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Reprinted from Ruovora, Vol. 45, March—-May, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 
——— 


CXLVII 


Pt a 


Poel —/o oO 


i 


THE BROAD-LEAVED SPECIES OF POTAMOGETON 
OF NORTH AMERICA NORTH OF MEXICO 


i. C. Ogpren 

Dares or Issun 
Pages 57-108 and Plates 746-748... gol ci 18 March, 1943 
SAT ei a ee ee ..10 April, 1943 


ROMO ROR E: oo ek eae ede oe a ees 1 May, 1943 © : 


Rhodora Plate 746 


Fruits or PotaMoGEtTon, X 5 


YUS Var. TENIFOLIUS, figs. 1-3; P. ALPINUS var. SUBELLIPTICUS, figs. 4—6; 

ig POLYGONIFOLIUS, figs. 7-9; peciteg ae gy 10-13; P. oe figs. 14-17; 

P. noposus, figs. 18-21; P. xa ANS, figs. 29-25; P. Oake ANUS, figs. 26- 29: 

. ILLINOENSIS, figs 30-- Sia og alanine figs. 36-38; eB PRAELONG us, figs. 

39 and a r; Ri ICH: ARDSONII, figs. 41-46; P. PERFOLIATUS Var. BUPLEUROIDES, 
47-5 


Reprinted from Ruopora, Vol. 45, March—May, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


CXLVII 


THE BROAD-LEAVED SPECIES OF POTAMOGETON 
OF NORTH AMERICA NORTH OF MEXICO 


EK. C. OGDEN 


Dates oF IssuUE 
Pages 57-105 and Plates 746-748... .....-- 5... c cee eee ees 18 March, 1943 
Pages TT re re ia: «hao ce ea ee 10 April, 1943 
Pages in CA ee Ren pt a piet aha vi kw 3 8 1 May, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM OF 
HARVARD UNIVERSITY—NO. CXLVII 


THE BROAD-LEAVED SPECIES OF POTAMOGETON OF 
NORTH AMERICA NORTH OF MEXICO 


E. C. OapEn 
(Plates 746-748) 


THE genus Potamogeton is of world-wide distribution. Be- 
cause it exhibits a great range of environmental variation, and 
specimens so often lack the fundamental diagnostic parts which 
might ally them with already known entities, the names that 
have been coined for its members are legion. 

Although a number of monographic treatments have dealt 
with the North American species in whole or in part’, the broad- 
leaved groups have been poorly understood. The really usable 
treatment of the troublesome linear-leaved species by Fernald 
lent new hope to the idea that possibly the broad-leaved plants 

1 Thomas Morong, The Naiadaceae of North America, Mem. Torr. Club 3: no, 2 
(1893). 

P. Ascherson and P. Graebner, = Engler, Pflanzenr. 4: fam. 11, Potamogetonaceae 
goign by Graebner) (190 

n Taylor, in N. Am. FI. ay, ‘pt. 1 (1909). 

: 0. Ha hone: Critical Re setae on the Potamogetons, Kungl. Svenska Veten- 
skapsakad. Handl. 55: no. 5 (1916 

Harold St. John, A Revision of ee North American Species of Potamogeton of the 
Section Coleophyl Ruopora, 18: 121-138 (1916). 

ald, The Linear-leaved North American Species of Potamogeton, Section 
Axillares, “Mem, Amer. Acad, Arts & Sci. 17: pt. 1 (1932). _As this work contains an 


la 


f th WOU Tit erely 


be repetitive and is unnecessary here. 


58 Rhodora [Marcu 


might be resolved into something approaching an expression of 
their true relationships. 

A fertile Potamogeton seldom causes great difficulty; but the 
great mass of sterile material, displaying a range of form un- 
known in the fertile plants, has made an understanding of the 
group difficult. To add to the confusion, the broad-leaved 
species, with their spikes buoyed up above the water, hybridize 
more or less freely, rendering it. difficult in many cases to de- 
termine the fundamental cause behind the abnormality. . 

The following treatment is offered in the hope that the species 
here considered may be better understood; and that specimens 
may be properly identified, even though they lack many of the 
important diagnostic characters and exhibit forms that are far 
from typical. 


DIAGNOSTIC CHARACTERS 


Fruits. When mature fruits are present there is seldom any 
difficulty in referring the specimen to the proper species. Un- 
fortunately the presence of fruit is not the rule but the less 
common occurrence. Most distinctive are those of P. alpinus, 
with a smooth, hard, eggshell-like exocarp, tawny-olive in color; 
fruiting specimens can never be mistaken for any other species. 
Neither can the minute reddish orbicular fruits of P. polygoni- 
folius with their beaks practically obsolete, nor the massive 
fruits of P. praelongus be confused with those of any other North 
American broad-leaved species. The long-beaked fruits of P. 
crispus are especially characteristic. Extremely important 
diagnostic characters are the prominence of keels and the color 
of the exocarp. If one makes a longitudinal section with a 
razor-blade, the amount of curvature of the seed becomes ap- 
parent and is of some slight diagnostic value. More important 
in some cases, and surprisingly consistent, is the presence or 
absence of a cavity in the endocarp tissue that projects as a fold 
into the center of the fruit. This projection (endocarp loop) is 
solid in all the broad-leaved species except P. polygonifolius, 
gh praelongus and P. Richardsonii. This is an important char- 
acter in the separation of P. Richardsonii from P. perfoliatus, 
specimens of which often strongly resemble each other. Because 
the fruits are of great taxonomic importance, but difficult ade- 


1943] Ogden,—The Broad-leaved Species of Potamogeton 59 


quately to describe, it has seemed desirable to illustrate them 
(see PLATE 746). 

FLowers. It is now generally considered that Potamogeton 
has no true perianth. The sepal-like structures are thought to 
be outgrowths from the connectives of the stamens, and are 
called sepaloid connectives.! In the species here treated they are 
of little or no diagnostic value. Careful observations on the 
shape and hundreds of measurements on the width and length 
of claw have been made, from which it has become evident that 
even species which in most characters are strikingly different 
have sepaloid connectives which are surprisingly similar. The 
anthers, also, are of about the same shape and size in all the 
broad-leaved species, except that in P. polygonifolius they 
average somewhat smaller. Pollen-grains should be examined 
when hybridism is suspected. Fertile species have pollen-grains 
well formed and abundantly produced, whereas hybrids invari- 
ably have a large percentage aborted. 

Spikes. While the flowering spikes are of little diagnostic 
value, differing chiefly in average size and in number and degree 
of crowding of the whorls of flowers, the diameter of the fruiting 
spikes can be used. This thickness is almost entirely dependent 
on the length of the fruits, but in many cases the fruit-measure- 
ments cannot be so easily obtained. While there are some dif- 
ferences as to length of the spikes of different species, these are 
mostly a direct correlation with the robustness of the species, 
and their diagnostic value is therefore secondary. 

PrepuncLes. Characters of the peduncle can usually not be 
used for the determination of broad-leaved species without 
supporting characters from other parts of the plant, due to the 
great variation within the same species, but they are sometimes 
of useful supplementary value. Most species seldom have 
peduncles over 10 or 15 em. long, but in P. amplifolius they may 
run up to 30 em. and it can then be distinguished from P. pulcher 
or P. nodosus, which at times it may superficially resemble. 
Similarly, P. Richardsonii may have peduncles up to 25 cm. in 
length, in contrast to its nearest relative, P. perfoliatus, with 
peduncles less than 10 em. long. The longest peduncles are 


t normal 


1 Phylogenists are not agreed on 
perianth with the anthers sessile on con claws of the segments, 


60 Rhodora [Marca 


found on P. praelongus, where they are often more than 30 cm. 
and may attain a length of 60 cm. The three species mentioned 
as sometimes having long peduncles (P. amplifolius, P. prae- 
longus, and P. Richardsonii) may also have them short; but 
whether long or short, usually with a tendency to be clavate. 
This tendency for the peduncles to be thickened upward is also 
somewhat characteristic of the subsection Lucentes. Most of 
the species here treated have their peduncles rather uniform in 
thickness throughout their length. Another character of slight 
value is the thickness of a peduncle in comparison with the 
thickness of the stem at the internode directly below. This has — 
some supplementary value in the subsection Natantes, where P. 
natans has peduncles about the same thickness as the stem, 
while P. Oakesianus, with its slenderer stem, has less reduction 
in the thickness of its peduncles, and as a consequence has them 
thicker than the internode below. Lastly, there are some dif- 
ferences in their internal anatomy. These are, however, few 
and inconstant and not used in this treatment. 

Stipuues. In all of the species here treated (with the excep- 
tion of P. crispus) the stipules are free from the leaves. They 
clasp the stem, but have their margins absolutely free and un- 
united. They are all obtuse, at least when young, but as there 
is often a tendency for them to be somewhat cucullate they may, 
at times, appear to be acutish. This is especially true of P. 
nodosus. All are more or less two-keeled (when mature) except 
in the subsections Crispi, Colorati, Praelongi, and Perfoliatt. 
The chief differences are in their length and coarseness, which 
are, for the most part, rather direct correlations with the robust- 
ness of the species. They furnish useful characters in the 
perfoliate-leaved species where one can usually recognize P. 
praelongus by its large white stipules, P. Richardsonii by its 
stipules reduced to strong white fibers, and P. perfoliatus with 
stipules delicate and inconspicuous. 

Leaves. Certainly some of the most important diagnostic 
characters for the identification of the broad-leaved species of 
Potamogeton are to be found in the leaves, for too often all that 
are available are stems and either submersed or floating leaves, 
or both. At least three fourths of the specimens in the herbaria 
lack not only mature fruits but any part of an inflorescence. 


1943] Ogden,—The Broad-leaved Species of Potamogeton 61 


The stipules often shrivel and decay, so that the few remnants 
(if any are left) are insufficient for diagnostic purposes. Rhi- 
zomes are seldom collected and when present are of but slight 
taxonomic value. Thus, unless one resorts to the internal 
anatomy of the stem, it is, in the great majority of cases, neces- 
sary to determine the specimens from the leaves. These may be 
either submersed or floating, or both, in all the subsections except 
Crispi, Praelongi, and Perfoliati, which have only submersed 
leaves. These three subsections always have sessile perfoliate- 
clasping leaves! with broad bases in contradistinction to the other 
subsections which have their submersed leaves petioled or at 
least narrow at base. 

First considering the submersed leaves: their shape is of 
primary importance. When typically developed, P. perfoliatus 
can be recognized immediately by its short-ovate leaves; P. 
Richardson by having them ovate-lanceolate; P. praelongus 
with them oblong and cucullate at the apex; the members of the 
Lucentes by the sharp-pointed apices; P. natans and P. Oakesia- 
nus by their coarse and delicate, narrowly linear leaves, respec- 
tively; P. nodosus by its long-petioled lanceolate leaves gradually 
tapering at both ends; P. pulcher having distinctive lanceolate 
leaves abruptly tapering to a petioled base; P. amplifolius with 
its massive leaves conspicuously arcuate; and P. alpinus with 
obtuse apex and sessile base. Unfortunately, however, it is not 
always as simple as that. These leaves exhibit such a wide 
range of variation? from the typical and are often in such a poor 
state of preservation, if not entirely absent, that one must 
resort to other parts of the plant for diagnostic characters. The 
leaf-margin of P. crispus is finely but evidently toothed; in the 
other broad-leaved species the leaf-margins are strictly entire, 
except in the Nodosi, which have 1-celled denticles so extremely 
fugacious as to be found only in the youngest leaves; the Per- 
foliati, in which the 1-celled denticles are more persistent; 
and the Lucentes, with strongly developed denticles. None of 
the indigenous North American species with free stipules has 
a dentation visible to the unaided eye, consequently they should 

1 The hybrid P. gramineus X seh ote var. bupleuroides may have perfoliate sub- 


mersed leaves and also floating leav 
2 See on under VARIATION. 


62 Rhodora [Marcu 


never be confused with P. crispus, a species naturalized from 
Europe, which has an evident serrulation. 

Turning to the floating leaves, we find at first glance a dis- 
couraging similarity. With the exception of P. alpinus, which 
has delicate, lacunate floating leaves the blades of which taper 
without sharp distinction into the petiole, all seem to possess the 
common ovate-elliptic shape and have the same blunt mucro 
at the apex. When typically developed, however, we find some 
help in the bases of the blades. Their shape ranges from cordate 
in P. natans and P. pulcher to rounded in P. Oakesianus and P. 
amplifolius to cuneate in P. nodosus to tapering without sharp 
distinction into the petiole in P. alpinus. Yet, the range of 
variation is so great in any one species that the shape of the leaf- 
base must be used with caution. Even P. natans with its strongly 
cordate leaf-bases may under certain conditions have them 
narrowly cuneate.! The number of nerves is of primary import- 
ance in the case of P. amplifolius, which has them very numerous 
and close together. The size and shape of the stomates appear 
to be rather uniform, and while there is some variation as to the 
number in a given field, this appears to be influenced by factors 
which are ecological rather than genetic. Examination as to 
number and condition (whether functional or closed by cuticle) 
may be useful in determining whether the abnormality of some 
floating leaves is due to genetic or to ecological factors. 

Stem. For a genus which varies so considerably due to 
ecological conditions, and which is so often found lacking parts 
that have the essential diagnostic characters, it is exceedingly 
fortunate that there are good characters in the anatomy of the 
stem. These were apparently first noticed and their importance 
realized by Raunkiaer? who used them in his studies on the 
Danish species and then extended his “anatomical investigations 
also to foreign species, intending to bring about a new and better 
basis for an eventual monograph of the genus Potamogeton.’” 
In 1907, Chrysler* elaborated further on the anatomy of the 

! See discussion under P. na 

2C. Rai pooanengh De danske iiiadiatatinialed Naturhistorie, Bind 1, Enkimbladede, 
p. -_ 10 (1895— 

unkiae, Anatomical Potamogeton-Studies and Potamogeton fluitans, Bot. 


mids, 28: 254 (1903). 
yb Bot. Gaz. 44: 161-188 (1907). 


1943} Ogden,—The Broad-leaved Species of Potamogeton 63 


central cylinder, explaining the phylogenetic relationships of the 
various types. agstrém in his monograph! emphasized the 
taxonomic value of the anatomical characters, especially of the 
stem, laying great stress on the types of endodermi, of which he 
described six kinds, and on the arrangement of bundles in the 
stele of which he described and illustrated six patterns. Other 
characters used by him include: the development of mechanical 
tissue, the presence or absence of a cortical layer directly be- 
neath the epidermis and acting as a supporting layer to it (hypo- 
derma), and the length in proportion to width of the cells of the 
epidermis. 

An examination of hundreds of stems has shown that a number 
of the characters have real diagnostic and phylogenetic value. 
These are discussed below.2 Although it has been suggested 
that the extent of mechanical tissue present is dependent on 
whether the plant grows in still or running water,? Raunkiaer* 
and Chrysler’ show that no such correlation can be drawn. 
Hagstrém, too, in his Critical Researches, repeatedly shows for 
many species that the habitat does not affect the essential char- 
acters of the stem-anatomy. An dquarium-plant of P. natans 
which I grew from a seed had its mechanical development not 
at all lessened. 

A morphological character of some importance is the degree 
of branching of the stem. The members of the subsections 
Lucentes and Perfoliati are usually much branched; P. praelongus 
of the Praelongi and P. Oakesianus of the Natantes are commonly 
branched; P. natans of the Natantes and the members of the 
Nodosi, Amplifolii, Colorati and Alpini are rarely branched, and 
then usually only slightly so in the upper part of the plant late in 
the season. a 

The rhizome appears to offer no important diagnostic char- 
acters, at least when dried. Some species tend to have dark- 
colored (usually reddish) spots on their rhizomes, while others 
are unspotted, but this is rather inconstant. 


1J.O. Hagstriém, Crit. Res. Pov. (1916). 


2 See EXPLANATION OF STEM-ANATOMY CHARACTERS USED IN THIS TREATMENT. 
. Schwendener, Das mechanische evinces in anatomischen Bau der Monocotylen, 
p. 122 (1874). 


*C. Raunkiaer, Bot. Tids. 25: 275 (1903). 
+ A Chrysler, Bot. Gaz. 44: 169 (1907). 


64 Rhodora [Marcu 


WintTeR Bups. With the exception of P. crispus, winter buds 
are so uncommon among the species here treated as to be of 
little or no diagnostic value. They have been so ably discussed 
by Professor Fernald! that further remarks are quite unnecessary. 


VARIATION 


It is well known that in vegetative characters any broad- 
leaved species of Potamogeton varies considerably. Those of the 
subsections Lucentes and Perfoliati have a wide range of varia- 
tion and all of the species here treated have a variability which 
is evident. Morong bewailed this characteristic of the group 
and felt that ‘“‘so protean are their forms, so eccentric their 
action, constantly changing under changed conditions of season 
and water’ that he must put forth his treatment “with great 
diffidence”’.2. Chamisso and Schlechtendal* had earlier reached 
the conclusion that, to use the translation of Emmeline Moore’, 
“Species of Potamogeton changing their habit seem often to pass 
into others, and feigning the habit of other species baffle re- 
search.” : 

Certain of these differences in form are undoubtedly genetic, 
but for many the evidence that they are ecological is very strong. 
Numerous observations and experimental studies have shown 
that individual plants of certain of the broad-leaved species 
exhibit a variation that is almost unbelievably wide, not only 
including many named varieties but even simulating other 
species. Noteworthy among such experimental studies are those 
of Esenbeck,® whose great contribution from the standpoint of 
taxonomy was in demonstrating that leaf-shape varies greatly 
under varying conditions of nutrition and changes of water- 
level. In general, poor nutrition appeared to give the following 
sequence of events: the broad floating leaves become increasingly 
narrower, finally disappear and are replaced by the submerged 
type, which in turn may be reduced to phyllodia. 

1M. L. Fernald, Mem. Am. Acad. Arts & Sci. 17: pt. 1: 18 (1932). 

2? Thomas Morong, Mem. Torr. Club 3: no. 2: 2 (1893). 

3’ Chamisso and Schlechtendal, Linnaea, 2: 159 (1827). 

‘Emmeline Moore, The e Potamogetons in Relation to Pond Culture, Bull. Bur. 
Fish. 33 (1913): 256 (1915 


5’ Ernst Esenbeck, pales zur Biologie der Gattungen Potamogeton und Scirpus, 
Flora pent 151-211 (1914). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 65 


Fryer’s experiments with the broad-leaved species which he 
grew in tubs caused him to prefer a very conservative treatment 
of the group. 

Of outstanding importance in this connection is the work of 
the Pearsalls' who found that variation in size and shape of leaf 
was due to many factors, acting independently or together. 
Their results led them to the conclusion that a conservative 
treatment is the sane one. 

While it is known that many of the variations are strictly 
ecological and one can in many cases name the contributing 
stimuli, it must be admitted that certain of these differences in 
form may be genetic. It is often difficult to distinguish from 
herbarium specimens alone which is the primary cause. When 
such variants appear to be mere forms or states, without separate 
geographic ranges, and especially when it is reasonable to suppose 
that the variations might be caused by ecological factors, it seems 
wise to refrain from giving them formal rank in our system of 
nomenclature. In the words of Fryer, ‘‘we should not confound 
temporary states with permanent varieties . . . In looking 
over an extensive series of dried specimens of any Potamogeton, 
it is easy to select very. distinct looking forms, and as easy to 
give varietal names to them; this is the method that has been 
followed by most of the writers on the genus, and I regret to say 
is still being followed, with the result of confusing rather than 
elucidating an already too difficult subject.’” 


HyYBRIDISM 


In dealing with broad-leaved species of Potamogeton, one is 
dealing with plants that are usually sterile, which are highly 
susceptible to ecological variation and which hybridize freely. 

If one classifies all the fruiting specimens, then correlates with 
them all the sterile plants the vegetative parts of which show 
their relationships to the fruiting entities, one still has a great 
mass of material that fits nowhere. Any attempt to understand 
such material must determine for each specimen whether the 
variation is genetic or ecological or due to hybridism. Here the 
internal anatomy of the stem will prove of great assistance and 


1W. H. Pearsall & W. H. Pearsall, Journ. Bot. 61: 1-7 (1923). 
2 Alfred Fryer, Pot. Brit. Isles, p. 41 (1900). 


66 Rhodora [Marcu 


often will allow the specimen to be rather definitely placed. 
But many times the stem-anatomy shows an upset; this points 
toward hybridism, for the species based on fruiting specimens 
show a stem-anatomy that is remarkably uniform, and repeated 
observations have shown that ecological factors do not affect the 
anatomical characters of the stem; neither will minor genetic 
variations. But one cannot merely say “hybrid,” as an easy 
way out of a difficulty; one should name the parents, usually 
not an easy thing to do. 

Whether experimental studies will produce data of value in 
this connection, I am not prepared to say. Hagstrém thought 
not, saying, ‘“‘The hybrids must be studied according as Nature 
produces them. Cultivation and experiments in hybridization 
may not lead to great results as to the solution of this intricate 
question.”! To this, Bennett retorts, ‘‘having seen the results 
of the late Alfred Fryer’s work in this direction, I consider 
cultivation is a very great help—anyhow it affords a negative to 
some of the proposed hybrids given.’”* 

Very little seems to be known about the chromosome number 
of Potamogeton. Wiegand reported a haploid number of seven, 
or possibly eight, for P. foliosus. Wisniewska‘ states that the 
haploid number for P. perfoliatus is probably (‘‘wahrscheinlich”) 
twenty-four. The only other reported count is in Tischler,° 
where the haploid number for P. fluitans (P. nodosus?) is given 
as twenty-six, referring to Kuleszanka 1934, without citing any 
publication. Such data promise interesting results from counts 
on the species in this genus. 

Cytological material (young pollen) of several species which 
I have examined shows that the chromosome number varies 
(at least four different numbers) in different species, although 
absolute counts could not be made. 

In the present work, the hybrids are taken up at the end of the 
general treatment. Only those that strongly simulate the sub- 
sections here specially treated are discussed. 


1J. O. Hagstrém, Crit. Res. Pot. 12 (1916). 


4 Ewa Wisniewska, Acta Societatis Botanicorum Poloniae 8: 157 (1931). 
8G. Tischler, Tab. Biol. Period. 5 & 6, Nachtrag no. 2, Teil no. 2, p. 99 (1935-36). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 67 


ACKNOWLEDGMENTS 


In a study of this nature I have naturally found it convenient 
to accept the assistance and suggestions so kindly offered by 
many people. I am especially grateful to Prof. M. L. Fernald 
under whose guidance the study was conducted and who, by his 
clear knowledge of the difficulties involved, offered invaluable 
suggestions. Also to Mr. C. A. Weatherby for his helpful advice, 
especially concerning nomenclature, I am deeply thankful. 
Thanks are also due to the members of the Gray Herbarium 
staff who have so willingly helped with various details; to Prof. 
F. H. Steinmetz, of the University of Maine, who has taken a 
keen interest in the work and has assisted greatly in the collect- 
ing of material; to my wife, Edith B. Ogden, for assistance in 
many ways; and to the curators of the herbaria from which 
plants have been borrowed. 

Herbarium material has been seen from the following sources: 
Gray Herbarium (G), New England Botanical Club (NE), New 
York Botanical Garden (NY), United States National Museum 
(US), Field Museum (F), Missouri Botanical Garden (M), 
Dudley Herbarium, Stanford University (8), Michigan State 
College (MSC), National Museum, Ottawa, Canada (C), 
University of Montreal (MT), Private herbarium of Marie- 
Victorin (V), University of Maine (ME), Portland [Maine] 
Society of Natural History (P), University of New Hampshire 
(NH), C. C. Deam (D), private herbarium, E. C. Ogden (0). 


EXPLANATION OF STEM-ANATOMY CHARACTERS 
USED IN THIS TREATMENT 


Five sets of characters are used, combinations of which 
should determine the species. These are: the arrangement of 
bundles in the stele, the type of thickening of the cells of the 
endodermis, the presence or absence of bundles in the interlacunar 
spaces of the cortex (interlacunar bundles), the presence or 
absence of bundles just underneath the epidermis (subepidermal 
bundles), and the presence or absence of a layer or layers of cells 
directly underneath the epidermis (pseudo-hypodermis). See 
PLATE 747, 

One of the most important sets of characters is that of the 


68 Rhodora [Marcu 


arrangements of bundles in the stele. These fall into six types. 
The arrangement called the proto-type (because it appears to be 
the type from which the others are derived) is illustrated by 
PLATE 748, r1G. 1. Here the bundles are all free and un-united 
(each having one patch of phloem), four of them being median 
and with usually three to five on each side. When this arrange- 
ment is found, the specimen will belong to one of the subsections: 
Colorati, Amplifolii, Lucentes, or Praelongi. In the trio-type 
(Frias. 2 & 3) three of the median bundles have united to form a 
bundle ‘‘trio”’, giving only two median bundles, one of them 
with a patch of phloem on the outer face of the xylem canal and 
two patches of phloem on the inner face (Fic. 2), or by a further 
union having only one inner patch of phloem (ria. 3). The trio- 
type is always found in the subsections: Alpini, Nodosi, Natantes 
and Perfoliati. Members of the subsection Lucentes may also 
have the trio-type of stele. A further reduction in the number 
of bundles is brought about by the fusion of the lateral bundles 
and as this invariably results in an oblong-shaped stele it is 
called the oblong-type. The two median bundles may remain 
separate (ric. 4) and even the lateral bundles may at times be 
scarcely united but merely crowded together. The oblong-type 
with two median bundles is found in the subsection Lucentes, 
and is prevalent in hybrids having a member of the Lucentes as 
one of the parents. A fusion of the two median bundles is 
accompanied by the more or less complete fusion of the laterals 
on each side, resulting in a stele of an oblong (or elliptic) shape 
with but three bundles (ric. 5). This is the usual type in the 
subsections Crispi and Lucentes. The final reduction to the one- 
bundled-type (ric. 6) is not met with in the broad-leaved 
species, but is found in the hybrid P. Berchtoldi « perfoliatus 
var. bupleuroides. 

The cells of the endodermis give an important set of diagnostic 
characters. Although several types can be recognized, these 
have been grouped in the present treatment into two principal 
types, according to the type of incrassation of the cell-walls. 
Cells which are thickened evenly or not at all are called O-cells; 
those which are much thickened on the inner and lateral faces 
but remain thin on the face next to the cortex are called U-cells. 
This latter type of endodermis is always found in the Natanies, 


1943] Ogden,—The Broad-leaved Species of Potamogeton 69 


Lucentes, and Praelongi. All the others here treated have 
O-cells. 

The bundles of the cortex give exceptionally good sets of char- 
acters. The cortex of a Potamogeton stem is highly lacunate. 
At the junctures of the walls separating these lacunae may, in 
some species, be found bundles composed of vascular tissue and 
fibers or merely of fibers alone. These are called interlacunar 
bundles and are well developed in the Natantes, Lucentes, 
Praelongi, and in P. amplifolius of the Amplifolii. The other 
subsections are devoid of them, or rarely have one or two weakly 
developed. 

The cortical bundles found directly underneath the epidermis 
are called subepidermal bundles. They are seldom as strongly 
developed as the interlacunar bundles, yet they furnish a set of 
characters which is quite useful. The subepidermal bundles are 
present in the Colorati, Natantes, Praelongi, and usually in the 
Lucentes, but absent in the other subsections. 

The last set of characters used is the presence or absence of a 
layer or layers of cells directly beneath and adjacent to the 
epidermis. This is called the pseudo-hypodermis. It is always 
present in the Crispi, Amplifolii, Praelongi, and Perfoliati, but 
is not present in the Alpini and Nodosi, whereas the Colorati, 
Natantes, and Lucentes may or may not have it. 

A number of methods of preparing a section of stem for ana- 
tomical examination may be used. The following procedure has 
. been found to be simple and rapid and highly satisfactory. 
When a choice can be made as to the part of the stem to be 
sectioned it should be at about the middle of the second or third 
internode below the lowest peduncle. However, any portion of 
the stem, if not too close to a node, is usually quite satisfactory. 
The piece of stem is boiled in water for a few minutes to soften 
and expand the tissues, then dropped into cold water to reduce 
flabbiness. A clean transverse cut is made near one end with a 
sharp razor-blade and this end dipped into ordinary blue ink, or 
other stain, for a second or two. The stem is now rinsed briefly in 
cold water and rolled on a blotter to remove excess water. With 
the razor-blade a thin free-hand section is taken off at the 
stained end, a water-mount made (stained end up), and examined 
under a microscope. If the stem is badly flattened, it can be 


70 Rhodora [Marcu 


teased out with fine needles. A magnification of about 100 
diameters is highly satisfactory, although much lower powers 
will enable one to determine most of the essential characters. 
Such mounts need not be thinner than that obtainable by free- 
hand sectioning, for enough light will come through the lacunae 
to illuminate the tissues. 

The amount of time necessary for making an anatomical ex- 
amination of a stem need not be greater than that necessary for 
boiling and dissecting a flower. 

A number of improvements can undoubtedly be made on this 
method, but mostly at the sacrifice of simplicity and speed. 
Potassium hydroxide solution is not recommended for swelling 
the tissues as it destroys the delicate cortex. A weak lactic acid 
solution will soften tissues quickly and swell them slightly with- 
out destroying the cortex, but for most specimens water is more 
satisfactory. If cold water does not stiffen the material suf- 
ficiently for proper sectioning, a few drops of concentrated 
ammonia in the water are helpful. 

Ordinary blue fountain-pen ink is suggested as a stain because 
it is easily available and has proved very satisfactory. It acts 
quickly and as a differential stain: coloring the vascular tissue 
deep blue and the cortical and pith cells light blue; the mechanical 
tissue (sclerenchyma) usually remaining a bright yellow and the 
endodermis usually yellow or orange. The clearer results ob- 
tained by aniline dyes are not necessary for this work. <A two- 
solution differential stain is not recommended for speedy eX- . 
amination. 


THE CLASSIFICATION OF NortTH AMERICAN PONDWEEDS 


In order that the groups of Potamogeton here specially dis- 
cussed may be properly oriented, an outline (largely following 
Hagstrém and Fernald) of the relationships of North American 
species is here appended. The subsections treated in this paper 
are indicated by asterisks. 


Subgenus CoLEoGETON (Reichenb.) Raunk. 
Section Connati Hagstr. 
Subsect. Fitrrormes Hagstr. (P. filiformis Pers., P. lati- 
folius (Robbins) Morong). 


Rhodora 


Plate 747 


Subepidermal 


Epidermis 


Pseudohypodermis 


TPP 


asnttan 


Trio-bundle 


Lateral bundle 


Interlacunor bundle 


QR 
oP e'e, 
OP 


Bas 
a Ne. 
Q ie la fi 


<4 


g 
enae. 
() 
? 
so 


xo 


we’ 


Endodormis 


bundle 


axe 
ORR 
SOUS 
uP", 


Locuno 


PoTAMOGETON NATANS: CROSS-SECTION OF STEM, X 35 


Plate 748 


Rhodora 


' 
cy | o ; “| § 
: ial 
3 *) 
$ Sime Ay 
2 5 
= =) 
o 2 s 
z = 2 
co 
a) us S a 
o a3 p 
oe ae se 
s= rs 3 
oOo. Os 
=o zo 2 
ac OE FR 
BE S a fey 
oe Qo a8 
s= -_ = 
s§ . 4 5 
Z = a 2 
3 = a = 
z if] 
° 
3 » 3 
x= ee & 
g 2 . 
= 
°o n 
= 
3) 
4 
om © & 
¢ ” ae 
H if 
2 oe 
° ae 
: 3: 
= 
2 ea 
= = eS 
3 2 cf 
a ws ew 
as —— “& 
> P= ~ c i) 
o $s o> sah 
mee Se a 
- am pe as 
- <= @Qe me bg Sed 
Fi oo 55 
ree} £ a 3 
bi = , 
a = 
ae 
. 4 
5 =e 
ak 
£ <~ 
\s = Os 
OOS 5 
a 


1943] Ogden,—The Broad-leaved Species of Potamogeton 71 


Section Convo.uti Hagstr 
Subsect. VAGINATI Pata: (P. vaginatus Turez.). 
Subsect. Pectinati (Fries) Hagstr. (P. pectinatus L.). 
Subgenus EvpoTamMoGeton Raunk. 
Section Apnati Hagstr. 
ubsect. SerRuLATI Hagstr. (P. Robbinsii Oakes). 
Section Ax1LLARES Hagstr. 
*Subsect. Crispr Wallm. (P. crispus L.). 
Subsect. Monticou1 Hagstr. (P. confervoides Reichenb.). 
Subsect. CoMpREsSI eee) Panga 6 (P. zosteriformis Fern.). 
Subsect. OxypHyiui Hag (P. Porsildiorum Fern., per- 
haps also P. engieienta Fern. }. 
Subsect. Pusriu1 (Graebn.) ge 
Series PusILLI connaTI Hags 
ee Fo.uiosi Fern. (P. ‘foliosus Raf., P. fibrillosus 


Subserie PANORMITANI Fern. ee ie socal 
strictifolius Ar. Benn., P. pusillus L. 
Series PusILLi CONVOLUTI Hagstr. 

Subseries Acutt Hagstr. (P. gemmiparus Robbins, P. 
Hillit Morong; P. longiligulatus Fern., an atypical 
species, may belong in the Oxyphylli). 

Subseries Oprust Hagstr. (P. Porteri Fern., P. obtusi- 
folius Mert. & Koch, P. clystocarpus Fern., P. 
Berchtoldi Fieber). 

Subsect. Javanict Graebn. (P. Vaseyi Robbins, P. later- 
). 


alis 
Subsect. Hysripi (Graebn.) Hagstr., in part; 7 
Series Eunysripi Hagstr. Spiri illus Picken, F. 


wre Rati 2. capillaceus Poir., P. bicupu ulatus 


Subsect. Nae ee (Hagstr.) Fern. (P. epihydrus Raf., 
. tennesseensis Fern 
*Subsect. ALPINI (Graebn.) Hagstr. (P. alpinus Balbis). 
— Be hess nc o (Graebn.) Hagstr. (P. polygonifolius 
Fr 


ou 
*Subsect, keno Hagstr. (P. amplifolius Tuckerm., 
pulcher Tuckerm.). 
mduteees Novosti Hagstr. (P. nodosus Poir.). 
*Subsect. Natantes Graebn. (P. natans L., P. Oakesi- 
anus Robbins oe 
*Subsect. pine Graebn. (P. gramineus L., P. illino- 


ensis Morong). 
*Subsect. PrRaAELONGI Hagstr. (P. praelongus Wulfen). : 
*Subsect. Perroiiati (Graebn.) Hagstr. (P. Richardsonii 
(Benn.) Rydb., P. perfoliatus L.). 


72 Rhodora [Marcu 


Kerry TO THE BROAD-LEAVED NortTH AMERICAN SPECIES OF PoTAMOGETON 


1. Stem laterally compressed; leaves all submersed, margin, 
out thed; stipules slightly adnate to base of leaf; fruits with 
winter buds hard and horny; stele of 
che oblong-type pattern: endodermis of O-cells; interla- 
cunar bundles absent.! (subsect. yen hace ate eek capes 1. P. crispus. 
1. Stem terete; leaf-margins entire or minutely denticulate; 
stipules completely free from leaf- ‘nee. fruits with bea 
not more than 1 mm. long; winter buds rare, not op Goal 
horny; stele with the pattern various in t of the 
rags Ben lice 9 endodermis of U-cells and yaa Gi 
Fis a 
2. a ienabied: oxi petioled or tapering to . sessile base, 
scarcely clasping; floating leaves present or absent... . 
3. Submersed leaves ae " chin aa leaves Gapaally absent) 
delicate, giranenter t, tapering without sharp distinc- 
son into the p atlole: fruits ‘— exocarp hard and 
mooth, t Saarascltve (oubmect: ALPINE se i058 ibaa: 2. P. alpinus. 
3. Submersed le eaves sessile or tioled: floating leaves cori- 
aceous, opaque, cordate to cuneate at base, blade dis- 
tinct for Detidle: fruits with exocarp soft and porous, 
greenish, brownish, or reddish... .4. 
4. se ergy connectives 1-2m plang soit spikes 5-7 
mm. thick; mm 


00 
usually with a cavity; interlacunar bundles absent; 
subepidermal bundles present; eastern Newfoun 
land and Sable Island, Nova Scotia (subsect. 

_OOORAT eee. ee a ae a P. polygonifolius. 
. Sepaloid eens mostly 1.2-3 mm. mide: ‘rating 


keels vadiable,. beak evident, endocarp rere eceait 
interlacunar bundles absent or pap wr if geese 
cad 


5. Submersed leaves broadly linear, "lanoéolate, or 
ovate, less than 30 tim = as long as broad, nerves 
3.37: ‘stele with the pattern various in t if of 
the trio-type, then with but 1 patch = i itd on 
the i = face of the trio-bun 3 Na 

6. Submersed leaves with 11-37 an rves, margin 

strictly entire; floating leaves cordate to reeds 

at base, nerves mostly more than 21; stele 
with the proto-type pattern; endodermis of 

 aegy (subsect. AMPLIFOLII)... .7. 


Yq of the nerves more prominent than the 
rest as seen by transmitted light; eer 
often clavate; fruits cuneate at base, 3.5-4.5 
(-5) mm. long; interlacunar bisachen present : 
4. P. amplifolius. 
7. Stem usually Sonmpacuonaly age ney Byron sub- 
mersed leaves not arcuate, 1-2.5 cm. wide; 


' Anatomical characters refer to those of the stem. See p. 67 and keys on pages 
75 and 77-85. 


1943] 


floating leaves cordate or rounded at base, 
mostly with less than 30 nerves, all nerves 
of Shoch equal prom ae as seen trans- 
mitted light; peduncles not conspicuously 
clavate; fruits rounded or lobed at base, 
of (-4) mm. long; interlacunar bundles 


BARES sil a Whe bee ia), Keke ee 

6. Submersed cain with 3-29 nerves, margins with 

fugacious one-celled translucent denticles; 

floating toe cuneele to rounded at ies — 
nerves mostly less than 21; stele with t 

tern various in type; i if of the reckon ag 
tern, then endodermis of U-cells. . . .8. 

8. Pam ecees hcg vee petioles 2-13 em. long, 


apex ac 1 but not sharp-pointed nor mu- 


dish, keels mostly muricate; endodermis of 
O-cells; interlacunar bundles absent (subsect. 


ODOSI 
8. Submersed leaves sessile or with petioles up to. 


4 cm. long, apex acutish or ye age 
often. somewhat eee ronate; fru a -3.5 

. long, usually greeni ish, rao arcely 
muricate; endodermis of U-cells; catartheweax 


Stem usual] uch branched, .5-1 mm. in 
ameter; submersed leaves (—1.5) em 
wide, (5—) 7-12 (—30) times as long as wide, 
sessile, nerves tly 3-9; floating leaves 


1.5-5 (-7) em. long, 1-2 (-3) cm. wide, 
petioles mostly longer than_ the ook laren tt, 
stipules .5-3 cm. long, .1- wide, 
keels faint; peduncles clavate or gone fruit- 
ing spikes 1-2.5 em. lon ng, .6-.8 cm. ‘thick; 

paloid connectives mos y 1.2-1.6 mm 
wide; fruits 1.7-2.5 (-2.8) em. long, 1.6-2 

2.3 


d 1 lateral b 
— bundles only in the outer 
nterlacunar GOle <6. Fee HES es 


9. Stem sealed or once branched, vib 1.5-5 mm. 


in diameter; submersed leaves 1.5-4 cm 
wide, mostly times as one as wide, 
; 


stele with the pattern onlay in type; 

interlactinar bundles in the outer interla- 

sie — circle or sometimes So in the other 
GUMAY CIPCIOS oo es oe ok ca ws 


5. Sabana g wiz sear linear, less than 3 mm 


Ogden,—The Broad-leaved Species of Potamogeton 73 


5. P. pulcher. 


. P. gramineus. 


. P. illinoensis. 


74 Rhodora [Marcu 


wide and more than 50 times as long as broad, 
nerves 1-5; stele of the trio-type pattern with 2 
patches of ‘phloe m on the inner face of the trio- 
he ndle (subsect. NaTAN as 
. Stem .8-2 mm. in diameter; ‘ submersed leaves 
.8-2 mm. wide; floating leaf-blades 4-9 cm 
long, 2.5-6 cm. wide, usuall So eo at base, 
= petioles 1-2. 5 mm thick; stipules 4.5-11 
m. long, strongly keeled; pedu uncles as thick 
as s the Sign 7 fa are picenagy ie mostly 1.8- 
erty s .9-1.2 cm. thick; 
matire leva mostly mm. lon 
e, keels ma + developed, re kn 


strongly ——— wh mice than 1 interlacunar 

circle; pseu 8 present; epidermal 
cells 1.5-3 (-4) tinesas long aa bro. ad gE Ei weld 7. P. natans. 

10. Stem ge mm. in diameter; submersed leaves 

.25-1 mm. wide; floating leaf-blades 1.5-5.5 

em. lo tog 1-3 em. wide, roun or cuneate 

at base, on petioles .2-1 mm. thick; sti — 

1-5.5 cm. long, keels -gtocpa'sand onhy at base; 


sides but not pitted, apex seed pointing 

a little above bas basa end; gence with 2-3 

lateral bundles on each sic 4% toy ew 

bundles strongly developed oa t 1 interla- 

cunar circle; pseu usually ab- 

sent; epidermal cule 4-7 ie: as thee as broad 

8. P. Oakesianus. 

2. Leaves all submersed, cordate or img at — clasping 
Big Ae! 74 the circumference of the s 

. Rhiz saree tpg: ted with rust ae aves Ceidditlile 


gins entire 2 ‘gr ae aod persistent and conspicu- 


m. vn pind i chan 3 mm wide, dorsal = l 
strongly developed; mbit ca th the proto-ty patte 
of U-cells; interlacunar boy su pidermal 


ate, seldom ae 1-10 cm. long, apex not 
cucullate, margin with fugacious, translucent, one- 
celled denticles; suaies pry inconspicuous or 
disintegrated to fibers; cles 1-25 cm. long; 

mm. 


pe sect. Pu 
12, — ovate-lanceolate to pore lanceolate, 3-10 
ong, coarsely nerved; stipules coarse, disinte- 


1943] Ogden,—The Broad-leaved Species of Potamogeton 75 


ating to persistent whitish fibers; peduncles often 
siaenta: 1.5-25 cm. long; fruiting spikes about 1 
cm. rw any (2.5-) 2.7-3.2 (-3.5) mm. long, 
() yaks 6 (-3) mm. wide; seicsan doe with a 
vi 2. P. Richardsonii. 
12. Leaves “onbiculat to ovate-lanceolate, the Pine ‘ipal 
nes m. long, delicat — nerved; s ——s Api 
a, piri ous; uncles never iavate e, 
long; rating — dhiuk .8cm penny fotite ro 3) 
2.5- niga (1.7-) 2-2.1 (-2.3) mm. 


O06 6 6 be ee 6 hoe eae eo 6 Oe a eee ee ee oe 8 Se 


2. 
wide; vioubiien ME OS Se Oa 2a 13. P. perfoliatus. 


Key To THE SPECIES mee ON THE ANATOMY OF THE STEM 
a Poe Nes bundles p ara 
. Interlacunar pension: sucntns Shel throughout... .3. 
“* Stele of the proto-type pattern... .4. 
. Endodermis of O-cells; subepidermal bundles absent 
4. P. amplifolius. 
4. Endodermis of U-cells; subepidermal bundles ee Vy 
1. P. praelongus. 
3. Stele of ai trio-type pattern, with the phloem on a 
i e of the trio-bundle appearing as 2 distinct 
patches; aides of U-cells; ee bundles ‘ 
SOE OS i bed ce Pee es ee ; 


patches; epidermal cells 4—7 times as long as broad 
P. Oakesianus. 


oT peat Yo pattern; epidermal cells 1-4 times ‘a 
10. P. illinoensis. 


oe ee 6) eee ee, 8 ee ee oe ee ee oe 


5. a of the Obie pine hgh ey a 


7. Stele with 1 (rarely with p real bundle and usually 

with _ lateral bundle on each side; interlacunar bun- 

me only in ea outer premio circle; stem .5—1 
9. P. gramineus. 


a Stele + with 2 ‘Garely with 1) central bundles and 2 or : 
n each side; interlacunar bundles in 
the outer sriteniaodiie circle and sometimes also in 
the — on tid circles; stem (1-) 1.5-2.5 : 
Win ti ibieIer: - oo a a ee 10. P. illinoensis. 

i ggeroEy ws bundles sient endodermis of O-cells (rarely 


Pe ie ie: 


8. tel - tt stem laterally com- 
S ni ‘ae tis ape ee type pattern; y LP tiepus. 


$e Oe See ae & eee 
6 ee ee ae eek ee OO oe 8 6 Owe Oe, ee 


8. Stele of the erode or proto-type pattern; stem terete... .9. 
° 6 ee vial soa ales aan a aie one: S$. FP. oe aloher. 
9. 10. § with the trio-type patte ere ti 
1. Stele with the phloem on re inner face of the trio- 
—_ " appearing as 1 patch; pseudo-hypodermis ec bies 


76 Rhodora [Marcu 


11. eels — the phloem on the inner face of the tri 
appearing as 2 patches (rarely with pi a 
12, Rucudaheooderinie Pree! IRONS ” pickae tool 
and 43. P. Pe 
12. Pseudo-hypodermis absent.................5- nus. 


SELECTIVE KEY TO THE SUBSECTIONS 


The following key is offered as a practical one for determining 
the subsection to which a specimen belongs. It is based on 
characters available with sterile and otherwise incomplete 
specimens. It is selective in that the user may choose the order 
in which the characters may be applied. This is of decided 
advantage when certain parts of the specimen are absent or 
difficult to make out, for the other parts may be sufficient for 
determining the subsection. 

The key is extremely easy to use. Any character may be 
selected which, if not sufficient to name the subsection, will lead 
to a number. This number will be found heading a column on 
one of the following pages. On this page another character is 
now selected and opposite it in the designated column may be 
found asymbol or new number. This is continued until a symbol | 
appears, which will designate the proper subsection. At any 
time the subsections that are still possibilities may be ascer- 
tained by reference to page 86. If a specimen traces to zero 
(0) it may be suspected that a character has been selected in- 
correctly or the specimen is a hybrid with parents in more than 
one subsection. The subsection Crispi is omitted from this key 
for it is represented only by the introduced species P. crispus, 
which can easily be distinguished from all our indigenous species 
of Potamogeton by its definitely toothed leaf-margins. 

When the subsection is determined, reference to the portion 
of the general key, on pages 72 to 75, dealing with that subsection 
should give the name of the species. 

The following symbols are used to designate the subsections: 
AL Alpini AM Amplifolii NA Natantes PR Praelongt 
CO Colorati NO Nodosi LU Lucentes PE Perfoliati 


1943] Ogden,—The Broad-leaved Species of Potamogeton 


STEM 
STELE 
Jove hag 2 es ee, aun errr 
T 


© 0 0 0 0g ee Wee ee oe ere oie. eS 


Cee ie ee W ee ee 2: wy 6 e610 ee eee 


OUI oi 0, 5 a 
Lanceolat 


Meer. iN ea 
‘Clas 


(8 aS ee Pe ee ee ee 
ON 8 ee oe Fe Eee Ae ee 
i 


Cee Sk We at ie Oe ae Gk cs ee oe Sue) fe ee oe | 


2 ew eee Oe ee 8 ee 6 bee tre 
Fee wes eee ee ee Hae ee eee 
@¢ U6 6 6s 66.0 8) 6.6 2 8 eee OS 8 ee 


STIPULES 
Intact and well developed..... 
sintegrated to fibers. ....... 
Delicate or fugacious......... 
FLOATING LEAVES 
BasE 
Condate; 22:25. (23 ee 


63a See Oe ee Pee bale Se ee ee, eee 


CO AM 


Sie) 698 (08 6 6 ee eS oO ee 


CO AM LU PR 


$8 6b ee ee 6 Oe ee ee wee oe eee Wee 8 8 


iM) DOA Ta Pe axes 29 eon 
RM UA LU FR. i. cate 
AL CO AM 


AL IL PR 
AL CO AM NO LU PR PE... 
Bhi BOA ces eG As 
PEE DU TA is ie el kk DSS 


a ee ee Roe Coe acre He eee oe 


Bi A PMR Pe eee 


BOA TAD TA) | oe a eet 
AM LU 


a a a ae Oe Se ee Doe ee ee ee a a 


i te a oh to Bak oe 
ow we ee eee bee oe ee 


Ce 


AL CO AM NO NA LU PR PE. 
AM NA PR PE 


A BM PR PE init Bes 


+6 Oe 8 ke eee 6 6 Fo ee 


eek 6 6 ee eee ee ee eee Os Ol ee eee ewe 


Bs OO A is i end 
OE Bai oe ies oes oe 


78 Rhodora 
STEM 1 baer, Sane 4 
Pro’ 18 18 36 43 
nae EOS PN Oe See EE 
o-t “ia Pee et ae ee 
Binnie eee cx LU LU LU LU 
NDODERMIS 
CUO; 6.5 see sah Lt 41-54 78 
WRI. y 53 3s seh 17 <4} 31. 71 
LwrERLAcUsAn B 
ROG ic. Se ves 15 36 15 68 
AOR eo es Cee ees 1? Core Hoey” ee) | 
Prciceaecde 2 B 
OM can Ve Pees li <4) 31 -7i 
BiG a ee eee ee 10 10° 25 10 
SUBMERSED LEAVES 
“Orbicula ore aa et 66 66 66 66 
PRN acka see eee 18 18 18 35 
ion "PR Bae pegs 5 48 48 48 76 
Soaked: 25.5620 es, Rise 8 oe 
Oblane. = Qe os ae 65 55 55 55 
Fee oe aa eee eae 33 62 64 
Base 
Cleaettat sis ee 46 46 46 74 
Not es iis Geek eae SW ws. 
Rounded So Ae ae 20 46 20 74 
Acutisn 3352-2 Se Se Be a 
nba geht a ep pete sy EOD 40 40 68 40 
Mucroniie.. .. os cs 68 68 68 68 
IOLE 
Pree 3 oe ee, 19 19 68 19 
Adee. fo Rc ke. ey Ri ee 
ee 
Wel Gl sO EGS bse r 1 3.4 
__Dentiuite jie lees vet ee 32 32 §&8 16 
ice pu feces ay. SL oe Sz 2 =e 
Bh ie ea ee a NN Ge | ER | 
BP cs a a ke S 6 i316 
Oe ee ke, 34 34 34 66 
STIPULES 
Intact SPP any een ananean 22 Se 
Pe. oe a ae 14 34 14 66 
Bb. OF Pe oe 24 24 24 54 
FLOATING LEAVES 
Base 
Contates ook io cc ee 44 73 69 73 
Round. of GU8Z 62 = c:. 7 ie 3 8 
SONNE ck AL AL AL AL 
NERVES 
dD i a ee SS. 7s 77. 78 
198 ae 39. 36 
tides eee 38 68 38 68 


53 NO 65 53 
5 40 7 23 


1943] Ogden,—The Broad-leaved Species of Potamogeton 


STEM 11 
STELE 
Ws Cy, eet Pe 73 
eee es, Gay Ske 51 
peo <i a fg Gee NE 0 
NDODERMIS 
et i eres Ba, 11 
Bae es Ga ae grees CO 
Trenetaciaias B 
BONE B62 oi eS AM 
Ee © Gt ee are 1 
east AN B 
OSOnL fe a CO 
Aneents? 28 662.754 Pa f 
SUBMERSED LEAVES 
“Onbicular MT DCN eet fy SEE 66 
es ae ee 66 
vi bec oie eee AL 
eee. 2. ee 11 
Ln Oe Oss cs eS 79 
MARE 88 2) US es NO 
ASE 
Cepia: 2 SES 74 
Not clamping. < 0.0.46. 
PEX 
UNnMES 4305 ee 74 
a eee See At 
arp-pointed........... 70 
Macau es Hee ee AM 
IOLE 
pent Ak as oe a 45 
AbSONG ei ae tes 
sal 
NGS, aE Riper cs Uaea 11 
Dental el ae 63 
RVES 
TS Gn a ee ee 51 
1 a ee es 1l 
Pe oe eee 39 
Be ee ee a ee 66 
STIPULES 
Tnteet as oko. 2h mee 11 
Fiber .c 3.925.433 66 
Ded, of Pikes 54 
FLOATING LEAVES 
Base 
Condlate. (5 Goo ee, 73 
Round. or cun.. <...:).. 45 
TRpmie:. Ak. AL 
NERVES 
110. 2 ee 78 
G+ Fe ae ee Scene « 29 
ee ees a AM 


12 13 

43 36 
58 

LU 


16 
36 
6: 


Pe SF gre 2 
41 18 
62 
LU LU 


CO 73 
iy are 3 


31 36 
CO 73 


17 41 
LU 68 


14 
67 
60 


AM 72 7 
40 42 43 
0. 3.0 
NA NANO NA 0 
69 40 4 3 
69 38 68 62 68 


LU 64 


19 
43 
LU 


45 
71 


68 
45 


71 
40 


0 AM AM PE 
5 57 


68 

LU 75 
19 46 

68 AL 

64 NA 


77 
NA 


68 


oS 6 oe 6 ee Oe Chee 6 ee 


Ue Sok Ve Oy ia ae dee a ay ye ee ont et ea 


ASCHUC 54. 2. oS 
SUBMERSED LEAVES 


Linear. Se hae ee es 


Caen sk ue weeks oa 
Notclasping 0 3c. 45. 
PEX 


PETIOLE 
Pr 


Coe EO Oe See eee Be ee 


5 egw ee be Se ee ee ele 


CXC OCHO oe eee Oo ee 6 ew 


Rhodora [Marcu 
$1.28: 8f 24 26 236 87 38 30: 


LU LU LU 67 68 68 AM68 73 PR 
21 22 23 74 47 #50 51 52 78 60 
LU LU LU 0 LU LU O LU O O 


74 51 78 54 54 79 27 56 29 PE 
62 LU 62 PR LU 62 O LU CO 6l 


62 LU 62 67 68 38 AM68 AM 61 
74 51 78 54 54 79 27 56 20 PE 


62 LU 62 PR LU 62 0O LU CO 61 
47 22 52 54 93 55 27 28 56 PE 


PE PE 0 66 66 AM66 AM AM PE 
66 68 AM 57 


21 22 23 24 25 26 27 28 29 30 
PE LU 63 64 N 


50 51 52 78 60 

25 85 27 28 20 Fa 

58 32 64 34 35 68 39 40 70 57 
AM 66 AM AM 57 


21 22 23 24 25 26 27 28 29 30 
60 PE NA 34 66 69 66 AM AM30 
74 74 AL 24 54 79 54 79 79 57 


NA 0 NA AMAM69 AMAM73 NA 
62 64 33 AM6S 38 70 40 45 NA 
AL AL AL AL AL AL AL. AL AL 


77° 7] 63 AL AL 77 78 78.78 Ne 
50 52 23 79 55 26 56 28 29 NA 
62 LU 62 AM68 38 AM68 AMNA 


1943] Ogden,—The Broad-leaved Species of Potamogeton 


STEM 81): SF 
STELE 
Proto-type...../...5...0. 59 LU 
SO UGOS 5 ETS Ax 62 3 
Oblong-type............ LU LU 
ENDODERMIS 
oe Je Ae ans 63 
IM etek da on 3h) LU 
INTBRLACUNA B 
- Ace weeks 31: LU 
| ee pepe ok ye 63 


