dorsal petals papillose on the
outer surface. The lip 6-6.5 mm. long in the lower flowers,
5-6 mm. long in the upper flowers. Basal calli of the lip
stout, 0.8-1.0 mm. long, incurved. The lower surface and
central upper surface of the lip, and region of the calli short
pubescent or papillose.

TYPE: CANADA: Ontario: MUSKOKA DISTRICT: ca. 6 mi.
w. of Bracebridge along highway 118 in Monck township,
(45° 02’ N., 79° 29’ W.), on dry hillside in full sun, Catling
& Whiting (TRT 169205).

REPRESENTATIVE SPECIMENS

CANADA: Nova Scotia: SHELBURNE C0.: Hope's Lot Barrens,
Clyde River, 7 Sept. 1921, M. L. Fernald & B. Long (AMES 85418).
Ontario: FRONTENAC CO.: Salmon Lake, Frontenac Park, Bedford
Tp., 28 Aug. 1968, R. Hainault & I. MacDonald (QK 97607). SIMCOE
co.: lot 4, cone. 10, Innisfill Tp., 30 Aug. 1970, T. A. Reznicek
(Simcoe Co. Museum 470) Quebec: STANSTEAD CO.: Hatley, 11 Sept.
1924, Mousley (QFA 42012, AMES 63699). UNITED STATES: Michi-
Zan: MARQUETTE CO.: summit of Huron Mountain, 1-9 Sept. 1916,
C. K. Dodge (MicH). New Hampshire: coos Co.: Colebrook, 21 Aug.
1942, A. S. Pease 29802 (NEBC). Vermont: ORLEANS CO.: Sutton Rd.,
Willoughby, 13 Sept. 1899 (Mrs. Bruche) (NEBC).

We dedicate this species to Mr. Frederick W. Case II,
whose book on orchids of the western Great Lakes region
has been enjoyed and admired by amateurs and profession-
als alike. Mr. Case has indicated that the northern plants
placed with Spiranthes vernalis differed from the southern
plants, and he repeated Ames' suggestion of a hybrid origin
for the northern plants (Spiranthes lacera X Spiranthes
cernua) here described as S. casei.

While such a hybrid origin is quite possible, several char-
acteristics of this plant suggest that it is worthy of specific
status. All plants in populations studied have been found to
be fertile, and many maintain themselves in the absence of
one or both putative parents. There is no evidence of back-

532 Rhodora [Vol. 76

cross-swamping of the distinctness of Spiranthes casei.
Although S. casei is intermediate between S. cernua and
S. lacera in many characters, certain other characters are
hard to rationalize in a hybrid; for example, the flowers of
S. casei are creamy-colored but white in both S. cernua and
S. lacera.

Spiranthes casei (Figs. 1 and 2) has been much confused
with other species of Spiranthes growing within its range,
particularly with S. cernua (L.) Rich. and S. lacera (Raf.)
Raf., but also with S. romanzoffiana Cham.

Spiranthes romanzoffiana, which also has creamy or pale
yellowish-green flowers, is readily distinguished by the
pandurate lip that is sharply constricted beyond the middle
(distally), and also by the connivent nature of the sepals
and petals which form the hood. Further, the flowers of
S. romanzoffiana are usually larger (perianth 8-10 mm.
long) and arranged in two or more rows in a compact spike.
Where S. casei and S. romanzoffiana occur sympatrically,
the latter reaches its peak of flowering in late July and
early August, while the former is in full flower in late
August and early September.

Spiranthes lacera has a singly-spiralled elongate spike
like that of S. casei, but it differs in being less robust, in its
large, oval, widely spreading leaves in a basal rosette, and
its smaller flowers (perianth usually ca. 4-5 mm. long) with
green colouration in the lip. S. lacera reaches its peak of
flowering in Ontario in late July and early August. We
have noticed that some plants of S. lacera have gone to seed
by late August when the first flowers of S. casei are open-
ing. Spiranthes lacera frequently occupies the same habi-
tats as S. casei, and both species grow in close proximity.

Spiranthes cernua may be distinguished from S. casei by
its shorter stature (20-30 cm.), usually short and compact
Spike (ca. 5-7 cm. long), and its larger, pure white flowers
(perianth usually 9-11 mm. long). In early September we
have encountered S. casei growing on dry sandy roadside
banks only a few feet removed from moist ditches with
abundant plants of S. cernua. On several occasions the

1974] Spiranthes — Catling & Cruise 533

plants of S. casei were noted to be more advanced in flower-
ing, with all of the flowers open and the lower flowers
fading. The nearby plants of S. cernua were less advanced
with the lower flowers fresh and many of the upper flowers
still in bud. Although this may be a result of microclimate,
it has nevertheless been a consistent observation.
Spiranthes casei shares some characteristies with S. cer-
nua (L.) Rich. var. ochroleuca (Rydb.) Ames, particularly
the cream-coloured flowers and a preference for an upland
habitat. The var. ochroleuca is stated to differ from the
typical form principally in yellowish-white flower colour,
longer floral bracts, monoembryonic seeds, and preference
for an upland habitat. Ames (1921, p. 78) has indicated
that the only reliable means of distinguishing var. ochro-
leuca is through microscopic examination of the seeds.
Apparently in other respects it is very close to typical S.
cernua, as is obvious from the plate provided by Ames (loc.
cit. pl. 127, Figs. 12-13), which was subsequently used in
“The Orchids of North America” (Correll, 1950, pl. 70,
12-13). Clearly, this drawing does not illustrate S. casei,
and the distinctive features of each make the separation of
S. casei from either variety of S. cernua relatively simple.

DISTRIBUTION AND ECOLOGY

The range of Spiranthes casei extends from Michigan
through Ontario and southern Quebec to the New England
states and Canadian maritime provinces (Fig. 3). It is
fairly frequent in the Muskoka-Haliburton region of On-
tario, where there are 34 stations (and probably many
more) separated from one another by distances of at least
one mile. Here it is usually found in dry, open locations
growing in coarse sandy soil in areas that have been dis-
turbed but are not continually disturbed. Associated plants
in such habitats include Danthonia spicata (L.) Beauv.,
Pteridium aquilinum (L.) Kuhn., and Polytrichum spp.
The barren tops of ridges (Fig. 4) are the only naturally
occurring habitats in which this orchid has been frequently
encountered.

534 Rhodora [Vol. 76

D:

=
4 | | J Ex ^]
( f JN o 700 300
` Pa k MEES
m— — —Á M

Figure 3. Distribution of Spiranthes casei based on specimens
examined in the following herbaria: ACAD, AMES, CAN, DAO, HAM,
LKHD, MICH, MTMG, NBM, NEBC, NFLD, OAC, QFA, QK, TRT, UWO.

Figure 4. Barren ridge-top on the Precambrian Shield in Matche-
dash Township, Simcoe County, Ontario. Spiranthes casei was quite
frequent on this ridge, growing in coarse shallow soil in open loca-
tions.

1974] Spiranthes — Catling & Cruise 535

The earliest collection of Spiranthes casei from Ontario
was made in 1904. Prior to 1965 it had been collected only
eight times in the province. However, during the period
from 1965-1972 more than 25 specimens have been added
to local herbaria. It is true that some of this represents our
own field work, but we cannot overlook the possibility that
this orchid has become more abundant in Ontario due to
human activity, such as land-clearing and road-building.
Of 14 collections made since 1970, 10 were from formerly
disturbed habitats such as roadsides, sand pits, abandoned
fields, and hydro-line transects. Spiranthes casei becomes
apparent in these sites several years after the initial dis-
turbance.

As with many other plants that quickly colonize habitats
created by human activity, Spiranthes casei has proved to
be at least partially apomictic. Seeds obtained from flowers,
from which pollinators were excluded with cloth bags dem-
onstrated 10-2596 germination.

ACKNOWLEDGEMENTS

The authors wish to thank Dr. B. Boivin (Central Ex-
perimental Farm, Ottawa) for his helpful criticism of this
manuscript and for suggesting the new name. Dr. L. A.
Garay (Harvard University) was of considerable help dur-
ing a visit to examine types in the Oakes Ames Orchid
Herbarium, and also kindly criticized our work, Miss S. M.
McKay, Mr. T. A. Reznicek and Mr. R. E. Whiting assisted
with field work. We are also indebted to the herbarium
curators who made specimens available for study: Dr. J. F.
Alex; Dr. D. S. Christie; Dr. L. Cinq-Mars; Mr. W. J. Cody;
Mr. C. E. Garton; Dr. A. E. Garwood; Dr. J. M. Gillet;
Dr. O. A. Olson; Dr. J. B. Phipps; Dr. J. S. Pringle; Mr.
T. A. Reznicek; Dr. E. A. Shaw; Mr. D. Strickland; Dr. S.
Van der Kloet; Dr. E. G. Voss; and Dr. D. W. Woodland.
Financial assistance has been provided by the National
Research Council of Canada through grant no. A-2363.

536 Rhodora [Vol. 76

REFERENCES

AMES, O. 1903. Natural Hybrids in Spiranthes and Habenaria.

Rhodora 5: 261-264, pl. 47.
1904. Spiranthes neglecta. Rhodora 6: 27-81, pl. 51.
1905. Contributions toward a monograph of the Ameri-
can species of Spiranthes. Orchidaceae, fasc. 1: 113-156.
. 1921. Notes on New England Orchids, — I. Spiranthes.
Rhodora 23: 3-85, pls. 127-129.
1924. An enumeration of the orchids of the United
States and Canada. Am. Orchid Soc., Boston, 120 pp.

BoiVIN, B. 1967. Enumeration des plantes du Canada. Naturaliste
Canadien 94: 131-157.

CasE, F. W. 1964. Orchids of the Western Great Lakes region.
Cranbrook Inst. Sci. 147 pp.

CORRELL, D. S. 1950. Native orchids of North America north of
Mexico. Chronica Botanica Co., Waltham, Mass. 399 pp.
GLEASON, H. A., & A. CRONQUIST. 1967. A manual of vascular
plants of the northeastern United States and adjacent Canada.

Van Nostrand Co., Princeton, New Jersey. 810 pp.

MARIE-VICTORIN, FRERE. 1964. Flore Laurentienne. Les presses
de l'universitie de Montreal, 925 pp.

Mousey, H. 1924. Further notes on the orchids of Hatley, Stan-
stead County, Quebec, 1923. Canadian Field-Nat. 38: 62-63,
86-88.

——————. 1941. Two orchids new to the Province of Quebec and
the Dominion of Canada, Spiranthes vernalis Engelm. & Gray
and Spiranthes cernua (L.) L. C. Rich. var. odorata (Nutt.)
Correll. Canadian Field-Nat. 55: 79-80, + 2 pl.

1942. The spring ladies’ tresses (Spiranthes vernalis
Engelm. & Gray) in Canada. Canadian Field-Nat. 56: 1-2, +
2 pl.

ROLAND, A. E., & E. C. SMITH. 1963-1964. The flora of Nova Scotia,
Part 1. Proc. Nova Scotian Inst. Sci. 26(2): 1-238.

Voss, E. G. 1972. Michigan Flora — Part I. Gymnosperms and
Monocots. Cranbrook Inst. Sci. & Univ. Mich. 488 pp.

WHITING, R. E, & P. M. CATLING. 1971. Checklist of Ontario
orchids. Fed. Ontario Nat., Newsletter 12(3): 9-10.

DEPARTMENT OF BOTANY
UNIVERSITY OF TORONTO
TORONTO, CANADA MDS 1A1

INVESTIGATIONS OF
NEW ENGLAND MARINE ALGAE VI:
DISTRIBUTION OF MARINE ALGAE
NEAR CAPE COD, MASSACHUSETTS?

DOUGLAS C. COLEMAN AND ARTHUR C. MATHIESON

Cape Cod is a major phytogeographic boundary on the
northeast coast of North America, delineating a distinctive
northern and southern marine flora (Chapman, 1964;
Humm, 1969; Setchell, 1922; Stephenson & Stephenson,
1949). Harvey (1852-1858) and Farlow (1870, 1882) first
recognized its importance as a phytogeographic boundary.
Additional information regarding the uniqueness of the
Cape Cod flora was contributed by Collins (1900), Davis
(1913a, b) and Taylor (1937, 1957).

Although the marine flora of Cape Cod has received con-
siderable attention since the time of Harvey, no one has
ever conducted simultaneous year round studies of the algae
on both sides of the Cape. In addition, nothing is known
about the seasonal changes that occur in the Canal, which
joins Cape Cod Bay and Buzzards Bay. Conover (1958)
and Sears (1971) conducted seasonal studies of the algae
in southern Cape Cod, but they gave no consideration to the
Canal. The purpose of the present investigation was to
atudy the horizontal and vertical distribution of seaweeds
at seven locations from Scituate to Woods Hole, Massachu-
setts, and reproduction including the Cape Cod Canal. The
seasonal occurrence of seaweeds at the same sites will be
discussed in another paper.

Monthly collections of all the conspicuous algae at seven
stations (Fig. 1) were made from January to December,
1969. Severe winter conditions (heavy ice and surf) ex-
isted from December to March, and they restricted some

"Published with the approval of the Director of the University of
New Hampshire Agriculture Experiment Station as Scientific Con-
tribution Number 656.

537

538 Rhodora [Vol. 76

collections. Specimens were collected on foot in the inter-
tidal zone and by SCUBA diving to a depth of 60 feet in the
subtidal zone. Diving in the Canal was done at slack water
(Anon. 1969a). The collections were brought to the Uni-
versity of New Hampshire for identification and processing.

Taylor (1937, 1957, 1960) was the primary source of
identification and nomenclature. A number of additional
references were consulted for the identification of species
and the determination of recent geographical records along
the Atlantic Coast of North America subsequent to Taylor's
(1960) summary. These include: Adey, 1964, 1965, 1966;
Bel & McFarlane, 1933a,b; Blomquist & Humm, 1946;
Cardinal, 1964, 1965, 1966, 1967a,b,c, 1968; Collins, 1909;
Edelstein, et al., 1967, 1969, 1970; Edelstein & McLachlan
1966, 1967a,b, 1968a,b, 1969; Fritsch, 1935, 1945; Hoek,
1964; Hoyt, 1920; Lamb & Zimmermann, 1964; Lee, 1968,
1969; Lewis, 1914; MacFarlane & Bell, 1933; McFarlane &
Milligan, 1965; Mathieson & Fuller, 1969; Mathieson, Dawes
& Humm, 1969; Rhodes, 1970; South & Cardinal, 1970;
Stone, et al., 1970; Wilce, 1959, 1965; Williams, 1948, 1949;
Wulf, et al., 1968; Zaneveld & Barnes, 1965; Zaneveld, 1965,
1966a,b, 1972. The nomenclature of Parke & Dixon (1968)
was applied in most cases.

Surface water temperature and salinity information was
recorded when each station was investigated. Temperature
was determined with a laboratory grade mercury thermom-
eter; salinity was measured with a set of hydrometers ( G.
M. Mfg., Co.). In addition, hourly and daily surface water
temperature data were obtained for each end of the Canal
from thermographs monitored by the Fisheries Division of
the State of Massachusetts. Additional (daily) temperature
and salinity information was supplied by the Woods Hole
Oceanographic Institute, Falmouth, Massachusetts (per-
sonal communication with Dean Bumpus and Joseph
Chase).

Figure 1 illustrates the location of the seven study sites.
One station (Scituate) was located north of the east end
of the Canal, four were in the Canal proper and two (Woods

1974] Marine Algae — Coleman and Mathieson 539

LI LI
BS SCITUATE
42° L` 7
CAPE COD BAY Ü x
»
4147 en CAPE COD CANAL 7
V a L/^O *
v
CAPE COD À
GS
ECK g y
4 f á |
u d f
<2 ( f
ay
o PI WOODS HOLE
70°30' 70510

Figure 1. Map of Cape Cod, Massachusetts, showing the seven
stations.

Hole and Wings Neck) were located south of the west end
of the Canal. The land-cut of the Canal cuts across the
Cape in an east-west direction from the town of Buzzards
Bay (approximately 41°44’ N Latitude, 70°37’ W Longi-
tude) to Sandwich (approximately 41°46’ N Latitude,
70°30’ W Longitude). It is seven miles in length, with an
additional seven miles of approach channels.

Several differences are obvious between the two sides of
the Cape. To the north, the substrate consists of granite
outcrops, boulders and cobbles, while to the south the sub-

540 Rhodora [Vol. 76

22 TaN
Í N
201 7 N
18]
16)
. 14
O
o 12
[99]
c 10!
O
A 8
6}
4 Jg
| 4 i STATION 1 ——
2 Da STATION 3 -——-
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0} cy 2 STATION 7 ----
PI TFITMITATMI I TI TATSTOINTI DI
MONTHS

Figure 2. Mean monthly variation of surface water temperatures
at stations 1, 3, 5 and 7 during 1969.

strate is largely sand and mud with fewer boulders. The
Cape is also the dividing line between the Labrador Current
to the north and the Gulf Stream to the south. Thus, a
marked difference in water temperatures occurs during the
summer. The yearly temperature range south of the Cape
was approximately 22°C., while that to the north was about
17°C. (Fig. 2). The salinity ranges south of the Cape were
usually higher than those to the north, particularly during
spring and summer (Fig. 3). The tidal amplitude is much
greater on the north than the south side. Thus, a three foot
tidal amplitude exists in Buzzards Bay, while it is approxi-
mately ten feet at Scituate (Anon. 1969a). The waters of
Cape Cod Bay are over 100 feet deep while those in Buz-
zards Bay are shallower.

541

Marine Algae — Coleman and Mathieson

1974]

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542 Rhodora [Vol. 76

STATION 1 ——
STATION 3 ———
STATION 5 -:—.-
25; STATION 7 -——-

ee

20:

151

o e
Ome — see — + — —— -— ——— e — -~
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104
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JANUARY

LES
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es

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Figure 4. Mean daily variation of surface water temperatures at
stations 1, 3, 5 and 7 during January and July, 1969.

1974] Marine Algae — Coleman and Mathieson 543

The Canal has a limited amount of solid substrate, no
wave action and a reduced intertidal zone. The seasonal
temperature ranges are intermediate between the north and
south sides of the Cape, but daily fluctuations during the
summer are greater than on either side of the Cape (Fig. 4).
The summer temperatures in the Canal are somewhat lower
in the east than the west end (Fig.2). Seasonal and spatial
variations of salinity are minimal (Fig. 3). The tidal
amplitude in the Canal ranges from four feet in the west
end to ten feet in the east end (Anon. 1969b). There is
a difference of about two hours between the tides at the
two ends (Anon. 1969b). The current reaches a peak of
about six knots, three hours after slack water. Slack water
occurs uniformly throughout the Canal, and it does not
coincide with either high or low water (Anon. 1969b). The
substrate at each site in the Canal grades vertically from
granitic boulders to sand-silt. In addition, there is a general
decrease in stable substrate towards the east end.

The Canal is maintained by the Army Corps of Engineers.
A private access road along the banks was used to reach
the stations. Telephone poles are located at 100 foot inter-
vals along the Canal; they are numbered beginning at the
east end. Stations 2-5 were located in the Canal and eor-
respond to poles 10, 45, 245 and 385 respectively. Station 3
(pole 45) is located in front of the Cape Cod Canal Power
Plant. A detailed description of the Canal stations, plus the
Scituate, Wings Neck and Woods Hole sites is summarized
in the Appendix.

SPECIES COMPOSITION

A total of 106 taxa was collected at the seven stations,
including 52 Rhodophyta, 35 Phaeophyta and 19 Chloro-
phyta. Table I summarizes the species composition at each
station. Scituate and Woods Hole showed the greatest
diversity of species, while stations 2-6 showed lower num-
bers. No subtidal collections were made at station 2. It is
of interest to note that the relative percentages of Rhodo-
phyta, Phaeophyta and Chlorophyta were about the same
at each station.

544 Rhodora [Vol. 76

60: NORTHERN ----
| SOUTHERN -=
COSMOPOLITAN

50;

40:

NUMBER OF TAXA/ GROUP
w
Q

\
\
\
7
204 \
; \
\ 7
\ m ^
MEMBRE d
104 V Ll
4
7
P — : =. =. -e
] æ. —. Á
0 :
| 2 3 4 5 6 7
STATIONS

Figure 5. Number of northern, southern and cosmopolitan species
at each station.

1974] Marine Algae — Coleman and Mathieson 545

A listing of the known geographical distribution of each
taxon along the Atlantic Coast of North America is also
designated numerically in Table I. The species can be
grouped into three components (i.e., northern, southern and
cosmopolitan) according to their major centers of distribu-
tion. The northern species are found most commonly north
of the Cape, and they include groups 1, 2, 3, 4 and 15. The
southern species are more common south of the Cape; they
include groups 7, 11, 12 and 13. Cosmopolitan species are
found on each side of the Cape, and they include groups 5,
6, 8, 9, 10, 14 and 16.

Figure 5 summarizes the number of northern, southern
and eosmopolitan species at each station. Most of the spe-
cies were cosmopolitan. The northern component declined
towards the south, while the southern component increased
towards the south, reaching a maximum at Wings Neck and
Woods Hole. Cosmopolitan species included Chaetomorpha
melagonium, Rhizoclonium tortuosum, Ulva lactuca, Asco-
phyllum nodosum, Petalonia fascia, Ahnfeltia plicata and
Ceramium rubrum, while conspicuous northern components
included Codiolum petrocelidis, Monostroma spp., Chorda
filum, Fucus spiralis, Laminaria digitata, Choreocolaz poly-
siphoniae, and Porphyra umbilicalis. Representative species
more common to the south of the Cape included Codium
fragile subsp. tomentosoides, Sargassum filipendula, Agard-
hiella tenera, Champia parvula and Hypnea musciformis.

Scituate had a species composition typical of northern
New England, while Wings Neck and Woods Hole were
more typical of warmer water locations. Thus, Laminaria
spp., Gigartina stellata, Ralfsia fungiformis, Chaetomorpha
atrovirens, Urospora collabens, and Fucus vesiculosus were
common at the former station, while Sargassum filipendula,
Agardhiella tenera, Callithamnion baileyi, and Codium frag-
ile subsp. tomentosoides were common at the latter stations.
The Canal stations, 3-5, included representatives from both
sides of the Cape (e.g. Laminaria saccharina, Chondrus
crispus, Rhodymenia palmata, Agardhiella tenera, Codium
fragile subsp. tomentosoides and Sargassum filipendula.

546 Rhodora [Vol. 76

VERTICAL DISTRIBUTION

Figure 6 summarizes the vertical distribution of the con-
spicuous species at each station. The distributions were
recorded as follows: 1) species restricted to the intertidal
zone; 2) species restricted to the subtidal zone; 3) species
found in both the intertidal and subtidal zones. Most of
the plants at Scituate were collected from the intertidal
and subtidal zones, including tide pools. Species that were
common to both zones included Chondrus crispus, Chorda
spp., Ectocarpus siliculosus, Petalonia fascia, Ceramium
rubrum, Corallina officinalis, Dumontia incrassata, and
Chaetomorpha linum. Few species (e.g. Chaetomorpha
atrovirens, Cladophora flexuosa, Polyides rotundus, Aspero-
coccus echinatus, and Laminaria spp.) were restricted to
the subtidal zone or intertidal zone (e.g. Codiolum spp.,
Enteromorpha intestinalis, Rhizoclonium tortuosum, Ulo-
thrix flacca, Choreocolax polysiphoniae, and Fucus spiralis).
Subtidal substrate was lacking at station 2, and the lowest
number of species was found here.

The Canal stations (2-5) had the highest subtidal com-
ponent. Species restricted to the subtidal zone in the Canal
were also found in the intertidal zone at other stations. The
most common subtidal species in the Canal included Lami-
naria spp., Chondrus crispus, Agardhiella tenera, Chorda
spp., Chordaria flagelliformis, Ulva lactuca, Petalonia fas-
cia, and Scytosiphon lomentarius. Species restricted to the
intertidal zone included Urospora penicilliformis, Ascophyl-
lum nodosum, Fucus spiralis, and Choreocolax polysiphon-
iae; those common to both zones included Ectocarpus sili-
culosus, Fucus vesiculosus, and Sphacelaria cirrosa.

The majority of species collected at Wings Neck and
Woods Hole was found in the subtidal zone and the inter-
tidal-subtidal zones. Few species were restricted to the
intertidal zone. Species restricted to the subtidal zone in-
cluded Agardhiella tenera, Sargassum filipendula, Callitham-
nion roseum, Dasya pedicellata, Seirospora griffithsiana,
Leathesia difformis, Chordaria flagelliformis, and Clado-

1974] Marine Algae — Coleman and Mathieson 547

INTERTIDAL
INTER-SUBTIDAL
SUBTIDAL

NUMBER OF SPECIES

STATIONS

Figure 6. Vertical distribution of species at each station.

phora spp. Species found in both zones included Chondrus
crispus, Melobesia lejolisii, Ectocarpus siliculosus, Fucus
vesiculosus var. spaerocarpus, and Chaetomorpha linum,

548 Rhodora [Vol. 76

while those restricted to the intertidal zone included Bangia
fuscopurpurea, Porphyra wmbilicalis, Fucus spiralis, and
Ulothrix flacca.

DISCUSSION

Setchell (1917) was one of the first workers to emphasize
the role of temperature in determining the geographical
distribution of seaweeds. He divided the oceans into 5°C.
intervals or isotherms, according to the maximum summer
water temperatures. Hutchins (1947) also confirmed that
5°C. intervals either favored or inhibited growth and re-
population of marine organisms. Williams (1948), Parr
(1933), Well and Gray (1960), and Humm (1969) have
reported a similar relationship between temperature and
species composition. As suggested by Setchell (1917) Cape
Cod is a dividing line between the 15° and 20°C. maximum
summer isotherms. Hence, it is not surprising that it is a
major phytogeographic boundary, with distinct floras oc-
curring north and south of the Cape.

Several factors are responsible for the marked difference
in summer temperatures between the north and south side
of Cape Cod. A southern extension of the Labrador Current
carries cold water as far south as the Cape, where it re-
mains throughout the year. In addition, the deep waters of
Cape Cod Bay warm up slowly during the summer. Davis
(1913a, b) described Cape Cod Bay as a “holding pocket”
of cold water. He further described the islands of Martha’s
Vineyard and Nantucket as barriers protecting Buzzards
Bay from the intrusion of cold waters off Gay Head. The
high surface water temperatures in Buzzards Bay result
from a northern extension of the Gulf Stream into the
shallow confines of Buzzards Bay. During the winter, cold
air lowers the temperature on both sides of the Cape and
the sharp differential of temperature disappears. Winter
temperatures in Buzzards Bay, however, may be somewhat
lower than in Cape Cod Bay, because of the shallowness of
the water of the former location.

1974] Marine Algae — Coleman and Mathieson 549

The transitional nature of the Cape Cod Canal is docu-
mented by its species composition. The Canal is dominated
by cosmopolitan species; northern species increase toward
the east end of the Canal, while southern components in-
crease toward the west end. Setchell (1922) also recognized
a cosmopolitan group of plants common to both sides of the
Cape — in contrast to more northern and southern elements.
The completion of the Canal in 1914 presumably provided a
direct route for spores to pass from one side to the other.
Stephenson (1944) has shown a similar transition of flora
and fauna near the Cape of Good Hope in Africa; on the
west coast the waters are relatively cold, while on the east
coast the shore is bathed by the warm waters of the Indian
Ocean. Seven major components are evident near the Cape
of Good Hope; the major components are the cosmopolitan,
warm- and cold-water elements.

Physical factors, such as tidal amplitude, wave action and
availability of solid substrate, determine the local distribu-
tion of species at the seven sites. Salinity was not con-
sidered a major factor in the areas studied because of the
small range recorded. Scituate had the highest number of
species; it is the most exposed site having ample substrate.
The lack of wave action, lower tidal amplitude, and reduced
substrate in the Canal were probably responsible for the
low number of species at stations 2-5. Woods Hole had the
second highest number of species, even though tidal ampli-
tude, wave action, and substrate were reduced south of the
Cape. The abrasive action of sand at Wings Neck resulted
in low species numbers.

A variety of physical factors, such as temperature, salin-
ity, light intensity, substrate and exposure, determine the
vertical distribution of seaweeds (Stephenson & Stephen-
son, 1949; Chapman, 1964). Most algae at Scituate were
collected from the intertidal and subtidal zones, while in
the Canal and south of the Cape the majority of species was
restricted to the subtidal zone. The former location (Scitu-
ate) is characterized by greater tidal amplitude and wave
action, both of which are necessary for the development of

550 Rhodora [Vol. 76

an extensive intertidal flora — assuming substrate is not
limiting. The vertical position of species varied from sta-
tion to station. Species found in the intertidal and subtidal
zones at Scituate were often restricted to the subtidal zone
in the Canal or south of the Canal.

ACKNOWLEDGEMENTS

We would like to thank Dr. A. Hodgdon for his critical
review of the manuscript. In addition we express our grati-
tude to Dr. R. Fralick, Dr. R. Turner, and Mr. John Hans-
man for their assistance with diving and collection of
specimens, to the Army Corps of Engineers at Cape Cod
for diving access to the Cape Cod Canal, and to Dean
Bumpus and Joseph Chase for the use of unpublished tem-
perature and salinity data.

APPENDIX
DESCRIPTION OF STATIONS

Scituate (station 1) is located about 30 miles north of
the east end of the Canal at approximately 42°12’ N latitude
and 70°43’ W longitude (Fig. 1). It is a semi-exposed site,
consisting of massive granite outcroppings. There are
several tide pools and a well developed intertidal zone. The
substrate in the subtidal zone consists of large outcroppings,
boulders, cobbles and sand. The surface water temperatures
ranged from approximately 2-16°C. The highest tempera-
tures occurred in August, while the lowest occurred in
March. The daily temperature range was about 2°C. during
the summer and 3°C. during the winter. The average
monthly summer temperatures at Scituate were the lowest
of all the stations. Salinity values remained relatively con-
stant throughout the year, ranging from a low of 30.6°/,,
in the summer to a high of 32.5°/,, during the winter.

Station 2 is located at the east end of the Canal at the
Scusset Breakwater (Fig. 2). Collections were restricted
to the intertidal zone because of the lack of subtidal sub-
strate. An extensive intertidal zone is present. The water

1974] Marine Algae — Coleman and Mathieson 551

temperatures ranged from 12°C. to 19°C. The tidal ampli-
tude is 10 feet (Anon. 1969a).

Station 3 is located at the power plant (Fig. 2). Rip-rap
extends to about 6 feet below M.L.W., while a shelf extends
out 30-40 feet and to a depth of 15 feet. During operation,
the power plant discharges water of over 23.9°C. Substrate
is limited and a small intertidal zone is present. The tidal
amplitude is approximately 10 feet.

Station 4 is approximately 2.5 miles east of the west end
of the Canal proper (Fig. 2). The study site was on the
south bank of the Canal. The substrate grades from rocks
(boulders) in the upper shore to sand-silt in the lower sub-
tidal zone. The tidal amplitude is about 6 feet (Anon.
1969a). A limited intertidal zone is exposed during low tide.

Station 5 is located at the Engineer’s station at the west
end of the Canal. Collections were made in the vicinity of
the pier. Rip-rap extends to about 12 feet below M.L.W.,
while a shelf extends out to about 75 feet and slopes to
about 20 feet below M.L.W. The tidal amplitude is about
4 feet (Anon. 1969a).

Wings Neck (station 6) is located at approximately
41°31’ N latitude and 70°40’ W longitude in the town of
Bourne, Massachusetts (Fig. 1). It is a semi-exposed area
subject to southwest winds during the summer. A limited
intertidal zone is present; it is composed of small cobbles
and a few large boulders. A few shallow tide pools are
evident. The tidal amplitude is about 4 feet (Anon. 1969a).
The range of water temperatures was 0.2-22.19C. The mean
monthly winter temperature (2.2°C.) was lowest in Janu-
ary. At that time the temperature was lower than at Scitu-
ate, the northernmost station. The area is subject to severe
ice scouring during the winter. The salinity remained rela-
tively constant throughout the year, with a range of about
0.9°/oo (Fig. 4).

Woods Hole (station 7) is located at approximately
41°31’ N latitude and 70°40’ W longitude in the town of
Falmouth, Massachusetts (Fig. 1). Collections were made
at the jetty behind the U. S. Fisheries Building. The bottom

552 Rhodora [Vol. 76

is approximately 20 feet below M.L.W., at the deepest point,
and the bottom substrate is composed of sand and silt. Some
small rocks were present at one side of the jetty. The tidal
amplitude is about 2 feet (Anon. 19692). The temperature
ranged from a low of 0.2°C. in January to 22.5°C. in Au-
gust (Fig. 3). Daily temperature ranges were small during
the year (Fig. 5). The salinity remained relatively con-
stant throughout the year with a range of only 0.89/,
(Fig 4).

Footnotes to Table I

! Geographical distribution zones relating to Table I, right hand column.

T

Northern Massachusetts to Newfoundland and north.
Northern Massachusetts to Nova Scotia.

Southern New England-Long Island Sound to Gaspé and Labrador.
Southern New England-Long Island to Newfoundland and north.
New Jersey-Maryland to Newfoundland and north.

New Jersey-Maryland to Gaspé and Labrador.

New Jersey-Maryland to ‘Cape Cod.

North Carolina to Newfoundland and north.

North Carolina to Gaspé.

South Carolina to Newfoundland and north.

. South Carolina to northern Massachusetts.

Tropies to southern New England-Cape Cod.

. Tropies to northern Massachusetts.

Tropics to Newfoundland and north.

. Southern New England-Long Island Sound to northern New Eng-
land-Nova Scotia.

16. Tropies to northern New England-Nova Scotia.

| 9l st

Ree RH HM
NC CET

? — Neoagardhiella baileyi (Harvey ex Kützing) Wynne et Taylor.
(Wynne and Taylor, 1937).

3 — Phyllophora truncata (Pallas) Newroth et Taylor. (Newroth and
Taylor, 1971).

* — Phyllophora pseudoceranoides (Gmelin) Newroth et Taylor.
(Newroth and Taylor, 1971).

553

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yplesy ^f CT) suvowuiqum v4fiqdaoq
Ypresy ‘O (upiedy 9) omuru duhyd.og
1e1nug, 2222250912] n4fia[d.0q
AAD (ULMA LA 70074431T) 727]0224n v1uoudisfijoq
J0 AeL 20:28u0-20a0w muoydishjod
Əə (uospng) su22s2401w viwo1disfijoq
Apue[ (T) 2s0u»] nuoydishjog
Kepreq ?/iaa40u o1uowdisfijoq
Jesueidg (uospnpg) 2326u0]? Diuoydishjog
JexooH ut AoA1eH x2 o[[IA9.12)
(Usma) 9?vpnuop viuowudisfijoq
Ər (uospnH) snpungo4 saprhjog
uoxu]

‘quod e3&uydopoud — I a TAV iL

1974] Marine Algae — Coleman and Mathieson 559

REFERENCES

ADEY, W. H. 1964. The genus Phymatolithon in the Gulf of Maine.
Hydrobiologia, 24: 377-420.

1965. The genus Clathromorphum (Corallinaceae) in
the Gulf of Maine. Ibid. 26: 539-573.

1966. The genus Lithothamnion, Leptophytum (nov.
gen.) and Phymatolithon in the Gulf of Maine. Ibid. 28: 321-
370.

ANONYMOUS. 1969a. Tide tables, high and low water predictions
for 1969, North and South America including Greenland. En-
vironmental Sciences Services Administration, Coast and Geo-
detic Survey. 280 pp.

-.—————, 1969b. Tide tables, Cape Cod Canal. U. S. Army
Engineer Division, New England Corps of Engineers. 12 pp.