Present ke a 31 LU 
Abseat bea hw leer LU 32 
SUBMERSED LEAVES 
PE 
Be yt ile Se games OP oat cae 0 PE 
WACO SoS eG BEE 59 58 
Oblong Lape separated: Belg 59 LU 
Lanoemiate : 2... ak 59 32 
Oblane: or ello. SS, LU LU 
MEROOES Estes for es 62 64 
ASE 
CMe 8 a oe PR PE 
Ag ne ae Na tee 62 64 
Roendee A SS Gas 61 PE 
here Mere ‘ sigitibrs Get 5 cla a 32 
a erat Gas Ameer ae S28 
iewcohabe CE ey oe LU LU 
IOLE 
Preehs .5.5...6 3 tes LU 64 
Abene.. . As KAS 31° 58 
te etal 
RIG ARE 31.32 
Dentist RPh bee oenaa LU 32 
oe pray yw a re eee 62 32 
Ball ais ee a LU 32 
Ba Fe oa 59 32 
A a ee eee PR PE 
STIPULES 
TOS SO ES Fes 31 32 
Pile or a RRS 61 PE 
Del. of fuk... 26a. PR PE 
FLOATING LEAVES 
Basg 
Cordate: 3. iid NA 0 
Rodnd. oF 6th). 6.440% 62 64 
Tiger. 3. i ae 
NERVES 
IO oi es NA NO 
1738 is a 62 64 
_ 62 LU 


O 6 © o's se 0 8 6 6 6 80 oe 2 6 8 


81 
383 34 365 
LU 67 68 
33 PE 58 
LU 0 LU 


NO 66 66 
62 PR LU 


36 37 


36 67 68 AM = 
U NA 62 63 
LU 0 LU 0 LU 


AM AM AM 39 70 
59 61 62 0 LU 


36 37 38 AM 68 
AM AM AM 39 70 


59 61 62 0 LU 
68 AM68 39 40 


38 39 40 


e 


62 67 68 
NO 66 66 
62 PR LU 
64 66 35 


AM AM AM 66 
68 


AM 68 
LU NA 62 
PR. PR <0 
68 69 38 


PR 61 NA 9 0 
AM 


M 68 
NA 57 PE 
64 


36 37 38 39 
LU 0 LU 63 


LU NA 62 63 
68 AM 


Bose £5 BS 


>> 
ze> 


33 34 
NA 34 
34 


69 AM AM 
38 7 0 


ag NA NO NO 
69 38 AM 68 


NO ces ee ee 
Ohlaes-bype Poe cye oan 
ENDODERMIS 


(oes 6 Oke Ce ek ee 


O00 06s + 00.8 eo e 6 oo 8 


Seanad oh. ae 
Muerotate. 4 6.55. 25 A 


2 i ty a ee ee ge MeO 


eee eee «6 eee we Olas wie 
Core eo er ese er sew ewes 


206 6 68 e 6 8 Oe eee a ed 


Rhodora 


41 42 43 
43 


aad = de | 


73 
71 
68 
73 


71 
68 


68 
LU 
43 
68 
62 
0 
42 
NA 
71 43 
LU 68 
LU 68 
71 
62 


LU 
43 
U 68 


43 


2 LU 
U LU 


AM 70 
AM AM AM 57 


44 45 46 
73 73 PR 
NA NO 74 
oe: 029 


73 
72 


45 74 
CO PR 


AM PR 
45 74 


72 COP 
AM 70 


69 
73 


4 


AM AM AM PE 


AM AM 57 
0 O 75 
73 45 46 
AM AM AL 


0 0 O 46 
43 44 45 AL 


44 
NA NO 74 
73 45 74 

57 


44 45 46 


14 73 z 
44 45 
0 O AL 


NA NO AL 


hs 


47 
LU 
47 
3 


74 
LU 


LU 59 
74 


LU AMNO 0 -LU 


74 
76 


74 
47 
LU 
LU 


LU 
47 


AL 


45 AL 76 76 
69 AM 0 LU LU 


48 


59 
76 


U LU 
AL 74 


59 


AL 


[Marcu 
49 
0 LU 


49 
0 LU 


1943] Ogden,—The Broad-leaved Species of Potamogeton 


Pea ats ep? ee fel ee Dee So a 


0 6 68 Oe eb ace 


ro ht eat Oe eh ge Pe ge ace a 


os eet ee ee ee ms 


Sharp-pointed........... 
Mucronate 


ee eh a ee ee ee 


00a wt Eh eee Hee ele eee 


7 ee ew Sue ae eee ae 


53 64 55 656 57 


78 54 79 56 PE 
NAO. LU 0 PR 


NA AM 68 AM PR 
78 54 79 56 PE 


NA 0 LU O PR 
78 54 55 56 PE 


> 
oo 


76 
55. 79 
LU 


on 
or 


AL 
55 5 


NO AM 68 
77 54 55 


70 ~20—0 
79 57 


55 56 (57 
NO PE LU NO PE 


53 74 76 78 PE 
78 54 55 56 PE 


66 68 70 57 
0 66 AM AM 57 
i] 


53 54 55 56 57 
NA AM AM AM 57 
AL 54 79 79 57 


NA 66 AMAM 0 
65 AM68 70 0 
AL AL AL AL 0 


53 AL AL 78 0 
53 79 55 56 0 
NA AM68 AM 0 


AM AM PE 
M 5 


58 


58 
58 
PE 


PE 


0 


LU 


59 


NA 


84 : Rhodora [Marcu 


STEM 61 62 63 64 65 66 67 68 69 70 


STELE 
Prototypes. foie tee is PR LU 0 LU O AM67 68 AMAM 
PPA VOG 0. oa fe vs NA 62 63 64 65 PE 0O LU NA NO 
Oblanmtyne. 66... i O° LU 0°: LU 0: 0 0: DU -Oeg 
ENDODERMIS 
WN es a. Se 0 O 63 NO NO 66 AM AM AM _ 
8 61 62 O LU NA O PR LU NA 
Nae opyadeties B 
ee oe Bee ea 38 61 62 0 LU NA AM67 68 69 AM 
We as eS 0 O 63 NO NO 66 AM AM AM 70 
SUBEPIDERMAL B. - 
TOBEN G8 hes 62-61 0% bU NA PR LU NA 0 
i eo a si 0 LU 63 64 NO es AM 68 AM 70 
SUBMERSED LEAVES 
SHA 
Onbicular iat Mav ey 0 O PE 0 O 66 AMAMAM AM 
we Se se PR LU PE LU 0O 66 67 68 AMAM 
Obisas bed a PR LU 0 ae 0 
fandeclate.. oe. PR LU 63 64 NO 66 67 70 
Oblanc: or ell. 6... 20.32 0 LU 0 LU 0 AMAM68 AMAM 
RE ee ee NA 62 NO 64 65 0O 
ASE 
Clseningts 6s 86. Bs PRO PE 0 PE PR 0.0239 
oe Wee re ee NA 62 NO 64 Pr AM AM 68 69 70 
PEX 
Rownaed. 2 i ess 61 NA PE 0O NA PE PR O NA 0 
Atte 6 ee. LU 63 64 NO66 AM68 AM/70 
rece RP Sy Sore 0 LU NO 64 NO AMAM68 AM/70 
Mueronate .; «#6. oi i 0 LU 0 LU 0 AMAM68 AMAM 
PETIOLE 
Pree 3 AS 0 LU NO 64 NO cg AM 68 AM 70 
Absint 05 220 ee eS 61 62 PE LU NA 66 67 68 69 AM 
MarGIn 
PO) oes cal ee vs 61 62 63 64 65 66 67 68 69 70 
Dentioniate: (645 3... LU 63 64 NOPE 0 LU 0O NO 
s 
WD ser hod oes dh NA 62 63 64 65 PE 0 LU NA NO 
Dre ose Gk. ay 0 LU 63 64 NO 66 AM68 AM/70 
ee ee a LU 63 64 NO 66 67 68 AM70 
oa te aes oie PR 0-PE 60 0 67 AM AM 
STIPULES 
lett kk 62 64 67 68 69 70 
i Pie cd ee 61 NAPE 0 NA 66 67 AM69 AM 
Deb or fae... 5... PR 0 PE .0 0 66 67 AMAMAM 
FLOATING LEAVES 
Base 
Cordatm (64.524 ics... NANA 0 0 NAAMAMAM69 AM 
Round. or cun........... NA 62 NO 64 65 AMAM68 69 70 
Teper... 55 i. m0 .0.0 0 4:86 4 
NERVES 
fi ee NA NA NO NO 65 0 NA NO 
11-88 cee NO 64 65 AM AM 68 69 70 
Sa NA 62 0 LU NAAMAM68 69 AM 


1943) Ogden,—The Broad-leaved Species of Potamogeton 


80 6 O20 6 elem 4 + ee 6 8 


SUBMERSED LEAVES 
SHAPE 


od 


2 


eeereeeee ee se eevee eees 


OF 8 O10 9 Cb 8-8 Se eee eee eS 


OO Ue ew 6 be ee bere ors 


o 6 Orie 6.8 Sele eee ele SU ee 


Eee 6 O10 OO ie! Come. 6 


68 Se eee O we ee ee mee 


ee 


72 73 74 75 76 
Ue 0 Fe ee 


98 TOs. Fhe TB IO aT 
CO 73 0 PRLU 0 
NA 0 74 AL 76 77 
Pig G2 © 


O73 74 AL qt AL 


C A 
72 CO 0 PR LU NA 


NA AM 0 PR LU NA 0 
CO 73 74 AL AL AL 78 


0. © 74°76 

72 73 AL AL 76 77 
NA 0 74 7 77 
CO 73 74 AL 76 AL 
0 AM 0 O LU O 
0 AMO O LU 


CO 73 O O LU 
NA AM74 75 76 


72 73 74 7 
NA AM PE PR 0 NA 
0 AM 74 


78 


0 
78 
0 


85 


79 


79 
0 


75 AL AL AL 79 


0 


O 72 73 0 
72 73 O O LU NA NO AM 
a 


AM 


AL AL AL AL AL AL 


0 AL AL AL 77 
73 AL AL 76 77 
NAAM 0 0O LU NA 


78 
78 
0 


79 
AM 


86 Rhodora [Marcu 


The numbers used in the selective key and the subsections they represent 
are as follows: 


1. AL CO AM NO NA LU PR PE 41. CO LU PR 
2. AL CO AM NO LU PR PE 42. CO NA LU 
3. AL AM NA LU PR PE 43. CO AM LU 
4. AL CO AM NO LU PE 44. CO AM NA 
5. AL CO AM NO NA LU 45. CO AM NO 
6. AM NO LU PR PE 46. AL PR PE 
. CO AM NO NA LU 47. AL LU PE 
% AL NO. NA LU. PE 48. AL LU PR 
AL AM LU PR PE 49. AL NA PE 
10. AL AM NO LU PE . AL NA LU 
ll. AL CO AM NO PE AL NO PE 
12, AL CO AM NO LU AL NO LU 
13. AM LU PR PE AL NO NA 
4. AM NA PR PE AL AM PE 
5. AM NA LU PR AL AM LU 
AM NO LU PE AL AM NO 
i. CO NA: LU PR PR Fe 
,§ CO AM LU PR LU 2m 
. CO AM NO LU LU PR 
. AL NA PR PE . NA PE 
AL NA LU PE NA PR 


> 
ay 
Frat 
ee) 
ee 
om 
ae) 
e 3] 
vA 
> 
a | 


AL AM NA LU 66. AM PE 
AL AM NO PE 67. AM PR 
AL AM NO LU 68. AM LU 
AL CO AM NO 69. AM NA 
.) NA PR PE . AM NO 
NA LU PR CO LU 
NO LU PE CO NA 
NO NA LU CO AM 
AM PR PE AL PE 
AM LU PE 7 | ire 
AM LU PR 6. AL LU 
AM NA PR 7. AL NA 
AM NA LU 7g. AL NO 
. AM NO PE 9. AL AM 
40. AM NO LU 


SYNOPSIS OF SPECIES 
1. P. crispus Linnaeus 


submersed, linear-oblong to linear-oblanceolate, principal ones 
5-8 (-10) em. long, (.2—) .5-.8 (-1) em. wide, semi-clasping 
at base, broadly rounded at apex; nerves 3-7; lacunae of 1 or 


1943] Ogden,—The Broad-leaved Species of Potamogeton 87 


rows along the midrib; margins finely and irregularly dentate. 
STIPULEs .5—1 cm. long, ‘slightly adnate at base, upper part fraying 
early, leaving the papery or shreddy bases. ‘PEDUNCLES sea 
about same thickness as greatest dines of stem, 2—5 (—7 
long. Spikes in anthesis compact or moniliform, ‘of 3-5 whoite 
of flowers; in fruit 1-2 em. long, 1—-1.3 em. thick. Flowers sessile 
or on pedicels up to .4 mm. long; sepaloid connectives orbicular, 
(1.2- > 1.5-1.9 (—2.1) mm. wide, claws .3-.9 mm. fed anthers 
.7-1.1 (-1.3) mm. long. Fruits ovate, (2-) 2.5-3 (-3.6) mm. 
long (excluding beak), (1.6—) 2-2.5 (-2. 8) mm. wide; keels obtuse 
ut prominent, dorsal one strongly developed below and with a 
small projecting tooth near the base; beak etrogom straight 
or incurved, 2-3 mm. long; exocarp rather smooth, greenish or 
brownish; endocarp loop solid and near the base; Bh of seed 
pointing toward the basal end. WuinTer Buns hard and horny, 
1-2 em. thick. 
P. crispus L., Sp. Pl. 1: 126 (1753); C. & S., Linnaea 2: 186 
(1827) ; Morong, Mem. Torr. Club 3: no. 2: 36 (1893); Graebn. 
in Engler, Pflanzenr. 4: rast 11: 97 (1907); Taylor, N. Am. FI. 
7: pt. 1: 21 (1909); Hagstr., Crit. Res. Pot. 58 (1916). 
onds and streams. South ern Quebec to Virginia, west to 
Missouri, also in California. Map 1. Europe, Asia, Africa, 
Australia. The following are sapianeneee: QuEBEC: lle 
Sainte-Therése, St.-Jean Co., Victorin & Rolland 45189 and 49141. 
VERMONT: tributary of Lake Champlain, Charlotte, July 8-11, 
1911, Dutton; Charlotte, Chittenden Co., July 7, ou ee 
N, mixed with P. peng nites MASSACHUSETTS: 
Arlington, Middlesex Co., Sep 1880, herb. C. E. Talon: 
ot Fresh Pond, Cambridge, Middlesex Co., Fernald & Weather- 
lantae Exsiccatae Grayanae 111; Cambridge Bot. Gar 
1864, J. T. Rothrock; Sudbury R., Concord, Middlesex Co., 
Ogden 1502. Connecticut: Housatonic R., Derby, £. H. : 
ie 11502; Lake Zoar, Housatonic R.., Southbury, New Haven 
i. Eames 1 1485. New York: Lake Ontario, southwest 
a Chaumont, Jefferson Co., Fernald, Wiegand & Eames 1 4089; 
Pierrepont P., Woodville, Jefferson Co., rs ouse 10069; north end 
of Cossayuna L., Washington Co., Muenscher & Lindsey 2739; 
abandoned canal, Montezuma, Cayuga Ca, Eames & Wiegand 
14535; pool at Eldridge L., Chemung Co., uppe per waters of the 
Susquehanna, Lucy 10814; erwisk Flats, Ithaca, Tompkins 
Co., Wiegand 11182; Old Ice Pond, Ithaca, R. Jones 7471; Float 
Bridge, Rochester, Bazter 5888; Lake Washington Park, Albany, 
Albany Co., June 14, 1910, S. H. Burnham. New JERSEY: tide 
eee Camden, May 5, 1866, C. F. Parker; Camden, June 1879, 
C, Martindale; Clifton, Nash 786; Cedar Brook, Plainfield, 
Nay 1879, Frank Tweedy. PENNSYLVANIA: Conestoga, near 
Lancaster, June 19, 1861, T. C. Porter; Tiaintn R., Huntingdon 


88 Rhodora [Marcu 


Co., Aug. 1864, 7. C. Porter; Lehigh R., Easton, July 11, 1868, 
T. C. Porter; Sharon, July 7, 1886, F. T. Aschman; outlet of pond 
on tributary to Pennypack Creek, Willow Grove, Montgomery 
Co., Adams & Tash 512. Drtaware: Wilmington, 1863, Wm. 
M. Canby; Wilmington, June 1879, A. Commons; Wooddale, 
June 24, 1879, A. Commons; Faulkland, New Castle Co., June 
1879, A. Commons; Greenbank, Oct. 16, 1879, A. Commons. 
Mary anp: Spesutie Island, May 27, 1879, J. D. Smith; Cleft 
Island, Great Falls, McAtee 2953. District or COLUMBIA: 
Fish Ponds, Shull 15; Georgetown, VanEseltine & Moseley 201. 
West Virainia: Fish Hatchery, White Sulphur Springs, Green- 
brier Co., Berkley 1215. Virarnta: Fourmile Run, Chesapeake 
Bay Region, Shull 465; near Four-mile Run, near Alexandria, 
Blake 9472; 1 mile s. w. of Williamsburg, Grimes 3255; Dyke, 
Fairfax Co., Metcalf & Sperry 1631; Claremont Wharf, Surrey 
Co., Fernald & Long 7747. Ontario: Toronto, Scott 16439; 
pool above Niagara Falls, John Macoun 26830; Kingston, 
June 15, 1901, J. Fowler; Sault Sainte Marie, Algoma Co., 
Fassett 14746. MricuicaAn: Muskegon Lake, Muskegon State 
Park, 4% mi. w. of Muskegon, Muskegon Co., Hermann 8647; 
Kalamazoo R., Kalamazoo Co., Aug. 15, 1938, W. G. Erwin. 
Reported from Ottawa Co., and Van Buren Co. by’ Oosting.’ 
Inprana: Wolf L., June 7, 1913, E. D. Hull; east side of Wolf 
Lake, just w. of Whiting, Lake Co., Deam 56607. ILLINOIS: 
Wolf L., Chicago, June 10, 1911, EZ. E. Sherff; Stony Island, 
Chicago, Cook Co., Steyermark 4227; Lake Nippersink, accord- 
ing to Tehon, Torreya 29: 42 (1929), specimen not seen. MIN- 
nesoTA: Lake Minnetonka, Keck & Stilwill 428 and 430; Mis- 
sissippi R., Wabasha Co., Keck & Stevens 335; Mississipp! 
bottoms, below Winona, Aug. 6, 1931, H. J. Oosting. Missouri: 
Neosho, Metcalf 948; Blue Spring L. and Osage L., formed by 
Blue Springs, 2 mi. s. e. of Bourbon, Crawford Co., Steyermark 
16313. Sours Daxota: Edmunds Co., July 1896, D. Griffiths 
(specimen in Herb. Brooklyn Bot. Gard., not seen; see Torreya 
29: 43 (1929)). Wyomrina: Lakes Swastika and Irene, Medicine 
Bow mountains (specimens not seen; see Torreya 32: 5 (1932)). 
CaLirornia: cultivated at Pomona College, the plants brought 
from Santa Ana R., near Corona, ‘only known station in Cali- 
fornia,” San Bernardino Co., Parish 19248; pond in botany lath- 
house of Pomona College, planted by Johnston from Santa Ana 
R., Munz 2785. Ornaon: near Silvies, Aug. 19, 1901, Griffiths & 
Morris (specimen in Herb. Brooklyn Bot. Gard., not seen; see 
Torreya 29: 43 (1929)). 


P. crispus is distinct from all other species of Potamogeton 
found in North America, and should never be confused with any 
1 Henry J. Oosting, Pap. Mich. Acad. Sci., Arts and Letters 15: 165 (1932). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 89 


of them, for it is the only one with leaf-margins having a denta- 
tation evident to the unaided eye. Its fruits are distinctive in 
having long (2-3 mm.) beaks. Fruits are usually not produced 
when the plant grows in deep water, but in shallow, warm pools 
it fruits freely.. The winter-buds are especially distinctive, being 
large (1-2 cm. thick) and hard and horny. This species is not 
native to the Western Hemisphere but is introduced into North 
America where in some places it is growing profusely and acting 
as a weed. It evidently arrived here rather early for Pursh in 
1814 reports it from ‘‘Canada to Virginia.’”! Bennett says, 
“The oldest dated American specimen I can find in England is 
in Mr. Cosmo Melvill’s herbarium, ‘Philadelphia, 1841-2, Gavin 
Watson & Kilvington.’ One from Delaware in the British Mu- 
seum Herbarium is probably older: it was collected by R. Egles- 
feld Griffith of Philadelphia . . . ’% In 1913, Hull’ reports 
it as being abundant and acting as a weed around Chicago in 
waters connected to Lake Michigan. He believed the arrival 
of the plant to that region to have been quite recent. Tehon* 
believes the westward spread of the species in North America to 
be due to migrating water birds and suggests that its introduction 
to North America may be due to the same agents. 


2. P. aupinus Balbis (American varieties) 

RuxIzoME about same thickness as stem, branching and creep- 
ing, pinkish, not spotted. Stem terete, 1-2 mm. in dia meter, 
simple below, rarely branched above, often pressing very flat: 
stele with the trio-type pattern, the phloem on the inner face of 
the trio-bundle usually appearing as two distinct patches; en- 
dodermis of O-cells; interlacunar and subepidermal bundles 
absent; pseudo-hypodermis absent. SuBMERSED LEAVES (ex- 
cluding transition-leaves) thin, delicate, translucent, oblong- 
linear to linear-lanceolate, 4. 5-18 cm. long, 1-2 cm. broad, usually 
with 7 prominent nerves, sometimes also having 2 to 6 less promi- 
nent or incomplete ones, sessile and slightly clasping, apex obtuse 
or rarely acutish but never sharp-pointed; margin entire; lacunae 
along the midrib of rectangular cells, 4-6 rows near the base, 
2o0r3 Sg at the middle and 0-2 rows near the apex. FLOATING 

in, translucent, often poorly developed or absent, 
blades ‘aipabal or oblanceolate to obovate or oblong-linear, 

' Frederick Pursh, Fl. Am. Sept. 120 (1814). 

* Arthur Bennett, Journ. Bot. 39: 201 (1901). 

*E. D. Hull, Ruopora 15: 171 (1913). 

+L. R. Tehon, Torreya 29: 42—46 (1929). 


90 Rhodora [Marcu 


4—6.5 em. long, 1-2 cm. wid, corny tapering with no sharp 
distinction into a petiole 1-3 ¢ long, with (7—) 9-13 (15) 

rominent nerves, outer pair ha ate te Ae cadet a short distance 
back from the obtuse apex; lacunae along midrib of elongate 
cells near base and of oval cells near the middle, entire blade more 
or less lacunate with rounded lacunae. Srrpu.Es of submersed 
leaves thin, membranous, oblong, obtuse, faintly 2-keeled with 
3 lateral nerves on each side, which meet the keel-nerves back 
from the apex, (1.2-) 1.5-2.5 (-4) em. long, 2-8 mm. broad; 
those of the gocam rag and floating leaves similar but broader 
(up to 1 cm.) and with more nerves (15), some also between the 
more prominent keel-nerves. PEDUNCLES same thickness as the 
stem, 3-10 (-16) cm. long. Sprkes in anthesis more or less 
moniliform, especially at the bane with 5-9 whorls “4 fore” in 
fruit cylindric and crowded, 1.5-3.5 cm. long, . thick. 
FLowERs on short pedicels 5-1 mm. long; eer connec 
orbicular to reniform, blades 1. 1. mm. wide, claws .7—.8 mm 
long; anthers oblong, .6—1 mm. long. Fruits mostly obovate, 
(2.5-) 3-3.5 mm. long eaains beak), (1.7—) 2-2.4 mm. wide, 
lateral keels none or very low, dorsal keel thin, usually prominent 
and well developed upward, "beak lateral, short and curved to- 
ward the back; exocarp smooth, pitted when immature, tawny- — 
olive; endocarp with keels rounded, beak linear, 1-1.3 mm. long, 
curved toward the back, loop solid; apex of seed pointing toward 
basal end. Whole plant and especially the rachis of the spike 
usually suffused with re 

Two varieties in North America: 


Submersed leaves neta prised to linear-lanceolate, 7-25 ce 
long, usually more t oa 8 times as long as broad, tapering to ae 
obtuse or acutish a 2a. var. tenuifolius. 
itmaaredd leaves ublane to pega po 4-10 em. long, usually 
less than 8 times as long as broad, apex rounded an 
GAUTIOD MENTED CHOON 5s i sss i de ksi 2b. pe subellipticus. 


Ci OBR OO OS 68 ee wk ee ea oe 


2a. P. aupinus Balbis var. tenuifolius (Raf.), comb. nov. 


P. tenuifolius Raf. Pgs Repos. hex. 3, 2: 409 fog re. ron 
Ruopora 33: 210 (1931). P. lucens sensu Michx. Fl. Bor.-Am 

1: 101 (1803), in part, not L. (1753). P. microstachys *Wolfg. 
in Schultes & Schultes, Mant. 3: 360 (1827). P. rue 


(1827). P. obrutus Wood, Cl.-Bk. ed. 2: 525 (1847). P. lucens 
var.? fluitans (Roth) Gray, Man. ed. 2: 435 (1856), as to syno- 
nyms P. rufescens Schrad. and P. obrutus Wood, not P. fluitans 
Roth. P. alpinus sensu Morong, Mem. Torr. Club 3: no. 2:19 
(1893), in large part; sensu Graebn. in Engler, Pflanzenr. 4: fam. 
11:70 (1907), as to Am. plant in large part; sensu Taylor, N. Am. — 


91 


1943] Ogden,—The Broad-leaved Species of Potamogeton 


2 


” 
] 
°o 
& 

» ul 
‘a & 
aT) 
von 
NK 
s 


= 


RanGEs oF PoTAMOGETON 


92 Rhodora [Marcu 


Fl. 17: pt. 1: 19 (1909) in large part; sensu Hagstr., Crit. Res. 
Pot. 141 (1916), as to Am. plant, in large part. P. alpinus 
proles heer © (Wolfg.) Graebn. in Engler, Pflanzenr. 4: 
fam. 11: 72 (1907). P. montanense Gandog., Bull. Soc. Bot. 
France 66: 304 (1920). P. microstachys var. typicus Fernald, 
Ruopora 32: 80 (1930). P. alpinus subsp. tenutfolius (Raf.) 
Hultén, Fl. Aleut. Is. 65 (1937) and Fl. Alaska and Yukon 98 
(1941). P. rufescens sensu Am. authors, in large part, not Schrad. 

Streams and cold ponds, southwestern Greenland, Labrador, 
and Hudson Bay to Alaska and the Aleutian Islands, south 
to Newfoundland, Nova Scotia, western Massachusetts and 
Pennsylvania, Colorado and California. Map 2. e fol- 
lowing, selected from many specimens, are representative: 
GREENLAND: Itivnera, 64° 22’ N., PS apne Fjord, Aug. 19, 
1931, M. P. Porsild; ‘Qagssiarssuk, 60° 53’ N., Igaliko-Fjord, 
Aug. 5, 1925, A. E. re a bee Tpaliko” July 23, 1888, 
L. Kolderup Rosenvin ae LaBRADOR: Rama, A. St ecker 332; 
Seal L., n. Lab., Spreadborough 16429; Grand Falls of Hamilton 
Ry M. T. Doutt 3296. NEWFOUNDLAND: Little Quirpon, 
Quirpon Harbor, Wiegand & Hotchkiss 27337; Highlands Brook 
above pond, Crabbes, R. B. Kennedy 81; shores of Conception 
Bay, reid aon Fernald & Wiegan d 4473; Grand Falls, 
Valley of Exploits River, Fernald, Wiegand & Darlington 4474; 
Lookout Mountain, West Arm (South Arm of charts), Bonne 
Bay, Fernald, Long & Fogg 1207; Grand L., Bay of Islands, 
AG. Waghorne 6; St. Georges P., "between Bay St. George and 
Bay of Islands, Fernald & Wiegand 2443 (approaching var. 
subellipticus). QUEBEC: Nacaiiian, Saguenay Co., St. John 
90083 & 90084; Lac Duguay, Newport, Gaspé Co., Proulx 58; 
Riv. Mistassin, (Michaux Herb., Paris Museum, as P. lucens, 
TYPE of P. tenuifolius Raf., as to plant on right) ; Lac William, 
Mégantic Co., Victorin 11162; shallow lagoon, head of Sargents 
Bay, Lake Memphremagog, Aug. 3, 1903, Churchill; Bolton 
Center, Sherbrooke Co., Pease 26709; La Sarre, Abitibi, Louis- 
Marie 313; lac tourbeux, Ville Montel, Abitibi Region, Victorin, | 
Rolland & Meilleur 43 780. Anticostt: Anticosti I., Victorin & 
Rolland 27630. New Brunswick: Junction of Restigouche and 
Matapedia Rivers, Rousseau 32332; Bass R., Nepisiquit, July 30, 
1873, J. Fowler; St. John R., Connors, ’Pease 2907. Nova 
Scotia: Truro, Colchester Co., Bean & White 22963; Mahone 
Bay, Hamilton 64010. MAINE: White’s Brook, Seven Islands, 
T. 13, R. 14 & 15, valley of the St. John R., Aroostook Co., St. 
John & Nichols 21045; St. John R., Madawaska, Fernald 117; 
Piscataquis R., Dover, Aug. 27, 1894, Fernald; Mattagodus 
Stream, TT redwell School, Prctiting Penobscot Co., ‘Steinmetz $23; 
Haley Pond, Rangeley, Franklin Co., Sept. 1, 1894, K. Furbish; 
Pembroke, Washington Co., Fernald 1622; Sydney, Kennebe¢ | 


1943] Ogden,—The Broad-leaved Species of Potamogeton 93 


Co., Fernald & Long 12388. New Hampsuire: Colebrook, 
Cods Co., Pease & Fernald 17024 (app. v. reel ENE Horse- 
shoe P., Northumberland, Coés Co., Pease 17270 (app. v. subel- 
lipticus) ; Cherry P., Jdleeion; ‘Coés Co., July 1829, J. W. 

Robbins; bayou of the Connecticut R.., Hanover, July 12, 1910, 
E. tams (flaccid form). VER RMO ONT: Harve ey’s P., W. 
Barnet, Sept. 7, 1885, F. Blanchard; Windsor, Sept. 6, 1881, 
Geo. Leland; “fe ‘flumine Passumpsic,” A. Wood (original collec- 
tion of P. obrutus). MaAssacuusEtts: Richmond, Sept. 19, 1864, 

Robbins. New York: Preble, Cortland Co., July 31, 1886, 
Dudley; Paradox L., Muenscher & Lindsey 2712; Black R.., 
Dexter, Jefferson Co., Fernald, Wiegand & Eames 14082; n. of 
R. R. bridge, Cayuga, Cayuga Co., Oct. 1886, W. R. Dudley 
(mixed with P. illinoensis in US); Oxford, seo 30, 1886, F. V. 


Coville. PENNSYLVANIA: near Easton, Aug. 4, 1869, Thos. C: 
Porter. Ontario: Cross L. Portage, decaaeain Forest Roggio 
Krotkov 5142 (app. v. subellipticus); Current R., July 0, 1869, 
John Macoun; “Bruce Co., Peninsula,’ 1871, Joke Maw 


Micuican: Isle Royale, W. S. Cooper, no. “954 in part”; Dead 
R., Marquette, C. F. Wheeler 26; 3 mi. n. e. of Watersmeet, 
Gogebic Co., Bessey B & D 2781; Beaver R., Petoskey, July 2, 
1878, E. J. Hill; Alma, Aug. 11, 1893, C. A. Davis; Chatham, 
C.F. Wheeler 90. Wisconsin: White Sand L., Vilas Co. , Aug. 11, 
1930, J. P. E. Morrison; Minocqua, Oneida Co., Fassett fap 
between McNaughton and Rhinelander on Wis 
Cheney 1501. Manrrosa: Churchill, Polunin 1976, 1977, _ 
2062. Minnesota: Lake Kilpatrick, Cass Co., Aug. 1893, 
C. H. Bullard (app. v. subellipticus); n. end of Squaw L., Clear- 
water Co., Moyle 894 (app. v. subellipticus); SouTH DaKora: 
Boulder Creek, Boulder Canyon, Lawrence Co., Over 13817 (app. 
Vv. subellipticus). MAcKENZIE: Setidgi L., 68° 28’ N., 132°: 20° 
W., Aug. 21, A. BE. & R. T. Porsild; McTavish ey Great 
Bear L., 66° 23’ N., 117° 40’ W., A. E. & R. T. Porsild. Sas- 
KATCHEWAN: Cornwall Bay, te Athabasca, 59° 27’ 30” N., 
108° 27’ 30” W., Raup 6618, 6621, & 6622. Archibald R., 
vicinity of Wolverine Ptiyki Athabasca, 59° 9’ N., 108° 25’ W., 
aup 6741. ALBERTA: Slave Lake Dist., Brinkman 4541; Ver- 
million Lakes, Banff, Macoun 4365. MONTANA: eect 
Lakes, Maguire 472; Swift Current Creek, Maguire 474; Trou 
Lake, Maguire & Piranian 5442; Rost L. , Big Fork, Whitford 258: 
Swift Current Creek, below Lake McDermott, Glacier Nat’l 
Park, Standley 16855; Swan R. at Elbow (Lindberg) L., Mission 
Range, Missoula Co., G. B. & R. “ Rossbach 16. Ipano: 
Kootenai Co., Sept. 1887, J. H. Sandberg. Wvyomtna: Shoshone 
Lake, Yellowstone Nat’l Park, Padberg. & Bessey 3724 (TYPE no. 
of P. montanense Gandog.) & 3725; Heart Lake Creek, Yellow- 
stone region, Sept. 3, 1878, C. Richardson (mixed with v. subel- 


° 


94 Rhodora [Marcu 


lipticus in US). Cotorapo: Lake Eldora, Boulder Co., Clokey 
3118; Georgetown, M. E. Jones 734; Walton Creek, Routt Co., 
July 1891, A. Eastwood; vicinity of Twin Lakes, July—Aug., 
1902, C. Juday; Seven Lakes, F. E. & E. S. Clements 491; Grand 
Lake, Shear & Bessey 5328; Howe P. O., Larimer Co., Osterhout 
2885; Tomichi R., Parlin, Gunnison Co., Aug. 20, 1901, Benj. H. 
Smith. Urvan: Clayton Peak, Wasatch Mts., Aug. 12-26, 1903, 
S. G. Stokes; Twin Lakes, Alta, Wasatch Mts., M. E. Jones 1297; 
Silver L. at Brighton, Big Cottonwood Canyon, Salt Lake Co., 
Maguire & Richards 13156; Bridger L., Uinta Mts., Summit Co., 
Rollins 2319; Brighton, M. E. Jones 6606. CALIFORNIA: North 
Fork of Kings R., Hall & Chandler 563, region of Kaweah Peaks, 
Funston’s Meadows, Tulare Co., Dudley 2201; Webber L., Sierra 
Co., Aug. 3, 1894, Dudley; Silver Valley, Alpine Co., Brewer 1978. 
OREGON: in a warm spring, Harney Valley, June 10, 1885, T. 
Howell (mixed with P. illinoensis). Wasuineron: Trout Creek, 
w. Klickitat Co., Suksdorf 2172; Falcon Valley, Mt. Adams, Sept. 
1879, Suksdorf; Baker L., Whatcom Co., Muenscher 7657a & 
7658. British Cotumstia: Kicking Horse L., Rocky Mts., Aug. 
17, 1890, John Macoun 4162; Revelstoke, John Macoun 4163; 
Barkerville, McCabe 43; Nanaimo, Vancouver I., John Macoun 


78321. AuasKa: Goldstream Cr. and Pedro Dome, 65° 0’ N., 
147° 30’ W., A. E. & R. T. Porsild 114; Buckland R., 65° 55’ N., 
161° 0’ W., A. E. & R. T. Porsild 1496; Afognak L., Afognak 
Island, Shelikof Strait off Alaska Pen., Aug. 1931, W. H. Rich; 
Kodiak Island, Trelease 2870 & 2871; Shumagin Islands, Saun- 
ders 2869; False Pass, Unimak Island, Aleutian Islands, Zyerdam 
2141; Unalaska (type-locality of P. microstachys), Van Dyke 202, 
also Eyerdam 2360, and Hultén 7573; Seldovia, Piper) 4426; 
Kukak Bay, Alaska Pen., Coville & Kearney 1564; Olga Bay; 
Kodiak Island, Z. H. & H. B. Looff 1501. 


2b. P. atpinus Balbis var. subellipticus (Fernald), comb. nov. 
P. microstachys var. subellipticus Fernald, Ruopora 32: 
(1930). _P. alpinus, sensu Morong Mem. Torr. Club 3: no. 2:19 
(1893), in small part; sensu Graebn., in Engler, Pflanzenr. 4: 
fam. 11: 70 (1907), as to N. Am. plant in small part; sensu 
Taylor, N. Am. Fl. 17: pt. 1: 19 (1909), in small part; sensu 4 
ue ye nts a Pot. 141 (1916), as to Am. plant in or 
part. . tenuifolius var. subellipti RA dv: 
B11 (198i). pticus Fernald, RHopo 
Shallow pools and slow streams, Newfoundland, south to 
southern Vermont and eastern New York, sparingly westward tO — 
Wyoming and Montana. Map. 3. The following are referr 
here: NEWFOUNDLAND: Salmonier R., 1931, Agnes Ayre; Flowet — 
Cove, Straits of Belle Isle, Fernald & Long 26221, also. Hotchkiss — 


1943] Ogden,—The Broad-leaved Species of Potamogeton 95 


27338; Stephenville, region of Bay St. George, F yest Wiegand 
& _ Kittredge 2442; Port a Port, Mackenzie & Griscom 10047. 
EC: Ile & la Proie, Archipel_ de Mingan, Victorin ao Rolland 
20462: Locked Camp, Riviére Cap Chat, Matane Co., Victorin, 
Rolland & Jacques 44451; Maria, Bonaventure Co., Victorin, 
Rolland & Jacques 33315; between Baldé and the Baie des 
Chaleurs, Bonaventure R., "Bonaventure Co., Collins, Fernald & 
Pease 591 i Georgeville, "Pease 1919; Charlton Island, about 
52° N., 79° 30’ W., James Bay, A. E. Porsild 4296; Lac Philippe, 
Ladysmith, Pontiae Co., Gauthier 2446. Anticosti: Riviére 
Sainte-Marie, Victorin & Rolland 25938; Riviére au Saumon, 
Victorin, Rolland & Louis-Marie 20467. MaAGpALEN ISLANDS: 
East Cape and East Point, Coffin Island, Fernald, Long & St. 
John 6766, (TYPE in Gray Herb. ); Ile de Havre-au-Ber, Victorin 
& Rolland 9928. New Brunswick: Eel R., Dalhousie, Svenson 
& Fassett 3028. Nova Scotia: Baddeck Bay, Victoria Co., 
Fernald & Long 19687. Matne: Houlton, Aroostook Co., 1881, 
K. Furbish, ee Fernald & Long nahh begin? Piscataquis 
Co., Sept. 5, 1894, Sept. 15, 1894 & Aug. 31, 1896, Fernald. 
VERMONT: Little Leech Pond, Averill, oe Co., Eggleston 1656; 
Willoughby, Orleans Co., Aug. 4& 11, 1881, E. Fazon; “Nigger” 
P., Westmore, Orleans Co., Eames & Godfrey 9334; West Burke, 
Redfield 8014; Evart’s P., Windsor, Aug. 27, 1933, Weatherby & 
Griscom. New York: Minerva Brook, east of Minerva, Essex 
Co., House 15182; Niagara Falls, Aug. 21, 1886, Morong; Para- 
dox L., Essex Co., Killip 12605 (mixed with P. pacts Sree in 
US). ONTARIO: Howdenvale, Watson 3144 & 3146; River 
Trent, John Macoun 4168; Batchawana Falls, Algoma Dist., 
Taylor et al. 294. MicnigAn: Eagle Harbor, Keweenaw Pen., 
1863, Robbins; North Cliff, Keweenaw Co., Aug. 1, 1888, O. A. 
Farwell; west branch Ontonagon Ri. endaetoat| Li: Gogebic Co., 
Hotchkiss & Koehle r 4849. Wisconsin: Three Lakes, Oneida 
Co., Aug. 5, 1918, R. Hoffman; n. w. end of Big Arbor Vitae L., 
Vilas Co., Hotchkiss & so btig 4460. Minnesota: Lake il- 
patrick, Gass Co., Aug. 1893, C. A. Ballard; Gull L., Cass Co., 
Aug. 1893, A. P. prep ii SASKATCHEWAN: Archibald Re 
Meg of Wolverine Pt., L. Athabasca, about 59° 9’ N., 108° 
5’ W., Raup 6742. Montana: Lake Josephine, Glacier Nat'l 
Park Maguire 478. Ipano: west end of Fernan L., Coeur 
d’ Alene, Rust 384. Wyomina: Heart Lake Creek, Yellowstone 
region (mixed with v. tenuifolius in US; sheet in G not mixed) ; 
e. fork of Big Sandy, Wind River Mts., ‘all 1878, C. Richardson. 
British Cotumsia: Austey Creek, Shuswap L., J. M. Macoun 
4166. 


In 1930, Fernald showed that the American and eastern 
Asiatic plant which had been passing as P. alpinus Balbis of 


98 Rhodora [Marcu 


even stranded along the shore, while many specimens of var. 
subellipticus show evidences of having developed in deep water. 


3. P. POLYGONIFOLIUS Pourret 

Ruizome buff or pinkish, often with arate ete about 
same thickness as stem. Stem simple, .7—-2 m n diameter; 
stele with the proto-type pattern; endodermis came: of O-cells; 
interlacunar bundles absent; subepidermal bundles present; 
pseudo-hypodermis 1 or 2 cells thick. SuBMERSED LEAVES 
(usually absent) with blades lanceolate, 3-10 cm. long, .5-1.5 
cm. wide, aioe gradually at both ends, apex acutish but not 
sharp-pointed, petiole 1-3 cm. long; nerves 7-11, outer ones 
marginal; lacunae rather obscure, of 2—4 hdd each side of mid- 
rib; margin entire. FLOATING LEAVES coriaceous, ovate, apex 
tapering to an obtuse tip, base rounded or slightly cordate, peti- 
oles 1-15 cm. long, blades 3-8 (-9) em. long, 1-4 cm. wide, 
nerves (11—) 15-19 (—21), all of about equal prominence; lacunae 
none. StipuLes about 3 em. long, somewhat persistent, without 
keels or practically so. PEpuNCLES about same thickness as 
stem, 4-12 cm. long. Spikes with about 10 whorls, in fruit 2-3 


cm. long, .5-.7 em. wide. FLOowers sessile or nearly so; sepalany = 


connectives greenish, blades orbicular or elliptical, 1-2 m 
wide, claws .3-.7 mm. long; anthers .6—.8 mm. long. Fruits 4 
obovate to orbicular, rounded at base, sides with a depression, — 
especially if immature, (1.6—) 2-2.5 mm. long, (1.2—) 1.5-2.1 mm. 
wide; beak minute or obsolete; keels absent or nearly so; exocarp 
reddish; endocarp loop often ‘with a large cavity; apex of see 
pointing toward the basal end or slightly above. 

P. po soem foliue Pourr., Mém. Acad. Toulouse 3: 325 (1788); 
Benn., Bot. Gaz. 32: 58 (1901); Graebn. in Engler. Pflanzenr., — 
4: fam. 11: 65 (1907); paige N. Am. Fl. 17: pt. 1: 21 (1909); 
sca Crit. Res. Pot. 175 5 (1916). 


gt Arséne At Nov VA je pba Sable Tuand. 43° 59! N., 
Tow; St John 1121 & 1 


ae polygonifolius ranges Bibi Europe, Algeria, Morocco, q 
Madeira and the Azores', but appears in the western hemisphere — 


} The old world range of this species is taken from Hagstrém, Crit. Res. Pot. 178° - 


os It is also given a widespread Asiatic range by Bennett, Bot. Gaz. 32: 59 
1) 


1943] Ogden,—The Broad-leaved Species of Potamogeton 99 


only in eastern Newfoundland, the island of St. Pierre close by, 
and Sable Island off Nova Scotia. It may be presumed that this 
species in America, now confined to areas that largely escaped 
the last continental glaciation, once had a much wider range. 

This species has a strong tendency to produce floating leaves 
which often spring almost directly from the rhizome. Terres- 
trial forms are common and in such condition may survive in 
ponds that dry up for a part of the summer.! 


4. P. ampuirouius Tuckerman 


Ruwi1zoMeEs whitish or often with a reddish cast, 2-4 mm. in 
diameter, scales black, broadly obtuse. Stem terete, 1-3.5 mm. 
in diameter, simple or ‘forming short branches late in the season; 
stele with the proto-type pattern; endodermis of O-cells; inter- 
lacunar bundles present throughout; subepidermal bundles 
absent; pseudo-hypodermis 1 or 2 cells thick. SuBMERSED 
LEAVES (excluding transition leaves) of two more or less distinct 
types: those on the lower part of the stem lanceolate, dark green 
and usually badly decayed by the time the floating leaves appear, 
with 19-25 nerves: those of the upper part of the stem broadly 
lanceolate to ovate with margins much longer than midrib 
giving those leaves just below the floating ones a characteristic 
arcuate appearance, with 23-87 nerves; both types obtuse or 
acutish at apex, but never sharp-pointed, ‘and —— to sper 
1-6 cm. ae blades 8-20 cm. long, 2.5-7.5 em. wide; margins 
entire; lacunae 3-6 rows each side of midrib. rome ee 

with gradual transitions from the submersed ones, coriaceous, 
opaque, ovate to elliptical, apex rounded or bluntly mucronate 
base cuneate or rounded, aes 8-20 cm. long, bla 5-10 
em. long, 2.5-5 cm. wide, with (21—) 29-41 (-51) Covel, about 
\4 of the nerves more nacre than the others, as seen by 
transmitted light; lacunae none. Srtrputes of the oa inne 
leaves somewhat persistent, fibrous, triangular, obtuse when 
young, becoming twistedly stringy with age, 3.5-11 cm. long, 
obscurely 2-keeled; those of the floating leaves similar, 5-12 
(-18.5) em. long, with 2 fairly well marked keel-nerves and 
30—40 fine nerves. PEDUNCLEs at base about same thickness as 
stem, but usually thicker at the middle or upper half, 4-30 cm. 
long. Sprkes with 9-16 whorls; in fruit 4-8 cm. long, 1—1.5 cm. 
thick. FLowers sessile or on very short pedicels up to 1 mm. 
long; sepaloid connectives usually with a — ari sometimes 
greenish, blades orbicular to elliptical, (1.2—) 1 3 (- —3.5) mm. 
wide, claws .4-.8 (-1) mm. long; anthers .8-1. 5 2) mm. long. 


1 See Harold St. John, Sable Island, etc., Proc. Bost. Soc. Nat. Hist. 36: no. 1: 59 
(1921). 


100 Rhodora [Marcu 


Fruits obovate, rounded on the back, cuneate at base, sides 
flat, 3.5-4.5 (-5) mm. long (excluding beak), 2.5-3.3 (-3.7) mm. 
wide, beak often prominent, up to 1 mm. long; keels usually 
prominent but sometimes rounded or obscure, the dorsal one 
often strongly developed, especially at the middle; exocarp 
usually reddish or orange-brown if fully mature, otherwise 
greenish; endocarp with 3 prominent, acutish and often some- 
what muricate keels, beak linear, facial, about 1 mm. long, loop 
solid; apex of seed pointing .6-1.5 mm. above the basal end. 
Robust plants characterized by several large arcuate submersed 
leaves clustered just below the water surface, and floating leaves 
with numerous nerves. 

P. amplifolius Tuckerm., Am. Journ. Sci. ser. 2: 6: 225 (1848); 
Morong, Mem. Torr. Club. 3: no. 2: 16 (1893); Graebner in 
Engler, Pflanzenr. 4: fam. 11: 67 (1907); Taylor, N. Am. Fl. 17: 
pt. 1: 18 (1909); Hagstr., Crit. Res. Pot. 163 (1916). P. lucens 


var. ? fluitans (Roth) Gray, Man. ed. 2: 435 (1856), as to plants — 


included in part, not P. fluitans Roth. ?P. amplifolius forma 


amphibius Benn., Journ. Bot. 42: 70 (1904). ?P. amplifolius — 


var. ovalifolius Morong ex Benn., Journ. Bot. 42: 70 (1904); 
? Graebn., in Engler, Pflanzenr. 4: fam. 11: 68 (1907). ?P. 
amplifolius var. amphibius (Benn.) Graebn. in Engler, Pflanzenr. 


. fam. 11: 68 (1907). Spirillus amplifolius (Tuckerm.) Nieuwl., — 


m. Mid. Nat. 3: 16 (1913). P. amplifolius forma homophyllus 
Hagstr., Crit. Res. Pot. 163 (1916). 

Lakes and streams, usually in deep water. Newfoundland, 
south to Virginia and Tennessee, west to Missouri and Kansas, 
also in California, Oregon and Washington. Mar 5. The follow- 
ing, selected from many specimens, are representative: NEW- 
FOUNDLAND: Salmonier R., 1931, Ayre; Frenchman’s Cove, Bay 
of Islands, Mackenzie & Griscom 10045. QurBeEc: Lac Sainte- 


Bap il 


Anne, Victorin, Rolland & Jacques 33517; New Richmond, Bona- . 


venture Co., Victorin, Rolland & Jacques 33858; Lake Mem- 
phremagog, July 29, 1902, J. R. Churchill; Riviére-aux-Serpents, 

ka, Victorin 20457, 21201 & 25815; East Templeton, John 
Macoun 85584; East Templeton near Hull, Malte 118267 & 
118268. McGregor L., J. M. Macoun 86002; Lac Lepéche, 
Rolland 13044; Otter L., Pontiac Co., Gauthier 2422; Lac Donald- 
son, Buckingham, Rouleau 7231; Kondiaronk (Lac Creux); 
Victorin 16065. Nova Scotta: Mill Brook, Sheffield’s Mills, 
Kings Co., Aug. 24, 1902, Fernald; Young’s L., North Mt: 
Belle Isle, Annapolis Co., Fernald, Bartram, Long & Fassett 23139; 


Lily L., Sandy Cove, Digby Co., Fernald & Long 19688; Little _ 


Meteghan L., Digby Co., Fernald & Long 23140; Sloane L., 


Pleasant Valley, Yarmouth Co., Fernald, et al. 19689. MaiNE: 
Saint Francis R., Aroostook Co., Aug. 13, 1902, Eggleston & 


Fernald; St. John P., T. 6, R. 17, Somerset Co., St. John & 


1943] Ogden,—The Broad-leaved Species of Potamogeton 101 


Nichols 2108; Eagle L. Del wig Co., Ogden 1704; E. Edding- 
ton, Penobscot Co. , Aug. 2 , 1897, Fernald; Harvey’s P., Levant, 
Penobscot Co., Ogden t ’ Marston 1698, Plantae Exsiccatae 
Grayanae 904; Loon L., Franklin Co., 1894, Furbish; Middle P., 
. Poland, Androscoggin Co., Sept. 1893, Furbish. New Ha AMP- 
SHIRE: Lime Pond, Columbia, Coés Co., Pease 13883; York P., 
Kilkenny, Coés Co., Pease 17185; Burns P., Whitefield, Coés 
Co., Pease 14585, also Moore 5036; Wolfboro, Carroll CoRR: A. 
Ware 38326; Squaw Cove, Squam e , Sandwi ch, Carroll Co. , July 
21, 1923, Svenson; Dublin, Cheshire Co., Aug. 8, 1931, Eaton & 
Griscom; Johnson Creek, Madbury, Strafford Co., Hodgdon 2662. 
VERMONT: Little Leech P., Averill, Essex Co., ‘Eggleston 1655; 
Long Pond, Willoughby, Orleans Co., spi 26, 1896, Kennedy; 
Willoughby L., Aug. 31, 1917, E. J. Win low; Missisquoi Bay, 
Lake Champlain, Swanton, Franklin Co., Get 3 , 1909, A. E. 
Blewitt; Spectacle L., Rutland Co., July 21, 1907, EB. C. Kent; 
Big P., Rutland Co., Aug. 29, 1895, F.C. Kent; Chittenden, 
Aug. 18, 1895, Eggleston. Massacuuserts: Wenham , Wen- 
ham, Essex Co., July 18, 1895, J. H. Sears; Fresh P., Cambridge, 
Middlesex Co., many collectors: Concord R., Billerica, Middlesex 
Co. , Fernald & Abbe 2506; Sudbu ry R., Conco rd, Middlesex Co., 
Aug. 4, 1886, Morong, also Aug. 4, 1886, Deane, also Ogden & 
Wiggins 1729; Wood’s P. | Wellesley, Norfolk Co., Aug. 1, 1881, 
Morong; Middleboro, Plymouth Co., Aug. 12, 1 1901, J. Murdock: 
Quinsigamond P., Worcester, June 2 1864, Robbins; Great 
Brook, Southwick, Hampden Co., Seymour 249; Spectacle P;; 
Sandisfield, Berkshire Co., June 27, 1912, R. Hoffmann; 
mont, Standley & Killip 7648. Ruope Isuanp: Pawtuxet R., 
Cranston, eke nie Co., July 6, 1898, J. F. Collins; Apponaug 
" Kings ” TApponaug Pond is in the town of Wa rwick] 
Wankicue. yor Aug. 26, 1880, Morong. CoNNEcTICUT: 
Thompson, Windham Co., June 10, 1922, Eaton & Fassett; East 
Granby, Hartford Co., Weatherby D2409; Hamlins P., Plainville 
Hartford Co., Blewitt. 632; Quinnipaug P., Guilford, New ae 


— or 1886, W. R. Dudley; Seldens Cove, Lyme, A ae, 
, Citi Bissell; Mudge P., Sharon, Weatherby & ree erson 
son Colebrook, Aug. 13, 18 50, J. W. Robbins; West Goshen, 


Aug. 1892, L. M. Underwood; Twin Lakes Station, Salisbury, 
Litehfield Co., Aug. 20, 1935, Fernald & Ogden. New York: 
Mouth of Perch River, Jefferson Co., Muenscher & Maguire 1600; 
Spencer Lake, Spencer, Tioga Co., Eames & Wiegand 11172; 
Pierrepont P., Woodville, House 8838, also Ogden & Bolan 1583; 
Indian L., Franklin Co., Muenscher & Maguire 719; Eagle L. 
Essex Co., Killip 12610; Bemus Point, Lake Chautauqua, Aug. 
8, 1896, Churchill; Edick Creek near Sears P., Lewis Co., Hotch- 
kiss 2646; Rockl and L., July 17, 1892, Morong; Carpe nters r, 
Fabius, Onondaga Co., House 1329; Waldorf F., & Of Nor th 


102, Rhodora [Marcu 


Chatham, Columbia Co., House 21752; Pine P. (Gypsy L.), 
Putnam Co., Muenscher & Curtis 5423; Sodus Bay, Killip 12261; 
Lake Luzerne, Luzerne, Warren Co., Fogg 4934; Hudson R., 
Coveville, Saratoga Co., Muenscher & Lindsey 2723; North 
Harpersfield, Delaware Co., Topping 203; near Peekskill, West- 
chester Co., 1868, LeRoy. New Jrersry: Delaware R., Camden, 
Oct. 7, 1877, C. F. Parker; Pensauken, Camden Co., Adams 
& Trudell 378; Swartswood L., Sussex Co., Griseom & Mac- 
kenzie 10679; Cranberry L., Sussex Co., Mackenzie 2305; Sparta, 

ussex Co., Sept. 13, 1887, N. L. Britton; Hackensac R., 
Bergen Co., July 25, 1861, C. F. Austin. PENNSYLVANIA: Mar- 
tin’s Creek, Northampton Co., Aug. 29, 1906, C. S. Williamson; 
Lehigh above Easton, June 15, 1869, Thos. C. Porter; Brandy- 
wine Creek, Icedale, Chester Co., Sept. 18, 1927, H. E. Stone. 
District oF Cotumsia: near Washington, Coville 129; Eastern 
Branch below Navy Yard, Sept. 1, 1900, EZ. S. Steele. MARYLAND: 


Mouth of Mill Creek, Chesapeake Bay Region, G. H. Shull 97. 4 


West Virainia: Minnehaha Springs, Pocahontas Co., July 31, 


1930, W. V. U. Bot. Exped.; also E. E. Berkley 1887; Huttons- — 


ville, Randolph Co., Moore 2488. Vriretnta: Four Mile Run, 
Shull 413 & 473; June 6, 1899, E. S. Steele; Dyke, Fairfax Co., 
Metcalf & Sperry 1621, 1634, & 1639; Hunting Creek, Dist. of 


Columbia and vicinity, McAtee 2237 & 2279. Grorata: Lafay- — 
ette, Walker Co., Wilson 189. Ontario: Plevna, Aug. 19, 1902, — 
J. Fowler; Epilobium Bay, Bear Island, Watson 1178; Whitney — 


L., 12 mi. n. of Temagami, Kane 1030; Bass Creek, Franks Bay, 
Lake Nipissing, Chitty 319; McGregor Bay, Manitoulin District, 


Ogden & Bolan 1647; Gore Bay, Manitoulin Island, Pease & — 
Ogden 25032; Old Woman’s R., Bruce Pen., Krotkov 8629; Gull L., 


Addington Co., July 15, 1870, John Macoun; Belleville, Hastings 


Co., July 1876, John Macoun; Lac Meach, Ottawa, Rolland 8697. : 
Micutean: Isle Royale, Cooper 93 & 215; Lake Manganese, Cop- — 
per Harbor, Keweenaw Co., Hermann 8233; Carp R., Porcupine 


Mts., Aug. 16, 1923, H. T. Darlington; Crystal Falls, Iron Co., 


Metcalf 2242, 2250, 2254 & 2255; Michigamme L., Iron Co., — 
Metcalf 2209 & 2210; Whitefish L., Mackinac Co., Metcalf 2321; _ 
Chassell, Houghton Co., Pease & Ogden 25167; Black L., Che- — 
boygan Co., Aug. 3, 1935, Gleason; Little Manistee R., Manistee, — 
Aug. 8, 1882, Morong; Washtenaw Co., July 17, 1838, Houghton; — 
Mill-pond, Alma, Aug. 11, 1893, C. A. Davis; Kalamazoo R., — 
Allegan Co., Wight 97 & 109; Kimble L., Vicksburg, Kalamaz00 — 


Co., July 5 & 28, F. W. Rapp; Sister Lakes, Van Buren Co.; 


De Selm 24 (F, mixed with P. natans); Papaw L., Berrien C0. _ 
Dodge 172; Haslet, Yuncker 361. Onto: Geauga L., Portage Co. — 


Webb 546; Phalanx, Trumbull Co., Webb 452; Cowles Creek 


Marsh, Ashtabula Co., Aug. 15, 1894, Goodrich 209; East Harbor, 


Ottawa Co., Aug. 10, 1898, E. L. Moseley; Put-in-Bay, Aug 


Se a aera aman SR RN RE abel FERRIER sa Cn eR SE em A ROR eS em 


23 STt 


1943] Ogden,—The Broad-leaved Species of Potamogeton 103 


1898, A. J. Pieters. InpiaAnNa: Lake Maxinkuckee, Scovell & 
Clark 1223, also Clark 5 & 10, also Scovell 26 (US, mixed with P. 
natans) & 28, also Evermann 1223; 4.3 mi. south of Hayden, 
Jennings Co., Deam 48399; Loon L., Whitley Co., Deam 49399; 
Lake Manitou, Rochester, Fulton Co., Deam 56490; Wolf L., 
Whiting, Lake Co., Deam 56616; Wolf L., Lake Co., Aug. 18, 
1920, D. C. Peattie; Hamilton L., Hamilton, Steuben Co., 
Deam 56683; Pine L., 2 mi. n. w. of La Porte, La Porte Co., 
Deam 52425. Kentucky: a specimen without locality data 
except ‘‘C. W. Short, M.D., Lexington, Kentucky.” aplee | 
TENNESSEE: Mill-Sharp Lane, Lost Creek, Union Co., 

1933, J. K. Underwood. ALABAMA: Larkin Fork of Peat oat 
RB: , Jackson Co., June 25, 1932, Harper. Wisconsin: Green Bay, 
June 29, 1890, Schuette; "Madison, July 30, 1889, Trelease; Lake 
Mendota, Madison, Dane Co., July 26, 1905, A. B. Stout; Lake 
Mendota, Aug. 20, 1912, R. H. Denniston; Hill ee Minocqua, 
Oneida Co., Fassett 5362: Deer L., Polk Co., Aug. 1892, Pei. 
Burglehaus; Pell L., Bloomfield Twp., Walworth Co., Hotchkiss 
& Koehler 4192; Kakagon Slough, Odanah, Sanborn Twp., Ash- 
land Co., Hotchkiss & Koehler 4353; Pewaukee L., Milwaukee 
Co., Aug. 1884, H. E. Hasse; near Lac Vieux Desert, Cheney 
1785. Inutnors: Lake Villa, Lake Co., Gleason & Shobe 176; 
Wolf L., Chicago, Chase 1710; Chicago R., Cook Co., July 21, 
1906, F. ¢. Gates; Englewood, June 15—July 22, 1880, herb. C. E. 
Faxon, also July 22, 1880, E. J. Hill; Dupage ss; Warrenville 

at 


Oosting 3275; Cedar L., Rice Co., Oct. 22, 1891, E. Douglas; 
Lake Koronis, Stearns Co., Kubichek 115b (US, mixed with P. 
natans) & 120; Centre City, Sandberg 658; Chisago L., Metcalf 
1140 & 1160; Center City, Sandberg 656; Basswood is, Lake Co.; = 
Metcalf 151 1; Tait L., Kandiyohi Co., ’ Metcalf 2024: Leech L., 

Cass Co., Shunk & Mc anning 353: Mille Lacs L., Mille Lacs Co., “4 
Keck & Stevens 332; Dudley L., Keck & Stilwill 377; L. Minne- 
tonka, Keck & Stibwill 433; ae Louis L., Stearns Co., Linsdale & 
Keck 5h: Map Wright Co., Linsdale & Keck 115; Lake 
Francis at Hiya, ‘Le ie Co., M ae 4 2261. Iowa: Clear is 


May. 7, 1893, H. Buen (M, raid with P. pulche r); Ha-ha- 
tonka, Camden Co., Metcalf 929; Gravois Mills, Morgan Co., 
Metcalf 888, also Steyermark 28071; Brice, Laclede Co., Geo. 

oore 81; Nian ngua R., near Bennett Spring ete Park, Dallas 
Co., Steyermark 13782: Eleven Point R., abot mi. n. e. of 
Peace Valley, Howell Co., Steyermark i448? eek’. Fork of 
Current R., 5 mi. s. e. of Arroll, Texas Co., paseo 14624; 
near Double Spring (Rainbow Spring ), 4 mi. n. e. of Dormis and 

mi. s. e. of Dora, Ozark Co., Steyermark 15560. ARKANSAS: 


104 Rhodora [Marcu 


Paragould, Green Co., June 27, 1893, H. Eggert; Powers Creek, 
2 mi. s. w. of Batesville, Coville 189. Norta Daxota: Antler 
Creek, Antler, Bergman 2484. Sours Daxora: Lake Hendricks, 
Brookings Co., Griffiths & Schlosser 4; Little Minn. R., Roberts 
Co., Over 15425 & 17142; Pickerel L., Day Co., Over 14459 & 
17141. Nepraska: Lincoln, Webber 4; Greenwood, T. A. 
Williams 344; Antelope Creek, Cedar Co., Clements 2979. 
Kansas: Comanche Co,, A. S. Hitchcock 1000; Ellis, July 1876, 
S. Watson; Decatur Co., A. S. Hitchcock 1099; Spring P., 
Cottonwood Springs, Chase Co., Aug. 1870, E. Hall. Oxkta- 
HOMA: Sapulpa, Bush 1318; Cache, Comanche Co., Stevens 1364 
US, mixed with P. nodosus). Montana: Fish L., Glacier 
Nat’l Park, Standley 18528. Ivano: Priest L., Piper 3684, also 
MacDougal 303; Alturus L., Evermann 493. Ca.irornta: Red 
L. above Soda Springs of the San Joaquin, Aug. 16, 1899, J. W. 
Congdon; Satcher L. near Devils’ Post Pile, Madera Co., Ferris 
8845; Webber L., Sierra Co., Aug. 1894, Dudley. OREGON: 
Salem, E. Hall 487, 488 & 488a; Ten-mile L., near Lakeside, 
Coos Co., Peck 9026; Sauvies Island, Thomas Howell 1498. 
WasuIncTon: Olympic Mts. near the coast, Clallam Co., A. D. 
E. Elmer 2798; Pullman, Henderson 2474; 28 mi. south of Tacoma 
on the Mt. Rainier Rd., Pierce Co., Abrams 9241; Blakeley 
9. San Juan Islands, S. M. & E. B. Zeller 1238; Lake 


P. amplifolius is a robust plant of ponds and streams, usually 
preferring the deep, clear water. It grows vigorously in either 
acid or alkaline water.. The submersed leaves persist in acid oF 
neutral waters, but if lime is present, it precipitates on the leaves 
to such an extent that they are soon broken off. This precipitate 
forms on P. amplifolius even when such minute traces of lime 
are present that other species show no evidences of it. 

Although for its typical development this species prefers the 
deeper water, whenever its rhizomes creep up onto the muddy ~ 
banks, terrestrial forms are produced. These are always sterile. 

The variety ovalifolius that Bennett credits to Morong 18 
presumably a state of P. amplifolius, but the meager description 
sounds suspiciously like P. pulcher. In view of the fact that — 
Bennett reports P. amplifolius from Arkansas and from Florida’, 
where it is not found, and P. pulcher from Maine? where it does — 


1 Arthur Bennett, Journ. Bot. 42: 77 (1904). 
?—_______—, Journ. Bot. 40: 146 (1902). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 105 


not seem to be, it is possible that he had the identities of these 
plants reversed. His forma amphibius is based in part on re- 
marks of Morong', which definitely refer to P. pulcher. Graebner 
evidently copied Bennett’s names and descriptions without any 
inquiry as to upon what they were based. 
1 Thomas Morong, Mem. Torr. Club. 3: no. 2: 17 (1893). 
(T'o be continued) 


THE BROAD-LEAVED SPECIES OF POTAMOGETON 
OF NORTH AMERICA NORTH OF MEXICO 
E. C. OGDEN 
(Continued from page 105) 
5. P. putcHer Tuckerman 
Ruizome buff, often with dark red spots, .5-1 mm. in diam- 
eter. Stem simple, terete, 1-2.5 mm. in diameter, usually con- 


120 Rhodora [APRIL 


- spicuously ba awed: stele with the proto-type pattern; 
endodermis of O-cells; interlacunar and subepidermal bundles 
absent; pseudo-hy podermis 1 cell thick. StuBmMERsED LEAVES 
(excluding transition leaves) of two more or less distinct types, 
those of the lower part of the stem semi-opaque, oblong with 
rounded apices; those of the upper part of the stem translucent, 
lanceolate to lance-linear, not arcuate, apex acutish but not 
sharp-pointed, ate types tapering at base to short petioles 
(up to 1.5 em. long) or sometimes practically sessile; blades 
8-14 (-18) em. long, 1-2.5 (—3.5) em. wide, nerves (9-) 1 11-21, 
outer ones marginal; margins entire; lacunae 4-8 rows each side 
of midrib. FLoatinGc LEAvEs coriaceous, ovate to rotund; apex 
gegen or bluntly mucronate; base cordate or rounded; petioles 
4-18 iat blades 2-7 (-11) em. long, 1.5-4 (-8.5) cm. wide, 
with (19-) 2 —29 (-35) nerves, all of about equal prominence, 
as seen by Scenes light; lacunae none or very faint. STI- 
PULES of the submersed leaves decaying early, those of the 
floating seen persistent, narrowly triangular, obtuse when 
young, becoming acutish with age, 2-5 cm. long, 2-keeled. 
PrepUNCLEs of rather even thickness ‘throughout, 5-8 (—11) cm. 
long. Spikes with about 10 whorls, in fruit 2-3.5 em. long, . 
1.1 em. thick. FLowers sessile or nearly so; sepaloid connec- 
tives usually with a tigers cast, blades sehiecltat to elliptical, 
(1.2-) 1.5-2.5 (-3) mm. wide, claws, .4-.8 mm. long; anthers — 
.8-1.4 mm. long. Gucie obliquely ovate, sande) or cuneate at — 
base, sides flat or slightly concave, (2. 7-) 3-3.5 (-4) mm. long, 
(2.3- -) 2 .6-3.2 (-3.4) mm. wide; beak often prominent, up to 8 
mm. long; keels usually prominent, acutish, the dorsal one often 
strongly developed, and sometimes with a basal lobe projecting 
below the point of attachment; exocarp mostly light brown, 
sometimes olive-green; endocarp with 3 prominent, acutish an 
somewhat muricate keels, beak linear, facial, about 1 mm. long, — 
loop solid; apex of seed pointing 5-1.2 m m. above the basal end. 
Plants characterized by a ete aye spotted stem, with 
large cordate floating leaves and lanceolate submersed leaves 
which taper rather abruptly to - gt . 
P. pulcher Tuckerm., Am. Jou : 45: 38 (1843); 


Morong, Mem. Torr. Club 3: no. 9: “16 1808), aaa in Engler, — 


Pflanzenr. 4: fam. a 67 (1907); Taylor, N. Am. Fl. 17: pt. 1:21 — 
(1909); Hagstr., Crit. Res. Pot. 152 (1916). P. natans sensu 
Bigel., Fl. Bost. 41 (igi4), according to Tuckerm., Am. Journ. 
Sci. ser. 1: 45: 38 (1843). P. lucens var. ? fluitans (Roth) Gray, — 
Man. ed. 2: 435 (1856), as to plants included in part, not P. 
fluitans Roth. ?P. amplifolius forma amphibius Benn., Journ. — 
aa ee 70 (1904). ?P. amplifolius var. ovalifolius Morong ex | 


, Journ. Bot. 42: 70 (1904); ?Graebn. in Engler, Pflanzent. 


rs tara 11: 68 (1907). ?P. amplifolius var. amphibius (Benn.) - 


1943] Ogden,—The Broad-leaved Species of Potamogeton 121 


Graebn. in Engler, Pflanzenr. 4: fam. 11: 68 (1907). Spirillus 
pulcher Nieuwl., Am. Mid. Nat. 3: 16 Hers P. pulcher forma 
amphibius Hagstr., Crit. Res. Pot. 153 (1916). 

Shallow muddy ‘pools, peaty pond-holes and sluggish streams 
chiefly of the Coastal Plain and Mississippi embayment, southern 
Nova Scotia, southern New Hampshire, south to Georgia, Texas, 
Oklahoma, Arkansas, Missouri and Minnesota. Map 6 The 
following, selected from many specimens, are referred here. 
Nova Scorta: Sears L., New Tusket, Digby Co., Fernald & 
Long 23137; Rhodeniser L., e: of Ridacounters Lunenburg Co., 
Fernald & Long 23138. New Hampsurre: Contoocook R.., 
Jaffrey, Rand & Robinson 1013. MassACHUSETTS: Foster’s Pond, 
Andover, Essex Co., Pease 2011; Waushakum P., Ashland, Mid- 
dlesex Co., June 1879, July 1, 1881, eee 1882 & Aug. 7, 1882, 
Morong; Spot Ps Stoneham, ‘July 852, Robbins; Whitman FP. 
Weymouth, Norfolk Co., ppg Perry ’Sampson’s P., Carver, 
Plymouth Co., Fernald 750; Uncatena, Elizabeth Islands, Dukes 
Co., Fogg 2 997; L. Neeseponset, Dana, Worcester Co., Goodale, 
Markert & Piper 96988; Readville, Suffolk Co., June 16, 1878 & 
June 16, 1880, C. E. Faxon, also June 23, 1879, Morong; Natick, 
June 1, 1881 & Sept. 27, 1881, Morong; Nantucket, Nantucket 
Co., July 1887, Morong. RuopE ISLAND: ponds between Pilot 
Hill iak: Southeast Point, Block Island, Newport Co., Fernald, 
Hunnewell & Long 8443; Apponaug P., Apponaug, Aug. 26, 1880, 

orong. CoN dae: Fairfield, E. H. Eames 8740 & 8746; 
Killingworth, Middlesex Co., Aug. 19, 1915, C. H. Bissell. New 
York: River Head, Wading R.., Long Island, May 25, 1878, E. S. 
Miller; Long Island, May 1890, nf a Tillinghast; Valley Stream, 
Queens Co. Roepe Island, July 1886, J. A. Bisky; Arden, Staten 
Island, Oct. 1886, A. Ho lick: Hockland L., July 17, 1872, 

orong. Nex JERSEY: Molly Wheaton Run, e. of Greenwich, 
Cumberland Co., Fogg 2077; Elmer, Salem Co., Redfield 7996; 
Cape May Co urt House, Killip 30845; Atlantic City, oa, © 
1868, C. F. Parker. Stage Ley ston Tullytown, Bucks , Ma ay 
24, 1930, W. M. Ben DetawarkE: Record’s P., Lau ang Sussex 
Co. , Fogg 1840; Glendaniel (Hudson) P., 2 mi. s. of Lincoln, aay 
sex Co. , Fogg 4504; Indian R.., Millsboro, Sussex Co. 23, 
1876 6, A. Commons; cedar swamp, New Castle Co. , Sept. 20, “1867, 
Commons; Cherry Island Marsh, below Edgmoor, Wilmington, 
July 27, 1896, Commons; Canterbury, July 1874, Wm. M. Canby. 
MaryLAND: Marshyhope Creek, Federalsburg, Caroline Co., 
Shreve 1622; Blackwater R., Dorchester Co., Shreve 1597; 
Willards, Wicomico Co., Aug. 12, 1910, J. J. Carter. VircINta: 
4 miles n. w. of Waverly, Sussex Co., Fernald & Long 5977; 
brook entering Nowney Creek, Back Bay, Princess Anne Co. 
Fernald, Griscom & Long 4535; near Cornland, Norfolk Co., 
Fernald & Griscom 1296; pond near Luray Caverns, Luray, 


122 Rhodora [APRIL 


June 1, 1909, HZ. B. Bartram; near Elko, Grimes 4196; vicinity 
of Cape Henry, Killip 6896; Great Dismal Swamp, Kearney 1626; 
Washington Canal, Dismal Swamp, Boettcher 9; Dahlia, Greens- 
ville Co., Fernald & Long 8588. NortTu CAROLINA: Cape Fear 
R., Wilmington, herb. Hexamer & Maier 466; Hendersonville, 
Henderson Co., Biltmore Herb., 5980*. Groraia: near Huguenin, 
Sumter Co., Harper 1402; Brier Creek, Screven Co., Harper 
2088. Fuoria: Miccosukee L., Sperry 509. Onto: Baumgart- 
ner’s L., Jackson Twp., Fra nklin Co., Aug. 16, 1929, H. 

Flint. INDIANA: pond about 3 mi. n. w. of Grayville, Sullivan 
Co., Deam 25704; Pine Station, Lake Co., June 1884, E. J. Hill; 
Pine, Lake Co., June 21, 1897, A. M. Chase. KENTUCKY: 
Lexington, 1836, "C. W. Short; s. e. of Mammouth Cave, Edmon- 
son Co., Svenson 156. TENNESSEE: Goose P., Pelham, cma 
Co., Svenson 9108 & 10160. ALABAMA: Montgomery, es 


loosa, Tuscaloosa Co., Svenson 9427. ILLINOIS: Mason "Coe 

Aug. 1860 & Aug. 1861, HE. Hall; Athens, Menard Co., 1861, — 
E. Hall. Minnesota: Colby li; Taylors’ Falls, Metcalf 1291. 
Missourt: St. Louis, 1838, N. Riehl, also Aug. 1847, Geo. Engel- 
mann; Montier, June 8, 1890 0, Bush; north of Flatwoods, Ripley 
Co., Steyermark 14245; Little Black River, Pleasant Grove, 
Ripley oe Mackenzie 359; between Gladden & Timber, Dent 


r & Steyerm ark 41417; Hogan, Iron Co., July 15, — 


Pal 

1898, C; eek Nettleton, Caldwell Co., May 7, 1893, H. Eggert 
(M, mixed with P. amplifolius; F, NY, US, not mixed). ARKAN- 
sas: Nettleton, Craighead Co., May 1893, agent; Greene Co., 
May 7, 1893, "Eggert; Judsonia, June 13, 1877, H. S. Reyne 
Lourstana: Calcasieu R., St: Martinville, Oct. vi 1893, B. 
Langlois (US). Oxuanoma: Page, Leflore Co., Blakley “ee 
also E. J. Palmer 33310. Texas: Lindale, April 23, 1901, 
Reverchon (M). 

Tuckerman’s original description of P. pulcher is brief but 
leaves little doubt as to what plant he referred to this species. 
He was certainly in error however, when he stated that it has 
‘“‘much larger seeds’”’ than P. praelongus. He was quite familiar 
with the P. praelongus of Fresh Pond, Cambridge, where it grew : 
abundantly and was collected there by Boott, Tuckerman, 
Robbins, Morong, and the Faxons. However, all of the Fresh 
Pond material of P. praelongus seen by me lacks mature fruits, 
and it may be that Tuckerman did not at that time realize how 
large the mature fruits of that species really are. His supple- — 
mentary description! is absolutely conclusive as to the plant he . 
was describing. 

1 Edward Tuckerman, Am. Journ. Sci. ser. 2: 6: 224 (1848). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 123 


The forma amphibius and var. ovalifolius which Bennett. re- 
ferred to P. amplifolius may possibly be P. pulcher. They have 
been discussed under P. amplifolius. Hagstrém’s forma am- 
phibius is the terrestrial state so common to P. pulcher. 


6. P. Noposus Poiret 


RuizoMeE white, suffused or spotted with rusty red. Stem 
simple, terete, often pressing very flat, 1-1.5 (-2) mm. in diam- 
eter; stele with the trio-type pattern, ‘with the phloem on the 
inner face of the trio-bundle appearing as one patch; endodermis 
of O-cells (rarely more thickened on the inner face and appear- 
ing as U-cells); Hanah ree! and eH ai org bundles absent; 
pacudodenederiin absen SUBMERSED LEAvEs thin, linear- 
lanceolate to broadly janes lhipadaly ‘9-20 em. tie 1-3.5 cm. 
wide, tapering gradually at base into a petiole 2-13 cm. long, 
tapering gradually to an acutish (but not sharp pointed) apex; 
nerves 7-15; lacunae of 2-5 rows along the midrib; margin of 
young blades with fugacious translucent denticles. FLOATING 
LEAVES coriaceous, with long petioles; blades lenticular to 
elliptical, cuneate or somewhat sounded at base, apex acutish bo 
rounded (sometimes with an obtuse wierd (3) 5-9 (-1l1) ¢ 
long, (1.5-) 2-4 (-4.5) em. wide; n s (9-) 13-21; tania 
rarely present. StTrpuLEs of pabulenid eaves brownish, often 
delicate and decaying early, linear, acute or obtuse, 3— “a ) em. 
long; those of the floating leaves similar but usually broader at 
base and more or less 2-keeled. ine deb ory usually thicker than 
the stem, 1.5-2.3 mm. in diameter, 3-15 cm. long. Young 
SPIKES compact but | becoming eas at anthesis, of 10-15 (—17) 
whorls of hpi at maturity usually not densely fruited, 3-6 
(-7) em. long, 8-1 em. thick. FLOweErs sessile; sepaloid. con- 
nectives greenish or brownish, orbicular or elliptical, (1.4-) 
1.6—-2.2 (to 2.6 on basal flowers) wide; 366 1—-1.4 mm. long. 
Fruits obovate, 3.5-4 (—4.3) mm. long, 2.5-3 mm. wide; keels 
prominent, the dorsal bag develieed, especially upward, the 
laterals often muricate; beak facial, short; exocarp of mature 
fruits brownish or reddish; ‘eedtoaes with keels strongly devel- 
oped, dorsal often .5 mm. wide, the laterals strongly muricate, 
beak linear, erect, up to 1 mm. long, loop solid; apex of seed 
pointing a little above the basal end.—A va ariable species char- 
acterized by floating leaves cuneate at base, narrowly lanceolate 
submersed leaves tapering gradually to each end, and reddish 
fruits with strongly developed, often muricate, eels. 

nodosus Poiret in Lamarck, Ene. Meth. Bot., Suppl. 4: 535 
(1816). Hagstr. Crit. Res. Pot. 183 (1916). P. ey C.& 
S., Linnaea 2: 226 (1827); Benn., Journ. Bot. 31: 297 (1893); 
Taylor, N. Am. Fi, 7: pt. 3:19 (1909). ae s eidoatibie Sieber 


124 Rhodora [APRIL 


ex C. & S., Linnaea 2: 224 (1827); Taylor, N. Am. FI. 17: pt. 1: 
20 (1909). ?P. montanus Presl, Rel. Haenk. 2: 85 (1835). P. 
natans var. fluitans sensu Torr. Fl. N. Y. 2: 244 (1848). P. 
lonchites sensu Tuckerm., Am. Journ. Sci. ser. 2: 7: 350 (1849), 
and subsequent Am. authors, not Tuckerm., ibid. ser. 2: 6: 226 
(1848). P. lucens var. ? fluitans (Roth) Gray, Man. ed. 2: 435 
(1856), in part. ?P. plantagineus var. jamaicensis Grisebach, 
Fl. Brit. W. Ind. 506 (1861). ?P. mexicanus Benn., Journ. : 
25: 289 (1887); ? Morong, Mem. Torr. Club 3: no. 2: 22 (1893); 
? Raunk., Bot. Tidskr. 25: 266 (1903); ? Graebn. in Engler, 
Pflanzenr. 4: fam. 11: 57 (1907); ? Taylor, N. Am. FI. 17: pt. 1: 
18 (1909). P. lonchites var. novaeboracensis Morong, Mem. Torr. 
lub 3: no. 2: 20 (1893). P. americanus var. novaeboracensis 
(Morong) Benn., Journ. Bot. 31: 297 (1893). P. pennsylvanicus — 
var. portoricensis Graebn. in Urban, Symb. Antill. 4: 73 (1903), 
at least in part. P. Nuttallit var. portoricensis Graebn. in Engler, 
Pflanzenr. 4: fam. 11: 56 (1907), at least in part. P. fluitans 
subsp. americanus (C. & S.) Graebn. in Engler, Pflanzenr. 4: 
fam. 11: 60 (1907). P. fluitans subsp. americanus proles novae- 
boracensis (Morong) Graebn. in Engler, Pflanzenr. 4: fam. 1! 
62 (1907). ?P. coloratus var. jamaicensis (Griseb.) Graebn. in 
Engler, Pflanzenr. 4: fam. 11: 69 (1907). ?P. insulanus Hagstr., 
Crit. Res. Pot. 154 (1916). P. rotundatus Hagstr., Crit. Res. 
Pot. 153 (1916). P. fluitans sensu Am. authors, ? not Roth, — 
Tent. Fl. Germ. 1: 72 (1788). Spirillus lonchites (Tuckerm.) 
Nieuwl., Am. Mid. Nat. 3: 16 (1913). i 
A widespread species of ponds and streams, generally in 
flowing water, southern Quebec and New Brunswick to southern 
British Columbia, south to Virginia, Tennessee, Louisiana, and 
California. Map7. Mexico and the West Indies, South America 
(rare), Eurasia, Africa. The following, selected from a large 
series, are representative: QueBEc: St. Lawrence River, 5t- 
Jean-Port-Joli, L’Islet Co., Svenson & Fassett 934; Saint-Lam- 
bert de Lauzon, Levis Co., Victorin, Rolland & Meilleur 43858; — 
Sainte-Rose, Laval Co., Victorin & Rolland 43565 & 49304, also — 
St. Cyr 3030; Longueuil, Chambly Co., Rolland 43359; Angers, — 
Ottawa R., Rolland 19269; Wakefield, John Macoun 62015 & 
62016; Sainte-Sulpice, L’Assomption Co., Ricard & Boivin 342. — 
New Brunswick: St. John R., Lincoln, Sunbury Co., Fassett 
2149. Maine: Houlton, Aroostook Co., Aug. 26, 1897, Fernald; 
Pushaw Stream, Old Town, Penobscot Co., Ogden, Steinmetz & 
Prince 1596, also Steinmetz 326. Androscoggin R., Gilead, OX 
ford Co., Oct. 1, 1897, Furbish; Sydney, Kennebec Co., Fernald — 
& Long 12389; St. George R. near Indian Garden, Warren, Knox — 
o., Aug. 15, 1913, Norton. New Hampsnire: Connecticut — 
R., vicinity of Hanover, Grafton Co., Aug. 17, 1876, H. G. Jesup; 
Cornish, Sept. 2, 1886, F. H. Knowlton. Vermont: Ferrisburg . 


1943] Ogden,—The Broad-leaved Species of Potamogeton 125 


and Vergennes, a collectors; Lake Champlain, Orwell, Addi- 
son Co., Cushman 6007; Hydeville, Rutland Co., July 21, 1892, 
Eggleston; Winooski R. Barngton, Aug. 25 & 27, 1885, Morong. 
MASSACHUSETTS: Concord, Aug. 12, 1887, E. S. Hoar; Mystic P., 
Oct. 1, 1865, herb. Wm. Boott; Winchester, Sept. 1, 1880, Aug. 
8 and ‘Aug. 29, 1881, Morong (the latter mixed with P. ee 
X epihydrus in Gray Herb.); Connecticut R., Springfield, Sept. 
6, 1864, Robbins; Hanon P., Sheffield, Ber kshire Co. , Aug. 12, 
1914, R R. Hoffmann; Pauls Bridge, Neponset KK. Readville, May 
30, 1881, herb. LE. & C. E. Faxon; Egremont, Standley & Killip 
7649. Connecticut: New Hartford, Driggs 40; Twin Lakes Sta- 
tion, Salisbury, Litchfield Co., Aug. '20, 1935, Fernald & Ogden; 
L Whitney, New Haven, Sept. 24, 1886, W. A. Setchell; 
Housatonic R., Stratford, 1845, Ro bbins; Housatonic R., New- 
town, Fairfield Co., A. E. Blewitt 3657. New Yorx: N. Beaver 
Creek, Haynes Hill, W. Fort Ann, Washington Co., Aug. 26, 1914, 
Burnham; Guildenland, Albany Co., House 21774 & 22044; 
Ellisburg, Jefferson Co., ’ House 19852 & 20002; Ithaca, Tompkins 
Co., R. Hitchcock 11167 & 11168; Lake Erie, Buffalo, Aug. 20 
& 25, 1886, Morong; Niagara Rapids, E. Tuckerman (P. lonchites 
of Tuckerman’ s supplementary description, but not of original 
description); pool near White Creek, DeKalb Co., Phelps 1091; 
Grass R.., Canto on, Phelps 1665; Raquette ae above Potsdam, 
St. Lawrence Co., Muenscher & Clausen 3751 & 8752; Float 
Bridge, Rochester, Baxter 5889; Chemung R., Chemung Co., 
Lucy 424 & 10816. New Jersey: Oldmans- Creek, 1.5 mi. 
n. n. e. of Eldridges Hill, Salem Co., Fogg 6794. PENNSYLVANIA: 
Harrisburg, Sept. 1893, John K. ‘Small; Penn’s Creek at the 
“Swinging Bridge,” Selinsgrove, Snyder Co., Moldenke 4208; 
Sellersville, 1868, C. D. Fretz; vicinity of Soe Ferry, York 
Co., Rose & Painter 8205; Chester Co., July 1858-1864, S. P. 
Sharples 303. DELAWARE: Shelpot Creek, Wilmington, June 
= 1879, A. Commons; Brandywine, Wilmingto on, — 17, 1896, 
Commons; White Clay Creek, Stanton, Sept. 4 1896, A, 
Conn mons. MARYLAND: Chicomuxen Creek, Tidestrom 7637; 
Mill Creek, Chesapeake Bay region, Shull 95; Cabin John, 
Montgomery Co., Painter 1189, also Dowell & Painter 5385; 
Chesapeake Canal above Cabin John, near Lock 13, Leonard & 
Killip 603; Spesutie Island, Harford Co., Moldenke 9396; Great 
Falls, House 617. District OF CoLuMBIA: Arlington Junction, 
Sept. 28, 1897, E. S. Steele; C. & O. Canal above Georgetown, 
Aug. 9, 1897, T. H. Kearney; Georgetown, Van Eseltine 
Moseley 202; Fish are Shull 39. West Virarnta: Cacapon 
ardy Co. , Aug. 13, 1930, W. V. U. Bot. Exped.; Tygart R., 
Boas Randolph Co., Millspaugh 457. VIRGINIA: Four-Mile 
Run, Chesapeake Bay region, Shull 474; near Leedstown, Trde- 
strom 7741; Potomac R., The Dyke, Tidestrom 7183; Dyke, 


126 Rhodora [APRIL 


Fairfax Co., Metcalf & Sperry 1622 & 1630 Hunting Creek, 
McAtee 2374. Ontario: Rideau R., at Billings Bridge, Ottawa, 
Malte 11 ie oe m Galetta, Carleton Co., Ogden & 
Bolan 1629 & - Chatham, Cain 935; Nation R. , Casselman, 
Aug. 21, 1884, ste Macoun, also Malte 118275; Nation R., 
Russel, Macoun 221 iss Maitland R., Goderich, Macoun 26839; 
Golden L., Renfrew Co., Macoun 221 7, Dunnville, John Ma- 
coun 26841; near Lake Muskoka, Aug. 29, 1899, D. LeRoy T op- 
ping; L’Original, Rouleau 304. Micuican: Kalamazoo R., Alle- 
gan Co., he 86, 87a, 87b, 87c, 88, 98 & 123; Grand Rapids, 
July 21, 1895, W. E. Mu liken; Vandercooks L., Jackson Co., 
July 21, 1898, ex herb. S. H & D. R. Camp; Pine L., Aug. 20, 
1892, C.F. Wheeler; Gun H. Barry Co., July 1926, Oosting; 
Freemont L., Newaygo Co., July 9, 1926, Oosting; Black L., 
Ottawa Co. , Aug. 27, 1926, Oosting; Huron R., 31% mi. s. e. of 
Ann Arbor, Washtenaw Co., Hermann 9383. Outo: Brady L., 
Portage Co., July 29, 1913, Red Webb; Put-in-Bay, Aug. 1898, 
ow, Pieters; Presque Isle Point, Sandusky Bay, Aug. 19 & 29, 
189 5, E. L. Moseley; Buckeye Creek, Liberty Twp., Jackson Co. 


Pontius & Bartle ey 18. InpIANA: Walnut Creek, 2 mi. n. e. Oba 


Bainbridge, Putnam Co., E. J. Grimes 594; Aberdeen, Ohio Co., 
Deam 56783; Lake Maxinkuckee, Scovell 32; Tippecanoe Ruy 
mi. n. and 1 mi. e. of Winemac, Pulaski Co., Welch 2106; Calu- + 
met R., Clarke, Lansing 1059; n. of Spencer, Owen Co., Deam 
38978; mouth of John’s Creek, Wells Co., July 2, 1905, ’ Deam. 


KENTUCKY: 3 mi. s. of Richmond, Madison Co., Svenson 7216; = 


eer R., June—July, Dr. Short; Ohio R.., Louisville, Sept. 16, 


4, C. Mohr. TENNESSEE: n. fork of Holston R., near Kings- 


port, Hawkins Co., Sharp & Underwood 33521; Reelfoot L., Lake 

o., Demaree 7061 & 7182. ALABAMA: Mobile R. near Piute — 
Island, May 28 & July, 1884, Chas. Mohr; East L., near Birm- — 
ingham, Jefferson Co., Biltmore Herb. 5806. | WISCONSIN: 
between Duck Creek R. and Bars Channel, Green Bay, July 28, 
1891, Schuette; Wisconsin R., near Newport, Delton, Sauk Co., — 
Aug. 26, 1906, A. B. Stout; Pickerel Slough, Prairie du Chien, — 
Crawford Co., Fassett 43650; Lake Mendota, Middleton, Dane — 
Co., Fassett t 3148; near Tomahawk, Cheney 1086. ILLINOIS: — 
Calumet L., Chicago, Chase 1420; Wolf L., Chicago, June 10, — 
1911, Sherff: Du Page R., Naperville, June 22, 1895, L. M. Um — 
bach; Ogden Ditch, Summit, Hill 169, 1909; Lyons Twp. 
Cook Co., Hill 151, 1901; Fox R., Richland Co., Ridgway 3318; 
Lake Lawrence,’ Lawrence, Ridgway & Eaton 34265; Swan L., — 
near Grafton, Calhoun Co., Metcalf 1105; Oquawka, 1879, 
H. N. Patterson. Minnesota: Wabana L., Itasca Co., Metca alf 
1471, also Kubichek 148 & 149; Borden L., Lic tsheons Twp. ., Crow = 
Wing Co., Hotchkiss & Jones 480 & 4109; Ft. Snelling, June — 
1895, E. P. Sheldon, also Mearns 805; Minn. R. bottoms, Dakot? — 


1943] Ogden,—The Broad-leaved Species of Potamogeton 127 


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RANGES OF PoTAMOGETON 


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128 Rhodora [APRIL 


Co., June 1895, EZ. P. Sheldon; Green L., Kandiyohi Co., Met- 
calf 2046; Mazaska L., Rice Co., Keck & Stilwill 401; Courtland, 
Nicollet Co.. July 1892, C. A. Ballard. Iowa: Fayette, July 
1893, B. Fink; Estherville, Aug. 7, 1897, R. I. Cratty; Granite, 
Lyon Co., Aug. 4, 1896, B. Shimek; Des Moines R., July 1881, 
R. I. Cratty. Missouri: along Current R., near Doniphan, Rip- 
ley Co., Steyermark 9233, 14257 & 14259; Gasconade R.., s. e. of 
Hazel Green, Pulaski Co., Steyermark 25102 & 25103; n. w. of 
Waynesville, Pulaski Co., Steyermark 25249; Gasconade R., n. e. 
of Vienna, Maries Co., Steyermark 25603; Buffalo Creek, s. e. of 
Louisiana, Pike Co., Steyermark 25876; Sect. 6, w. of Lynchburg, 
Laclede Co., Steyermark 27148; Osage R., Mary’s Home, Miller 
Co., Steyermark 13083 & 13097; Iron Mountain L., Metcalf 842; 
Killarney L., East Arcadia, Metcalf 845; Ice P., Unionville, 
Metcalf 1071; Duck L., Platte Co., Metcalf 1024; Goose P., 
Springfield, Standley 9780; Gascondy. Emig 224; e. of Ashland, 

oone Co., Drouet 3028; n. w. of Joplin, Jasper Co., Palmer 
21526; Meramec, Sept. 2, 1886, Eggert; Meramec R., St. Louis 
Co., Sept. 12, 1886, Eggert; Atherton, Jackson Co., Bush 630; 
Sheffield, Jackson Co., Aug. 4, 1896, Mackenzie; White R., 
Forsythe, Taney Co., Trelease 817. ArxKansas: Big L., Horners- 
ville, Metcalf 636; Saline R., Ozment Bluff, Drew Co., Demaree 
17893. Soutu Daxora: Sioux R., near Brookings, Sept. 1, 
1893, T. A. Williams; Medicine Creek, near Canning, Aug. 16, 
1892, T. A. Williams 1; s. of St. Pierre, Stanley Co., Over 17432. 
Nesraska: Niobrara R., southwest of Valentine, Tolstead 637; 
Middle Loup R., near Norway, Thomas Co., Rydberg 1421; 
Anselmo, July 6, 1889, Weber 6; Lake Manawa, near Omaha, 
Lawton 50. Kansas: Topeka, Aug. 1870, E. Hall. OKLAHOMA: 
near Cache, Comanche Co., Stevens 1864 (G, US, mixed with P. 
amplifolius, NY, not mixed); Sapulpa, Bush 1207. TExas: 
Comanche Spring, New Braunfels, Lindheimer 1234; Little 
Aguja Canyon, Jeff Davis Co., Moore & Steyermark 3077; Haley 
Ranch, Brewster, Cory 9198; Victoria, Lindheimer 393; Lake Polk, 
near Temple, Bell Co., Wolff 3290; Dallas, June 25, 1929, Mary 
R. Stephenson; Lubbock, Reed 3168; Nueces R., Uvalde, Uvalde 
Co., Palmer 33707; Houston, Harris Co., Palmer 11952; Syca 
more Creek, Fort Worth, Ruth 141. Montana: Great Falls, 2 
S. Williams 285. Ipano: Payette, Henderson 4882. WyYoMING: 
Ft. Steele, Carbon Co., Goodding 539. Cotorapo: Lee’s L- 
Crandall 2530; Alamosa, Clements 305; Rio Grande, Alamosa 
Shear 3745; Owen’s L., Boulder, Daniels 683; Gunnison R» — 
Grand Junction, Mesa Co., Biltmore Herb. 5806%. Utau: Hills — 
Park, Salt Lake City, June 29, 1908, Mrs. J. Clemens; Hill — 
Creek, Uinta Basin, Uinta Co., Graham 9821; near Goshen’s 
fixed sand-dunes, Utah Co., Garrett 3958; Corinne, Wetmore 395. — 
Nevapa: Hot Creek, near Gold Creek, Elko Co., P. B. Kennedy — 


1943] Ogden,—The Broad-leaved Species of Potamogeton 129 


4476; Glendale, Truckee valley, Washoe Co., P. B. Kennedy 
8041; Sparks, near Reno, A. E. Hitchcock 444; North Fork, A. 
E. Hitchcock 1034; Wadsworth, Tidestrom 10655. New Mexico: 
Santa Fe, Fendler 837; San Jose, near Santa Fe, Arséne & Bene- 
dict 16636; Albuquerque, Oct. 13, 1894, C. L. Herrick. Arizona: 
Camp Verde, W. W. Jones 432; Lower Oak Creek, Fulton 9703; 
Granite Reef Dam, Maricopa Co., Peebles 14190; Pinal Creek, 
Toumey 496; Beaver Creek, M acDougal 548. CALIFORNIA: 
Pit R., at Lookout, Modoc Co., Aug. 24, 1899, M. S. Baker; 
Big R.., Mendocino Co. Hy McMurphy id: Russian th Oe 
Cloverdule! Mendocino Co., Heller 5824; Mormon Creek, Tuo- 
lumne Co., Williamson 309; Santa Cruz, M. E. Jones 2310; 
Visalia, Tulare Co. v5 Coville & Funston 1278; Bakersfield, 
Kern Co., Coville & Funston 1244; Laguna Lakes, ee Co., 
Street & Williams, 2689; Colton, San Bernardino Co., Parish 
2106 & 2128; Deep Creek, San Bernardino Mts., jie Bernardino 
CO: d. C. Wheeler 1974. WASHINGTON: Okanog n R., Sereno 
Watson 898. British CotumsBia: Kamloops, ese 26, 1889, 
John Macoun 2970. 

P. fluitans of European authors consists of two quite inde- 
pendent plants: one a fruiting plant with an endodermis of 
O-cells and no bundles in the cortex; this occurs in North Ameri- 
ca. The other is a sterile plant with an endodermis of U-cells 
and with numerous bundles in the cortex; this plant is thought to 
be a hybrid, P. lucens X P. natans. Roth’s original description, 

P. foliis inferioribus longissimis, lanceolatis, acuminatis, membrana- 
ceis; superioribus ouali-lanceolatis, coriaceis: omnibus petiolatis. 
Habitat in fossis oe fundis lente fluentibus et in Hunte fluuio Du- 
catus Oldenburgensis 
would include both odes The fact that no fruits are described 
does not at all mean that Roth’s plant lacked them, for at that 
time vegetative characters were given prominence and the 
fruits often ignored. Of the thirteen species of Potamogeton 
described in Roth’s flora where fluitans is proposed as new, not 
one of them has any mention of fruit. Even if Roth’s specimens 
lacked fruit, it does not follow that they were the non-fruiting 
entity with bundles in the cortex. Bennett thought the name 
fluitans ought to be kept for the hybrid, saying, ‘‘We have no 
certain knowledge of any specimen of Roth’s species being 
preserved in any herbarium; but there are at Munich specimens 
in Schreber’s herbarium, named as such and gathered ‘In Seebach, 
1775,’ and others, ‘In Seebach, 1782.’ It seems to me a reason- 


130 Rhodora [APRIL 


able inference that these specimens are from (or seen by) Roth; 

the more so because there are other species in the same collection — 
actually received from Roth, and signed by him. They are the 

plant we call fluitans in England (hybrid?)”!. Later he adds, 
“The following extract from Roth’s Catalecta Botanica (fase. 1, 
p. 31, 1797) will show that Schreber’s specimens in the Munich 
Herbarium are, as I supposed, the plant of Roth: Ai 
‘Prope Erlangam etiam sued te Ill. Praes. de Schreber’ 2, To 
this Raunkiaer answers, ‘‘that because a herbarium contains 
plants actually from Roth it can not be necessary that other 
plants in the same herbarium should be from him. The speci- 
mens from the Munich herbarium mentioned have been exam- 
ined by me and that they belong to the barren form of P. fluitans 
is true enough but they can not in the least be considered original 
specimens.’? He then attempts to show that in the Bremen 
herbarium there is a specimen which has a good chance of being 
the original; and it is the plant with O-endodermis and lacking 
bundles in the cortex. He also states that in the “‘ Petersburg 
herbarium and examined by me . . . three specimens += 
may well be original specimens.”4 As these specimens also — 
proved to be the fertile plant he concludes that we should retain = 
the name P. fluitans for the fertile species which lacks strengthen- _ 
ing tissue in the cortex. Hagstrom agrees that “This proof 

would be very strong, if those specimens examined also really — 
corresponded with the original description by Roth ‘foliis 
inferioribus longissimis’ . . . which they can scarcely be — 
said to do.”®> From Roth’s later and more detailed description’ — 
it does appear that he then was at least including the barren form _ 
with his P. fluitans. Also, it would seem,if Roth had in mind the — 
specimens in the Bremen and Petersburg herbaria when drawing — 
up his original description, that he would have mentioned them 
in his Catalecta Botanica, as he does mention the Schrebet q 
specimens. It thus appears that the evidence that P. fluitans — 
should be retained for the fruiting plant is not strong, and I am 


Pel aged Bennett, Journ. Bot. 31: 296 Shes 
——__—___—, Journ. Bot. 39: 198 (190 

‘e. Raunkiaer, Bot. Tidskr. 25: 278 ian 
——_—_—_——, Bot. Tidskr. 25: 278 (1$03). 
an ag Hagntrtien, Crit. Res. Pot. 184 (1916). 

6 A. G. Roth, Tent. Fl. Germ. 2: 202 (1789). 


1943) Ogden,—The Broad-leaved Species of Potamogeton 131 


inclined to agree with Hagstrém in treating it as a nomen con- 
fusum. Hagstrém takes up for this plant P. nodosus Poiret. 

P. nodosus Poiret is based on a plant from the Canary Islands, 
collected by Broussonet. This specimen should be in the Paris 
Museum. A photograph of it was, some years ago, requested by 
the Gray Herbarium but not received. The original description 
agrees with the specimens here placed under that name, but is 
not conclusive. Until it can be shown that the Canary Island 
plant is not the wide-ranging species correlated with it, it is best 
to retain Poiret’s name, as taken up by Hagstrém, for the 
American plant. 

The species in North America is a rather variable one. The 
floating leaves especially vary greatly in size. Morong’s var. 
novaeboracensis (of P. lonchites) covers the large-leaved form, 
which in America is less frequent than the narrow-leaved form. 
In Europe the broad-leaved plants are the more common. Which 
form is represented by the Broussonet specimen I am at present 
unable to say, but at any rate this size-variation appears not to 
be worth nomenclatorial distinction. The robustness of the 
plant does not correlate with other variations and the inter- 
mediate forms are the most common. Even the fruits of this 
species show a diversity in the prominence of the keels, but this, 
too, does not correlate with other differences, and sometimes a 
marked variation is found on an individual plant. When typi- 
cally developed, the keels are strongly prominent; their lack of 
development is probably mostly due to a rapid maturation—a 
ripening before the endocarp is fully formed. Unkeeled fruits 
invariably have aborted embryos. 

From Missouri westward, this species tends to have smaller 
floating leaves of a more yellowish green than those typical of 
the east. Correlated with this is a smaller fruit with less strongly 
developed keels (P. rotundatus Hagstr.). However, the typical 
large green leaf is also common in the west, as well as all degrees 
of intermediate forms. Also, some specimens with small yellow- 
ish leaves may exhibit fruits as strongly keeled as those of the 
typical eastern plants. 

In 1848, Tuckerman described as P. lonchites a plant with 
“stem . . much branched, . . . Submersed leaves 

with 6-8 prominent nerves,” (making no mention of a 


132 Rhodora [APRIL 


petiole) and ‘‘floating leaves delicate, . . . always more or 
less tapering above and waved above, . . ._ stipules 
shortish . . .. Nutlets small ... . obscurely tricasii 


ate.”’ He stated that it was “near to P. heterophyllus of authors, 
(P. gramineus, Fr., Koch.).’! The following year he remarked 
further on his P. lonchites and spoke of ‘‘a remarkable state of 


this species . . . In this the stem is simple . . . an 
the . . leaves are either all coriaceous and floating, or only 
the haere submembranaceous, . . . the whole habit of 


which accords, often strikingly with that of P. fluitans; but its 
strongly marked fruit at once refers it to the present species. The 
published description of the fruit of this species was from imma- 
ture nutlets. The following is taken from perfectly ripe ones 
The lateral keels are conspicuous when dry 
The exocarp being removed, the back appears acutely carina 
and a little alate, especially above.’? It is quite evident from 4 
comparison of the two descriptions and an examination of speci- 
mens labeled ‘ P. lonchites’”” by Tuckerman that he was dealing 
with two separate and distinct species. Plants in the Gray 
Herbarium which, though not fruiting, otherwise fit his original 
description perfectly and are labelled ‘‘ Potamog. lonchites”’ in 
Tuckerman’s hand, have no close relationship to the plants 
associated with that name by Robbins, Morong and, following 
them, some other American authors, but are flowing-water 
forms of P. gramineus var. maximus. Tuckerman’s supplement 
ary description was based on P. nodosus, which he mistook to be 
a state of his P. lonchites, as a specimen in the Gray Herbarium 
clearly shows. 

P. rotundatus was based by Hagstrém on four specimens: one 
each from Nebraska, California, New Mexico and Mexico. 
Although referring it to the subsection Amplifolii, he states: 
“I have scarcely met with a species corresponding so nearly t0 
P. nodosus as regards the stem-anatomy as this. It differs by 
the smooth leaf margin, the long lower petioles, the prasinous 
leaf-colour, the ligules, the styles, and chiefly by the character- 
istic fruits.’’? In the Gray Herbarium are specimens from three — 
of the collections cited. All lack true submersed leaves. The — 


' Edward Tuckerman, Am. Journ. Sci. ser. 2: 6: 226 (1848). 
* Edward Tuckerman, Am. Journ. Sci. ser. 2: 7: 351 (1849). 
* J. O. Hagstrém, Crit. Res. Pot. 154 (1916). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 133 


Nebraska plant (Rydberg, no. 1421) has no fruit; the specimen 
from New Mexico (Fendler, year 1847) has fruit which is definite- 
ly keeled. The Mexican collection (Pringle, no. 1390) alone has 
the fruits as described by Hagstrém. That they lack keels is 
true enough, but that they are quite immature is also evident. 
Numerous specimens of P. nodosus show immature spikes the 
fruits of which vary greatly in the development of keels. As to 
the leaf-margin, the denticles on the submersed leaves of P. 
nodosus are so extremely fugacious that they are seldom found 
in any but the youngest leaves. Young leaves of plants approach- 
ing the appearance of P. rotundatus have denticles as freely as 
those of typical P. nodosus. The stipules (“‘ligules’’) of typical 
P. nodosus are often scarcely if at all keeled, so that the lack of 
keels on those of P. rotundatus is not sufficient for its separation. 
The species proposed by Hagstrém as P. insulanus is based on 
a single specimen from Puerto Rico: Sintensis, no. 2537; in the 
herbarium at Stockholm. Originally identified as P. pensyl- 
vanicus Willd. (P. epihydrus Raf.) by Bennett, this collection 
(at least the specimen in the Berlin herbarium) became, with 
Sintensis, no. 1025, the type material upon which P. Nuttalliz 
var. portoricensis Graebner was based. Hagstrém refers Sinten- 
sis, no. 1025 to P. nodosus, though saying, ‘dubious 
probably . . . whatI call P. insulanus.” The two sanabenis 
before me show clearly that neither has any affinity to P. epihy- 
drus (P. pensylvanicus or P. Nuttallii), as has already been point- 
ed out by Fernald.!. The number 1025 is rather definitely P. 
nodosus. The number 2537 in the Gray Herbarium is sterile and 
with small abnormally developed submersed leaves. It appears 
to be either an ecological form of P. nodosus or possibly a hybrid 
between P. nodosus and some member of the subsection Lucentes. 
Just what P. mexicanus Ar. Benn. is cannot be definitely 
determined. Bennett’s description agrees rather well with P.. 
nodosus, except: “Fruit 3/16 in. long by 1/4 in. broad.” How- 
ever, fruits seen by Morong were described by him as being “2 
lines long, 114 lines wide,’? less than half as wide, so it can be 
concluded that Bennett’s ‘(1/4 in.” is in error. Graebner, who 
evidently found it convenient to compute his measurements from 


1M. L. Fernald, Mem, Am. Acad. Arts & Sci. 17: pt. 1: 119 (1932). 
? Thomas Morong, Mem. Torr, Club 3: no. 2: 23 (1893). 


134 Rhodora [APRIL 


Bennett’s description, gives “‘5 mm longi et 6 mm lati.’’! Speci- 
mens in the Gray Herbarium and cited by Graebner are sterile, 
but referable to P. nodosus. : 

P. occidentalis, described by Chamisso and Schlechtendal and 


credited to Sieber, appears, from the description and excellent 


illustration of the fruit, to be typical P. nodosus. 