BELL, H. P., & C. MACFARLANE. 1933a. The marine algae of the
maritime provinces of Canada, I. List of species with their
distribution and prevalence. Can. Jour. Res. 9: 265-279.

1933b. The marine algae of the maritime provinces
of Canada, II. A study of their ecology. Ibid. 9: 280-293.

BLoMQUIST, H. L., & H. J. HUMM. 1946. Some marine algae new
to Beaufort, North Carolina. Jour. Elisha Mitchell Sci. Soc. 62:
1-8.

CARDINAL, A. 1964. Étude sur les Ectocarpacées de la Manche.
Beihefte zur Nova Hedwigia. 15. 86 pp.

1965. Liste préliminaire des algues benthiques de la
baie des Chaleurs. Rapp. Ann. 1964, Sta. Biol. mar. Grande
Riviere: 41-51.

1966. Additions a la liste benthiques de la baie des
Chaleurs. Rapp. Ann. 1965, Sta. Biol. mar. Grande Riviere:
85-43.

1967a. Inventaire des algues marines benthiques de la
baie des Chaleurs et de la baie de Gaspé (Québec), I. Pheo-
phycees. Nat. Can. 94: 233-271.

1967b. Ibid. II. Chlorophycees. Ibid. 94: 4471-469.

1967c. Ibid. III. Rhodophycees. Ibid. 94: 735-760.

1968. Répertoire des algues marines benthiques de l'est
du Canada. Cah. Inf., St. Biol. mar. Grande Riviere, Que. No.
48. 213 pp.

CHAPMAN, V. J. 1964. The Algae. Macmillan and Company Ltd.,
London. 472 pp.

CoLLINS, F. S. 1900. Preliminary lists of New England plants V.
Marine algae. Rhodora 2: 41-52.

1909. The green algae of North America. Tufts Col-
lege Studies. (Sci. Ser.) 2: 79-480.

560 Rhodora [Vol. 76

CONOVER, J. T. 1958. Seasonal growth of benthic marine plants
as related to environmental factors in an estuary. Publ. Inst.
Mar. Sci. 5: 97-147.

Davis, B. M. 1913a. General characteristics of the algal vegeta-
tion of Buzzards Bay and Vineyard Sound in the vicinity of
Woods Hole. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries
31: 443-544.

1913b. A catalogue of the marine flora of Woods Hole
and vicinity. Dept. Comm. and Labor, Bull. (U.S.) Bur. Fisheries
31: 795-833.

EDELSTEIN, T. L. CHEN, & J. MCLACHLAN. 1970. Investigations
of the marine algae of Nova Scotia, VIII. The flora of Digby
Neck Peninsula, Bay of Fundy. Can. Jour. Bot. 48: 621-629.

EDELSTEIN, T., & J. MCLACHLAN. 1966. Investigations of the marine
algae of Nova Scotia I. Winter flora of the Atlantic Coast. Can.
Jour. Bot. 44: 1035-1055.

& . 1967a. Ibid. III. Species of Phaeophy-
ceae new or rare to Nova Scotia. Ibid. 45: 203-9210.

& . 1967b. Ibid. IV. Species of Chlorophy-
ceae new or rare to Nova Scotia. Ibid. 45: 211-214.

& . 1968a. Ibid. V. Additional species new
or rare to Nova Scotia. Ibid. 46: 993-1003.

& . 1968b. Ibid. VI. Some species new to
North America. Ibid. 47: 555-560.

& . 1969. Petroderma maculiforme on the

coast of Nova Scotia. Ibid. 47: 561-563.

; , & J. S. CRAIGE. 1967. Investigations of the
marine algae of Nova Scotia II. Species of Rhodophyceae new
or rare to Nova Scotia. Ibid. 45: 193-202.

, .& . 1969. Preliminary survey of
the sublittoral flora of Halifax County. J. Fish. Res. Br. Canada
26: 2703-2713.

FanLow, W. G. 1870. XVII. List of sea-weeds or marine algae of
the south coast of New England pp. 281-294. In: U. S. Comm.
of Fish and Fisheries. Commissioner’s Report 1871-1872. Wash-
ington. Government Printing Office.

. 1882. I. The marine algae of New England. Ibid.,
report 1879. pp. 1-210.

FRITSCH, F. E. 1935. The structure and reproduction of the algae,
I. xviii + 791 pp. Cambridge Univ. Press.

1945. Ibid., II. xiv + 939 pp. Cambridge Univ. Press.

Harvey, W. H. 1852-58. Nereis Boreali-Americana. I. Melano-
spermae. Smithsonian Contrib. Knowl., 3: 1-150, pl. 1-12.
1852; II. Rhodospermae. Ibid. 5: 1-258, pl 13-36. 1853; III.
Chlorospermae. Ibid. 10: ii + 1-140, pl. 37-50. 1858.

1974] Marine Algae — Coleman and Mathieson 561

HoEK, C. VAN DEN. 1963. Revision of the European species of
Cladophora. xi + 248 pp. Leiden.

Hovr, W. D. 1920. Marine algae of Beaufort, N. C. and adjacent
regions. Bull. U. S. Bur. Fish. 36: 367-556.

HuMM, H. J. 1969. Distribution of marine algae along the Atlan-
tic Coast of North America. Phycologia 7: 43-53.

Hutcuins, L. W. 1947. The bases for temperature zonation in
geographical distribution. Ecol. Monographs 17: 325-335.

LAMB, M. & H. ZIMMERMANN. 1964. Marine vegetation of Cape
Ann, Essex County, Massachusetts. Rhodora 66: 217-254.

LEE, R. K. S. 1968. A collection of marine algae from Newfound-
land, I. Introduction and Phaeophyta. Nat. Can. 95: 957-978.

1969. Ibid., II. Chlorophyta and Rhodophyta. Ibid. 96:
123-145.

Lewis, I. F. 1914. The seasonal life-cycle of some red algae at
Woods Hole. Plant World 17: 31-35.

MACFARLANE, C., & H. P. BELL. 1933. Observation of the seasonal
changes in the marine algae in the vicinity of Halifax with
particular reference to winter conditions. Proc. Nova Scotian
Inst. Sci. 18: 134-176.

, & G. M. MILLIGAN. 1965. Marine algae of the Maritime
Provinces of Canada. A preliminary checklist. Nova Scotia Res.
Found., Seaweed division. Halifax. Mimeographed, 24 pp.

MATHIESON, A. C., & S. FULLER. 1969. A preliminary investigation

of the benthonic marine algae of the Chesapeake Bay Region.

Rhodora 71: 524-534.
, C. Dawes, & H. J. HvMM. 1969. Contributions to the

marine algae of Newfoundland. Rhodora 71: 110-159.

NEwRoTH, P. R., & A. R. A. TAYLOR. 1971. The nomenclature of
the North Atlantic species of Phyllophora Greville. Phycologia
10: 93-97.
PARKE, M., & P. S. Drxon. 1968. Checklist of British marine algae
— second revision. J. Mar. Biol. Assoc. U. K. 48: 783-832.
Parr, A. E. 1933. A geographic-ecological analysis of the sea-
sonal chenges in temperature conditions in the shallow water
along the Atlantic Coast of the U. S. Bull. Bingham Oceanogr.
Coll., 4. 90 pp.

Ruopes, R. G. 1970. Seasonal occurrence of marine algae on an
oyster reef in Burston's Bay, Virginia. Chesapeake Sci. 2: 61-63.

Sears, J. R. 1971. Morphology, systematics and descriptive ecology
of the sublittoral benthic marine algae of southern Cape Cod and
adjacent islands. Ph.D. Thesis, 295 pp. University of Massa-
chusetts.

562 Rhodora [Vol. 76

SETCHELL, W. A. 1917. Geographical distribution of the marine
algae. Science 45: 197-204.

. 1920. Stenothermy and zone-invasion. Am. Nat. 54:
385-397.

. 1922. Cape Cod in its relation to the marine flora of
New England. Rhodora 24: 1-11.

SovrH, G. R., & A. CARDINAL. 1970. A checklist of marine algae
of eastern Canada. Can. J. Bot. 48: 2077-2095.

STEPHENSON, T. A. 1944. The constitution of the intertidal fauna
and flora of South Africa. Part II. Ann. Natal. Mus. 10: 261-358.

, & A. STEPHENSON. 1949. The universal features of
zonation between tide-marks on rocky coasts. Jour. Ecol. 3T:
289-305.

STONE, R. A., E. HEHRE, J. CONWAY, & A. C. MATHIESON. 1970. A
preliminary checklist of the marine algae of Campobello Island,
New Brunswick, Canada. Rhodora 72: 314-338.

TAYLOR, W. R. 1937. The marine algae of the northeastern coast
of North America. 405 pp. Univ. of Michigan Press, Ann Arbor.

1957. Marine algae of the northeast coast of North
America. 509 pp. Ann Arbor Press.

1960. Marine algae of the eastern tropical and sub-

tropical coasts of the Americas. 870 pp. Ann Arbor Press.

WELL, H. W., & I. E. Gray. 1960. Summer upwelling off the
northeast coast of North Carolina. Limnol. Oceanogr. 5: 433-
468.

WiLCE, R. T. 1959. The marine algae of the Labrador Peninsula
and northwest Newfoundland (ecology and distribution). Nat.
Mus. Canada 158: 1-103.

1965. Studies in the genus Laminaria III. A revision
of the North Atlantic species of the simplices section of Lami-
naria. Botanica Gothoburgensia III. pp. 247-256.

WILLIAMS, L. G. 1948. Seasonal alternation of marine floras at
Cape Lookout, North Carolina. Am. Jour. Bot. 35: 682-695.

1949. Marine algal ecology at Cape Lookout, North
Carolina. Bull. Furman Univ. 31: 1-21.

WULFF, B. L., E. M. T. Wurrr, B. H. RoBINSON, J. K. Lowry, &
H. J. HuMM. 1968. Summer marine algae of the jetty at Ocean
City, Maryland. Chesapeake Sci. 9: 56-60.

WYNNE, M. J., & W. R. TAvLoR. 1973. The status of Agardhiella
tenera and Agardhiella baileya (Rhodophyta, Gigartinales).
Hydrobiologica 43: 93-107.

ZANEVELD, J. S., & W. D. BARNES. 1965. Reproductive periodicities
of some benthic algae in lower Chesapeake Bay. Chesapeake
Sci. 6: 17-32.

1974] Marine Algae — Coleman and Mathieson 563

ZANEVELD, J. S. 1965. The benthic marine algae of Virginia. Vir-
ginia Jour. Sci. 1965: 346.
1966a. The marine algae of the American coast be-
tween Cape May, N. J. and Cape Hatteras, N. C., I. The Cyano-
phyta. Bot. Mar. 9: 101-128.
1966b. The benthic algae of Delaware. A preliminary
checklist. 35 pp. Scientific Series, Publ. no. 2, Instit. of Ocean-
ogr., Old Dominion College, Norfolk, Virginia.
. 1972. The benthic marine algae of Delaware, U.S.A.
Chesapeake Sci. 13: 120-138.

JACKSON ESTUARINE LABORATORY
UNIVERSITY OF NEW HAMPSHIRE
DURHAM, NEW HAMPSHIRE 03824

A REVISION OF NORTH AMERICAN POTAMOGETON
SUBSECTION PUSILLI (POTAMOGETONACEAE)'

RoBERT R. HAYNES

The importance of the aquatic environment is becoming
more evident with our present ecological concerns. Many
aquatic plants have been shown to be indicators of water
quality and of changes in the aquatic environment (Volker
& Smith, 1965; Lind & Cottam, 1969; Harman & Doane,
1970; Stuckey, 1971; Wentz & Stuckey, 1971). Certain
species, e.g. Najas minor, Potamogeton foliosus, P. pusillus,
and Vallisneria americana, are tolerant of polluted waters.
Other species, e.g. Najas flexilis, Potamogeton amplifolius,
P. friesii, and P. strictifolius are intolerant.

Early workers, e.g. Pond (1905) and Shelford (1918,
1937), considered aquatic vascular plants of little value in
the food chain of aquatic animals. Later workers, e.g.
Moore (1915), Moyle and Hotchkiss (1945), and Welch
(1935, 1952), however, considered the aquatic vascular
plants an important source of food for aquatic animals. In
a survey of literature on the significance of aquatic vascular
plants in the nutrition of animals, Gaevskaya (1966) con-
sidered the aquatic vascular plants as a primary source of
food for animals of the aquatic environment.

According to Martin (1951), the pondweeds (Potamoge-
ton) are probably more valuable to ducks than any other
group of plants in the United States. Trautman (1957)
indicates that the distribution in Ohio of certain species
of fish is contingent upon that of associated pondweeds.
Gaevskaya (1966) lists 124 species of animals (89 inverte-
brates and 35 vertebrates) that feed upon Potamogeton.

'Contribution from the Botany Department, The Ohio State Uni-
versity (Paper No. 867). Portion of a Ph.D. dissertation written
at The Ohio State University. Field work for two summers was
supported by The University of Michigan Biological Station and for
one summer in part by a Research Grant from the Ohio Academy
of Science and Sigma Xi Grant-in-Aid for Research.

564

1974] Potamogeton — Haynes 565

The plants also serve as shelter for panfish and young fish
of other species. Many aquatic insects have at least one
stage of their life cycle, if not their entire life cycle, upon
the vegetative structures of the pondweeds (Welch, 1952).
These insects are, in turn, preyed upon by fish and some
birds. Müller-Liebenau (1956) found 294 invertebrates in
beds of five species of Potamogeton in the lakes of eastern
Holstein. The pondweeds are, therefore, a major factor in
the food cycle of the aquatic environment. In fact, among
the submerged and floating-leaved species of aquatic vas-
cular plants, Gaeyskaya (1966) states that as regards
distribution, productivity, and trophic relations of aquatic
animals, Potamogeton is the most important genus.

Potamogeton Linnaeus is a cosmopolitan genus with ap-
proximately 100 species of submerged and floating-leaved
aquatic plants. The genus has been separated into two
subgenera (Raunkiaer, 1896) and numerous sections and
subsections (Graebner, 1907; Hagstrom, 1916). The North
American representatives of one of the latter, Potamogeton
subsection Pusilli Graebner (narrow-leaved pondweeds),
are the subject of this paper. The Pusilli can be separated
from the other taxa of the genus by their 3-7(-9)-nerved
leaves that are 2-5 mm broad, by their stipules free from
the blade, and by their small (to 4.0 mm long) fruits that
normally are produced terminally.

Although the Pusilli have been included in at least four
revisionary treatments (Morong, 1893; Ascherson, 1907;
Hagstrom, 1916; Fernald, 1932), uncertainty still exists as
to the number of taxa to be included in the group and at
which rank these taxa should be recognized. Taylor (1909),
for example, accepted only six taxa; Ogden (1957) divided
the group into 18 taxa; Muenscher (1944) recognized only
12 taxa; Fernald (1932) acknowledged 21 taxa.

The Pusilli have been considered difficult taxonomically
for many years (Fassett, 1940) and, in fact, many of the
recently collected specimens from North America have
been extremely difficult, if not impossible, to identify. The
reasons for these difficulties may have been: 1) the Pusilli

566 Rhodora [Vol. 76

are morphologically small and, thus, the characters are
difficult to observe; 2) vegetatively (the condition in which
much of the material is encountered) the species super-
fieially resemble each other; 3) ecological variability has
not been fully understood; 4) the nomenclature has been
confused. Many authors in the past have not recognized
the vegetative variability of the species. As a result, more
than 60 names have been proposed for the taxa of North
American Pusilli.

Because of the apparent economic importance of the
group, it is essential to understand the taxonomic relation-
ships within the genus. As stated above, this understanding
has not previously been accomplished. Therefore, I con-
sidered that the subsection Pusilli was in need of a thorough
revision. It is understood that problems still exist within
the Pusilli, but it is hoped this re-evaluation will clarify
some of the confusion within the group.

TAXONOMIC HISTORY

Potamogeton was first described by Tournefort (1719)
who recognized 13 species. Linnaeus (1753), basing his
treatment upon that of Tournefort, accepted 12 species.
Since that time, a proliferation of names has occurred, both
at the generic and infra-generic levels, resulting in con-
siderable chaos of the nomenclature. However, because of
the nomenclatural confusion and the fact that I am treating
only the Pusilli, a brief outline of the major events is pre-
sented rather than a detailed discussion of the history.

1788. Walter applied Potamogeton to two species of
Myriophyllum (Haloragaceae).

1790. Loureiro named Hydrogeton, based on P. hetero-
phyllum.

1819. Peltopsis was created by Rafinesque, based on
Potamogeton perfoliatus.

1837. Koch divided the genus into five sections: Chloe-
phylli, Coleophylli, Enantiophylli, Heterophylla,
Homophylla.

1974] Potamogeton — Haynes 567

1845. Reichenbach recognized five major groups, with-
out giving these groups formal rank: Antiphyl-
logeton, Chloegeton, Coleogeton, Heterophylloge-
ton, Homophyllogeton.

1845. Spirillus was named by Gay, without any species
having been listed.

1858. Irmish divided the Homophylla of Koch into two
sections, naming the new one Batrachoseris.

1896. Raunkiaer placed all Danish species into two sub-
genera: Coleogeton and Hupotamogeton.

1903. Raunkiaer divided the genus into 16 groups, which
were not given any formal taxonomic rank.

1907. Graebner (in Ascherson & Graebner) gave formal
rank to most of Raunkiaer’s groups (mostly at the
subsectional level) and named several other sub-
sections.

1913. Nieuwland transferred 20 species of Potamogeton
to Spirillus.

1916. Hagstrom named several new sections and numer-
ous subsections.

Potamogeton subsection Pusilli was first recognized as a
natural unit by Raunkiaer (1903). He gave no formal
name to the group, but referred to it as “P. pusillus-group.”
Graebner in Ascherson and Graebner (1907) combined two
of Raunkiaer’s groups — P. pusillus-group and P. confer-
voides-group — into the subsection Pusilli. Based upon
characters of the stem anatomy, Hagstrom (1916) removed
P. confervoides from the subsection. Hagstrom's concept
of the group was the one followed by Fernald (1932; 1950)
and accepted in the present work.

MORPHOLOGY

HABIT: The Pusilli are rooted linear-leaved obligate
aquatic plants that grow totally submerged except for the
spikes, which are usually extended above the surface of the
water. When fruiting, the spikes are normally withdrawn
below the surface so that the plant is again totally sub-

568 Rhodora [Vol. 76

merged. The mature plants, at least in the temperate re-
gions, are mostly functionally annuals, which overwinter
as seeds, winter buds, or turions. The winter buds are
produced in such abundance that Fernald (1932) and
Hagstrom (1916) were of the opinion that the plants
rarely reproduced by seeds, but instead reproduced by
winter buds. These conclusions were based upon a high
percentage of plants having been collected with the winter
buds still attached. Muenscher (1938), however, has shown
that for Potamogeton pusillus [var. tenuissimus], at least,
and possibly for other Pusilli, the young seedlings are very
similar to a winter bud and these could easily be mistaken
for old winter buds.

STEM: The stems of the Pusilli are erect to decumbent
or prostrate, depending upon the water depth and rate of
water flow, and are round to compressed in cross section.
Ogden (1972) has suggested that no species of the Pusilli
from New York possess a rhizome. This statement implies
that the stem arises from a winter bud or a similar struc-
ture formed from the seedling as illustrated by Muenscher
(1936). This situation probably represents the majority
of the cases. However, some taxa, such as Potamogeton
hillii, may produce a rhizome if the stem becomes decum-
bent and subsequently becomes covered by debris. Rooting
at the nodes may then occur, and a rhizome-like structure
several centimeters in length would be formed. Ogden
(1966), in fact, illustrated a rhizome of P. foliosus from
Texas.

Slight variation in the shape of the stem in cross-section
and in the number and sizes of ridges along the stem has
been observed. Certain species, e.g. Potamogeton foliosus
and P. friesii, may have compressed stems, whereas other
species, e.g. P. pusillus and P. clystocarpus, usually have
terete stems. However, at certain times the stem of all
species may be slightly compressed. Therefore, stem char-
acters used for identification are helpful only as supple-
mental evidence.

1974] Potamogeton — Haynes 569

Raunkiaer (1903), Hagstrom (1916), and Ogden (1943)
have utilized stem anatomy for distinguishing certain spe-
cies of other sections of Potamogeton. However, the stem
anatomy of the various species of the Pusilli is so similar
that this character is of little or no use to separate the
species.

At the nodes of some species, a pair of translucent oil
glands may be present; in other species, they are extremely
uncommon. The glands range in color from white, cream,
gold, brown, or green, and in diameter from 0.1-1.0 mm.
When present, by variations in color and size, the glands
can be useful supplemental evidence. As with the lacunae,
their presence (or absence) should not be relied upon abso-
lutely, as they are apparently a physiological response to
unknown ecological factors.

LEAF BLADES: The blades of the Pusilli are rufes-
cent, green, pellucid, to nearly black, setaceous to linear,
non-lunate, sessile, stipulate, and acute to obtuse or apicu-
late. Venation is parallel with 1-9 veins (nerves). The
lateral nerves join the midrib at or just below the apex.
In some species, one to several rows of lacunae parallel the
midrib. To my knowledge, no one has studied the develop-
ment or structure of the lacunae of Potamogeton. Singh
(1964) studied the vegetative structures of various species
of Potamogetonaceae, including P. berchtoldii [P. pusillus
var. tenuissimus], which is characterized by having lacunae
bordering the midrib. If Singh noticed any lacunae, he
made no mention of them. Fernald (1932) suggested they
are rows of nearly empty and colorless cells, presumably
giving buoyance to the leaves. To my knowledge, the origin
and functions of the lacunae are not known. However,
regardless of their origin, the lacunae appear as light lines
on each side of the midrib. The individual rows can be
observed with 10X magnification, which would indicate
that each row is several cells in width, rather than one
cell wide. The lacunae are useful for identification, as some
taxa rarely possess them, whereas others commonly have 1-.

570 Rhodora [Vol. 76

several rows on each side of the midrib. This character usu-
ally serves as good supplemental evidence when used in con-
junction with other features, preferably reproductive ones.

Recognition of many taxa of Potamogeton, especially of
the Pusilli, has been based upon slight morphological dif-
ferences in the leaves (Hagstrom, 1916; Fernald, 1932).
Dandy and Taylor (1940) suggested that, due to different
ecological conditions, the vegetative variability of the spe-
cies is too great to be useful as the only criterion for
naming a new taxon. During the present study I have ob-
served that several populations undergo a seasonal di-
morphism, e.g., producing two morphologically dissimilar
sets of leaves in one growing season. Often, plants of
Potamogeton pusillus var. tenuissimus produce in the early
summer a set of obtuse leaves that are near the limits of
width for the species and which possess four to five rows
of lacunae on each side of the midrib. Later in the growing
season, these leaves die and a new set is formed. The leaves
of this second set are usually acute, quite narrow, and pos-
sess only one to three rows of lacunae. If one were to
follow Fernald's (1932) taxonomy of this group, one could
correctly identify the plant as one variety in the early sum-
mer and another variety in the fall. The cause of this sea-
sonal variation is unknown. Since the same genotype may
produce two sets of leaves, each morphologically distinct,
these vegetative characters should be considered with cau-
tion when attempting to distinguish taxa.

STIPULE: Associated with each leaf blade and sur-
rounding the stem of the Pusilli is a tubular or convolute
stipule. The stipules are free from the leaf blades and vary
in color from white to brown to green. The number and
coarseness of the veins varies from one species group to
another. This vein character, along with the stipule color,
can be useful for identification of species. In some taxa,
e.g. Potamogeton friesii, the stipules are white with many
coarse veins, With age the interveinal tissue near the apex
decays leaving the coarse veins extending as fibers past the

1974] Potamogeton — Haynes 571

remaining interveinal tissue. In other species, e.g. P. pusil-
lus, the stipules are usually brown to green with only a few
veins, these being quite delicate. With age, the vein tissue
decays along with the interveinal tissue.

Hagstrom (1916) considered the tubular versus convolute
condition of the stipules to be of great evolutionary signifi-
cance. He, in fact, divided the Pusilli into two series —
Pusilli Convoluti and Pusilli Connati — based on no char-
acters other than the connation of the stipules. Later
workers, Dandy and Taylor (1938), Ogden (1966), and
Voss (1972) have accepted this character as important for
determining species but have not accepted its use in sepa-
rating the subsection into two series. For one of the species
complexes in which the stipule character is considered most
useful, according to Hagstrom and Fernald, — that being
P. pusillus and P. panormitanus [P. pusillus var. tenuissi-
mus and var. pusillus, respectively], I have compared the
character with other vegetative and reproductive characters
considered important by these workers. The data obtained
from this comparison indicate only about a 70-80% corre-
lation of this character with the other characters. In my
opinion, when the situation is either the presence or ab-
sence of a character, for that character to be given such
high value in a classification system, there should be little
variability of that character within a taxon.

WINTER BUDS: The winter buds (turions) are veg-
etative reproductive structures that function as agents for
multiplication of the individuals and as perennating organs
during the unfavorable season. Yeo (1966) planted one
winter bud of Potamogeton crispus on 1 Apr. 1963. At the
end of the growing season, 23,520 winter buds had been
produced. This is obviously a very effective method of
propagule production. In some species, the winter buds
are produced early in the growing season and may, there-
fore, germinate to produce new plants that year. In other
species they are produced only at the end of the season,
and do not germinate until the following year.

572 Rhodora [Vol. 76

The winter buds consist of a short stem apex with short-
ened internodes. The leaves of the structures can be di-
vided into two types: outer and inner. The outer leaves
usually resemble the vegetative leaves, but the inner leaves
may not. The inner leaves may be numerous, shortened,
and oriented in a plane perpendicular to that of the outer
leaves, thus giving a fan-shaped appearance, as in P. friesii.
On the other hand, the winter buds may be few to several,
compacted, and rolled into a hardened structure, thus ap-
pearing fusiform, as in P. pusillus. Finally, they may be
various in number and similar to the vegetative leaves, as
in P. obtusifolius.

As indicated above the winter buds may be useful as
characters for the identification of the Pusilli. Hagstrom
(1916) and Fernald (1932) have placed emphasis upon the
location on the plant of the structures. I consider the loca-
tion of the structure too variable to be of taxonomic value.
Instead, the size of the structure and the modification of
the inner leaves are of significant taxonomic value.

INFLORESCENCE: The inflorescence of the Pusilli is
either a capitate or cylindric spike consisting of from one
to five whorls of four flowers each. Some authors have
considered this structure to consist of a reduced compound
spike (see comments under discussion of the flower). The
inflorescence is a very good character for distinguishing
species of the Pusilli. Important features are the length,
shape, and number and separation of the whorls of the
inflorescence. In some taxa, e.g. Potamogeton pusillus var.
pusillus, with an inflorescence of more than three whorls,
the whorls will usually be separated; however, in other
taxa, e.g. P. foliosus, the whorls are rarely separated.

PEDUNCLES: The peduncles of the Pusilli are either
terminal on the stem or axillary to vegetative leaves. Often,
axillary peduncles are reflexed, whereas the terminal ones
usually are erect. In longitudinal section, the peduncles
either have parallel sides or are clavate. In cross sections,

1974] Potamogeton — Haynes 573

they range from nearly terete to so compressed that they
are three to four times as broad as high.

The peduncle is often a very good character for dis-
tinguishing taxa of the narrow-leaved pondweeds. In
Potamogeton foliosus for example, the peduncle is short,
clavate, usually reflexed, and axillary. In P. pusillus var.
pusillus, on the other hand, the peduncle is elongate, paral-
lel-sided, erect, and usually terminal.

FLOWER: The flowering structures of Potamogeton
have been interpreted variously. First, Eames (1961) in-
terpreted each perianth segment as a sepaloid bract sub-
tending and adnate to the corresponding stamen. He,
therefore, regarded the flower of Potamogeton as an in-
florescence comprising four staminate flowers, each of a
single stamen and its adnate bract, and four apetalous
carpellate flowers, each of a single carpel with one cam-
pylotropus ovule. By this interpretation, the spike repre-
sents a reduced compound inflorescence. Second, based upon
vascular anatomy and floral morphology of seven species of
Potamogeton, Singh (1965) chose to consider the flower
of Potamogeton as a normal flower with four perianth seg-
ments. Finally, Sattler (1965) considered it impossible to
classify the structure as a true flower or as an inflorescence,
because it displayed characters of both. He questioned the
classical use of flower and inflorescence as two separate
categories and emphasized the need for more developmental
work in the “Helobiales” (an order that is now considered
to be the Alismatidae). For the sake of convenience,
Singh's concepts are followed in this revision.

Controversy has also existed as to the real nature of the
perianth-like structures that are adnate to the stamens.
Ascherson (1889) and Rendle (1930) considered the struc-
tures to be outgrowths of the stamens. They then called
the structures stamen connectives. Sattler (1965) and
Singh (1965) have reached a different conclusion. Sattler,
for example, demonstrated in Potamogeton richardsonii
that these structures are initiated on the floral apex prior

574 Rhodora [Vol. 76

to stamen initiation. Thus, they cannot be outgrowths of
the stamens. He instead considered them to be perianth
segments. After inception of the stamen primordia, growth
occurs between the perianth primordia and that of the
stamen. Thereby, the base of the developing stamen and
perianth become united. The perianth is usually light green
to greenish brown in color and consists of two parts — a
claw and an expanded upper portion — each of approxi-
mately equal length.

The gynoecium consists of four separate, superior, uni-
ovulate carpels with parietal placentation. The androecium
consists of four bithecate stamens. Anthesis usually is
approximately at the time of embryo sac maturity (Law-
rence, 1951).

Among the various taxa of the Pusilli, only slight mor-
phological variation exists in the flowers. These structures,
therefore, are of little value for distinguishing species.
However, the size of the structures often is quite useful
for separating various subsections of the genus.

FRUITS: The fruits of the pondweeds have been vari-
ously classified — e.g., as a drupe (Reichenbach, 1845), as
a drupelet or drupaceous (Morong, 1893; Fryer and Ben-
nett, 1915), as a drupe or an achene (Clapham et al., 1962)
and as a druplet or nutlet (Lawrence, 1951). The use of
such an array of terms suggests that no one term could
adequately describe the fruits. Although the fruits resemble
an achene in size and by having a thin, often leathery meso-
carp, they do not conform with the definition of an achene
in that the seed coat is not adnate to the pericarp, and
under natural conditions the fruits are rarely ever dry.
Muenscher (1936), in fact, has shown that when the fruits
are subjected to drying, a very high rate of mortality oc-
curs, The fact that the fruits rarely dry also rules out the
possibility of their being a nut or nutlet. According to Aalto
(1970) the fruits are histologically drupes since the peri-
carp is differentiated into an exocarp, mesocarp, and stony
endocarp. The fruit is, however, unlike a drupe because

1974] Potamogeton — Haynes 575

the dorsal area of the pericarp wall appears to open and
then be covered by a lid (Aalto, 1970). At the junction of
the lateral walls and the lid, a ridge is formed. This ridge,
in many species, is pronounced into two lateral wings (one
on each side of the fruit). The lid can be removed exposing
the seed coat. Because of the histological similarity, Aalto
chose to call the fruits of the Fennoscandian species drupes.
Until a better term is proposed, I am considering the fruits
drupe-like.

The fruits are the single most important character for
distinguishing the taxa of the Pusilli. Several fruit char-
acters are useful for identification: 1) color — usually
brown or green; 2) size; 3) shape — whether widest above
the middle, at the middle, or below the middle; 4) presence
of dorsal or lateral wings or ridges — one species, P. folio-
sus, has a dorsal wing, three others two lateral and one
dorsal ridge, others have no ridges; 5) amount of depres-
sion on the sides; 6) length and position of the style (or
beak).

Aalto (1970) has demonstrated that for some species of
Potamogeton the endocarp can be useful for identification.
The majority of his work was done with subfossil endo-
carps. He also exposed recent endocarps by boiling the
fruits in 5% potassium hydroxide solution for 10 to 15
minutes. The exocarp and mesocarp disintegrate leaving
the endocarp exposed. These were examined for shape and
size of the cells and for shape, size, and appendages of the
endocarp.

VEGETATIVE DIMORPHISM

Vegetative morphology has been used as a basis for sep-
aration of certain taxa of Potamogeton. In Fernald’s study
(1932), P. pusillus was divided into six varieties based on
shape of the leaf apex and number of lacunae bordering the
leaf midrib. The species was divided into two informal
groups using the shape of the leaf apex. These groups
then were further divided into the varieties with the num-

576 Rhodora [Vol. 76

ber of lacunae being the only character used for separa-
tion. During the summer of 1878, E. S. Miller made two
collections of P. pusillus from presumably the same locality
of the Wading River, Suffolk County, New York: 1) sterile
plants on 25 May 1878, and 2) fertile plants on 12 Jul. 1878.
A sterile and fertile plant together were mounted on a
herbarium sheet and these sheets were distributed. I have
seen six such sheets and on every one the sterile plant had
leaves with obtuse apices and two to three rows of lacunae
each side of the midribs. Leaves of the fertile specimens,
on the other hand, were acute and lacked lacunae. Fernald
examined some of the specimens and consistently identified
the sterile plants as P. pusillus var. mucronatus and the
fertile plants as P. pusillus var. tenuissimus. This situation
led me to consider the hypothesis that the taxonomic char-
acter of the leaf apex and lacunae were environmentally
controlled. I, therefore, began visiting populations several
times throughout one year and, often, in subsequent years
to see if any morphological changes occurred. Data gath-
ered from these field studies indicate that P. pusillus and,
also, P. friesii may undergo a dimorphism in which two
morphologically dissimilar phenotypes may be observed in
the same population.

During the summer of 1969, I made a collection of Pota-
mogeton pusillus (Haynes 3251) from Pelee Island, Ontario.
These specimens had obtuse leaves with five to six rows of
lacunae on each side of the midrib (Fig. 1). The popula-
tion was visited separately by Dr. Ronald L. Stuckey and
Mr. Marvin L. Roberts, both of The Ohio State University,
several times during the next two years. In late August
1971, the population was found in fruit (Roberts 1562).
The leaves from this collection were acute and had at most
two rows of lacunae each side of the midrib (Fig. 1).
Therefore, plants of this one population could be correctly
identified, according to Fernald (1932), as var. mucronatus
when sterile and var. tenuissimus when fertile. I also have
observed this phenomenon in a population from New York
(Haynes 3326). Here both types of leaves were found on

1974] Potamogeton — Haynes 577

the same plant. The obtuse, lacunate leaves were black and
appeared dead at the time of collection and the acute, non-
lacunate leaves were green and alive at the time of collec-
tion. During the course of herbarium work, I have exam-
ined many specimens in which both types of leaves were
present on one plant, usually with the obtuse, lacunate ones
appearing to have been dead at the time of collection.
These data appear to indicate that plants with obtuse lacu-
nate leaves are always sterile. This is not the case. Some
are fertile; however, as compared with the number of fer-
tile plants with non-lacunate, acute leaves, the former are
few in number.

A similar situation can be observed in Potamogeton
fries. However, rather than two distinct sets of leaves
being produced by one plant, apparently only one set of
leaves is produced a year, but these leaves undergo a mor-
phological change during the growing season. According
to Voss (1972), P. friesii tends to have leaves without
revolute margins, whereas P. strictifolius normally has
ones with revolute margins. Data gathered during my
field work indicate that in the early summer P. friesii does
have leaves without revolute margins (Fig. 1). A popula-
tion in Emmet County, Michigan, visited by me on 15 Jun.
1971 (Haynes 3694) had such leaves. However, on 30 Jul.
1971, I again visited the same population and found most
plants with leaves having revolute margins (Haynes 3747).
Thus, one population may produce both phenotypic expres-
sions in one year.