Just what P. plantagineus var. jamaicensis Grisebach repre- 
sents is not determined. From the description and judging from 
the plants found in Jamaica, it appears to be P. nodosus. Further 
study is needed on the subsection Nodosi of Central America and 
the West Indies. 

7. P. NaTANs Linnaeus 

Ruizomes white with reddish spots when fresh, buff with 
dark red spots when dry (these spots often with lighter centers). 
SreM simple or rarely branched, terete, .8-2 mm. in diameter, 
with transverse ridges (these also on the rhizome and submersed 


leaves); stele with the trio-type pattern, the phloem on the inner — 


face of the trio bundle appearing as 2 distinct patches; endoder- 
mis well developed, of U-cells; interlacunar and subepidermal — 


bundles present; pseudo-hypodermis 1 cell thick. SuBMERSED 


LEAVES coriaceous, semi-terete, narrowly linear (excluding a 
transition leaves), no differentiation between blade and petiole, 


tapering at the apex to an obtuse tip, 10-20 cm. long, .8-2 mm. 


with about one-third of them prominent; lacunae none or ob- 


? 


scure. STiPULES of submersed leaves clasping the stem, whitish, — 
fibrous, persistent, linear to lanceolate, cucullate at apex in the 


cia a aie 


bud, splitting on maturity and becoming raggedly obtuse, OF 


twisting and becoming acutish, 4.5-9 (-11) em. long, about 


mm. wide at base, with 2 well-developed keels and many fine — 


nerves; those of the floating leaves similar but usually broader 
(up to 12 mm. wide at base). Prpuncuxs as thick as or slightly 
thicker than the stem, 3-8 cm. long. Sprxxs in anthesis com- 
pact, with 8-14 whorls; in fruit 3-5 em. long, .9—1.2 em. thick. 


FLOWERS sessile or nearly so; sepaloid connectives greenish, — 


! P. Graebner, in Engler, Pflanzenr. 4: fam. 11: 57 (1907). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 135 


what prominent if dried before fully mature; beak short and 
broad; exocarp sack-like, wrinkled, bright orange drying to buff 
(greenish when immature) ; endocarp more or less pitted on each 
side, and with 2 longitudinal sinuses on the back forming 3 
rounded keels, beak linear, about 1 mm. long, loop solid; apex of 
seed pointing toward the basal end. Plants mostly with strongly 
developed cordate floating cathe and with the narrowly linear 
submersed leaves i 

P. natans L., Sp. Pl. 1 126 (i759); Saree eae Torr. Club, 
13: 145 (1886), Mem. Torr. Club 3 13 (1893); Graebn. in 
En gle ee Space 4: fam. 11: 42 (1907): reste N. Am. Fi. 17: 
pt. 1:16 (1909); Hagstr., Crit. Res. Pot. 191 (1916). P. natans 
var. Pat Hi sensu Am. authors; an Koch? 

A common species of lakes and. streams, Newfoundland, south 
to Pennsylvania, west to California, and north to southern 
Alaska. Map 8. Eurasia. The following, selected from a large 
series of specimens are representative: NEWFOUNDLAND: Bishop 
Falls, Valley of Exploits R., Fernald, Wiegand & Darlington 4461; 
Highlands P., Crabbes, Kennedy 80; 4 miles northeast of Port a 
Port, Mackenzie & Griscom 10043. QuesBec: Mingan Islands, 
Saguenay Co., St. John 90081; Maria, Bonaventure Co., Victorin, 
Rolland & Jacques 383316; marly pond, Grand R.., Gaspé Co., 
Collins, Fernald & Pease 52965; ed Pore-Epic, Saint-Fabien, 
Rousseau 30003; Black L., Megan e Co., Fernald & Jackson 
11986; LacTremblant, Labelle Co., "Viclorin & sep ete 44070; 
Lake Memphremagog, Sargent’s Bay, Aug. R. 
Churchill; Ile Verte, Longueuil, Chambly om ‘Ralland 43363; 
McGregor L., John Macoun 855380; North Wakefield, J. M. 
Macoun 4358. MaAGDALEN ISLANDS: between E. Cape & E. 
Point, Coffin Island, Fernald, Long, & St. John 6763. Dance 
EDWARD IsLAND: east of Britain P., Kings Co., Fernald & St. 
John 10894. Nova Scorta: Pottle’s L., North Sydney, Cape 
Breton Co., Bissell & Linder, cit west. of Ingonish, Cape 
Breton Island, Nichols 749; Salm n R., Truro, Colchester Co., 
Bean & White 19675; Middleton, rppmoer tos Co., Fernald & Pease 
19676; Wentworth i; Digby Co., Fernald & ’ Long 23130; St. 
John (Wilso on’s) L., Yarmouth Co., Fernald, Bartram & Long 
23129; Charcoal, valley of the East R., St. John 1372. MAINE: 
Portage Li; Aroostook Co., 1881, Kate Furbish; Great Works 
Stream, Clifton, Penobscot ve Fernald 2756; Foxcroft, Pisca- 
taquis Co., Fernald 475; Bak “a T 7 R 17, Somerset Co. GSE 
John & Nichols 2106; Swan "P., Oxford Co., July: 1892, J. C. 
Parlin; Torrey P., — r Isle, Hanco k Co., "A. F. Hill '2560a; 
Stevens P., Liberty, Waldo Co., Seanibiieh 60; Black Duck P., 
Matinicus, Knox Co, July 13, 1919, C. A. B. Long; Sydney, 
Kennebec Co., Fer nald & Long 12881; n. of Perley P., Sebago, 
Cumberland Co., Fernald, Long & N orton 1 2382; Wells, York Co. 


136 Rhodora [APRIL 


July 1881, J. Blake. New Hampsuire: Cherry P., Jefferson, 
Coés Co., Pease 20073; Long (Stacy) P., Washington, Sullivan 
Co., Fernald & Svenson 745; Frost P., Jaffrey, Cheshire Co., B. 
L. Robinson 494; Derry, Rockingham Co., Aug. 3, 1926, C. F. 
Batchelder; West Lebanon, Sept. 7, 1891, G. G. Kennedy; 
Bellamy R., Madbury, Strafford Co., Hodgdon 2640. VERMONT: 
Pelot’s Bay, Lake Champlain, North Hero, Grand Isle Co., 
Aug. 2, 1899, Nellie Flynn; West Barnet, Caledonia Co., Aug. 
20, 1884, F. Blanchard; Lowell L., Londonderry, Windham Co., 
L. A. Wheeler; Dead Creek, Ferrisburg, Aug. 15, 1881, EZ. Faxon. 
MassacuusetTts: Long P., Tewksbury, Middlesex Co., Aug. 24, 
1865, herb. Boott, also L. B. Smith 632; Lower P., Wakefield, 
Middlesex Co., Collins 937; Plymouth, Plymouth Co., Aug. 26, 
1913, S. N. F. Sanford; Eastham, Barnstable Co., Collins 3171; 
Sutton, Worcester Co., Anderson, Smith & Weatherby 1166; 
Robinson Creek, Pembroke, Fernald & Svenson, Gray Exsic. 409} 
Lake Buel, New Marlboro, Berkshire Co., July 20, 1920, J. RB. 
Churchill. Ruopr Isuanp: Providence, July 1866, G. Thurber. 
Connecticut: Twin Lakes, Salisbury, Litchfield Co., Hames & 
Godfrey 8679; Dog P., Goshen, Litchfield, Aug. 24, 1913, Bissell 
& Weatherby; Farmington R., Hartland, Hartford Co., Ogden & 
Bolan 1565; Long P., Thompson, Windham Co., Weatherby 4364; 
Mahoney Meadow, Franklin, New London Co., July 27, 1908, 
Rk. W. Woodward. New York: Pierrepont P., Woodville, — 
Jefferson Co., House 16979; Lake Canandaigua, Woodville, Aug. — 
19, 1884, Morong; Spencer L., Spencer, Tioga Co., FE. Moore — 
1488; Tioughneoga R., Riverside Park, Cortland Co., EF. L. — 
Palmer 37; Carpenters P., Fabius, Onondaga Co., House 1338; — 
Sodus Bay, Wayne Co., Killip 6204 & 12258. New JERSEY: — 
Black R., Chester, Morris Co., Mackenzie 4377; Swartswood L., — 
Sussex Co., Griscom & Mackenzie 10685. ONTARIO: Ko-Ko-Ko _ 
Heh L. Timagami, Cain 1045; Franks Bay, Lake Nipissing, — 


1926, Oosting. Onto: Buckeye L., e. of Columbus, Morris A4li — 
Put-in-Bay, Aug. 1898, A. J. Pieters. InpIANA: Wolf L., Agnes 


1943] Ogden,—The Broad-leaved Species of Potamogeton 137 


Chase 1459; Wolf L., Lake Co., Lansing 4274; Bear L., Noble Co., 
Deam 493891; Cheeseborough ie Flint, Steuben Co., Deam 49360: 
Lake Maxinkuckee, Evermann. 1032 (US ), also Scovell & Clark 
1032 (271), under direction of Evermann (F), also Scovell 26 (US, 
mixed with P. amplifolius). Wisconsin: Elkhart L., Aug. oa 
1887 and Aug. 4, 1892, F. H. Schuette; Pell L., Bloomfield Tw 
Walworth Co., H otchkiss & Koehler 4193; Lauderdale, Bebb 996 
& 1008; Valley of the Wisconsin R., near Rainbow Rapids, 
Cheney 1420. Inurnots: Lake Villa, Lake Co., Gleason & Shobe 
179; Grass L., Lake Co., Gates 1752.2; Cedar L., 50 mi. n. of 
Chicago, Roush 812; Ringwood, Geo. Vas asey. Minnesota: Lake 
Itasca, Clearwater Co., Grant & Oosting 3203; Cass L., Pammel 
100; Minnesota R., Dakota Co., June 1895, E. P. Sheldon; doses 
Crab L., St. Louis Co., Sept. 3, 1919, Butters; Green L., 
City, Metcalf 12965; Swan L., Nicollet Co., "Metcalf 50; Schultz 
, Kandiyohi Co. , Metcalf 2113; Koronis L., Stearns Co., Metcalf 
1388, also Kubichek 115b (US, mixed with P. amplifolius) : Little 
Pine L. , Aitkin Co., Over 17139; Lizzie L., Ottertail Co., Kubichek 
190; Bear dss Freeborn Co., Shunk & Manni ing 83; German dis, 
Le Sueur Co., Shunk & Man ning 225; Lake Charlotte, Wright 
Co., Linsdale & Keck 127; Silver L., Mille Lacs Co., Aug. 1892, 
E. P. Sheldon. Iowa: Spirit is, Dickinson Co., July 31, 1896, 
B. Shimek, also July 29, 1897, R. T. Cratty; Round je Lake Tw wp., 
Clay Co., Hayden 823. Nort Daxora: Upsilon L., Turtle Mts., 
St. John, Rolette Co., Mabbott 459; Metigoshe L., Turtle Mts., 
Bottineau Co., Mete tcalf 544. NesBrRASKA: Hann ah’s L., Cherry 
Co., Smith & Pound 228; Hackberry L., Cherry Co., July 0, 
1912, Pool & Folsom, also Tolstead 638; Niobrara Game Reserve, 
near Valentine, Tolstead 428; Swan ae Grant Co., Rydberg 1 652: 
Shafer L., Garden Co., Uhler & Martin 1660; South Cody L., 
Ray Thomson Bee. ALBERTA: n. of Lake Louise, Rocky Mts., 
Macoun 68425 (C). Montana: Lake McDowell, Glacier Nat'l 
Park, Maguire & Piranian 5439; Avalanche L., Glacier Nat'l 
Park, Standley 18500; Rost L., Big Fork, Whitfor d 254, also 
M acDougal 676; Whitefish L., ‘Aug. 24, 1892, R. S. Williams. 
Ipano: Lake Pend Oreille, near Hope, Sandberg, MacDougal & 
Heller 939; Lake Pend Oreille, Aug. 1891, J. B. Leiberg; valley of 
Lake Tesemini, Kootenai Co., Sandberg, MacDougal & Heller 
697; Paradise Creek, Moscow, Henderson 271 7; Priest L., Piper 
3765, also MacDougal 240; Potlatch R., Nez Perce Co., St. John 
et al. 9740; Warm L., 25 mi. n. e. of Cascade, Valley Co., Rollins 
& Chambers 2590; Fernan L., Coeur d’Alene, Rust $85. Wro- 
MING: Jackson’s Hole, Lincoln. Co., E. B. & Lois B. Payson 2251; 
Grand EDcaePnen ioe Nelson 4145. CoLorapo: Laramie 
r Co., Aug. 4, 1891, C. S. Crandall (NY); ahi 
Butte, lak 1891, Cal. Acad. Sci. Herb. (S). Uran 
Sereno Watson 1131 (G, see also next citation). av att Ruby 


138 Rhodora [APRIL 


L., Sereno Watson 1131 (G, NY, US, see also previous citation). 
New Mexico: Long L., Chusca Mts., San Juan Co., A. Wetmore 
541 (US). Arizona: Marsh L., White Mts., Goldman 2458 (US); 
Walker L., San Francisco Mts., Knowlton 288 (US). CALIFOR- 
ntA: Fletcher Creek at Pease-Place, Devil’s Garden, Modoc Co., 
L. C. Wheeler 3973; near Lassen Buttes, Plumas Co., H. E. 
Brown 644; Upper Mud L., Coal Mine Ridge, San Mateo Co., 
R. S. Ferris 2043; Mather, Tuolumne Co., Keck 1188; Lakeside, 
Eldorado Co., June, July 1912, H. D. Geis; Lily L., near Fallen 
Leaf, Lake Tahoe ‘region, Eldorado Co., Wiggins 6757, 6777 & 


Lamb 1259; Nooksack R., Lummin Indian Reservation, What-_ 
com Co., Muenscher 7643; Seattle, Piper 758. British CoLUM- 
BIA: Revelstoke, John Macoun 3019; Colquitz R., near Victoria, — 
John Macoun 88248; San Juan L., Dist. of Renfrew, Rosendahl — 
790; Chilliwack Valley, J. M. Macoun 26814; Griffin L., Macoun — 
2971 (C, mixed with P. epihydrus v. Nuttallii), also Macown 3020. — 
Auaska: Prince of Wales Island, Klawak L., Mr. & Mrs. E. P. 
Walker 994; Ketchikan, Cowles 1405; Dundas Bay, J. P. Ander- 
son 1344; Sitka, J. P. Anderson 21, also Evans 781. 
P. natans is a familiar species over all the northern half of the — 
United States. Because of its wide range and tendency to fruit — 
freely it is one of the primary foods for wild water-fowl. 
Although the American plant seldom attains the robustness 
so typical of the European plant, there seem to be no characters 
fundamental enough to separate the two, even as varieties: 
The fruit of the American plant has a weaker endocarp-beak — 
than that of the European, a fact noted for the American P- — 
alpinus, but. unlike the case of that plant, the drying of the — 
mesocarp of P. natans does not cause any appreciable difference — 
in the shape of the fruits on the two hemispheres. The endocarP 
loop is invariably solid in the American plant and sometimes — 
shows a cavity in European specimens. : 
P. natans, like the other broad-leaved species, responds : 
markedly to ecological conditions. Many of these forms have — 
been given names. In fact, some of the names on the labels fot _ 


1943] Ogden,—The Broad-leaved Species of Potamogeton 139 


the European plants make habitat-notes quite superfluous. 
When in quiet water, the floating leaf-blades become broad and 
definitely cordate; when in a current, the blades are narrower 
and rounded or cuneate at the base. Flowing water also causes 
an elongation of the internodes and a marked reduction in the 
production of inflorescences. 

An aquarium plant of P. natans, which grew from a seed in my 
laboratory, showed an interesting sequence of development. 
The first shoot produced only the linear submersed leaves; the 
second shoot produced some broad leaves, which were much 
narrower than normal and narrowly cuneate at base; the third 
shoot produced the typical broad floating leaves which were 
cordate at base. Then a number of shoots were sent up at about 
the same time, the floating leaf-blades of which, however, re- 
verted to the narrow type with cuneate bases. Finally, the 
branching rhizome sent up numerous shoots, all of which pro- 
duced submersed leaves only, or a few leaves with slightly dilated 
tips. Thus from one seed were produced forma submersus 
Gliick, var. prolixus Koch, and var. vulgaris Koch & Ziz (var. 
typicus). The aquarium was not so constructed that var. ter- 
restris S. F. Gray might appear. 

An interesting form of P. natans which grew in the tidal water 
of Robinson Creek, Pembroke, Massachusetts, has been observed 
on several occasions by Prof. Fernald, and as it appeared to re- 
main the same, was collected by Fernald and Svenson and dis- 
tributed from the Gray Herbarium. With its reduced floating 
leaf-blades, narrowly cuneate at base, on long petioles, and its 
production of winter buds (collected in October), this is obviously 
an ecological state. Mr. Weatherby kindly drove me to the 
locality, but so many changes, attendant on the building of a 
paved road and a new bridge, have so altered the locality as 
described by Prof. Fernald that no P. natans was found. In 
such a habitat, where the tidal water rises and lowers twice a 
day and perhaps at times becomes slightly brackish, no fresh- 
water species of Potamogeton can be expected to lead a normal 
life. That the floating leaves of this plant were submersed at 
times is evidenced by the non-functional and reduced number of 
stomates. 

Fryer’s remarks on ‘some beautiful seedling forms of P. 


140 Rhodora [APRIL 


natans, with lanceolate, oval, and round floating leaves, sufficient 
to afford examples of several named ‘varieties’, but unfortunately 
all growing on one rootstock in the instance in which the ‘varieties’ 
were most marked!” are of interest here. 

American plants labeled ‘var. prolixus’’ include juvenile and 
elongated forms of P. natans and elongated forms of P. Oakesia- 
nus. They are always sterile. 


8. P. OaxestaNnus Robbins 


Ruizomes whitish with red spots. Srem often branched, 
terete, .5-1 mm. in diameter; stele with the trio-type pattern, 
the phloem on the inner face of the trio-bundle appearing as 2 
patches; endodermis well developed, of U-cells; interlacunar 
and subepidermal bundles present; pseudohypodermis absent or 
1 cell thick. SuBMersepD Leaves delicate, narrowly linear, 
obtuse, 5-16 cm. long, (.25—) .3-1 mm. wide; nerves 3. FLOATING 

EAVES coriaceous, with long petioles .2-1 mm. thick; blades 
ovate-elliptical to oblong-elliptical, rounded or tapering at base, — 
obtuse, (1.5-) 2-4 (-5.5) em. long, 1-2 (-3) cm. wide; nerves — 
(7—) 9-19 (-23), about one-third of them prominent; lacunae 
none or obscure. StrputEes of the submersed leaves clasping 
the stem, whitish, delicately fibrous, persistent but beconsay 4 
shreddy, linear, acutish when dry, about 1-3 cm. long; those of _ 
the floating leaves larger, 2-4 (—5.5) em. long, linear or narrowly 
triangular, strongly fibrous, 2-keeled, at least at base. PEDUN- 


cLEs thicker than the stem, .9-1.6 mm. in diameter, 2.5-6 cm. _ 


long. Sprkes with 3-8 whorls; in fruit 1-3.5 cm. long, .7-.9 cm. 
thick. FLowers sessile or nearly so; sepaloid connectives 1.3- _ 
1.8 (-2.2) mm. wide; anthers about .8 mm. long. Fruits ob0- 
void, 2.5-3.5 (3.7) mm. long, (1.6) 2-2.4 mm. wide; lateral keels _ 
rounded, dorsal keel usually prominent and acutish; beak short — 
and broad; exocarp smooth or nearly so, greenish or rarely buff; 
endocarp with smooth sides, and with 2 rather deep sinuses 0D 
the back forming 3 obtuse keels, beak linear, about .8 mm. long, — 
loop solid; apex of seed pointing a little above the basal end. — 
Plants similar to P. natans but smaller. 


1943] Ogden,—The Broad-leaved Species of Potamogeton 141 


Maine, south to New Jersey, west to central New York, and 
local in Michigan, Wisconsin, and western Ontario. Map 9 
NEWFOUNDLAND: Quirpon Island, Straits of Belle Isle, ee 
Gilbert & Hotchkiss 27339; Grand Falls, Fernald, Wiegand & 
Darlington 4464 & 4465; Blomidon (BlowemedDown’: Mts.. 
Fernald & Wiegand 2436; Lookout Mt., Bonne Bay, Fernald, 
Long & Fogg 1208; McCleman’s P., Crabbes , R. B. Kennedy 543; 
Port aux Basques, Fernald, Long & Dunbar 26217. QUEBEC: 
Les Trois Lacs, Laurentides, Victorin, Rolland “a - — 33639; 
Riviére Noire, Portneuf Co., Rousseau 25814; Matamek R. Dist 
North Shore, Bowman 392. Antrcostt: Ellis Bare “M acoun 9008. 
MaGpAteEN Isuanps: Coffin Island, Fernald, Long St. John 
6764 & 6765; Cap-de-l’Est, Ile de la Grande-Entrée, Victorin & 
Rolland 9922. Nw Brunswick: Lac Fox Creek, Westmorland 
Co., Victorin, Rolland & Jacques 44749; Lily P., Southern Head, 
Grand Manan, Charlotte Co., Knowlton & Weatherby pint 
Nova Scorta: Taylor’s L. Bes ver Pictou Co., St. 
1378; Clyde R.., Shelbourne Co., Prince & Atwood 1318 (s): 
Goose L. , Argyle, Yarmouth Co., Sey & White 19680; Petpe- 
wick, Musquodoboit Harbour, ’ Halifax Co., Rousseau 35293; 
Kiroiaed L., Hants Co., Fernald, Bartram & Long 23131; Lena 
St. Paul Island, Perry & Roscoe 38. Ma se Haley P 
Rangeley, Franklin Co., Sept. 1, 1894, Furbish; Gilead, Oxford 
Co., 1897, Furbish; Jordan gt Hancock Co., Sept. 10, 1898, E. 
Rand; "Hackmatack Swamp, Isle au Haut, Knox Co., A. 
Hill 1222; Southport, Lincoln Co., Fassett 18803; Perley P., 
Sebago, Cumberland Co., Fernald, Long & Norton 12384; Lily 
P., East Limington, Limington, York Co., Fernald, Long 
Norton 12383. New HAMPSHIRE: Connecticut ee Northumber- 
land, Coés Co., Pease 12171; Wheeler P., Shelbourne, Cods Co., 
Aug. 31, 1918 , De eane; Merrimack, Hillsboro Co., June 19, 1918, 
Batchelder; Stonehouse P., Barrington, Straff ord Co., Hodgdon 
599. Vermont: Grout P., Stratton, Windham Co., Eggleston 
2111; also Sept. 1, 1931, ae 2 Eaton. Massacuuserts: So. 


Co., Seymour 1487; Nantucket, Nantucket Go.; 886, L. L. 
Dame; Uxbridge, Worcester Co., Aug. 28, 1851, Beicny in herb. 
. Bot. Gard., cotypes = F, G, NE) and. Aug. 18, 1870, 
Robbins, also June 5, 9, & 2 . 1876, Morong; Lake Chaubuna- 
gungamaug, Webster, Worcester Co. ., Ogden & Bolan 1562; 
Spectacle P., Gheendar! Berkshire Co., raed 29, 1912, R. Hoff- 
mann. Connecticut: Middlebury, New Haven Co., Sept. 14, 
1901, arse: “Stafford, Aug. 1897, herb. E. L. Morris. New 


142 Rhodora [APRIL 


York: Quiver P., Fourth L., Fulton Chain, Adirondack Mts., 
Killip 12574 (US, mixed with P. epihydrus v. Nuttallii, G, not 
mixed); Brandy Brook Flow, Cranberry L., St. Lawrence Co., 
Muenscher & Maguire 1711; Big Moose L., Herkimer Co., 
Muenscher & Maguire 1716; Fall Creek, Tompkins Co., Dudley; 
Deep P., Wading R., Long Island, EZ. S. Miller; Rock P., Adiron- 
dacks, Aug. 5, 1884, Morong; McDonough, July 26, 1886, F. V. 
Coville. New Jersey: Pump Branch of Albertson Brook, 
Ancora, Camden Co., J. W. Adams 511; Magnolia L., Ocean 
View, Cape May Co., Sept. 29, 1921, H. B. Meredith; Estellville, 
Atlantic Co., July 4, 1883, C. A. Gross. Onrarto: Sand Pt., 
Algoma Dist., Lat. 47° 00’ N., Long. 84° 45’ W., Taylor et al. 29 


Micuiaan: bog near Rock R., Alger Co., Fernald & Pease — 


3066; Au Train, Alger Co., Pease & Ogden 25135; Crooked L., 
Clyde Twp., Allegan Co., Aug. 18, 1937, D. L. Allen; Crooked L. 
Marsh, Allegan Co., Aug. 4, 1938, W. G. Erwin; 4% mi. s. w. of 

est L., Portage Twp., Kalamazoo ©o., Hanes 407. WISCONSIN: 
Potter’s Cranberry Farm, Cutler, Juneau Co., Sept. 23, 1932, 
J. H. Steenis (G); Valley of the Wisconsin R., near Grand Rapids, 
Cheney 3610 (NY). 


P. Oakesianus has the general appearance of P. natans except | 


that it is uniformly smaller in all its parts. There are funda- 
mental differences, however, chiefly in regard to the fruit. The 


fruit, besides being smaller, lacks the puckered, buff epicarp of = 


RE eae cars 


P. natans and is, instead, stretched and smooth and usually — 


green. The fruits also differ from those of P. natans by having 3 


prominent keels. The other differences are mainly those corre 


lated with size. While the submersed leaves of P. natans are 


generally borne on the single main stem, those of P. Oakesianus : 


are on branches. 


9. P. GraminEevus Linnaeus (American varieties) 


RuizomeE buff, often suffused or spotted with red, variable ine 
thickness. Stem much branched, terete .5-1 mm. in diametel; — 


stele with the oblong-type pattern with but 1 central bundle — 


(rarely with 2) and usually but 1 lateral bundle on each side; 
endodermis of U-cells strongly thickened on the inner and 


lateral faces; interlacunar bundles strongly developed but only — 


in the outer interlacunar circle; subepidermal bundles present — 
or absent; pseudo-hypodermis absent or 1 cell thick. SUB — 


MERSED Leaves linear to linear-lanceolate or lance-elliptical 
(sometimes oblanceolate), 1-9 (-13) em. long, (.1-) .2-1 (-1.5 
em. wide, tapering gradually to a sessile base; apex acute, usually 
sharp-pointed; nerves 3-9 (—11); lacunae of 1 or 2 rows along 


1943] Ogden,—The Broad-leaved Species of Potamogeton 143 


midrib, mostly obscure; margins with fugacious 1-celled trans- 
lucent denticles. FLOATING LEAVES coriaceous, blades ovate to 
elliptical (rarely subrotund), 1.5—5 (—7) em. long, 1-2 (-3) cm. 
wide; apex obtuse or bluntly mucronate; base cuneate or rounded; 
petioles 2-10 (-15) cm. long, mostly longer than the blades; 
nerves 13-17 (—23); lacunae obscure. STIPULES persistent, ob- 
tuse and slightly cucullate at apex, those of the submersed 
leaves and branches .5-3 cm. long, 1-2 mm. wide at base, faintly 
2-keeled, with 8 to 30 igs nerves, those of the floating Anon 
broader. PrpuNcLEs at base about same thickness as 
sometimes clavate, 2-10 (-30) ecm. long. Sprkes in aitheas 
usually rather compact, of 3-10. whorls of yt in fruit 
canta and crowded, 1—2.5 em. long, .6—.8 cm. 
sessile or on pedicels up to .56 mm. long; peace connectives 
orbicular to oval, blades (.7—) 1.2-1.6 (—2.3) mm. wide, claws 
.2-) .4~.8 (-1) mm. long; anthers oblong .6-1 (-—1.1) mm. long. 
Fruits mostly obovate, 1.7—2.5 (—2.8) mm. long foxaledlne beak), 
(1.4-) 1.6—2 (-2.3) mm. wide, keels usually strongly evident, but 
often obscured by the loose exocar p, beak facial, short and 
curved toward the back; exocarp usually loose, green or rately 
tawny; endocarp with keels low and obtuse, beak linear, be 
mm. long, loop solid; apex of seed pointing 3-7 m 
' above the basal end. A v ariable species characterized by a pedal 
with many lateral pottipoltid branches bearing numerous small 
leaves. Among the many variants of P. gramineus, the following 
seem worthy of recognition 


i Ber rate ae ieornias leaves narrowly elliptic to oblanceolate, 
(1—) 1.5-9 (-13) em. long, .2-1 (-1.5) cm. wide, eg times 
long as broad, or if more than 10 times, =) t less 
than 6 em. long, sides not parallel; actly (3- 
2. Principal submersed leaves (1-) 1.5-4.5 (-6. c. cm. long, 
—-.6 (—.8) em ahecaes WOVEN He Oise aw as 
2: Principal submersed leaves (3-) fe (-13) em. long, 6-1 
(-1.5) em. wide; nerves 7-9 (-11).........-..-. 9b. var. maximus. 
i. Principal submersed leaves eer 1) 1.5- os 5 (-5.5) em. 
long, .1-.25 (—.3) em. wide, 10-20 (-30) times as long as 
broad, sides caaieiaatty set ni most of their eek 
tapering at apex to an acute tip; nerves 3...... 9c. var. myriophyllus. 


te 


. var. typicus. 


9a. P. Gramrnevs L. var. typicus 

P. gramineus L., Sp. Pl. 1: 127 (1753); Graebn. in Engler, 
Pflanzenr. 4: fam. 11: 34 (1907). P. Proteus see ten C. & 
S., cp end 2: 202 (1827). P. gramineus var. graminifolius 
Fries, Novit. Fl. Suecicae 36 (1828), and aethantiuiaind Am. 
authors. P. Piceschetiok sensu Morong, Mem. Torr. Club 3: no. 2: 
23 (1893): Taylor, N. Am. Fl. 17: pt. 1: 19 (1909); not Schreb. 
P. heterophyllus forma gramini ifolius Morong, Mem. Torr. Club 3: 
no. 2:24 (1893). P. asebiben ie forma sdupsaduacthatts Morong, 


144 Rhodora [APRIL 
Mem. Torr. Club 3: no. 2: 24 (1893), at least in part. P. gra- 


mineus var. longipedunculatus Graebn. in Engler, Pflanzenr. 4: 


Fern., Gray’s Man., ed. 7: 74 (1908). Spirillus heterophyllus 
Nieuwl., Am. Mid. Nat. 3: 17 (1913). P. gramineus forma 
longipedunculatus House, Bull. N. Y. State Mus. 254: 53 (1924), 
at least in part. P. gramineus forma terrestris Carpenter, Fl. 

t., 3rd_rev. ed.: 25 (1937). P. gramineus var. lacustris sensu 
Hultén, Fl. Alaska and Yukon, pt. 1: 100 (1940). 

Lakes and streams, southern Greenland to Alaska, south to 
New Jersey, Ohio, Indiana, Illinois, Iowa, Nebraska, New 
Mexico, Arizona, and California. Map 10. Eurasia. Of the 
very numerous collections of this variety the following are the 


most typical in the areas cited: GREENLAND: Igaliko, 1828, J. — 


. 


Vahl, also July 23, 1888, Rosenvinge 2990; Igaliko-Fjord, Qags- 
siarssuk, Aug. 5, 1925, A. E. 


Rolland & Jacques 33714; New Richmond, Bonaventure Co. 
Victorin, Rolland & Jacques 33855; Lake Temiscouata, Victorin 


$26; Farm Point, Gatineau R., J. M. Macoun 80929; Ottawa R. 
at Gatineau Point, Malte 118257; Blue Sea L., Malte 118265; 


Aug. 17, 1917, Victorin 4196. Macpatmn ISLANDS: Amherst 
Island, St. John 1758 (toward var. maximus). Nova Scotia: 


e - New Hampsuire: Dead R Berlin, Coés C 
22754; Lake Winnipisaukee near Melvin Village, Aug. 


M. P. Porsild; Frederiksdal, — 


ie Se Riglaeseos. 


: 
i 
; 

G 


1943] Ogden,—The Broad-leaved Species of Potamogeton 145 


M. A. Day. Vermont: Little Otter Creek, Lake Champlain, 
Ferrisburg, Aug. 7, 12, and 20, 1880, C. EF. Faxon; Queechee 
Gulf, July 29, 1890, G. G. Kennedy; Fairfield P., Fairfield, 
Franklin Co., Blake 3071. Massacuusrrts: Mystic L., July 4, 
1852, Robbins; Fresh P., Cambridge, August 6, 1883, Morong; 
Sandy P., Lincoln, Sept. 15, 1868, herb. Wm. Boott; Natick, Aug. 
14 and 17, 1883, Morong. Connecticut: Selden’s Cove, Lyme, 
Aug. 31, 1900, C. B. Graves; Pistapaug P., Durham, Weatherby 
3389; several collections from Lake Saltonstall, E. Haven are 
typical P. gramineus, others from the same lake approach var. 
myrtophyllus, still others approach var. maximus or are perhaps 
P. gramineus X P. illinoensis. New York: Butterfield L., 
Jefferson Co., Muenscher & Maguire 1690; Osgood P., Franklin 
Co., Muenscher & Maguire 778; Otsego L., Otsego Co., Muenscher 
& Curtis 4880; Bullhead P., Minerva, Essex Co., House 15193; 
Myers Pt., Ludlowville, Tompkins Co., Aug. 13, 1884, W. R. 
Dudley. New Jersey: Morris P., Sept. 13, 1887, N. L. Britton; 


maximus; NY, not mixed); Swartzwood L., Sussex Co., Griscom 


3 


McAtee 3076. Ounto: Sandusky Bay, July 20, 1895 and Sept. 2, 
1898, E. L. Moseley; Put-in-Bay, Aug. 1898, A. J. Pueters. 


ee Oe I 
28, 1897 and June 29, 1898, L. M. Umbach; Lake Maxinkuckee, 
Scovell 44, also Scovell & Clark 1221, also Evermann 1221. Wis- 
CONSIN: Pell L., Bloomfield Twp., Walworth Co., Hotchkiss & 
Koehler 4194; near State House, Trout Lake, Vilas Co., Fassett 
9067 & 9069; Green Bay, Big Suamico, Aug. 28, 1891 and July 
31, 1893, J. H. Schuette. Iniinots: Edgewater, June 7, 1890, 
L. N. Johnson; Rogers Park, June 7, 1890, herb. W. H. Dunham. 
ManiTospa: 4 mi. w. of Hamiota, Macoun & Herriot 76868; 
Killarney, Macoun 16441. Mrnnesota: Green L., Kandiyohi 
Co., Metcalf 2050; Itasca Park, De Soto L., Becker Co., Grant & 
Oosting 3272 & 3276; Snail L., Ramsey Co., Oosting 28166; Horn 
L., Anoka Co., Oosting 291 & 28100; Muskeg Bay, Lake of the 
Woods, Warroad, Roseau Co., Hotchkiss & Jones 417; Long L. 
near Ely, St. Louis Co., Hotchkiss & Jones 4083; Pleasant L., 


146 Rhodora [APRIL 


Stearns Co., Linsdale & Keck 1; Dudley L., Rice Co., Keck & 
Stilwill 373 & 379; Birch L., Sherburne Co., Kubichek 101; Fish 


L., Chisago Co., Kubichek 66; Borden L., Garrison Twp., Crow 
Wing Co., Hotchkiss & Jones 4112. Iowa: Armstrong, Emmet 
Co., Aug. 8, 1891, June 20, 1897, and Aug. 21, 1897, R. I. Cratty; 
Lost Island L., Freeman Twp., Clay Co., Hayden 821. NortT# 
Daxorta: King Slough, s. of Bismark, Metcalf 345; Doctor L., 
Drake, Mabbott 423; Spiritwood, Bergman 443; Leeds, Benson 
Co., Aug. 2, 1899, Aug. 21, 1899, and Aug. 16, 1915, J. Lunell. 
SoutH Dakota: South Bass P., Cottonwood L., Spink Co., 
Over 17138; eastern Day Co., Over 14466. NeprasKa: Pelican 
L., Thomson 153; Red Willow L., Thomson 361 & 365; Enders 
L., Thomson 16; Dewey L., Tolstead 615; Shafer L., Garden Co., 


Uhler & Martin 1656. Mackenzie: Mosquito Creek and Drift- — 


wood R., Great Bear L., 66° 55’ N. 121° 20’ W., July 6-8, 1928, 
A. E. & R. T. Porsild; Edna Travers Bay, Great Bear L., 66 
25’ N. 117° 40’ W., A. E. & R. T. Porsild. SaskaTCHEWAN: 
vicinity of William Pt., Lake Athabasca, 59° 7’ 30’” N. 109° 19 


Oa: 


W., Raup 6849; Little Buffalo L., J. M. Macoun 2975; s. of — 


Battleford, Macoun 2981. ALBERTA: east end of Crow’s Nest 


Pass, Rocky Mts., Macown 23180; Sand Pt., n. shore of Lake — 


Athabasca, Raup & Abbe 4614. Monrana: Mud L., Bigfork, | 


Flathead L., M. E. Jones 9293; Flathead L., Big Fork, Flathead — 

0., G. B. & R. P. Rossbach 17; Whitefish L., Aug. 24, 1892, R. — 
S. Williams; Lower Two Medicine Lakes, Glacier Nat’l Park, — 
Maguire 484; Echo L., MacDougal 639. Ivano: Priest Ly | 
MacDougal 241; Lake Pend Oreille, Sperry & Martin 719, also — 
Henderson 4576; “Lake Pend Oreille, near Hope, Sandberg, — 


MacDougal & Heller 955 & 1026. Wyomina: Shoshone Creek, 


Yellowstone, Aug. 23, 1878, C. Richardson; Yellowstone Ly — 
Yellowstone Park, Tweedy 411; Leighs L., Jacksons Hole, Mer — 


rill & Wilcox 902. Couorapo: Estes Park, Lorimer Co., Beé 


2341; South Park, Wolf 961; near Boulder, Boulder Co., Tweedy — 
4978. Uvan: Grassy L., Goodman Ranch, Bear R. valley, 


Ranch, Lake Co., Coville 602. Wasnineron: Lake Washingtod: — 


Mercer Island, King Co., Thomson 7589; Ozette L., Clallam er Af 


Otis 1584; Blakeley Island, San Juan Islands, S. M. & E 


Zeller 1237; Falcon Valley, Aug. 1, 1885, W. N. Suksdorf (toward 


ae 
4 


1943] Ogden,—The Broad-leaved Species of Potamogeton 147 


Si 6 gt 
{24 og: - 


faa } P. GRAMINEUS 


RANGES OF PoTAMOGETON 


148 Rhodora [APRIL 


var. myriophyllus). British Cotumpia: Kamloops, Macoun 
2974; Wellington, Vancouver Island, John Macoun 88254; 
Sproat L., Albernie, Vancouver Island, Carter 505. ALASKA: 
Yes Bay, Howell 1668; Fairbanks, L. J. Palmer 1866. 


9b. P. GRAMINEUS L. var. MAxIMUS Morong ex Bennett 


Var. maximus Morong ex acim? Journ. Bot. 19: 241 (1881). 
P. lonchites Tuckerm., Am. Journ. Sci. ser. 2: 6: 226 (1848), not 
Tuckerm., ibid ser. 2: 7: 350 shoe and subsequent Am. authors. 
at gramineus var. maximus Morong, Bull. Torr. Club 13: 155 
(1886), without description. P. heterophyllus forma maximus 
Morong, Mem. Torr. Bot. Club 3: no. 2: 25 (1893). P. gramineus 

var. maximus Graebn. in Engler, Pflanzenr. 4: fam. 11: 88 (1907). 
a: heterophyllue in part, Taylor, N. Am. Fl. 17: pt. 1: 19 (1909). 
P. gramineus f. Wolfgangit sensu Hagstr., Crit. Res. Pot. 209 
(1916), as to American citations. P. gramineus f. jemtlandicus 
ee Hagstr., Crit. Res. Pot. 209 (1916), as to American cita- 
io 

Lakes and streams, often in flowing water, with the typical 
variety and having essentially the same range in North America. 
Mar 11. Among many collections, the following are the most 
representative from the areas cited: LaBrapor: 18 mi. .up 

kaupi R., Lake Melville Dist., R. H. Wetmore 103096. 
NEWFOUNDLAND: Rushy P., cpuaiee R., Fernald, Wiegand, 
Bartram & Darlington 4477; Lewisport, Notre Dame Bay, Fer- 
nald, Wiegand & Darlington 4480. Quesec: Lac des Quinze 
(Baie Gilies), Temiscaming-Abitibi, Victorin 8194 & 8196; 
Roberval, July 16, 1892, Geo. G. Kennedy; Deschenes, near Hull, 
Malte 11 8263; Lac Saint-J. ean, Victorin 16064. ANTICOSTI: 
Riviére McKane, Victorin & Rolland 27095; Pointe de 1|’Est, 
Victorin & Rolland 27094. NEw BRUNSWICK: Titusville, Brit- 
tain 2987; near St. John R., Connors, Pease 2589. Nova Scotia: 
Salmon R. , Truro, Colchester Co., Bean & White 22962. MAINE: 
Aroostook Pe Ft. Fairfield, July 18, 1893, eden St. John R., 
Ft. Kent, Mackenzie 3613; Dead R., Somerset Co., Fernald & 
Strong 477: stream below Dwinall P:, Winn, Penobscot Co., 
Steinmetz 365; Stillwater R., Old Town, ’ Penobscot Co., Ogden & 
Steinmetz 1602; Orland R., Orland, Penobscot Co., Ogden & 
Marston 1694. New HAMPsnine: Connecticut R., Walpole, 
Fernald 436; Connecticut R., near Hanover, July 26 and Aug., 
1876, H. G. Jesup (not typical). Vermont: Little Otter Creek, 
Lake Champlain, Aug. 7, 1880, herb. E. & C. E. Faxon (not 
typical). MASSACHUSETTS: Charles R., Dedham, July 14, 1879 
and July 16, 1880, Morong, also Aug. 2, 1880, C. E. Faxon; 
Charles R., S. Natick, July 14, 1879 and Sept. 5, 1882, Morong; 
Charles R., Needham, Tuckerman; Ashland, July 9, 1879, herb. 
Morong. ‘ConNECcTICUT: Quinnipiac R. at Old Turnpike, 


a Ne 


SS 


ee 


1943] Ogden,—The Broad-leaved Species of Potamogeton 149 


Southington, Aug. 17, 1900 and Aug. 17, 1901, C. H. Bissell; 
Housatonic R., near Lake Zo oar, Southbury, E. H. Eames 11 745. 
New York: Saranac pe Adirondacks, July 31, 1884, Morong; 
French Creek, Clayton, Jefferson Co., Muenscher & Maguire 
1693; Hudson R., below Glen Falls, Warren Co., Muenscher & 
Lindsey 2769; Song L., Cortland Co., Muenscher & Curtis 4841; 
Buffalo, Clinton 6. New Jersey: Delaware R., Hunterdon Co., 
Sept. 19, 1885, T. C. Porter; Lake Hopatcong, C. F. Austin 
(C, mixed with P. gramineus var. typicu us). PENNSY Daeg 
Penn’s Creek at “‘Swinging Bridge,” Selinsgrove, Snyder 
Moldenke 4207. Ontario: Dog R., above Michipicoten, pee 
Macoun 97; Ottawa R., Rockliffe, John Macoun 85536; Ottawa 
R., Harrington 99086 & 99102; Templeton, Scott 16444. Mucuti- 
GAN: St. Clair R., near Port Huron, Dodge 155; Sault R., near 
Sault Sainte Marie, Aug. 11, 1910, J. R. Churchill (not typical). 
Onto: Sandusky Bay, Aug. 19 and Aug. 31, 1898, A 
(not typical, perhaps P. gramineus X P. illinoensis). rcs: 
SIN: Wisconsin R., near Lac Vieux Desert, Cheney 683; Green 
Bay near Big Suamico shore, Brown Co., Jul 1 gee ee 
Schuette (not typical, perhaps | gramineus ae ek illinoensis). 
Minnesota: Vermilion ol July 28, 1886, L. H. Bailey, also 
Arthur, Bailey & Holw y B46, B69 & B403; Garden Taina: 
Lake of the Woods, M. acMillan & Sheldon 1332; near mouth of 
Brule R., Cook Co., Rosendahl & Butters 4638. Iowa: Armstrong, 
Emmet Co., July 11 and Aug. 21, 1897, R. I. Cratty. Nort 
Dakota: Leeds, Benson Co., July 2, 1906, J. Lunell. Macken- 
zie: Edna Travers Bay, Great Bear L., Aug. 8, 1928, A. E. & R. 
T. Porsild. SasKaTCHEWAN: along Grand Trunk Pacific R. R., 
Yorkton, Macoun & Herriot 76869. AuBerta: Murdock Creek 
Dist.. Wood Buffalo Park, Raup 1546; L. Mamawi, Wood 
Buffalo Park, Raup 1548. Montana: Midvale, Umbach fas 
Ipano: Moose Creek, near Big Springs, Fremont Co., 
P. Rossbach 20. Wyomina: Obsidian Creek, Yellowstone Neti 
Park, Aven & Elias Nelson 6061. Couorapo: Trout L., near 
Lizard Head Pass, San Miguel Co., Maguire, Piranian & Rich- 
ards 12771. Uran: Salt Lake City, PP E. Jones 1304; Dry L., 
Cache Co., Maguire 138149. New Mexico: Chusca Mts., San 
Juan Co., Wetmore 549. Anizona: Crater L., San Francisco 
Mts., Aug. 1886, Lemmon Herbarium. CALIFORNIA: Lily L., 
near ’ Fallen Leaf, Lake Tahoe region, Eldorado Co., Wiggins 
6775. Wasnincton: Tumwater Canyon, Wenatchee R., Sand- 
berg & Leiberg 524; Camas Land, Wenatchee Mts., Chelan Co., 
Thompson 11768. BritisH ConuMBIA: Shawnigan L., Vancouver 
inne: John Macoun 88258. Awuaska: Sitka, Evans 780; Selawik 
a J. Palmer 638 (US, mixed with P. Richardsonit). 


150 Rhodora [APRIL 


9c. P. GRAMINEUS L. var. MYRIOPHYLLUS Robbins 


Var. myriophyllus Robbins in A. Gray, Man. ed. 5: 487 (1867). 
Fr, ee re forma myriophyllus Morong, Mem. Torr. Club 
: no. 2: 24 (1893). P. heterophyllus forma minimus Morong, 
Mane Torr. Giub 3: no. 2: 25 (1893). P. gramineus var. myrio- 
phyllus Graebn. in Engler, Pflanzenr. 4: fam. 11: 87 (1907); 


Graebn. i in a Pflanzenr. 4: fam. 11: 89 (1907). 'P. hetero- 


53 aged 

Quiet water; local in New Hampshire, Massachusetts, Rhode 
Island, New York, Michigan, Indiana, Wisconsin, Minnesota, 
and Montana. oe 12. Although many collections approach 
this variety, only the following are considered typical: NEW 
Hampsuire: Lake Winnepesaukee, 1876, W. F. Flint. Massa- 
CHUSETTS: Spot P., Stoneham, Aug. 20, 1865, herb. Wm. Boott, 
also July 27, 1876, Morong, also Aug. 13, 1880, herb. E. & C. E. 
Faxon; Spot e Wyoming, Aug. 13, 1880, herb. C. E. Faxon; 
Spot P., Melrose, Aug. 13, 1880, "Edwin Faxon; Winter P., 
Wihehestcr Middlesex Co., Fernald & Svenson Thh Wakefield, 
ee ba ble J. W. Chickering; Lake Pattaquatic, Ware, Aug. 
25,1 E. L. Morris. Ruope Istanp: Apponaug P., Apponaug, 
Oct. vy "1865, Robbins (type in NY), also Aug. 1879, Sept. 29, 
1879 and Aug. 26, 1880, Morong, also Aug. 26, 1880, "E. Faxon, 
also Aug. 26, 1880 herb. E. & C. BE. Faxon (“ Kingston” appears 
on the labels, which is an error, for the collection was made at 
Apponaug in the town of Warwick), also Aug. 24, 1881, E. Faxon, 
also Aug. 25, 1881, Morong; Gorton’s P. (Apponaug P. of early 
botanical collectors), Apponaug, town of Warwick, E. C. & E. 
B. Ogden 1765. New Yorx: Mud P., Pattens Mills, Warren 
Co., Aug. 25, 1918, S. H. Burnham; Friends L., Warren Co., 
M uenscher & Lindsey 2776. MIcHIGAN: Sister Lakes, Van 
Buren Co., De Selm 22. Inpiana: Dune Park, Peattie 2305. 
Wisconsin: Oneida Reservation, Sept. 8, 1881, ‘J. H. Schuette; 
Devils L., Sauk Co., Fass.tt t 14262. Minnesota: Lake Mora, 
Kanabec Co., July 1892, E. P. Sh: ldon; Milaca, Mille — Co., 
July 1892, E. P. She Idon. Montana: Bitterroot Ri; 
Missoula, Missoula Co., Barkley 1996 


The extremely variable P. gramineus i aehick often approaches 
in appearance its near relative, P. illinoensis, is further compli- 
cated by the fact that it hybridizes with most (perhaps all) of the 
other broad-leaved species of the genus and even one (perhaps 
more) linear-leaved species. In North America three varieties 
can be recognized which, though distinct in their extreme de- 


1943] Ogden,—The Broad-leaved Species of Potamogeton 151 


velopments, intergrade freely. The variety that appears to be 
identical with the one upon which the species is based (var. 
typicus) is by far the most common, and is rather widespread in 
both Europe and North America. Its much branched stem has 
usually an abundance of small elliptic-lanceolate submersed 
leaves which distinguish it from all other species of Potamogeton 
as well as from the other varieties of the same species. 

The variety myriophyllus has a stem even more branched and 
the leaves still smaller but with parallel margins. This would 
seem to be but an ecological form were it not found in somewhat 
different habitats, according to data on labels. When I collected 
it in Apponaug Pond in 1938 it was exactly like the plants 
Robbins collected there in 1865. 

The variety maximus is perhaps the most variable one. It 
differs from the other variants of the species in the larger size of 
the submersed leaves, less branching of the stem, and usually 
longer internodes. While for most collections this variety seems 
to be genetic, for many the condition appears to be ecological. 
It is oftentimes very difficult to distinguish it from hybrids 
having P. gramineus for one of the parents. It can usually be 
distinguished from P. gramineus X P. illinoensis, which appears 
‘most like it, by the more flaccid stem and leaves of the variety. 

Plants appearing like typical P. gramineus or var. maximus, 
except that the submersed leaves are oblanceolate or spatulate, 
with the apex rounded, mucronate, or cuspidate (often with 
many variations on the same plant) occur. These have been 
called P. gramineus var. spathulaeformis. The type material 
upon which this name is based is from the Mystic Lakes, Med- 
ford, Mass. and proves to be P. gramineus X P. illinoensis. 
Plants with submersed leaves somewhat similar but usually 
smaller and with floating leaves usually orbicular or very short- 
elliptic occur in Newfoundland, Quebec, New Brunswick, Ver- 
mont, New York, Ontario, Michigan, Minnesota and Iowa. 
While the usual stem-anatomy pattern is that of P. gramineus, 
occasionally O-cells are found in the endodermis, thus pointing 
toward a parent (or grandparent) with an endodermis of O-cells. 
The broad rounded apex of the submersed leaves and the nearly 
orbicular floating leaves point to an ancestor with broad rounded 
leaves. Some of these plants are P. gramineus X P. perfoliatus, 


152 Rhodora [APRIL 


others are P. gramineus X P. Richardsonii, in both cases prob- 
ably backcrossed with P. gramineus. At Moran, Mackinac Co., 
Michigan, in the shallow water at the northwest end of Brevort 
Lake, I found typical P. gramineus (Ogden & Bolan 1680) and 
obvious P. gramineus &X P. Richardsonii (Ogden & Bolan 1681, 
with Richardsonii-like submersed leaves and gramineus-like 
floating leaves) and clearly intermediate between the two a plant 
with oblanceolate submersed leaves (Ogden & Bolan 1681 in 
Gray Herbarium). See discussion of these hybrids on later 
pages. Not all oblanceolate-leaved plants are hybrids, however, 
for leaves of the typical variety or of the variety maximus which 
have been produced at the surface of the water may tend to form 
a transition to the floating type, which may narrow the lower half 
and produce a mucro at the apex; in such cases the lower leaves, 
if submersed, will have the normal shape. Then, too, submersed 
leaves often have a tendency for the edges of the lower part to 
curl under, giving the leaf a false oblanceolate shape which close 
examination will detect. 