The two examples of vegetative dimorphism outlined
above illustrate the vegetative plasticity of Potamogeton
explained by Dandy and Taylor (1940). These data indicate
that within the Pusillii and probably other groups of
aquatic plants, the effects of differing environmental con-
ditions upon the vegetative characters should be under-
stood before one uses these characters in a classification.

578 Rhodora [Vol. 76

A B
e D
E

K^ a»

F

Fig. 1l. Potamogeton pusillus leaf (A-D). A. Midsection, early
summer. B. Apex, early summer. C. Midsection, late summer. D.
Apex, late summer. Cross-section of Potamogeton friesii leaf (E-F).
E. Early summer. F. Late summer.

1974] Potamogeton — Haynes 579

REPRODUCTIVE ECOLOGY

REPRODUCTION: The reproductive ecology of Pota-
mogeton is poorly known. According to Sculthorpe (1967)
certain species of Potamogeton reproduce by rhizome frag-
ments, tubers, and winter buds or turions. Also, Moore
(1915) studied seed germination of Potamogeton and con-
cluded that once a seed germinates and develops into a
mature plant, several periods of vegetative reproduction
must occur before the plant will produce flowers and fruits.
In the Pusilli, Hagstrom (1916) and Fernald (1932) were
of the opinion that the plants rarely reproduced by seeds,
but instead, reproduced mostly by winter buds. For the
species in which winter buds are rare or unknown, this
idea of reproduction by winter buds may not be valid.
However, for the species which prolifically produce winter
buds, reproduction by these structures is probably the most
common type, since fruiting plants often are found with
winter buds attached to the underground parts.

POLLINATION: According to Sculthorpe (1967) and
to Cronquist (1968), the pollination trend in the Alismati-
dae is from insect to wind, and ultimately, to water pollina-
tion. Members of the genus Potamogeton including the
Pusilli are, for the most part, wind pollinated (Sculthorpe,
1967). However, according to Voss (1972) and data based
on personal field experience, many inflorescences of the
Pusilli never become emergent. In such instances, if pol-
lination does occur, the medium for the transport of the
pollen must be the water. However, the mature, viable
fruits might be formed as a result of apomixis.

GERMINATION: As shown by Muenscher (1936), the
germination rate of Potamogeton fruits is greatly reduced
when the fruits are subjected to drying. Many species,
according to Muenscher, require near-freezing tempera-
tures for a period of one to three months. However, as
shown by Sullivan (1967) for many of the broad-leaved
species of Potamogeton, the fruits can be induced to germi-

580 Rhodora [Vol. 76

nate without a previous cold treatment if the exocarp and
mesocarp are split, thus exposing the endocarp.

As stated earlier, the fruits of the pondweeds are among
the most common foods of waterfowl. Often only the exo-
carp and mesocarp are digested, leaving the endocarp with
the intact seed to be passed from the digestive system.
Lohammar (1954) has shown that a high percentage of
germination may occur after the fruits have passed through
the digestive system of the waterfowl.

CYTOLOGY

Several workers, e.g. Palmgren (1939), Harada (1942;
1956), Lóve (1954), Lóve and Lóve (1961), and Stern
(1961), have studied Potamogeton cytologically. However,
according to Bolkhovskikh et al. (1969), these studies have
produced chromosome counts for only 50 of the approxi-
mately 100 species of the genus. Stern (personal commu-
nication) indicates that the pondweeds are difficult to
collect for meiotic material since the plants undergo meiosis
quite rapidly in very early bud stage. The above-mentioned
factor added to the fact that the aquatic environment is a
difficult one for field work probably are the major reasons
that few cytological studies have been undertaken on the
pondweeds. During the summers of 1970, 1971, and 1972,
I collected flower buds with the intention of obtaining
chromosome counts from meiotic material. The buds were
preserved in the usual manner with a 3:1 solution of ethyl
alcohol and acetic acid. In some instances, the buds were
examined within two or three days of collection; in others,
the buds were examined after several months. In all in-
stances I failed to have success in obtaining countable
meiotic configurations. Therefore, all counts included in
this paper are based upon previous reports in the literature.

According to Stern (1961), the base number for Pota-
mogeton is x — 13 or 14. The majority of the chromosome
numbers reported for the floating-leaved species are multi-
ples of these figures; e.g. P. amplifolius Tuckerm., n — 26;
P. gramineus L., n= 26; P. illinoensis Morong, n = 52

1974] Potamogeton — Haynes 581

(Stern, 1961). On the other hand, most reports for the
submersed-species indicate an n number of 13 or 14. A
summary of the reported chromosome numbers for the
Pusilli is included in Table 1.

These reports of chromosome numbers in Potamogeton
should be considered with caution, as the nomenclature has
been considerably confused and most workers, excluding
Stern (1961) and Taylor and Mulligan (1968), did not cite
voucher specimens from which the counts were taken.

TABLE 1

Published chromosome numbers for Potamogeton
subsection Pusilli.

Taxon Chromosome number

P. berchtoldii Fieber n = 13 (Taylor & Mulligan, 1968)
[P. pusillus
var. tenuissimus] 2n — 26 (Taylor & Mulligan, 1968)

P. foliosus Raf. 2n — 28 (Stern, 1961)
P. groenlandicus

Hagstrom 2n = 26 (Jørgensen, et al., 1958)
P. mucronatus Schr. 2n — 26 (Palmgren, 1939)

[P. friesii]
P. obtusifolius M. & K. 2n = 26 (Palmgren, 1939)
P. panormitanus Biv. 2n — 26 (Palmgren, 1939)
[P. pusillus
var. pusillus]
P. strictifolius
Ar. Benn. 2n — 52 (Lóve, 1954)

PHYLETIC RELATIONSHIPS

GENERIC RELATIONSHIPS: According to Cronquist
(1968), Potamogeton is classified in the Class Liliatae
(monocots), Subclass Alismatidae, Order Najadales, and
family Potamogetonaceae. He considers the Alismatidae
to be “a near-basal sidebranch, a relictual group which
has retained a number of primitive characters." The group

582 Rhodora [Vol. 76

is, therefore, not considered to be in the main line of evolu-
tion of the monocots.

In studying the relationships of the Potamogetonaceae
with other families of the “Helobiae,’ Chrysler (1907)
came to the conclusion that Potamogeton was the most
primitive genus of the group and that Potamogeton-like
organisms gave rise to most of the other genera. He con-
sidered the floating-leaved species, especially P. pulcher
with its submersed leaves very similar to its floating leaves,
to be more primitive than the submersed-leaved species.
The submersed species were assumed to have been derived
from the floating-leaved species as “a stage in the assump-
tion of the aquatic life by the genus." One would conclude
that Chrysler considered the Helobiae (Alismatidae) to
have been derived from terrestrial species.

Cronquist (1968), on the other hand, considered the
primitive monocots to have been derived from a Nym-
phaeales-like dicot, thus aquatic in origin. He considered
the typical parallel-veined leaf of the monocot to be a
modified, bladeless petiole. From this modified, flattened
petiole, the expanded blade of, say Sagittaria, could have
been derived by the spreading of the veins farther apart
near the tip of the petiole. By applying this concept to
Potamogeton, one could infer that the floating leaves might
have been derived from the submersed leaves by an increase
in the amount of tissue between the veins near the tip of
the petiole. This would imply that possibly the floating-
leaved species were derived from some ancestral stock of
submersed-leaved plants.

Published cytological data (cf. cytological section) sup-
port this interpretation. The submerged-leaved species are,
for the most part, diploid, whereas the floating-leaved spe-
cies are, for the most part, tetraploid. Although Raven and
Thompson (1964), DeWet (1965), and Anderson (1972)
have demonstrated instances of polyhaploidy, based upon
our present interpretation of cytological data as evidence
for evolutionary relationships, the published chromosome

1974] Potamogeton — Haynes 583

numbers indicate that the primitive condition probably was
that of total vegetative submergence.

As stated earlier, pollination in the Alismatidae is of
three types — insect, wind, and water. According to Cron-
quist (1968), the reduction of the perianth in the Najadales
reflects the abandonment of insect pollination, with water
pollination, as in Najas and Zostera, being the most special-
ized type. One would suspect, therefore, the primitive
species of Potamogeton to be wind-pollinated. The water-
pollinated species, then, would have been derived from some
wind-pollinated ancestral stock. Again, this interpretation
is supported by cytological data. Chromosome number
reports for the P. filiformis subsection are the highest of
all Potamogeton. This group is the only one thought to be
totally water-pollinated. Palmgren (1939) reports numbers
of 2n — ca. 66 for P. filiformis Pers. and 2n — ca. 88 for
P. vaginatus Turcz.

Based upon the scanty cytological, pollination, and mor-
phological evidence, I here propose that the primitive spe-
cies of Potamogeton were similar to P. zosterifolius in that
they possessed narrow submersed leaves with many fine
nerves and that they were wind pollinated. From the Pota-
mogeton zosterifolius-like ancestor, several lines of differ-
entiation probably occurred. One of these, featuring a
reduction in vein number and retaining the wind pollina-
tion, eventually gave rise to the Pusilli. As the scope of
this paper is only the Pusilli, I will not postulate on the
systematic relationships of the entire genus. Instead, my
further comments on this subject will be restricted to the
Pusilli.

SUBSECTIONAL RELATIONSHIPS: Potamogeton
subsection Pusilli is treated here to consist of 15 species.
The species morphologically most similar to P. zosterifolius
is P. friesii. The latter species, as does P. strictifolius,
resembles P. zosterifolius in fruits, peduncles, inflorescence,
leaves, and stipules. Potamogeton friesii, therefore, is prob-
ably the most primitive species of the Pusilli. Potamogeton

584 Rhodora [Vol. 76

strictifolius, being a tetraploid, probably was derived from
a P. friesii-like ancestor by polyploidy. One can postulate
two main lines having developed from the ancestral Pusilli
(Fig. 2): 1) dorsal and lateral keels absent; 2) dorsal
and/or lateral keels present.

The first line is represented by Potamogeton pusillus and
P. groenlandicus. Potamogeton pusillus is widespread
throughout the world, whereas P. groenlandicus is re-
stricted to the coastal areas of Greenland. The latter spe-
cies probably represents a population of P. pusillus that
was isolated on Greenland and has differentiated from
those of P. pusillus.

The second line is represented by Potamogeton obtusi-
folius, P. hillii, P. clystocarpus, and P. foliosus. Potamoge-
ton foliosus, having a chromosome number of 2m — 28,
probably represents a population which was separated
from a P. obtusifolius-like ancestor by aneuploidy and has
become widespread. In fact, it is so widely distributed in
North America that it is difficult to draw any relationships
as to its possible place of origin. However, other species
of the line as well as P. strictifolius and P. friesii have
very distinct distributions. These ranges will be discussed
and used as supplemental evidence for the phyletic relation-
ships. Potamogeton obtusifolius, P. strictifolius, and P.
friesii, have distributions limited almost entirely to geo-
graphical areas that were covered with ice during Pleisto-
cene glaciation. These species probably survived glaciation
elsewhere and migrated into the once glaciated areas fol-
lowing the retreat of the ice. Apparently, species or vari-
eties could have evolved in a matter of 10,000 years. One
need only examine a few of the Great Lakes endemics, e.g.
Tris lacustris (Guire & Voss, 1963), Cirsium pitcheri (John-
son & Iltis, 1963), and Calamovilfa longifolia var. magna
(Thieret, 1960). However, in comparison to these sand
dunes endemics, most of the Pusilli are wide ranging cir-
cumboreal species. If these Pusilli were to have evolved
recently, then rapid circumpolar dispersal also would have
had to occur.

1974] Potamogeton — Haynes 585

Hultén (1937) proposed that there were two main ref-
ugia in North America — the Rocky Mountains and the
continental shelf outside eastern America — where plants
survived during continental glaciation. From these refugia,
he stated, the plants probably spread in an easterly and
westerly direction, respectively, toward the center of the
continent. Some taxa probably survived in only one area,
while other taxa possibly survived in both areas. Some of
the taxa which survived in both refugia migrated until
their ranges overlapped; thus, they now have a continuous
range across North America. For others, however, migra-
tion ceased before the ranges overlapped. Hultén conceded
that species surviving in other areas south of the glacial
boundary did migrate north to some extent into the once
glaciated lands. However, he suggested that a much smaller
proportion of the species now found in the glaciated areas
survived in the vast areas south of maximum glaciation
than in the two other refugia.

Iltis (1965) and Stuckey (1972) have suggested that cer-
tain species of Gentianopsis and Rorippa, respectively, sur-
vived glaciation in the Rockies and migrated eastward
following the retreat of the Wisconsin Ice sheet. The dis-
tributions of the taxa discussed by these two workers are
similar to that of Potamogeton obtusifolius, P. strictifolius,
and P. friesii in that all have a more or less continuous
range across northern North America and are mostly known
from south of the glacial boundary only in the Rockies.
Marie-Victorin (1938) proposed an explanation of the east-
ward migration of such species by suggesting that ‘the
unforested belt that must have existed along a receding
icefront [served] as a kind of sidewalk extending from the
Rockies to the Gulf of St. Lawrence." He cited no pond-
weeds which demonstrated such distributions, but Stuckey
(1972) indicated that P. friesii and P. strictifolius have
ranges similar to a distribution pattern that Marie-Victorin
probably had in mind when proposing this idea.

At the time of Hultén's article (1937) Potamogeton ob-
tusifolius from western America was represented by few

586 Rhodora [Vol. 76

collections, all from localities in glaciated areas. This
exiguous number of specimens from western America led
Hultén to regard the taxon as a cireumpolar species which
survived in the eastern refugium. However, recent collec-
tions indicate that the taxon is widespread, although uncom-
mon, throughout this geographical area, including the Rocky
Mountains. Therefore, I include P. obtusifolius as a taxon
which survived continental glaciation in the western ref-
ugium and then migrated eastward.

Potamogeton hillii, on the other hand, survived conti-
nental glaciation in eastern America just south of the glacial
boundary, in the area of Pennsylvania, and migrated north
and west with the retreat of the ice. In fact, the morpho-
logical similarity of the species with P. obtusifolius would
indicate that prior to glaciation, the two currently recog-
nized taxa were perhaps represented by one large hetero-
geneous population. Glaciation would have separated this
large population into two smaller allopatric ones, one to
the east and one to the west. During glaciation, the two
populations differentiated, resulting in each population
evolving into a different species. Once the period of glaci-
ation was complete and the distributions overlapped, the
reproductive barriers that had been established during
glaciation, evidently now prevents gene flow from one popu-
lation to the other. Iltis (1965) indicated that Gentianopsis
procera and G. crinita possibly had a similar history. Sim-
ilarly, P. clystocarpus is known from only one locality in
Jeff Davis County, Texas. This species probably represents
a population of P. obtusifolius in the southern mountains
which was isolated and has differentiated from the parental
population.

TAXONOMY

Of the more than 60 names that have been proposed for
the North American Potamogeton subsection Pusilli, most
of these are based on differences in venation, texture, color,
lacunae number, and apex shape of the leaves. Dandy
(1937), Dandy and Taylor (1940), and Clapham et al.

1974] Potamogeton — Haynes 587

groenlandicus

var. pusillus

P. pugtllus ar. tenuissimus

ar. gemmiparus

P. friesii-like ancestor friesii

tricttfolius

Lystocarpus

obtueifolius

hillii

var. foliosus

P. foltosu

ar. fibrillosus

Fig. 2. Hypothetical phylogenetic relationships of North Ameri-
can Pusilli.

(1962) have shown that the leaf form and anatomy of
Potamogeton may vary widely with age, water depth, cur-
rent speed, nutrient supply, light intensity, and perhaps
other factors. Data from classical and experimental studies
of other genera of aquatic vascular plants indicate that this

588 Rhodora [Vol. 76

heterophyllous condition is common among the submerged
species. Fassett (1951) illustrated no fewer than ten quite
distinct individuals of Callitriche heterophylla and drew
attention to the close morphological similarities of ecological
forms which actually belong to different species. Bostrack
and Millington (1962) found in Ranunculus flabellaris that
the shape of the leaf may be affected by temperatures, pho-
toperiod, light intensity, and submergence acting directly
upon the distribution and frequency of cell division in the
primordium so that the inherent zones of growth initiated
on the primordium are modified in their development.
Cook (1966), in fact, after recognizing 20 taxa of Ranun-
culus subgenus Batrachium listed 302 names which applied
to these taxa. Williams (1970) found that by transplanting
individuals of Nymphaea tuberosa to localities where N.
odorata occurred, the individual would lose its “tuberosa”
characters and obtain those of N. odorata. For reasons just
outlined, these workers have found it necessary to compile a
classification based primarily on reproductive characters.
My approach to the systematies of Potamogeton subsec-
tion Pusilli has been primarily that of combined herbarium
and field studies. The data presented are based on three
summers field work in northern United States and southern
Canada and an examination of over 8000 herbarium speci-
mens from the following 22 herbaria (Acronyms from Lan-
jouw and Stafleu, 1964) : C, CAN, CGE, DAO, F, GH, ILL, K,
LAF, MICH, MO,MU, NY, NYS, OS, PH, S, UC, UMBS, UPS, US, W.
Evidence from the field work has indicated that the
Pusilli are as morphologically plastic as Callitriche and
Ranunculus. Just as workers in these other genera have
resorted to fruiting characters as a basis of a classification,
I think that a classification based for the most part upon
the reproductive structures is the only one which would be
workable. Therefore, one should always attempt to collect
flowering or fruiting specimens (preferably fruiting) ! Just
as identification of a sterile Aster or Solidago is nearly
impossible, it is difficult at best to identify a sterile pond-
weed (and then one can never be sure of his determination).

1974] Potamogeton — Haynes 589

I have thus refrained from mapping any specimen in which
the determination was questionable.

For one to completely adhere to the biological species
concept, he should demonstrate, either experimentally or
naturally, if hybridization between two taxa does occur.
However, in genera such as Potamogeton, in which the
flowers are so small as to render difficult the artificial cross-
ing of individuals, one hardly can demonstrate whether two
individuals are capable of crossing and producing fertile
offspring. Ogden (1943) has suggested that the broad-
leaved species of Potamogeton hybridize quite readily. He,
however, did not demonstrate this phenomenon experi-
mentally; rather he depended wholly upon the hybrid in-
dividuals possessing characters intermediate between those
of the two putative parents. With the Pusilli, however, it is
not so easy. Vegetatively, the condition one would expect
to find a hybrid, the species of the Pusilli are quite similar;
thus, intermediates would be difficult to detect. Therefore,
the techniques used by Ogden are not applicable to the
Pusilli.

Because of the difficulties outlined above with hybridiza-
tion, I have chosen to stress a morphological species con-
cept. I have followed the premise that species should be
separated by discontinuities between the morphological
ranges of the fruits, peduncles, inflorescence, and, to a lesser
extent, the winter buds. Therefore, for the 60 names which
have been applied to the North American Pusilli, I am
accepting only eight species, with one of these, Potamogeton
foliosus, being represented by two varieties and another,
P. pusillus, being represented by three varieties. In an
attempt for consistency, I have regarded varieties as mor-
pho-geographic subdivisions of a species (Kapadia, 1964)
that presumably reflect genetic differences. As will be
noticed, I do not recognize subspecies or forms. It is my
contention (as that of Raven, 1969) that only one infra-
specific unit should be recognized. I have chosen to use
variety over subspecies because of tradition in eastern
North America.

590 Rhodora [Vol. 76

In the treatment that follows, keys are based upon fruit-
ing material. Dimensions of leaves are taken from the fully
expanded, longest leaves of a specimen. Measurements of
the width were taken approximately at the widest point of
the leaf. Dimensions of the spike and peduncles are based
upon both flowering and fruiting specimens. Descriptions
of the fruits are taken strictly from mature structures.

POTAMOGETON Linnaeus, Species Plantarum 126. 1753

Hydrogeton Loureiro, Flora Cochinchinensis 244. 1790.
Type Species: Hydrogeton heterophyllum — Potamogeton
octandrus Poir.

Patamogeton Honckeny, Syn. Plan. Germ. 2: 110. 1793.
(orthographic variant).

Potamogiton Rafinesque, Med. Repos. 5: 354. 1808. (or-
thographic variant).

Potamogetum Clairville, Man. d'Herb. Suisse & Valais
34. 1811. (orthographic variant).

Peltopsis Rafinesque, J. Phys. Chim. Hist. Nat. Arts 89:
102. 1819. Type Species: Potamogeton perfoliatus. L.

Spirillus J. Gay, Compt. Rend. Hebd. Seances Acad. Sci.
38: 703. 1854. (name without any listed species) ; emend.
Nieuwland, Amer. Midl. Naturalist 3: 14. 1913. Type
Species: (lectotype here designated) Potamogeton diversi-
folius Raf.

Plants herbaceous, aquatie, submerged in fresh or rarely
brackish water, annual or perennial, propagated from seeds,
winter buds, or rhizomes. Stems variable in length accord-
ing to water depth, branched or unbranched, terete or com-
pressed, rooting at the nodes. Leaves all submersed or both
submersed and floating, alternate or subopposite; sub-
mersed leaves pellucid, sessile or petiolate, linear to orbicu-
lar, subulate to obtuse at the apex, the margins entire to
serrate, rarely crimped, the nerves 1-35; floating leaves
coriaceous, mostly petiolate, rarely subsessile, elliptic to

1974] Potamogeton — Haynes 591

ovate, acute to obtuse at apex, cuneate to rounded or cor-
date at base, the margins entire, the nerves 3-51. Stipules
tubular, sheathing the stem and young inflorescences, con-
nate or convolute, either free or adnate to the base of sub-
mersed leaves, free from base of floating leaves. Winter
buds present or absent, with extremely shortened nodes,
divided into inner and outer leaves; inner leaves few to
numerous, either shortened and oriented at 90? angles with
respect to outer leaves, rolled into a fusiform structure, or
unmodified; outer leaves 1-5 per side, mostly similar to
vegetative leaves, rarely corrugated near base. Inflorescence
a capitate or cylindrie spike with 1 to 20 whorls of flowers,
compact or moniliform, with 2 to 4 flowers each whorl,
mostly buoyed above surface of water. Flowers bisporangi-
ate, Perianth of 4, free, rounded, short-clawed, greenish
segments. Androecium of 4 stamens: filaments adnate
to the perianth claw; anthers bithecate, extrorse. Pollen
spherical monoaperturate. Gynoecium of 4, free, unilocular,
uniovulate carpels ; ovule campylotropus; placentation pari-
etal. Fruit drupe-like; dorsally rounded or keeled ; embryo
coiled; cotyledon one, endosperm absent. Chromosome base
number: z — 13 or 14 (Stern, 1961). (Name from the
Greek potamos, a river and geiton, a neighbor.) Type Spe-
cies: Potamogeton natans L. (fide Taylor, 1909).

POTAMOGETON Subsection PUSILLI Graebn. in Aschers.
& Graebn. Das Pflanzenreich 4(11): 260. 1907.

Plants submersed in fresh water, annual or perennial.
Stems branched or unbranched, terete or compressed.
Leaves all submersed, pellucid, sessile, linear, subulate to
obtuse at apex, entire, 1-9-nerved. Stipules connate or con-
volute, free from the base of the leaves. Winter buds pres-
ent or absent. Inflorescence mostly emersed, a capitate or
cylindric spike with 1-5 whorls of flowers, compact or
moniliform, mostly with 4 flowers at each whorl. Fruit
dorsally rounded or keeled, to 4.0 mm long. Chromosome
base number x = 13 or 14. Type Species: Potamogeton
pusillus L.

592 Rhodora [Vol. 76

KEY TO THE TAXA OF NORTH AMERICA
1. Leaves 7-9-nerved and inner-leaves of winter buds un-
modified; plants of Greenland. .. 8. P. groenlandicus.

1. Leaves with up to 9 nerves, but if 7-9-nerved, then with
inner-leaves of winter buds modified; plants from areas

other than Greenland. ............. 0.000002 0 ee 2.
2. Fruits with a dorsal keel or ridge to 0.4 mm high,
often with 2 lateral keels. .................... 3.

3. Inflorescence cylindric, 8 mm or more long; basal
glands usually present, mostly 0.5 mm or larger
in diam; leaves obtuse or apiculate, rufescent.
Lecce eee ee ee ee ee eee eee 3. P. obtusifolius.

3. Inflorescence capitate or rarely cylindric, 7.5 mm
or shorter; basal glands when present, mostly
smaller than 0.5 mm diam; leaves acute, rarely
obtuse, olive or green. .............0000005 4.
4. Peduncle cylindrie, mostly terminal, erect;

basal glands present; fruit with basal tuber-
cles, ....... 0.00. less 6. P. clystocarpus.

4. Peduncles usually clavate, axillary, often re-
curved; basal glands uncommon; fruit with-
out basal tubercles. ................00-. 5.
5. Fruits rounded on sides, 3-keeled, 3-4 mm

long; dorsal keel ridge-like, to 0.2 mm
high. ..............2000050. 4. P. hillii.

5. Fruits with concave sides, 1-keeled, to 2.7
mm long; dorsal keel wing-like and undu-
late, to 0.4 mm high. ................ 6.
6. Spike rarely interrupted; fruit olive to

green-brown, 1.5-2.7 mm long, 1.2-2.2
mm wide; keel mostly 0.2 mm high or
higher; beak 0.2-0.6 mm long ; stipular
veins decaying with age; basal glands
rare, to 0.3 mm diam, ..............
NEM 5a. P. foliosus var. foliosus.

1974] Potamogeton — Haynes 593

6. Spike mostly interrupted; fruit pale-
green, 1.4-1.7 mm long, 1.1-1.2 mm
wide; keel mostly less than 0.2 mm
high; beak 0.2 mm long or less; stipu-
lar veins with age remaining as long
fibers; basal glands common, to 0.5 mm
diam. .. 5b. P. foliosus var. fibrillosus.

2. Fruits with dorsal surface rounded; lateral keels
IO ec ese ne eed ve ERI eee T.

7. Stipules more or less coarsely fibrous, whitish,
the oldest tending to disintegrate into shreds;
bases of winter-buds usually indurated and corru-
gated; peduncles mostly clavate. ............ 8.

8. Leaf tips rounded to apiculate; leaves light
green to rufescent, 5-7 (-9)-nerved, 1.2-3.2
mm wide; winter buds 1.5-4.0 mm wide, inner
leaves modified into a fan-shaped structure,
outer leaves corrugated at base; peduncles
compressed; stem compressed. . 1. P. friesii.

8. Leaf tips acute, rarely obtuse, mostly deep
green to olive, 3-5 (-7)-nerved, 0.6-2.0 mm
wide; winter buds 0.8-2.2 mm wide; inner
leaves modified into a fusiform structure,
outer leaves rarely with corrugations at base;
peduncles mostly terete; stem mostly terete.
LOUER UERTASEEAITT RN FRA 2. P. strictifolius.

7. Stipules mostly delicate, whitish, green, or brown-
ish, usually disintegrating with age; bases of
winter-buds without corrugations; peduncles cy-

Hndrie ees se es ce eg RE ee 9.

9. Fruits 2.5-3.6 mm long, 1.7-2.4 mm wide;
winter buds 3.5-7.8 cm long, 2.3-5.1 mm wide,
inner leaves unmodified. .. 3. P. obtusifolius.

9. Fruits 1.5-2.2 mm long, 1.2-1.6 mm wide;
winter buds 0.9-3.2 cm long, 0.3-1.8 mm wide,
inner leaves modified into a fusiform struc-
INPUNE esc icem e iq EM RD 10.

594 Rhodora [Vol. 76

10. Leaves uninervate, subulate, 0.2-0.7 mm
wide; plants of New England and south-
ern Quebec. ............ 002 eee eee
EN Tc. P. pusillus var. gemmiparus.

10. Leaves 1-5-nerved, acute to obtuse, 0.2-
2.5 mm wide; widespread throughout
North America. ............. sess 11.

11. Mature fruit widest above the mid-
dle, sides concave, beak central; pe-
duncle filiform to cylindrical, usually
1-3 per plant; inflorescence usually
of 2-4 distinct verticels; leaves with
up to 2 rows of lacunae along mid-
rib, apex acute, rarely apiculate;
stipules mostly connate. ..........
MEM (a. P. pusillus var. pusillus.

11. Mature fruit mostly widest at or be-
low the middle, sides rounded, beak
mostly forward, peduncles cylindri-
eal, usually more than 3 per plant;
inflorescence mostly of 1-2 adjacent
verticels; leaves with 1-5 rows of
lacunae along midrib, apex acute to
obtuse; stipules mostly convolute. ..

.. Tb. P. pusillus var. tenuissimus.

1. Potamogeton friesii Rupr. in Beitr. Pflanzenk, Russ.
Reiches 4: 43. 1845. TYPE: (type not located), application
of the name is from the illustration of Potamogeton com-
pressus auct. non Linn.: In Reichenbach. Icones Florae
Germanicae et Helveticae 7: pl. 24. 1845!

Spirillus friesii (Rupr.) Nieuwl. Amer. Midl. Naturalist
3: 17. 1913.

Potamogeton pusillus var. major Fries, Novitiae Florae
Suecicae, ed. 2, 48. 1828. P. mucronatus Schr. ex Sonder,
Flora Hamburgensis 99. 1851. P. major (Fries) Morong,
Mem. Torrey Bot. Club 3: 41. 1893. TYPE: (type not lo-

1974] Potamogeton — Haynes 595

cated), application of the name is from the illustration of
Potamogeton compressum auct. non L.: In Smith, English
Bot. 2: pl. 418. 1797!

Potamogeton pusillus var. latifolius Meyer, Chloris Han-
overana, 525. 1836. P. oederi Meyer, Flora Hanoverana
Excursoria 536. 1849. TYPE: (type not located), applica-
tion of the name is from the illustration of Potamogeton
compressum auct. non L.: In Oeder, Flora Danica 2: (4)
t. 203. 1765!

Stem pale green to pale brown, simple to profusely
branched near the apex, compressed, slightly ridged, 10-
135 em long, 0.3-1.1 mm diam. Leaves usually light green,
rarely olive-green to rufescent, delicate to rigid, 5-7 (-9)-
nerved, 2.3-6.5 cm long, 1.2-3.2 mm wide; apex acute to
apiculate, glands green, greenish-brown, or gold, to 0.7 mm
diam; lacunae absent or 1 narrow row each side of the
midrib; lateral nerves joining the midrib 0.2-0.9 mm below
the apex. Stipules white, fibrous, shredding at apex, 5.5-
21.0 mm long, 0.7-2.5 mm diam. Winter buds common,
terminal or lateral, 1.5-5.0 cm long, 1.5-4.0 mm wide; inner
leaves reduced, arranged into a fan-shaped structure and
oriented at right angles to the outer leaves; outer leaves
2-3 per side, apiculate to acute, corrugated at base. Pe-
duncles usually slightly clavate, terminal or axillary, rarely
recurved, 1.2-4.1 (-7.0) em long, (0.1-) 0.5-1.2 mm diam.
Spike cylindrical, 7.0-16.0 mm long, 2.6-8.0 mm diam; ver-
ticels 2-5, 1.5-5.0 mm apart. Perianth segment 1.2-1.5 mm
long, 0.5-1.5 mm wide. Fruit olive-green to brown, without
dorsal keel, 1.8-2.5 mm long, 1.2-2.0 mm wide; beak central,
0.3-0.7 mm long, 0.2-0.5 mm diam; sides rounded, rarely
centrally depressed; wall texture smooth, rarely reticulate.
Chromosome number, 2n — 26 (Palmgren, 1939).

Distribution: Central Alaska to Newfoundland, south to
northeastern Utah and southeastern Pennsylvania, Fig. 3.
Fruiting from late June to September.

Illustrations: Fernald (1932, pl. 6; 29, fig. 3; 33, fig. 1;
and 39, fig. 8).

596 Rhodora [Vol. 76

Nomenclaturally, Potamogeton friesii has been poorly
understood. The taxon was long referred to P. compressum
L. (Smith, 1797; Oeder, 1765; Reichenbach, 1845). Fries
(1828) regarded the taxon as a variety of P. pusillus and,
therefore, proposed P. pusillus var. major. However,
Mertens and Koch (1828) had made the same combination
for a different taxon. Thus, Fries! name was a later homo-
nym. The first combination at specific rank which appar-
ently applied to this entity was P. mucronatus Schrader.
However, to my knowledge, this name was never published
by Sehrader, but first appeared in Romer and Schultes
(1818) as a nomen nudum — “Qid P. mucronatus Schrader."
I have been unable to locate the name again until used by
Reichenbach (1845), definitely placing the name in synon-

Fig. 3. Map of North America showing the documented distribu-
tion of Potamogeton friesii. The barred line represents the southern
limit of Wisconsin Glaciation in the United States east and west
of the Rocky Mountains (after Flint, 1957; Flint, et al., 1959).

1974] Potamogeton — Haynes 597

omy under P. compressum — “P. mucronatus Schrader,
R. S. III. p. 517!" Neither of these references to P. mucro-
natus constitute publication, since a name is not considered
validly published when it originally appears in synonomy.
Apparently, the name was not validly published in specific
rank until Sonder (1851) applied a description, giving
credit to Schrader.