10. P. 1LLINOENSIS Morong 


epidermal bundles present or absent; pseudo-hypodermis absent 
or 1 cell thick. Suspmursep Leaves thin, elliptic or oblong- 
elliptic (sometimes ovate-elliptic) to lanceolate or linear (by the 
further reduction of the blade to the midrib), often somewhat 
arcuate; blades 5-20 cm. long, (.2-) 1.5-4 (-4.5) cm. wide, 
sessile or tapering (except when reduced to midrib) into petioles 
up to 4 cm. long; apices acute, usually somewhat mucronate; 
nerves (7—) 9-17 (-19); lacunae of 2-5 rows along midrib and 
larger nerves; margin with fugacious 1-celled translucent 
denticles. FLoatinec Leaves (often absent) more or less coria- 
ceous, transition to submersed leaves usually gradual; blades 
elliptic, ovate-elliptic, or oblong-elliptic, 4-13 (-19) cm. long, 
2-6.5 cm. wide; apices obtuse, ending in a blunt mucro; bases 
cuneate or rounded; petioles 2-9 cm. long, shorter than the blade; 
nerves 13-29; lacunae of 2 or 3 rows of cells along midrib, some- 


1943] Ogden,—The Broad-leaved Species of Potamogeton 153 


IL 


P. ILLINOENSIS 


a 
- 
bY “yy a 
~~ Aa eh 
ee ie ee 
y eye 


aa 
aw 


RANGES OF POTAMOGETON 


154 Rhodora [APRIL 


times obscure. SvrpuuEs persistent, divergent and conspicuous, 
obtuse, those of the submersed leaves (1—) 2.5-8 em. long, (.3- 
.2 cm. wide at base, prominently 2-keeled, with 15-35 finer 
nerves; those of the floating leaves broader. PEDUNCLES as 
thick as or thicker than the stem, 4-15 (-30) cm. long. Spikes in 
anthesis more or less compact, of 8-15 whorls of flowers; in 
fruit cylindric and crowded (2.5—) 3-6 (—7) cm. long, .8-1 cm. 
thic LOWERS sessile or on pedicels up to .5 mm. long; sepaloid 
connectives orbicular to oval (or reniform), blades (1.3—) 1.6-3 
(-3.2) mm. wide, claws 1-1.5 mm. long; anthers oblong, .6-2 
mm. isis Fruits obovate to lan or ovate (2.5—-) 2.7-3.5 
(-3.6) mm. long (excluding beak), (2.1-) 2.2-3 mm. wide; sides 
flat; keels prominent and acute, the dorsal ad developed 
above and below, the laterals less Antari developed but often 
each with a projecting knob at the base; beak facial, short, 
erect or curved toward the back; aarp ‘eray-green to olive- 
7 (rarely brownish) ; endocarp with keels low but prominent, 
r with dorsal keel thin and very weak, beak deltoid, very weak, 
neti .5 mm. long, loop solid; apex of seed pointing at about the 
middle of the opposite side (or between middle and _ base). 
A highly variable species in which the extremes appear distinct, 
but transitional specimens, which are not a exception but the 
rule, indicate that they are a confluent ser 
P. illinoensis Morong, Bot. Gaz. 5: 30. (1880); Mem. Torr. 
Club 3: no. 2: 27 (1893); Graebn. in Engler, Pflanzenr. 4: fam 
a as (1907); Taylor, N. Am. FI. 17: pt. 1: 20 (1909); Hagstr., 
Res. Pot. 198 a et P. lucens ae Am. authors; not L., 
Sp. ‘pL 126 (1753). P. Zizii of Am. authors, in part; no t M. & K. 
in Roéhling, Deutschl. Fl. 1: 845 (1823) ; not Koch ex ety Enum. 
Plant. Germ. 1: 531 (1827). P. Proteus lucens C. & S., Linnaea 
2: 197 (1827) . e Pi Saoige Zizii C. & §., Linnaea 2: 201 (1827), 
in part. var. conmecticutensts Robbins in A. Gray, 
Man. ed. 5: “488 (1867): Morong, Mem. Torr. Club 3: no. 2: 31 
(1893). P. angustifolius var. connecticutensis Bennett, Journ. 
Bot. 39: 199 (1901). P. Zizii var. connecticutensis Morong ex 
Bennett, Journ. Bot. 39: 199 (1901); Graebn. in Engler, Pflan- 


Bennett, Journ. Bot. 45: 374 (1907). ?P. Ziett var. 
porrectifolius Bennett in Graebn. in Engler, Pflanzenr. 4: fam. 
11: 83 (1907). ?P. Zizii var. gracilis Bennett in Graebn. Pflan- 
zenr. |. ce. (1907). Spirillus lucens Nieuwland, Am. Mid. Nat. 
3: 17 (1913), as to plant, not as to source of name. seh 
Zizit Nieuwland, Am. Mid. Nat. 3: 17 (1913), in part 
illinoensis forma rosulatus Hagstr., Crit. Res. Pot. ee 1916). 
P. illinoensis forma homophyllus Hagstr., Crit. Res 

(1916). X P. pseudolucens Hagstr., Crit. Res. Pot. ‘00 (1916). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 155 


ae Pas lideres Benn., Trans. & Proc. Bot. Soc. Edinburgh 29: 
58 (1924 

Lakes and streams, southwestern Quebec to southern British 
Columbia, a to Florida, Texas and California, Map 
Mex., Centr. A , and W. I. the very numerous collections, 
the following are es most typical from the areas cited: QUEBEC: 
Gatineau R., Wakefield, Macoun 62021. Vermont: Lake Dun- 
more, Salisbury, Aug. 11 and 31, 1896 and Sept. 2, 1899, Ezra 
Brainerd; Lake Champlain, Sept. 1, 1880, C. £. Faxon; Lake 
Hortonia, Aug. 15, 1896, Ezra Brainerd; Barton ri 1829, herb. 
Robbins. MASSACHUSETTS: Fresh P., Cam bridge, many col- 
lectors; Wenham P., Essex Co., ae 2 fa 1867, J. W. Robbins; 
Leverett P., July 1874, dH. G. Jes Connecticut: Lake 
Saltonstall, E. Haven 1845, Robbins, ni 1850, Robbins (G, NY, 
type collection a lucens Vv. connecticutensis) ; Lake Kenosha, 

anbury, H. H. Eames 11351; Housatonic R., Aug 
Robbins; Twin Lakes, Salisbury, Litchfield Co., Ogden é Bolan 
1569. New York: Rockland L., July 17, 1892, Morong; Lake 
Cayuga, Cayuga, Aug. 20, 1884, Morong; Du ck L., Conquest, 
Cayuga Co., Hames, Ra ndolph & Wiegand 11175; bayou back of 
Renwint Park, Cayuga L., Ithaca, Tompkins Co., Hitchcock 
11175a; n. of R. R. bri ridge, Cayuga, Cayuga Co., Oct. 1886 
Wek. Dudley; Lake Oneida, July 28, 1880, Miss M. T. Hotch- 
kiss; Lake Cossayuna, Dobbin 11 12; Atuskatong Bay, Jefferson 
Co., Muenscher & Maguire eee Ballston L., Saratoga rat 
Muenscher & Clausen 4232 & 42 233; Warde: i. Albany Co., 
: uenscher & Clausen iy Little York L., Preble, Cortland Co., 

. N. Jones 7465; Greens L., Greene Co., Muenscher & Curtis 
ries PENNSYLVANIA: pres 1876, E. A. "Rau (ME, “ Hanover, 
_ ” printed on the label). DELAWARE: Stanton, Sept. 4, 1896, 

. Commons. VrireGinta: Four Mile Run, Alexandria, June 29, 
isos, I. Tidestrom; Dyke, Fairfax Co., Metcalf & Sperry _ 
Norra Carona: Orton L., Orton Plantation, 10 mi of 
Southport, Brunswick Co., Sept. 8, 1941, R. K. Gaitrey &, 
FLoripa: near the south New R. canal, beyond head of New R 
J. K. & G. K. Small 4437; near the Miami canal, J. K. & G. K. 
Small 4486; between Cutler and Longview Camp, Small & 
Carter 1007; Royal Palm Park, Dade Co., Moldenke 752; near 
Eustis, Lake Co., Oklawaha R., Nash 859 & 1 750, also Curtiss 
6692; Caloosa R., Simpson 386; ’ Everglades, Miami R., Small & 
Carter 1118 (F, NY , US, type collection of P. lucens v. floridanus). 
Ontario: Golden Le Renfrew Co., July 28, 1899, L. M. Umbach; 
Pelee Island, Lake Erie, Macoun 3023; River Trent, Macoun 
29965; Elziver, Hastings Co., Macoun 2996; Stokes Bay, Tober- 
mory, Bruce. Co., Krotkov 7027. Micuican: Whitefish L., 
Mackinac Co., Metcalf 2317; n. w. of St. Ignace, Mackinac Co., 
Pease & Ogden 24964; Lake Orion, Farwell 900; Houghton L., 


156 Rhodora [APRIL 


Roscommon Co., June 1876, herb. C. F. Wheeler; — : 
Allegan Co., Wight 8; Papaw L., Berrien Co., C. K. Dodge 171; 


, Whitley Co., Deam 49428; Tippecanoe Ls " Scovel 
58; Take i teeces Scovell 54, also Evermann 1079 & 1222, 
also Clark 6; Lake Maxinkuckee, Culver, Marshall Co., Aug. 27, 
1926, J. R. Churchill; Pine Station, Lake Co. , Aug. 8, 1876, E. J. 
Hill; Clarke, Umbach, also Lansing 1079; Atwood ii n. w. of 
Wolcottville, Lagrange Deam 553850; Lake Wawasee, Kosci- 
usko Co., Deam 56396 & 56401; Lake James, w. of Pokagon 
State Park, Beaches Co., Deam 56588 & 56539. WISCONSIN: 
Lake Wingra, Dane Co., June 20, 1892, L. A Cheney; White L., 
near Weyauwega, Waupaca Co., Hotchkiss & Martin 4432; 
Pickerel L., Nashville Twp., Forest Co., Hotchkiss & Koehler 
4331; Big Muskego L., Muskego bees Waukesha Co., Hotchkiss 
& Koehler 4245; Wind L., Norw y Twp., Racine Co., Hotchkiss 
& Koehler 4226 & 4227; Pike hed Hartford Twp., Washington 
Co., Hotchkiss _ Koehler 4262; Shawano L. , Washington Twp., 
Shawano Co., Hotchkiss & Koehler 43804; ’ Lake Noque Bay, 
Lake Twp., tives Co., Hotchkiss & Koehler 4826. I.ui- 
nots: Mississippi River Bottoms near Oquawka, July 23, 1873, 
Aug. and Sept. 1881, H. N. Patterson (vyPpE material); Pitts- 
burger L., Centerville, June 29, 1878, H. Eggert; Lake Villa, 
Sak 28, 1887, M. B. Waite. Manrtosa: Souris, July 7, 1883, 
J: acoun. Mrnnesota: Elk L., Itaska Park, Clearwater 
Co., Grant & Oosting 3184; Evans ee Kandiyohi Co., Metcalf 
2377; Green . a ie ohi amie Metcalf 2039, 2048 & 2045; Lake 

tcalf 1563; Ric i 


calf 1427 & 1432; Koronis L., Stearns Co. , Metcalf 1379, 1883 & 
1391, also Kubichek 110, 114, 115 & 121; Swan L., Nicollet Co., 
Metcalf 8; Martin L., Anoka Co., M etcalf 1829; Birch L., Sher- 
burne Co., Metcalf 1370; Horseshoe L., Chisago Co., Kubichek 
71 & 72; Prior L., Scott Co., Oosting 2938; Lake Ida, Douglas 
Co., Uhler & Warren 858; Lake Julia, Sherburne Co., Keck & 
Stevens 280; Borden L., Garrison Twp., Crow Wing Co., Hotch- 
kiss & Jones 3s 4113; Center City, Aug. 1892, B. C. Taylor; Two 
Inlet L., Becker Co., sian & Manning 237. Iowa: Armstrong’s 
Grove, Emmet Co. , July 1882, R. I. Cratty (cotype collection 
of P. illinoensis) : : Pou duly 189 4, B. Fink 191; Fremont Co., 
Aug. 2, 1898, herb. T. J. & M. F. L. Fitzpatrick; Clear L., 

Cerro Gordo Co., July 11, 1896, B. Shimek. Mussourt: Fish L., 

Sibley, Jackson Co., Mackenzie 293; Gasconade R., between 
Falcon and Nebo, Laclede Co., Steyermark 13910; Current R., 
n. of Buffalo Creek, e. of Bennett, Ripley Co., Steyermark 14285; 
Phillips Spring, s. e. of Van Buren, Carter Co., Steyermark 21236; 


1943] Ogden,—The Broad-leaved Species of Potamogeton 157 
Ae Creek, s. of French Mills, St. Francois Purchase Unit, 
21087. N 


SoutH Dakota: water Bs near Missouri R., Clay Co., Over 
4008. NEBRASKA: Rat rien 538; Swan L. mS of Whitman, 
Grant Co., Rydberg 1 rie Shafer Ris Garden Co., ‘Uhler & Martin 


Dewey L., Cherry oi Tolstead 614. Texas: Guadalupe R., 
Kerrville, Kerr Co., E. J. Palmer 12217; 16 mi. n. of Leakey 
Real Co., Cory 8502; San Antonio, Thurber 48; Montell Crete 
non Cory 9800; McKittrick Canyon, Guadalupe Mts., Cul- 
berson Co., Moore & Steyermark 3667 — typical). Wyomine: 
cur R., Elias Nelson 3386. Cou o: Tabequache Basin, 
Payson 1 v4: Cerro Summit, Pr ae of af ee Watershed, 
Baker 427. Uvan: Twelve Mile Cafion, Wasatch Mts., Tides- 
trom 517. New Mexico: Bah Augustine oo Organ Mts., 
Dona Ana Co., July 10, 1909, E. O. Wooton. CALIFORNIA: 
Marine Hospital, San Francisco, June 27,1892, 7. W. Blankinship; 
Mission Dolores, Bolander 274; near Sebastopol, Sonoma Co., 
Aug. 1900, M. S. Baker; Mountain L., San Francisco, Aug. 22, 
1920, Alice Eastwood (not typical). OrEGoN: creeks, western 
Oregon, 1880, T. J. Howell; in a warm spring, Harney Valley, 
June 10, 1885, card Howell (F, mixed with P. alpinus v. 
tenuifolius, C, G, US, n xed). Wasurnaton: Lake Chelan, 

ashington Forest fee aw Gorman 703; Headly ig ee 
Spokane Co., Sperry & Martin 731. Bririsn CoLu : Sum 1 
Li. Chilliwack Valley, J. M. Macoun 26815 Ghats a Dehn 
type of X P. perplexus in C, isotypes in G, NY). 

The highly complex species, P. illinoensis, has been variously 
treated by students of the genus. Morong! restricted the name 
to two collections of those known to him: Oquawka, II1., collected 
by H. N. Patterson, and Armstrong’s Grove, Iowa, collected by 
R. I. Cratty. Both plants are very robust and evidently grew in 
shallow, quiet, rich, muddy water. He complained that some 
botanists doubted the validity of P. illinoensis as a distinct 
species, but argued that it “. . . is evidently allied to lucens 
in habit, and with that species, P. angustifolius, P. spathulae- 
formis and P. heterophyllus, forms a very natural group, but it is 
clearly distinct from all of them in its vigorous growth, its 
abundant foliage, its ample floating and submerged leaves, and 
its large, strongly 3-keeled fruit.’’? Of these characters, the only 
One that can be considered of specific importance is that per- 

1 Thomas Morong, Mem. Torr. Club, 3: no. 2: 27 (1893). 

* Thomas Morong, Mem. Torr. Club 3: no. 2: 28 (1893). 


158 Rhodora [APRIL 


taining to the fruit, and examination shows that the fruits of the 
plants in this country called P. lucens or P. angustifolius are as 
strongly 3-keeled as those of P. illinoensis. Graebner’s treat- 
ment of this group is quite artificial. He followed Morong’s 
grouping rather closely but recognized numerous subspecies, 
varieties, subvarieties, and forms to account for the minor 
divergencies. The name P. illinoensis was, however, confined to 
the two original collections. Hagstrém evidently made a genuine 
attempt to understand the complex. Lack of sufficient material 
greatly handicapped him. He was misled by previous treat- 
ments into believing that P. lucens occurs in this country, but 
cited no specimens. Morong, who had realized that the plants 
he referred to P. lucens did not match any of the European forms 
of that species, treated them as var. connecticutensis. Hagstrom 
ignored this, except to question its being a variety of P. lucens, 
and does not even include it asasynonym. Some sterile speci- 
mens in North America resemble P. lucens of Europe but they 
lack the tendency for the lower leaves to have the blades reduced 
at the apex so that the midrib extends well beyond as a cusp. 
When mature fruits are obtained, the keels are seen to be promi- 
nent and acute (those of P. lucens are low and rounded) and the 
beak is more facial. Interestingly enough, specimens with ma- 
ture fruit do not have the compact, bushy habit of P. lucens and 
would not be confused with that species. 

Hagstrém considered P. angustifolius (P. Zizii) to be the hy- 
brid P. gramineus X lucens, a belief held or suspected by many 
students of the genus. He cites specimens from North America 
but calls attention to the fact that “. . many Zizii-like 
North American plants are not at all this dbad. but of another 
origin, and great carefulness is necessary when considering these 
difficult forms.’ If P. lucens does not occur in this country, the 
hybrid P. gramineus X lucens must be absent also. It may be 
suspected that the hybrid P. gramineus X illinoensis is not un- 
common with us for it is known that each of these closely related 
species hybridizes with species of other subsections. Many inter- 
mediate forms occur, but it is usually difficult to determine which 
are the results of crossing and which are ecological variants of 
one or the other species.? . The plants of the subsection Lucentes 


1J. O. Hagstrém, Crit. Res. Pot. has (1916). 
2 See discussion under Hysripis 


1943] Ogden,—The Broad-leaved Species of Potamogeton 159 


fruit as freely as the average species of Potamogeton, and I think 
that nearly all of them must be considered to belong to one or 
the other of the two species. 

The earliest specific name for the large-leaved plant is P. 
illinoensis Morong. This species has several variants, the recog- 
nition of which is made difficult in that ecological forms of one 
may simulate normal states of another. The Lucentes need to 
be studied cytologically and, until the hybrids having P. illino- 
ensis as a parent are better understood, a grouping of the variants 
into varieties would be merely an artificial one. In order not to 
delay this treatment further, P. illinoensis is treated in the 
broad sense, including its many variations. 


11. P. praELONGUS Wulfen 


RuizoMeE whitish, suffused or spotted with rusty red, as thick . 
as the stem or often much thicker. Stem simple or branched, 
whitish or olive-green, often zigzag, 1.5-4 mm. in diameter; 
stele with the proto-type pattern; endodermis of ‘Uecels inter- 
lacunar and subepidermal bundles present; pseu do-hypodermis 
1-3 cells thick. Leaves all submersed, ovate-oblong, (5—-) 10-20 
(—36) em. long, 1-3 em. Nags 13-25 nerves, 3-7 of them more 
prominent than the others, cordate or rounded at base and 
clasping 14 or % the siredietaneate of the stem; apex cucullate, 
splitting when pressed, or rounded; margin entire; lacunae of 
2-4 rows of cells each side of the midrib. SrTrPUuLEs white, 
nerves moderately OAL, oblong to oblong-linear or ovate- 
lanceolate, rounded at apex, (3-) 5-10 cm. long, sliniieen baa 


usually persistent and Bey agra PEDUNCLES varia in 
thickness, clavate, (5-) 15-60 cm. long. Berk with 6-12 
whorls, not crowded at anthesis, sometimes moniliform; in fruit 
3-5 cm. long, 1.1-1.4 em. thick. FLOWERS scanils or on ’ pedicels 


up to 5mm. long; sepaloid connectives cpr blades orbicular 
or elliptical, (1.7—) 2-2.6 (-2.9) mm. wide, claws 1-2 mm. long; 
anthers 1-2 mm. long. Fruits obovate, aunied on the back, 
cuneate at base, (4-) 4.3-5 mm. long (excluding beak), 3.2—4 
mm. wide; beak prominent, short and thick; dorsal keel acute 
and strongly developed, especially upward; lateral keels rounded 
or none; exocarp dark green; endocarp ‘with rounded lateral 
keels and a dorsal keel which is alate, thin and weak, beak linear, 
facial, about .8 mm. long, loop with a linear cavity; apex of seed 
pointing toward the basal end or slightly above. Plants om 
acterized by large ovate-oblong leaves, cucu ullate at the 
whitish stem, large phe stipules, and with long seth 
cles bearing large fruit 


160 Rhodora [APRIL 


P. praelongus Wulf., Arch. Bot. Roem. 3: 331 dae i 
Journ. Bot. 19: 241 (1881); Morong, Mem. Torr. Club 0. 2: 
32 (1893); Graebn. in Engler, Pflanzenr. 4: fam. 11: a (4907); 
Taylor, N. Am. Fl. 17: pt. 1: 22 (1909); Hagstr., Crit. Res. Pot 
250 (1916). Spirillus praelongus Nieuwl., Am. Midl. Nat. 3: 17 
oo 


Deep, cold water, southern Labrador, Newfoundland, Gaspé 
Menon Prince Edward Island, New England, and New Jersey, 
west to California, and north to Alberta, Mackenzie and the 
Aleutian Islands. Map 14. Eurasia. Reported from Mexico. 
The following, from a large series of specimens, are representa- 
tive: LaBrapor: Trout P., Blanc Sablon R., Straits of Belle 
Isle _— on the Quebec side of the river), Fernald & Long 27347. 

NDLAND: Cook Point, Pistolet Bay, Fernald, Gilbert & 
H otchkiss 27346; Tilt Cove, northern shores of Notre Dame Bay, 
no? Wiegand & Darlington 4482; Birchy Cove (Curling), 
Region of Humber Arm, Bay of Islands, Fernald, Wiegand & 
Kittredge 2441; Middle Birchy Ps eastern drainage area of the 
Humber R. system, Fernald & Wiegand 2444 (C, G, flaccid 
ppm Morley’s P., Humber Arm, Bay of Islands, Fernald, 

Long & Fogg 1211; Junction P., Whitbourne, Avalon Peninsula, 
Fernald, Long & Dunbar 26223. QUEBEC: Lac Pleureuse, Gaspé 
Co., Fernald, Dodge = Smith 25428 & 25424; West Branch of 
Mont Louis R., Gaspé Co., Fernald, Dodge & Smith 25422; Lac 
Sainte-Anne, Camps. Co., Victorin, Rolland & Jacques 33518; 
New-Rich mond, Bonaventure Co., Victorin, Rolland & Jacques 
83838; Lac Pore-Epic, Saint-Fabien, Rimouski Co., Rousseau 
30004. Prince Epwarp IsLaAnp: Dundee, oae Co., Fernald, 
Long & St. John 6767; French River, Queens Co., Fernald, Long 
& St. John 6768. New Brunswick: ‘Woodstock, Aug. 1884, Geo. 
U. Hay 4131. Nova Scotia: Earltown Lakes, Aug. 10, 1883, 
Mekay. 4129. Maine: Houlton, Aroostook Co., 1881, Furbish; 
5th Lake Musquacook, Piscataquis Co., G. B. Rossbach 82; St. 
John P., Somerset Co., St. John & Nichols 2107; Pushaw L., 
Orono, Penobscot Co. , Aug. 1891, Furbish; Hammond P., pee 
den, Penobscot Co. , Ogden & Steinmetz 1604; Hermo Her- 
mon, Penobscot Co.. , Ogden, Babel & Kozicky 1880 doesid form); 
Pleasant Ps Stetson, Penobscot Co., E. C. & E. B. Ogden & F. H. 
Steinmetz 1882, Gray Exsic. no. 1105; Androscoggin R., Liver- 
more Falls, Androscoggin Co., 1894, Furbish; Torsey P., Kent’s 
Hill, Readfield, Kennebec Co., "1892, Furbish. New HampsHIre: 
Ladd Py Stewartstown, Pease 1 4012: Streeter’s P., Lisbon, 
June 18, 1887, ex herb. E. & C. E. Faxon; Round P., Connecticut 
Lakes, Kendall, Gouldsborough & Doolittle 17. pi coma Shel- 
burne P., June 28, 1878, Pringle; Harvey’s P., West Barnet, 
Sept. 7, 1885, F. Blanchard; Willoughby, Orleans Co., Aug. 31, 
1904, A. Lorenz. MASSACHUSETTS: Chadwick’s P., Essex Co., 


: 

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1943] Ogden,—The Broad-leaved Species of Potamogeton 161 


S. K. Harris 589; Pleasant L., Hamilton, Essex Co., Fernald & 
er 749; Fresh P., Cambridge, Middlesex Co., many collectors; 
Harmon P., New Marlboro, Berkshire Co., June 29, 1912, R. 
Hoffmann; Stockbridge Bowl, Stockbridge, Berkshire Co., Aug. 
9, 1914, R. Hoffmann. Connecricur: Bantam L., Litchfield, 
Litchfield Co., July 25, 1883, Morong; Lake Saltonstall, Sept. 23, 
1880, LE. & C. EB Faron; Twin Lakes, North Bra nford, New 
Haven Co., June 22, 1887, FE. B. Harger; Twin Lakes, Salisbury, 
Litchfield Co., Eames & Godfrey 8685. New York: Southeast 
Bay, Saratoga L., Saratoga Co., Muenscher & Lindsey 2819; 
Dexter L., St. Lawrence a. Muenscher & M aguire 829; Lake 
Placid, Essex Co., Muenscher, Manning & Maguire 141; Cayuga 
L., Tompkins Co., July 29, 1884, Dudley; Pierrepont P., inlet 
from Lake Ontario, Woodville, Jefferson Co., House 10070; 
Ballston L., Saratoga Co., Muenscher & Clausen 4205. New 
JERSEY: Swartswood L., Sussex Co., Griscom & Mackenzie 10681. 
Ontario: Lake Hannah, Nipigon R., July 21, 1884, John Ma- 
coun; Lake Scugog, Wm. Scott 16451, also Cain 972; Inner Long 
Point Bay, Lake Erie, Cain 1050; Almonte, July 6, 1898, J. 
owler. MricuiGcan: Isle Royale, Cooper 312; Lake Manganese, 
Copper Harbor, Keweenaw Co., Hermann 8234; Tahquamenon 
R., Luce Co., Metcalf 2291; Douglas L., Cheboygan Co., Ehlers 
5383 & 1756, also June, July 1924, J. R. Swallen; Fremont L., 
Newago Co., July 7, 1926, Oosting; Crystal L., Montcalm Co., 
C. F. Wheeler 273; Pine L., vicinity of Agr’l Coll., C. F. Wheeler 
10; Haslet, Yuncker 713; Vicksburg, Kalamazoo Co., July 5, 1937, 
F. W. Rapp; Clear L., Jackson Sip. Hermann 6281. Onto: Put- 
in-Bay, July 16, 1898, A. J. Pieters. Inp1tana: Bear L., Noble 
Co., Deam 49394; Wolf L., Agnes Chase 1466; Lake Maxinkuckee, 
Scovell 45 & 66; near Lake Maxinkuckee, Scovell & Clark 1321. 
Naber chit Green Bay, 1892, Schuette; Twin Lakes, Marquette 
Co., Uhler & Warren 1079; Big Muskego L., Muskego Twp., 
Wanksaa. Co., Hotchkiss & Koehler 4246; Wind L., Norway Twp. 
Racine Co., Hotchkiss & Koehler 4228; Rice gh Nashville 
Twp., Forest Co., Hotchkiss & Koehler 4340; valley ‘of the Wis- 
consin R., near Lac Vieux Desert, L. S. Cheney pag nee 
Bebb 1009. MINNESOTA: Burntside L., July 25, 1886, L. 
Bailey; Lake Itasca, Clearwater Co., Grant py Swan a 
Far ja & Jones 3928; Center City, Chisago Co., July 1892, 
B. C. Taylor; Lake Chisago, Metcalf 1229; James L., Kandiyohi 
Co. , Metcalf 2387; Geneva L., Freeborn Co., Shunk & Manning 
80; ‘German Li LeSueur Co., ., Shunk & Manning 229; Leaf L., 
Becker Co., Shunk & Manning 368; Christine L. , Douglas Co., 
Shunk & Manning 410; Pelican L. , Wright Co., Linsdale & Keck 
153, also Uhler & Warren 797 6: Child L., Cass Co., Metcalf 
2371; Lake Johanna, Pope Co., Keck & Stilwill 455. Iowa: 
Clear L., Cerro Gordo Co., July 11, 1896, B. Shimek. Nortu 


162 Rhodora [APRIL 


Daxora: Roland Twp., Turtle Mts., Bottineau Co., M etcalf 522; 
Pelican L., Turtle Mts., Bottineau Co., Metcalf 546. NEBRASKA: 
Hackberry L., Cherry Co., July 18, 1912, Pool & Folsom; Dewey 
L., near Valentine, Tolstead 430; Enders L., Thomson 2. Mac- 
KENZIE: 2nd Eskimo L., 68° 10’ N., 132° 55’ W., Aug. 19, 1927, 
A. E. & R. T. Porsild. Auserta: Moose Lake district, Wood 
Buffalo Park, Rawp 1570 & 1571; Murdock Creek district, Wood 
Buffalo Park, Raup 1572. Montana: Lower Two Medicine 
Lakes, Maguire 485; Lower St. Mary’s L., Maguire 486. IDAHO: 
Henry L., Fremont Co., E. B. & L. B. Payson 2024; Gray’s L., 
Sperry & Martin 696. Wvyomina: Swastika L., Medicine Bow 
Mts., Albany Co., A. & R. A. Nelson 973, also R. J. Davis 378-W; 
Heart L., Yellowstone Nat’l Park, Clifford Richardson. CoLo- 


Peak, July 1879, Mrs. R. M. Austin. OREGON: Wallowa L., 
Cusick 2484; Klamath marsh, Klamath Indian Reservation, 
Coville 1254. Wasutnaton: Wiser L., Whatcom Co., M uenscher 
7647; Lake Leland, Jefferson Co., Otis 1767. British COLUMBIA: 

efor uver I., John Macoun 41382a & 88257; 
Beaver L., near Victoria, John Macoun 88258 & 88259. ALASKA: 
Atka, Aleutian Islands, Eyerdam 948 & 1122. 

P. praelongus is a beautiful species with large bright green ob- 
long leaves, and it often has extremely long peduncles terminated 
by huge spikes with massive fruit. It can scarcely be mistaken for 
any other species. Bennett! quotes remarks from Morong to 
the effect that this species fruits very late (Nov. to Dec.). How- 
ever, specimens in herbaria with fruit are mostly collected in 
July and August. Robinson and Fernald say, ‘Fruiting in June 
and July, withdrawing the stems to deep water to mature the 
fruit,’’? and this seems to be the case. More June collections of 
this plant are desirable. An extremely flaccid form (forma 
elegans Tiselius) was collected in Newfoundland by Fernald and 
Wiegand (Fern. & Wieg. 2444); one much like it grows in Hermon 
P., Maine (Ogden, Babel & Kozicky 1880). 


12. P. RicHarpsonu (Bennett) Rydberg 
RuizoMe whitish, yellowish or pinkish, not spotted; scales 
blackish, rounded at apex. Srem often branched, unspotted, 
1 Arthur Bennett, Journ. Bot. 19: 241 (1881). 
2? Robinson & Fernald, Gray’s Man. ed. 7: 74 (1908). 


1943] Ogden,—The Broad-leaved Species of Potamogeton 163 


1-2.5 mm. in diameter; stele with the trio-type pattern, the 
phloem on the inner face of the trio-bundle appearing as 2 
patches; endodermis of O-cells; interlacunar and subepidermal 
bundles absent; - pseudo-hypodermis 1 cell thick, at least partly so. 
LEAVES all submersed, coarse, mostly ovate-lanceolate, often 
ovate at the lower part of the stem and Serm tg narrowly 
lanceolate above, (1.5-) 3-10 em. long, (.5-) 1-2 em. wide, 
nerves (7—) 17-29 (—33), all rather prominent, 3-7 Fr them more 
prominent than the others, cordate at base and clasping 4% or 34 
the circumference of the stem; apex acutish but never sharp- 
pointed, sometimes rounded; margin with fugacious 1-celled 
translucent denticles, which are usually more or less appressed, 
lacunae of 2 or 3 rows of cells each side of the midrib. StrpuLEes 
whitish, coarsely nerved, ovate to lanceolate, obtuse when 
young, 1-2 cm. long, without keels, early disintegrating into 
whitish fibers. PrpuNcLES at base about same thickness as 
stem, often clavate, 1.5-25 cm. long. Sprkes with 6-12 whorls, 
not crowded at anthesis, sometimes moniliform; in fruit 1.5-4 
cm. long, and about 1 cm. thick. FLowenrs sessile or on pedicels 
up to .5 (-1) mm. long; s pe connectives hag se blades 
orbicular or allinteal, (13 ) 14-2 (-2.3) mm. wide, claws 
usually 1-1.5 mm. long; anthers (38>) 1.1-1.3 vot 4) mm. long. 
Fruits obovate, rounded on the back and at the base, sides 
plump or with a shallow a ry, (2.5-) 2.7-3.2 (-3. 5) mm. 
long (excluding beak), (2-) 2 6 (-3) mm. wide; beak prom- 
inent, up to 1 mm. long; vscis saree or none, rarely acutish; 
exocarp usually gray-green or olive-green; endocarp with rounded 
keels, beak linear, facial, about .8 mm. long, loop with a cavity, 
or at least a weak area ; apex of seed pointing toward the basal 
end or slightly above. Plants characterized by numerous 
coarsely-nerved pee submersed leaves, with stipules per- 
sistent Bes whitish fiber 
gabe (Bai) Rydb., Bull. Torr. Club 32: 599 
(1905). P. perfoliatus var. lanceolatus Robb. in Gray, Man. ed. 
5: 488 (1867); not ees Sapre ¥: soolanie: var. Richardsonit 
enn., Journ. Bot. 27: 25 (1 889); Morong, Mem. Torr. Club 3: 
no. 2: 33 (1893); Ga in Engler, Pflanzenr. 4: Tb ‘ae - 
(1907). P. perfoliatus L. sensu Taylor, N. Am. Fl. 17: 
Sah in par ; sensu Hultén, Fl. Alaska & ‘Yukon 101. (1940). 


Nini , Am. Mid. Nat. 3:17 (19 13). af petits ssp. Richard- 
sonii Hultén, Fl. Alaska & Yukon 102 (1940). 

Deep water of lakes and rivers, Labrador, Quebec, northern 
Maine, Vermont and western Massachusetts, west to Utah an 
California, north to British om Mackenzie, Alaska and 
the Aleutian Islands. Mar 1 


(To be continued) 


1943] Ogden,—The Broad-leaved Species of Potamogeton 171 


THE BROAD-LEAVED SPECIES OF POTAMOGETON 
OF NORTH AMERICA NORTH OF MEXICO 


E. C. OapEN 
(Continued from page 163) 

The following, selected from a very large series of specimens, 
are representative: LaBrapor: Grand Falls of Hamilton R., 
Doutt 3323. Quxrserc: Lake Mistassini, J. M. Macoun 3055; Lac 
au Saumon, Matane Co., Svenson & Fassett 38045; Roberval, July 
16, 1892, Mantods Lac William, near St. Ferdinand, Louis- 
Me arte, Laporte & Dudemaine 501; Odanak, Yamaska Co., Adrien 
1806; Longueuil, Chambly Co., Victorin 18462, also "Rolland 
43354, 483855, 483856 & 48481; Isle-aux-Noix, Richelieu R., 
Victorin 8179. Maine: Beau Lac, valley of St. Francis R., Aug. 
14, 1902, Eggleston & Fernald; Fish River Lake, Aroostook Co., 
Ogden 1717; Round P., Aroostook Co., Fellows 2007; Cross L., 
Aroostook Co., Fellows 4887. Vermont: Lake Champlain, 
Orwell, Addison Co., Cushman 6009; Big Otter Creek, Ferris- 
burgh, Aug. 12, 1887, Morong; Ferrisburg, Aug. 16, 1896, Eggles- 
ton; Swanton, Franklin Co., Blewitt 2137; mouth of Winooski R., 
Burlington, Chittenden Co., Aug. 30, 1903, N. F. Flynn, also 
Blake 2308; Rescue L., Ludlow, hates Co., July 12, 1906, 
Burnham. | MAassacHUsEtTs: Ponto 6 oL., Lan esboro, Berk- 
shire Co., Sept. 9, 1915, J. R. Churchill: ‘Mill’ R., New Marlboro, 
Berkshire Gai July 24, 1912, R. Hoffmann. New York: Muska- 
longe Bay, Lake Ontario, Jefferson Co., Muenscher & Maguire 
1782; Salmon R., Selkirk, Oswego Co., Fernald, Wiegand & 
Eames 1 4087; Rapids eg Niagara Falls, Aug. 21, 1886, 
Morong; Tioughnioga R., paoene Cortland Co., E. L. Palmer 
60; Silver L., Perry, Wyo ming Co. oy. Burkholder & Muenscher 
1 6424; Myers Point, Poa Tompkins Co., Jones & Hoffmann 
7467; ee a July 15, 1879, and Aug. 28, 1884, Dudley, also 
Aug. 15, 1893, Hermann von Schrenk, also Muenscher & Bechtel 
459; Bemus Point, Lake Chautauqua, Aug. 06, Ja. 
Churchill; Shushan, Dobbin 1072 & 1327. “sme thane of 
Brandywine Creek, Chester Co., no date, ex Herb. . M, 
Canby (F, on a sheet with P. perfoliatus var. Sntacaidie! per- 
7 the label refers only to P. perf. v. bup.). ONTARIO: Spawn- 

ing L., Temagami Forest Reserve, Cain 1051; Lake Nipissing, 
Chitty 306; Agawa R., Lake Superior, Pease 17983; Lake Hannah, 
Nipigon R., July 21, 1884, John Macoun; Mississippi R.., Galetta, 
Carleton Co., Ogden & Bolan 1631; Hay Bay, Tobermory, Bruce 
Co., Krotkov 7041; Golden L., Renfrew Co., Macoun 22176. 
MicHican: Ontonagon R., 1862, J. W. Robbins; Sault Ste. 
Marie, Aug. 6, 1881, E. J. Hill, also Aug. 25, 1882, Morong; Lake 
George, Homestead, Sugar Island, Chippewa Co., Hermann 7153 
& 7224; Douglas L., Cheboygan Co., Gates 258; Lake Charlevoix, 


172 Rhodora [May 


Ironton, Charlevoix Co., Ogden & Bolan 1678; Thread P., Flint, 
Aug. 11, 1909, Sherff; Detroit R., Gillman 65; Black L., Ottawa 
Co., July 14, 1926, Oosting. Onto: Sandusky Bay, Sept. 2, 1898, 
E. L. Mosely; Painesville, Aug. 21, 1887, herb. W. C. Werner, 
also Biltmore Herbarium 688. INnp1ANA: Wolf L., Whiting, Lake 
Co., Deam 55274A; Hamilton L., Hamilton, Steuben Co., 
Deam 56686; Long L., Wolcottville, Lagrange Co., Aug. 20, 
1930, Johnson & Myers; Culver Bay, Scovell & Clark 1057; Lake 
Maxinkuckee, Scovell 27a, also Evermann 1057. WISCONSIN: 
University Bay, Lake Mendota, Aug. 19, 1912, R. H. Denniston; 


Pease & Bean 26389; Lake Itasca, Clearwater Co., Grant 
Oosting 3206; Lake Augusta, Arno Twp., Cottonwood Co., 


Co., Rydberg 1792; Enders L., Thomson 5: Crescent L Thomson 
299; Shafer L., Garden Co., Uhler & Martin 1659; Lake Manawa, 


ee eee ee ee eT eS ee ee 


1943] Ogden,—The Broad-leaved Species of Potamogeton 173 


Omaha, Amy C. Lawton 53 (F, semi-narrow-leaved form). 
MAcKENzIE: Setidge L., Aug. 21, 1927, A. EF. & R. T. Porsild 
8098; rea Bay, Aug. 15, 1928, A. E. & R.T. Porsild; Big 
Point, Aug. 24, 1928, A. E. & R. T. Porsild. SASKATCHEWAN: 
Yorkton, hana &: Herriot 76872; Eagle L., Macoun & Herriot 
86871; Methye L., J. M. Macoun 4368; near mouth of McFarlane 
Ri; Lake Athabasca, Francis Harper 98; Cornwall Bay, Lake 
Athabasca, Rawp 6617. Atsperta: Bow River Valley, Steward- 
son Brown 694; near Banff, Macown 4383; Vermilion Lakes, 
Banff, John Macoun 4380, also Malte & Watson 960, also McCalla 
2370; near Chipewyan, n. shore of Lake Athabasca, Laing 147; 
also Raup & Abbe 4666. Montana: Missoula Valley near 
Frenchtown, S. Watson 397; Flathead L., Polson, Flathead Lake 
Co., Muenscher 10218; Flat Top L., Maguire 488; Lower Two 
Medicine Lakes, Maguire 4965; Swift Current Ridge L. (Bath 
Tub), Maguire 490; Trout L., Glacier National Park, Maguire 


805. Ipauo: Lake Pend Oreille near Hope, Sandberg, Mac- 
Dougal & Heller 956; Clarks Fork Valley below Weeksville, Lez- 
berg 1574; Blackfoot R., Trail Ranger Station, Bannock Co., 
Eggleston 9974; Henry La Fremont Co., E. B. & L. B. Payson 
2023; Mud L., Sperry & Martin 671; ‘Snake R., St. Anthony, 
Merrill & Wilcox 868; Portneuf R., McCammon, Bannoch Co., 
Ray J. Davis 328-37; Alturus L., Hvermann 492. Wyomina: 
Goose Creek L., Aven Nelson 2276; Laramie R., Elias Nelson 87 
& 3362; Lewis ify Yellowstone region, 1878, C. Richardson; N. 
Yellowstone L , Tweedy 413. COLORADO: vicinity of Twin Lakes, : 
July—Aug. 1902, C. Juday; Tomichi R., Parlin, Gunnison oN 
Aug. 20, sot. ’ Benjamin H. Smith; Kremmling, Grand C 

Beetle 1462. Uran: Lehi, Aug. 1883, M. E. Jones; Fish L., Mw 
E. Jones 5788, also Rydberg & Carleton 7522. NEvADA: Truckee 
R., Sereno Watson 1135. Ca.tFornta: Pit R., at Lookout, Modoc 


576; "Heather i. Mrs. R. M. Austin 1177; Truckee R., alt. 
4000 ft., W. W. Bailey 1135. Orrcon: Des Chutes R.., Lupine, 
Crook Co., Peck 9619; Ten-Mile L., near Lakeside, Coos Co., 
Peck 9009; ’ Willamette R., Salem, Hall 489 & 490; Willamette 
R., June 1890, Drake & Dickson, also Henderson 1009; Straw- 
berry L., Blue Mts., Cusick 3620; U. S. R. S. Main Canal, Kla- 
math Falls, Rian Co., Applegate 3416; Laidlaw, Crook Co. s 
Kirk & Whited 3141. Wasuincton: Lake Washington, Mercer 
Island, King Co., J. W. Thompson 7594; Lake Whatcom, What- 
com Co., Muenscher 7658; Seattle, June 26, 1889, E. C. Smith 
on also Aug. 1892, C. V. Piper; Quiniault, Conard 318; Prosser, 

ima Co., Cotton 809; Stevenson’s Ranch, Lake helan, 
Comma 677; Lake Sammamish, King Co., Otis 1678; also Aug. 


174 Rhodora [May 


1936, G. N. Jones. British CoLtumsia: Sicamous, John 
rt Mie 4127; Sumas L., Chilliwack Valley, J. M. Macoun 26817; 
Cowichan L., Vancouver Island, J. T. Howell 7679; Shuswap L., 
og Macoun 3049; Howser i; Shaw 771 & 777; Kamloops, 
John Macoun 3048. YUKON: Lewis R., Lat. 60°, Sept. 6, 1887, 
Dawson 3046. AwaskKaA: Unalaklet, A. E. & R. T. Porsild 1102; 
Votlik, June 10-14, 1926, A. E. & R. T. Porsild; Pastolik, 
July 16-20, 1926, A. E. d& R. T. Porsild; Holy Cross, July 5, 
1926, A. E. & R. T. Porsild; False Pass, Unimak Island, Aleutian 
Islands, Eyerdam 2156; Fairbanks, L. J. Palmer 1862; Ankow 
R. Piper r 4430; Matanuska, J. P. Anderson 838 & 1622; Pyramid 
Creek, compet Jepson 238; Selawik L., L. J. Palmer 638 
(US, mixed with P. gramineus var. maimus) 5 ta Ri; 


Lake Iliamna, Gorman 233; Naknek L., Point 795, Katmai ; 


Region, Alaska Pen., July 22, 1919, A. E. Miller; “Kotlik, Yukon 
R. delta A. E. & R. T. Porsild 847. 

og Richirdsondt j is closely related to P. perfoliatus and is con- 
sidered by many to be merely a variety or state of that species. 
Often, on vegetative characters alone, the two species are difficult 
to distinguish, but with fruiting specimens there is never any 
question. The larger fruit, borne on clavate peduncles, and 
always with a cavity in the endocarp loop, is quite distinctive 
from that of P. perfoliatus. In the western part of its range, 
this plant exhibits the characteristic ovate-lanceolate leaves with 
the strong white stipule-fibers, and only in the region east of 
New York state do its leaves and stipules approach those of P. 
perfoliatus. As intermediate specimens are nearly always sterile 
and scarcely found except where the ranges of P. Richardsonit 
and P. perfoliatus var. bupleuroides overlap, it is reasonable to 
suppose that they are hybrids of these two closely related plants. 

In many respects P. Richardsonii is intermediate between 
P. praelongus and P. perfoliatus and Hagstrém suggested that 
it “has arisen by cooperation between these two species.’ It is 
however a fertile species (for a Potamogeton) and with its greatest 
development falling outside the range of P. perfoliatus it cert- 
tainly cannot be considered a recent hybrid. The anatomy ° 
its stem is exactly that of P. perfoliatus and strikingly different 
from that of P. praelongus. It would seem that if P. praelongus 
were involved in its origin it would retain some of the funda- 
mental anatomical characteristics of that species, especially the 


1 J. O, Hagstrém, Crit. Res. Pot. 254 (1916). 


175 


1943] Ogden,—The Broad-leaved Species of Potamogeton 


\ rs aN = 
“ i 5 z 
ow . “le 2 \ *. ° 
NAN : Ge fa, “ip } 7) 
gi “NM OF ONO > ) > ; 9° 
AN y A = Ts eS “ Sy 
£ = ats OTe r fl \ 


RANGES OF PoTAMOGETON 


Sat Sy ia STN i alk i a lala 


176 Rhodora [May 


cortical bundles and a tendency to the prototypic stele. It 
would seem more logical to suppose that P. praelongus evolved 
from a perfoliatus-like ancestor, with P. Richardsonii as an inter- 
mediate step, except that P. Richardsonii is confined to North 
America (reported from Scotland!), while both P. perfoliatus and 
P. praelongus have a much wider distribution. Also such a 
postulate would involve the development of a proto-type stelar 
pattern from a trio-type pattern. The fossil record is of little or 
no help here, for both P. perfoliatus and P. praelongus appear for 
the first time in the Amber flora (Upper Eocene or Lower Oligo- 
cene) of Denmark,? and P. Richardsonii is not distinguished from 
P. perfoliatus in fossil literature. In America, members of the 
perfoliate-leaved species are reported only from the Pleistocene, 
near Ottawa, Canada.* 

A narrow-leaved form of P. Richardsonii is found in Big Stone 
Lake, between northern South Dakota and Minnesota. It was 
collected there by T. A. Williams in 1892 and in 1895 and by 
W. H. Over in 1922 (no. 14464). Another collection from the 
same lake (Over 3367) and collections from Sand Lake, South 
Dakota (Griffiths 6) and Lake Manawa, near Omaha, Nebraska 
(Lawton 53) appear to be nearly the same thing, and several 
other collections approach it. This form might be considered a 
variety were it not that typical P. Richardsonii frequently pro- 
duces narrow-leaved branches, especially in deep or moving 
water. 


13. P. perrFoLiaTus Linnaeus 
Migs whitish or pinkish, not spotted. Srem terete, .4-2 
. in diameter, often much bra nched; stele with the trio-type 

pasion, the phloem on the inner lane of the trio-bundle usually 
appearing as 2 patches; endodermis of O-cells; interlacunar an 
subepidermal bundles absent: pseudo-hypodermis 1 cell thick, 
at least partly so. Leavers all submersed, delicate, orbicular to 
ovate-lanceolate, usually ovate, 1-6 (7) em. long, .5-3 em. wide, 
nerves 7-21, not ‘strongly developed, only 1—7 of them prominent, 
cordate at base and oe about 34 the circumference of the 
stem; apex rounded or at least broadly obtuse; margin with 
fugacious 1-celled peal eiens denticles which are usually di- 

vergent, lacunae of 1-3 rows of cells each side of the midrib. 

' Arthur Bennett, Journ. Bot. 19: 241 (1881). 

? N. Hartz, Danmarks geologiske Untersogelse, 2: no. 20: 121 (1909). 

' J. W. Dawson, Can. Nat: n. ser., 3: 72 (1868). 


en athe ak ee ee 


1943] Ogden,—The Broad-leaved Species of Potamogeton 177 


STIPULES delicate, membranous, translucent, often appressed to 
the stem and seemingly a part of it, fugacious, ovate-oblong, 
rounded at apex, .4—-2 cm. long, without keels. PEDUNCLES 
about same thickness as stem, 1-9 cm. long. SprKkes with 2-8 
whorls, not crowded at anthesis, sometimes moniliform; in 
fruit 1-2 cm. long, and about .8 em. thick. FLowxErs sessile or 
on short pedicels up to .5 mm. long; apie connective greenish, 
a ae or elliptical, (1.2—) 1.4-1.9 (-2.3) mm. wide, 
claws .7-1.3 mm. long; anthers (.7—) .9-1.2 (-1.3) mm. long. 
Faas jue rounded on the back and at base, sides plump 
when nee, (2.3- ) 2.5-2.7 (-3) mm. long (excluding beak), 
(1.7-) 2 Te —2.3) mm. wide; beak usually prominent, up to .7 
mm. Fits keels rounded or none; exocarp usually gray-green or 
pbuh endocarp with rounded keels, beak linear, facial, 
out .6 mm. long, loop solid; apex i seed pointing toward the 
basal end. Plants characterized by erous, ovate, perfoliate, 
submersed leaves, with delicate weakly-nerved focneece stipules. 
Two varieties differing only in size but strongly marked as to 
their extreme variation 


Stem 1-2 mm. in diameter; leaves 1.5-3 cm. wide, 11-21 nerves, 


5-7 of them conde nn MAE ss chcges ot bees Se aa 13a. var. typicus. 
Stem .4—1.5 mm. in diameter; leaves .5-2 em. wide, 7-17 nerves, 
1-5 of heats prominent eta ts PRE A Sa ee eV 13b. var. bupleuroides. 


13a. P. perrouiatus L. var. typicus 
tee Pas L. ie Pl. 1: 126 (1753); Morong, Mem. Torr. 
Club 3: 33 (1893), in small part; Graebn. in aaa 
Planseny ry eat 11: 92 (1907), in part; Taylor, m. Fl. 17: 
pt. 1: 22 (1909), in small part; Hagstr., oa ig oe Pot. 254 (916). 
P. perfoliatus var. ae cilis Am. auth., art; perhaps C. & S8., 
Linnaea 2: 190 (1827). Spirillus ectoheiins Nieuwl., Am. Mid. 


13 
Rivers and lakes, southern Labrador, Quebec, and New 
Brunswick. Map 16. Eurasia, northern rica, Australia. 
The following are referred here: LaBRapor: Blanc Sablon R., 
Straits of Belle Isle (also on the Quebec side of the river), Ferna d, 
Wiegand & Long 27348; Blane Sablon R., Straits of Belle Isle, 
Pernali & Wiaaail 2450. QUEBEC: Blane Sablon K., Brest, 
Saguenay Co., St. John 90087; Blane Sablon R., Blane ‘Sablon, 
Saguenay Co., Lewis 130397; Lac Pleureuse, Gaspé Co., Fernald, 
Dodge & Smith 25421; L’'lle Plate, Longueuil, near Montreal, 
Victorin 20454. New Brunswick: Kennebecasis R., Lakeside, 
Kings Co., Svenson & Fassett 933. 


13b. P. perroutiatus L. var. BUPLEUROIDES (Fernald) Farwell 


P. bupleuroides Fern., Ruoopora 10: 46 (1908); Hagstr., Crit. 
Res. Pot. 258 (1916). 2P. crispus sensu Darlington, F'. Cestr. 


178 Rhodora [May 


23 eet not L., BD. Ph, .22<126:(1753). s perfoliatus sensu 
Morong, Mem. Torr. Club 3: no. 2: 33 (1893), in great part; 
sensu Ceasbr: in ack Pflanzenr. “ fam. 11: 92 (1907), in 
ale sensu Taylor, N. Am. Fl. 17: 1: 22 (1909), in part. 
P. perfoliatus var. bupleuroides foe. nes Mid. Nat. 8: 264 
(1923). 
Brackish or fresh ponds and quiet rivers, Newfoundland to 
Florida, west to Ontario, Ohio, and Louisiana, common only in 
the northeastern part of its range. Map 17. Also Guatemala 
(Lake Atitlan, 1906, C. M. Barber) and probably elsewhere in 
Central America. The following are selected from a large series 
of specimens: NEWFOUNDLAND: Killigrew’s, Conception Bay, 
Avalon Peninsula, Fernald & Wiegand 4484; Holyrood, gle 
3 Schrenk 207 (G, TYPE of P. bupleuroides, C, F, M, NY, US, 
sotypes); near Frenchman’s Cove, Bay of Islands, M Hehe & 
Csiseoh 10049; St. Georges, Fernald, Wiegand & Kittredge 2461; 
Highlands P. peer — 661; Topsail, Williamson 472. 
St. PIERRE ET Mique Mig uelon, Louis Arséne 40 & 48. 
QUEBEC: Percé, Victorin, teak Rolla nd & Rousseau 17286; 
Newport, Gaspé Co., Victorin, Rolland & Jacques 44467; Dart- 
mouth R., Gaspé Co., Co llins, Fernald & Pease 5578; Saint- 
Augustin, ‘Portneuf Co., Victorin, Rolland & Jacques 33879; St. 
Lawrence R. , st. Vallier, Bellechasse Co. , Svenson & Fassett 3031; 
Ile Plate, St. Lawrence R., Longueuil, 'Victorin 8178 & — 
near Ottawa, Victorin 1 10104. MAGDALEN IsLaNps: Cap-aux- 
Meules, Ile de ’Etang-du-Nord, Victorin & Rolland P9924; 
Etang-du-Nord wharf, Grindstone Island, Fernald, Long & St. 
John 6771. Prince Epwarp ISLAND: Tignish John Macoun 


Svenson & Fassett 932; Nipisiquit R., Bathurst, Gloucester Co., 
Blake 5483; St. John R., Westfield, Kings Co., Fernald 1610 & 
1611; also ene 2148; ’ Bass i, Kingston, Aug. 13, 1874, J. 
Fow Nova Scotia: South Ingonish, Cape Breton Island, 
Nichols 694; Mira Bay, Cape Breton Island, Macoun 20756; 
Bevis Brook, Port Bevis, Victoria Co., Fernald & Long 19694; 
Salmon R.., Truro, Colchester Co., Bean & White 19696: Truro, 
cn rr Co. , Prince & Atwood 1405; Port ow Shelburne Co., 

ernald & Long 23143; Midway (Centreville) L., Centreville, 
Di gby Co., Graves & Linder 19693; Sable Island, St. John 1124. 
Maine: Mattawamkeag L., Island Falls, Aroostook Co., Sept. 
7, 1897, Fernald; also Pease & Hopkins 22692; also Steinmetz 374; 
Pushaw P., Glenburn, Penobscot Co. , Ogden & Steinmetz 1544; 
Bradley, July 29, 1905, Ora Knight; Souadabscook Stream, 
Hampden, Penobscot Co., Fernald & Long 12392; Eagle L., 
Piscataquis Co., Ogden 1701; Indian P., St. Albans, Somerset 


1943] Ogden,—The Broad-leaved Species of Potamogeton 179 


Co., E. C. & E. B. Ogden 2008; South Paris, Oxford Co., 1897, 
Furbish; Roque Bluffs, Washington Co., Aug. 6, 1932, Knowlton; 
Mt. Desert I., Hancock Co., many collectors; Winterport, Waldo 
Co., Rossbach 77; Thomaston, Knox Co., Aug. 15, 1913, FE. B. 
Chamberlain; also Aug. 14, 1915, A. H. Norton; Sheepscot R.., 
Alna, Lincoln Co., Fassett 456; Sydney, Kennebec Co., Fernald 
& Long 12391; Topsham, Merrymeeting Bay, Sagadahoc Co., 
Steinmetz 526; Scarboro, Cumberland Co., Fellows 4679; Old 
Orchard, York Co., Fernald 1612; also Chamberlain & Knowlton 
571; also Parlin 1073; also Fellows 2966. New HAMPSHIRE: 
Connecticut R., Northumberland, Coés Co., Pease 12143; Mel- 
vin, Merrimack Co., Aug. 15, 1904, M. A. Day; also Sargent 29; 
Rye Beach, Rockingham Co., Aug. 24, 1886, Walter Deane; 
mith R., Danbury, Merrimac Co., Aug. 23, 1933, Charles 
Bullard; Rollinsford, Hodgdon 11. Vermont: Maidstone, Essex 
Co., Sanford 1225; Willoughby L., Aug. 31, 1917, E. J. Winslow; 
Missisquoi R., Swanton, Franklin Co., Blewitt 4319; Missisquoi 
R., Sheldon, Franklin Co., Blewitt 4320; Pelot’s Bay, Lake 
Champlain, Aug. 2, 1899, Nellie F. Flynn; also Carpenter 6069; 
Winooski R., Essex Junction, Chittenden Co., Blake 2212; 
Winooski R., Montpelier, Washington Co., Svenson & Fassett 
3040. Massacnusertrs: Chadwick’s P., Haverill, Essex Co., 
Harris 540; Mystic P., Medford, Middlesex Co., many dates, 
Morong; Fresh P., Cambridge, Middlesex Co., many collectors; 
Wareham, Plymouth Co., Svenson & Smith 822; Mashpee, Barn- 
stable Co., Fogg 3606; Tashmoo L., Tisbury, Dukes Co., Marthas 
Vineyard, Seymour 1027; Marthas Vineyard, McAtee 1060; Wash- 
ing P., Nantucket Island, Bicknell 118; also Sept. 8, 1909, J. R. 
Churchill; Congamond P., Southwick, Hampden Co., Seymour 
267; Horse P., Westfield, Hampden Co., Markert 76909; Housa- 
tonic R., Stockbridge, Berkshire Co., Aug. 20, 1902, R. Hoffmann. 
RuopeE Isuanp: Roger Williams Park L., Providence Co., July 
23, 1908, 7. Hope; Little Compton, Newport Co., Sanford 10192; 
Wash P., Block Island, Newport Co., Fernald, Hunnewell 
Long 8445. Connecticut: Selden’s Cove, Lyme, New London 
Co., Sept. 6, 1911, EZ. B. Harger; Paton Brook, Southington, 
Hartford Co., Aug. 28, 1900, C. H. Bissell; Fenwick, Middlesex 
Co., Sept. 2, 1884, C. Wright; East Haven, New Haven Co., 
Blewitt 1677 & 1978; Pistapaug P., Wallingford, New Haven Co., 


0, L. M. Underwood; Schroon L., Warren Co., Muen- 
scher & Lindsey 2728; Round L., Saratoga Co., Muenscher & 
Clausen 4210; n. of Canoga Marshes, Cayuga L., Seneca Co., 
Eames & Wiegand 9102; Riverhead, Southampton, Suffolk Co., 
St. John 2542. New Jersry: Mantoloking, Ocean Co., Edwards 


180 Rhodora {May 


d- ). Nortn Carona: North Channel, Currituck 
Sound, Shull 456 (G, M, NY, US). Ftortpa: Apalachicola, no 
date, herb. Chapman (G, NY). Ontario: Almonte, July 7, 1898, 


July 1884, C. Mohr (S, US). Lovtstana: Chifuncte Lighthouse, 
Lake Ponchartrain, Aug. 16, 1838, Riddell ( US). 