However, Ruprecht (1845) had validly published Pota-
mogeton friesii and accurately applied the name to the en-
tity which we now consider to be this taxon. Although I
have not seen any of his original specimens, he referred to
the illustration cited above in Reichenbach (1845) and also
to one in Oeder (1765), both of which are Potamogeton
friesii,

REPRESENTATIVE SPECIMENS

CANADA: Alberta: Bow River Valley, Brown 692 (GH, NY, PH);
Banff (Vermilion Lakes), Malte & Watson 961 (CAN, GH). British
Columbia: Blackwater Lake, ca. 32 mi. N. of Golden, Hitchcock &
Martin 7645 (NY, UC). Manitoba: Cross Lake, 45 mi N of Lake
Winnipeg, Scoggan 3573 (CAN). New Brunswick: Tidehead, near
Campbellton, Malte 784 (CAN). Newfoundland: St. Georg’s Pond,
near Bay St. George, Fernald & Wiegand 2456 (CAN, GH, PH, NY).
Northwest Territories: MacKenzie River delta, East branch, 68°40'N,
Porsild 7243 (GH, US). Nova Scotia: GOLDCHESTER CO.: Salmon River,
Truro, Bissell & Linden 19698 (GH). Ontario: HASTINGS CO.: no
locality, Macoun s.n. (F, K, MO, NY, s); Charlton Island, James Bay,
52°N, 79°30’W, 4292 (CAN, US, photo at DAO). Prince Edward Island:
KINGS CO.: Black Pond, Fernald & St. John 6772 (CAN, GH, K, NY,
US); Cove Head, Macoun 3000 (CAN, GH, K, MO, NY). Quebec: Coffin
Island, Magdalen Islands, Fernald et al. 6775 (CAN, GH, K, NY, PH,
US); Ile de la Grand-Entree, Magdalen Islands, Marie-Victorin &
Rolland-Germain 9921 (F, GH, NY, PH, US). Saskatchewan: Barrier
River, ca. 4 mi. W, 3 mi. S. of McKague, Breitung 414 (CAN, DAO,
NY, UC). Yukon: Craig Lake, vicinity of Carcross, Porsild 18418
(CAN). UNITED STATES: Alaska: vicinity of College, near Fair-
banks, Argus 1097 (pao). Connecticut: NEW HAVEN CO.: Derby,
Robbins (GH, NY). Idaho: JEFFERSON CO.: Mud Lake, Sperry &
Martin 670 (us). Illinois: cook co.: South Chicago, Hill 79-1881
(ILL). Indiana: WHITLEY CO.: Goose Lake, ca. 7 mi. NW of Columbia
City, Deam 48794 (GH). Iowa: DICKINSON CO.: East Lake Okoboji,
Shimek (DAO, F, GH, MO). Maine: AROOSTOOK CO.: Washburn, Aroo-

598 Rhodora [Vol. 76

stook River, Ogden & Chamberlain 2692 (os). Massachusetts: MID-
DLESEX CO.: Fresh Pond, Cambridge, Faxon (GH, NY, US). Michigan:
CHEBOYGAN CO.: Black River between mouth and Alverno, Haynes
3377 (LAF, os); Haynes 3385 (Mo, 0S); Haynes 3698 (MO, NY, OS);
Hook Point, Douglas Lake, Haynes 3361 (F, GH, MO, NY, OS, PH, UC,
US); mouth of Bessey Creek, Douglas Lake, Haynes 3887 (os); Marl
Bay, Douglas Lake, Haynes 3711 (ASU, GH, LAF, OS). EMMET CO.:
small arm of Carp Lake, Haynes 3376 (os); Haynes 3694 (os);
Haynes 8747 (os); Haynes 3889 (08S). MANISTEE CO.: Little Manistee
River, Manistee, Hill (F, GH, ILL, MO, US). Minnesota: CASS CO.:
Cullen Lake, Ballard (ILL, MICH, MO, UC, US). MARTIN CO.: Silver
Lake, Cratty (GH, ILL, MO, NY, PH, UC, US). Montana: GLACIER CO.:
marshes above Lower Two Medicine Lakes, Glacier National Park,
Maguire 482 (GH). Nebraska: CHERRY CO.: Dewey Lake, near Valen-
tine, Tolstead 477 (GH). New York: CORTLAND CO.: Upper York
Lake, near Little York, Haynes 3347 (MICH, MO, OS, US); Haynes
3348 (F, GH, MO, NY, OS, PH, UC, US); Haynes 3349 (MICH, OS);
Haynes 3350 (MICH, 08). ONTARIO CO.: Lake Seneca, Geneva, Morong
(CAN, GH, NY, S). SARATOGA CO.: Saratoga Lake, Haynes 3311
(MICH, MO, OS). WARREN CO.: Dunham Creek near S. end of Lake
George, Haynes 3330 (GH, MICH, MO, OS, UC, US); Haynes 3331 (MICH,
MO, OS, US). North Dakota: cass co.: Fargo, Whitney (MU).
Ohio: ERIE C0.: East Bay, Sandusky, Osburn & Williamson (GH).
Pennsylvania: LEHIGH CO.: Bethlehem, Noble (GH, PH). South
Dakota: FAULK CO.: ponds and rivers, near Jacques, Geyer (US).
Utah: suMMIT co.: Lyman Lake, Blacksfork Creek, Hobson 52 (GH).
Vermont: ADDISON CO.: Little Otter Creek, Ferrisburg, Faron (GH,
US). Washington: OKANOGAN CO.: Bonaparte Lake, St. John, et al.
5298 (GH, MO, UC). Wisconsin: BROWN CO.: Point Sable, Green Bay,
Schuette (F, GH, K, MICH, UC). MILWAUKEE CO.: Milwaukee, Lapham
(GH, K, MO, PH). Wyoming: TETON CO.: shallow lake, 10 mi. s. of
Moran, Porter & Porter 7869 (DAO, UPS, UC).

2. Potamoveton strictifolius Ar. Benn. J. Bot. 40: 148. 1902.

Potamogeton pusillus var. pseudo-rutilus Ar. Benn. J.
Bot. 39: 201. 1901. P. foliosus X P. rutilus? Hagstrom,
Kongl. Svenska Vetenskapsakad. Handl. 55(5): 91. 1916.
P. strictifolius var. typicus Fern., Mem. Amer. Acad. Arts
17: 56. 1932. Type: E. J. Hill [146], Wolf Lake, Ham-
mond, [Lake Co.] Indiana, 3 Sept. 1900, (lectotype, fide
Fernald, 1932, at K, but not located; photo of lectotype,
CAN! NY!, isolectotype, ILL !).

1974] Potamogeton — Haynes 599

Potamogeton strictifolius var. rutiloides Fern. Mem.
Amer. Acad. Arts 17: 57. 1932, P. pusillus var. rutiloides
(Fern.) Boivin, Naturaliste Canad. 94: 527. 1967. TYPE:
F. P. Metcalf 1453, Molly Lake, 10 mi. N of Brainerd, Crow
Wing Co., Minnesota, 26 Aug. 1921, (holotype, GH!; iso-
type, US!).

Stem pale green to green-brown, branched or simple,
more or less rounded, slighty ridged, 27-95 cm long, 0.4-
0.8 mm diam. Leaves green to olive, usually rigid, 3-5 (-7) -
nerved, 1.2-6.3 cm long, 0.6-2.0 mm wide; apex acute to
nearly bristle-tipped; glands white, green, greenish-brown,
or gold, to 0.3 mm diam; lacunae usually absent; lateral
nerves joining the midrib 0.8-1.5 mm below the apex.
Stipules usually white, fibrous, shredding at tip, connate,
0.6-1.6 cm long, 0.6-2.0 mm diam. Winter buds common,
terminal or lateral, 2.5-4.8 cm long, 0.8-2.2 mm wide; inner
leaves undifferentiated; outer leaves 3-4 per side, acute,
mostly without, rarely with, corrugations at base. Pedun-
cles usually cylindric, rarely slightly clavate, terminal,
mostly erect, rarely recurved, 1.0-4.5 em long, 0.3-0.9 mm
diam. Spike cylindrical, 0.6-1.3 em long, 1.5-5.2 mm diam;
verticels 3-4, 1.5-3.2 mm apart. Perianth segments 0.9-1.7
mm long, 0.8-1.5 mm wide. Fruit green-brown, without
dorsal or lateral keels, 1.9-2.1 mm long, 1.3-1.8 mm wide;
beak central, 0.3-0.5 mm long, 0.2-0.4 mm diam; sides
rounded, often centrally depressed; wall texture smooth.
Chromosome number, 2n == 52 (Lóve, 1954).

Distribution: Northwestern Northwest Territories to
eastern Quebec, south to northeastern Utah and north-
western Connecticut. Fruiting from early July to late
September. Fig. 4.

Illustrations: Fernald (1932, pl. 7; 8; 29, fig. 4, 5; and
33, fig. 2, 3).
Bennett, in the original description of Potamogeton pusil-

lus var. pseudo-rutilus, cited two collections: *Lake Scugog,
Ontario, Canada, 1897, W. Scott, ex Prof. Macoun;" and

600 Rhodora [Vol. 76

*Wolf Lake, Indiana, U.S.A., 1900, Rev. E. J. Hill." After
receiving more material from Hill, Bennett later raised the
taxon to specific rank. Hagstrom (1916) examined addi-
tional material sent to him by Hill and ascertained this to
be a hybrid between P. foliosus and P. rutilus. Hagstrom
cited the original specimen as being from Lake George,
East Chicago, rather than from Wolf Lake. I think Hag-
strom made an error in citing this specimen, or possibly
Lake George and Wolf Lake refer to the same lake. The
original most certainly was from Wolf Lake. Hagstrom did
not examine material from Lake Scugog, but based his
decision on the 1902 description of P. strictifolius by Ben-
nett and other Canadian material he had examined from
Sea-Cow-Pond, Crane Lake, Assiniboia, Picanok River,
Quebec, and Ottawa River, Quebec [which he said “seem
excellently to answer to the description of the fruiting plant
of Lake Scugog"]. He concluded that the Lake Scugog ma-
terial was probably the same as P. panormitanous (P.
pusillus var. pusillus). Fernald examined Bennett's collec-
tion but overlooked the point of whether the Lake Scugog
plant was the same taxon as the Wolf Lake plant. As he
could not be certain that the two were the same, he did not
deny or accept Hagstrom's decision. He did, however, desig-
nate Hill's collection from Wolf Lake, Indiana, September
3, 1900, as the Type [lectotype] of P. strictifolius. Fernald
did say, with which I concur, that Hagstrom had erred in
calling P. strictifolius a hybrid of P. foliosus and P. rutilus.

I also am unable to state with assurance that the speci-
men from Lake Scugog is conspecific with the one from
Wolf Lake. I have corresponded with curators of the
herbaria (K, BM, & CGE) where the majority of Bennett's
material is located. There is no sheet in Bennett's collection
at these herbaria filed under Potamogeton pusillus, P. berch-
toldii, or P. strictifolius. As I have photographs from CAN,
NY, & US of a sheet with only the two specimens mounted
on it, I am certain that it is not correctly filed under the
name of another specimen on the same sheet. The photo-
graph from CAN has a paragraph in Bennett's handwriting

1974] Potamogeton — Haynes 601

stating that the sheet was in his collection and that the
photograph was taken by his daughter.

I have, however, located a duplicate of Hill's collection
from Wolf Lake, and also a second specimen collected by
Seott at Lake Seugog on 3 August 1897 (the same day as
the one in Bennett's collection). I can not ascertain if
Scott's collection is a duplicate of the one sent to Bennett.
With its convolute stipules, characteristic winter buds, and
one row of lacunae along each side of the midrib, this
specimen is rather characteristic of P. pusillus var. tenuis-
simus. It is, however, unusual in that the stipules are white
rather than the usual greenish-brown of P. pusillus var.
tenuissimus. Until the specimen in Bennett’s collection can
be located, I am following Fernald in that I cannot be sure
to which taxon the Lake Scugog collection should be re-
ferred.

Fernald (1932) separated Potamogeton strictifolius into
two varieties, var. strictifolius & var. rutiloids, based on
the leaf apex shape, rigidity of the leaves, and the coarseness
of the stipules. After examining many sheets and hundreds
of plants in the field, I have concluded that this variability
is both within and between populations. The leaf tips of the
isotype of var. strictifolius at ILL, for example, vary from
very gradually tapering acute to obtuse. Because of this
variability within populations and of no apparent range
distinctions, I consider P. strictifolius to be one taxon. The
rigidity of the leaves, I believe to be of ecological rather
than of genetic origin.

Morphologically, Potamogeton strictifolius is most similar
to P. friesii. The best way to distinguish the species is by
the winter buds. In P. strictifolius, the inner leaves of the
winter buds are only slightly modified, whereas in P. friesii
the inner leaves are much shorter than the outer leaves and
are oriented at 909 to the outer leaves. Differences in the
leaves are difficult to assess. Fernald (1932) and Ogden
(personal communication) stress the number of veins —
three to five in P. strictifolius and five to seven in P. friesii.

602 Rhodora [Vol. 76

Street L0 bee.

~?

vex = EN

Fig. 4. Map of North America showing the documented distribu-
tion of Potamogeton strictifolius. The barred line represents the
southern limit of Wisconsin Glaciation in the United States east and
west of the Rocky Mountains (after Flint, 1957; Flint et al., 1959).

1974] Potamogeton — Haynes 603

For both species, I find, as does Voss (1972), that the usual
number is five. If specimens are sterile and lacking winter
buds and I must use vegetative characters, I then rely upon
the apex shape and whether the leaves are revolute. In P.
strictifolius, the apex is usually acute and the leaves are
revolute. In P. friesii, the apex is obtuse to apiculate and
the leaves are usually flat. However, as I indicated earlier
and pointed by Voss (1972) one must use these characters
with caution. In the late summer the margins of P. friesii
roll under, thus the leaves become revolute.

REPRESENTATIVE SPECIMENS

CANADA: Alberta: Large pond, Lacombe, Dixon 691 (DAO, NY).
Manitoba: York Factory, Drummond (GH). Northwest Territories:
MACKENZIE DIST.: E branch of Mackenzie River, 68°40’ to 68°55’,
Reindeer Station, Porsild 7234 (CAN). Ontario: THUNDER BAY DIST.:
Black Fox Lake, Haynes 3750 (CAN, GH, LAF, MO, 0S); Nipigon, Lake
Superior, Macoun (CAN, NY, S). Quebec: GATINEAU CO.: Danford
Lake, 45 mi. N of Ottawa, Dore 9032 (DAO, MO). Saskatchewan:
Pike Lake, Saskatoon, Fraser 8 (DAO, GH). UNITED STATES:
Connecticut: LITCHFIELD CO.: Indian Pond, Sharon, Eames 11868
(GH). Illinois: cook co.: Wolf Lake, Chicago, Chase 1709 (CGE, F,
GH, ILL). Indiana: LAKE C0.: Lake George, East Chicago, Hill 144-
1903 (GH, ILL, NY). Michigan: CHARLEVOIX CO.; Cunningham Lake,
ca 6 mi SW of Ironton, Voss & Haynes 13602 (MICH). CHEBOYGAN
co.: Black River between its mouth and Alverno, Haynes 3379 (GH,
MO, OS, US), Haynes 3384 (Mo, os, US), Haynes 3743 (GH, 0S); Black
Lake, near mouth of Upper Black River, Haynes 3777 (os); Cheboy-
gan River, Haynes 3798 (os); Douglas Lake near source of Maple
River, Haynes 3739 (GH, MICH, MO, NY, OS, UMBS, US). WAYNE CO.:
Detroit River near Belle Isle, Wheeler (GH, ILL, NY, US). Minnesota:
BECKER CO.: DeSota Lake, Grant 3269 (GH, Ny, UC, US). Nebraska:
CHERRY CO.: Willow Lake, Thomson 122 (Us). New York: MONROE
co.: Clausen & Hinkey 4196 (GH, NY). North Dakota: MCHENRY
co.: Red Willow Lake, Mabbott 372 (US). Ohio: ERIE CO.: Sandusky
Bay, Pieters (US). Pennsylvania: ERIE CO.: Presque Isle, Garber
(GH). Utah: BOX ELDER CO.: Bear River, Watson 1136 (GH, K,
NY, US). Vermont: CALEDONIA CO.: Sarah Moor Pond, 2 mi NW of
Barnet, Hotchkiss 7804 (Us). Wisconsin: WASHINGTON CO.: Big
Cedar Lake, Hotchkiss & 'Hoehler 4264 (GH, US). Wyoming: ALBANY
CO.: reservoir ca 6 mi. NE cf Lookout, Porter 7370 (DAO, NY, UC,
USP). PARK CO.: Firehole River S of Madison Junction, Porter 6385
(DAO, MO, NY, UC, UPS).

604 Rhodora [Vol. 76

Potamogeton strictifolius X zosteriformis

Fernald (1932) named Potamogeton longiligulatus based
upon flowering specimens from Newfoundland. The entity
was said to resemble P. hillii by its bristle-tipped leaves
but differed from the latter species by having numerous-
nerved leaves, Voss (1967) reported that, at least in north-
ern Michigan, the entity appeared to be a result of hybridi-
zation between P. strictifolius and P. zosteriformis, a spe-
cies in the subsection Compressi of Hagstrom. Data gath-
ered during my field studies in northern Michigan support
Voss' conclusions. I have seen several instances where the
putative hybrid was growing between populations of P.
zosteriformis and P. strictifolius. The plants of the putative
hybrid were morphologically intermediate between the two
species. Therefore, until the putative hybrid can be studied
in detail, I accept Voss' concept of the taxon.

SPECIMENS EXAMINED:

CANADA: Newfoundland: Straits of Belle Isle, pond in barrens
S of Flower Cove, Fernald & Long 27330, (holotype, GH!; isotype,
PH!, photo at DOA!). Ontario: BRUCE CO.: Bruce Peninsula, Stokes
Bay, Krotkov 8627 (Us). LAMBTON C0.: Sarnia Bay, Dodge (GH,
photo at DAO). UNITED STATES: Connecticut: LITCHFIELD CO.:
Salisbury, Twin Lakes, Bissell (GH). Illinois: LAKE CO.: Grayslake,
Dolbeare 1444 (MicH). Michigan: CHEBOYGAN CO.: Black River
between its mouth and Alverno, Bedell & Prescott 55 (UMBS), Haynes
3380 (GH, Mo, os, us), Haynes 3381 (GH, MO, os, US), Haynes 3702
(os), Haynes 3744 (os), Majors (GH, NY, UMBS), Voss 11118 (GH),
Voss 11359 (UMBS); Black Lake, bay at mouth of Black River,
Stuckey 2411 (os), Stuckey 3275 (os, CAN), Voss 11745 (GH, NY),
11760 (UMBS); Cheboygan River, Haynes 3799 (OS). ST. CLAIR CO.:
Algonac, Big Ditch, Dodge s.n. (US); Port Huron, Dodge s.n. (NY).
SCHOOLCRAFT CO.: Lake McDonald, SE of Whitedale, Uhler 96 (Us).
Minnesota: BECKER C0.: Cotton Lake, Hotchkiss 6341 (GH). HEN-
NEPIN C0.: Lake Minnetonka, Keck & Stilwill 416 (GH). New York:
COLUMBIA CO.: Beebe Pond near Queechy Lake, Haynes 3344 (os),
Haynes 3801 (os), Smith & Ogden 45590 (os).

3. Potamogeton obtusifolius Mert. & Koch, in Róhling,
Deutschland Flora 855. 1823. TYPE: (type not located),
application of the name is from the illustration of Potamo-

1974] Potamogeton — Haynes 605

geton gramineus auct. non L.: In Smith, English Bot. 11:
pl. 2253. 1811.

Spirillus obtusifolius (Mert. & Kóch in Róhling) Nieuwl.
Amer. Midl. Naturalist 3: 19. 1913.

Potamogeton compressum. var. tenwius Wahl. Flora Up-
saliensis 60. 1820. TYPE: G. Wahlenberg, Upsala Sraklen,
Ekebysja near Quam, 22 Aug. 1818, (holotype, UPS!).

Potamogeton obtusifolius var. angustifolius Fieber in
Berchtold, Oekon.-tech. Flora Bohmens 275. 1838. TYPE:
(not located).

Potamogeton obtusifolius var. latifolius Fieber in Berch-
told, Oekon.-tech. Flora Bohmens 275. 1838. TYPE: (not
located).

Stem green to green-brown, slightly compressed, usually
without ridges, 35-90 cm long, 0.3-0.7 mm diam. Leaves
light green to rufescent, usually flaccid, 3-nerved, 3.0-8.2
em long, 1.0-3.5 mm wide; apex round to round apiculate;
glands yellow-green to gold, 0.2-1.0 mm diam; lacunae of
1-3 rows each side of midrib; lateral nerves joining the
midrib 0.2-1.2 mm from apex. Stipules usually white,
slightly fibrous, rarely shredding at tip, convolute, 0.6-1.8
em long, 0.5-1.2 mm diam. Winter buds abundant, terminal,
3.5-7.8 em long, 2.3-5.1 mm wide; inner leaves undifferen-
tiated; outer leaves 3-4 per side, apiculate to obtuse, with-
out corrugations at base. Peduncles cylindrical, axillary,
rarely recurved, 0.8-1.9 (-4.2) em long, 0.5-1.0 mm wide.
Spike cylindrical, 0.8-1.3 cm long, 4.6-7.0 mm wide; verti-
cels 3, crowded or to 2.0 mm apart. Perianth segments 1.4-
1.5 mm long, 1.1-1.3 mm wide. Fruit olive-green to brown,
2.5-3.6 mm long, 1.7-2.4 mm wide; keels absent or present,
when present, ridged, to 0.2 mm high; beak mostly central,
rarely forward, (0.2-) 0.6-0.7 mm long, 0.5-0.7 mm diam;
sides rounded; wall texture smooth or rough. Chromosome
number 2n — 26 (Palmgren, 1939).

Distribution: Eastern Yukon to eastern Quebec, south to
Washington, Wyoming, and New Jersey. Fruiting from
early July to late September. Fig. 5.

606 Rhodora [Vol. 76

Illustrations: Fernald (1932, pl. 14; 30, fig. 3; 34, fig. 4;
39, fig. 1).

In its winter buds, fruit, and peduncles, Potamogeton
obtusifolius most closely resembles P. hillii. When in fruit,
however, it can easily be separated from the latter species
by its spikes being cylindric, whereas those of P. hillii are
capitate. Sterile specimens of P. obtusifolius can usually
be distinguished from P. hillii by the former having obtuse
leaves and the latter having acute leaves. However, plants
of P. hillii occasionally possess obtuse leaves.

o. uo CASH u T RRA;

` FAR AUTRES (CON, TA NES

>; mw YIN AG

= f 4d: Mac 2.* Jj Ys Y
BANNA
Fig. 5. Map of North America showing the documented distribu-
tion of Potamogeton obtusifolius. The barred line represents the
southern limit of Wisconsin Glaciation in the United States east and
west of the Rocky Mountains (after Flint, 1957; Flint et al., 1959).

h.

REPRESENTATIVE SPECIMENS

CANADA: Alberta: ca. 2 mi. W of Glenevis, Moss 12449 (CAN,
DAO). British Columbia: VANCOUVER ISLAND: Cowichen, Glendenmire
(CAN, GH). Manitoba: Fort Churchill, 58*67'N, 94^10'W, Schofield
& Crum 6787 (CAN, F). Northwest Territories: MACKENZIE DIST.:
small pool ca. 45.5 mi. WNW of Yellowknife, Thieret & Reich 7947

1974] Potamogeton — Haynes 607

(F, os, US). Nova Scotia: CAPE BRETON CO.: Mira Bay, Macoun
20755 (F, GH, CAN). Ontario: THUNDER BAY DIST.: Black Fox Lake,
Haynes 3749 (CAN, GH, LAF, MO, OS, UC); Sibley Twp., Ravine Lake,
Taylor et al. 232 (CAN, GH, uc). Quebec: Lake Memphremagog,
Sargents Bay, Churchill (GH, K, MO). GASPE CO.: Riviere Bonaven-
ture, Marie-Victorin et al. 33764 (F, GH, NY, PH). Saskatchewan:
MACKENZIE DIST.: Meridian Creek near Amisk Lake, Hudson 1550
(DAO). Yukon: Sheldon Lake, Canol Road mi. 222, Porsild & Breitung
11511 (CAN). UNITED STATES: Connecticut: FAIRFIELD CO.:
Norwalk River, Ridgefield, Eames 11717a (GH). Maine: PENOBSCOT
co.: Pushaw Stream, Old Town, Ogden et al. 1617 (CAN, GH, MO, NY,
US). PISCATAQUIS CO.: cold clear stream, Foxcroft, Fernald 478
(GH, MO, US). Massachusetts: ESSEX CO.: ditch in Wenham, Morong
(F, GH, ILL, PH); Pleasant Pond, Wenham, Faxon (GH, NY, US),
Ogden & Ogden 1763 (CAN, DAO, F, GH, ILL, MICH, MO, NY, PH, UC,
UPS, US). Michigan: KEWEENAW CO.: Beaver Pond, ca. 4 mi, S of
Central Mine, Robbins (GH, MO, NY, PH). LUCE CO.: Bodie Lake,
Haynes 8787 (0S). SCHOOLCRAFT CO.: Canoe Lake, Haynes 3741la
(os). Minnesota: sT. LOUIS CO.: West Two Rivers, Tower, Hill 218-
1889 (CAN, ILL). Montana: GLACIER CO.: Howe Lake, Glacier Na-
tional Park, Hazzard 480 (GH). New Hampshire: GRAFTON CO.:
pond, Enfield, Kennedy (GH). New Jersey: MORRIS CO.: Rockaway
River near Milton, Williamson (GH, PH). New York: ALBANY CO.:
Alcove, Shear (GH, NY, UC). Pennsylvania: WAYNE CO.: Howell’s
Pond, Twining (GH, PH). Rhode Island: PROVIDENCE CO.: Fountain
Spring Brook, Smithfield, Lovewell (GH). Vermont: ADDISON CO.:
Little Otter Creek, Ferrisburg, Eggleston & Grout (F, GH, PH, US).
CALEDONIA CO.: spring-fed pond, ca. 2 mi. N of Walden Village,
Haynes 3829 (os); Coles Pond, Walden Twp., Haynes 3837 (os).
Washington: SAN JUAN CO.: Summit Lake, Orcas Island, Sutherland
& Kern 1182 (CAN, NY, UC). Wisconsin: ONEIDA CO.: stream, Three
Lakes, Hoffman (GH, MO). Wyoming: TETON CO.: Beaver ponds
near Moose, Porter & Porter 9405 (GH, UC).

4. Potamogeton hillii Morong, Bot. Gaz. (Crawfordsville) 6:
290. 1881. TYPE: E. J. Hill, Stagnant pools, Manistee,
[Manistee Co.] Michigan, (holotype, NY!; isotypes, F[2
sheets] !, aH [2 sheets] !, ILL[4 sheets] !, K!, PH!).

Potamogeton porteri Fern. Mem. Amer. Acad. Arts 17:
73. 1932. TYPE: T. C. Porter, Run in Dillerville Swamp,
near Lancaster, [Lancaster Co.] Pennsylvania, (lectotype,
PH!; isolectotypes, F[2 sheets]!, MO !, NY[2 sheets] !, PH[2
sheets] !).

608 Rhodora [Vol. 76

Stems green to olive, slightly compressed, heavily ridged,
30-60 em long, 0.5-1.0 mm diam. Leaves pale-green to olive-
green, delicate, 3-nerved, 2.0-6.0 cm long, 0.6-2.5 (-4.0) mm
wide; apex apiculate to bristle-tipped; glands present or
absent, brown to green, 0.1-0.3 mm diam; lacunae of 1-2
rows each side of midrib; lateral nerves joining midrib
(0.4-) 0.7-1.7 mm from apex. Stipules white to light brown,
slightly fibrous, rarely shredding at tip, convolute, 7.0-16.0
mm long, 0.6-2.2 mm diam. Winter buds rare, terminal,
2.8-3.0 em long, 1.5-3.0 mm wide; inner leaves undifferenti-
ated; outer leaves 3-4 per side, acute to apiculate, without
corrugations at base. Peduncles slightly clavate, axillary
or terminal, rarely recurved, 6.0-13.5 mm long, 0.3-1.0 mm
diam. Spike globose, (2.0-) 4.0-7.0 mm long, 4.6-7.0 mm
diam; verticels 1-2, when 2, these crowded, 0.5-1.0 mm
apart. Perianth segments 1.3-1.5 mm long, 1.1-1.5 mm wide.
Fruit brown to light greenish-brown, dorsally and laterally
keeled, 2.3-4.0 mm long, 2.0-3.2 mm wide; keels forming
ridges, without undulations, to 0.2 mm high; beak central,
rarely forward, 0.3-0.7 mm long, 0.2-0.6 mm diam; sides
rounded rarely centrally depressed; wall texture rough.
Chromosome number unknown.

Distribution: In cold stagnant or slow moving, often
brown water, from northern Lower Peninsula Michigan to
Vermont and south to northeastern Ohio and southeastern
Pennsylvania. Fruiting from late June to late August.
Fig. 6.

Illustrations: Fernald (1932, pl. 13; 30, fig. 2; 34, fig. 2).

In the original description, Morong cited only the follow-
ing data: “excellent specimens obtained in August 1880, by
Mr. Hill, at Manistee, Michigan . . ." He included no indi-
cation as to the location of the specimen. Hill prepared,
from Manistee in August 1880, no less than 14 herbarium
specimens on at least two separate days. Fernald (1932,
p. 169), also giving no specimen location, designated as the
type collection the one gathered by Hill on 10 August 1880.

1974] Potamogeton — Haynes 609

However, in the collection at NY there is a specimen col-
leeted on 5 August 1880 by Hill and labeled in Thomas
Morong's handwriting as “Type Specimen." Of the many
other specimens examined by Morong, none of these others
were indicated as types by Morong. Therefore, the 5 Au-
gust specimen, having been designated by the author, then
is the holotype and the one cited by Fernald then is a topo-
type.

Fernald (1932) named Potamogeton porteri based on
several sheets collected by Thomas Porter in mill ponds
near Lancaster, Pennsylvania. In the body of the original
description Fernald did not cite any one specimen as the
type. The data he gave are “PENNSYLVANIA: Cold
streams or rivulets in Dillerville Swamp, Lancaster Co.,
October 5 and 6, 1860, Thos. C. Porter, distributed as P.
pusillus and P. obtusifolius (TYPE in Herb. Acad. Sci. Nat.
Phila.; duplicate in Herb. Gray; fragmentary (sterile) du-
plicates in Herb. Field Mus., Dudley, and Mo. Bot. Gard.).”
In a footnote on the same page he states ‘“Porter’s labels,
all of material collected on October 5, with the exception of
one sheet marked October 6, indicate somewhat different
habitats: ‘Cold rivulets near Lancaster,’ ‘Near Lancaster,’
‘In a rivulet, Dillerville swamp, Lancaster,’ ‘Stream in Dil-
lerville Swamp, near Lancaster,’ ‘Run in Dillerville Swamp,
near Lancaster.’ ”

I have seen the specimens at PH, GH, F, MO, and none of
the sheets have the locality data given by Fernald in the
body of the text. There are, in fact, three sheets at PH
labeled by Fernald as P. porteri. He indicated “Type Col-
lection” on two of these sheets, “Run in Dillerville Swamp,
near Lancaster, Penn. October 5, 1860” and “Stream in Dil-
lerville Swamp, near Lancaster, Penn. October 5, 1860.”
On the other sheet, ‘Cold rivulets near Lancaster, Pa.
October 5," Fernald did not indicate “Type Collection.”
The two specimens labeled as “Type Collection” by Fernald
are obviously the same taxon and both match the descrip-
tion. However, the one with data of “Stream in Dillerville
Swamp, near Lancaster, Penn. October 5, 1860” is fertile

610 Rhodora [Vol. 76

while the other is sterile. Hence, I am here designating the
fertile one as the lectotype.

Fernald states (p. 74), “In its large fruit with prominent
keel and recurved beak and its essential lack of glands at
the base of the stipules P. porteri is close to P. hillii, but
the latter species is wholly different in its acute, attenuate
and thinner stipules; and in its attenuate almost bristle-
tipped leaves with a single row of lacunae each side of
the midrib and with the lateral nerves confluent with the
midrib well below the tip." The specimens annotated by
Fernald as P. porteri vary from ones with narrow leaves
with apiculate leaf tips to very broad ones with obtuse leaf
tips. The broad leaved specimens are quite distinct from
characteristic P. hillii. However, the ones with the nar-
rower leaves closely resemble P. hillii in all characters men-
tioned by Fernald. A locality of P. hillii in Cheboygan
County, Michigan, first located by Dr. E. G. Voss in 1967,
was visited by me during the summers of 1970, 1971, and
1972. I noticed a variability within the population from
individuals with the characteristic narrow attenuate leaves
with delicate stipules to ones with broad obtuse leaves with
coarse stipules. Within this one population, a complete
gradation exists from characteristic P. hillii to character-
istic P. porteri. For this reason I believe P. porteri is a
morphological extreme of P. hilli.

With the capitate inflorescence, short axillary peduncles,
and fruits with a small dorsal keel, Potamogeton hillii
closely resembles P. foliosus. However, it can be distin-
guished from the latter species by the large 2.3-4.0 X 2.0-
3.2 mm fruits (the largest of any Pusilli), by the nearly
bristle-tipped leaves and by the stipules being convolute.
By the nearly bristle-tipped leaves, some individuals re-
semble P. longiligulatus, a proposed hybrid, but can be dis-
tinguished from this putative hybrid by the leaves having
only 3 veins, whereas those of P. longiligulatus have mostly
7-9 veins. From all other Pusilli, P. hillii can be separated
by its large fruits, short peduncles, and capitate inflores-
cences.

1974] Potamogeton — Haynes 611

Fernald (1932) states that the Pusilli reproduce, for the
most part, not by seeds, but by winter buds. He also states
that winter buds are unknown in Potamogeton hillii. I have
seen one sheet with well developed winter buds and several
with ones that were immature. The winter buds, when
present, with the inner leaves being unmodified, resemble
small ones of P. obtusifolius in shape and structure, even
in having the leaves round apiculate.

Potamogeton hillii has been considered to be extremely
local throughout its range. This distribution is becoming
more apparent with the destruction of the aquatic environ-
ment by pollution and dredging. In fact, Voss (1967) re-
ported that the type locality of P. hillii has been destroyed.

UN

2 WARES

Fig. 6. Map of eastern North America showing the documented
distribution of Potamogeton hillii. The barred line represents the
southern limit of Wisconsin Glaciation (after Flint et al, 1959).

612 Rhodora [Vol. 76

Around 1900, the species was present in northeastern Ohio
and northwestern Pennsylvania. This area has become
heavily populated, with few if any natural areas remaining.
The two Ohio counties, Portage and Ashtabula, have been
rather heavily collected over the past 20 years. Some col-
lectors thought they had collected the species, but after
examining the specimens, I found them all to be P. pusillus.
On the other hand, intensive field work in northern Michi-
gan has shown the species to occur in no fewer than four
localities within a radius of 30 miles of Cheboygan County.
The same situation may prove to be true in other less heav-
ily populated areas of its range, especially northeast New
York state, as botanists continue to search for the species.

REPRESENTATIVE SPECIMENS

CANADA: Ontario: ELGIN CO.: near St. Thomas, James (DAO).
UNITED STATES: Connecticut: LITCHFIELD CO.: Indian Pond,
Sharon, Eames 11867 (GH). Massachusetts: BERKSHIRE CO.; pond
fed by Karner Brook, South Egremont, Weber 1626 (CAN, F, GH,
MICH, MO, NY, UC, US). Michigan: CHEBOYGAN CO.: Elliot Creek,
4 mi. E of Cheboygan, Haynes 3389 (GH, MO, OS, UC, US), Haynes
3713 (os). EMMET CO.: Cecil Bay marsh, 4 mi. SW of Mackinaw
City, Voss 14061 (CAN, GH, MICH, MSC, NY, OS, UMBS, WIS). MAC-
KINAC CO.: small stream ca. 1 mi. S of Engadine, Haynes 4025 (ASC,
DAO, F, FSU, GH, K, MICH, MO, MSC, NY, OS, PH, SMU, TENN, TEX, UC,
US, VDB, WIS, WVA). MANISTEE CO.: Manistee, Morong (GH, MICH,
NY, US). OTSEGO CO.: outlet of Grass Lake, Stuckey 1300 (GH, MICH,
NY, OS, UMBS). PRESQUE ISLE CO.: mouth of Black Mallard River,
Haynes 3390 (F, GH, MO, NY, OS, PH, UC, US), Haynes 3714 (08),
Haynes 3740 (ASU, GH, LAF, MO, NY, OS, PH, QFA, UC, VDB, US), Haynes
3890 (os). New York: THOMPKINS CO.: Ithaca, Lake Cayuga, Dudley
(CAN, F, GH, MICH, MO. US). WASHINGTON CO.: creek 2 mi. N of
Dresden Station road, Haynes 3342 (GH, MICH, MO, OS, UC, US).
Ohio: ASHTABULA CO.: pools, Ashtabula, Hill (F, ILL). Pennsylvania:
BEDFORD CO.: millpond S of Woodbury, Hotchkiss 6003 (GH, US).
Vermont: WINDSOR CO.: Evarts Pond, Windsor, Dudley (GH, NY,
UC, US).

5. Potamogeton foliosus Raf., Med. Repos. Hexade 2, 5:
354. 1808.

Stems green to olive, slightly compressed and ridged,
4-75 em long, 0.2-1.2 mm diam. Leaves pale green to olive,

1974] Potamogeton — Haynes 613

rarely rufescent, delicate, 1-3 (-5)-nerved, 1.3-8.2 cm long,
0.3-2.3 mm wide; apex acute to apiculate, rarely with a
bristle; glands present or absent, black to gold, to 0.5 mm
diam; lacunae absent or to 2 rows each side of midrib;
lateral nerves joining the midrib 0.5-1.2 mm below the
apex. Stipules greenish to brown, rarely white, delicate to
fibrous, shredding or not at the tip, connate or convolute,
0.2-2.2 em long, 0.2-1.7 mm diam. Winter buds uncommon,
lateral or terminal, 0.9-2.5 em long, 0.6-2.0 mm wide; inner
leaves rolled into a hardened fusiform structure; outer
leaves 1-3 per side, acute to apiculate, without corrugations
at base. Peduncles clavate, mostly in axiles of lower, rarely
upper, leaves, usually recurved, 0.3-1.1 (-3.7) cm long, 0.3-
1.4 mm diam. Spikes capitate to cylindric, 1.5-7.0 mm long,
1.0-6.0 mm diam; verticels 1-2, when 2, these usually
crowded, 0.6-1.2 (-2.0) mm apart. Perianth segments 0.4-
1.4 mm long, 0.3-1.0 mm wide. Fruit pale green to olive or
brown, dorsally keeled, 1.4-2.7 mm long, 1.1-2.2 mm wide;
keel undulate winglike, to 0.4 mm high; beak central, rarely
forward, 0.2-0.6 mm long, 0.1-0.4 mm diam; sides rounded
to centrally depressed; wall texture smooth.

Potamogeton foliosus is most similar to P. hillii in its
short clavate peduncles and capitate inflorescence. How-
ever, when in fruit, P. foliosus can be separated from the
latter species by the presence of a dorsal undulate wing
on the fruit. The fruit of P. hillii, on the other hand, is
three-keeled — two lateral and one dorsal ridge which does
not appear as a thin undulating wing. Vegetatively, P.
foliosus is most similar to P. pusillus. From P. pusillus var.
tenuissimus, P. foliosus often can be distinguished by the
near lack of lacunae. From P. pusillus var. pusillus, how-
ever, the separation is not that simple. Fernald (1932)
suggests that P. foliosus may be separated from P. panor-
mitanus [var. pusillus] by the absence of nodal glands.
Voss (1972), on the other hand, has questioned this char-
acter as reliable. In most fruiting specimens of P. pusillus
[var. pusillus] from Michigan, Voss could not find evident
nodal glands. Data gathered by me from examination of

614 Rhodora [Vol. 76

hundreds of herbarium specimens and many populations in
the field support Voss's conclusions. Therefore, I give little
value to the absence of glands as a character for separating
these two taxa. Although intermediates occur, the only
character which I have been able to consider with any re-
liance is the coarseness of the stipular veins. Those of
P. foliosus, for example, are usually evident, appearing as
ridges extending the length of the stipules. The veins are
not usually so evident in P. pusillus.

5a. Potamogeton foliosus var. foliosus

P. foliorum Raf., Med. Repos. Hexade 3, 2: 409. 1811.
(orthographic variant). P. pauciflorus Pursh, Flora Amer.
Sept. 121. 1814. P. foliosus Raf. var. genuinus Fern., Mem.
Amer. Acad. Arts 17: 43. 1932. TYPE: Michaux, in rivis
affluente mari inundatis Carolinae inferioris [South Caro-
lina] (holotype, P; photograph of holotype, aH !).

Potamogeton niagarensis Tuckerm., Amer. J. Sci. Arts
Series 2, 7: 354. 1849. P. pauciflorus Pursh var. niagaren-
sis (Tuckerm.) Robbins in Gray, Man. Bot. North. U. S.,
ed. 2. 435. 1856. P. foliosus Raf. var. niagarensis (Tuck-
erm.) Morong, Mem. Torrey Bot. Club 3: 39. 1895. Spiril-
lus foliosus (Raf.) Nieuwl. var. niagarensis (Tuckerm.)
Nieuwl., Amer. Midl. Naturalist 3: 18. 1913. P. foliosus
Raf. f. niagarensis (Tuckerm.) Hagstrom. Kongl. Svenska
Vetenskapsakad. Handl. 55(5): 91. 1916. TYPE: Tucker-
man, Hogback, Niagara Falls, [Niagara Co.] New York,
(holotype, NY!; isotypes, GH !, K!).

Potamogeton pauciflorus Pursh var, californicus Morong,
Bot. Gaz. (Crawfordsville) 10: 254. 1885. P. foliosus Raf.
var. californicus (Morong) Morong, Mem. Torrey Bot. Club
3: 40. 1895. P. californicus (Morong) Piper, Contr. U.S.
Natl. Herb. 11: 98. 1906. P. foliosus Raf. f. californicus
(Morong) Hagstrom, Kongl, Svenska Vetenskapsakad.
Handl. 55(5): 91. 1916. TYPE: S. B. & W. F. Parish 940,
submerged aquatic, streams, San Bernardino, San Ber-
nardino Co., California, (holotype, NY!; isotypes GH !, MO!,
PH!).

1974] Potamogeton — Haynes 615

Potamogeton foliosus Raf. var. macellus Fern., Mem.
Amer. Acad. Arts 17: 46. 1932. TYPE: E. & C. E. Faxon
Fresh Pond, Cambridge, [Suffolk Co.] Massachusetts, (holo-
type, GH ; isotype, US!).

Stems 4-72 cm long, 0.2-1.2 mm diam. Leaves 1-8 (-5)-
nerved, 1.3-8.2 cm long, 0.3-2.3 mm wide; apex acute to
apiculate, rarely with a bristle; glands rare, to 0.3 diam;
lacunae absent or to 2 rows each side of midrib; lateral
nerves joining the midrib 0.5-1.2 mm below the apex.
Stipules greenish to brown, delicate to slightly fibrous,
rarely shredding at the apex, connate, 0.2-2.2 cm long, 0.2-
1.7 mm diam. Winter buds lateral, 0.9-1.8 cm long, 0.6-1.0
mm wide. Peduncles usually clavate, recurved, 0.3-1.1 (-3.7)
em long, 0.3-1.4 mm diam. Spikes usually capitate, rarely
cylindric, 1.5-7.0 mm long, 1.0-6.0 mm diam; verticels 1-2,
when 2, these crowded, 0.6-1.2 (-2.0) mm apart. Perianth
segments 0.4-1.4 mm long, 0.5-1.0 mm wide. Fruit olive to
green-brown, 1.5-2.7 mm long, 1.2-2.2 mm wide; keel 0.1-
0.4 mm high; beak 0.2-0.6 mm long, 0.2-0.4 mm diam.
Chromosome number, 2n — 28 (Stern, 1961).

Distribution: Widespread, in waters of lakes, springs,
streams and rivers, from central Alaska to Nova Scotia and
south to southern Guatemala and Jamaica. Fruiting from
late May to late October. Fig. 8.

Illustrations: Fernald (1932, pl. 3; 29, fig. 1; 32, fig. 3;
38, fig. 1).

Potamogeton foliosus was first recognized by Michaux
(1803) when he reported P. gramineus? L. for North
America. Rafinesque (1808), noticing that many of the
Western Hemisphere plants represented different species
from those of Europe, listed several species of Michaux
which he later intended to describe. Among these was “P.
gramineum to be described later as P. folioswm." Indeed,
he did later describe the taxon (Rafinesque, 1811), but it
was spelled as P. foliorum rather than as P. foliosum.
However, the name of P. foliosum [foliosus] was validly
and effectively published in 1808 when he made direct ref-

616 Rhodora [Vol. 76

erence to the published description of P. gramineum of
Michaux.

Pursh (1814) applied a superfluous name — Potamogeton
pauciflorus — to Michaux’s specimen. Other botanists
either overlooked or neglected Rafinesque’s name and ac-
cepted instead the one of Pursh. Tuckerman (1849) recog-
nized that two forms — a large-leaved form and a small-
leaved form — of P. pauciflorus Pursh [foliosus] existed.
He divided the group into two species, applying P. niaga-
rensis Tuckerm. to the larger-leaved individuals and retain-
ing the name P. pauciflorus for the smaller-leaved ones.
After noticing considerable morphological integration, Fer-
nald (1932) chose to consider the two taxa at the varietal
rank. After having examined Michaux's specimen at Paris,
Fernald determined the specimen to be the large-leaved
entity. Thus, P. foliosus and P. pauciflorus actually applied

-
Zz
c
EF]
Lal
m
z
Q
P Dos e
^9
a e oœ
z oe oo 9 © 0e L
= 9 9 M ao 9 e
C Oo O 9 o 9
9 99 c

+
+
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4
- Tt
P

LEAF LENGTH (CM)

Fig. 7. Potamogeton foliosus var. foliosus. Scatter diagram com-
paring leaf length, fruit length, and number of nerves per leaf.
1 nerve, open circle; 3 nerves, half-closed circle; 5 nerves, dot.

1974] Potamogeton — Haynes 617

to this growth form. Therefore, the smaller-leaved plants
were unnamed. Fernald then named the smaller-leaved
specimens P. foliosus var. macellus Fern. and called the
larger river form var. genuinus [now correctly as var.
foliosus]. I have examined thousands of specimens in the
field and herbarium. Many of these have been plotted on
scatter diagrams (see Fig. 7). Although only three char-

Fig. 8. Map of North America showing documented distribution
of Potamogeton foliosus var. foliosus.

618 Rhodora [Vol. 76

acters are used here, other characters were plotted and
similar patterns developed. Data gathered from these
studies indicate that a continuous gradient exists from
ones with the largest leaves to those with the smallest
leaves. The two groups do not appear to be separated into
distinct geographical areas, with the intermediates in areas
of overlap. Instead, these data indicate that the larger-
leaved individuals occur in rapidly flowing waters, the inter-
mediates occur in slow moving streams, and the smaller-
leaved plants occur in lakes or ponds. Sculthorpe (1967)
indicates that the size and shape of the leaves of many
species of Potamogeton vary greatly with different rates of
water flow. This is probably the situation with P. foliosus.
I am, thus, considering var. macellus to be an ecological
growth form of P. foliosus, and therefore, am not formally
recognizing it.

REPRESENTATIVE SPECIMENS

CANADA: Alberta: Wood Buffalo Park, Raup 1562 (CAN, NY,
uc). British Columbia: Revelstoke, Macoun 3037 (CAN, NY).
Manitoba: Little Saskatchewan, Rapid City, Macoun 16482 (CAN,
GH). Northwest Territories: small lake, 38.3 mi. WNW of Yellow-
knife, Thieret & Reich 8356 (F, os). Nova Scotia: KINGS CO.: Canard
River, Smith et al. 11446 (CAN, DAO). Ontario: FRONTENAC CO.:
Kingston, Smith Falls, Fowler (CGE, MO, US). Prince Edward Island:
QUEENS CO.: ponds, Southport, Fernald & St. John 6782 (CAN, K,
NY, US). Quebec: NEW-RICHMOND CO.: Bonaventure River, Marie-
Victorin et al. 33839 (CAN, F, MU, NY, PH). Saskatchewan: MOOSE
JAW DIST.: Mortlach, Hudson 1143 (Dao). GUATEMALA: Dept.
Esquintla: mountain stream, Finca San Filipe, Muenscher 12074
(cH, os). JAMAICA: Portland: Rio Grande, Maxon & Killip 265
GLL). MEXICO: Baja California: Rancho San Jacinto, 45 mi. S.
of Eusenada, Wiggins & Demaree 4749 (F, MICH, MU, NY, US);
stream, Santiago, Wiggins 5662 (MICH, NY, UC, US). Chihuahua: DEPT.
DE GUERRERO: stream near Minaca, Shreve 7993 (F, GH). Guanajuato:
DEPT. ATOTONILCO: just S of village of Santuario, between San Miguel
Allenda and Dolores Hidalgo, Ogden 51149 (os). Michoacan: Rio
Duero at Zamora, Ogden 51183 (os). Puebla: DIST. DE TEPEACA:
San Hipalito, vicinity of Puebla, Arsene 2860 (ILL, NY, us). Sonora:
Fronteras, Hartman 992 (GH, NY). Veracruz: Orizaba, Engenio,
Rio Blenio, Muller 1330 (NY). UNITED STATES: Alabama: JACK-
SON co.: Paint Rock River, Larkin Fork, Harper 2998 (GH, MO, NY,

1974] Potamogeton — Haynes 619

PH, US). Alaska: 2 mi. SW of College, small pond, Smith 1911 (CAN).
Arizona: Colorado Havasupai Canyon, Clover 5218a (MICH, US).
Arkansas: FULTON C0.: Mammoth Springs, Spring River, Thomas
& Smith 17094 (os). California: Los ANGELES CO.: Lancaster, Elmer
3504 (GH, K, NY, UC, US). SISKIYOU CO.: near Mt. Shasta, Abrams
Lake, Heller 13908 (F, ILL, MO, NY, PH, UC, US). Colorado: DENVER
co.: Denver, Platte River, Jones 600 (F, NY, US). Connecticut:
LITCHFIELD CO.: Litchfield-Morris Wildlife Sanctuary, Bantam Lake,
Dwyer 2150 (NY). Delaware: NEW CASTLE CO.: near Wilmington,
Brandywine Creek, Commons (Pn). District of Columbia: Wash-
ington, Ward (GH, UPS). Georgia: CHATTOOGA CO.: 2.8 mi, E 30^N
of Trion, small pond below Greeson Springs, Duncan & Harris 12842
(GH, LAF, MICH, US). Idaho: BOISE co.: 20 mi. S of Idaho City,
marsh, Hitchcock & Muhlick 9947 (Ny, vc). Illinois: COOK CO.:
Englewood, ditches, Hill (F, GH, ILL, US). ST. CLAIR CO.; East St.
Louis, Eggert (CAN, F, ILL, K, MICH, NY, PH, OS, US). STARK CO.:
near Wady Petra, prairie pond, Chase 1148 (ILL, MO, NY, PH).
Indiana: LAKE CO.: Wolf Lake, West Bay, Chase 1461 (F, ILL, MO,
PH, US). Iowa: CLAY CO.: Lake Twp, Dan Greene’s slough, N end,
Hayden 820 (MO, NY, PH, US). EMMET CO.: Estherville, running
water, Cratty (GH, ILL, MO, NY, US). PALO ALTO CO.: Highland Twp,
2 mi. S of Ruthven, Virgin Lake, Hayden 10125 (Mo, NY, PH, US).
Kansas: RILEY CO.: ponds, Pond 1100 (MICH, MO, NY, US). Kentucky:
FAYETTE CO.: near Lexington, brook, Peter (K, MICH, NY, PH).
Maine: PISCATAQUIS CO.: Dover, river margin, F'ernald 479 (Mo, Us).
Massachusetts: MIDDLESEX CO.: Cambridge, Fresh Pond, Morong (F,
MO). Michigan: CHEBOYGAN CO.: Douglas Lake, Marl Bay. Haynes
3923 (os). EMMET CO.: small arm of Carp Lake, Haynes 3374 (GH,
os, UC, US), Haynes 3705 (ILL, os). Haynes 3745 (os), Haynes 3888
(os). Minnesota: CLEARWATER CO.: Bohal Lake, Grant & Oosting
3228 (NY, UC, US). Missouri: BARRY CO: Eagle Rock, river, Bush
511 (CGE, GH, MO, US). BUTLER CO.: Poplar Bluff, Eggert (CAN, F, K,
MO, OS, US). OREGON CO.: Thomasville, spring pond, Steyermark &
Palmer 41701 (MO, NY. PH, US). TEXAS CO.: SW of Plato, Roubidoux
Creek, Steyermark 25006 (DAO, F, NY. os). Montana: MISSOULA CO.:
ca. 6 mi. E of Lolo Hot Springs, small slough, Hitchcock 23978 (DAO,
ILL, NY, US). Nebraska: LINCOLN co.: North Platte, slough, Shear
4445 (GH, NY, US). New Hampshire: GRAFTON CO.: Hanover, Count
River, Jesup (MO). New Jersey: MORRIS CO.: Oak Ridge, running
water, Mackenzie 3736 (MO. Ny, US). New Mexico: LINCOLN CO.:
Bonita River, ca. 616 mi. NW of Alto, Haynes 2923 (LAF). New
York: COLUMBIA CO.: 1% mi. SSW of West Copake, Lower Rhoda
Pond, Haynes 3802 (os). DUTCHESS CO.: 14% mi. SW of Pine Plains,
Stissing Pond, Haynes 3804 (0s). ESSEX CO.: near Ticonderoga, outlet
of Lake George, Haynes 3333 (os), Haynes 3336 (0S). NIAGARA CO.:

620 Rhodora [Vol. 76

Niagara Falls, mill race near Cataract House, Hill 26-1887 (ILL,
MICH, MO). PUTNAM CO.: Carmel, Glendia Lake, Muenscher & Curtis
5447 (MO, NY, PH, UC, US). WARREN CO.: Lake Luzerne boat launch,
Haynes 3814 (os). Nevada: ELKO CO.: Copper Mountains, 3 mi. S
of Coon Creek Pass, Maguire & Holmgren 22411 (CAN, NY, US).
North Carolina: CHEROKEE C0.: N of Marble, limestone quarry,
Radford 4812 (NY). North Dakota: CASS CO.: Fargo, ditch, Stevens
1376 (CAN, UC, US). Ohio: AUGLAIZE CO.: Lake St. Marys Fish
Hatchery, Haynes 3427 (GH, MICH, OS, us); 2 mi. SE of Wapakoneta,
borrow pit, Haynes 3420 (0s, US). CHAMPAIGN CO.: Kaiser Lake
State Park, Haynes 3435 (MICH, OS, UC, US). ERIE co.: 4 mi. NW of
Castalia, Haynes 3247 (MICH). LOGAN CO.: 1.9 mi. due ESE of
Russells Point, Haynes 3415 (MICH, OS, PH, us). OTTAWA CO.: Lake
Erie, North Bass Island, Fox’s Marsh, Haynes 3145 (08). PORTAGE
co.: 5% mi. NW of Ravenna, borrow pit, Haynes 3452 (GH, MICH,
MO, OS, US). Oklahoma: OTTAWA CO.: Hattenville, overflow pond,
Stevens 2481 (GH, ILL, K, MO, NY, US). WOODS CO.: Waynoka pond,
Stevens 1768 (GH, ILL, K, MO). Oregon: pond on the Willamette
River, Hall 492 (F, GH, ILL, MO, NY, us). Pennsylvania: BERKS CO.:
1 mi. SE of Pine Forge, Manatawny Creek, Wilkens 5619 (GH, PH).
South Carolina: GEORGETOWN CO.: Georgetown, Hobeaw Plantation,
Alexander 148 (Us). South Dakota: PENNINGTON CO.: Sheridan
Lake, Porter 6593 (DAO, UPS). Tennessee: FRANKLIN CO.: Sher-
wood, Eggert (GH, MO, US). Texas: JEFF DAVIS CO.: Limpia Canyon,
Tracy 288 (GH, NY). Utah: CACHE CO.: Bear River Range, E side
of Tony Grove Lake, Maguire & Snell 16048 (DAO, ILL, MICH, NY).
GRAND CO.: Hill Creek near Weaver Reservoir, Holmgren et al, 2358
(DAO, ILL, NY, UPS). Vermont: ORLEANS CO.: Ca. 1 mi. W of Crafts-
bury Common, backwater of Black River, Haynes 3839 (os); 1 mi.
S of Craftsbury Common, backwater of Black River, Haynes 3840
(os). WINDSOR CO.: Plymouth, Eglleston (F, MO, PH, us). Virginia:
BOTETOURT CO.: Cloverdale, Tinker Creek, Wood 6183 (GH). Wash-
ington: GRANT CO.: Pond SW of Moses Lake, St. John et al. 4960
(Mo, UC). West Virginia: GREENBRIAR CO.: Dunlap Creek, Berkley
1194 (mo). Wisconsin: WAUSHARA CO.: Poy Sippi, Pine River, Hill
(CAN, F, GH, ILL). Wyoming: LARAMIE CO.: Laramie Mountains,
Middle Fork of Crow Creek, Porter 6320 (DAO, MO, NY, UPS).

5b. Potamogeton foliosus var. fibrillosus (Fern. Haynes
& Reveal, Rhodora 75: 76. 1973.

Potamogeton fibrillosus Fern. Mem. Amer. Acad. Arts
17: 51. 1932. TYPE: W. C. Cusick 2598, in warm spring,
margin of Harney Valley, “P” Ranch, [Harney Co.] Ore-
gon, (holotype, GH!; isotypes, F! K!, MO!, NY!, uc!, us!).

1974] Potamogeton — Haynes 621

Stems 20-75 cm long, 0.3-1.2 mm diam. Leaves (1-) 3-
nerved, 2.1-5.5 mm long, 0.5-1.7 mm wide; apex acute apicu-
late; glands usually present, black to gold, to 0.5 mm diam;
lacunae absent; lateral nerves joining midrib 0.6-0.8 mm
below apex. Stipules brown, rarely white, fibrous, shred-
ding at tip, often convolute, 0.4-12.1 em long, 0.5-1.2 mm
diam. Winter buds lateral or terminal, 1.2-2.5 cm long,
0.8.2.0 mm wide. Peduncles clavate, recurved, 6.5-9.2
(-22.0) mm long, 0.4-1.0 mm diam. Spike capitate to cylin-
dric, 1.7-5.5 mm long, 1.0-4.0 mm diam; verticels 1-2, when
2, then 0.6-0.8 mm apart. Perianth segments 0.5-0.7 mm
long, 1.1-1.2 mm wide. Fruit pale-green, 1.4-1.7 mm long,
1.1-1.2 mm wide; keel to 0.2 mm high; beak ca. 0.2 mm long,
0.1-0.2 mm wide. Chromosome number unknown.

Distribution: In waters, often warm, of shallow lakes,
springs, streams and rivers, from southwestern Washing-
ton southward through southeastern Oregon, hence east-
ward to southeastern Wyoming. Fruiting from mid-June
to mid-September. Figure 9.

Illustrations: Fernald (1932, pl. 5; 28. Fig. 5; 32, Fig. 5).

In 1932, Fernald proposed Potamogeton fibrillosus based
on specimens collected in Wyoming, Idaho, Oregon and
Washington. He noted that the new species closely resem-
bled P. foliosus Raf. in its foliage, peduncles and small
dorsally keeled fruits. Fernald said that P. fibrillosus dif-
fered from P. foliosus by its stipules which disintegrate
into “rope-like” fibers and are thus open or convolute, and
by having fruits with a less developed keel and more nearly
median beak. Fernald (1932, p. 52) stated his reservations
in assigning specific rank to this new entity: “The plant
will doubtless be found to have a broader range, when it
may prove to be a marked geographic variety of the wide-
spread P. foliosus."

Several collections, especially from Yellowstone National

Park of Wyoming and from southeastern Oregon, are dis-
tinct from Potamogeton foliosus. In general, these speci-

622 Rhodora [Vol. 76

mens differ from P. foliosus not only in having the stipules
disintegrating into fibers and fruits with poorly developed
keels, but also in having cylindric, interrupted inflorescences
and leaves with basal glands — the last two characters not
mentioned by Fernald, but are present even on the holotype.

Other collections from the Yellowstone region are not so
distinct. One such collection (Haynes 3849) has fruits,
including the keel, which are identical with those of Pota-
mogeton foliosus, but the inflorescence is capitate and the
stipules only rarely disintegrate into fibers. Yet, basal
glands are present. Flowering specimens from Albany
County, Wyoming, taken by C. L. Porter (3473) are similar
to P. fibrillosus in having basal glands and short, clavate
peduncles. However, in this collection, the stipules only
rarely disintegrate into fibers. Flowering specimens ob-
tained by Maguire (21578) in northern Utah commonly
produce fibers when the stipules decay, but basal glands
are rarely found.

Hitchcock (1969) regarded Potamogeton fibrillosus as a
distinct species, but conceded its close relationship with P.
foliosus. Porter (1963) reduced P. fibrillosus to synonymy,
regarding it only as a local form of P. foliosus produced by
the rather warm-water conditions of “geyser formations
and hot springs" in the Yellowstone area. Porter stated
(19683, p. 9) “The warm water hastens disintegrations of
the sheaths by bacterial action." 'This is not always the
case, however, in northern Utah or southeastern Oregon
where the water is often cool. Here the stipules still form
fibers. In the same general region, P. foliosus is known to
occur in similar environmental conditions and at approxi-
mately the same degree of development. Therefore, the
possibility that mere bacterial activity (or some other type
of mechanical breakdown) being the only cause of the
fibrous stipules seems unlikely. For this reason and that
the extremes can be distinguished quite easily, Haynes and
Reveal (1973) thought that P. fibrillosus should be taxo-
nomically recognized. However, since many intermediate

1974] Potamogeton — Haynes 623

forms can be found, they could not justify the specific rank
and therefore ranked the taxon at the varietal level.

From var. foliosus, the var. fibrillosus may be distin-
guished by its poorly developed keel, cylindric and inter-
rupted inflorescence, fibrous stipules and glands present at
the base of most leaves.

Fig. 9. Map of northwestern United States showing documented
distribution of Potamogeton foliosus var. fibrillosus.

624 Rhodora [Vol. 76

REPRESENTATIVE SPECIMENS

UNITED STATES: Idaho: BLAINE Co.: Russian John Ranger
Station, Macbride & Payson 3790 (Uc). Oregon: HARNEY CO.: warm
spring, near Burns, Henderson 8867 (GH); irrigating ditch at French-
glen, Thompson 12068 (NY). KLAMATH CO.: N end of Klamath Marsh,
Peck 21954 (UC). MALEUR CO.: Otis Creek, Leiberg 2340 (GH, US).
Utah: CACHE CO.: stream, 3 mi. NE of Logan, Maguire 16685 (CAN,
MO, NY); stream, 2 mi. NW of Logan, Maguire 21578 (DAO, F, GH,
MICH, MO, NY, US). Washington: PIERCE CO.: outlet, Lake Spanaway,
near Tacoma, Thompson 9657 (GH, NY); unknown locality, Wash-
ington Territory, Brandegee 1127 (GH, UC). Wyoming: ALBANY CO.:
N. fork of Pole Creek, Porter 3474 (Ny, US). PARK CO.: Firehole
River, Jepson 2540 (GH); Firehole Canyon, Firehole River, Haynes
3848 (ILL, OS, PH, UC, US); Midway Geyser Basin, Firehole River,
Haynes 3850 (GH, MICH, MO, NY, 0S); Upper Geyser Basin, Richard-
son (GH); Upper Nez Perce Creek, Haynes 3849 (MICH, os).

6. Potamogeton clystocarpus Fern., Mem. Amer. Acad. Arts
17: 79. 1932. TYPE: J. A. Moore & J. A. Steyermark 3088,
pool in rock, Little Aguja Canyon, Davis Mts., 1575 ft. Jeff
Davis Co. Texas, (holotype, GH!; isotvpes, MICH!, MO!, NY!,
PH!, ua,

Stem light green to brown, terete to slightly compressed,
rarely ridged, to 57 cm long, 0.5-0.7 mm diam. Leaves
green, 3 (-5)-nerved, 3.2-7.8 cm long, 0.7-1.7 mm wide; apex
acute; glands usually present, white to gold, 0.2-0.3 mm
diam; lacunae to 4 rows each side of midrib, rarely absent;
lateral nerves joining midrib 0.2-0.4 mm below apex. Sti-
pules brown, delicate, not shredding at tip, usually convo-
lute, to 6.2 mm long, 0.5-0.8 mm diam. Winter buds un-
known. Peduncles cylindrical, axillary to terminal, erect,
3.2-4.8 cm long, 0.3-0.5 mm diam. Spike capitate to cylin-
dric, 5.5-7.5 mm long, 3.0-5.7 mm diam; verticels 3, 1.5-1.7
mm apart. Perianth segments 1.7-2.0 mm long, 1.5-1.8 mm
wide. Fruit brown to yellow-green, dorsally and laterally
keeled, 2.0-2.2 mm long, 1.7-1.8 mm wide; keels ridged,
without undulations, to 0.2 mm high; beak central, 0.5-0.6
mm long, 0.2-0.4 mm diam; sides depressed, with 1-3 tuber-
cles near base; wall texture rough. Chromosome number
unknown.

1974] Potamogeton — Haynes 625

Distribution: Known only from the type locality. Fruit-
ing from early May to mid-June. Fig. 10.

Illustrations: Fernald (1932, plate 15.30, fig. 5) ; Ogden
(1966, plate 51).

Potamogeton clystocarpus was named by Fernald (1932),
based on the “gibbous-tuberculate-based fruits.” He sug-
gested the fruits were “so similar to those of the western
Eurasian and African P. trichoides C. & S. in having basal
bosses that they might easily pass as fruits of that species.
P. trichoides, however, as its name implied, has very bristle-
tipped leaves . . . P. clystocarpus cannot be referred to it."
No other North American pondweed, to my knowledge, has
fruits with the basal “bosses.” In this character, P. clysto-
carpus is quite distinct. But, as with other Pusilli, it is
almost impossible to identify with sterile material!

Fig. 10. Map of southwestern North America showing docu-
mented distribution of Potamogeton clystocarpus.

626 Rhodora [Vol. 76

REPRESENTATIVE SPECIMENS

UNITED STATES: Texas: JEFF DAVIS CO0.: Little Aguja Canyon,
Davis Mts, Palmer 34526 (Mo, NY, PH), Correll & Ogden 25070
(NYS, UC).

7. Potamogeton pusillus L., Species Plantarum 1: 127. 1753.

Stems pale green to olive, terete to slightly compressed,
smooth to slightly ridged, 18-150 cm long, 0.1-0.7 mm diam.
Leaves pale green to olive, rarely rufescent, delicate to
coarse, 1-3 (-5)-nerved, 0.9-6.5 cm long, 0.2-2.5 mm wide;
apex subulate to obtuse; glands present or absent, green,
gold, brown, or rarely white, to 0.5 mm diam; lacunae
absent or to 5 rows each side of midrib; lateral nerves,
when present, joining midrib 0.1-1.0 (-2.0) mm below apex.
Stipules brown to green or white, delicate, rarely appearing
fibrous, not shredding at tip, connate or convolute, 3.1-9.2
mm long, 0.2-0.7 mm diam. Winter buds common, lateral
or terminal, 0.9-3.2 em long, 0.3-1.8 mm wide; inner leaves
rolled into a hardened fusiform structure; outer leaves 1-3
per side, subulate to obtuse, without corrugations at base.
Peduncles filiform to slightly clavate, axillary or terminal,
mostly erect, rarely recurved, 0.5-6.2 (-6.6) cm long, 0.2-
0.7 mm diam. Spikes capitate to cylindric, 1.5-10.1 mm
long, 0.7-5.2 mm diam; verticels 1-3 (-4), 1.2-4.7 mm apart.
Perianth segments 0.7-1.7 mm long, 0.5-1.2 mm wide. Fruit
green to brown, without keels, 1.5-2.2 mm long, 1.2-1.6 mm
wide; beak forward or central, 0.1-0.6 mm high, 0.2-0.5 mm
diam; sides rounded; wall texture smooth or rough.

Potamogeton pusillus has been interpreted variously by
botanists. Bennett (1890-1894; 1900-1904) and Morong
(1893), for example, considered the taxon to consist of two
species, one being divided into several varieties. Hagstrom
(1916), on the other hand, divided the taxon into several
poorly defined species, with two of these — P. panormitanus
[var. pusillus] and P. pusillus [var. tenuissimus] — being
split into many infraspecific taxa. These two species were
separated by the connate versus convolute character of the
stipule. Fernald (1932) took the intermediate position and

1974] Potamogeton — Haynes 627

recognized three species — P. panormitanus [var. pusillus],
P. gemmiparus [var. gemmiparus], and P. pusillus [var.
tenuissimus]. He divided P. panormitanus into two vari-
eties and P. pusillus into six varieties. These varieties were
based on the size of the leaf, apex shape, and number of
lacunae bordering the midrib. The latter concept of the
taxon was accepted until Dandy and Taylor (1938), after
studying the type specimen of P. pusillus in the Linnaean
herbarium, decided that the name P. pusillus actually should
be applied to the taxon which had been passing as P. panor-
mitanus. For the taxon which had been passing as P. pusil-
lus, the next available name in the specific rank was P.
berchtoldii Fieber. Fernald (1940) finally made the com-
binations of various varieties to conform with the revised
nomenclature. This is the concept that appeared in the
eighth edition of Gray’s Manual (Fernald, 1950). Gleason
and Cronquist (1963), however, did not accept P. pusillus
and P. berchtoldii as two distinct species. They, instead,
suggested that although this character may be good for
distinguishing certain species, there was no reason to as-
sume that both states of the stipule and gradations be-
tween could not oceur within one species. As I indicated
earlier, à single plant may produce two sets of leaves
during one growing season. The first set may have obtuse
leaves with 4-5 rows of lacunae; the second, instead, would
probably have acute leaves with 1-3 rows of lacunae. In
the early summer, this plant would resemble var. mucro-
natus [of Fernald, 1932] and in the late summer, it would
resemble var. tenuissimus [of Fernald, 1932]. For this
reason and because of the ecological variability as explained
by Seulthorpe (1967), I have decided to recognize only
three taxa.

Fernald (1950) suggested that useful characters in dis-
tinguishing Potamogeton pusillus from P. berchtoldii were
length of inflorescence, branching of the plant, and whether
the fruit is wider at, above, or below the middle. Voss
(1972) had noticed that fruiting specimens with connate
stipules [P. pusillus var. pusillus] from Michigan often

628 Rhodora [Vol. 76

lacked glands. Ogden (personal communication) suggested
that the presence or absence of lacunae along the midrib
often is a reliable character for distinguishing the species.
Eighty specimens were examined for the characters men-
tioned above and other features. Some correlation, although
slight, was noticed between the stipule character, the diam-
eter of the glands, and the number of lacunae bordering
the midrib. However, for the majority of the characters
measured, little or no correlation was observed. Regardless
of the inconsistency of the stipule character, the shape of
the inflorescence and of the fruit, and the length of peduncle
seem to be associated. This is not to say that intermediates
do not occur, because they do! In their extremes, the two
entities can be separated by the length of the inflorescence,
the shape of the fruit, the length of the peduncle, the number
of laeunae bordering the midrib, the presence and size of
nodal glands, and, finally, the connation of the stipules. No
one character can definitely be depended upon, but by using
a combination of as many of the above mentioned cnarac-
ters as are present on the plant, one can place the specimen
into the correct taxon 70-80% of the time. The remaining
20-30%, however, are almost impossible to identify without
depending upon one character taxonomy. Also, var. tenuis-
simus has a more northerly distribution, e.g. into Alaska,
and var. pusillus has a more southerly distribution, e.g.
into Mexico. Because of the morphological integradation
and the slight range differences, I have decided to place the
two taxa at the varietal level.

When in fruit, Potamogeton pusillus should not be con-
fused with any other pusilloid. Its fruit, lacking a keel, a
wing, or basal warts, and its delicate stipules separate it
from all other species. However, in the sterile state, identi-
fieation is not so easy. Vegetatively, it closely resembles
P. foliosus, P. clystocarpus, and some individuals of P. hillit.
I can give no morphological characters which serve to sep-
arate P. pusillus from P. clystocarpus if fruits are lacking.
If the specimen is from areas other than western Texas, one
can assume the specimen is not P. clystocarpus, although

1974] Potamogeton — Haynes 629

this may not be a valid assumption. The stipules of P.
pusillus, usually lacking slightly fibrous veins, often serve
to distinguish the species from P. hillii and P. foliosus.
However, the stipules of the two latter species may, on
occasions, appear to lack the fibrous veins. In this instance,
the species are impossible to separate. Therefore, one
should always make an effort to collect fertile material!