P. perfoliatus exhibits such an amount of variation that one 
might suppose it to include many varieties, yet these variants 
grade so insensibly into each other that lines can searcely be 
drawn. Here, however, one*is justified in refusing to give the 
variants formal rank, for they are sometimes found coming from 
the same rhizome, or as branches of the same plant. 

In America, two extremes appear to deserve varietal recogni- 
tion. One of these, a plant sparingly found in the cooler parts of 
northeastern North America, cannot be distinguished from P. 
perfoliatus of Europe. It approaches var. gracilis C. & S., which, 
being based entirely on variable foliage-characters, scarcely 
merits recognition as a genetic entity. Our plant is considered 
to be var. typicus. The few specimens from North America are 
all sterile. The other variety in North America is a smaller plant, 
with slender stem and delicate leaves having few prominent 
nerves. It is abundant from Newfoundland to New York and 
fruits freely. This latter plant has been separated from P. 
perfoliatus as P. bupleuroides. In its typical development in the 
brackish ponds of eastern Massachusetts, this plant differs so 
strikingly in size from the robust typical variety that, were it 


FOE ee eee ee ee ew 


Ye eee eee 


1943] Ogden,—The Broad-leaved Species of Potamogeton 181 


not for the transitional forms so common in Maine, Quebec, and 
New York, it might well be regarded as a separate species. 

In view of the fact that typical P. perfoliatus of Europe may 
have branches with foliage matching that of the variety bupleur- 
oides, and that in patches of the latter variety there may occa- 
sionally be found plants bearing robust leaves approaching those 
of the typical variety, it seems best to treat them both as variants 
of the same species. Any differences in the fruits of var. bupleur- 
oides and var. typicus of Europe are too slight and inconsistent 
to be used for distinguishing the varieties. 

The remarks of Fryer as to the variability of P. perfoliatus in 
Europe are of importance here: 

pic ssc perfoliatus has so considerable an amount of variation both 
in the shape and colour of the leaves that at first sight it seems easily 
separable ete distinct varieties, “but the examination of a good series, 
even of dried specimens, shows that the most extreme forms pass into one 
another so gradually that they are probably nothing more than states 
produced by local and often temporary conditions. An isolated plant 
growing in a newly dug clay pit afforded unusually good opportinuties for 
Families hei several years, and this plant changed so much in the 

shape, colour, and even texture of the leaves, that I am further induced to 
think that all ae British forms are mere states, not true varieties. When 


Mrs. Arber took a typical shoot of P. perfoliatus in July and 
kept it floating in a rain-water tub. “By October 1 most of the 
large perfoliate leaves had decayed and those on the new shoots 
were so much narrower and less perfoliate as to make it difficult 
to believe that they belonged to the same species.’ 

The work of the Pearsalls* with this species is very illuminating. 
Their ecological observations and experiments with plants of 
lakes and on those grown in tanks, led them to the conclusion 
that light intensity played a very great part in the shape of the 
leaf. However, leaf-shape was also influenced by other factors; 

* Alfred Fryer, Pot. Brit. Isles 41 (1900). 


ston ee Arber, hfe Plants 58 (1920). 
. H. Pearsall & W. H. Pearsall, Journ. Bot. 61: 2 (1923). 


“182 Rhodora [May 


the nature of the soil probably being the primary one. Plants 
on calcareous soil tended to have broader leaves than those of 
acid waters. In America var. bupleuwroides appears to vary ac- 
cording to the chemical nature of the water. Plants of brackish 
water invariably have small ovate leaves; those of neutral or 
acid waters tend to have their leaves larger and of a more elon- 
gate shape. 
HYBRIDS 


As nearly all Potamogeton-hybrids lack fruit and as flowers 
among the broad-leaved species are of slight, if any, value in 
distinguishing entities, it is necessary to base one’s opinions as to 
the probable parents on the characters of the leaves and the 
anatomy of the stem. It is important that both parents be 
found in the general vicinity of the hybrid they are supposed to 
have produced, though there is some question if this needs to be 
so, for such a hybrid may reproduce vegetatively and persist for 
long periods of time, sometimes, perhaps, after a parent has 
ceased to exist in that vicinity. Although interspecific hybrids 
(except P. gramineus X illinoensis) seem not to form mature 
fruit, they often produce pollen, some of which appears to be 
viable. It is suspected that in this way they backcross with the 
parents, producing plants which strongly simulate a given species, 
but show evidences of influence from some other species. In 
some such cases one parent can be named with assurance, the 
other with less confidence. In many such cases it seems wisest 
to correlate the plant with the species to which it shows the 
strongest affinity, even though in some minor details it does not 
agree with the normal development of the species with which it 
is placed. Unless this be done, the number of hybrids might be 
so great as to make a treatment unwieldy. It may well be that 
hybrids backcrossing with the parents will eventually supplant 
one or both parents, which will then not be found in the immedi- 
ate vicinity of the hybrid. . 

Hybrids should be considered, not as taxonomic units in them- 
selves, but as blends of two (or more) distinct entities. Most 
hybrids cannot be described, except by saying “intermediate 
between the parents.’ No attempt is made, except where quite 
obvious, to determine what variety of a species is the parent, nor 
is attempt made to designate which is the female and which the 


RRR CO = RRNA Fo rere, 


1943] Ogden,—The Broad-leaved Species of Potamogeton 183 


male parent. For many collections both parents could not be 
definitely determined. Most of these are, however, cited and in 
some cases discussed under the hybrid that is most probable, as 
determined from herbarium sheets. I hope that collectors who 
are acquainted with some of the cited localities will observe the 
plants in the field, for it is among the growing plants that hy- 
brids can be more easily recognized and their parents determined. 


P. ALPINUS X NODOSUS 

P. alpinus X nodosus (X P. subobtusus) a Crit. Res. 
Pot. 147 (1916). Not P. alpinus & ameri s (X P. recti- 
folius) Bennett, Journ. Bot. 40: 147 (1902): ei Graebn. in 
Engler, Pflanzenr. 4: fam. 11: 132 (1907). Not P. alpinus X 
americanus (X P. Faxont) Robinson & Fernald in Gray, vin " 
7:73 (1908). Not P. Faxoni Morong, Mem. Torr. Club 3:n 
22 (1893); nor Taylor, N. Am. i 17: pt. 1: 20 (1909). Not x p 
Faxoni Bennett, Journ. Bot. 46: 248 eae Not P. alpinus var. 
Fazxoni Stevens, Ill. Guide to FL. Pl. 96 (1910). 

This hybrid is extremely difficult to recognize for seldom are 
both parents obvious and the stem-anatomy of the two is so 
similar that it is of little help except to exclude from the parent- 
age all species with dissimilar anatomy of the stem. Hagstrém 
cites but one specimen, which, then, is the type of X P. subob- 
tusus: ‘Nov. Ebor.’, E. Tuckerman Jr (hb. Upps.).”” Material 
in the Gray Herb. and New York Bot. Gard. Herb., which is 
probably isotypic, may well be this hybrid; both are mixed with 
typical P. alpinus var. tenuifolius. Other collections which may 
be this hybrid are: New York: in a canal or raceway at Niagara 
Falls, Aug. 21, 1886, Morong (G); Niagara R., Aug. 23, 1886, 
Coville (US); Normans Kill, Kenwood, C. H. Peck 2 (US). 


P. ALPINUS X GRAMINEUS 

?P. alpinus * gramineus (X P. nericius) Hagstr., Crit. Res. 
Pot. 145 (1916). 

No American specimens can be cited that are undoubtedly this 
hybrid. However, plants growing in Mattagodus Stream, 
Prentiss, Penobscot Co., Marne, which I studied and collected 
(Ogden & Wright 2342) may possibly be this hybrid. It grew in 
close proximity to P. alpinus v. tenuifolius and P. gramineus. 
It is, however, more likely to be P. gramineus X P. perfoliatus 
var. bupleuroides; the latter species was not found in the im- 


184 Rhodora [May 


mediate vicinity, but is plentiful in the region. Other collections 
that might belong here are: Churchill, Manrrosa, Polunin 2070 
(C, G, this might be P. gramineus X some linear-leaved species. 
P. gramineus is quite obviously a parent, but there are no inter- 
lacunar bundles in the stem, and the reddish color suggests P. 
alpinus); Okanogan R., WasHincton, Sereno Watson 396 (G); 
Catfish L., Algonquin Park, Onrar1o, Macoun 22220 (C). Col- 
lections from QuEBEc: Hull Brook, Harrington 99100; Brigham’s 
Creek, Hull, John Macoun 85531; also Malte 118259, 118260, 
118261 & 118262, may be this hybrid. 


P. AMPLIFOLIUS X ILLINOENSIS 
?P. amplifolius X illinoensis (X P. scoliophyllus) Hagstr., 
Crit. Res. Pot. 164 (1916); ?x P. scoliophyllus forma barensis 
bid. 


Hagstr., ibid 


Hagstr6ém cites a specimen from Bear Lake, Manistee, MIcHI- 
GAN, collected by Morong in 1882 as the type for x P. scoliophyl- 
lus, and on the supposition that a form might someday be foun 
differing from this, further named the Bear Lake plant: forma 
barensis. None of the collections of Morong from Bear Lake, 
Manistee, Mich. that I have seen appears to be this hybrid, 
though both P. amplifolius and P. illinoensis are represented. 

agstrém reports this hybrid also from Cedar Lake, Ont. and 
Buckingham, Que., but cites no specimens. I have seen no 
specimen that I would refer here. 


P. AMPLIFOLIUS X PRAELONGUS 

Collections that may be this are: Wisconstn: Green Bay; 
sloughs between Halfway Slough and Bass Channel, June 29, 
1890, J. H. Schuette (F); Green Bay, between Duck Creek Chan- 
nel and Bars Channel, nearer to the latter, July 17, 1890 (or 
1899?), J. H. Schuette (F); Green Bay, June 23, 1891, J. H- 
Schuette (F). New York: very abundant, Southeast Bay, 
Saratoga L., Saratoga Co., Muenscher & Lindsay 2814b (G). 
ao Tinmouth P., Tinmouth, Aug. 5, 1907, G. G. Kennedy 


P. AMPLIFOLIUS X RICHARDSONII 
A collection from the north end of Tupper L., Franklin Co-, 
New York, Muenscher & Clausen 3784 (G), may be this hybrid. 


P. EPIHYDRUS Var. NvutTTALuu > PULCHER 


Virainia: outlet of Lee’s Millpond, Isle of Wight Co., Fernald 
& Long 12230 (G). 


1943] Odgen,—The Broad-leaved Species of Potamogeton 185 


P. NATANS X NODOSUS 
x P. perplexus ye X natans), Benn., Trans. & Proc. 
Bee we Edinburgh 29: 53 (1924). 

Bennett considers a collection from: British CoLuMBIA: 
Sumas L., Chilliwack Valley, J. M. Macoun 26815 to be this hy- 
brid and makes it the type of & P. perplerus. It may be this, 
but from the three sheets in the National Museum of Canada 
(vyPE), the Gray Herbarium, and the New York Botanical 
Garden, it appears to me to be a typical P. illinoensis. 


P. GRAMINEUS X NODOSUS 


?X P. argutulus Hagstr., Bot. Not. 106 (1908); Hagstr., Crit. 
Res. Pot. 220 (1916). 2x P. argutulus formseries nodosifolius 
Hagstr., Bot. Not. 106 (1908); ? Hagstr., Crit. Res. Pot. 220 

916). 


The following may belong here: MAINE: a eens Bay, 
Topsham, Sagadahoc Co., Steinmetz se! (G, ME, O); Sunkhaze 
Stream, Milford, Penobsco t Co., Ogden & Steinmetz 1607 (Q). 

EW HAMPSHIRE: Goonctese i. Corsiah Sept. 2, 1886, F. H. 
Knowlton (US). New York: Spencer : ee Spencer, Tioga Co., 
Thomas 1494 (G); east shore of Hudson R., above Coveville, 
Washington Co., M uenscher & Lindsey 2716 (O). DistTRICT OF 
COLUMBIA: in the vicinity of Washington, Aug. 5, 1879, L. F. 
Ward (US). West Virernta: Greenbrier R., Talcott, Summers 
Co., Morris 1343 (F, US, possibly P. illinoensis x nodosus). 


P. ILLINOENSIS X NODOSUS 


P. alpinus X pensylvanicus Benn., Journ. Bot. 28: 301 (1890). 
P. Faxoni Morong, Mem. Torr. Clu b3: no. 2: 22 LET 
in Engler, Pace 4: fam. 11:75 (1907); Taylor, N. Am. Fl. 17: 
pt. 1:20 (1909). ? P. es X rufescens Morong, Mem. Torr. 
Club 3: no. 2: 22 (1893). ? P. alpinus X Claytonii, Graebn. in 
Engler, Pflanzenr. 4: fam. 11:75 & 132 (1907). xX P. Faxon = 
P. americanus X pensylvanicus Benn., Journ. Bot. 46: 248 (1908). 
x champlainii = P. alpinus ce penayloanicus Benn., Journ. 
Bot. 46: 248 (1908). xX P. Faxoni = P. alpinus X americanus 
Robinson & Fernald, Gray Man. ad. 7: 73 (1908). Pp. alpinus var. 
Faxoni Stevens, Ill. ‘Guide to Fl. Pl. 96 (1910). P. illinoensis X 
nodosus Hagstr., Crit. Res. Pot. 200 (1916). P.lucens X nodosus 
Be P; subrufus) onto Crit. Res. Pot. 241 (1916). 

e following are referred here: QueseEc: Ile Bizard, Adrien 
1442 (G, MT). Vermont: Little Otter Creek, Ferrisburg, Aug. 
10, 1880, C. E. Faxon (G, MSC, NE, NY, US, type of P. Faxoni 
in NY); ‘Aug. 20, 1880, C. E. Faxon (G, NE); Little Otter Creek, 
near Barnum’s Mill and Ferrisburg Station, Ferrisburg, Aug. 19, 


186 Rhodora [May 


1882, Edwin Faxon (G, NE, NY), Aug. 23, 1882, Edwin Faxon 
(F, G, NE, NY): Otter Creek, below Vergennes, Addison Co., 
Aug. 24, 1882, Edwin Faxon (NE); Otter Creek, Middlebury, 
Sept. 5, 1896, herb. Ezra Brainerd (G); Little Otter Creek, near 
Lake Champlain, Ferrisburg, Addison Co., Ogden & Bolan 1589% 
(G); Lewis Creek, Ferrisburg, Aug. 20, 1881, FE. & C. E. Faxon 
(G, US), Aug. 18, 1882, Edwin Faxon (F, G, NY, isotypes of X 
P. champlainii). New Yorx: Lake Champlain, Plattsburg, 
Aug. 19, 1885, Morong (C, G, NY); outlet of Lake Seneca, 
Geneva, Aug. 14, 1884, Morong (C, G, NY, cotype collection of 
x P. subrufus). Vrrernta: northeast part of Four-mile Run, 
Chesapeake Bay region, Shull 471 (G, NY, US). Mussourt: 
Cooley L., Clay Co., Metcalf 1055 (US). Texas: Cebelo Creek, 
near Boerne, Kendall Co., H. J. Palmer 12907 (G, NY, US, pos- 
sibly not this hybrid). Ipano: Altmas L., Evermann 490 (F;, 
possibly not this hybrid). 


P. Noposus X RICHARDSONII 

x P. rectifolius (americanus < alpinus) Benn., Journ. Bot. 
40: 147 (1902); Graebn. in Engler, Pflanzenr. 4: fam. 11: 132 
(1907); Robinson & Fernald, Gray Man. ed. 7: 73 (1908). P. 
americanus Taylor, N. Am. Fl. 17: pt. 1: 19 (1909), in part. 
nodosus X Richardsonii Hagstr., Crit. Res. Pot. 148 (1916). 

Inurnors: Railroad ditch, Stoney Island, Chicago, Sept. 14, 
1900, E. J. Hill 171.1900 (NY, isotype material of X P. recti- 
folius), Aug. 30, 1901, Hill 179.1901 (C, isotype material of X 
P. rectifolius), Sept. 6, 1902, Hill 191.1902 (G), also Agnes Chase 
1477 (F) & 1994 (F). Orecon: Sauvie’s Island, Thomas Howell 
1497 (US). 

P. EPIHYDRUS X NODOSUS 

? P. americanus X pensylvanicus Benn., Journ. Bot. 46: 250 
(1908); not Benn., Journ. Bot. 46: 248 (1908). P. nodosus X 
Nuttallii (X P. subsessilis) Hagstr., Crit. Res. Pot. 190 (1916). 

Bennett mentions “. . . specimens gathered in New York 
and eastern Massachusetts by the late Dr. Morong . . - e 
and Hagstrém cites what may be the same Massachusetts col- 
lection. From Hagstrém’s description and an examination of 
the specimens, it is obvious he refers to Morong’s collection from 
Winchester, Massacuuserts, Sept. 1, 1880 (NY), Aug. 8, 1881 
(C, NY), Aug. 29, 1881 (G, 2 sheets, one mixed with P. nodosus) 
which is quite probably this hybrid. 


P. GRAMINEUS X NATANS 


? P. natans X gramineus (X P. sparganifolius) Hagstr., Crit- 
Res. Pot. 197 & 217 (1916). : 


1943] Ogden,—The Broad-leaved Species of Potamogeton 187 


Hagstrém cites: Pine Plains, New York, Hoysradt. What 
may be the same collection appears to me to be P. gramineus. 


P. GRAMINEUS X ILLINOENSIS 


P. Proteus Zizii C. & §., Linnaea 2: 201 (1827), in part. P. 

gramineus var. ? spathulaeformis Robbins in Gray, Man., ed. 5: 
487 (1867); not sensu Fern., Ruopora 23: 190 (1921), nor 
Ruopora 35: 130 (1933). P. heterophyllus forma maximu 
Morong, Mem. Torr. Bot. Club 3: no. 2: 25 (1893), in sone 
P. spathulaeformis Morong, Mem. Torr. Club 3: no. 2: 26 (1893); 
Graebn. in Engler, Pflanzenr., 4: fam. 11:91 (1907), as P. spathu- 
liformis. P. spathaeformis Tuckerm. in es opie in Gray's 
Man. ed. 5: 487 (1867), as a synonym; , Journ. Bot. 
130 (1900). xX P. spathaeformis Fern., ES 8: 294 (1906) ; 
Robinson & Fern. in Gray, Man., ed. "1: 74 (1908). P. varians 
Fryer, Journ. Bot. "97: 308 (1889), as to American specimens. 
? P. Zizit var. porrectifolius Benn. ex Graebn. in Engler, Pflan- 
zenr. 4: fam. 11: 83 (1907). ? P. Zizit var. gracilis Benn. ex. 
Graebn. in Engler, Pflanzenr. 4: fam. 11: 838 (1907). ? P. 
gramineus X i (X P. spathuliformis) Graebn. in Engler, 
Pflanzenr. 4: day ne 136 (1907). P. angustifolius Taylor, N. 
Am. Fl. 17: pt. 1: 18 (1909), in part. Spirilus Zizii Nieuwland, 
Am. Mid. Nat. 3: 17 (1913), in part. P. illinoensis gs lucens (x 
P. pseudolucens) Hagstr., Crit. Res. Pot. 199 (1916). P. gra- 
mineus X illinoensis (X 'P. de minutus) Hagstr., Crit. Res. Pot. 
209 (1916). P. gramineus X illinoensis X lucens (X P. pseudo- 
Ziziit) Hagstr., Crit. Res. Pot. 210 (1916). P. gramineus X 
lucens (X P. Zizii) Hagstr., Crit. Res. Pot. 210 (1916). xX P. 
Zizii forma pulcherrimus Hagstr., Crit. Res. Pot. 214 (1916). 
P. angustifolius of Am. auth., in part; not Bercht. & Presl, 
Rostlin 2: Alismac. 19 $ (1eaiy P. Zizii of Am. auth., in part; not 
M. &. K. in Rohling, Deutschl. Fl. 1: 845 (1823); nor Koch ex 
Roth, Enum. Plant. Germ. 1: 531 (1827). 

Of the numerous collections, the following may be mentioned: 
Vermont: Lake Champlain, Alburgh, Aug. 22, 1885, Morong 
(NY, isotype of X P. Zizii forma i Sane Windmill 
Point Reef, Alburg, Grand Isle Co., Muenscher, Manning & 
Maguire 73 (F); Lake Bomoseen, Castleton, Oct. i, 1898, W. W. 


type of P. gramineus var. cade eran. G, isotype); 
Meelis P., Medford, Oct. 8 & Sept. 24, 1865, some sheets without 
date, Robbins (G, MS SC, NY, cotype material of P. gramineus Var. 
spathulaeformis) ; also July 10, 1879 (G), wy, 25, 1879 (G, NY), 
Aug. 5, 1879 (NY), Aug, li, 1879 (F, NY), Sept. 1, 1879 (NY), 
Sept. 15, 1879 (F, NY), Aug. 5, 1881 (MSC, NY), July Septem- 
ber (C), all by Morong; also July 27, 1880, E. . E. Fax 


188 Rhodora [May 


(G, US); Fresh P., Cambridge, Sept. 5, 1853, Sept. 27 and Oct. 
24, 1865, herb. Wm. Boott (G); also Sept. 21, 1867, Robbins (NY); 
also Aug. 1876 (F), July 25, 1879 (F, NY), Aug. 2, 1879 (US), 
July 13, 1880 (MSC) and Sept. 14, 1886 (NY), Morong; also 
Sept. 17, 1880, C. E. Faxon (G); Pleasant P., Wenham, Sept. 13, 
1880, E. & C. E. Faxon (C, F, G, NY, US); also Sept. 9, 1881, 
Morong (NY); also E. C. & E. B. Ogden, 1762 (O). CONNECTI- 
cut: Lake Saltonstall, Sept. 22 & 23, 1880, EH. Faxon (G, US). 
New York: Cossayuna L., Washington Co., Muenscher & 
Lindsey 2786 (F, G); Perch L., Jefferson Co., Muenscher & 
Maguire 1684 (G); Lake Champlain, s. of Fort Ticonderoga, 

ssex Co., Muenscher, Manning & Maguire 75 (F); Junius, near 
Geneva, Bazter 5399 (US); Phillips P., Junius, Seneca Co., 
Eames & MacDaniels 3463 (G); Pine Plains, L. H. Hoysradt (G). 
Ontario: McKay’s L., near Ottawa, Malte 118258 (G, V); cove 
above Grass Creek Island, 7 miles below Kingston, Frontenac 
Co., Pennell 16333 (NY). Mucuican: Crystal Falls, Iron Co., 
Metcalf 2253 (US) & 2256 (G, US); Pine L., Ingham Co., 1893, 
C. F. Wheeler (C, mixed with P. illinoensis); Cassidy L., near 
Chelsea, Washtenaw Co., Hermann 9294 (NY). INDIANA: 
Grassy Creek, near Tippecanoe L., Scovell 37 (US); bed of inlet 
to Gage L., Steuben Co., June 17, 1903, C. C. Deam (NY); e. side 
of Lake James and w. side of Pokagon State Park, Steuben Co., 
Deam 56587 (G); Fletcher L., near Fulton, Fulton Co., Deam 
56524 (G); Lake Maxinkuckee, Evermann 544a (US). WiscoNn- 
stn: Green Bay near Suamico’s shore, July 11, 1886, J. H. Schuette 
(G); n. side of Puckaway L., Green Lake Co., Uhler & M cLaugh- 
lin 375 (F, G) & 1091 (US). Mrnnesorta: Green L., Chisago 
Co., July 1892, B. C. Taylor (NY); Pelican L., Koochiching:Co., 
Metcalf 1582 (G, US); Deep L., Cass Co., Metcalf 2373 (US); 
Snail L., Ramsey Co., Oosting 28164 (NY). Iowa: Armstrongs 
Grove, Aug. 1881, R. I. Cratty (NY). Nepraska: West Chain 
L., Thomson 73b (US); Dewey L., Tolstead 616 (G). IDAHO: 
Priest L., MacDougal 304 (C, NY); also Piper 3761 (G, NY, US); 
Pend Oreille L., Bonner Co., Muenscher 10208 (G, O) & 10212 
ratU Eh gesean Montezuma Well, Jackson 52 (US); also Taylor 


P. GRAMINEUS X RICHARDSONII 


(1924). 


_ The following appear to be this hybrid: OnTarto: Lake Nipis- 
sing, Chitty 265 (G). Micuican: St. Mary’s R., Sault Ste. Marie, 


1943] Ogden,—The Broad-leaved Species of Potamogeton 189 


EK. J. Hill 133.1881 (C); northwest end of Brevort L., ree 
Mackinac Co., Ogden & Bolan 1681 (G, O); Detroit R, Oct. 4 
1892, Farwell 4780 (G); also Aug. 25, 1892, Farwell (G); also Oct. 
4, 1892, Farwell (NY). Iturnots: Wolf L., Chicago, Agnes Chase 
1718 (F): also June 10, 1911, Earl E. Sherff (F). ALBERTA: 
Indian Graveyard, Peace R., Wood Buffalo Park, Rawp 1547 (C, 
NY). Conorapo: Carleton L., near Grand L., Shear & 
Bessey 53833 (NY). BritisH Coiumsra: Griffin Li; Macoun 4179 
(C, G, isotypes of & P. subdentatus @ petiolatus) ; Sumas Sy 
M. Macoun 26816 (C, type of P. Hagstrémii, G, NY, isotypes). 


P. GRAMINEUS X PERFOLIATUS Var. BUPLEUROIDES 


? eo abe sensu Morong, Mem. Torr. Club 3: no. 2: 34 (1893). 
mineus var. spathulaeformis sensu Fern., Ruopora 23: 
190 “(1921) and Pee i 35: 131 (1933); not Robbins in Gray, 
Man. 5: 487 (1867). X P. nitens var. subgramineus f. restrictus 
Hagstr., Crit. Res. Pot. 223 (1916). Xx P. subnitens Hagstr., 
i (1916). 


The following are referred here: NEWFOUNDLAND: Highlands 
Brook above bridge, Crabbes, Kennedy 78 (G); Otter P., Brig 
Bay, Fernald, Wiegand, Long, Gilbert & Hotchkiss 27341 (F, 
probably backcrossed with P. gramineus) ; also 27342 (G, prob- 
ably backcrossed with P. gts eus). QuEBEC: Matane R., 
Matane, Matane Co., Aug. 5 1904, Forbes 156 (C, G, one sheet 
of this collection in G is rather typical Fr. perfoliatus var. bu- 
pleuroides); Riviére Ouest, Gaspé Co., Vzctorin, Rolland & 
Jacques 44459 (G, MT); Tle du ae Iles de Boucherville, St. 
Lawrence R., Victorin & Rolland 44738 (F, G); aor ae 
St. Lawrence R., Rousseau 20444 (G); also 20445 (G, US, V); 
Ile Plate, St. Lawrence R., Longueuil, Chambly Co., Viclrn 
8176 (G); also stan (C, G); also © Rolland ‘wirie (C, F, G); Il 
Verte, St. Lawre ., Longueuil, Chambly Co., Rolland 13867 
yee “ako 13368. an “also 43376 (G); also 43378 (G); also 

2 (F, G); Lac Aylmer, Wolfe Co., Louis-Marie, Laporte & 
putea 301 (G). ANTICOSTI: Becscie R., Macoun 2983 (C); 
Salt Macoun 2986 (C). New Brunswick: St. John R., 
Ingleside, Westfield, Fernald 1609 (G); Hammond R., Hammond, 
Kings Co., Svenson & Fassett 3025 (G); Jacquet R.., ‘Restigouche 
Co., Svenson & Fassett 3030 (G); St. John R.., Lincoln, Sunbury 
Co, ; Fassett 2161 (G); gered, July 30, 1892, J. Fowler (US): 
Campbellton, Aug. 26, 1905, J. Fowler (US). Nova Scotia: 

“plaster hole lake’? near Danes Cape Breton Island, Nichols 
1037 (G); Shinimikas R., Northport, Cumberland Co., Fassett 
2150 (G); Midway (Centreville) L., Centreville, Digby Co., 
Graves & Linder 19691 (G, perhaps ‘backcrossed with P. gra-~ 
mineus). Matne: St. John R., Madawaska, Aroostook Co., July 
28, 1893, Fornell (G, NE, NY); Pushaw P., Oldtown, Aug. 21, 


190 Rhodora [May 


9, L. H. Harvey (US); Pushaw P., Glenburn, Penobscot Co., 
Deak Sie week % (G, 0); Pushaw P,, Hud son, Penobscot 
Co., Ogden & Steinmetz 2195 (ME, QO); Mattagodus Stream, 
Brontize, Penobscot Co., Stecnmetz a (G, ME, O, possibly P. 
alpinus X prawns); also Ogden & Wright 2342 (ME, O, possi- 

ly P. alpinus X gramineus). New Rata eee Connecticut 
R. Walpole, Fernald 440 (G); mouth ‘of brook, Melvin, Sargent 31 
(G). ont: Winooski R., Essex x Junction, Blake 2216 (US); 
Conileetiout R., Brattleboro, 1865, C. C. Frost (G). MassacHu- 
seTrs: Wenham P. , Sept. 13, 1880, Edwin Faxon (C, G, NY, US, 
see next citation) ; also Sept. 5, 1882, Edwin Fazon (G, NY, type 
collection of X P. subnitens; some of the Wenham Pond collec- 
ions appear to be this hybrid, probably backcrossed with 
perfoliatus var. bupleuroides; some cannot be distinguished from 
ecological forms of P. lana var. veaigiy hy CoNNECTI- 
cut: Taunton P., Newtown, E. H. Eames 11733 (G); Beardsley 
Fy Hp a E. H. ce 11741 (G, Sonate not this hybrid). 
New York: Hudson R., Mechanieville, Rensselaer Co., Muen- 
scher & Lindsey 2770 (G) & 2865 (G, O); Myers Pt., Cayu ga L., 
Aug. 13, 1884, herb. W. R. Dudley (NY); Old en ‘Conia 
near South Oxford, July 1, 1886, F. V. Coville (US); Chemung R. 
near Wellsburg, Lucy 1 0843 (F); Chenango R., Brisben, June 28, 
1887, F. V. Coville (US); slow stream, 4% mi. s. of the Jefferson 
Co. line, on lg No z. n. W. ts Sandy Creek, Oswego Co., 
Ogden & Bolan 1578 (G, QO). w JERSEY: river-edge above 
Phillipsburg, Taly 23, 1886, T. o Poet (NY). PENNSYLVANIA: 
near Easton, Sept. 8, 1868, T. C. Porter (F). Vrrernta: Dyke, 
Fairfax Co., Metcalf & Sperry 1684 (US). Ontario: Almonte, 
oa at c 7, 1806, J. Fowler (F, G); Missinaibi R., J. M. Macoun 


ais cross is between species of relatively unrelated subsections 
and the progeny are extremely variable and often very odd. As 
with most hybrids among the broad-leaved species of Potamogeton, 
no description can be given that is both inclusive and limiting. 
The hybrid must be distinguished as an intermediate between 
the two parents. It often appears with floating leaf-blades 
tapering gradually into the petiole and with clasping submersed 
leaves having sharp-pointed apices. Mature fruit is unknown 
in interspecific crosses, but may occasionally appear in back- 
crosses. The anatomy of the stem shows a blend between the 
two parents, presenting all the possible combinations with a trio- 
type or oblong-type of stele, O-cells or U-cells in the endodermis, 
and cortical bundles present or absent. Next to P. gramineus X 
illinoensis, this is probably the most frequent Potamogeton- 


} 


1943] Ogden,—The Broad-leaved Species of Potamogeton 191 


hybrid in North America. It is one of the easiest to recognize. 
The nearest approach to it is P. gramineus X Richardsonii which 
can usually be distinguished by its coarser stipules. 


P. GRAMINEUS X P. sp. 

P. angustifolius var. methyensis Benn., Journ. Bot. 29: 151 
(1891). P. Zizii var. methyensis Benn. i n Macou n, Cat. Can. PI. 
370 (1890); Graebn. in Engler, Phausent. 4: fain. 11: 83 (1907). 
‘A yay oe Benn., Trans. & Proc. Bot. Soc. Edinburgh 29: 50 
Methye L., near Methye Portage, meter noe July 18, 
1888, J. M. Macou n 4178 (C, rypx of P. methyensis). 

A number of collections are hybrids with Pr gramineus quite 
evidently one of the parents. Among these is the Methye Lake 
plant upon which P. methyensis is based. From the single sheet 
seen by me I cannot determine the other parent. P. illinoensis 
might seem a good guess, but it is not reported so far north. P. 
alpinus and P. Richardsonii abound in the region, but Iam unable 
to see marked influence of either of these species. Hagstrém 
may not have seen this collection; at least he mentions the name 
only in the index to his Critical Researches. He is, however, re- 
ferring to it (among others) when he says, “. . . but many 
Zizii-like North American plants are not at all this hybrid, but of 
another origin, and great carefulness is necessary when con- 
sidering these difficult forms.’”! 


P, ILLINOENSIS X RICHARDSONII 
INDIANA: Bass L., Starke Co., Aug. 7, W. S. Blatchley (Deam). 


P. ILLINOENSIS X PERFOLIATUS Var. BUPLEUROIDES 

? x P. subdentatus « sessilis Hagstr., Crit. ee Pot. 201 (1916); 
not X P. subdentatus @ petiolatus Hagstr. s 

New York: Bie hg Bay, rae 12264 ic, U8); Onondaga L., 
Aug. 1, 1890, L. M. Underwood (NY). FLoripa: Apalachicola, 
herb. 1 aaa (G, NY, US);mouth of Choctawhachee R., June 
18, 1880, C. Mohr (US). AuaBaMa: estuary of Mobile R., fely 
22) 1884, C. Mohr (US). 

Hagstrém cites a specimen from Queenston, ONTARIO (without 
mentioning collector’s name and date), which he considers to be 
P. illinoensis X perfoliatus. I have not seen this collection. 


} Hagstrém, Crit. Res. Pot. 216 (1916). 


192 Rhodora [May 


The collection from Griffin L., Bririsa Cotumsta, which he also 
cites as having these parents, is P. gramineus X Richardsonii. 


P. ILLINOENSIS X P. sp. 

s a peculiar form of P. natans,’’ Morong, Bull. Torr. 
Bot. Club 13: 145 (1886). P. tonsa Small, Fl. Se. U. § 
37 and 1326 (1903); Benn., Journ. Bot. 45: 376 (1907); Graebn. 
in Engler, Pflanzenr. 4: fam. 11: 62 (1907). ? P. Tepperi Benn., 
oe Bot. 45: 373 ay oad in Engler, hein jones 4; fam. 

2 (1907). : ently an immature f 
P natans, Taylor, N. Peg TL. 17: 28.1: 16 (1909). “prob- 
ably belonging to the group Amplifolii,” Hagstr., Crit. Res. Pot 
268 (1916). 

Fioripa: Blackwater R., May A. H. Curtiss (NY, 
TYPE and isotype of P. Wontaaiien): ae June 22, 1886, Curtiss 
(NY). 

Further collections and perhaps a study of the living plants 
will be needed to determine the exact nature of this plant. It is 
possible that it is a cross between P. illinoensis and a linear- 
leaved species. The stem-anatomy is: stele with the proto-type 
pattern, endodermis of U-cells, interlacunar bundles weakly 
developed and only in the outer interlacunar circle, subepidermal 
bundles absent or weakly developed, pseudo-hypodermis present 
or absent. Curtiss collected in the same river, and by the dates 
on the sheets presumably at the same time, fragmentary bits of 
P. foliosus Raf. var. macellus Fern. Other linear-leaved species 
of Potamogeton, subgenus Ewpotamogeton, found in the general 
region are: P. pusillus L. (P. panormitanus Biv.), P. Berchtoldt 
Fieber (P. pusillus of auth., not L.), P. diversifolius Raf., and P. 
capillaceus Poir. None of ‘thee linear-leaved species has inter- 
lacunar bundles and all have O-cells in the endodermis, so if one 
is a parent, the other parent must possess interlacunar bundles 
and a U-celled endodermis. The only broad-leaved species 
agreeing with this would be P. illinoensis. The stelar pattern of 
the plant in question leads me to consider P. pulcher as a possible 
parent, though it lacks interlacunar bundles and has an O-celled 
endodermis. In such case, the only linear-leaved species that 
can be considered to be the other parent is P. pectinatus, in the 
subgenus Coleogeton; it being the only linear-leaved species iD 
the region with interlacunar bundles and with U-cells in the 
endodermis. It is possible that the plant is a pronounced ec0- 


1943] _Ogden,—The Broad-leaved Species of Potamogeton 193 


logical form of P. Oakesianus or P. natans, neither of which has 
been otherwise found within 600 miles of Florida. 


P. PRAELONGUS X RIcHARDSONII 


MicuiGcan: Saulte Ste. Marie, Aug. 4, 1881, #. J. Hill (US). 
Uraun: Fish L., Sevier Co., Tanner 5786 (F). 


P. PERFOLIATUS Var. BUPLEUROIDES X RICHARDSONII 


It is reasonable to suppose that the two closely related species, 
P. Richardsonii and P. perfoliatus, would hybridize rather freely 
where their ranges overlap. Such appears to be the case, for 
these species remain quite recognizable (though exhibiting much 
ecological variation) throughout their separate ranges, but where 
these ranges overlap, all intergradations occur. One might 
consider the entities that bridge the narrow gap to be those from 
which the two species have evolved were it not that such plants 
are always sterile and often exhibit other evidences of hybridism. 
Such plants occur in Quebec, northern Maine, New Hampshire, 
and are especially abundant in New York. They occasionally 
occur elsewhere. Although the evidence is strongly in favor of 
numerous hybrids between P. Richardsonii and P. perfoliatus 
var. bupleuroides, it is practically impossible to be sure of this 
with individual collections, for ecological or other conditions of 
growth might cause a simulation of this in either of these closely 
related species. For this reason, no collections are cited. 


P. BERCHTOLDI X PERFOLIATUS Var. BUPLEUROIDES 


non-stratified O-cells; interlacunar and subepidermal bundles 
absent, or with a few weakly developed subepidermal ones; 
pseudo-hypodermis 1 cell thick. Lzaves all submersed, delicate, 
oblong to oblong-linear, .5-4 cm. long, .2-.6 cm. wide; nerves 3-7; 
sessile and cuneate at base, semi-clasping; apex obtuse or acutish, 
but not sharp-pointed; margin entire (with 1-celled denticles on 
the Nantucket specimen) ; lacunae of 1 or 2 rows of cells each side 
of the midrib, commonly with a pair of glands at the base. 
StrPuEs delicate, more or less persistent, ovate-oblong, rounded 
at apex, .3-1.5 cm. long, without keels. PspuNcLES about same 
thickness as stem, sometimes slightly incrassate, 1-4 cm. long. 
SPrkes with 1-4 whorls, not crowded at anthesis; in flower .2-.5 | 


194 Rhodora [May 


m. long, .2-.3 em. thick. FLowers sessile; sepaloid connectives 
Seon, blades orbicular or elliptical, 1-1. 8 mm. wide, claws . 
8 mm. long; anthers .7-1 mm. long. Mature Fruits unknown. 
Immature fruits obovate, about 9 mm. long (excluding beak) 
and .8 mm. wide; beak prominent, about .5 mm. long, curved 
toward the back; keels none; exocarp greenish; seed aborted, 
apex pointing toward basal end. Winter Bups abundantly 
produced late in the season, mostly near the rhizome, with 
leaves appressed to the shortened axis, or tightly appressed for 
about half their length and with the upper halves strongly 
divergent; basal glands prominent and forming adventitious 
roots. 

P. mysticus Morong, Bot. Gaz. 5: 50 (1880); Mem. ea oo“ 
3: no. 2: 34 (1893); Graebn. in Engler, Pflanzenr. 4: fam. 11: 95 
(1907); Robinson & Fern. in Gray, Man. ed. 7: 75 p08), with 
suggestion that it is ‘(probably a hybrid” of P. bupleuroides 
(perfoliatus var. bupleuroides) and P. pusillus (Berchtoldi); Tay- 
lor, N. Am. FI. 17: pt. 1: 22 (1909), “apparently a depauperate 
form” of P. perfoliatus, and ‘‘may be a hybrid’; Britton in 

t. & Brown, Ill. Fl. 1: 80 (1913) “apparently a depauperate 
form? of P. perfoliatus and “Perhaps a hybrid.” P. bupleuroides 
X pusillus (X P. mysticus) Hagstr., Crit. Res. Pot. 259 (1916). 

In brackish water, Maine, Massachusetts, and Maryland. 
The following are referred here: MAINE : Dunston Marshes, 
Searboro, Sept. 20, 1920, A. H. Norton; Stuart Brook, West 
Scarboro, Scarboro, Cumberland Co., Steinmetz & Marston 539, 
Plantae Exsiccatae Grayanae 905; also Steinmetz 617; also Ogden, 
Rollins & Wiggins 1731; Lily P., Fortune Rocks, Biddeford, 
Sept. 3, 1899, G. G. Ken nedy. Massacnusetts: Mystic P., 
Aug. 13 and Sept. 3, 1865, Wm. Boott; Mystic P., Medford, on 
several dates in 1879, 1880, and 1881 by Morong and the Fazons; 
Miacomet P., Nantucket, ‘July 13, 1887, Morong. MaryYLAND: 
near Ocean City, H. L. Clark 6 


This interesting hybrid was seiecth) first collected by Wm. 
Boott in Mystic Pond (now called Mystic Lakes) in 1865. In 
1879 Morong found the plant still there, and in 1893 wrote, 
“Since I obtained this . . . in 1879, I have visited the 
locality for several years in succession, and, though I have always 
found the plant growing vigorously, yet it has shown no signs 0 
prefecting [sic] fruit.”! In the summers of 1935, ’36 and ’37 I 
searched the Mystic Lakes carefully for it without success. The 
town of Medford extends only into the lower lake where P. per- 
foliatus var. bupleuroides now abounds. In this lower lake n° 


1 Morong, Mem, Torr. Club, 3: no. 2: 35 (1893), 


1943] Ogden,—The Broad-leaved Species of Potamogeton 195 


P. Berchtoldi was seen, but there is in the middle lake, now 
separated from the lower by a dam, a flourishing patch of P. 
Berchtoldi var. tenuissimus. This var. of P. Berchtoldi was 
found with the hybrid at Scarboro, Maine, and it appears safe to 
designate this variety as one of the parents of the Mystic Lakes 
and Scarboro plants. Whether it is this variety of P. Berchtoldi 
which is a parent of the other specimens cannot be said. A 
search at the Scarboro locality, where the plant thrives in abund- 
ance, failed to reveal any mature fruits, although flowering spikes 
were plentiful. One of the most striking characteristics of this 
plant in the latter part of the season is its abundance of winter 
buds. In its morphological characters as well as its stem- 
anatomy this hybrid is intermediate between the supposed 
parents. 
EXPLANATION OF PLATES 
Piate 746. FRuirs OF POTAMOGETON (all X 5) 
LPINUS Var. yar tae ee F1Gc. 1, Maine, Fernald 117; ria. 2, Newfound- 
iad ‘Kennedy sey a. 3, basa ee we 2172. 
er Sv area LLIPTICUS: FIG. 4, e, Sept. 5, 1894, Fernald; Fic. 
os Quebee, Fernald, Long & St. nine 6766 gue iy 6, New York, House 


ne pec Fias. 7, 8, 9, Sitch gym ge Fernald & Wiegand 4467. 

P. AMPLIFOLIUS: 10, Ohio, ’Goodric h 209; Fics. 11 & 12, New York, 
Muenscher & Magu e 723 (12 with mesocarp removed) ; FIG. 13, Ontario, 
Aug. 19, vlontns Fowler, 

P. puLcuEr: Fic. 14, Rhode Island, Fernald, Long & Torrey 8444; Fic. 15, 
Mussachieetia, July 1887, Morong; Fics. 16, 17, Manachiiastis, June 16, ‘1878, 
C with pene remov: 

P. Noposus: FIG. sot hig + sota, Hotchkiss & Jones 480; r1a. 19, Connecti- 
cut, 1845, Robbins; 20, ‘Wisconsin, July 28, 1891; Fie. 21, District of 
Columbia, Sept. 25, 1897, Stee 

P. NATANS: FIG. 22, Ontario, ae & Bolan 1646; ria. 23, Minnesota, Grant & 
Oosting 3203; Fias. 24, 25, New York, Ogder & Bolan 1580 (25 with mesocarp 
removed). 

P. Oakes SIANUS: FIG. 26, Massachusetts, Aug. 28, 1851, Robbins; Fia. 27, 

uebec, Fernald, Long & St. John pete FIG. = Nova Scotia, Long ’& Linder 

n 


P. ILLINOENSIS: FIG. 30, Iowa, July 21, i883, F Cedi ria. 31, Illinois, Sept. 
1881, Patterson (ryPE); Fics. 32, 33, Fl lorida, Curtiss 6692; ria. 34, Indiana, 
ge Sohn FIG. 35, New York, R. Hitche 


MINEUs: FIG. 36, Was hington, J. W. Thom on 7589; Fic. 37, New- 
foundland, Fernald & Long 1210; Fria. 38, gvamend Pete if 2187. 
LONGUs: Fas. 39, 40, New Yo rk, Mue er & Maguire 1747 


Aug. 21, 1886, Mororg; ria. 43, Quebec, Viclorin 18462; 1a. 44, Michigan, 
— Robbins; ric. 45, Washington, Aug. 1892, Piper; ric. 46, Montana, 
se oben 

OLIATUS Var. BUPLEUROIDES: FIG. 47, Newfoundland, Robinson 


P. 
Sechrenk "207 TYPE); FIG. 48, Maryland, Coville 118; Fic. 49, Nova Scotia, 
Graves & Pd ed Fig. 50, nabs , Fernald, ‘Long & "St. John 6771 


Gaeaeigs removed). 


196 


Rhodora 


[May 


Puate 747. PoraMOGETON NATANS: CROSS-SECTION OF STEM, X 35. 


Camera-lucida drawing fee contente not shown) by Francis T. Horne from 


preparation by Edith B. Ogd 


LATE 748. CAMERA-LUCIDA DRAWINGS Biren carga NOT SHOWN) 
FROM STEMS OF POTAMOGETON SHOWING Typ 


Pia; 1, F; cab went x 40 (Ogd 

S var. BUPLEUROID 
detach 1105); Fra. fe ? GRAMINE 
1552); Fia. 5, P. cr 
OLDI bang TENUISSIMUS 


US var. rid dei 
MINEUS Var. TYPICUS, X 


den & aor 09) FIG. 2, P. PERFOLIA- 
X 75 (Steinmetz 


526); Fic. 3, P. noposus, X 75 
a eT gden & Steinmetz 
x 75 (Og =. & Bolan 1644); FIG. § 


RFOLIATUS var. BUPLEUROIDES (XE 


‘ a X PE 
mysticus), < 100 (Ogden, Rollins & Wiggins 1731). 


List of NUMBERED EXSICCATAE 


Abrams, L. R. 6094, 8677 natans, 
O241 a amplifolius. 

Adams, J. W. 293 amplifolius, 511 
Oakesianus. 

Adams & Tash. 512 crisp 

— & Trudell. 378° cane 
ius 

drien, F. 1316 perfoliatus 
da idl A 1442 illinoensis x 

1750 perfoliatus v 


ardsonii), 1809 nodosus, 1975 per- 
foliatus v. i ge gis 


er 

Allen, J. A. 55. sg 149, 180, 
181 gramineus 

Anderson, J gramineus ap- 


8 Ri oe 1344 natans, 

1522 Richardsonii. 

Anderson, R. M. 118276 Richard- 
sonii. 

— & Fassett. 19349 ampli- 
olius 

Anderson, Smith & Weatherby 
1166 natans. 

Ans Im, Bro. M. 18 natans, 20 


gramineus v., 298 perfoliatus v 


approaching v. ma na- 
ee 3416 icharie 4489 
nata 
Asati. L. 38 pulcher, 40 perfolia- 
tus v. bupleuroides, 41 polygoni- 
i 43 perfoliatus v. bupleu- 
roi 
re & Benedict. 15429, 16636 
no 
‘Seiie & Holway B46, 
B69, B403 gramineus v. Maximus. 


Austin, Mrs. R. M. 575, 1177 
ichardsonii, 1672 natans. 
Bailey, L. H. 86 natans. 
Bai 1135 Richardsonii. 
427 illinoensis. 
Ballard, C. A. 731 ot. D dete 
approaching v. Mahe 
Barkley, F. 783. ‘granineds 
1996 gramineus v. myriophyllus, 
1997 is” yar omy 
axter, S. 5382 praelongus, 
5385 Riel chardsonii i, 5388 crispus, 
5389 nodosus, 5391 illinoensis, 5392 
us, 2395 natans, 5396 1 il- 
linoensis, 5399 gramineus illino- 
ensis, 5402 illinoensis, 5403 ampli- 
— 
& White. 19675 natans, 
19696 perfoliatus v. bupleuroides, 


gr X1 ’ 

22963 alpinus v. tenuifolius. 
Bebb, R. illinoensis, 995 na- 
tans, 1007 illinoensis (perhaps 


gramineus  _ illinoensis), 1 


natans, 1009 praelongus, 1538 il- 
linoensi 

Beetle, A. A. ‘aes Richardsonii, 
2341 gramineu 

Bell, R. 2968 amplifoius 

Benson natans, 4013 
gramineus 


Bergman, H. F. 443 gramineus, 
2484 ~~. 
E. 


Berkle 1215 crispus, 1387 
a 
Be . A. B & D2781 alpinus 


v. de reds 
Bicknell, E. P. 91 gramineus, 92 
illinoensis, 112 Oakesianus, 118 
iatus v. bupleuroides, 122 
raelongus. 
Biltmore Herbarium. 688 Rich- 
ardsonii, 4413a illinoensis, 58068 


1943} Ogden,—The Broad-leaved Species of Potamogeton 197 
et pe 5980a pulcher, 8806b ain, 935 nodosus, 938 
no gramineus, 940 Richardsonii, 941 

Bissell, C. H. 633 natans ties tml 972 praelongus, 978 


Bissell & Linder. 19678 natans, 
19692 gramineus, 19695 perfoliatus 
vy. bupleuroides. 

Blake, BS. F. 2212 a v. bu- 


ardsoni i, 3071 gra mineus 
folitus ¥. ae, ait 9472 cris- 


Biakiey; O. W. 1453 pulcher 

Blewitt, A. E. 632 amplifolius, 
1677 perfoliatus v. bupleuroides, 
1977 gression’ ‘1978 nantomatn Vv 
bupleuroides, 2137 Richardsonii, 
3651 rfoliatus v. bupleuroides, 
36 ramineus, 3657 nodosus, 
4319, 4320 perfoliatus v. bupleur- 
oides. 

Bodin, J. E. 


264 illinoensis. 
Boettcher, F. oe J. 9 pulcher. 

oivin 294 alpinus v. 
tenuifolius, 670 ‘perfoliatus v. bu- 
paced 1344 alpinus v. tenui- 

lius : ig v. subellipticus, 
2139 inus y. tenu — 2446 
Muinis v. subelli ine 


Bolander 274 illinoensie 

Bowman, 303 gramineus, 
392 Oakesianus. 

Breed, Jeffre , Loveless, 


y, Je 
Philli ps, Stauffer & Stebbins 20 


- 1783  gramineus, 
1978 alpinus v. tenuifolius. 
Bridges, T. 359 natans. 
Brien, C. 306 nodosus. 
Brinkman 73 ichard- 
sonii, 2277, 2289, daca mtine 4541 
alpinus y. tenuifo 
tain. 2987 avait v. maxi- 


us 
wn, H.E. 644 natans. 
Brown, 8. 694 ape sau 


Burgess. mage 
Boreas lder & Mu =a aches 16404 
amplifolius, 16424, 16425  Richard- 
Burkholder & Tressler. 16888 
Richardsonii. 
Burnham ; a egy 
— B.F. 389 pulcher, 112 ampli- 
folius 7 nodosus, 1318 en 
folius’ 6947 illinoensis. 
utler, B. T. 25 —— 255 
Siete nieh atk 257, 296 g neu 
Butler, G. D. 142 Richardsonii. 


illinoensis, 1030, 1031 amplifolius, 
1048 gramineus v. bare per- 
haps gramineus 1045 
“aint 1049 Richardt 1050 
praelongus, 1051, 1052 Richard- 


13 praelongus. 
337 gramineus v., 
505 gramineus. 