Ta. Potamogeton pusillus var. pusillus

Potamogeton pusillus L., Species Plantarum, 128, 1753.
P. pusillus var. typicus Fern., Mem. Amer. Acad. Arts 17:
81. 1932. TYPE: Europe. (holotype, LINN; photograph of
holotype, os!).

Potamogeton panormitanus Ant. Biv. Bern. in And. Biv.
Bern. Nuove piante inedite del barone Ant. Bivona Bernardi
6. 1838. (publication not seen). P. pusillus var. panormi-
tanus (Ant. Biv. Bern. in And. Biv. Bern) Morong, Mem.
Torrey Bot. Club 3: 46. 1893. TYPE: (location unknown).

Potamogeton panormitanus var. major Ant. Biv. Bern.
in And. Biv. Bern, Nuove piante inedite del barone Ant.
Bivona Bernardi 6. 1838. (publication not seen). TYPE:
(location unknown).

Potamogeton panormitanus var. minor Ant, Biv. Bern. in
And. Biv. Bern. Nuove piante inedite del barone Ant.
Bivona Bernardi 6. 1838. (publication not seen). P. pusil-
lus var. minor (Ant. Biv. Bern. in And. Biv. Bern.) Fern.
& Schubert, Rhodora 50: 154. 1948. TYPE: (location un-
known).

Potamogeton pusillus var. vulgaris Fries, Novitiae Florae
Suecicae 2nd ed. 49. 1828. TYPE: (type not located), ap-
plication of the name is from the illustration of Potamoge-
ton pusillus L. In Smith, English Bot. 2: pl. 215. 1794!

Potamogeton pusillus var. vulgaris subvar. interruptus
Robbins in Watson, Bot. U. S. Geol. Expl. Fortieth Parallel
338. 1871. TvPE: S. Watson, 1137, Parley's Park in the
Wasatch, 6,000 ft. [Wasatch Co.], Utah, (holotype, NY!;
isotype, US!).

630 Rhodora [Vol. 76

Stems green to brownish, mostly terete, slightly ridged,
18-150 em long, 0.2-0.7 mm diam. Leaves pale green, rarely
olive, delicate, 1-3-nerved, 1.4-6.5 cm long, 0.5-1.9 mm wide;

Fig. 11. Map of North America showing documented distribu-
tion of Potamogeton pusillus var. pusillus.

1974] Potamogeton — Haynes 631

apex acute or rarely apiculate, rarely with a bristle; glands
often present, green, gold, brown, or rarely white, to 0.3
mm diam; lacunae absent or to 2 rows each side of midrib;
lateral nerves joining midrib 0.1-0.7 mm below the apex.
Stipules greenish to brown, delicate, rarely fibrous, usually
connate, 2.2-9.2 mm long, 0.2-0.7 mm diam. Winter buds
uncommon, lateral or terminal, 0.9-1.5 em long, 0.3-1.8 mm
wide; outer leaves 1-2 per side, acute to apiculate. Pedun-
cles cylindric, axillary, to lower or upper leaves, rarely re-
curved, 1.0-6.2 em long, 0.2-0.7 mm diam. Spikes cylindric,
2.5-10.1 mm long, 1.2-4.7 mm diam; verticels 2 or 3 (-4).
usually interrupted, 1.4-4.5 mm apart. Perianth segments
1.0-1.5 mm long, 0.7-1.2 mm wide. Fruit green to brown,
usually wider above the middle, 1.5-2.2 mm long, 1.2-1.5 mm
wide; keel absent; beak forward, rarely central, 0.1-0.6 mm
high, 0.2-0.5 mm diam; sides rounded, usually centrally
depressed; wall texture smooth. Chromosome number 2n —
26 (Palmgren, 1939).

Distribution: From central British Columbia to Nova
Scotia, south to northern Guatemala and western Florida.
Fruiting from early May to late September. Fig. 11.

Illustrations: Fernald (1932, pl. 9; 10; 29, fig. 7, 8; 33,
fig. 4, 5; 39, fig. 10).

REPRESENTATIVE SPECIMENS

CANADA: Alberta: Wood Buffalo Park, Raup 1568 (CAN, GH,
NY, UC, US). British Columbia: pond near Similkameen River, Macoun
70317 (F, GH, NY). Manitoba: Desford, pothole, Love & Love 6092
(DAO, ILL, US). New Brunswick: KINGS CO.: Westfield, Ingleside,
cove in St. John River, Fernald 1618 (GH, NY). Northwest Territories:
MACKENZIE DIST.: Norman Wells, MacKenzie River, Cody & Gutte-
ridge 1631 (F, NY, US). Nova Scotia: CAPE BRETON CO.: Fort Louis-
bourg, Smith et al. 8281 (CAN, DAO). Ontario: OTTAWA CO.: Ottawa,
Rideau Canal, Fletcher (MO, NY, PH). Prince Edward Island: Tig-
nish, pools and ponds, Fernald et al. 6776 (CAN, K, NY). Quebec:
Magdalen Islands, Grindstone Island, Fernald et al. 6780 (GH, K,
NY, US), Fernald et al. 6781 (GH, NY, PH, US). Saskatchewan: MAPLE
CREEK DIST.: 14.7 mi. N of Govenlock, middle Creek, Bird 1022
(Dao). GUATEMALA: Quiche Dept.: Quiche, pond, Muensclher

12085 (uc). MEXICO: Baia California: La Encantada, Sierra San
Pedro Martir, Wiggins & Demaree 4906 (GH, MU, vs). Hidalgo:

632 Rhodora [Vol. 76

DIST. MOLANGO: Molango, Lake Atexca, Moore 3470 (uc). Morelos:
Zempoala National Park, Lake Zempoala, Moore 3435 (GH, US).
Puebla: vicinity of Puebla, Laguna de San Baltasar, Arsene 1328
(US). San Louis Potosi: San Louis Potosi, Schaffner 533 (K, NY).
Tamaulipas: vicinity of San Jose, Sierre de San Carlos, Bartlett
10397 (F, MICH, US). Veracruz: Los Molinos, Perote, pond, Balls
& Gourlay (K, MICH, UC, Us). UNITED STATES: Alabama: 7-8
mi. NE of Mobile, Chuckfey Bay, Hotchkiss & Ekvall 3902 (us).
Arizona: GILA CO.: Young's Ranch, Lemmon (K, MO, NY, UC, US).
Arkansas: MISSISSIPPI C0.: S. of Hornersville, Missouri, Metcalf
644 (GH, NY, US). California: HUMBOLDT CO.: Eel River, near Fern-
bridge, Harris 808 (CAN, DAO, F, GH, ILL, MICH, MO, NY, PH, UPS, US).
Colorado: ROUTT CO.: ponds between Hayden and Craig, Porter 6563
(CAN, DAO, NY, UC, UPS). TELLER CO.: base of Pike's Peak, Lake
Osborne, Parry (GH, MO, NY). Connecticut: MIDDLESEX CO.: Grove
Beach, creek, Enquist 366 (NY). Florida: JACKSON co.: N of
Sneads, Lake Seminole, Adams 445 (us). Idaho: CUSTER CO.: 15 mi.
N of Challis, ditch, Hitchcock et al. 23839 (DAO, ILL, NY, UC). Illinois:
COOK CO.: South Chicago, ditches, Hill 89-1875 (F, ILL). Indiana:
CASS CO.: Lake Cicott, Deam 49306 (GH). Iowa: CLAY CO.: Lake
Twp, Round Lake, Hayden 10120 (GH, MO, NY, PH, UC, US). Kansas:
DOUGLAS CO.: 14% mi. E and % mi. N of Lawrence, ponds, Richards
3079 (NY). Louisiana: CAMERON PAR.: Sabine National Wildlife
Refuge, Valentine (os). Maine: AROOSTOOK CO.: Oxbow, Aroostook
River, Oaden 2661 (os). Maryland: CECIL co.: Blairs Shore, Elk
River, Long 57093 (pu). Massachusetts: BERKSHIRE C0.: New Marl-
boro, Lake Buel, Churchill (Gn. MO). Michigan: EMMET CO.: small
arm of Carp Lake, Haynes 3370 (CAN, FSU, GH, ILL, MICH, MO, MSU,
NY. OS, PH, UC, US, WVA), Haynes 3706 (ILL. os). Minnesota: sT.
LOUIS CO.: Armstrong, near Ely, Jones 18543 (ILL, NY. US). Missouri:
LACLEDE CO.: Gasconade River, between Falcon and Nelio, Steyer-
mark 13913 (GH, MO). Montana: GALLATIN CO.: ca. 12 mi. NW of
West Yellowstone, Hegben Lake. Haynes 3841 (os). LAKE CO.: 0.2
mi. SE of Nine-Pipes Reservoir, pot-hole, Hitchcock et al. 23980
(CAN, F, ILL, NY, UC). Nebraska: CHERRY CO.: 30 mi. S of Valentine,
small pond, Porter 6445 (DAO, NY, UPS). Nevada: STOREY CO.: vicinity
of Reno, Sparks, irrigation ditch, Hitchcock 443 (GH, US). New
Jersey: SUSSEX CO.: Catfish Pond, Griscom 9750 (GH). New Mexico:
SAN MIGUEL CO.: Montezuma, Peterson Reservoir, Drouet & Richards
3308 (F, GH). New York: ST. LAWRENCE CO.: Lisbon, Sucker Brook,
Phelps 269 (CAN, GH, NY, US). North Carolina: DARE CO.: Currituck
Sound, inlet S of Duck, Radford 5758 (pAo, Uc). North Dakota:
BURLEIGH CO.: 7 mi. E of Bismark, slough, Metcalf 356 (US). Ohio:
PORTAGE CO.: ca. 54% mi. NW of Ravenna, small pond, Haynes 3450
(os). Oklahoma: COMANCHE co.: Wichita Mountains Wildlife

1974] Potamogeton — Haynes 633

Refuge, Lake Quanah Parker, McMurry 1219 (NY). Oregon: WHEELER
co.: 15 mi. E of Mitchell, roadside pond, Hitchcock et al. 23779
(CAN, DAO, F, ILL, NY, UC). Pennsylvania: BERKS CO.: 1 mi. NW of
Moselem Springs, Brwmbach 2841 (PH). South Dakota: CODINGTON
co.: outlet of Lake Kampeska, Over 17149 (US). Tennessee: OBION
co.: Walnut Log, Sharp 7933 (os). Texas: TRAVIS co.: Colorado
River, ca. 5 mi. below Austin, Barkley 13313 (F, GH, Mo, PH, UC).
WALLER CO.: Hempstead, Hall 620 (F, K, MO, NY, US). Utah: CACHE
co.: 1 mi. W of Logan, marsh, Maguire 13883 (CAN, GH, DAO).
Vermont: WINDSOR CO.: Windsor, mill pond, Hellquist 1818 (Boston
State College). Virginia: FAIRFAX co.: Dyke, Metcalf & Sperry
1688 (GH). Washington: GRAYS HARBOR CO.: slough near ocean
beach, Otis 2116 (GH, NY, PH, UC). Wisconsin: LA CROSSE CO.: Lake
Onalaska, Hartley 1826 (DAO, F, ILL, Us). Wyoming: ALBANY CO.:
Laramie River E of Lookout, Porter 6352 (DAO, MO, NY, UPS).

7b. Potamogeton pusillus var. tenuissimus Mert. & Koch in
Rohling, 857. 1823. TYPE: (location unknown).

Potamogeton tenuissimus (Mert. & Koch in Róhling)
Reichen. Icones Florae Germanica Helveticae 7: 14. 1845.
P. berchtoldii var. tenuissimus (Mert. & Koch in Rohling)
Fern. & Schubert, Rhodora 42: 246. 1940.

Potamogeton berchtoldii Fieber, in Berchtold, Oekon.-
tech. Flora Bohmens, 277. 1838. TYPE: (location unknown).

Potamogeton berchtoldii var. mucronatus Fieber in
Berchtold, Oekon.-tech. Flora Bohmens 277. 1838. P.
pusillus var. mucronatus (Fieber) Graebner in Ascherson
& Graebner, Das Pflanzen. 4(11): 115. 1907. TYPE: (loca-
tion unknown).

Potamogeton berchtoldii var. acuminatus Fieber in
Berchtold, Oekon.-tech. Flora Bohmens 278. 1838. TYPE:
(location unknown).

Potamogeton pusillus var. polyphyllus Morong, Bot. Gaz.
(Crawfordsville) 5: 51. 1880. P. berchtoldii var. polyphyl-
lus (Morong) Fern. Rhodora 42: 246. 1940. TYPE: T.
Morong, oozy pool in South Natick, [Middlesex Co.], Mas-
sachusetts, (holotype, NY!).

Potamogeton pusillus var. elongatus Ar. Benn., in Macoun,
Cat. Canadian Plants 4: 371. 1888. TYPE: J. Macoun 4140,
Spullamasheen River at and above Enderby, British Colum-
bia, Canada, (holotype, K[?]; isotype, GH!). P. pusillus

634 Rhodora [Vol. 76

var. capitatus Ar. Benn., nom. illeg., J. Bot. 39: 201. 1901.
(Superfluous when published).

Potamogeton pusillus var. cuspidatus G. Fisher, Ber.
Bayer. Bot. Ges. 11: 116. 1907. TYPE: (location unknown).

Potamogeton lacunatus Hagstrom, Kongl. Svenska Vet-
enskapsakad. Handl. 55(5): 120. 1916. P. pusillus var.
lacunatus (Hagstrom) Fern. Mem. Amer. Acad. Arts 17:
85. 1932. P. berchtoldii var. lacunatus (Hagstrom) Fern.
Rhodora 42: 246. 1940. TYPE: T. Morong, Ashland, in lacu
Wauhakune, [Middlesex Co.], Massachusetts, (lectotype
here designated, UPS!).

Potamogeton turionifera f. tenuis Hagstrom, Kongl.
Svenska Vetenskapsakad. Handl. 55(5): 91. 1916. TYPE:
J. Macoun, Algonquin Park, Cache Lake, Ontario, Canada,
(lectotype here designated, c !; isolectotypes, C ! CAN I, GH!).

Potamogeton turionifera f. mucronulatus Hagstrom,
Kongl. Svenska Vetenskapsakad. Handl. 55(5): 91. 1916.
TYPE: J. Macoun, Ottawa, Brigham’s Creek, Ontario,
Canada, (holotype, C!: isotypes, C!, CAN], GH!).

Potamogeton pusillus var. colpophilus Fern. Mem. Amer.
Acad. Arts 17: 90. 1932. P. berchtoldii var. colpophilus
(Fern.) Fern. Rhodora 42: 246. 1940. TYPE: J. F. Collins,
M. L. Fernald, & A. S. Pease, brackish pools and dead
waters near the mouth of Dartmouth River, Gaspe Co.,
Quebec, Canada, (holotype, GH!; isotypes, CAN!, NY!, us!).

Stem light green to olive, terete to slightly compressed,
usually smooth, 22-74 cm long, 0.1-0.5 mm diam. Leaves
pale green to olive, rarely rufescent, delicate, 1-3 (-5)-
nerved, 0.9-5.4 cm long, 0.2-2.5 mm wide; apex acute to
obtuse; glands usually present, green to brown or white,
to 0.4 mm diam; lacunae usually present, 1-5 rows each side
of midrib; lateral nerves, when present, joining midrib
0.3-1.0 (-2.0) mm below apex. Stipules brown to green,
rarely white, delicate, mostly convolute, 3.1-5.0 mm long,
0.2-0.5 mm diam. Winter buds common, lateral or terminal,
1.2-3.2 em long, 0.7-1.2 mm wide; outer leaves 1-3 per side,
acute to obtuse. Peduncles cylindrical to slightly clavate,

1974] Potamogeton — Haynes 635

axillary or terminal, rarely recurved, 0.5-4.6 (-6.6) em long,
0.2-0.7 mm diam. Spikes capitate to cylindric, 1.5-7.2 mm
long, 0.7-5.2 mm diam; verticels 1-8, when of 2 or 3, then
these crowded, 1.2-2.5 mm apart. Perianth segments 0.7-
1.0 mm long, 0.5-0.7 mm wide. Fruit green to brown, usu-
ally wider at or below the middle, 1.6-2.1 mm long, 1.2-1.5
mm wide; keels absent; beak usually central, rarely for-
ward, 0.2-0.5 mm long, 0.2-0.4 mm diam; sides usually
rounded, rarely centrally depressed; wall texture smooth.
Chromosome number, n = 13; 2n = 26 (Taylor & Mulligan,
1968).

Distribution: Central Alaska to Labrador, south to Cali-
fornia and northern Florida. Fruiting from early June to
late September. Fig. 12.

Fig. 12 Map of North America showing documented distribu-
of Potamogeton pusillus: var. tenuissimus.

636 Rhodora [Vol. 76

Illustrations: Fernald (1932, pl. 16-20; 30, fig. 6-9; 31,
fig. 1; 35; 39, fig. 9).

REPRESENTATIVE SPECIMENS

CANADA: Alberta: CRAIGMYLE DIST.: Craigmyle, Brinkman 538
(us). British Columbia: QUEEN CHARLOTTE ISLANDS: Graham Island,
2 mi. NW of Tlell, n — 13, Calder & Taylor 35688 (DAO); Moresby
Island, White Swan Lake, 2n — 26, Calder & Taylor 35297 (DAO).
VANCOUVER ISLAND: Lost Lake near Victoria, Macoun (CAN, GH, NY,
us). Labrador: Goose Bay, 53°20'N, 60°09'W, Gillett & Findlay
5910 (DAO, NY). Manitoba: CHURCHILL DIST.: Gillman Lake, Schofield
1524 (DAO, GH). New Brunswick: CHARLOTTE CO.: Grand Manan,
Weatherby & Weatherby 7284 (CAN, GH). Newfoundland: Harry’s
River, near Bay of St. George, Fernald & Wiegand 2453 (CAN, GH,
NY). Northwest Territories: small pool 45.5 mi. WNW of Yellow-
knife, Thieret & Reich 7944 (CAN, F, OS, us). Nova Scotia: Sable
Island, St. John 1125 (CAN, GH, NY, PH, us). Ontario: THUNDER BAY
DIST.: Black Fox Lake, 12 mi. E of Terrace Bay, Haynes 351a
(os); small lake 0.2 mi. NE of Coldwell, Haynes 3773 (ASC, CAN,
FSU, GH, ILL, MICH, MO, MSU, NY, OS, PH, UC, US, WVA). ESSEX CO.:
Fox’s Pond, Haynes 3251 (0s), Roberts 1567 (oS). Prince Edward
Island: QUEENS C0.: Watervale, Erskine (DAO). Quebec: MAGDALEN
ISLANDS: ridges back of the Narrows, Alright Island, Fernald et al.
6778 (CAN, GH, K, NY, US). Saskatchewan: Cornwall Bay, Lake
Atabaska, 59?27'N, 108°27'W, Raup 6633 (CAN, DAO, F, GH, NY).
Yukon: Lewes River, Whitehorse, Porsild & Breitung 10644 (CAN).
UNITED STATES: Alaska: Goldstream Creek, 51 mi. N of Fair-
banks, 65°N, 147?30'W, Porsild & Porsild 115 (CAN, GH, US).
Arkansas: DREW CO.: Taxodium swamp, Tillar, Demaree 17394 (F,
MO, NY, UC). Arizona: COCONINO CO.: Crater Lake, N of Mt. Agassy,
Lemmon 8244 (GH). California: BUTTE CO.: cold spring, Jonesville,
Copeland 604 (F, GH, K, MICH, MO, MU, NY, uc, us). Colorado:
JACKSON CO.: Big Creek Lake, Porter 6433 (DAO, ups). Connecticut:
TOLLAND CO.: Stafford, Morris (F, GH, ILL, us). Florida: FRANKLIN
co.: 6 mi. NE of Carrabelle, Adams 442 (GH, UC, US). Idaho:
SHOSHONE CO.: St. Maries River, 7 mi. SE of Clarkia, Cronquist &
Jones 6038 (CAN, DAO, GH, ILL, MICH, NY, UPS, UC, us). Illinois: COOK
co.: South Chicago, Hill 88-1880 (ILL). Indiana: STEUBEN CO:.
Crooked Lake, Williams (MO). Iowa: DICKINSON co.: Manhattan
Pond, Lakeville Twp. Thorne 13015 (uc). Louisiana: CALDWELL PAR.:
Horseshoe Lake, 8 mi. NNW of Columbia, Thomas 4224 (os). Maine:
PENOBSCOT CO.: Stillwater River, Old Town, Ogden & Steinmetz 1600
(CAN, GH, NY, us). Maryland: HARFORD CO.: Spesutie Island,
Moldenke 9398 (NY). Massachusetts: NORFOLK CO.: Charles River,
Dedham, Fernald & Svenson 410 (CAN, DAO, F, GH, ILL, MICH, MO,

1974] Potamogeton — Haynes 637

NY, UC, UPS, US). Michigan: CHEBOYGAN CO.: Black River, between
its mouth and Alverno, Haynes 3378 (GH, ILL, os, US), Haynes 3387
(os); Marl Bay NW corner of Douglas Lake, Haynes 3712 (GH, ILL,
MO, 0S, UC), Haynes 3924 (os). LUCE CO.: Bodie Lake, Haynes 3790
(os); bog lake, 612 mi. NE of Newberry, Haynes 3718 (08). SCHOOL-
CRAFT CO.: Seney Wildlife Refuge, Haynes 3741 (os). Minnesota:
KOOCHICHING CO.: ditch 2 mi. E of Tilson Bay, Rainy Lake, Moore
& Moore 11778 (DAO, GH, ILL, NY, UC, US). Montana: MISSOULA CO.:
Ft. Missoula, Hitchcock 23968 (DAO, NY, UC). Nevada: ELKO CO.:
Hot Creek Field, NE of Golliher Pasture, Holmgren 1401 (DAO, NY,
uc). New Hampshire: CHESHIRE CO.: East Jaffrey, Deane (GH).
New Jersey: CAMDEN CO.: Newton Creek, West Collingswood, Adams
370 (GH, MO, PH). New York: CORTLAND CO.: borrow pit just N of
Little York, Haynes 3353 (0S). ESSEX CO.: outlet of Lake George,
near Ticonderoga, Haynes 3336 (08). SARATOGA CO.: Saratoga Lake,
Haynes 3301 (os), Haynes 3307 (0S). SUFFOLK CO.: Wading River,
Riverhead, Miller (F, GH, US[4]). WARREN C0.: Dunham Bay, S end
of Lake George, Haynes 3326 (GH, MICH, MO, NY, OS, US), Haynes
3329 (CAN, GH, ILL, MICH, MO, NY, OS, PH, UC, US); Lake Luzerne,
Haynes 3812 (os). North Carolina: CRAVEN CO.: 2 mi. SW of Blades,
Radford 5856 (DAO, GH, NY). North Dakota: STUTSMAN CO.: Jim
Lake, Pingree, Mabbott 316 (NY). Ohio: JACKSON CO.: Sec. 10,
Jefferson Twp., Roberts 795 (CAN, LAF, MICH, OS, PH, US). LOGAN
co.: Indian Lake State Park, Haynes 3418 (08). OTTAWA CO.: Mid-
dle Bass Island, Lake Erie, Haynes 3134 (os); Squaw Harbor,
Put-in-Bay, Lake Erie, Haynes 3127 (MICH). PORTAGE CO.:
Twin Lakes, Haynes 3448 (oS). VINTON CO.: Lake Alma State Park,
Haynes 3468a (os). Oklahoma: COMANCHE C0.: Creek near Cache,
Stevens 1369 (GH, ILL, K, MO, NY, US). Oregon: Porter Lake of
Willamette River, 5 mi. S of Corvallis, Wentz 344 (MICH, MO, NY,
os, vs). Pennsylvania: LUZERNE CO.: Lily Lake, Small (F, ILL,
MU, NY). Rhode Island: NEWPORT CO.: Block Island, Fernald et al.
8448 (PH). South Carolina: GEORGETOWN CO.: Georgetown, Forster
(NY). Texas: BOWIE COo.: Club Lake, 3 mi. W of New Boston,
Correll & Ogden 25248 (LAF). Vermont: CALEDONIA CO.: beaver
pond, Walden Twp., Haynes 3828 (MICH, OS). ESSEX CO.: Bruns-
wick, Eggleston 1654 (GH, NY, US). LAMOILLE CO.: Wolcott Pond,
Wolcott. Haynes 3826 (os). Virginia: SUSSEX CO.: Chappell’s Mill-
pond, W of Lumberton, Fernald & Long 12236 (GH, PH). Wash-
ington: KLICKITAT CO.: Columbia River, Bingen, Suksdorf 2570
(F, GH, Mo, US). Wisconsin: JUNEAU CO.: Glacial Lake Wisconsin,
Cutler Twp. Hartley 8276 (GH, US). Wyoming: ALBANY CO.: Swas-
tika Lake, Medicine Bow Mountains, Porter 6190 (DAO, GH, UC, UPS).
CROOK CO.: pool 7 mi. N of Moorcroft, Porter & Miller 5974 (DAO,
MO, UC, UPS).

638 Rhodora [Vol. 76

Potamogeton perfoliatus X pusillus var. tenuissimus

Morong (1880) named Potamogeton mysticus based upon
a collection from Mystic Pond, Massachusetts. The entity
was said to have the habit of P. perfoliatus, a species of
subsection Perfoliati of Hagstrom, but differed from this
species by its delicate stature. Morong noted that the entity
had never been known to fruit. Ogden (1943), however,
based upon the extreme rarity of the entity and that it was
not known to fruit, proposed it to be a hybrid with P. per-
foliatus as one of its parents. He examined the stem anat-
omy of the entity, as well as studying the plants in the field.
The stem anatomy appeared to be intermediate between
P. perfoliatus and the Pusilli. At the one locality in which
Ogden found the putative hybrid, P. berchtoldii var. tenuis-
simus [P. pusillus var. tenuissimus] was present. He thus
suggested the entity to be a result of hybridization of P.
perfoliatus and P. berchtoldii var. tenuissimus [P. pusillus
var. tenuissimus]. Again, until the entity can be studied
in depth, I am accepting the concept of Ogden.

SPECIMENS EXAMINED:

UNITED STATES: Maine: CUMBERLAND CO.: Scarboro, West
Scarboro, Scarboro River, Ogden et al. 1731 (CAN, MO, NYS, US);
Scarboro, West Scarboro, Stuart Brook, Steinmetz & Martson 539
(CAN, F, ILL, UC, US). Massachusetts: MIDDLESEX CO.: Medford,
Mystic Pond, Booth (US), Faxon (ILL, US), Morong 11 Aug. 1879
(lectotype here designated, Ny!; isolectotypes, K!, MICH!, NY[3]!),
Morong (K, NY). North Carolina: DARE CO.: Currituck Sound, inlet
S of Duck, Radford 5757 (GH, NY, UC).

7c. Potamogeton pusillus var. gemmiparus Robbins in A.
Gray Man. Bot. North. U. S. 5th ed. 489. 1867. TYPE:
Robbins, Valley of the Blackstone, Uxbridge, Massachusetts,
(lectotype here designated, NY!; isolectotypes, GH!, NY!,
PH !, uS!).

Potamogeton gemmiparus (Robbins in A. Gray) Morong,
Bot. Gaz. (Crawfordsville) 5: 51. 1880.

Stems pale green, terete, smooth, 30-140 em long, 0.1-0.3
mm diam. Leaves pale green, uninervate, 1.1-6.0 cm long,
0.2-0.7 mm wide; apex subulate; glands gold, to 0.8 mm

1974] Potamogeton — Haynes 639

diam; lacunae absent or to 2 rows each side of the midrib.
Stipules white, convolute, 4.1-7.2 mm long, 0.2-0.4 mm diam.
Winter buds abundant, terminal or lateral, 1.2-2.8 cm long,
0.7-1.5 mm wide; outer leaves 1-2 per side, subulate, much
longer than inner. Peduncles cylindric, axillary or terminal,
1.1-2.8 em long, 0.2-0.5 mm diam. Spike cylindric, 2.5-5.5
mm long, 1.2-3.0 mm diam; verticels 2-3, to 3.5 mm apart.
Perianth segments 1.0-1.7 mm long, 0.7-0.8 mm wide. Fruit
rare, greenish-brown, 1.9-2.0 mm long, 1.5-1.6 mm wide;
beak central, 0.1-0.2 mm long, 0.2-0.3 mm diam; sides cen-
trally depressed; wall texture rough. Chromosome number
unknown.

Distribution: South-central Quebec south to central Mas-
sachusetts and Rhode Island. Fruiting late August to late
September. Fig. 15.

Illustrations: Fernald (1932, pl. 12; 30, fig. 1; 34, fig. 1).

o
b \ y X 4
em Y
~y N P , x
I: v om \
g Y \
"4
iu ws
S
\
" AD | 4
D 3 g Y
e.
. T
, c ‘As P \ art
o 9 At uU e a
d D j i" à >Ê
c - \ Q
4 [e] - A p

` E l t e q
Pa i bo ; " © Le
Eon \ [*] i
^ A A P OE c i n- 1
Fig. 13. Map of northeastern United States and southeastern
Canada showing documented distribution of Potamogeton pusillus

var. gemmiparus.

640 Rhodora [Vol. 76

REPRESENTATIVE SPECIMENS

CANADA: Quebec: Saint-Tite, comte de Champlain, Lake Pierre-
Paul Gauthier 2273 (GH, photo at DOA). UNITED STATES: Maine:
CUMBERLAND CO.: Horseshoe Pond, North Fryeburg, Hellquist 4457
(MICH). KENNEBEC CO.: Belgrade Stream, Belgrade, Hellquist 4501
(MICH). PENOBSCOT CO.: Upper Stillwater, Fernald (uc, us), Fer-
mald (GH). Massachusetts: HAMPSHIRE CO.: Amherst, Jesup (GH,
NY). MIDDLESEX CO.: Charles River, South Natick, Morong (NY).
NORFOLK CO.: Charles River, Dedham, Fernald & Svenson (GH).
WORCESTER CO.: Uxbridge, Morong (CGE, F, ILL, MICH, MO, NY, US).
New Hampshire: CARROLL CO.: Conway Lake, Eaton, Hellquist 4469
(MICH). HILLSBOROUGH CO.: Pine Island Pond, Manchester, Kroch-
mal 265 (GH). Rhode Island: PROVIDENCE CO.: Central Pond, Ten
Mile River, East Providence, Collins (GH).

8. Potamogeton groenlandicus Hagstrom, Kongl, Svenska
Vetenskapsakad. Handl. 55(5): 127. 1916. TYPE: Hartz,
Kingua Orpiksnit, 68°30’, Greenland, (lectotype here desig-
nated, C!).

Potamogeton pusillus ssp. groenlandicus (Hagstr.) Boch.
1952. Meddel. Groenland 147 (9) : 44.

Stems light green, terete, smooth, 20-50 cm long, 0.2-0.3
mm diam. Leaves pale to deep green, 7-9 nerved, 2.6-8.8
mm long, 0.7-1.7 mm wide; apex acute to apiculate; glands
usually present, white, 0.1-0.7 mm diam; lacunae absent or
to three rows present each side of the midrib; lateral nerves
joining the midrib 0.7-1.2 mm below the apex. Stipules
brown, delicate, convolute, 4.5-19.1 mm long, 0.2-0.4 mm
diam. Winter buds common, terminal or axillary, 4.2-8.4
em long, 1.4-2.5 mm wide; inner leaves unmodified; outer
leaves 2 per side, acute to apiculate, without corrugations
at base. Peduncles rare, cylindric, terminal, ca 18.2 mm
long, ca 0.5 mm diam. Spikes cylindric, ca 5.0 mm long,
ca 3.5 mm diam; verticels 3, crowded, ca 1.0 mm apart.
Perianth segments ca 1.1 mm long, ca 0.7 mm wide. Fruit
unknown. Chromosome number unknown.

Distribution: Western Greenland. Flowering in late
August. Fig. 14.

Illustrations: (apparently none).

1974] Potamogeton — Haynes 641

In 1916, Hagstrom named Potamogeton groenlandicus
from several sterile specimens. He distinguished this spe-
cies from other Pusilli by its possessing 5-9 nerves per leaf.
“In this respect,” he stated, “this species constitutes an
interesting link between the Pusilli and the Oxyphylli and
would be ranked with the latter as well as with the former."
He added that the ligules and leaf-apices, however, unite it
with the Pusilli. Fernald (1932), after examining as many
specimens as available [but not the Type], concluded the
taxon to be conspecific with P. pusillus var. mucronatus
[var. tenuissimus]. He stated that he had been unable to
convince himself that the leaves were more than three-
nerved. He added that this character alone does not war-
rant specific rank. I have examined the material at Copen-
hagen and at all the large herbaria of the United States and
Canada. In my opinion, the leaves are 5-9 nerved. I agree
with Fernald, however, that one vegetative character is not
enough to separate two Species, especially when one is

Fig. 14. Map of Greenland showing documented distribution of
Potamogeton groenlandicus.

642 Rhodora [Vol. 76

known only from non-fruiting material. However, both
Fernald and Hagstrom overlooked the winter buds. In P.
groenlandicus, the inner leaves of the winter buds are not
modified into hardened fusiform structures, as in P. pusillus.
Instead, the winter buds are similar to those of P. obtusi-
folius, in that the inner leaves are unmodified. To my
knowledge, only one population of fertile individuals has
been found, this being in flower. The flowers are most
similar to P. pusillus. Because the winter buds and number
of nerves are so different from P. pusillus, I am currently
accepting P. groenlandicus at the specific level. However,
material should be gathered and cultivated. If fruiting does
occur, then the correct affinities of this endemic taxon can
be ascertained.

REPRESENTATIVE SPECIMENS

GREENLAND: Christianshaab Dist.: Christianshaab, Hartz (c).
Egedesminde Dist.: Egedesminde, 67?44'N, Porsild (CAN, GH, MO,
NY, US). Frederikshaab Dist.: Frederikshaab, 62^00'N, 49°40'W,
Jorgensen & Larsson (CAN); Ivigtut, Berlin (c). Godhavn Dist.:
Disko, near Arktisk Station, 69?15'N, Porsild (CAN, F, GH, US) ; Disko,
Godhavn, Laegaard 350 (CAN, Us); Disko, near Godhavn, 69?14'N,
Porsild 228 (GH, US); Disko, Sinigtik, 69°25'N, Porsild (CAN, GH);
Disko, Skauseklippen, 69°25'N, Porsild (F). Godthaab Dist.: vicinity
of Godthaab, 64°10'N, 51°43’W, Porsild 8273 (CAN) ; Godthaab Fjord,
Itivnera, 64°22'N, Porsild (GH) ; Godthaab Fjord, Jersiutilik, Trap-
nell 402 (GH, K). Jakobshavn Dist.: Jakobshavn, Sorensen (C).
Julianehaab Dist.: Igaliko Fjord, Qagssaarssuk, Polunin 10779
(CAN); vicinity of Julianehaab, 60^43'N, 46°05'W, Porsild 8120
(CAN). Ritenbenk Dist.: Ritenbenk, 69°44'N, Hansen (C); Sarkak,
pool on gmeiss, Seward & Holttum (cGE, K); 5 km E of Sarkak,
Beschel 12163 (CAN, photo at DAO). Sukkertoppen Dist.: Ungarsivik
Lake, 65?25'N, 52^50'W, Porsild 8870 (CAN). Upernavik Dist.:
Laksefjord, Proven, Porsild 8837 (CAN).