E. B 2 nse 
folius, “B16 1 perfoliatis v. bupleu 
oides, na , 913 sericea 

need Arar sis natans, 1033 
perfoliatus v. bupleuroides 


Chamberlain, GD. 1750, 1770 


gramineus, 2011 alpinus v. tenui- 
folius, 3123 Biaburaeion v. be eo 
oides, 9 gramin 2240 p 
longus, 2276 saeitlohancs: 
Chamberlai Collins. 616 per- 
foliatus v. bupleuroides. 
Chamberlai Knowlton. 571 
sie v. bupleuroides. 
Chamber. Morris. 11741 
Oakesian 
has . cone: 1420 nodosus, 1421 
Richardsontt; 1459 natans, 1466 
praelongus, 1477 nodosus X Rich- 
ardsonii, 07 gramineus, 1710 
amplifolius, 1713 gramineus X 
Richardsonii, 1994 nodosus X 
Richardsonii. 
Cheney, L. 8. 499 praelongus, 683 


imus, 866 Rich- 


ardsonii, 3610 
Richaeoe 4921 illinoensis. 


v. bupleuroides 
Richardson, 319 amplifolius, 328 


: 2008 Richardsonii. 
Clark, H. L. 6 0 ade x per- 
foliatus v. bupleuroide 
Clark, H. W. 5 ampliolius, 6 il- 
linoensis, 10 ge ge 
Clements, F. 305 | 2627 nodosus, 
3979 amplifolius. 
ements, F. E. & E. 8. 491 
alpinus v. ten uifoliu 
Cléonique. 72351 aenlifotion: 


198 Rhodora [May 


Clinton, oi W. 5 gramineus v. 
maximu 

Clokey, I. W. 3118 alpinus v. 
tenuifolius. 

Collins, F. 8. 300 Oakesianus, 421 
perfoliatus v. bupleuroides, 937 


amon Fernald & Pease. 401314, 
5200, 5200A, 5287 gramineus, 5295 
natans, 5314 _AgSeenins 5510 gra- 
mineus Wiyid atus v. bu- 
leuroides, ge alpinus v. subel- 
oteus nus vy. subellip- 
ticus, 6125 eounicain 


onard, H. 8. 182 natans, 318 
Richardsonii 

Cooper, 9 gramineus, 93 
amplifo gramineus v., 215 


ee 260 natans, 312 prae- 


lon 
Score & Andrews. 14 gramineus. 
Copeland, E. B. 406 ise mineus 
approaching v. maximu 
Copelan 3482 osknsiinitis ap- 
cate v. myriophyllus. 


Corbett & Williams. 11 Richard- 
sonii. 
Cory, V. L. 8502 illinoensis, 9198 


aoa 20906, 23819 illinoensis, 
243 2 nodosus, 24335, 27999 il- 
samen, 29711 nodosus. 


v. bupleuroides, 
hes gra mineus, 1254 prae- 


Coville & Applegate. 44 gramin- 
Coville & Funston. 1244, 1278 


85 gramin- 
eus, 1564 alpinus v. tenuifolius. 
ph toma & Leiberg. 224, 228 na- 


Conta. H.C. 1405 natans. 
Crandall, C.S. 2530 nodosus. 
Crickmay H. 5 Richardsonii. 


gramineus, 
imus. 
Cusick, W. C. 2484 praelongus, 
2597 natans, 3620 Richardsonii. 
Danie , F. 683 nodosus. 
n,H.T. 1311 illinoensis, 
1323, 3 & D2401, B & D2692 
Richardsonii. 


6825 gramineus  v. 


Davis, R. : 


378-W ——— 
. 38046 Rinhardsoes 


illinoensis, 49205 
linoensis, 49248 il Aegon 49 68 


ens 49314 amplifolius, 493 

gramineu illinoensis, 49347, 
49355 _ illinoensis, 3 ns, 
49363, 49369, 49372, 49389 illino- 
ensis, atans, 49394 prae- 
longus, 49395  illinoensis, 4939 
sciptitolios, 49402, oe illinoen- 
i8s:. B26 amplifolius 52334 
natans, 52341 illinoensis, 52385 


55176 nodosus, 55313 am 
55370, 55410 illinoensis, 56274 
Richardsonii, 56 illinoensis, 
56398 praelongus, 56401, 56441, 
56448 illinoensis, 56490 amplifolius, 
56498, 56501 illinoensis, 56502 
amplifolius 56524 gramineus 

illinoensis, “56534 dee if lig pl 


gramineus X inoensis, 56538, 
56539 Tiieeskis, 5 41 ans, 
66545 illinoensis, 56546 natans, 
5 » 965 oensis, 


» 56686 
linoensis, 56692, 56704, 56783 
nodosus, 57139 praelongus, 57149 
tans, 57195 illinoensis, 57259 


s. 
an & Thomas. 3465 Richard- 


elamare, M. 341 ae hime ap- 
roachin v. bupleuroide 
en °D. 7051, 7132, 17893 
yedieea 

Selm, A. W. 22 gramineus 
myriophyllus, 24 amplifolius suit 
natans, 


Dobbin, nde 860 Richardsonii, 863 


erfoliatus v. bupleuroides, 1072 
schandooull, 1112 illinoensis, 11a 
amplifolius, 1207 nodosus, 
iittaceraia, 1327 Richardsonii, 1328, 
1330 natan 


Dodge, C. K. 4 Richardsonil, | 7 il- 


Sona 8 grami — 115 natans, 

Richardsonii, prasad 
133 fig nsis ad Richardsonii, 
146 Richardsonii, 147 gramineus, 


1943] Ogden,—The Broad-leaved Species of Potamogeton 199 


153 gramineus v. maximus and Eggleston, W. W. 1 nodosus, 1655 


gramineus Spprosceo v. maxi- amplifolius, 1656 Prats y. subel- 
mus, 154 aap gramineus X il- lipticus, 2111 Oakesianus, 9974 
linoensis, TBS amineus v. maxi- ichardsonii. 

mus, 156 n es 2 171 set tn Ehlers, J. H. 533, 1756, 7957 
172 amplifolius, 4 pean neus, praelongus. 
Foes fG 1038 Richardsoni Elmer, A. D. E. 2798 oe incl 


outt, M.T. 2145, 2258 Richard- Emig, 'W. H. 224 nodos 
sonii, 2490 praclongus 3296 alpinus Empain, Rousseau & "Wavean. 


v. tenuifolius, 3323 Richardsonii. 50925 Richardsonii. 

Dowell & Painter 5385 nodosus. Evans, W. H. 780 gramineus v. 

riggs, osus. maximus, 781 natans. 

Drouet, F. 3028 nodosus. Evermann, B. W. 490 perhaps il- 

Drouet & Richards. 3309 gra- linoensis X nodosus, 492 Richard- 
mineu sonii, 493 amplifolius, 544a pemare 

Grammoud, T. 250 nodosus, 272 gramineus X illinoensis, 1032 n 
illinoensis. ans, 1057 Richardsonii, 1079 

Drushel, J. H. 6069 perfoliatus v. illinoensis, 1221 gramineus, 1222 
bupleuroides. illinoensis, 1223 amplifolius. 

Dubois, A. 660 perfoliatus v. bu- Eyerdam, W. J. 1122 praleongus, 
pleuroides. ae gramineus, 1316 gramineus X 

Dudley, W. R. 18, 19 natans, 20 141 alpinus v. tenuifolius 
illinoensis, 2030 natans, 2201 al- 2136 Richardsonii, 2360 alpinus v. 
pinus v. tenuifolius, 2202 natans. tenuifolius. 

Eames, A.J. 1498 ramineus, 1500 Farwell, O. ay 473 gramineus, 473a 
gramineus Sc sertoliat atus v. bu- gramineus Richardsonii, 505 
pleuroides, 3460 nodosus 3467 natans, 514 apne v. subellipticus, 
crispus, 9099 gramin 900 illinoensis. 

Eames, E.H. 8707 WMinceesia: 8746 Fassett, N. C. 8 gramineus v., 85, 
pulcher, elongus sp., 56 oliatus v. bupleuroides, 468 

1351  illinoensis, 11485, 11502 amplifolius, 2148 perfoliatus v. bu- 
crispus, 11 neus XX per- pleuroides, 2150, 2151 gramineus 
foliatus v. bupleuroides, 11741 x v. bupleuroides, 2152 
perhaps gramineus X liatus perfoliatus v. bupleuroides (thi 
v. bupleuroides, 11742 illinoensis, number also a linear-leaved species 

11745 gramineus maximus, according to Fernald, . Am 


11749, T1750 illinoensis, 11856 Acad. Arts & Sci. 17: pt. 1: 62 & 


crispus. 127), 3148, 4850 nodosus, 5343 al- 
Eames & Gershoy. 9101 Richard- pinus v. tenuifolius, 5362 ampli- 
ens folius, 7535 Richardsonii, 9014 


Eames & Godfrey. 8685 praelong- praelongus, 9061 Richardsonii, 
ah ‘834 alpinus v. subellipticus. 9067, 9069 gramineus, 14262 gra- 
Eam & MacD aniels. 3463 gra- ineus, v. myriophyllus, 14746 
‘ pared x ieee rena 18731  illinoensis, 18803 
ames, Randolph & Wiegand. akesi 
11175 semana i 11181 Richard- Fassett & “Wilson. 4349 nodosus. 
sonii (perhaps perfoliatus v. bu- Fellows, D. W. 2006 gp pub 
pleuroides X Richardsonii). 2007 7 Richardson 2966, 4679 p 
Eames & Thomas. 3459 nodosus. foliat v. buple uroides, 43s 
Eames & Wiegand. 9100 Richard- Hichardsoni 5656 gramineus X 
sonii, 9102 perfoliatus v. bupleur- ie 
oides, 11162 natan ns, 11172 a mpli- Fendler, A. 132 amplifolius, 837, 


ns 
xe 
a7 


folius, 11173 oe 1117 839 nodosus 
Richardsonii (perhaps perfoliatus Fern ald, M. tL. 116 gramineus ap- 
upleuroides Rishardeonil), proaching ¥. maximus, 117 alpinus 
14535 crispus. v. tenui ror 436 gramineus v. 
Eaton, A. A. 335 illinoensis. maximus, ramineus per- 
Edwards & Clausen. 1400 perfoli- een Vv. ae euroides, pase: na- 


atus v. bupleuroides. tans, 477 gramineus v. maximus, 


200 
509 Oakesianus, 750 pulcher, 778 
amineus x perfoli- 


rnal 
natans, 23131 Oakesianus, aril 
eo v. nec pa 
rnald, Bartram ng & Fas- 
pins 23139 fe Pa 
Fernald, Bartram, Long St. 
John. 67 ~Oakesianus 6769 
perfoliatus . Be 


Fernald, Bis: "raven, Long & 
Linder. 19689 amplifolius 


Fernald & Collins. 327 alpinus v. 
subellipticus, 965 alpinus v. tenui- 
folius. 


ernald, Dodge & Smith. 25421 

beens 25422, 25423, 25424 
praelon 

F ernald & Fogg. 505, 747 Oakesi- 


Fer maaid. Gilbert & Hotchkiss. 
27346 praelongus. 

Fernald & Griscom. 4295 pulcher. 

Fernald, Griscom & Long. 4535 


toe er. 
rnald, Hunnewell & Lon ~~ 
8443 pulcher, S445 perfoliatus v 
bupleuroides 
Fe rnald & Jackson. 11986 natans, 
11987 alpinus v. tenuifolius, 11988 
11989 Richardsonii 


—— & Linder. 19682 Oakesi- 

anu 

F ernaid & Lon 1210 ae 
5977 puller, *: 747 cri , 8440 

844 


alpinus v. eubelli jae 
12388 alpinus v. tenuifolius, 12389 
nodosus, 12391, 12392 perfoliatus 
v. bupleuro oides, aa pulcher, 
15967 perfoliatus bupleuroides, 
17805 Oakesianus, 17807 perfolia- 
tus v. bu nat roide - 19677 natans, 
19687 subellipticus, 
19688 aaiiyiinething "19690 gramine- 


Rhodora 


[May 


bupleuroides, 

, 27345 gramineus, 
27347 gi nem 

Fernald, Long & Dunbar. 26217 
ove ate _— 18 polygonifolius, 


26221 alpinus v. lipti ou 
26222 soca gram x_ per 
foliatus v. bt Fear tats 26223 
a. : 
Fernald, Long & Fogg. 1207 alpi- 


nus v. ienuifolius 1208 Oakesianus, 
1209 gramineus v. maximus, 
praelongus, “1212 perfoliatus v. bu- 
PY canst 
Fernald, to ong & Nort 12382 
natans, 12383, 12384 F Oalcesianus 
Fe d, Lon g & St. Joh 6763 
natans, 6765 A locke 67 
alpinus v. subellipticus, 6767, 6768 
praelongus, 6770, 6771 perfoliatus 
v. bupleuroides. 


Fernald, Long & Torrey. 8444 


pulcher. : 
ern & Pease. 3066 Oakesi- 
anus, 16957, 17056 gramineus, 
19676 natans 


- 10894 natans. 
m 25420 alpinus 
bas. subellipticus. 

riguae a oe 477 gramineus 


Fernald & ‘Svenson: 744 gramine- 
us v. myriophyllus, es natans, 748 
Oakesianus, 749 praelongus, 1 
perfoliatus v. bupleuroides, 

Fe _— & White. 19680 Oakesi- 


& Wiegand. 2435 natans, 
2436 Oakesianus, 2443 alpinus 4a 
a approaching v. sube 


2444 praelongus, 


4478, 4479 gramineus, 4483, 4484 
4485 perfoliatus v. bupleuroides, 
14533 at Wiecas 
Fernald, Wiegand & Bartram. 
4463 Cui vented & 
Fernald, Wiegand, Bartram 
Darlington 4477 gramineus V: 


maximus. 
Fernald, Wiegand & Darlingto”- 


1943] 


4461, 4462 natans, 4464, 4465 
ae 4474 alpinus v. bees 


folius, 4475 ~ ear ot v. maximus, 
476. gramineus, 4480 pets v. 
maxi per dese 4482 


ngus. 

Fernald, hide gros & 

us V. pre pe 14084, 
14085 cnc, mplifoli- 
us, 14087 Ri acuont = P4088 
Richardsonii and oliatus v. 
bupleuroides, 14089 er “nies 

ere Wiegand & Kittredge 


7 gramineus, 24 
foliatus v. oe ides 

Fernald, ela and & poe 27348 
perfoliat 

Fernald, Wieser Long, Gilbert 
af ei chkiss. 
vides (probably backcrossed with 
gramineus) 
tris, Roxana S. 2043 natans, 


Fiker, C. B. 1455 natans. 


Fink, B. 191 illinoensis. 
Fitch, A. 7778 natans. 
Fletcher. 2969 amplifolius, 3025, 


3028 natans, 3044 Richardsonii. 
g 710, 1840, 2077, 2997, 

3505 pulcher, 3606 perfoliatus VY. 
bupleuroides, 3867 akesianus, 
4504 pulcher, 4934 amplifolius, 
4935 ad ig 6794 nodosus, 
12254 crispus 

Forbes, F. F. 156 eerie v. 
bupleuroides and gramineus 


perfoliatus v. ay gee ide 

oster, A. S. 866 natans, 1992 

amplifolius 
Fredholm, A. 6183 illinoensis. 
Fulton, H. J. 970 s 
Garrett, A.O. 529 alpinus v. tenui- 


folius approaching v. subellipticus, 
3958 nodosus. 

Gates, F.C. 258 oe 261 
praelongus, 1752.2 n 0 
a 10644 ‘libeoeseaks, 12217 


Gauthier, R. 100 amplifolius, 135 
rfoliatus v. bupleuroides, 242 
amplifolius. 


Gillman, H. 40 Ri gee oap hn (per- 
sp.), 65 


o . ° 
661 amplifolius, 562 ‘Biskaniaont. 


Ogden,—The Broad-leaved Species of Potamogeton 


201 


Gleason, H. A. & H. A. ad 166 
ye 302, 310 gramine 


Gleason & Shobe. 176 seuplitetis: 
179 natans 
Glendenning, R. 92616, 92617 


gramineus, 92618 Richardsonii 
oldman, E. A 


oodale, Mar ciet iper 
76038, 76945, Ted natans, 76948, 
951 Oakesianus, 76973 gramine- 
a 96988 : 


rrioeeiats Piehs hematin x 
illinoensis). 
i 207 Richard- 


a 33, 677 Richard- 
sonii, 703 illinoensis, 781 Richard- 


am, E. H. 9821 nodosus. 


3207 illinoensis, 
3275 amplifolis, 3276 gramineus. 
Linder. 19681 Oakesi- 


s v. bupleuroides (perhaps back- 

crossed wit ineus), 19693 

perfoliatus v. bupleuro oides. 
illinoensis 


crime, D. ‘ 6 Richardsonii. 
oe & Schlosser. 4 ampli- 


, E. J. 594 nodosus, 3255 
10679 am- 


P0685. i 10686 gram 
486 natans, 487 (this num- 


ii. 

andler. 563 alpinus v. 

tenuifolius. : 

amilton. 64010 alpinus v. tenui- 
folius. 

327 nodosus, 367, 


Hanes, C. R. 
Sti, 397, 4 477 gramineus X sp., 
r 


amineus x 
757 eee 
re dru 


497 illinoensis, § 

sp., 517 illinoensis, 

928 Oakesianus an 

Nuttallii, oe 1988 n 
I. 


er, 4 sae aoe = “maxi- 


202 


mus (perhaps ath <_ alpi- 
nus), 98, 141 Ri gt vl 
er, R. M. Seat (not 
typical), bie 2088 ty ulcher 
Harri , Ww. H. 2441 perfolia- 


tus em pip barciies, 99085 gramin- 
eus, 0. gramineus v. are 
pil ees perfoliatus ve bupleu 

oides, 99089 ardsonii, 


Harris, S. K. 539 praelongus, 540 
perfoliatus v. bupleuroides 
Harrison, A 


m, A. e365 natans, 16 
eam 
weg, T. 2016 nodosus (this 


number also a linear-leaved s preg 
ernald, Mem. Am. 
. 1:66 & 


oe ; 50 Richardsonii. 
U. 280° gramineus, 4131 


n da. 751 Richardsonii, 

752 nodosus, 804 Richardsonii, 806 

nodosus, 807 illinoensis, 811 Rich- 

mineus, 823 natans, 

10 13 Richardsonii, 

10114, 10115, 10116, 10117, 10123, 
dosus. 


Haydon, W. 254n 
Heller, A. A. 939. nae 5824 
osus. 


FE 
=] 
es 
ja 


-» ¥. 1007 natans, 
1009 Richardsonii, 2473 illinoensis, 
2474 amplifolius, 2475 eben, 
ae oe My 2ekeon 


781 gramineus, 


ermann, °F. J. § 
6281 praelongus, 7153, 7224 Rich- 


ardsonii, gramineus 
praelongus, 8286 gramineus, 8647 
crispus, 9294 gramineus < illinoen- 
sis, 9383, 9728 n us 

Herriot, W. 78019 78020 


a 
»E.5. 46. 54 pulshan | 92. 1888 
alpinus v. baliidenn 133.1881 
ichardsonii, 
159.1909 nodosus 
162.1900 __ illinoensis, 171, 1900, 
179.1901, 191.1902 nodosus x 
ichardsonii. 


, 


Rhodora 


[May 
Hitchcock, A. E. 260 gramineus, 


us. 

1000, 1099 am- 
plifolius, 12358 natans, 15655 no- 
dosus. 

Hitchcock, R. 11167, 11168 nodo- 
sus, 11175a illinoensis. 

Ho on, A. R. 11 perfoliatus v. 
bupleuroides, 530 Batis 40 
natans, 2647 perfoliatus v. bu- 
pleuroides, 2652 amplifolius. 

Hodgdon & Heale 2977 natans. 

Hodgdon, a es & Har- 

oO 


folius approa pening 3 v. subellipticus, 
2646 hapa tone matte 31 fine ineus 
Melia gram 

inus ‘“ “subelli ip 


yee weit ii, 4033 il- 
eben 4058 H. Ehonianue, 4059 


gramineus, 4060 praelongus, 4083 
ramineus, 4109 nodosus, 4110 
natans, 4112 gramineus, 4133 il- 
linoensis. : 
Hot schias & Koehler. 4179 il- 
linoensis, 4192 amplifolits, 4193 
natans, gr s, 4222, 
4223, 4226, 4227 illinoensis, 4228 
elongus, 4245 i nsis, 4 


307 gramineus, 4326, 4331 illino- 
ensis, 4340 praelongus, 4349 alpi- 
a v. subellipticus, 4353 a mpli- 

olius. 

Hotchkiss & Martin. 4432 illino- 
ensis, 4460 alpinus v. subellipticus. 
ouse, H. D nodosus, 138 

a 


15193 gramineus, 

19852, 20002 nodosus, 21752 am- 

plifolius, 21774, 22044 nodosus, 

23174 gramineus. 
owe & _ Lang. 768 natans, 1040 
gramin 

Howell, I ‘T. 7679 Richardsonii- 


chen danty 1498 
lifolius, 1668 gram 
Huitén, EB. 7573 sane. ee " tenui- 
folius. 


1943] 


Hylan, D. R. hg natans. 
Innes & Moon. 1093 nodosus. 
T. 52 gramineus 


Jepson, W.L. 147 alpinus v. tenui- 
folius’ 238 Sidisssen at 

Jes i. gramineus ap- 
proaching v. myriophyllu llus 

Jo H. N. 2719 ‘nodosus, 
D795 5 perfolatus Vv. cr at 

Johnston, I. M. 600 ilicianate, 

cee G. N. 3511 illi ecko eae 
— hi tenuifolius, 5227, 7851 
gram 

Jones, M. z. 

1304 gramineus _ v. 


Richardsonii, aelongus, 
606 alpinus v. taniiitelin: 9293, 
9295 gramineus, 9299 Richardsonii. 
Jones, 465 illinoensis, 7470, 
7471 crispus. 
Jones, W. W. 432 nodosus. 
— & Hoffman. 7467 Richard- 


J caval '& Tene 7 7462 illinoensis. 


Kearn T 74 nodosus, 1626 
pule nf 

Keck, D.D. 1188 nata 

Keck & Stevens 280. jllinoensis, 


323 Richardson, 332 amplifolius, 
335 eri 

Keck & “Stilwill. 368 natans, 373 
gramineus, 377 amplifolius, a 
gramineus, 401 nodosus, 
tans, 412 praelongus, 428, 43 
crispus, 433 amplifolius, 454 gra- 
mineus, 465 praelongus, 458 Rich- 
ardsonii. 

Kellogg & Harford. 949 illinoensis. 

Kelsey & Jor 8 gramineus, 


, oldsborough & Doo- 
12 amplifolius, 17 prae- 

gramineus, 111 perfoli- 
atus v. bupleuroides (perhaps per- 
cranes v. bupleuroides XX 
ar 

Kennedy, Rac hel B. 78 gramineus 
x perfoliatus v. rig oaetion 80 
a ae 81 alpinus tenuifolius, 

5 gramineus, 478 a aiseyete-9 

v. ’ bupleuroide es, 543 vores 
551 — v. pe age 

Keno oe © ‘itinobabla 
( cathians ott ty  illinoensis), 
139 praelongus. 
illip, E. P. 931 amplifolius, 6203 
Richardesall: 6204 natans, 6896 


Ogden,—The Broad-leaved Species of Potamogeton 


203 


pulcher, 12258 natans, 12261 am- 
lifolius, 12264 illinoensis x _ per- 
oliatus v. bupleuroides, 
Richardsonii (perhaps 
x Richardsonii), 
mi 


u 
= amplifolius, 1253 
12574 Oakesianus and epihyd 
Nuttallii, 12605 alpinus v. subel- 
lipticus and aephitalias us, 12610 am- 
plifolius, 13379 Oakesianus, 30845 
pulcher. 
Kimball, R. H. 70 Oakesianus. 
dle, E. M. vee 93572 gra- 
mineus, 93684 na 
Knowlton, F. O88 n 
Knowlton & Weatherby. * 6632 
Oakesianus. 
Kreager, F.O. 441 amplifolius. 
Krotkov, P. V. 5142 alpinus v. 
tenuifolius approaching v. subel- 
lipticus and epihydrus v. Nuttallii, 
caer Syren 5145 gramineus 
5147 natans, 5150 praelongus, 
7027 eg ee 7033, 7034, 7035, 
7036 gramineus, 7038 natans, 7041 
Richardson, "9629 amplifolius, 


Kubichek, W. F. 7 natans, 13 gra- 


eus, 109 R 
115  illinoensis, 
and natans, 120 amplifolius, 121, 


1 
nodosus, 172 gra ramineus, 173, 184 
illinoensis, 190 natans, 192 gra- 
mineus, 196 illinoensis. 


untze, Herbarium of Otto. 3105 

perh geamineus x illinoensis. 
Laing, pS M. 147 Richardsonii. 
Lake, 614 gramineus. 


Lansin Ei. 
ea illinoensis. “74 natans. 
C. 50 nodosus, 53 
"Hichasdeonit 
Leiberg, J. a 751 gramineus, 1574 
foc 


nodosus, 
‘63 pices v. bupleuroides. 
nard & Mannakee am- 
lifolius. 
Lewi 


wis, H. . 180388  natan 
130390, 130391, 130393 alpinus v. 


204 


tenuifolius, 130394, 130395, 130396 
gramineus, 7 erfoliatus. 

Lindheimer, F. 116, 311, 393 
poker 513, 547 illinoensis, 1234 


nodos 
‘inedals & Keck. 1 gramineus, 54 
amplifolius, 59 illinoensis, 115 = 
en oO ange 127 natans, 153 pra 


Sean , G. A. E. 937 natans, 1012 
19679 ae aoe or 
H. & H 1501 


tan 

ouis-Marie, 313 “alpinus Vv. 
air er 1255 nodosu 

aporte & Dude- 
sry 1 as eg « x perfoli- 
atus v. bupleuroides, 302 gramin- 
u ichardsonii 1403 gra- 

e ; 


: pie 
oides X Richardsonii), 10843 gra- 
—— x perfoliatus v. bupleur. 


Lun 150 I 
Mabbos, D.C. 1, 252, 317, 334, 
350, 390 Richardesrti, 459 natans, 
Richardsonii. 
feos ot uk L. 977 nodosus, 1060 
pe v. bupleuroides, 1707 
inoensis, 2279, 2337 psa Ral 
ab n us, 3953 pus 74, 
neus, 3078 natans, 3082 
olen 3201a, 3205 Oakesianus, 
2409 illinoensis, 3420 gramineus 
McCabe, T. T = alpinus v. tenui- 
folius, 47 nata 
cCalla, W. C. "2370 rp mbna 
aniels, L.H. 3464 illinoensis. 
al, Me ae MS Sidaad. 
amineus ail SP gen v. 
atans, 241 gramin- 
eus, 302 Richardson 303 ampli- 
folius, 304 a us X illinoensis, 
462 natans, 543 nodosus, 57 
alpinus v. tenuiolus, 638 gramine- 
ing Vv. maximus, 639 
natans, 805, 956 


oa 3 illinoensis, 
359 pulcher, 2305 acapliigitae 3613 
gramineus v. maximus, 4377 na- 
tans 


Rhodora 


[May 


sages & Griscom. 100435 na- 
044 Oakesianus, 10045 am- 

pifelius, 10047 alpinus v. subellip- 
ticus, 10048 cna 10048a 
gramineus approaching v. maxi- 
sate 10049 pertoliatis v. bupleur- 


McLouth, C.D. 4 illinoensis. 
MacMillan & Sheldon. 488 gra- 
mineus, 571 ar oniaggaat 1332 
gramineus v. maxim 
McMurphy, J. 192 cnodoeee:. 193 
an 


natans 

Macoun 5 graminens 
2979 gr. x perfoliatus v 
bupleuroides, 2980, 2984, 2 92 
gramineus, 3020, 3021, 3022, 3026 
natans, ichardsonii, 3047 
Richardsonii (perhaps Richard- 
sonii 3049, 3052, 3054, 


p.), 9, 3052, 
3055, 3056 Richardsonii, 4162, 4165 
alpinus v. tenuifolius, 4166 alpinus 
v. subellipticus, 4178" gramineus X 
p natans ae gramineus 
v. petra 4368, 381 Richard- 
sonii, 26814 se a 36815 illino- 
ensis (perhaps a hybrid), 26816 
gramineus X Richardsonii, 26817 
Richardsonii, 80929, 18 gra- 
86002 am plifolius. 
Macoun, 9 pra 
longus, 14 illinoensia (this fa 
also a linear-leaved species accord- 
ing to Ferna em. Am. Acad. 
Arts & Sci. 17: pt. 1: 55 & 131), 94 
alp inus v. tents. 95 Richard- 
mineus 


achin aximus, 1730 am- 
plifotiur, 1733 der rryg yt approach- 
, 1740 Ri chardsonii 


pleuroides, 2991 u : 
akesianus, 2995, 2996 illinoensis, 
3019 tate. 3023 illinoensis, 3024, 
3 natans, 30 erfoliatus V- 
bupleuroides, 3048 Richardsonii, 
3053, 3057 Richardsonii, ’ 
4130, 4132, 4132a praelongus, 4163; 
41 65, 4167, 4167a alpinus v. te nui- 
folius, 4168, 4169 alpinus v. subel- 
lipticus, 4177 illinoensis, 4179 gt@ 
mineus < Richardsonii, 4180 ‘llino- 
ensis, 4357 natans, 4360 gramineus 


1943] Ogden,—The Broad-leaved Species of Potamogeton 205 


maximus >X_ nodosus, —- 118249 gore v. bupleuroides, 


ee 4365 alpinus v. tenu 118250, 118251, 118252 Richard- 
folius, 4379, 4880, 4382 Richard sonii, 118257 gramineus, 118258 
as ” 3006 ‘al inus v. tenuifolius, perhaps gramineus _illinoensis, 
1644 aa 16458, 16459, 118259, 118260, 118261, 118262 
16460, 16461 Ri chardsonii, 20748, perhaps alpinus gramineus, 
20749 Oakesianus, 20751 alpinus v. 118263, 118264 gramineus v. maxi- 
subellipticus approaching v. tenui- u 826 oman 118266 
folius, 20756 perfoliatus v. - gramin Vv. ximu 267, 
pleuroides, 22176 Richardsonii, 118268 eester Evy 118269, 118270 
22177 nodosus, 22211, 22212 na- ee, 118273, 118274, 118275 
tans, 22216, 2 gramineus, 

22220 perhaps alpinus < gramine- Malte ir Watson. 960, 1276, 1818 
us, 22221 amplifolius, 22227 per- sacra i 


foliatus v. bupleuroides, 22228, Markert, W. C. 76909 perfoliatus 

22229 amplifolius, 23173, 23175 v. puplearoies. 

rte gate 23180 gramineus, Martin, A.C. 158 natans 

26824, 26826 pence em 26830 Martindale, 1 12003 ampli- 

crispus, 26832, 26833 Richardsonii, foliu 

26834 amplifolius, 26839 nodosus, Mathias, Mildred E. 265 natans. 
natans, 26841, 62 O15, 62016 Maxon, William R. 4617 Richard- 

nodosus, 62021 illinoensis, 62! sonii. 

natans, 62023, 62024 gramineus, Mearns, E. A. 805 nodosus. 

68425 natans, 68919, 78320, 78321 Merrill & Wilcox. 863 Richard- 


alpinus v. tenuifoliu 8, 85530 na- sonii, 902 gramineus. 
tans, 85531 annie alpinus < Mertie, J.B. 66 yg ose 
gramineus, 85534 amplifotius, Metc | oe pein Bi 
85535, 85536 gramineus v. max Richardsonii, 14 aelon 
imus, 85537 Richardsonii, 85550 natans, 193, 304, 329, 339 Michard. 
nodosus, 88248, 88249, 88250, sonii, : gramineus, 404 
88251 natans, 88252 amplifolius, Richardson i, 423 gramineus, 432, 
8825: i ximus, 451 Ri “wie Berd 471 gramineus, 
88254, 88255 oe aia — 512 Richardsonii, 5 raelongus, 
gramineus approaching 542 Richardsonii, 544 natans, 546 
us, 88257, 88258, $8259 pe nage See 547 Richardsonii, 636, 
longus. 819, 845 nodosus, 888, 9 
Macoun & orageeate De es re snide, ~ crispus, 1018, 1024 
anise 76869 ¢g nodosus, 10 inoensis X nodo- 
s, 76870, 7687 L 76872 Richard- one 1071, 1105 nodosus, 1139 na- 
snd. tans, — amplifolius, 1145 Rich- 
Maguire, B. 472 alpinus v. tenui- ardso 1147 praelongus, 1160 


pra 
folius, alpinus v. subellipticus, am lifolius, 1162 gramineus, 1176 
74 alpinus v. tenuifolius, 483, 484 Ric ardsonii, 1229 praelongus, 
barton 485, 486 praelongus, 1291 pulcher, 1295 natans, 1297, 


488, 490, 491, 495, es 497 Rich- 1298, 1329, 1370 illinoensis, 1373 
ardsonii, 13149 gramineus v. maxi- gramineus, " _illinoensis, 
, 16201 Richardeonii, 16220 1388 natans, 1390 Richardsonii, 
gramineus v. maximus, 1636 Rich- 1391 illinoensis, 1412 gramineus, 
gle ony, 1413 praelongus, 1415 gramineus 
aguire & Pir 5439 na- v., 1420, 1427 illinoensis, 1428 gra- 
tans, 5440 Biber new) 5442 alpi- mineus, 1432 illinoensis, 1463 Rich- 
nus v. ten uifoli ius. ardsonii, gramineus, 1470 il- 
aguire, ian = = linoensis, 1471 nodosus, 1472 gra- 
12771 gramineus v. m mineus, 1511 amplifolius, 1520 
aguire & Richards. 13136 alpi- gramineus, 1532 gramineus il- 
nus v. " tenaaitciiag linoensis, 1542, 1548, 1551, 1563, 
Maguire, Richards, Maguire & 1564 illinoensis, 1780, 1794, 1839, 
Hammond. 4367 illinoensis. 1908, 1937 Ric hardsonii, 1962 na- 


Malte, M. O. 118247, 118248, tans, 1963, 1979, 1991 Richard- 


206 


sonii, 2014 illinoensis, 2024 ampli- 
folius, 2039 iRingeua, 2044 Rich- 
ardsonii, 2046 n sus, 2048 illino- 
ensis, 2049 natans, 2050 oe ineus, 
2055 illinoensis, 2065 natans, 2076 


illinoensis, 2093 praelongus, 2105 
gramineus, 2107 i ensis, 2113 
natans, hardsonii, 2132 


gramineus, 2201 natans, 2209, 2210 
amplifolius, 2211 natans, 2226 
yee as 2 2242, 2250 ampli- 


2321 

2341 Sinnanaa, 2342 natans, 2345 
illinoensis, 2352 nodosus, 
apelin a fea 2370 sm 


2371 praelo eus 
vs 2 Ae et 5377, 2 79 {llinoensis, 
2381, 2387 ce 2388, 2396 
illinoensi 

Metcalf & Sperry. 1603 perfolia- 


tus v. bupleuroides, 1621 ampli- 


4 gramineus Eee 
ros peer 1690 il wien 


Michel. oliatus v. bu- 

pewoids 1993 amplifolius. 
29 amineus. 
Sotaks, i 457 nodosus. 

Motte W.S. 294 amplifolius. 

Mold 3 - 752 illinoensis, 
4207 gramineus v. maximus, 4208, 
9396 Ae 9397 ane i v. 
a oiags tos 

Moo A. H. 89 natans, 945 
ae 2488 amplifolius, 5022 
gramineus, 5036 amplifo 

Moore, E. 1488 natans. 

+ ,G. 81 amplifolius 


oore & Steyermark. 3667 il- 

linoensis. 

rris, E. L. A41 natans, _ 
nodos 


perhaps 
saber 8 Hinoensi x nodosus). 
Mosier, Piety noensis. 
oyle, J. B Richardsonii, 894 
alpinus v. vaalaiion 2052 illino- 
Hee amp ifolius. 
, W. C. 2739 crispus, 
7632 ac, 7633, 7634, 7635 am- 


Rhodora 


[May 


eee fo TY approach- 
llus, 7642 gramin- 

eus, 7643, 76 

longus, 7653, 7654, 

Piette 7657a, 765 

foliu , 10202 natans, 10204 Ri 


gra  illinoensis, 10209, 
10210, 10211 gramineus, 10212 
a ts ensis, 10216 
gramineu ichardsonii, 
17098 umennia 71 , 17106 gra- 

By Ltd tichardsonii, 17545 
illinoensis, 17550 gramineus, 17556 
praelongus, 17671 natans, 17676 


sus, 434 gramineus, 459 Richard- 


cher & Burkholder. 16401 
is, 16846 crispus, 16885 


Richardsonii. 
Clausen. 3751, 
illinoensis, 


SS 
CH ON 
& 


Sparing) 3770 
772 gramineus (perhap 


eras x Richardsonii, 4161, 

4164 amplifolius, 4167 opevfolintus 
v. bupleuroides, 4190 gramineus, 
4205 tigen 7 4209 Richardsonit 


(perhap: rfoliatus v. bupl 
oides x ichardsonii, 4210 ser 
foliat v. leuroides, 4232, 


Muenscher & Curtis 4827, 4828 


5 
plifolite, ata le 
ge 4876 gra. 
Vv. oe 4877, 
ie raelongus, 49 
perhaps gramineus X perfoliatus 


& Lindsey. 2712 al- 
pinus v. tenuifolius, 27 7 ps 

ineus a nodosus, 2 nodo- 
23, 2724 ae lifolius, 


atus v. bupleuroides, 2739 ones 
2769 gramineus v. maximus, 7 


1943] 


gramineus >_perfoliatus v. bu- 
pleuroides, 2775 gramineus Vv. myri- 
oP on 2777 gr stig ice a 

proac v. maxim 2780 gra- 
mineus approaching * myriophyl- 
lus, 2782 gramineus, 2785 gramin- 


atus v. bu- 


713, 715 


perfo 


Ipinus v. tenuifolius, 729 Richard- 


esata ee Bae 


608, 1609 perf 

bupleuroides, 1618 crispus, 1676 
illinoensis, 16 gramineus, 
1680 ilinoensis, 1683 gramineus, 
1684 gramineus illinoensis, _ 
1689, 1690 gramineus, 1691 g 
mineus approaching v maximus, 
oe 1693 g santiees Vv. maximus, 
1 


approaching v. maximus, 1699 gra- 
m s, , 1701 illinoensis, 
1703, 1705, 1706, 1709, 1710 na- 
tans, 1711, a ; 16.041 
Oakesianus, 7, 1748 praelongus, 
1751 illinoensis, 1753, 1754, 1756 
praelongus, 1773, 1775, 1776, 1779, 
1781, 1782 Richardsonii, este 
haps perfoliatus v. bupleuro 
Richardsonii, 1787 ichardaeall, 
1806 em ensis. 

Muensc 


g & Maguire. 

67, 71 Yamplifolius, 73, 75 gramineus 
illinoensis, 120 gramineus ap- 
mee v. sraeset 141 prae- 
157 perhaps perfoliatus v. 

ri bea bic x Richardsonii. 


Ogden,—The Broad-leaved Species of Potamogeton 


10] & . 
v. bupleuroides, — 
Ogden n. 


207 
Muenscher & Wiegand. 14538 
perhaps gramineus X s 
unz, P. A. 2785 crispus, 10805 
illinoensis. 
Murdoch, J. 
pleuroides, 939 amplifolius, 
Oakesianu 
Nash, G. v. "786 crispus, 859, 1750 


524 perfoliatus v. bu- 
2062 


illin sis. 

Nelson, A. 2276 oF perione, 2406 
gramineus v., 4145 n 

Nelson, A. S Vv. 
maximus, 6 atans eoubsarsa 
form, perhaps a hybrid), 6771 per- 
haps gramineus X illinoensis, 6807 
Richardsonii. 

fap ecole . & Ruth A. 973 prae- 
on, 

Ne ; 87, 3362 Richardsonii, 
3386 illinoensis. 

Nelson, Cc 


eel yk gr amineus 


Ogden, na Cc. 914, O64 eel EN 
us, 1502 crispus, 1620 na- 
tan ns, 1701 perfoliatus v. bupleur- 
oides, 1702 alpinus v. tenuifolius, 
1704 amplifolius, 1705 natans, 1716 
gramineus, 1717 Richardsonii, 1718 
natans, ee — v. tenuifolius, 


akesi: 
Ogden & Babel. "2166 Oakesianus, 


g = 


2203 gramineus 
Ogden, Babe Chamberlain. 
2242 alpinus v. tenuifolius. 
Ogden, Babel & Kozicky. 1880 
praelongus. 

e & Bolan Bors 1562 
Oakesianus, 1565. 1568 
rede bige o llmoensis, 1571 . 
foliatus v. bupleuroid - (perhaps 


v. bupleuroides xX 


n 
a lifolius, — 1584 1589 


gramineus X 
1537 = tus 


& Marsto 432 go oli- 


208 Rhodora [May 


atus v. bupleuroides, 492 prae- 
orn 493 perfoliatus v. bupleur- 
oides, 1691 gramineus, 1692, 1693 


Og ee Aa i 1762 
gramineus X illinoensis, 1765 gra- 
Ss Vv. my phyll 5 8 per 


ra 

8, amplifolius, 2020 

praelongus, 2032 perfoliatus v. bu- 

pleuroides 

ee Ogden & Babel. 2271 na- 

ans, 2272 ee 2294 Rich- 

haere 2308 gramineus 

Ogden, naire de Stetnmnete: 1882 
praelo 

Ogden & Paine. 1505 A eas tration 
v. tp ug 1506 n 

Ogden, Rollins & W Wiegins. 1731 
Berchtoldi > _ perfoliatus v. a 
shu, 1732 patfotinges v. bu- 
pleuroi 

Ogden & Steinmetz. 
1543 gramineus v. maxim 
perfoliatus v. biplentoiden a 1544 yey 
gramineus ™X_ perfoliatus v. bu- 
pleuroides, 1545 gramineus, 1547 
amplifolius, 1548 natans, 1549 — 
mineus, 1552 amineus v. i- 
mus, 1556 amplifolius, 1592 ‘prae- 
longus, 1593, 1602, 1603 gramineus 
Vv. maximus, 1604 - 

cheese =F v. bupleu 


1542 Hager at 


Ogden, sige gh & Prince. 1596 
nodosus, 1597 gramineus v. maxi- 


m 
Ogden & Trask. 2073 Oakesianus. 
Ba 978 & Wiggins. 1729 ampli- 


perfoliatus — v. bupleu 
( Seige alpinus X grami ass, 
z inus v. tenuis, 2344 
gramin 2 Vv. maxi 
Oosting, H. J. 391 gramineus, 302 
pr aipreepe oe 2945 illin 
, 2971 Richardsonii, 28100 gra 


n- 


mineus, 28164 —  illino- 
neus. 


“ 553 Richardsonii. 
E. 


Osterhout, G 2885 alpinus v. 
tenuifolius. 
, IT. C. 1565. natans, 1583 


Richardsonii, 1584 gramineus, 1678 
Richardsonii, 1 praelongus, 
1768, 1769 amplifolius. 
ver, W. H. 3366, 3367 Richard- 
sonii, 4008 illinoensis, 4044 gra 
mineus, re alpinus v. tenui- 
oaching v. subellipticus, 
13818 Roheniand, 14459 ampli- 


1 
171 re a 17141, 
mean 17432 nodosus. 
alm E. J. 11952 nodosus, 
12217. illinoensis, 12907 perhaps 
rege x nodosus, 21526 n 

i: — Tadehes 33707, 43586 


Palmer, 5: L. 36, 37, 38 natans, 50 
a ie 

Palm oo J. 638 gramineus V. 

dai Richardsonii, 1862 

Richardeoati 1866 gramineus. 

Palmer & Steyermark. 41417 
pulcher. 

Pammel, L. H. 
illinoensis. 
arish, S. B. 2106, 2128, 3350 


4°] 


100 natans, 776 


n 
Parish, S. B. & W. F. 1435 natans. 
1073 perfoliatus v- 


ides. 
Parlin & Fernald. 924 natans. 
174 illinoensis. 
Payson, E. B. & Lois B. 2023 


ov aad ig 2024 =praelongus, 
2251 na 

Pease, A. ie 1919 alpinus v. subel- 
lipticus, 2011 pulcher, 2064 crispus, 
2500 amineus fine — eus V- 
maximus, 2907 alpi v. tenui- 
folius, 3795 ak 4699A Oakesi- 
anus, 14 ramineus, 11982 
Richardsonii, 12143 perfoliatus V 


bupleuroides, 12171 Oakesianus, 
138 


amine amplifoli- 
us, 14012 praelongus, 14582 gra- 
—— 14585 amplifolius, 16591, 


16943 perfoliatus v. bupleuroides, 


1943] 


17024 alpinus v. tenuifolius_ap- 


i ng v. subellipticus, 
17983 Richardsonii, 20073 natans, 
22752, 227538, 227 gramineus, 

25217 natans, 26709 alpinus v. 


Polio ag. 
Bean. 26092 gramineus, 
mpli 


Richardsonii, 26199 am 
folius, 2624 ichardsonii, 26304 
gramineus, 26389 ardsonii 


Pease & Edgerton. 27169 gramin- 


eus. 
Pe wore & Fernald. 16958 amplifoli- 
17024 alpinus v. tenuifolius ap- 
prosching v. subellipticus, 17047 


at air ane perfoli- 


as 24828 Richard- 
sonii, 2491 Seer: 24918 
Richardsonii, 24964 _ illinoensis, 
— eg atans, 25013, 25031 Rich- 
ardsonii, 25032 amplifolius, 25135 
Oakesiabus, 25167 amplifolius. 
Peattie C. 2385 “Richardsonii, 
2305 gramineus v. myriophyllu 
Peck, C. H. perhaps ce te D4 
nodosus, 3 se nsis. 
Peck, M. E. 6284 Richardsonii, 
8514 natans, 900) ichardsonii, 
9026 dmaptifolias 9619 Richard- 


sonii. 

Peebles, R. H. 14190 nodos 

Pennell, F. W. 3351 oetitiation Vv. 
bupleuroides, 16220 Richardsonii, 


Pepoon, 5 

899 praelongus 

erry & Roscoe. 38 Oakesianus. 
ay O. P. 1091 nodosus, 1092, 


655 perfoliatus v. bupleuroides, 
1883 nodosus. 


Piete A. J. 3 Richardsonii, 6 
pis 7 illinoensis. 
Piper, C. V 57 Richardsonii, 758 


natans, 3684 amplifolius, 3761 gra- 
mineus - illinoensis, 3765 natans, 
4426 alpinus v. tenuifolius, 4430 
Rishardarats 4431 _— ap- 
proaching v. maxim 

Polunin, N. 1976, 1977, 2062 “en 
nus v. tenuifolius, 2070 per 


lpinus X gramineus. 
Pontious & Bartley. 18 nodosus. 
Porsi 4295 precines,. ap- 


proaching v. maximus (perhaps 


Ogden,—The Broad-leaved Species of Potamogeton 


209 


gramineus X sp.), 4296 alpinus v. 
subellipticus. 
Porsild, A. E. & R. T. 
v. tenuifolius, 847 Ri ere 
943, 1102 Richardsonii, 1131 g 
mineus, 1496 alpinus v. tenuis hu i. 
gramineus, 29 aelongus, 
3098 alpinus v. eabiie ea an 
Richardsonii, 3099, 5174 gramineus 
Vv. maximus. 
Atwood. 1318 Oakesi- 
Pa 1405 perfoliatus v. bupleur- 
oi 
ra & Hyland. 649 gramineus. 
Proulx, T. 58 pesca tenuifolius. 
Rand & Robinson. 1013 Sheil 
Randolph & Wiegand. 9094 


1977, 1979 praelong- 
lifolius, 


114 alpinus 


. amplifolius, 
2251, 2262 a 2267, 2273, 2299 
amplifolius. 

, E. A. 88 perfoliatus v. bu- 


1545 perhaps alpinus 
ramineus Vv. 
i — * = on 


m 1552, 1553, 1554, ‘1555, 

1556, 155i, 1558, 1559, 1 60 Rich- 

a 1570, aby 1572 peer 
6352, 64 Ric 


7002 ramineus 

Raup Abbe. 4038 gramineus, 

4313 eusven 4614 gramineus, 4 
Richardso 

Redfield, J. "H. 7996 pulcher, 8002 
perfoliatus v. bupleuroides, 8014 

inus v. subellipticus, 15341 per- 

lst = bupleuroi sts 


cena stony J. 1640. ee Sg 
~2502 pulcher. 

Ricard & Boivin. 342 nodosus. 

Ricker, natans 

Rider, Sadie L. 349 Richardsonii 

Ridgway, R. 3318 nodosus. 

Ridgway ana 3425 nodosus. 

Riehl, N. 128 er. 

Robinson B.L 494 natans. 


© per- 
foliatus v. bupleuroides, 231 poly- 


210 Rhodora [May 


gonifolius, 232 sarees ae ap- 
proaching v. maximu 

binson & Webb. 1087 perfoli- 
atus v. bupleuroides (perhaps 
hybrid). 

Rolland-Germain. 6231 gramin- 
eus, 6233, 6235 gramineus v. maxi- 
mus, 6280 amplifolius, 8696 per 

. foliatus v. bupleuroides, 8697, 13044 

amplifolius, 1 chardsonii, 
58 alpinus v. tenuifolius, 1 


T alpinus x perfoliatus v. 
a pasa 43359 nodosus, 43360 
er 


aps gramineus 4 perfoliatus 

Vv. ggeor eo nae 43382 gramineus 
iatus v. bupleuroides, 

43481 | Richandeoidt 
R R. C. 2319 alpinus v. 


b 51 us V., 
55 natans 59 alpinus v. subelli 
icus, 60 1 natans, 61, 62, 63 ampli- 
folius, 7 if gramineus, 73 


amineus v. maximus, 74 per- 
ne v. bupleuroides, 75 gra- 


: a . “P ps le rfoliatus v. 
upleuroides, raelongus 
Ro b ) B. & 16 alpi- 


nus v. tenuifolius, ss gramineus, 20 


T 67 Richardsonii. 
304 nodosus, 1185 per- 
foliatus v. bupleuroides. 

M.F. 812 natans. 


Oakesianus, 25817, 30003 nata ans, 
30004 praelongus, 31207, 32297 


boueian, 35812 perfoliatus v. 
bu 
Rugel, F, 613 illinoensis. 
Ru 384 alpinus v. subellip- 
re 385 natans. 
uth, A. 49, vi 141, 772 nodosus. 
Rydberg, P. A. 1421 ‘nodosus, 1440 


y 
illinoensis, 1652 natans, 1792 Rich- , 


ardsonii, 1846 nodosus. 

Rydberg & Bessey. 3724, 3725 
alpinus v. tenuifolius. 

Rydberg & Carlton. 7522 Rich- 
ardsonii. 

St. Cyr, D. N. 2973 gramineus, 
2994 amplifolius, 3030 nodosus, 
3050 Richardsonii. 

St. John, H. 1121, 1122 polygoni- 
folius, 1124 perfoliatus v. bupleur- 
oides, 1372, natans, 1373 iar 
anus, 1756 alpinus v. subellipticus, 
1 


bupleuroides, 11919 gramineus V. 
myriophyllus, 90081 natans, 90083, 
90084 alpinus v. tenuifoliius, 90085 
alpinus v. Oey petolet 90086 gra- 


mineus, 9008 oliatus. 
St. John, English, Moore & 
Palmer 


St. pea & Nichols. "D104 alpinus 


636 rareatewnnens 656, 658 ae 
find Richardsonii, 4927 
praelon 


Seridbere ; é asap 524 gramin- 
eus V. ma 

Sandberg, ‘Mastoural & Heller. 
697, 939 natans, 955 gramineus, 
956 Richardsonii, 1026 gramineus- 

Sanford, S.N.F. 1225, 10192 per- 
foliatus v. bupleuroides 

Sargent, H. 29 perfoliatus v. 
Saalee cies: 31 gramineus X per- 
foliatus v. bupleuroides, 33 per- 
foliatus v. bupleuroides. 


ulz, Ellen D. 797 illinoensis. 
Scott, W. 16208 crispus, 16431 n 
dosus, 1 amplifolius, 16433, 


16434 illinoensis, 16438, 16439 
crispus, 16443 gramineus, 1 
gramineus v. maximus, 


16446, ~ 


1943] 


16447 gramineus, 16448 a oa 

16450 gramineus, 16451 praelon 

16454, 16455, 16457 Richardson 
Scovel we T. 26 amplifolius and 
4 Bishasdaehit, 28 ne li- 


linear-leaved species oan to 
Fernald, Mem. Am. Acad. 
Sci: 17: pt. 1: 136), 54 Hissar: 66 


1057 reg 


ineus, 1 amplifolius, i321 
praelong 
hee M. 100 = 
Seymou 249 amplifolius, 


267, 1027 perfoliatus v. bupleur- 
oides, 1487, 1567, 1568 Oakesianus, 
570 per oliatus. v. bupleuroides, 


approaching 


. M. 3 Richardsonii, 11 
a 73, 100 natans, 101 
Sh <a & ‘Dndecwoolds 33521 no- 


us. 
Sharples, 8S. P. 303 nodosu 
Shaw, C. H. 771 Richardson 
1206 natans and gram 
745 piles 3804 
uraclongur "4524 alpinus v. tenui- 
oO 


us 
& Bessey. 5328 alpinus v. 


tenuifolius, 5333 gramineus XX 
Richardsonii. 

Shreve, F. 1597, 1622 pulcher. 
ull, G. H. 15 crispus, 39, 95 no- 


—— perfoliatusv. v. bupleur- 
osus, 


70 recon 
sonii, 80 pra gus, § S na 
Ri chnieceee 225 na 


473 py er 474 nodo eee: 
Shunk & vasinnde ng. 


us, 51 Shan nii, 
folius, 560 Richardsonii. 


Ogden,—The Broad-leaved Species of Potamogeton 


211 


Simpson, J. H. 386 illinoensis. 
ie J. K. 7241, 8170 illinoensis. 
"& Carter. 1007, 1118 illino- 
is 
ll, J. K. & G. K. 4148, 4487, 
eh illinoensis. 
iley, F. J. 330 natans, 749 no- 
mith, E. F.. 87 praelongus. 


Sones, G 307 nodosus 
Sperry, C. C. 509 pulcher. 
Sperry Martin. 671 Richard- 


sonii, 696 praelongus, 718 Richard- 
sonii a gramineus, 731, 735 il- 


linoens 

Sieaadbor rough, W. 16429 alpi- 
nus v. tenuifolius, 20750 gramineus, 
62662, — Richards onii. 

Standley 7557 gramineus V., 
9780 nN 16855, i alpinus 
y. tenuifolius, 18500 na , 18528 
amplifoliue, 40488 ciate, 40649 
dee ene 

& or 7648 ampli- 

bra 7649 nodos 

Stecker, A. 332 apis v. tenui- 
— 


tee J. H. 4741 alpinus v. 
tenuuifolius, 4781 eee 
mplifolius, 


2 


Pp 
bupleuroides, 527 per nin 
eus < nodosus, vont Berchtoldi x 
e 


ardsoni i, 830 natans, 

60 ramplifolis, 957 einen 1127 
Oakesia 

Steinmetz “& tet . 888 gramine- 

us perfoliat bupleuroides, 

889 sectelintur ag Na a etary 905 

nodosus. . 


teinmetz & Marston. 539 Be 
toldi X perfoliatus v. bupleuroides. 