ACKNOWLEDGMENTS

I am grateful to Dr. Ronald L. Stuckey, who supervised
this study, for his advice and constant encouragement dur-
ing the course of my work. Appreciation is also owed to
Dr. Edward G. Voss for help with nomenclature; to Miss
Judy Canne, Mr. Robert C. Gardner, and Mr. David Keil

1974] Potamogeton — Haynes 643

for their helpful suggestions; to the curators of the herbaria
from which specimens were borrowed; to the staffs at Kew
Botanical Gardens and the British Museum for their pains-
taking search for the holotype of Potamogeton strictifolius;
to the Linnean Society of London for the photograph of the
holotype of P. pusillus; to my colleagues at the many librar-
ies who supplied xerox copies of many original descriptions;
to Mr. Robert C. Gardner, Dr. Pete Hostetter, and Mr.
Robert Kalinsky for their help with the photographs; to
Dr. Eugene C. Ogden, Mr. Stanley Smith, Dr. Edward G.
Voss, and Dr. Steven Young for help in locating populations
of Potamogeton in the field; and to Mr. Marvin L. Roberts,
Dr. John A. Schmitt, and Dr. Tod F. Stuessy for their
critical reading of the manuscript.

LITERATURE CITED

AALTO, M. 1970. Potamogetonaceae fruits. I. Recent and sub-
fossil endocarps of the Fennoscandian species. Acta Bot. Fenn.
88: 1-85.

ANDERSON, L. C. 1972. Flaveria campestris (Asteraceae): A case
of polyhaploidy or relic ancestral diploidy? Evolution 26: 671-
673.

ASCHERSON, P. 1889. Potamogetonaceae. In: A. Engler & K.
Prantl. Die natiirlichen Pflanzenfamilien. 2(1): 194-214. Verlag
von Wilhelm Engelmann, Leipzig.

ASCHERSON, P., & P. GRAEBNER. 1907. Potamogetonaceae. In: A.
Engler. Das Pflanzenreich Regni vegetabilis conspectus 4(11):
1-183. Wilhelm Engelmann, Leipzig.

BENNETT, A. 1890-1894. The nomenclature of Potamogeton. Jour.
Bot. 28: 297-302; 29: 150-152, 307; 30: 227-230; 31: 132-134, 294-
297; 32: 208-205.

1900-1904. Notes on Potamogeton. Jour. Bot. 38: 125-
130; 39: 198-201; 40: 145-149; 42: 69-77.

BOLKHOVSKIKH, Z., et al. 1969. Chromosome numbers of flowering
plants [in Russian]. Acad. Sci. of the USSR, Leningrad. 926
pp.

BosTRACK, J. M., & W. F. MILLINGTON. 1962. On the determination
of leaf form in an aquatic heterophyllous species of Ranunculus.
Bull. Torrey Bot. Club. 89: 1-20.

CHRYSLER, M. A. 1907. The structure and relationships of the Pota-
mogetonaceae and allied families. Bot. Gaz. (Crawfordsville)
44: 161-188.

4

644 Rhodora [Vol. 76

CLAPHAM, A. R., et al. 1962. Flora of the British Isles, 2nd ed.
Cambridge Univ. Press, Cambridge. 1296 pp.

Cook, C. D. K. 1966. A monographie study of Ranunculus sub-
genus Batrachium (DC) A. Gray. Mitt. Bot. Staatssamml.
München 6: 47-237.

CRONQUIST, A. 1968. The evolution and classification of flowering
plants. Houghton Mifflin Co., Boston. 396 pp.

Danpy, J. E. 1937. The genus Potamogeton in tropical Africa.
Jour. Linn. Soc. Bot. 50: 507-540 + 2 pl.

Danby, J. E. & G. TAYLOR. 1938. Studies of British Potamogetons.
— I. The typification of Potamogeton pusillus. Jour. Bot. 76:
89-92.

— —————24 1940. Studies of British Potamogetons. — XIII. Pota-
mogeton berchtoldii in Great Britain. Jour. Bot. 78: 49-66.
DEWET, J. M. J. 1965. Diploid races of tetraploid Dichanthium

species. Amer. Naturalist 99: 167-172.

Eames, A. J. 1961. Morphology of angiosperms. McGraw-Hill,
New York. 518 pp.

Fassett, N. C. 1940. A manual of aquatic plants. McGraw-Hill
Book Co., New York, 382 pp.

1951. Callitriche in the New World. Rhodora 53:
187-155, 161-182, 185-194, 209-222.

FERNALD, M. L. 1932. The linear-leaved North American species of
Potamogeton section Azillares. Mem. Amer. Acad. Arts 1T:
1-183. (Also, Mem. Gray Herb. No. 3)

1940. Some spermatophytes of eastern North America.
Rhodora 42: 239-302.

1950. Gray's manual of botany. 8th ed. American
Book Co., New York. lviv + 1632 pp.

FniNT, R. F. 1957. Glacial and pleistocene geology. John Wiley
& Sons, New York. 553 pp. * 5 pl

Funt, R. F., et al. 1959. Glacial map of the United States east
of the Rocky Mountains. Geol. Soc. Amer.

Fries. E. M. 1828. Novitiae florae Suecicae, ed. 2, Lund.

Fryer, A., & A. BENNETT. 1915. The Potamogetons (pondweeds)
of the British Isles. L. Reeve and Co., Limited, London. 94 pp.
+ 60 pl.

GAEVSKAYA, N. S. 1966. The role of higher aquatic plants in the
nutrition of the animals of fresh-water basins. [Transl. from
Russian by] Maitland Muller, D. G. 1969. National Lending
Library for Science and Technology. 3 vols.

Gav, M. J. 1854. Botanique — Etudes organographiques sur la
famille des Potamees. Premier Memoire: Sur les genres Pota-
mogeton Spirillus et Groenlandia. Compt. Rend. Hebd. Seances
Acad. Sci. 38: 702-705.

1974] Potamogeton — Haynes 645

GLEASON, H. A., & A. CRONQUIST. 1963. Manual of vascular plants
of Northeastern United States and adjacent Canada. D. Van
Nostrand, New York. 810 pp.

GUIRE, K. E. & E. G. Voss. 1963. Distributions of distinctive
shoreline plants in the Great Lakes region. Michigan Bot. 2:
99-114.

Hagstrom, J. O. 1916. Critical researches on the Potamogetons.
Kongl. Svenska Vetenskapsakad. Handl. 55(5): 1-281.

HaRADA, I. 1942. Chromosome studies on Potamogeton. Jap. Jour.
Bot. 18: 92.

1956. Cytological studies in Helobiae I. Chromosome
idiograms and a list of chromosome numbers in seven families.
Cytologia 21: 306-328.

HARMAN, W. N., & T. R. DoANE. 1970. Changes in the aquatic
flora of Otsego Lake between 1935-1969. New York Fish Game
Jour. 17: 121-123.

Haynes, R. R, & J. L. REVEAL. 1973. A re-evaluation of Pota-
mogeton fibrillosus Fern. (Potamogetonaceae). Rhodora 75: 75-
77.

HiTCHCOCK, C. L. 1969. Potamogetonaceae. pp. 158-175. In: C. L.
Hitchcock et al. Vascular plants of the Pacific Northwest. Univ.
Wash. Publ. Biol. 17(1).

HULTÉN, E. 1937. Outline of the history of arctic and boreal biota
during the quaternary period. Bokforlags Aktiebolaget Thule,
Stockholm. 168 pp. + 43 pl.

IRMISH, T. 1858. Ueber einige arten aus der naturlichen pflan-
zenfamilie der Potameen. G. Bosselmann, Berlin. 56 pp. +
3 pl.

Irris, H. H. 1965. The genus Gentianopsis (Gentianaceae): trans-
fers and phytogeographic comments. Sida 2: 129-153.

JOHNSON, M. F., & H. H. IrTis. 1963. Preliminary reports on the
flora of Wisconsin No. 48. Compositae I — Composite family 1.
Trans. Wisconsin Acad. Sci. 52: 255-342.

JØRGENSEN, C. A. T. SØRENSEN, & M. WESTERGAARD. 1958. The
flowering plants of Greenland. A taxonomical and cytological
survey. Biol. Skr. 9(4): 1-172.

KAPADIA, Z. J. 1964. Varietas and subspecies: a suggestion towards
greater uniformity. Taxon 12: 257-259.

KocH, D. G. D. J. 1837. Synopsis florae Germanicae et Helveticae.
Frankfurt. 844 pp.

LANJOUW, J., & F. A. STAFLEU. 1964. Index Herbariorum I. The
herbaria of the world. Reg. Veg. 31: 1-251.

LAWRENCE, G. H. M. 1951. Taxonomy of vascular plants. The
MacMillan Co. New York. 823 pp.

646 Rhodora [Vol. 76

LIND, C. T, & G. COTTAM. 1969. The submerged aquatics of Uni-
versity Bay: A study in eutrophication. Amer. Midl. Nat. 81:
353-369.

LINNEAUS, C. 1753. Species plantarum, Stockholm. 2 vols.
LOHAMMAR, G. 1954. The effect of digestion on the germination of
Potamogeton seeds [in Swedish]. Fauna & Fl. 49: 17-32.

LOUREIRO, J. DE. 1790. Flora cochinchinensis. Lisbon, 2 vols.

LóvE, A. 1954. Cytotaxonomical evaluation of corresponding taxa.
Vegetatio 5-6: 212-224.

Love, A., & D. Live. 1961. Chromosome numbers of central and
northwest European plant species. Opera Bot. 5: 1-581.

MARIE-VICTORIN, F. 1938. Phytogeographical problems of eastern
Canada. Amer. Midl. Nat. 19: 489-558. (Also, Contr. Inst. Univ.
Montreal No. 30).

Martin, A. C. 1951. Identifying pondweed seeds eaten by ducks.
Jour. Wildl. Manage. 15: 253-257.

MERTENS, F. K, & W. D. J. KocH. 1823. In: J. C. Rohling.
Deutschland flora, 3rd ed., vol. 1. Frankfurt. 891 pp.

MICHAUX, A. 1808. Flora boreali-americana. Paris. 2 vols.

MooRE, E. 1915. The Potamogetons in relation to pond culture.
Bull Bur. Fisheries 33: 251-291.

MonoNG, T. 1893. The Naiadaceae of North America. Mem. Torrey
Bot. Club 3: 1-65.

1880. New species of Potamogeton, with notes upon
some published forms. Bot. Gaz. (Crawfordsville) 5: 50-53.

Movie, J. B, & N. Horcnkiss. 1945. The aquatic and marsh
vegetation of Minnesota and its value to waterfowl. Minnesota
Dept. Conserv. Tech. Bull. No. 3: 1-122.

MUENSCHER, W. C. 1936. The germination of seeds of Potamogeton.
Ann. Bot. (London) 50: 805-821.

1944. Aquatic plants of the United States. Comstock
Publ. Co. Ithaca. 374 pp.

MULLER-LIEBENAU, I. 1956. Die besiedlung der Potamogeton-zone
ostholsteinischer seen. Arch. Hydrobiol. 52: 470-606.

NIEUWLAND, J. A. 1913. Notes on our local plants. II. Amer.
Midl. Nat. 3: 14-22.

OEDER, G. C. 1765. Flora Danica, vol. 1. Copenhagen. 180 pp.

OGDEN, E. C. 1943. The broad-leaved species of Potamogeton of
North America north of Mexico. Rhodora 45: 57-105, 119-163,
171-214.

1957. Revision appendix, p. 363-384. Im: N. C. Fas-
sett. A manual of aquatic plants. Univ. Wisconsin Press,
Madison. 405 pp.

1966. Potamogetonaceae. In: C. L. Lundell, Flora of
Texas 1: 369-282, + 9 pl.

1974] Potamogeton — Haynes 647

1972. Flora of New York. Potamogeton. Unpubl.
manuscript [New York State Museum].

PALMGREN, O. 1939. Cytological studies in Potamogeton. Bot. Not.
1939: 246-248.

Ponp, R. H. 1905. The biological relation of aquatic plants to the
substratum. Rep. U. S. Commiss, Fish and Fisheries 1903: 483-
526.

PonTER, C. L. 1963. 13. Najadaceae, p. 6-12. In: A flora of
Wyoming. Part II. Wyoming Agric. Exp. Sta. Bull. 404.
PursH, F. T. 1814. Flora Americae Septentrionalis. London. 2 vols.
RAFINESQUE, C. S. 1808. Prospectus of Mr. Rafinesque Schmaltz's
two intended works on North-American Botany; the first on the
new Genera and species of plants discovered by himself, and
the second on the natural history of the funguses, or mushroom-

tribe of America. Med. Repos. series 2, 5: 350-356.

1811. Botanical information concerning two families
of plants. Med. Repos. series 6, 2: 407-409.

1819. Prodrome des noveaux generes de plantes ob-
serves en 1817 et 1818 dans l'interieur des Etats-Unis d'Amerique.
J. Phys. Chim. Hist. Nat. Arts 89: 96-107.

RAUNKIAER, C. 1896. De Danske Blomsterplanters Naturhistorie
I. Helobieae. Copenhagen.

1903. Anatomical Potamogeton-studies and Potamoge-
ton fluitans. Bot. Tidsskr. 25: 253-380.

RAVEN, P. H. 1969. A revision of the genus Camissonia (Onagra-
ceae). Contr. U. S, Natl. Herb. 37: 162-396.

RAvEN, P. H., & H. J. THoMPSON. 1964. Haploidy and angiosperm
evolution. Amer. Naturalist 98: 251-252.

REICHENBACH, L. 1845. Icones florae Germanicae et Helveticae.
vol. 7. Leipzig. 40 pp. + 71 pl.

RENDLE, A. B. 1930. The classification of flowering plants. Cam-
bridge Univ. Press, Cambridge. 2 vols.

ROEMER, J. J., & J. A. SCHULTES. 1818. Systema vegetabilium. vol.
3. Stuttgart.

RUPRECHT, F. J. 1845. Distributio cryptogamarum vascularium in
imperio rossico. Beitr. Pflanzenk. Russ. Reiches 4: 1-56.
SATTLER, R. 1965. Perianth development of Potamogeton richard-

sonii, Amer. J. Bot. 52: 35-41.

SCULTHORPE, C. D. 1967. The biology of aquatic plants. Edward
Arnold Ltd., London [also, St. Martin Press, New York]. 610 pp.

SHELFORD, V. E. 1918. Conditions of existence, pp. 21-60. Im:
H. B. Ward and G. C. Whipple. Fresh-water biology. John
Wiley & Sons, Inc. London.

1937. Animal communities in temperate America.
Univ. of Chicago, Chicago. 362 pp.

648 Rhodora [Vol. 76

SINGH, V. 1964. Morphological and anatomical studies in Helobiae.
I. Vegetative anatomy of some members of Potamogetonaceae.
Proc. Indian Acad. Sci. 40: 214-231.

1965. Morphological and anatomical studies in Helo-
biae. II. Vascular anatomy of the flower of Potamogetonaceae.
Bot. Gaz. (Crawfordsville) 126: 137-144.

SMITH, J. E. 1797. Potamogeton compressum. English Bot. 2:
pl. 418.

SONDER, W. 1851. Flora Hamburgensis. Hamburg. 601 pp.

STERN, K. R. 1961. Chromosome numbers in nine taxa of Pota-
mogeton. Bull. Torrey Bot. Club 88: 411-414.

Stuckey, R. L. 1971. Changes of vascular aquatic flowering plants
during 70 years in Put-in-Bay Harbor, Lake Erie, Ohio. Ohio
Jour. Sci. 71: 321-342.

1972. Taxonomy and distribution of the genus Rorippa
(Cruciferae) in North America, Sida 4: 279-430.

SULLIVAN, V. I. 1967. The effect of water depth on the germina-
tion of Potamogeton seeds. Unpubl. manuscript [Univ. Michi-
gan].

TAYLOR, N. 1909. Zannichelliaceae. N. Am. Fl. 17: 13-35.

TAYLOR, R. L., & G. A. MULLIGAN. 1968. Flora of the Queen Char-
lotte Islands. pt. 2. Cytological aspects of the vascular plants.
Res. Branch, Canada Dept. Agr. Monogr. no. 4, pt. 2. 148 pp.

THIERET, J. W. 1960. Calamovilfa longifolia and its variety magna.
Amer. Midl. Nat. 63: 169-176.

TOURNEFORT, J. P. 1719. Institutiones rei herbariae. Paris. 3 vols.

TRAUTMAN, M. B. 1957. The fishes of Ohio. The Ohio State Univ.
Press, Columbus. 683 pp.

TUCKERMAN, E. 1849. Observations on American species of the
genus Potamogeton L. Am. Jour. Sci. Arts. series 2, 7: 347-360.

VOLKER, R., & S. G. SMITH. 1965. Changes in the aquatic vascular
flora of Lake East Okoboji in historic times. Proc. Iowa Acad.
Sci. 72: 65-72.

Voss, E. G. 1967. The status of some reports of vascular plants
from Michigan. Michigan Bot. 6: 13-24.

1972. Michigan flora. pt. 1. Gymnosperms and mono-
cots. Cranbrook Inst. Sci. Bull. No. 55. Bloomfield Hills, 488

pp.
WALTER, T. 1788. Flora caroliniana. London. 263 pp.
WELCH, P. S. 1935. Limnology. McGraw-Hill Book Co., New York.
471 pp.
1952. Limnology, ed. 2. McGraw-Hill Book Co., New
York. 538 pp.

1974] Potamogeton — Haynes 649

WENTZ, W. A. & R. L. Stuckey. 1971. The changing distribution
of the genus Najas (Najadaceae) in Ohio. Ohio Jour. Sci. 71:
292-302.

WILLIAMS, G. R. 1970. Investigations in the white waterlilies
(Nymphaea) of Michigan. Michigan Bot. 9: 72-86.

Yro, R. R. 1966. Yields of propagules of certain aquatic plants.
I. Weeds 14: 110-113.

DEPARTMENT OF BIOLOGICAL SCIENCES
LOUISIANA STATE UNIVERSITY IN SHREVEPORT
SHREVEPORT, LOUISIANA 71105

SARRACENIA ALABAMENSIS,
A NEWLY RECOGNIZED SPECIES
FROM CENTRAL ALABAMA'

FREDERICK W. CASE, JR. AND ROBERTA B. CASE

In February, 1953, while collecting pitcher plants at
Elmore, Alabama, for taxonomic studies, we located a
colony of Sarracenia with frozen leaves. As we were seek-
ing plants of S. oreophila (Kearney) Wherry at Wherry's
published range extension for that species at Elmore, and
since the frozen leaves were mostly large and widely ex-
panded at the orifice, much resembling the published photo-
graphs of the yellow flowered S. oreophila, which we had
not at that time seen, we assumed our discovery to repre-
sent that species. When the plants flowered in our green-
house, however, the flowers were small and red. This
seemed to place the plant in S. jonesii Wherry, which was
widely accepted at that time. After observing and growing
the plant for two seasons, we became convinced that either
the published description of S. jonesii was too narrow, or
that the plant was not S. jonesii at all. Our next step was
to visit the type locality of S. jonesii in North Carolina to
observe and obtain plants for comparison. They proved
not to be the same, so we began the studies leading to this
paper.

With the help of the late Dr. R. M. Harper, who made
his field notes available to us, and who, in 1955, guided us
to his colonies of pitcher plants in Central Alabama, we
were able to obtain a wide sampling of the population and
to locate other colonies of our own. Because there has long
been controversy and confusion among Sarracenia taxon-
omists over the treatment of the taxa related to S. rubra
Walter, we visited the Elmore and Chilton County colonies
eight times at various seasons in the period from 1953 to
1973 to better understand this related species.

'Manuscript received January, 1973.

650

1974] Sarracenia — Case & Case 651

Since 1953, we have grown this plant, which we call the
Alabama Canebrake Pitcher Plant, continuously, both in
our greenhouses, and, since 1966, outdoors as well, at Sag-
inaw, Michigan. Although its small red flowers with obo-
vate petals indicate a close relationship to S. rubra, it shows
a number of differences. In this connection, those interested
in significant taxonomic characteristics in Sarracenia may
wish to refer to the remarks of MacFarlane (1908), Wherry
(1929, 1935), Bell (1949), and MeDaniel (1966).

Unlike Sarracenia rubra (MacFarlane, 1908) our plant
produces ftvo types of pitchers and occasional phyllodia
each season. Its spring pitchers, similar in size to the
leaves of S. rubra, differ markedly in developing a curved
form and in lacking the external dark maroon coloring of
the veins, pitcher rim and hoods found in that species. The
summer pitchers, straight and usually much larger and
more tapered than those of S. rubra (excepting S. rubra
ssp. jonesii), differ from those of both of those taxa in
possessing a softer texture, a visibly fine pubescence, and a
paler yellow-green color. The major veins are maroon col-
ored mainly only on the inside of the pitcher tube. The
hoods of the summer pitchers are from 2-8 times larger
than those of summer leaves produced in S. rubra, The
greater expansion of the hood, the stronger reflexion of
the hood neck margins, the strongly rolled orifice rim, the
leaf size, and overall yellowish green coloration give the
summer leaves a profile more suggestive of the pitcher of
S. flava or S. oreophila than of S. rubra.

The flowers, while less useful than the leaves for species
delineation within this genus (c.f. Bell, 1949, p. 140), differ
from Sarrecenia rubra in their larger size, lighter red
coloring, petal shape, in their erose petal margins, and in
their fainter fragrance (Fig. 1). In addition, the plant
produces flower scapes abundantly; often two or three
scapes develop from one terminal growth in vigorous plants,
making large flowering clumps spectacular. The regular
production of multiple blooms from one terminal bud is an
unusual feature in Sarracenia.

652 Rhodora [Vol. 76

Figure 1. Comparison of flowers of Sarracenia alabamensis, Chil-
ton Co. Ala. (left) and S. rubra, Crestview, Florida (right). Note
differences in color tone, shape of petal lobe, and detail of petal
margins.

Since, in addition to its structural differences, its range
is disjunct from Sarracenia rubra and probably from that
of all other species except S. oreophila, we consider it a dis-
tinct species.

MacFarlane (1908) has pointed out that all immature
Sarracenia leaves tend to resemble closely those of Sar-
racenia minor Walter and do not show good specific dif-
ferences. While he spoke mostly of very young pitcher

1974] Sarracenia — Case & Case 653

leaves, we believe that all specific differences are best dis-
played by the largest leaves of the season. In our descrip-
tion we give the usual size ranges, but we urge readers to
standardize their comparisons, when possible, by examining
the largest seasonal leaves of related taxa. Because we
believe that vague size references without standardized
points of measurement have contributed to the problems
in understanding the taxa related to S. rubra, we have tried
to be quite exact in our measurements. Pitcher length was
measured from the point of attachment to the rhizome to
the rim of the pitcher orifice; it does not include the hood.
The length of hood refers to the distance from the narrow-
est part of the hood-constriction (neck) to the tip of the
hood, and hood width refers to measurement across its
widest point. To facilitate measurement comparisons of
the tubular pitchers in fresh or herbarium material, we
give width figures for pressed pitchers rather than diam-
eters for inflated pitchers.

The panduriform petals present problems for accurate
measurement comparisons, being divided into a cuneate
basal portion, an isthmus, and an obovate distal portion.
For purposes of comparison, the length of the basal cuneate
portion of the petal was measured as the distance from the
petal attachment along the median line to the widest point
of the segment; the length of the distal petal lobe we
measured from the widest point of the basal portion to the
tip of the petal.

We have based the following description upon both culti-
vated specimens and those collected in the wild.

Sarracenia alabamensis Case & Case, sp. nov.

Rhizoma valde ramosum, ramis confertis; folia trimor-
pha; folia vernalia ascidiformia, maxima plerumque plus
minusve sigmoidea, tubo basi angusto, sursum gradatim
ampliato, 17.7-49.5 cm. longo, 0.7-3 cm. lato; ala ventralis
[adaxialis] ad medium expansa, 0.4-1.5 em. lata, versus
orem et basin attenuata; tubi venae extus plerumque in-
coloratae, intus atropurpureae; operculum cordatum, acu-

654 Rhodora [Vol. 76

tum, subundulatum vel convexum, 1.9-7 cm. longum, 1.7-6
cm. latum, venis intus prope basin coloratis; collum 0.8-3
em. latum.

Folia maxima ascidiformia aestivalia quam vernalia
multo majora, foliis S. flavae vel S. oreophilae similia,
suberecta, recta vel subsigmoidea, extus dense pubescentia,
24.5-48.5 em. longa, ad basin longe angustata, in quarta
parte superiore parum ampliata, 3.1-6.7 em. lata; peristom-
ium valde revolutum, adaxiale incisum; ascidii textura
tenuis, mollis, ex salmoneo flavo-viridis, demum pallide
flavo-viridis; tubus prope orem extus obscure areolatus,
areolis albidis, venis intus valde purpureis, extus plerum-
que (maximis interdum exceptis) incoloratis; operculum
magnum, expansum, cordatum, fere tam latum quam
longum, 3.7-9.5 cm. longum, 3.4-8.6 cm. latum; collum 1.3-
3.4 cm. latum; textura inter venulas convexa; operculum
salmoneum, demum pallide flavo-viride, intus (necnon
praecipue collum peristomiumque) iridescens.

Phyllodia laminaria pauca, interdum e gemmis terminali-
bus majoribus oriunda, ascidia vernalia sequentia, aestivalia
praecedentia, prope basin valde recurvata, ultra medium
latiora, apicem versus denuo recurvata, 4-6.5 cm. longa,
0.3-0.8 em. lata.

Folia squamiformia pauca, triangularia, amplitudine
admodum variabilis.

Scapi numerosi, interdum aliquot per ramulum termina-
lem, 34.5-57.6 em. longi.

Calyx applanatus 4.5-6 cm. latus.

Sepala ovata, rotundata vel subacuta, 1.7-2.6 cm. longa,
1.1-2.2 cm. lata, sub anthesi faleato-incurvata, ovarium
mox cingentia, demum patentia vel reflexa, marginibus
vetustate valde revolutis.

Petala panduriformia, 3.4-4.6 cm. longa, extus sanguinea
(anglice “Oxblood Red"); pars basalis cuneata 0.7-1 cm.
longa, basi 0.3-0.4 em. lata, maxime 0.9-1.1 cm. lata; con-
strictio mediana marginibus reflexis; lamina ultra constric-
tionem in isthmum brevem parallelilateralem tum in lobum

1974] Sarracenia — Case & Case 655

obovatum erose marginatum 2.7-3.6 cm. longum, 1.8-2.5 cm.
latum expansa.

Ovarium tuberculatum, paullo compressum, 0.8 cm.
longum, 0.9 cm. latum. Stylus umbraculiformis, umbraculo
valde convexo, 3.5-4.2 em. lato.

TYPE: Alabama: ELMORE CO.: very rare and local in wet
gravelly or peaty soil and around springheads and rills in
open grassy swales, and in thickets of Rhododendron, Alnus,
Magnolia virginiana, Rhus vernix and Arundinaria tecta;
along the railroad between Elmore and Speigner. Type
plant collected in June 1971, Case & Case S-500 (us), but
all specimens prepared from clonal divisions grown in our
greenhouse and taken May 1, 1972 (flowering) and July
27, 1972 (large summer leaves).

REPRESENTATIVE SPECIMENS EXAMINED

Alabama: CHILTON CO.: Clanton, Pollard & Mason (us); sloping
gravelly bog near Jasmine, 28 Apr. 1921, Harper (NY, Us); Jasmine,
18 Apr. 1921, Harper (NY); gravelly bog, near Adams, 17 June
1955, F. & R. Case, C-100 (MicH), C-101, C-102 (NY); boggy rill
feeding Mulberry Creek, Pletcher, 17 June 1955, F. & R. Case et al.,
C-108 (us); cultivated material, Canebrake-Magnolia glauca bog,
Adams, grown at Saginaw, Michigan, F. & R. Case 8-520, S-522 (us),
S-524, S-527, S-528 (Ncvu), S-521, S-525, S-526 (MICH); boggy soil
around springhead, with Arundinaria tecta, Magnolia virginiana, Jas-
mine, 19 Apr. 1973, F. & R. Case C-104 (NcU), C-105 (US). ELMORE
co.: Slightly damp sandy soil 112 miles S. of Speigner, 14 Apr. 1934,
Harper (PH); boggy depression along railroad near station, Elmore,
13 July, 1935, Wherry (PENN); approximately 114 miles north of

Elmore along railroad, damp boggy ground, specimen from a culti-
vated plant (Oct. 1955), collected 16 Feb. 1953, F. & R. Case, C-106
(us).
DISTRIBUTION AND ECOLOGY

This extremely rare plant is known to occur only in
small, very localized colonies among the Fall Line Sand
Hills of Chilton, Autauga, and Elmore Counties, Alabama,
north of the Alabama River in an area roughly bounded by
the Coosa River to the east, and the Mulberry River to the
west. The soils there derive from rocks generally regarded
as Upper Cretaceous in origin (Harper, 1922). Except for
Wherry’s report of Sarracenia oreophila at Elmore, Ala-

656 Rhodora [Vol. 76

bama, no other Sarracenia is reported to grow within the
range of S. alabamensis.

Sarracenia alabamensis occurs in somewhat sandy-grav-
elly bogs, usually on sloping ground, or in damp peaty or
mucky soil around small spring-heads and tiny rills. It is
not usually a plant of sphagnum beds although species of
Sphagnwm may occur in isolated patches nearby. The
colonies we have observed are extremely small compared
to those of most of the other southern species, and range
from single clumps to one group of perhaps 100 plants in
a 20 by 50 foot area. According to Harper's (1922) ob-
servations and our own, the following plants commonly
grow with the Canebrake Pitcherplant: Magnolia virgin-
tana, Alnus sp. Arundinaria tecta, Acer rubrum, Rhus
vernix, Smilax laurifolia, Osmunda cinnamomea, Pteridium
aquilinum var., Juncus sp., Drosera capillaris, Pinguicula
sp., Aletris aurea, Rhexia sp., Pogonia ophioglossoides,
Rhododendron sp., and Calopogon tuberosum.

The pitcher plant grows most often in open stands of
Magnolia virginiana, Arundinaria tecta, and Alnus, in the
“Canebrakes” of the region, or in open boggy swales. Tol-
erant of more shade than most of the trumpet-leaf sarra-
cenias, the species nevertheless reaches its most vigorous
and distinctive development when growing in full sunlight.
Although few of the large, distinctive summer leaves de-
velop on heavily shaded plants, the species is able to flower
sparingly even when so shaded that not even the curved
spring pitchers can develop without severe deformity.

Hardest to detect of the sarracenias, this species probably
has always been very limited in distribution in modern
times, for its habitat is very restricted and by nature it
occurs in small stands among other plants. But recent de-
velopments within its range have caused further restriction.

In 1921, Harper (1922) reports, the bogs in which Sar-
racenia alabamensis grew were open, with woody plants
rather small and scattered, the bulk of the vegetation sun-
loving. It was still generally thus at both Elmore and in
Chilton County when we first saw this species in 1953 and

1974] Sarracenia — Case & Case 657

1955. At that time the swamps were shrubby but open,
and small rills lined with pitcher plants were frequent. The
entire area was sparsely inhabited, remote, and the roads
were limited, temporary, and unditched. In contrast, on
visits in the past few years, we found much change. The
bogs have not burned in some years. No longer are they
open ; dense stands of magnolia and maple, solid canebrakes,
tangles of Smilax, and alder, rob the bogs of light. The
sarracenia is much rarer today, and seedlings scarcely can
be found. Good roads reach the area, and farms, cattle
raising operations, use of herbicides, ditching and draining,
all have taken considerable toll of the habitat. Perhaps the
greatest change in the pitcher plant habitat here has been
the invasion of the bogs and openings by the Japanese
honeysuckle, Lonicera japonica Thunb., which destroys vir-
tually all native species in the bogs it invades.

DISCUSSION

The Alabama Canebrake Pitcher Plant has many features
of interest. It is the only trumpet leaved Sarracenia to
produce pitchers which differ significantly in size and form
with the season (Fig. 2).

The usually recurved, shorter, spring pitchers are less
voluminous than the often very much larger, erect, summer
pitchers. In these spring leaves, the laminar wings expand
at or just below the middle, pushing the pitcher into a
loosely sigmoid curve. The upper, sharply expanded part
of the tube and hood is erect. In effect, these early leaves
seem almost deformed, as in a plant growing under dras-
tically shaded conditions or in one which shows the effects
of an application of an herbicide. This leaf form, produced
in other species only under conditions of ecological duress,
is entirely normal and comprises the usual spring pitcher
in this species, even on plants growing in full sun (Fig. 2).
But occasionally, on extremely vigorous, uncrowded plants
in very wet, sunny situations, the plant produces the large
summer type pitchers without first producing the recurved
leaves.

658 Rhodora [Vol. 76

j2 328 is 15

ie i
z

4

4

Figure 2. Sarracenia alabamensis. Three spring type, curved
leaves (left) and three of the larger, straight summer leaves (right).
Note the obscure white, areole-like markings near summit of pitcher
tube on the right-hand specimen.

1974] Sarracenia — Case & Case 659

A single clone may produce either type of pitcher in the
first wave of growth in different years, but the trend is
clearly toward recurved spring pitchers. In a group of
about 50 study clones grown in our greenhouses, roughly
90% of the plants will produce the recurved pitchers in a
given season. In the wild, in more competitive (and per-
haps darker) situations all except two of several hundred
clones we have observed have produced the recurved spring
pitchers. These two plants had produced only the large
summer-type pitchers that season.

The variable form of the early leaves of the season, some-
times as recurved, reduced smallish pitchers with an un-
usual development of wing tissue, sometimes as fully de-
veloped phyllodia, and occasionally as erect, summer-type
pitchers, suggests to us that these variations occur at least
partly in response to a combination of degree of light in-
tensity and increasing daylength. Phyllodia and the re-
curved, reduced pitchers do not reappear in the autumn;
at that time only smaller, fully erect, summer-type pitchers
develop.

MacFarlane (1908) has stated that phyllodia represent
an advanced leaf type, developed by reduction of the su-
perior pitchered portion of the wing while the inferior
lamina remains full sized or enlarged. He notes that oc-
casional leaves intermediate between normal pitchers and
phyllodia appear in all the phyllodium-forming species. We
suggest that the variable structure of the spring leaves in
Sarracenia alabamensis represents a stage in the evolution
of a laminar phyllodium from a pitcher, one in which the
exact degree of pitcher reduction in response to daylength
and light intensity has not yet become stabilized. The fact
that these variable leaves occur only in spring and that in
other species which regularly produce phyllodia, such leaves
are strictly seasonal, seems to strengthen this interpreta-
tion.

The summer pitchers of the Canebrake Pitcherplant are
fully erect, remarkable for their relatively large size, and
great volume. The tissue of these large summer pitchers

660 Rhodora [Vol. 76

possesses a soft substance lacking the waxy-thickened qual-
ity of the other trumpet leaf species. This, together with
the fine, dense pubescence, gives to the pitcher a unique
"feel" prompting Dr. Wherry to remark that this is the
only “rubra type" Sarracenia one can readily identify in
the dark.

Other features of the summer pitcher include its very
light green color when grown in bright light, and the pres-
ence in many leaves of a few rather obscure, whitish areole-
like areas scattered on the outer pitcher-tube surface near
its orifice. These cannot be seen from within. These faint
areoles seem to be the consequence of the extreme tissue
expansion characteristic of the summer pitchers. They are
not structurally quite the same as those in the truly areolate
species, but they may offer interesting evidence as to the
mode of origin of areoles within this genus. Because all
taxa related to Sarracenia rubra have become controversial
and because other taxa in the rubra alliance do not show
areoles at all, some botanists may seize upon this feature
as evidence that S. alabamensis represents a hybrid popu-
lation of some sort, not a species.

We cannot regard these areole-like tissues as evidence of
any recent hybridization with an areolate species for sev-
eral reasons, The areolae do not appear on the spring
leaves in this taxon; in the known hybrids between other
areolate and non-areolate species, the areolae occur in all
leaves of mature plants and even on the larger seedling
leaves. In known areolate hybrids, the white transparent
areoles extend from outer to inner tube tissues. Also, no
areolate species occurs in contact with the known range of
this species. If these areole-like formations be of hybrid
origin, it is ancient hybridization and the whitish markings
have become a fixed although obscure characteristic of the
larger summer leaves.

Although we have already stated our interpretation of
the nature of the recurved spring pitchers above, the inter-
mittent production of full phyllodia in Sarracenia ala-
bamensis, coupled with Wherry's report of S. oreophila

1974] Sarracenia — Case & Case 661

within its range at Elmore, may lead some to say that S.
alabamensis represents a hybrid swarm of S. oreophila with
S. rubra. Several points ought to be made in this connec-
tion. Since S. oreophila is a large yellow-flowered species,
and S. rubra a small red-flowered one, any hybrid popula-
tion ought to produce orange, or yellow and red mottled
flowers of intermediate size as are reported for all other
hybrids between S. rubra and yellow flowered species. Not
one plant of our more than 50 study clones produces any-
thing but medium to dark oxblood red small flowers. S.
rubra Walter does not occur in the region of S. alabamensis.
The population of S. alabamensis is an extremely uniform
one in structure and behavior: even the variability of the
spring leaf structure and the variability in occurrence of
the areole-like markings on the larger summer leaves are
uniform features of the species. There is none of the type
of variation in leaf structure or coloration so characteristic
of Sarracenia populations where hybridization can be dem-
onstrated.

In an effort to ascertain whether this plant might repre-
sent a population of direct hybrid origin between Sarracenia
oreophila and S. rubra, we hand pollinated a plant of S.
rubra from Fruitdale, Ala., with pollen of S. oreophila from
De Soto Falls, Ala. The resulting hybrid plants, grown in
our greenhouse, were intermediate between the parents in
leaf structure and size and produced the orange-red flowers
typical of such hybrids. The hybrids did produce laminar
phyllodia, but, surprisingly, these were of an erect type as
in S. leucophylla and did not resemble closely those of either
S. oreophila or S. alabamensis. Leaf coloration patterns
and pitcher hood conformation were unlike S. alabamensis.
No recurved pitcher leaves of the S. alabamensis spring
pitcher type have appeared. We could not find justification
in the structure of these hybrids for suspecting a recent
hybrid origin for S. alabamensis involving these two possi-
ble parents.

The tendency toward phyllodia, the presence of very large
summer leaves resembling more in profile those of Sar-

662 Rhodora [Vol. 76

racenia flava, S. oreophila, or S. leucophylla than those of
S. rubra which it resembles in flower and spring leaf, and
the tendency to produce tissue areas resembling faint, ran-
dom areolae, could be interpreted as placing this species in
a central primitive position from which differing selective
factors have led in one direction toward the S. rubra types
and in another toward the phyllodium-producing groups.
Its presence as a relict in an area of ancient soil derivation
(Harper, 1922) in an area largely devoid of other pitcher
plants suggests to us that this is a possibility.

HISTORY

In the earliest published reference to this plant, R. M.
Harper (1918) referred the plant to Sarracenia sledgeii
MacF. [S. alata (Wood) Wood], although he was certainly
skeptical, for in his discussion of the distribution of S.
sledgeti he says: “East of Mobile Bay, I have seen a few
specimens [of true sledgeii] in the southern part of Bald-
win County, Alabama, and what appears to be the same
thing (though it may be a variety of the next [S. flava])
in boggy places among the long-leaf pine hills in Chilton
and Autauga Counties, near the center of the State.” At
that time Harper had not seen the plant in flower. In April,
1921, Harper found the flowers to be small and dark red, so
he identified the plants as S. rubra Walt., although his
locality was farther inland than the known distribution of
that species (Harper, 1922).

In 1932, Wherry, guided by Harper, visited some of these
colonies but stated: “We located a colony of a red flowered
pitcher plant but it was not in good enough condition to
establish its identity" (Wherry, 1934) [Possibly Wherry's
uncertainty was because of the presence of the peculiar,
recurved spring pitchers which simulate pitchers deformed
by shading]. In the September, 1973 issue of the Carnivo-
rous Plant Newsletter, Dr. Wherry refers to this plant
which he saw in our greenhouse in September 1955, after
having seen it with Harper in the wild, as an *undescribed
species from Alabama."

1974] Sarracenia — Case & Case 663

Wherry's discovery of Sarracenia oreophila at Elmore,
Ala. represents the only known contact of the range of S.
alabamensis with another species. At this station both
species apparently grew in proximity, for both have been
collected in summer leaf there and determined by various
taxonomists as S. oreophila. The difficulty of determining
certain summer plants of S. alabamensis from S. oreophila
is emphasized by the fact that Wherry's July 13, 1935,
Elmore, Ala. specimen (PENN), determined by both Wherry
and McDaniel as S. oreophila, is in our opinion S. ala-
bamensis. This is indicated by the small size of the flowers,
dark sepals, the shape and size of the phyllodia, and in this
particular specimen, smaller pitcher size than is typical of
flowering-sized S. oreophila (see Fig. 3). The difficulty of
separating all specimens of the two species without the
presence of petals to indicate flower color, is further pointed
up by McDaniel's comments (1966, pp. 45, 46) on the plants
of the Elmore area: “... the Harper collection from Elmore
County, Ala. differs in certain details of which the much
reduced phyllodia seem to be the most significant. Addi-
tional collecting both in Elmore County and in the lower
Coosa River Valley is needed to determine the status of this
central Alabama population."

Indeed, the dissimilar spring and summer pitchers on
well developed plants of Sarracenia alabamensis have led
to much historical confusion and to determinations of or
references to it in the literature in the past as S. sledgeii,
S. flava, S. jonesii, S. rubra, and S. oreophila. Yet those
who have seen vigorous living specimens agree it is distinct.

ACKNOWLEDGEMENTS

We wish to thank the curators of the herbaria whose
specimens we have studied. We are grateful to Dr. W. H.
Wagner, Jr., University of Michigan, and to Dr. James
Wells, Cranbrook Institute of Science, for their valuable
counsel and for securing the specimen loans for us at vari-
ous times in this study.

7
Rhodora [Vol. 76
ka 1 Scan NUS E i oo SN mm a. A@NVOOW A3NQN$ 430
s *$cót T PONE c i QU J ÉXisQ (aeuiwe,) WTTITTTIS VYITSTUTIVI
"wT W
‘pret zweu PRT Tt yoye woperaidəp Cog

è facea (ÉmAnwey) vipudoeio wrusdwicer

poc
uA sigh T
o iu oin

Figure 3. A specimen of Sarracenia alabamensis from Elmore,
Ala. showing the fully reduced laminar phyllodia occasionally pro-
duced in this species which, in combination with the large summer
leaves, give to this specimen a strong resemblance to S. oreophila.

1974] Sarracenia — Case & Case 665

To Dr. Rogers McVaugh, University of Michigan Herbar-
ium, we are especially grateful for examining our manu-
seript, for help with several difficult points, and for pre-
paring the Latin description.

To G. L. Burrows, IV, Mr. and Mrs. Charles F. Moore,
Mr. and Mrs. Bernard Klein, and to David Case go our
thanks for help in the field search for pitcher plant stations.

Over the years we have received special help from the
late Dr. R. M. Harper, Tuscaloosa, Ala., who furnished us
with transcripts of his field notes and who guided us in
1955, and Dr. Edgar T. Wherry, Philadelphia, Pa. No
words could express our gratitude to Dr. Wherry, nor
acknowledge adequately all of the assistance and kindness
he has extended to us.

LITERATURE CITED

BELL, C. R. 1949. A cytotaxonomic study of the Sarraceniaceae
of North America. Jour. Elisha Mitchell Sci. Soc. 65: 137-166.
pls. 8-14.

Harper, R. M. 1918. The American pitcher plants. Jour. Elisha
Mitchell Sci. Soc. 34: 110-125.

. 1922. Some pine-barren bogs in Central Alabama.
Torreya 22: 57-59.

MACFARLANE, J. M. 1908. Sarraceniaceae. In: A. ENGLER, Das
Pflanzenreich IV. 110(34): 1-38.

McDANIEL, S. T. 1966. A taxonomic revision of Sarracenia (Sar-
raceniaceae). Unpubl. Ph.D. Diss., Florida State Univ. (Avail-
able from Univ. Microfilms, Ann Arbor, Mich.)

Ripeway, R. 1912. Color standards and color nomenclature. A.
Hoen & Co., Baltimore.

Wuerry, E. T. 1929. Acidity relations of the sarracenias. Jour.
Wash. Acad. Sci. 19: 379-390.

. 1934. Exploring for plants in the southeastern states.
Sci. Monthly 38: 80-85.

1935. Distribution of the North American pitcher
plants. Im: M. V. WarcoTT, Illustrations of North American
pitcher plants. Smithsonian Inst., Washington, D. C.

1973. Reminiscences on carnivorous plants. Carnivo-
rous Plant Newsl. 2: 35, 36.

7275 THORNAPPLE LANE,
SAGINAW, MICHIGAN 48603

666 Rhodora [Vol. 76

INSTRUCTIONS FOR CONTRIBUTORS TO RHODORA

Manuscripts should be submitted in duplicate and should
be double-spaced or preferably triple-spaced (not on cor-
rasable bond), and a list of legends for figures and maps
provided on a separate page. Footnotes should be used
sparingly, as they are usually not necessary. Do not indi-
cate the style of type through the use of capitals or under-
scoring, particularly in the citations of specimens, except
that the names of species and genera may be underlined to
indicate italics in discussions. Specimen citations should
be selected critically especially for common species of broad
distribution. Systematic revisions and similar papers should
be prepared in the format of “The Systematics and Ecology
of Poison-Ivy and the Poison-Oaks," W. T. Gillis, Rhodora
73: 161-237, 370-443. 1971, particularly with reference to
the indentation of keys and synonyms, Papers of a floristic
nature should follow, as far as possible, the format of
“Contribution to the Fungus Flora of Northeastern North
America. V.," H. E. Bigelow & M. E. Barr, Rhodora 71: 177-
203. 1969. For bibliographic citations, a recommended list
of standard journal abbreviations is given by L. Schwarten
& H. W. Rickett, Bull. Torrey Bot. Club 85: 277-300. 1958.

Volume 76, No. 808, including pages 497-680, was issued Feb. 28, 1975.

Dodova

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Conducted and published for the Club, by
ALFRED LINN BOGLE, Editor-in-Chief

ROLLA MILTON TRYON )
STEPHEN ALAN SPONGBERG
Associate Editors
GERALD JOSEPH GASTONY
RICHARD EDWIN WEAVER J
VOLUME 76

1974

The Neu Lngland Botanical Club, Ine.

Botanical Museum, Oxford St., Cambridge, Mass. 02138

INDEX TO VOLUME 76

Entries include: authors; title words such as geographic area; and
new records, systematic revisions, or ecological studies by taxa. Lists
or tables of taxa and maps are also classified separately under “Lists”
and “Maps”. Note that only one page number per article is given for
each taxon entered. New scientific names and combinations are printed

in bold face type.

Adoxa moschatellina 43

Alabama, Sarracenia alabamensis,
A Newly Recognized Species
From Central 650-665

Algal Ecology, The Role Of Some
Haloragaceae In 446-459

Alternanthera brasiliana 476

Anaphalis margaritacea 44

Andrus, Richard E., Significant
New Distributional Records For
The Genus Sphagnum In The
Northeastern United States 511-
518

Antirrhinum orontium 467

Arkansas, Lindera melissaefolium
In 525

Ascophyllum mackaii 408; nodo-
sum 59-63

Asiatic Species Of Desmodium: A
Review 142-144

Asparagopsis hamifera 410

Asperococcus echinatus 407

Authority For Cardamine clema-
tis (Cruciferae), The Correct
53-57

Averett, John E., Typification of
Chamaesaracha coronopus 311

Bahama Flora, Name Changes
For The Seed Plants In The
67-138

Bahama Flora, Range Extensions
And New Records For The 471-
ATT

Bald Cypress (Taxodium disti-
chum (L.) Richard) In Suffolk
County, Long Island, New
York, The Occurrence Of 25-26

Baldwin, Henry I., The Flora of
Mount Monadnock, New Hamp-
shire 205-228

Barkley, T. M. and Arthur Cron-
quist, Cacalia rugelia: A New
Combination For A North
American Senecionoid 48-50

Bauhinia variegata 476

Bidens beckii 44

Biosystematic Revision Of Lupi-
nus lyallii 422-445

Black Spruce, Environmental Con-
trol Of Needle Characteristics
In Subalpine 519-524

Blanchard, Orland J., Chromosome
Numbers In Kosteletzkya Presl
(Malvaceae) 64-66

Bogle, A. Linn (see Lane, David

M.)
Bollard, Peter M. (see Hehre,
Edward J.)

Bryophytes of Southern Manitoba,
Studies On The III. Collections
From Grand Beach Provincial
Park 27-38, list 29-34

Cacalia rugelia: A New Combina-
tion For A North American
Senecionoid 48-50

Calamagrostis lapponica 40

669

670

Caltha natans 42

Campanula rotundifolia var. velu-
tina 468

Cape Cod, Massachusetts, Investi-
gations Of New England Ma-
rine Algae VI: Distribution Of
Marine Algae Near 537-563

Capsosiphon fulvescens 401

Cardamine clematis (Cruciferae),
The Correct Authority For 53-
57

Carex adusta 40; exilis, Geocaulon
lividum And Other Plants Of
Interest In Pittsburg, New
Hampshire 307-309; rufina 41

Carya pallida 462

Case, Frederick W. and Roberta
B. Case, Sarracenia alabamen-
sis, A Newly Recognized Spe-

cies From Central Alabama
650-666
Case, Roberta B. (see Case,
Frederick W.)

Catling, Paul M. and James E.
Cruise, Spiranthes casei, A New
Species From Northeastern
North America 526-536

Central Plains States, New and
Interesting Plants From The
489-490

Chabot, Brian F., Environmental
Control Of Needle Character-
istics In Subalpine Black Spruce
519-524

Chamaesaracha coronopus, Typi-
fication Of 311

Chenopodium 462; bushianum
463; pallescens 463; strictum
var. glaucophyllum 463-464

Chromosome Numbers In Koste-
letzkya Presl. (Malvaceae) 64-
66

Cladophora laetevirens 403

Coleman, Douglas C. and Arthur
C. Mathieson, Investigations Of

Rhodora

[Vol. 76

New England Marine Algae VI:
Distribution of Marine Algae
Near Cape Cod, Massachusetts
587-563

Color-Forms From
Maine, Two New 470

Colt, L. C., Jr. and C. Barre Hell-
quist, The Role Of Some Halo-
ragaceae In Algal Ecology
446-459

Combination For North Ameri-
can Senecionoid, Cacalia ru-
gelia: A New 48-50

Conrad, Melvin L., Panicum ensi-
folium Baldw. — New For
Mexico 509-510

Corallorhiza wisteriana 489

Cox, B. J., A Biosystematic Revi-
sion Of Lupinus lyallii 422-445

Crataegus marshallii 465

Crinum asiaticum 476

Cronquist, Arthur (see T. M.
Barkley)

Crotalarias Of North America
(Leguminosae), A Systematic
Treatment Of The Native Uni-
foliate 151-204; Crotalaria bre-
vipedunculata 164-166, fig. 3,
map 166; bupleurifolia 194-198,
map 197; var. bupleurifolia
198, fig. 11; var. robusta 198,
fig. 12, map 197; mexicana 184-
186, fig. 9, map 166; nayari-
tensis 198-200, fig. 13, map 197;
nitens 161-164, map 159; var.
nitens 164, fig, 2; var. gracilis
164, fig. 2, map 159; pilosa
157-161, fig. 1; map 159; poly-
phylla 166-169, fig. 4, map 166;
purshii 177-181, fig. 7, map 179;
quercetorum 181-184, fig. 8,
map 166; rotundifolia 186-194;
X purshii 202; var. rotundi-
folia 189-193, fig. 10, map 191;
var. vulgaris 193-194, fig. 10,

Southern

1974]

map 192; sagittalis 172-177,
fig. 6, map 174; XX quercetorum
202; X rotundifolia var. vul-
garis 202; X stipularia 202;
stipularia 169-172, fig. 5, map
159

Cruise, James E. (see Catling,
Paul M.)

Cucurbita pepo var. ovifera 468

Datura inoxia 476

Desmodium: A Review, The
Asiatic Species Of 142-144

Digitaria sanguinalis, A New
Form; f. illinoensis 57-58

Distributional Records For The
Genus Sphagnum In The North-
eastern United States, Signifi-
cant New 511-518

Draba reptans var. micrantha 464;
verna var. boerhaavii 464

Drosera From Venezuela, A New;
felix 491-493, fig. 1

Dudley, T. R., The Correct Au-
thority For Cardamine clematis
(Cruciferae) 53-57

Ebinger, John E., A New Form
Of Digitaria sanguinalis 57-58;
A Systematic Study Of The
Genus Kalmia (Ericaceae) 315-
398

Enteromorpha ahlneriana 401;
groenlandica 402
Environmental Control Of

Needle Characteristics In Sub-
alpine Black Spruce 519-524
Eryngium prostratum var. pros-

tratum 490
Euphrasia arctica var. dolosa 43
Evans, Dan K. (see Mohlenbrock,
Robert H.)

Fissidens adianthoides 32

Index to Volume 76

671

Flora Of Mount Monadnock, New
Hampshire, The 205-228, list
211-221

Florida, A. New Species Of Lechea
(Cistaceae) From Peninsular
478-483

Fogg, John M., Jr., Stone’s Plants
Of Southern New Jersey 139-
141

Form Of Digitaria sanguinalis, A
New 57-58

Fucus vesiculosus 59-63

Fuirena scirpoidea 460

Geocaulon lividum And Other
Plants Of Interest In Pittsburg,
New Hampshire, Carex exilis
307-309

Gillis, William T., Name Changes
For The Seed Plants In The
Bahama Flora 67-138

Grindelia lanceolata f. latifolia
469

Growth Of Some New England
Perennial Seaweeds 59-63

Habenaria orbiculata 41
Haloragaceae In Algal Ecology,
The Role Of Some 446-459
Harman, Willard N., Phenology
and Physiognomy Of The Hy-
drophyte Community In Otsego
Lake, N.Y. 497-508

Harms, Vernon L. and John H.
Hudson, Some New Or Note-
worthy Vascular Plant Records
From Northwestern Saskatche-
wan 39-44

Harriman, Neil A., Pimpinella
saxifraga L. (Umbelliferae) In
Wisconsin 483

Haynes, Robert R., A Revision Of
North American Potamogeton
Subsection Pusilli (Potamoge-
tonaceae) 564-649

672 Rhodora

Hehre, Edward J. and Peter M.
Bollard, The Occurrence Of
Bald Cypress (Taxodium dis-
tichum (L.) Richard) In Suf-
folk County, Long Island, New
York 25-26

Hellquist, C. Barre, A White-
Flowered Form Of Utricularia
purpurea In New Hampshire
19; (see Colt, L. C., Jr.)

Hepper, F. N., Parasitic Witch-
weed: Striga asiatica Versus S.
lutea (Scrophulariaceae) 45-47

Hill, Steven R., Range Extensions
And New Records For The Ba-
hama Flora 471-477

Hodgdon, A. R., Carex exilis,
Geocaulon lividum And Other
Plants Of Interest In Pittsburg,
New Hampshire 307-309

Holosteum umbellatum 489

Hudson, John H. (see Harms,
Vernon L.)

Hydrophyte Community In Ot-
sego Lake, N.Y. Phenology
And Physiognomy Of The 497-
508

Illinois Field and Herbarium
Studies 460-470

Jacquemontia tamnifolia 466

Kalmia (Ericaceae), A Syste-
matic Study Of The Genus
315-398; angustifolia 367-381;
var. angustifolia 368-378; f.
candida 377-378; var. carolini-
ana 378-381; cuneata 381-383;
ericoides 386-390; var. ericoides
388-389; var. aggregata 389-
390; hirsuta 383-386; latifolia
352-366; f. angustata 363; f.
fuscata 363-364; f. myrtifolia
364; f. obtusata 365; f. poly-

[Vol. 76

petala 365-366; microphylla
333-342; var. microphylla 334-
340; var. occidentalis 340-342;
f. alba 342; polifolia 343-351;
f. leucantha 351

Kitfield, Aminta, Range Exten-
sion Of Woodwardia areolata
312

Kosteletzkya presl (Malvaceae),
Chromosome Numbers In 64-66,
table 66

Lactuca saligna f. ruppiana 490

Lagenaria siceraria 468

Laminaria digitata, 59-63; sac-
charina 59-63

Lane, David M. and A. Linn
Bogle, A Second Report Of The
Prothallia Of Lycopodium in-
undatum In North America
484-488

Lechea (Cistaceae) From Penin-
sular Florida, A New Species
Of; lakelae 478-483

Lindera melissaefolium In Ar-
kansas 525

Lists (Tables): Bryophtes From
Grand Beach Provincial Park
29-34; Chromosome Counts In
Kosteletzkya 66; Chromosome
Numbers In Perityle Section
Perityle 236; Distribution Of
Marine Algae Near Cape Cod
553-558; Flora Of Mount Mo-
nadnock 211-221; Marine Algae
Of Rhode Island 400-416; Name
Changes In The Bahama Flora
10-124; Range Extensions And
New Records For The Bahama
Flora 473-477

Lithophyllum lenormandi 411

Lobelia dortmanna 43

Longleaf Pine, On The Scientific
Name Of 20-24

Lophocolea heterophylla 29

1974]

Lóve, Áskell, A Magnificent Swiss
Flora Manual: A Review

Lupinus lyallii, A. Biosystematic
Revision Of 422-445, map 444;
subsp. lyallii 426-435; var.
lyallii 426-429; var. danaus
429-431; var. fruticulosus 431-
432; var. macroflorus 432-4535;
var. roguensis 433-435; var.
villosus 435-436; subsp. alcis-
temporis 436-438; subsp. mi-
nutifolius 438-439; subsp. sub-
pandens 439-442; subsp. washo-
ensis 442-443

Lycopodium inundatum In North
America, A Second Report Of
The Prothallia Of 484-488, fig.
3, 4, map 488

Magrath, Lawrence K. and Ronald
R. Weedon, New And Interest-
ing Plants From The Central
Plains States 489-490

Maine, Two New Color-Forms
From Southern 470

Malphigia glabra 476

Manitoba, Studies On The Bry-
ophytes Of Southern III. Col-
lections From Grand Beach
Provincial Park 27-38

Maps: Crotalaria spp. 159, 166,
174, 179, 191, 192, 197; Lupinus
lyallii 444; Lycopodium inun-
datum In New England 488;
Marine Algae Stations On Cape
Cod 539; Perityle Section Peri-
tyle spp. 251, 257, 267, 281, 285,
300; Potamogeton Section Pu-
silli spp. 596, 602, 606, 611,
617, 623, 625, 630, 635, 639,
641; Sphagnum spp. In North-
eastern United States 512;
Spiranthes casei 534; Xanthis-
ma texanum 3, 5

Index to Volume 76

613

Marine Algae Near Cape Cod,
Massachusetts, Distribution Of,
Investigations Of New England
Marine Algae VI: 537-563, list
553-558

Marine Algae Of Rhode Island
399-421, list 400-416

Matelea decipiens 467

Mathieson, Arthur C. (see Cole-
man, Douglas C.)

Mentzelia decapetala 465-466

Mexico, Panicum ensifolium
Balw. — New For 509-510

Mohlenbrock, Robert H. and Dan
K. Evans, Mlinois Field And
Herbarium Studies 460-470

Monostroma leptodermum 402

Mount Monadnock, New Hamp-
shire, The Flora Of 205-228

Myriophyllum alterniflorum 43

Name Changes For The Seed
Plants In The Bahama Flora
67-138, list 70-124

New And Interesting Plants From
The Central Plains States 489-
490

New England Perennial Sea-
weeds, The Growth Of Some
59-63

New Hampshire, A White-Flow-
ered Form Of Utricularia pur-
purea From 19

New Hampshire, Carex exilis,
Geocaulon lividum And Other
Plants Of Interest In Pittsburg
307-309

New Hampshire, New Station
For Nuphar microphyllum In
Durham 310

New Hampshire, The Flora Of
Mount Monadnock 205-228

New Jersey: A Reprint, Stone’s
Plants Of 139-141

674

New York, The Occurrence Of
Bald Cypress (Taxodium dis-
tichum (L.) Richard) In Suffolk
County, Long Island 25-26

Newly Recognized Species From
Central Alabama, Sarracenia
alabamensis, A 650-665

North America, A Second Report
Of The Prothallia Of Lycopo-
dium inundatum In 484-488

North America (Leguminosae), A
Systematic Treatment Of The
Native Unifoliolate Crotalarias
Of 151-204

North American Potamogeton
Subsection Pusilli, A Revision
Of 564-649

Northeastern North America,
Spiranthes casei, A New Species
From 526-536

Northeastern United States,
Significant New Distributional
Records For The Genus Sphag-
num In The 511-518

Nuphar microphyllum In Dur-
ham, New Hampshire, New
Station For 310

Opuntia rubescens 476

Otsego Lake, N.Y., Phenology
And Physiognomy Of The
Hydrophyte Community In 497-
508

Panicum ensifolium Baldw. —
New For Mexico 509-510
Penstemon cobaea 467-468
Percursaria percursa 402
Perityle Section Perityle (Com-
positae-Peritylinae), Taxonomy
Of 229-309, table 236; aglossa
289-291, map 285; aurea 262-
264, map 251; californica 298-
302, map 300; canescens 275-
276; ciliata 276-277, map 267;

Rhodora

[Vol. 76

cordifolia 291-292, map 285;
coronopifolia 277-279, map 267;
crassifolia 248-253; var. crassi-
folia 248-250, map 251; var.
robusta 250-253, map 251; cu-
neata 302-305; var. cuneata 302-
305, map 300; var, marginata
304-305, map 300; emoryi 255-
262, map 257; feddemae 283-
284, map 281; hofmeisteria 279-
280, map 267; incana 264-266;
jaliscana 280-282, map 281;
leptoglossa 292-294, map 285;
lineariloba 268-269, map 267;
lloydii 294-295, map 285; lobata
295-296, map 285; microcephala
269-270, map 267; microglossa
270-275; var. microglossa 270-
273, map 267; var. saxosa 273-
275, map 267; palmeri 296-298,
map 285; parryi 284-288, map
285; rosei 282-283, map 281;
socorrensis 253-255; trichodonta
283, map 281; turneri 266-268,
map 267; vaseyi 288-289, map
285

Phenology And Physiognomy Of
The Hydrophyte Community In
Otsego Lake, N.Y. 497-508

Phleum pratense f. viviparum
460

Physalis macrophysa 467

Phytogeography And Systematies
Of Xanthisma texanum DC.
(Asteraceae): Proper Usage Of
Interspecific Categories 1-19

Picea mariana 519-524

Pimpinella saxifraga L. (Umbelli-
ferae) In Wisconsin 483

Pinus palustris 20-24

Poa bulbosa 460

Polygonum ramosissimum f. alba
470

Polypremum procumbens 466

Populus X smithii 461-462

1974]

Porphyra miniata 409

Potamogeton Subsection Pusilli
(Potamogetonaceae), A Revi-
sion Of North American 564-
649; clystocarpus 624-626, map
625; foliosus 612-624; var.
foliosus 614-620, map 617; var.
fibrillosus 620-624, map 623;
friesii 594-598, fig. 1, map 596;
groenlandicus 640-642, map 641;
hillii 607-612, map 611; obtusi-
folius 604-607, map 606; pusil-
lus 626-640, fig. 1; var. pusillus
629-633, map 630; var. gemmi-
parus 638-640, map 639; var.
tenuissimus 633-637, map 635;
X perfoliatus 638; strictifolius
598-603, map 602; X zosteri-
formis 604

Powell, A. Michael, Taxonomy Of
Perityle Section Perityle (Com-
positae-Peritylinae) 229-306

Prothallia Of Lycopodium inun-
datum In North America, A
Second Report Of The 484-488

Pyrus pyrifolia 464-465

Quercus velutina f. missouriensis
462

Range Extension Of Woodwardia
areolata 312

Range Extensions And New Rec-
ords For The Bahama Flora
471-477, list 473-477

Ranunculus hyperboreus 42; tes-
ticulatus 490

Records From Northwestern Sas-
katchewan, Some New Or Note-
worthy Vascular Plant 39-44

Reviews: Asiatic Species Of Des-
modium 142-144; Stone’s Plants
Of Southern New Jersey 139-
141; Swiss Flora Manual 145-
149

Index to Volume 76

675

Revision Of North American
Potamogeton Subsection Pusilli
(Potamogetonaceae) 564-649

Reynolds, Norman B., The Growth
Of Some New England Peren-
nial Seaweeds 59-63

Rhizoclonium tortuosum 404

Rhode Island, Marine Algae Of
399-421

Rhodymenia palmata 59-63

Salix X myricoides 461

Sargassum natans 409

Sarracenia alabamensis, A Newly
Recognized Species From Cen-
tral Alabama 650-665

Saskatchewan, Some New Or
Noteworthy Vascular Plant
Records From Northwestern
39-44

Schubert, Bernice G., The Asiatic
Species Of Desmodium: A Re-
view 142-144

Scirpus americanus 51-52; olneyi
51-52; pungens 51-52

Scleranthus annuus 489

Seed Plants In The Bahama
Flora, Name Changes For The
67-138

Semple, John C., The Phytogeog-
raphy And Systematics Of Xan-
thisma texanum DC. (Astera-
ceae) 1-19

Senecio jacobaea 469

Sherardia arvensis 468

Smith, L. B. (see Steyermark,
J, A2)

Sparganium chlorocarpum 39

Spergula arvensis 42; media 464

Sphagnum In The Northeastern
United States, Significant New
Distributional Records For The
Genus 511-518, map 512; henry-
ense 511-513; jensenii 516; por-
toricense 513-514; pulchrum

676

515-516; riparium 515; strictum
514; subfulvum 516-517; tenel-
lum 516; torreyanum 514

Spiraea latifolia f. rosea 470

Spiranthes caseii, A New Species
From Northeastern North
America 526-536, fig. 1, 2, map
534; gracilis 41-42

Spongomorpha lanosa 404; spines-
cens 404

Steyermark, J. A. and L. B.
Smith, A New Drosera From
Venezuela 491-493

Stone's Plants Of Southern New
Jersey: A Reprint 139-141

Striga asiatica Versus S. lutea

(Scrophulariaceae), Parasitic
Witchweed 45-47
Stringer, Muriel H. L. (see

Stringer, Paul W.)

Stringer, Paul W. and Muriel H.
L. Stringer, Studies On The
Bryophytes Of Southern Mani-
toba III. Collections From
Grand Beach Provincial Park
277-38

Suaeda depressa 464

Swiss Flora Manual, A Review,
A Magnificent 145-149

Systematies Of Xanthisma texa-
num DC. (Asteraceae): Proper
Usage Of Intraspecific Cate-
gories. The Phytogeography
And 1-19

Systematic Study Of The Genus
Kalmia (Ericaceae), A 315-398

Taxodium distichum 25-26

Taxonomy Of Perityle Section
Perityle (Compositae-Perityli-
nae) 229-306

Thelesperma gracile 469

Thlaspi perfoliatum 490

Trailliella intricata 415

Triadenum virginicum 465

Rhodora

[Vol. 76

Tucker, G. E., Lindera melissae-
folium In Arkansas 525

Typification And Application Of
The Names Scirpus americanus
Pers., S. olneyi Gray, And S.
pungens Vahl 51-52

Typification Of Chamaesaracha
coronopus 311

Utricularia purpurea From New
Hampshire, A White-Flowered
Form Of; f. alba 19

Venezuela, A New Drosera From
491-493

Villalard-Bohnsack, Martine (see
Wood, R. D.)

Vinca major 466

Ward, Daniel B., On The Scientific
Name Of The Longleaf Pine
20-24

Weedon, Ronald R. (see Magrath,
Lawrence K.)

Wells, Theodore W., Two New
Color-Forms From Southern
Maine 470

Wicks, Johonet C., New Station
For Nuphar microphyllum In
Durham, New Hampshire 310

Wilbur, Robert L., A New Species
Of Lechea (Cistaceae) From
Peninsular Florida 478-483

Windler, Donald R., A Systematic
Treatment Of The Native Uni-
foliolate Crotalarias Of North
America (Leguminosae) 151-
204

Wisconsin, Pimpinella saxifraga
L. (Umbelliferae) In 483

Witchweed: Striga asiatica Ver-
sus S. lutea (Scrophulariaceae)
Parasitic 45-47

1974] Index to Volume 76 677

Wood, R. D. and Martine Villa- geography And Systematics Of
lard-Bohnsack, Marine Algae 1-19, fig. 3, map 3; subsp. texa-
Of Rhode Island 399-421 num 14; var. texanum 14; var.

Woodwardia areolata, Range Ex- orientale 14-16; f. orientale 15;
tension Of 312 f. rubrum 15-16; subsp. drum-

mondii 16

Xanthisma texanum DC. (Astera-
ceae): Proper Usage Of Infra- Zephyranthes citrina 476; puerto-
specific Categories, The Phyto- ricensis 476

STATEMENT OF OWNERSHIP, MANAGEMENT AND

CIRCULATION
Title of Publication: RHODORA
Date of Filing: September 27, 1974
Frequency of Issue: Quarterly

Location of Known Office of Publication:
Botanical Museum, Harvard University, Oxford Street,
Cambridge, Mass. 02138

Location of the Headquarters of the Publishers:
Botanical Museum, Harvard University, Oxford Street,
Cambridge, Mass. 02138

Publisher:

New England Botanical Club, Inc., Botanical Museum,

Harvard University, Cambridge, Mass. 02138

Editor:
Dr. Alfred L. Bogle, Dept. Botany,
University of New Hampshire, N.H. 03824

Managing Editor:
Dr. Herman R. Sweet, Botanical Museum,
Harvard University, Cambridge, Mass. 02138

Owner: New England Botanical Club, Inc., Botanical Museum, Oxford

Street, Cambridge, Mass. 02138
No stockholders

Known Bondholders, Mortgagees, and other Security Holders: None

Average No. Copies
each issue during

preceding 12 months

Total No. copies printed 850
Paid Circulation

Sales through dealers, etc. none

Mail subscriptions 778
Total paid circulation 778
Free distribution 12
Total distribution 790
Office use, left-over, etc. 60
Total 850

Single issue
nearest to
filing date

850

none
785
785
12
797
53
850

I certify that the statements made by me above are correct and
complete. Herman R. Sweet, Business Manager and Treasurer.

ui

175

CONTENTS: — continued

Lindera Melissaefolium in Arkansas
CORE MEME I ears acta a a ecg I IU 525

Spiranthes casei, a New Species from Northeastern North
America
Paul M. Catling and James E. Cruise .............. eee 526

Investigations of New England Marine Algae VI: Distribu-
tion of Marine Algae near Cape Cod, Massachusetts

Douglas C. Coleman and Arthur C. Mathieson ................ 537

A Revision of North American Potamogeton Subsection
Pusilli (Potamogetonaceae)

BE SEET a AEE EI A ett reno nnt entente pn e N I A 564

Sarracenia alabamensis, a Newly Recognized Species from
Central Alabama
Frederick W. Case, Jr., and Roberta B. Case .................. 650

Instructions for Contributors ................. eene 666