212 Rhodora [May 
Steinmetz & Ogden. 98 ampli- & Leslie. 280 amplifolius, 291 

folius . . gramineus, 293 Richardsonii, 294 
Stei & Quimb v- 581 gra- alpi ious ¥ papain 295, 296 


anson. Ee natans, 
31 nodosus, 55 gramineus. 
Stevens, G. W. 1364 amplifolius 


rmark, ‘J. A. 1003 natans, 

$297 ori crispus, 4327 pulcher, 4473 

4609 illin noensis, 4639, 4688, 

5410 nodosus 

inoensis on dosus), 

BALI, 9233 fe pa 11328, 11939 

ees, 11946 amplifolius, 
12031, 12428 open 3782 

13869 amplifolius, 13910, 13938, 

14226 awn it. 14245 _pulcher, 

14257, 9 nodosus, 14985 - 

ees 14487, 14624, 15560 am- 


plifolius, 16: 13 pus, 21087, 
21138, 21145 illinoensis, 21208 a 
plifolius, 21235 illinoensis, 2126 


9 
22 us, 22773, 

22840, 22873, 23321, 23365 ampli- 
folius, 23494, 23543, 23546 ampli- 
folius, 24650 ‘nodos 
pia 
103 nodosus, 25104, 25142 il- 
linoensis, 25249, 25250, 25317 no- 
dosus, 25365 illinoensis, 25367 no- 
osus, 2537 inoensis, 25425 
nodosus, 25533 mnplifclitn 25534, 
ont 25876, 27148, 27634, 27657, 


Street & Williams. 2689 nodosus. 

728 nodosus, 2172 
alpinus v. tenuifolius 

Svenson, H. K. 156 pulcher, 7216 
nodosus, 9108, — ae 

Svens: Fas 932 


5786 praelongus 
bie soi bod P. 78 gramineus x il- 
Pasice, ee Fitzpatrick, Losee 


natan Oakesia gra- 
mi oan. 300 Ri chardsonit, te gra- 
mineus, 304 gramineus X ard- 


sonii (proba bly backerossed with 
gramineus), 305 gra 
arp, B. C 


Cc. 2181 sisi naaeiaie. 
horns Cc. C. aps gra- 
us X nodosus, 1499 gramine- 
on "3466 perfoliatus v. bupleuroides 
(perhaps perfoliatus v 
oides X Se 1 


eictamecn Richardsonii. 
Thompson, 589 gramineus, 
594 rdsonii, 8620 ampli- 


7 c 

folius, 942214 natans, 11768 gramin- 
eh Vv. sie “pase 14046g ramineus. 

Richardson, 7 natans 

mineus, 53 illinoensis, 69 gramine- 

us, 73b gramineus > _ illinoensis, 

142 Richardsonii, 145_ illinoensis, 
ichardsonii, 
ns, 253 ainionrn 99 


Thurber, G. i co ; 
Tidestrom, I. 517, 1861 illinoens!s, 
7171 nodosus, 7175 perfoliatus V. 


bupleuroides, 7183 P7637, 7741, 
10 
Tolst 428 natans, 429 


W. L. 
Richardsonii, 430 praelongus, 473 
Richardsonii, 614 se nig 615 
gramineus, 616 gramineus ™ il- 
linoensis, 637 nodosus, 638 natans, 


us. 
Topping, D. L. 203 amplifolius. 
Toumey, J. W. sea nce s. 


D; 867 gree 
isk 2870, 2871 alpinus v. tenul- 
lius 


eedy, F 


physio 


. 46 illinoensis, 411 gr@ 
413 Richardsonii, 4978 


a 62, 373 
cog caer 375, 1091 gramineus 
x illino 

e mite. 1645 Richard- 
sonii, 1656 gramineus, 1659 Rich- 
ardsonii, 1660 natans, 1662 line: 
ensis gramineus (perhap 
gramineus > illinoensis). 

Uhler arren. 793 natans 
797: % ono 858 jllinoensis, 
1079 praelon 


1943] 

Umbach, L. M. 31 gramineus, 433 
Richardsonii, 457 gramineus v. 
eg a 
an ,» EB. C. 202 alpinus v. 
tenuifoliv, 


aa & wenn 201 crisp- 


202 
Victo Mar 526 Ey 
565 Test lint v. bupleur 
1124 tries 1152, 1505 alain 
v. tenuifolius, 1537 ‘natans, 1539 
gramineus, 3177 “paige cores . bu- 
chardsonii, 


oides, 8177, 8178 perfoliatus y. bu- 
pleuroides, sohiardanati, 


cus, 1116 ps sec per- 
foliatus v. bupleuroides, 11169 
perhaps gramineus X _ perfoliatus 


bu ides, s, 
16056 $060, eoee” 16057 — amin- 
ie . natans, 16064 

mi ae ximus 16065 am- 


plifolius, 18460 alpinus v. tenui- 
olius, 18462 Richardsonii, 20451 
2 gramineus v. 
maximus, 20454 perfoliatus, 20456 
perfoliatus v. a 20457, 
21201, 25815 amplifolius, 2 27897 
perfoliatus v. bupleuroides, sa 
gramineus, gramineus 
rfoliatus v. bupleuroides, 23005 
Richardsonii (perhaps —— 


Victorin & peieenoneen 9922 Oakesi- 


v. sube 
25810 gramineus, 25938 alpinus 
subellipticus, 25939, 27094, 27095 
gramineus v. maximus, 27 630 alpi 
nus v. tenuifolius, 29088 peafolintas 
v. kag tebe 33178 Richard- 


sonii (perha oliatus v. bu- 
pleuroides < Richardsonii), 33180 
gramineus v. maximus, per- 


foliatus v. arr ait 43565 no- 
dosus, 43576, 43796 perfoliatus v. 
bupleuroides, to70 —— 44738 
gramineus X_ perfoliatus v. bu- 


Ogden,—The Broad-leaved Species of Potamogeton 


213 


pleuroides, 45184 ree ns 
45189 crispus, 4 osus, 
49141 crispus, 49148 Richardsonii, 
49304 nodosus, 49366 amplifolius. 
Victo on olland, Brunel & 
Rous 17278 perfoliatus v. 
bupleuroides, 17285 gramineus ap- 
sine v. een 17286 per 
siren v. bu es. 
Victorin, Rol aa “& Jacqu 
33315 alpinus v._ subellipticus, 
ae ee 33393 perfoliatus ap- 
aching v. bupleuroides, 33477 
vena Pi 351 amplifolius, 
33518 praelongus, 33609 Richard- 


sonii, 9 Oakesianus, 33 
praelongus, 33854, ae natans, 
33858 a mr 9 perfoli- 
atus v. bup oides, 33714, 33855 
asic ig “441 praelongus, 
44451 . subellipticus, 
44452 gramineus, 44453 gramineus 


Lee oliatus — v. 
bly backerossed with gra- 
mineus), 44454, 44458 - ear 
44459 gramin eus x perfoliatus v. 
bupleuroides, 44460 natans, 44463 
ramineus, 


44468 alpinus subellipticus, 
44749 Oakesianus. : 
Victorin, Rolland & Louis-Marie. 


Frio alpinus v. subellipticus. 
torin, Rolland & Méeilleur. 
"2B 730 alpinus v. tenuifolius, 13858 
2 grami 


nodosus, 44382 gramineus v. 
ramineus X sp., 44728 aie, 
45446 Richardsonii. 
rin, d, Michel & 
oe peta 43633 esianus 
Waghorne, A. C. 6 alpinus v. 
senuitalincs, 40 perfoliatus v. bu- 
pleuroides. 
Wahl, H. A. 174 a sh ae 204 


perfoliatus, v. bupleuroides, 240 
perfoliatus v. indedroides (perhaps 
perfoliatus v. bu “gg Pa — 
alker, Mr. & Mrs. E 


atans. 
Ware, R.A. 2446 perfoliatus v. ed 
pleuroides, 3324 natans, 3326 a 


plifo 
arnock, B. H. 1557 nodosus. 
Watso on, S. 396 perhaps alpinus X 
a patie 397 Rianlacst, 398 
, 1131 natans, si alpinus 
v. rane Sa Se 
haps illinoensis secsisaean, 1134 
gramineus, 1135 Richardsonii. 
Watson wT. R. 969 gramineus, 973 


214 


——. approaching v. 
s, 979 natans, 1006 amplifies, 


ramineus, 36 


Weatherby, C. A. D2409 ~— 
folius, 3389 gramineus, 4364 n 
tans 

Weatherby & Anderson. 5901 
amplifolius, 5904 natans. 

Weatherby, C.A.& UnaF. 5625 
gramineus stk haste Vv. maxi- 
mus, 6624 gra 


Webb, R. J. 452, "546 amplifolius. 
Webber, H. J. 4 amplifolius, 5, 6 
dosus. — 


Welch, Winona H. 2106 nodosus. 


A. 395 541 
natans, fd bhp Vv. maximus, 
550 gram 


Wetmore, R. ‘#. 103096 gramine- 
axim 


Richardsonii, 90 alpin 

—_ 231 illinoensis, 973 Pelee soie 
Wheeler, L. C. 1974 nodosus, 3973 
Whited, z 3141 ge tnegnniees 
Whitford, H. gr cape tiny 

—o natans, 258 Spe v. tenui 


Whitney, Elsie G. 4673a ampli- 
folius. 


Rhodora 


[May 


Wiegand, K. M. 
11182 crispus. 
Wiegand, Gilbert & Hotchkiss. 

27339 Oakesianus. 


11178 illinoensis, 


Wiegand & Hotchkiss. 27337 alpi- 
y ore 27343 gramineus 
up bes 


(probably gies eee with g 
mineu 
Wiegand & Pease. 27340 gramine- 
ight, W. F. 1 praelongus, 8 il- 
linoensis, 9 natans, 59 illinoensis, 


tans, 6796 eo. 6797 
Williams, S andosaas ‘804 
Richardsonii $00 pocbarse i 
illi T. A. 1 nodosus, 344 
amplifolius 
Williamson, C.8S. 418 Oakesianus, 
455 chyeoeitolick: 472, 1026 per 


foliatus v. bupleuroides. 
illiamson, W. T. 309 no- 


189 amplifolius. 
alpinus v. tenuifolius, 


. 8290 nodosus. 
Wood, F. F. 4 Richardsonii. 
Wright, C. 675 nodosus, 676 il- 
ong 1893 nodosus. 
. P. 542 illinoensis. 
amplifolius, 


G. & E. C. 7011 


E. B. 1237 gra- 
mineus, 1238 amplifolius. 


INDEX 


New scientific names are printed in full-face type 


iar gp na subg. Coleogeton, 71, 
1 ect. Connati, O, sect. on- 


A 
69, 71, 72, 76; subsect. Amplifolii, 
63, 68, 69, ei "72, 2, 76, 132; subsect. 


73, 76, 133, 158, 


: . 6 
71, 74, 76; subsect. Pusilli, 71, ser. 
Pusilli connati, 71, va pots, 
71, subser. Pano ormitani, 71, se 
Pusilli convoluti, subser. ‘Acuti, “1, 


9-62, 69, 71, 72, 75, 95, 99, 
104, 122, 123, 127, 128, 137, ise 
9 Is. 


ensis, 184; amplifolius X_ prae- 


215 


longus, 184; am plifoli ius X Rich- 
ardsonii 


jamaicensis, 124; confervoides, 71; 
crispus, 60-62, 64.°71,:72, 75, 
76, 86-88, 91, Aa deminutus, ‘187; 
diversifolius, ; epihydrus, 


186; epihydrus, var. Nuttalii, 138, 
142: epihydrus, var. Nuttalii 


maximus, 
153, he 196, pl. 748, var. mi inimus, 
50, myrioph yllus, ior 145, 
147, 150, 151, var. spat thulaeformi 
151, 187, 189, forma pertoctvie 144, 

var. typicus, 143, 149, 151, 196, 
pl. 748, forma Wolfgangii, 148; 
gramineus X illinoensis, 145, 149, 
51,158, 1 


1 1 87, 2 us X 
illinoensis x lucens, 187; grami- 
s X lucens, 158, 187 mineus 


xX natans, 186; gramine us xX 
nodosus, 185; gramineus X_per- 


216 


foliatus, 151; gramineus X_ per- 
foliatus, var. bupleuroides, 61, 183, 


peduncul — 143, forma maximus, 
pa 187, minimus, 1 0 
myri tenbyilne 150, 
toreeaeeee 144; Hillii, 71; Ginseng 
21, ta, 4, 93, 94, 150, 152-154, 
156-159, sine 188, 191, 192, 195, 
ma homophyllus, 154, 
s; 154; illinoensis X 


158, var. ke at 1 


=) 
< 
ro 
= 
D 
=} 
= 
O. 
As] 
ss; 
‘et 
< 
S 
— 


var. ter- 


7 ‘ ; panor- 
mitanus, 192; pectinatus, 71, 192; 


INDEX 


pennsylvanicus, var. portorioemaa 
Mf Lt 


bupleuroides, 171, 1 

178, 181, 182, 189, 190, 193-196, 
pls. 746, 748; ‘perfoliatus, var. bu- 
pleuroides xX Ri i 5 
amen var. gracilis, 163, 177, 
180, var. lanceolatus, 1 3, SSP. 


oe polygonifolius, 58, 59, 12; 
8, 195, p 746; Porsildior- 


a — 


1, 195, a 131-133; rufes- 
cens, 90, 92, § 
90; x seolioph is 
barensis, ee 

186; spathaeformis 187; * 8 spath- 
aeformis, spathulaeformis 


siietitolias, a1; subdentatus @ 
ag 191, p petiolatus, 188, 189, 
1 x su 


gracilis, 154, 187, 
og? Var. ss ierentifoltad, 154, 187; 
Zizii, forma pulcherrimus, 187; 
zosteriformis, 
spirillus amp lifolius, 100; hetero- 
phyllus, 44; er 124; lucens, 
154; perfoliat us, 177, va Richard- 
sonii, 163; pulcher, i21; Zizi, 154, 
187 


Reprinted from Ruopora, Vol. 45, June-August, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 


I. Five common Rhizomatous Species of Muhlenbergia 221 
ii... Netes on Danthoiiia. 3.5.0 oo tt ee 239 
III. Erianthus brevibarbis and other Species........--. 246 
IV. Why not Andropogon Gerardi?...........:..+:.. 255 
7, Studies in North American Species of Scirpus... ... 279 
VI. The Identity of Scleria setacea of Poiret.........: 296 
VII. What is Angelica triquinata?..............+:-.+- 298 
Vill. Notes on Mierschim 53.5 ee ee 317 
By M. L. FERNALD 
Dates or Issuz 
Pages 221 to 258, plates 749 to 761... .--..- 6. +e. ee eee ee 29 May, 1943 
Pages 279 to 303, plates 762 to 769. .-.------+ +--+ 225+-257 - July, 1943 : 


- 
Deas ee 10 August, 1 


OF HARVARD UNIVERSITY 


CXLVI / 


Reprinted from Ruopora, Vol. 45, June—August, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


CXLVIII 


I. Five common Rhizomatous Species of Muhlenbergia 221 


Hi. Notes of Peathonsa eo ee 239 
III. Erianthus brevibarbis and other Species.......... 246 
IV. .. Why not Andropogon Geraidit. oo 255 

V. Studies in North American Species of Scirpus...... 279 
VI. The Identity of Scleria setacea of Poiret.......... 296 

Vit. What is Angetica triguinatal. ...........,.55.. 3. 298 
Nill. : Notes on Hieracnii., ... - aa ee 317 


By M. L. FERNALD 


Dares or IssuB 


Pages 221 to 258, plates 740 to 761. ~~... 5 +e eee ee ee. 29 May, 1943 
Paces 279 to 303, pinta TOR 06 700s, tas ee tk: 6 July, 1943 
Pages 317 to B26... 656s oes hee ce re as ee be eae 10 August, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM OF 
HARVARD UNIVERSITY—No. CXLVIII 


M. L. FERNALD 
(Plates 749-769)! 


During the studies necessary in a thorough revision of the flora 
of northeastern America and, especially, a checking with photo- 
graphs of the types of Linnaeus, Michaux and other authors of 
American species much new matter has accumulated. Some of 
the studies, with photographs by Dr. Bernice G. Schubert, are 
here presented. 


I. FIVE COMMON RHIZOMATOUS SPECIES OF 
MUHLENBERGIA 


(Plates 749-757) 


In eastern North America five species of Muhlenbergia stand 
out as the most common representatives of the rhizomatous 
members of the genus, the plants passing, mostly erroneously, 
as M. mexicana (L.) Trin., M. sylvatica Torr., M. foliosa (R. & S.) 
Trin. and the two very distinct species included under M. 
racemosa (Michx.) BSP. These five species have many technical 
differences (in glumes, lemmas, anthers, grains, internodes, 
nodes, sheaths, etc.) and habitally they are distinctive. The 
names currently applied to them, however, need most careful 
scrutiny for, when the types (all but one in Europe) are studied, 
it is apparent that those who have easily recognized the species 
involved have largely guessed at their names and, too often, 


1The cost of pmanperly blocks has been met in part from an appropriation for 
original research from the Department of Biology, Harvard Universi 


222 Rhodora [JUNE 


have not consulted the original descriptions. Sixteen years ago 
I undertook a clarification of our species, at that time securing 
from friends in London, Paris and Berlin photographs of many 
of the types involved; and in 1930, while in Europe, I checked 
many points not shown in habit-photographs. 

As a result of this study it was evident that we must adopt 
many unfamiliar names or change the significance of those now 
familiar but, disliking the upheaval necessitated, I have refrained 
from publication, with the hope that specialists on the Gramineae 
would take the responsibility of correcting the many errors. 
Having now reached a stage in my work when the facts can no 
longer be dodged, I am presenting in key-form the diagnostic 
characters of the five species. This will be followed by consider- 
ation of the names now in use and those which, it seems to me, 
must be taken up. In some instances, however, inability at 
present to consult many critical specimens abroad may result in 
slight future changes. In his Manual Hitchcock! adopts for 


‘Tam lana g informed that Hitchcock strenuously and rightly objected to these 
es, which were forced into his book through the ruling of his Chief (see 
Ww 


to upset when studied by scientists. If the time used in inventing ‘‘English” names 
for these plants had gone into study of their proper typification some of the upsets 
in ye sinc pages might be unnecessary. ‘‘C’est a rire!”’ 
pos the reference to dictatorship by some in power over young (sometimes 
eee pectoris have recently received copies of correspondence from a botanist 
formerly in siti ernment employ, in which the inventor of “‘English’’ names ruled that 
the y n should not publish a new plant in Ruopora because the type is not 
in the Nasi! Herbarium and because the author had followed the Internationa 
Rules of Plant Nomenclature, though in this case the dictator was overruled by 
higher officials. Probably no more severe blight upon scholarly aah gp vot 
taxonomy under federal dictatorship has existed. The inventor of ‘English’ names 
himself once made a wi effort in taxonomy, That he nes not ve is clear 
from his greatest nap hata with its delightfully naive key 
Leaves obova’ 
Leaves not i on one characters given — etc. 
Leaves not erie ecir marge 


1943] Fernald,—Rhizomatous Species of Muhlenbergia 223 


members of Muhlenbergia the stupid ‘English’ name ‘“‘Muhly”’ 
(“Marsh Muhly’’, ‘‘Wire-stem Muhly’’, ete.). There would be 
some propriety in dropping the h and then applying the word as 
an adjective to the current nomenclature of the misinterpreted 
five. In tabulating the characters I am numbering the species, 
the name in current use being bracketed. Beautifully clear il- 
lustrations of four of the plants will be found in Hitchcock’s 
Manual, figs. 768, 770, 774 and 776. I am greatly indebted to 
Mr. Jason Swallen for an opportunity to study fragments of the 
types of Agrostis mexicana L. and of A. foliosa R. & S. Photo- 
graphs made from them will be found in pLates 749, Frias. 1-3 
and 751, FIG. 2). 


a. a and stolons 2-6 mm. thick, with cucullate-arching 
vate scales; nodes of culm thick ‘and ever enlarged; 
oF 


mm. long; 
— anthers 0.3-0.6 mm. 


ranches soon a the me ants thus 
habit and topheavy, the p mptly uncovered internodes 
lustrous and glabrous ; leat-ehew = loose, r com- 
pressed, promptly opening to base; terminal panicle ex- 
serted, or with included base, at first lax and open, lan- 


very rey pa both awned, membranaceous, green 
; lemma ape Te ; grain easily Cronies 


linear-cylindrc, 16-3 ih 0 os a as a ees l, (M. mexicana). 
. Culms erect or ascending, lower nodes not rooting, un- 
bra ee a0 


from middia: node es, internodes largely covered | 2 sheaths, 
opaque and puberulent; leaf-sheaths tight, tere te, tardily 


aly glumes ni rat (if "rarely awned, sub- 

Rivet, with few simple ae ssed-ascending branches; 
leaf-blades sovaudien or sli re! spreading, firm; pani 

rather stiff, the subse art amici sive spikelets 
imbri ases of the glom 


ulate appressed-ascending branches and _ branchlets; 
glumes subequal, firm and herbaceous, usually green or 


Leaves not with all ag iss aude given above. 
Leaves orbicular to te. 
Leaves or other Beer lem as described a! 
and so on, including the informative calls in proay cap ‘Leaves not as described It 
is no uaa that one who 60 little deeper than that into an exacting field did not 
fully understand its importance 


224 Rhodora [JUNE 


purplish; lemma firm, usually awnless (rarely awned) ; 
mature florets persistent; grain slenderly ellipsoid, 
1.3-1.6 mm. long, tightly embraced by lemma and palea. 

2. [M. foliosa]. 
Ascending to erect, with many ascending simple or forking 
branches from middle nodes; leaf-blades spreading, 
rather soft and pliant; panicles slender, loosely flowered, 
exuous, the longer branches mostly without crowded 
basal spikelets, the spikelets often long-pedicelled; 
umes very unequal, scarious or hyaline, usually whitish 
or silvery; lemma scarious, silvery, often long-awned; 

mature florets promptly dropping; grain linear-cylin- ; 

ic, 1.4-2.1 mm. long, easily remove 3. [M. sylvatica]. 
a. Rhizomes and stolons slender (2-4 mm. thick), greatly pro- 
longed and forking, with appressed lanceolate to narrowly 


8b Owe eee hee 


ty) , 
Culms mostly branching from middle nodes, the leaves and 
branches there approximate, the internodes lustrous and 


3 .8 mm. long; grain tightly 
embraced, linear-cylindric, 1.8-2.2 mm. long; species of 
dry prairie, rocks and bluffs from Wisconsin to Saskat- 
chewan, and west to Oregon, south to Illinois, Missouri, 
Kansas and New Mexico; casual al ilroad tward 


5 


4. [M. racemosa]. 


True M. mexicana (Puates 751 and 752) 


Muhlenbergia mexicana (L.) Trin. Gram. Unifl. 189 (1824) 
rests upon Agrostis mexicana L. Mant. i. 31 (1767). Agrostis 
mexicana was raised by Linnaeus in the garden at Upsala from 
seeds sent him from Vienna by Jacquin and erroneously thought 
to have come from tropical America. By most recent authors 
it is considered to be species no. 1 of the preceding key: “the 
plants becoming topheavy and bushy”, to use Hitchcock's 
phrase, with abundant decumbent and rooting or ascending ofte? 
forking lustrous branches; the sheaths compressed and loose, 
promptly opening to the base; the branches bearing numerous 
partly included axillary panicles; the terminal panicles relatively 


1943] _Fernald,—Rhizomatous Species of Muhlenbergia 225 


soft and loose upon expanding, many of the spikelets long- 
pedicelled; the glumes very unequal and both awned; the easily 
removed grain linear-cylindric, etc. How different was the 
unusually full description by Linneaus of his Agrostis mexicana: 


mexicana. 20. _AGROSTIS panicula lounanien ealycibus [glumis] 


Hl. citar in n America calidiore. D. J —- a 
Culmi numero dales, laeues, er any Ramis indiuisis. 
nea, 


inatae 
mata atropurpurea, ramosa. Stamina alba. Difficillime hoc 
Gramen determinatur. Altero anno floret. Facies Cinnae. 
H.U: 


Passing for the moment the ‘‘panicula oblonga congesta’”’ or 

“congesta coaceruatis’’, it should be clear that the ‘‘culmi j 
erectr”’ the “Ramis Snddiciane?” and the “Calyces [glumae] apice 
subulati, subaequales”’ do not belong to species no. 1, which has 
often decumbent or lopping culms with mostly forking and very 
abundant basal and median branches, the glumes very unequal 
and both slenderly awned. The ‘‘Aristae nullae’’ could belong 
to no. 1 or to no. 2. Linnaeus said nothing about the very nu- 
merous axillary panicles partly included in the subinflated 
sheaths of the branches which so generally characterize no. 1. 
His unusually full description is clearly not a good one for no. 1 
and, in view of the almost universal recent error of so identifying 
it, the original comment of Linnaeus still has force: ‘‘Dzfficillime 
hoc Gramen determinatur’’. Linnaeus had two sheets of the type 
material, both clearly marked by him Agrostis mexicana. Beau- 
tiful photographs of the two sheets, sent to the Gray Herbarium 
in 1927 by the late Dr. B. Daydon Jackson, are before me. One 
is of a badly crumpled specimen, the other (our PLATE 751, Fria. 1) 
better prepared. They both show erect culms, with few simple 
erect branches, tight sheaths, terminal panicles, no definitely 
included axillary panicles, subequal awnless glumes (PLATE 751, 
FIG. 3) and awnless lemmas; and the broken-off culms are 5 dm. 
(20 inches) high. Linnaeus’s “‘culmi . . . pedales’ was too 
conservative. PiatTe 752, ric. 2 is from a portion of the panicle 
of the type, now preserved at the United States National Herbar- 


226 Rhodora [JUNE 


ium and most generously sent me for study by Mr. Swallen. 
The type specimens of Agrostis mexicana are of species no. 2, 
the plant passing as Muhlenbergia foliosa. 

As he stated, Linnaeus received his seed from Jacquin and 
raised the plant at Upsala. Jacquin’s plant at Vienna was 
clearly described, and illustrated by a life-size colored plate, by 
the younger Jacquin in his Eclogae Plantarum Rariorum— 
Gram. Fase. ter. et quart., 44, t. 30 (1813). Jacquin filius 
departed little from Linnaeus, his fuller account including 
“Cutmi . . ._ bipedales, : erecti, . . .. teretes 
ik ; ramis axillaribus, secs, Meticlite: adpressis 
vaginis . . . apertis . . . Ftorssin paniculis suis 
nalibus culmi primarii et ramorum, ante et post anthesin coarc- 
tatis, . . . 3 famis Tamulisque ... .. erectis adpressis 

Calyx . . . Glumae subaequales, lanceolatae, acu- 
vitae ; and his wonderful plate (our PLATE 752) showed a 
characteristic plant of our species no. 2, even to the slenderly 
ellipsoid grain. The material cultivated by the Jacquins at 
Vienna was, then, like that sent to and cultivated by Linnaeus 
at Upsala and described by him as Agrostis mexicana. When he 
transferred A. mexicana to Muhlenbergia, Trinius, Gram. Unifl. 
189 (1824) held the diagnostic characters of the Linnean species 
“Panicula contracta densiuscula; Glumis lineari-lanceolatis acu- 
tissimis subaequalibus perianthio acutissimo vix brevioribus’’. 
There seems to be no way to avoid taking up for the plant erro- 
neously passing as M. foliosa the clearly described and typified 
M. mexicana (L.) Trin. 


M. MEXICANA OF RECENT AuTHORS (PLaTEs 749 and 750) 


When we consider the earlier available names for the plant 
recently but erroneously passing as Muhlenbergia mexicana, our 
species no. 1, the name seemingly available, if we accept current 
bibliographies, is M. lateriflora (Michx.) Trin. ex Kunth, Enum. 
i. 207 (1833) in synonymy of Cinna lateriflora Kunth, Rev. 
Gram. i. 67 (1829), both names based on Agrostis lateriflora 
Miehx. Fl. Bor.-Am. i. 53 (1803) from rock-cliffs of the Missis- 
sippi and shores in [southern] Illinois. Michaux’s description 
is not at all good for a plant with geniculate and bushy-branched 
stems, relatively large terminal panicles and long-awned glumes; 


1943] Fernald,—Rhizomatous Species of Muhlenbergia 227 


and the photograph of the type sent me by M. Cintract is very 

inconclusive. The description was as follows: 

LATERIFLORA. A. culmis erectis, nodosis: foliis linearibus, planis: panic- 
ulis lateralibus et terminalibus, pusillis, coarctatis, densifloris: 
glumae muticae et scaberulae. valvis acutissimis; interioribus 
majoribus, basi barbulatis 

Hab. praesertim in praecipitibus saxosis fluminis Mississipi et ripariis 
Illinoensibus. 

Authors immediately following Michaux seem not to have known 

the plant, Beauvois shifting it without discussion to Vilfa, and 

Kunth, likewise with no discussion, transferring it to Cinna. It 

was not until Kunth’s Enumeratio that the plant, as Cinna 

lateriflora (Michx.) Kunth, with ‘‘Muehlenbergia lateriflora 

Trin.” as a synonym, was again accorded a diagnosis, that copied 

directly from Michaux. Michaux’s erect culm, densely flowered 

small panicles and muticous glumes are not satisfactory for the 
common plant known as M. mexicana. The photograph is of 
mere fragments: broken-off tips of a stiffly ascending plant with 
long internodes; leaf-blades erect, only 2-4 mm. broad and long- 
tapering; the linear-filiform panicles only 1.5-4 mm, thick, with 
spikelets only 2.5-3 mm. long. These fragments look like small 
bits picked from a plant of M. glabrifloraScribn. Only Michaux’s 

“slumae . . . interioribus . . ._ basi barbulatis’” would 

seem to separate it from that species of dry or baked soils, gravels 

or rocky slopes, from southwestern Indiana (perhaps also Ohio) 
and Illinois to Texas. MM. glabriflora is reputed to have the 
lemma glabrous. It would not now be justifiable to reduce 

M. glabriflora to M. lateriflora; when the florets can be actually 

studied that may be inevitable. In fact, Steudel, Synop. PI. 

Gram. 182 (1854), describing in detail the species he took to be 

Cinna lateriflora (Michx.) Kunth, from Ohio — of Frank’s, 

said “glumis . ye @mubesqualibus( <>. 6; valyulis-. 

glabris.”’ It is, however, fairly clear that we Ms not scopeny 
take up M. lateriflora for the plant, no. 1, which has been passing 
as M. mexicana. 

Another name, ignored or waved aside by recent American 
authors, needs consideration, for it is with little doubt the first 
name for species no. 1, the plant generally passing as Muhlen- 
bergia mexicana and so passing for more than a century, the 
confusion dating back to botanists of a full century ago, when 


228 : Rhodora [JUNE 


nos. 1 and 2 were completely mixed in our floras, Torrey, Fl. 
N. Y. ii. 437, 438 (1843), for instance, saying ‘“M. foliosa Trin. 
(Agrostis filiformis, Muhl. gram.) seems to be only a variety of 
this species’’, in which, unintentionally perhaps, he was correct, 
for, as I later show, the type of Agrostis filiformis is the form of 
true M. mexicana with awned lemmas. The species (M. mezi- 
cana sensu Torrey) with “Culm . . . much branched . . - 
often geniculate; sheaths compressed, loose . . . is some- 
imes troublesome in gardens’, the latter points applying to 
ecies no. 1. The neglected name to which I refer is Agrostis 
frondosa Poir. in Lam. Encye. Suppl. i. 252 (1790). The 
description, noting fleruous stems, very leafy and compressed 
branches, purplish nodes, loose sheaths and oblong and often pur- 
plish panicles suggesting those of A. mexicana, are all indicative 
of species no. 1. Here is Poiret’s account, he giving the name 
because of the leafiness of the plant: 
46. Acrostis feuillé Agrostis frondosa. 
Agrostis culmo flexuoso, articulis ramosis, foliosis; panicults coarctats; 


mexicana; les ramifications en forme d’épis oblongs, de couleur verte, uD 
peu purpurine; les fleurs petites, oblongues; les valves calicinales plus 
courtes que celles de la corolle, aigués, mucronées A leur sommet; celles 
de la corolle étroites, oblongues, aigués. 

ette plante croit en Allemagne. (V. s. in herb. Desfont.) 

Although it was supposed that Agrostis frondosa came from 
Germany, the name was completely disregarded in the great 
German floras of Gmelin, Koch and others of their period; nor 
does it have any recognition in the more modern and voluminous 
treatments by Ascherson & Graebner and by Hegi. The source, 
like that of so many species early reaching European collections, 
was evidently misunderstood, as was that of A. mezicana of 
Linnaeus, Jacquin and others. Nevertheless, the weedy tend- 
ency of species no. 1 evidently enabled it to reach Europe some- 
what after Poiret’s time. For instance, Lejeune & Courtols, 
Comp. Fl. Belg. i. 61 (1828) refer to A. ‘mexicana’ having bee? 
collected in Belgium, “‘e semine exotico orta’’; and Hegi, Il. Fi. 


1943] Fernald,—Rhizomatous Species of Muhlenbergia 229 


Mittel-Eur. vii. 154 (1931), refers to the recent occurrence in 
Germany of M. ‘“‘mexicana.’’ Whether these are true M. mexicana 
(‘foliosa’’) or the very different plant (our no. 1) which generally 
passes as M. mexicana I can not now determine. 

But, returning to Agrostis frondosa Poir. Roemer & Schultes, 
Syst. 1. 373 (1817) made it an exact synonym of their newly 
defined A. foliosa, saying ‘‘certissime huc spectat’”’. Steudel, 
who certainly understood grasses, treated it unequivocally in his 
Nomenclator, ed. 2: 365 (1840) as identical with various plants 
now placed under Muhlenbergia (including M. foliosa, ete., but 
not M. mexicana, which Steudel retained in the true sense). 
Hooker filius & Jackson in Index Kewensis, placed it, also with- 
out question, in the all-inclusive M. mexicana of their period. 


As already noted, Poiret’s “‘culmo flexuoso . . . de chacune 
de ces articulations il sort des rameux feuillés, comprimés, égale- 
ment flexueux . . . Les feuilles . . . nombreuses; leur 
gaine liche, . . . Les panicules . . . les ramifications 


en forme d’épis oblongs” are reasonably good characterizations of 
our common so-called M. mexicana; no other species involved in 
the problem has lax and compressed sheaths. I am, therefore, 
subject to verification when the type can be studied, identifying 
M. mexicana of recent authors with AGrosTIs FRONDOSA Poiret. 

The name Muhlenbergia foliosa (Roemer & Schultes) Trin. 
Gram. Unifl. 196 (1824) rests for typification upon Agrostis 
foliosa Roem. & Schultes, Syst. Veg. ii. 373 (1817). As currently 
treated M. foliosa is an erect plant with few erect branches; 
scabrous-puberulent internodes; firm ascending leaves; ‘‘panicles 
mostly exserted, often rather long-exserted, narrow, of numerous 
short appressed densely flowered somewhat aggregate branches’”’ 
(Hitchcock), the branches densely flowered to base; the spikelets 
subsessile or short-pedicelled; glumes subequal and subulate- 
tipped (only rarely definitely awned), firm and herbaceous; the 
usually awnless lemma of similar texture; the mature florets 
tardily dropping and tightly embracing the slenderly ellipsoid 
grain (1.3-1.6 mm. long), the plant, my no. 2, which I identify 
with M. mexicana in the strict sense. Roemer & Schultes 
evidently supposed that Poiret’s Agrostis frondosa was the plant 
in cultivation under the equivalent name, A. foliosa, and their 
glumes unequal, with hispid awns (“calycibus inaequalibus aris- 


230 Rhodora [JUNE 


tatis hispidis”) might well have been derived from A. frondosa; 
but their lemma and palea unequal, the former awned at tip and 
the culm much branched, erect (“corollis . . . inaequalibus 
valvulé majore apice aristata, culmo ramosissimo erecto’’) sug- 
gest M. sylvatica! Roemer & Schultes had two varieties: ‘ee 
paniculis viridibus” to which they assigned A. filiformis Willd. ; 
and “8. paniculis rubicundis” to which they assigned A. frondosa 
Poir. The latter has already been discussed. 

The two subdivisions of their Agrostis foliosa were borrowed in 
part from Willdenow, who had described, also from cultivated 
plants, A. filiformis Willd. Hort. Bot. Berol. i. 95 (1809); but 
without including in it A. frondosa Poir. 


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Agrostis foliosa Hortulanorum. 
Habitat in America boreali. 2. D. ; 
Polypogone diversa calycibus corolla brevioribus. Habitu accedit 
ad A. mexicanam, sed calycibus, corollis, inflorescentia tenuiore, et 
florescentia serotina diversa. Varietas 8 robustior. 

A photograph of Willdenow’s type of Agrostis filiformis, most 
kindly presented to me by Professor Diels, shows the slenderest 
extreme of Muhlenbergia mexicana, the plant usually passing a 
A. foliosa; but the awns at the tips of the lemmas show that it 1s 
the form known as A. foliosa, forma ambigua (Torr.) Wieg. The 
exact identity of the plants of Roemer & Schultes must await 
opportunity to study them. In the mean time, however, the 
very liberal fragments of what was taken as the type of Roemer 
& Schultes have been sent me from the National Herbarium at 
Washington for study. Portions of this material are reproduced 
as PLATE 749, Fics. 1, 2 and 3. They certainly belong to the 
species which has erroneously passed as M. mexicana; and since 
the name Agrostis foliosa Roemer & Schultes was nomenclaturally 
a substitute for A. frondosa Poir. and, therefore, illegitimate, it 
is fortunate that the Roemer & Schultes type belongs so un- 


equivocally to the species which I take to be A. frondosa of 
Poiret. 


1943] Fernald,—Rhizomatous Species of Muhlenbergia 231 


M. sYLVATICA 


MUHLENBERGIA SYLVATICA TORR. IS, APPARENTLY, CORRECTLY 
* ° ° . * 

UNDERSTOOD! A. northeastern variety of it will be published on 
a succeeding page. ‘ 


M. raceMosa (PLatsEs 753 and 754) 


Muhlenbergia racemosa (Michx.) BSP., on the other hand, has 
been quite misinterpreted. This misinterpretation arose through 
the fact that most specimens reaching the European herbaria and 
the plant best known to botanists of Cambridge, New York and 
Washington is the very different M. setosa (Spreng.) Trin., with 
erect, slender and usually simple culms with scabrous-puberulent 
internodes, typical of bogs, wet meadows and shores and doubt- 
less suggesting to the manufacturer of “English” names the pseu- 
donym ‘‘Marsh Muhly”, commented upon on p. 222. Agrostis 
racemosa Michx., basis of the very recent and by its authors not 
understood combination M. racemosa (Michx.) BSP. Prelim. 
Cat. N. Y. 67 (1888), ‘“Presumably based on Agrostis racemosa 
Michx.”’ (Hitchcock), is a wholly different species of the interior of 
the continent and almost unknown in herbaria prior to 1850. It 
was finally recognized as M. glomerata, var. ramosa Vasey, Descr.. 
Cat. Grasses U. S. 40 (1886) but Vasey did not see the really 
distinctive characters. 

Muhlenbergia setosa (or glomerata) has the culms unbranched 
or with few erect basal branches; opaque and scabrous-puberulent 
internodes; leaf-sheath with rounded midrib and minute usually 
hidden ligule; conspicuous anthers 1-1.5 mm. long; oblong- 
cylindric grain easily freed and 1.2-1.5 mm. long; the species 
with transcontinental boreal range and intrusions southward, 
east and west, in paludal habitats. M. racemosa, on the other 
hand, is a plant of the dry interior, often ‘‘in the dryest of soil’, 
with culms usually stiffly branched at the middle nodes; the 
internodes lustrous and glabrous; the leaf-sheath keeled; the 
ligule prolonged; the anthers only 0.5-0.8 mm. long; the tightly 
embraced linear-cylindric grain 1.8-2.2 mm. long. 

Michaux’s original description was too general to note any of 
the specific points, and his habitat, “in ripis sabulosis inundatis 
fluminis Mississipi” inconclusive. The photograph of the type 
before me shows, however, the high-branching culm and the 


232 Rhodora [JUNE 


characteristic stiffly glomerulate-spiciform panicles of M. glomer- 
ata, var. ramosa; and examination by me of the Michaux speci- 
mens in 1930 revealed the glabrous internodes and tiny anthers. 
The name Muhlenbergia racemosa (Michx.) BSP. should be 
restricted to the plant of the interior later described as M. 
glomerata, var. ramosa. 


M. setosa (PLatTEs 755-757) 


Muhlenbergia glomerata (Willd.) Trin. Gram. Unifl. 191, pl. 5, 
fig. 10 (1824) rests upon Polypogon glomeratus Willd. Enum. PI. 
87 (1809), which was described in great detail, the simple diag- 
nosis and accompanying statements being 

*2, POLYPOGON glomer 

P. panicula pear ‘subspicata, ealycibus subulatis scabris, culmo 

adscendente bas os 

Agrostis festavaides ‘Mi ahlenbeng in litt. 

Habitat in America boreali 2. D. 

Since the plant was received from Muhlenberg, it presumably 
came from eastern Pennsylvania. The very long description 
following the diagnosis may be abbreviated. 

“Folia rigidiuscula Ligula brevissima Vaginae foli- 
orum solutae. Pantouta glomerato-spicata obtusiuscula, ramis brevis- 
simis adpressis flexuosi eae Gye... 4 ristatus corolla lon- 
gior, nervo medio viridi, una cum arista scabro . . . Basis corollae 
pilis aliquot est instructa Habitu accedit quodammodo 4 
Agrostidem mexicanam, sed recedit ab hoc genere calyce aristato, et est 
vera Polypogonis species.” 

The type, for a photograph (our PLATE 756, FIG. 1) of which I 
am indebted to Professor Diels, consists of four plants of the 
characteristic species (MAP 1) growing in bogs and wet meadows 
from western Nova Scotia and southern Maine to southern 
Ontario and Michigan, south to southern New England, northern 
New Jersey, Pennsylvania, mountains to North Carolina, an 
Indiana. There is no question about the identity of M uhlen- 
bergia glomerata; but a second plant (map 2), also of simple habit 
or branched usually from the base, with puberulent internodes, 
large anthers and short grains, occurs farther north, from New- 
foundland to Alberta, south to Nova Scotia and Maine, northern 
Massachusetts, Connattioat: New York, central Pennsylvania, 
Ohio, Michigan, Wisconsin, Minnesota, Wyoming, Nevada and 
Oregon. This is Daitylopiesene cinnoides Link, Hort. Berol. il. 


1943] Fernald,—Rhizomatous Species of Muhlenbergia 233 


Map 1 (upper), — Lipids of MUHLENBERGIA SETOSA; MAP 2 (lower), of 
M. SETOSA, var. CINNO 


234 Rhodora JUNE 


248 (1833), described in great detail from plants raised from seed 
from northwestern Canada sent by Dr. John Richardson. The 
genus Dactylogramma Link was based on this material, Link 
separating it from Cinna. From the very full description the 
following points are drawn: 
“Caulis erectus parum ramosus . . . Panicula tenuis glomerata. 
Valvae subaequales longissime oe quasi aristatae, ene longi- 
res. Valvu ongis adpressis ad margin 
ee sub repens, th pedes altus . . . inferne vilis brevis- 
imis pubescens gula brevis truncata. Panicula 2-4 poll. 


fone: sar subdistantibus Seeeviaits superne approximatis, ramulis 
conglomeratis 


The description is vivid; the photograph (our PLATE 757, 
FIG. 1) of Link’s type sent me by Professor Diels unequivocal. 

The more northern Dactylogramma cinnoides differs in several 
characters from the relatively southern Muhlenbergia glomerata. 
In the latter the flowering culms bear 7-15 leaves, many of them 
often crowded and overlapping at the middle nodes; in the former 
there are 5-8 (rarely —10) more scattered leaves. In M. glomeraia 
the panicle, as correctly defined by Willdenow, has round-tipped 
spikes mostly crowded to form a subcontinuous lobulate-spiciform 
inflorescence; in D. cinnoides the spikes or branches, except the 

upper, are more scattered or subdistant and less rounded at 
summit. In M. glomerata the inflorescence is often purplish, 
sometimes green; in D. cinnoides green, though sometimes 
purplish. The panicle of M. glomerata has a misty appearance, 
the awns and glumes not sharply visible. This comes from the 
copiously hispid keels and awns. In D. cinnoides the panicle 
has the glumes and awns more clearly visible, the keels and awns 
being merely scabrous with definitely shorter trichomes. These 
differences, with no appreciable difference in anthers and grains, 
seem to me varietal, rather than specific, and I am treating D. 
cinnoides as a northern geographic variety of the more southern 
plant. 

Unfortunately, however, there is an earlier valid specific name 
for true Muhlenbergia glomerata (Willd.) Trin., which was based 
on Polypogon glomeratus Willd. (1809). This is Polypoge” 
setosus Spreng. Mant. Fl. Hal. 31 (1807). That it is specifically 
identical with the Muhlenberg material which formed the basis 
of Willdenow’s species is clear from the description; the photo 


1943] | Fernald,—Rhizomatous Species of Muhlenbergia 235 


graph of Sprengel’s type (our pLATE 755, Fic. 1), for which I am 
also indebted to Professor Diels, is certainly of the plant of 
eastern Pennsylvania. Sprengel’s description was to the point: 

9. PoLypocon seETosus, panicula spicaeformi, glumis acuminatis 

aristatis scabris, foliis involutis rigidis. 

ulmus teres, glaber, foliosus, pedalis. Folia glabra, vaginantia, 
involuta, rigida. Panicula terminalis, parumper ex ultimo folio 
emergens, fusca, densé spicaeformis, ramulis glomerstis contractis. 

Gluma calycina valvulis acuminatis scabris aristatis, uniflora, corolla 

paullo maior. Gluma corallina valvulis acuminatis muticis. 

Pensylvania. Muhlen 

In his De Graminibus Unifloris et Sesquifloris, 195 (1824), 
after properly transferring several species to Muhlenbergia, 
Trinius took a short cut and merely stated that certain species 
belonged to that genus: “Ad sectionem a (Muiihlenbergiam) 
pertinent: . . . Polypogon setosus Spreng.’’, but the combi- 
nation, ascribed to Trinius at this place and with the correct 
synonym, M. glomerata, was made in Index Kewensis. The 
species with culms simple or sparingly branched at base and with 
scabrous-puberulent internodes, large anthers and small grains 
is, then, M. serosa (Spreng.) Trin. ex Hook. f. & Jackson, Ind. 
Kew. ii. 209 (1894), 

Summarizing this rather verbose but seemingly necessary dis- 
cussion, the five species of Muhlenbergia specially considered 
resolve themselves as follows (omitting M. sylvatica which, 
except for the designation of a variety, is unchanged). In order 
to clarify the complex questions I am reproducing photographs 
of types in so far as they are before me and sufficiently clear for 
reproduction. These I am supplementing by some of Dr. Bernice 
G. Schubert’s very clear enlargements of pertinent details. 

MUHLENBERGIA frondosa (Poir.), comb. nov. Agrostis frondosa 
Poir. in Lam. Enecye. Suppl. i. 252 (1790). A. foliosa 
Schultes, Syst. ii. 373 (1817), substitute-name for A. frondosa. M. 
foliosa (Roem. & Schultes) Trin. Gram. Unifl. 196 ve ioe a re- 
cent auth. “ML mexicana sensu most recent auth., not (L.) T 
For cee characters see no. 1 in key. PLaTEs 749 and 750. 

M. nposA, forma commutata (Scribn.), comb. nov. M. 
Bepeer ypeenanr (as subsp.) gee in Ruopora, ix. 18 
(1907), at least as to TrPE, Fernald, 522 , designated by 

Hitche. Man. 890 Ftcaide where Sedbnar’ 8 T tinal ““n. subsp 
is cited as ‘“‘var.’’ M. mexicana, var. ee (Scribn.) Fava 
in Rep. Mich. ata “Sei. xvii. 181 (1916). M. commutata 


236 Rhodora [JUNE 


(Seribn.) Bush in Am. Midl. Nat. vi. . tba M. mexicana, 
forma commutata gee Wiegand in Ruopora, xxvi. 1 (1924). 
M. mexicana (L.) Trin. Brae. Unifl. 189 (1824). Ageay 

mexicana L. Mant. i. 31 (1767); Jacq. f. Eclog. Pl. Rar.-Gra 
Fase. ter. et quart. 44, t. 80 (1813). For other synonyms based 
on A. mexicana see _Hiteheoe k. M. foliosa sensu most recent 
auth., not (R. & S.) Trin. For diagnostic characters see no. 2 in 
a7 ’PLates 751 and ue 

MEXICANA forma ambigua (Torr.), comb. nov. Agrost? 
fgorm acne een ap i. 95 (1809). Cinna filiformis (Willd.) 

Enum. i. 70 (1821). Se ery lateriflora, var. filiformis 
cwiia) Torr. Fl. N. Mid. U.S. 86 (1823). M. ambigua Torr. in 
Nicollet, Rep. Miss. 164 (1843). M. mexicana  fliformis (Willd.) 
Scribn. in Mem. oe rr. Bot. Cl. v. 36 (1894). M. foliosa ambigua 
(Torr.) Seribn. Ruopora ix. 20 (1907). M. ambigua, var. 
ere (W ilid.) Farwell in Mich. Acad. Sci. Rep. xx. 168 
(1919). M. foliosa, forma ambigua (Torr.) Wiegand in RHoDORA, 
xxvi. 1 (1924). 

M. MEXICANA, aes se (S. Wats.), comb. nov. M. 
sylvatica, var. setig lumis 8. Wats. in King, Geol. Expl. 40th Paral. 
v. 378 (1871). M. foliosa sete aries (S. Wats.) Seribn. in RHopo- 
ix. 20 (1907). <p setiglumis (S. Wats.) Nels. & Macbr. in 

t. Gaz. lxi. 30 (1916). 

mir SYLVATICA Torr., var. robusta, var. nov., culmis rigidis 
arcte adscendentibus; foliis firmis ad 5-9 mm. latis; paniculis 
densioribus; ~ is late lanceolatis vel lanceolato-ovatis; lem- 
matibus 3-4 longis eo aristatis; a 0.5-0.7 m 
longis; ppiiseet a 1.9-2.1 mm. longis.—Open woods and 
thickets, central Maine to western ee York and eastern 
Pennsylvania. The following are characteristic. MAINE 
Austin Stream, Moscow, August 27, 1902, Collins & Chamberlain; 
Sydney, August 18, 1916, Fernald & Long, no. 12,597 (TYPE in 
Herb. Gray). New HAMPSHIRE: by Merrimac River below Bed- 
ford, August 27, 1931, Fernald & Griscom, no. 24995; Gis 

Y 


Batchelder. Vermont: Grand Isle, July 24, 1935, Knowlton. 
Massacuusetts: Needham, August’ 31, 1884, 'T. O. Fuller; Blue 
Hills Reservation, September 22, 1895, E. F. Williams; Hunting- 
ton, August 17, 1912, Robinson, no. 358. RuopEe ISLAND: 
Lincoln, October 2, 1910, E. F. Williams. Connecticut: 
Sprague, September is 1905, Woodward; Reynold’s Bridge, 
pune 4, 1910, Blewitt; Bridgeport, September 22, 1890, 
Eames. New York: Starbuckville, fc, ok 26, 1932, Muenscher 
& Lindsey, no. 2954; Genoa, August 21, 1918, Wiegand, no. 9185; 
Ithaca, September ‘5 191 16, Be ag *Metealt no. 5613; Butler, 
October 5, 1916, Metcalf & "Wright, no. 5614. PENNSYLVANIA: 
Wayne, October 15, 1910, Bartram, no. 1313. 


1943] Fernald,—Rhizomatous Species of Muhlenbergia = 237 


Typical Muhlenbergia sylvatica has relatively weak and loosely 
ascending culms; the leaves subflaccid, the larger ones 2-7 mm. 
wide; the panicle very loose; the glumes linear-lanceolate to 
linear-attenuate, the 2d about equaling to slightly exceeding the 
blade of the lemma, the latter 2.3-3 mm. long; the anthers 
0.3-0.6 mm. long; the mature grain nearly or quite free, 1.4-1.8 
mm. long. It is widely distributed, from southwestern Quebec 
to Minnesota, south to North Carolina, Alabama, Arkansas and 
northeastern Texas. 

Var. robusta is stiffer and larger in all parts. Its leaves are 
firm and more ascending, the larger ones 5-9 mm. broad; the 
panicle fuller; glumes broadly lanceolate to lance-ovate, shorter 
than the blade of the lemma, the latter 3-4 mm. long; anthers 
0.5-0.7 mm. long; grain more firmly embraced and 1.9—2.1 mm. 
long. 

M. racemosa (Michx.) BSP. Prelim. Cat. N. Y. 67 (1888), by 
inference. Agrostis racemosa Michx. Fl. Bor.-Am. i. 53 (1803). 
Polypogon racemosus (Michx.) Nutt. Gen. i. 51 (1818). Cinna 
racemosa (Michx.) Kunth, Rév. Gram. i. 67 (1829). M. glom- 
— var. ramosa Vasey, Descr. Cat. Grasses U. 5. 40 (1885). 

. racemosa, var. ramosa Vasey in Beal, Grasses N. Am. ii. 253 
Cinees For diagnostic characters see no. 4 in key. PLaTEs 753 
and 7 vi 

M. seTosa (Spreng.) Trin. ex Hook. fil. & Jackson, Ind. Kew. 
ili. 209° pe 4). Polypogon setosus Spreng. Mant. Fl. Hal. 31 
(1807). poe Willd. Enum. 87 (1809). Alopecurus 
glomeratus (Willd.) Poir. in Lam. Encyel. v. 495 (1817). Agrostis 
setosa (Spreng.) Muhl. Descr. Gram. 68 (1817). Trichochloa 
glomerata Trin. Fund. Agrost. 117 (1820). T. calycina Trin. 
1. ec. (1820). M. glomerata (Willd.) Trin. Gram. Unifl. 191, pl. 
5, fig. 10 (1824). M. calycina Trin. 1. c. 193 (1824). Podosaemum 
glomeratum (Willd.) Link, Hort. Berol. i. 84 (182 7). Cinna 
glomerata (Willd.) Link, l. ¢. ii. 237 (1833). For specific char- 
acters see no. 5 in key. —Typical M. setosa has the leaves of 
flowering culms 7-15, many of them often browded and over- 
lapping at the middle nodes; panicle purplish, fuscous or green, 
with all but the lowest of the densely flowered ellipsoid to 
rounded-obovoid branches closely crowded, the panicle thus 
appearing densely lobulate-spiciform; keel and awns of glumes 
copiously hispid, thus giving the inflorescence a ‘“‘misty”’ aspect.— 
Meadows, bogs and wet shores, western Nova Scotia and south- 
ern Maine to southern Ontario and Michigan, south to southern 
New England, northern New Jersey, Pennsylvania, mountains 
to North Carolina, and Indiana. PLaTEs 755 and 756; Map 1. 


238 Rhodora [JUNE 


Var. cinnoides (Link), comb. nov. Dactylogramma cinnoides 
Link, Enum. Hort. Berul. ii. 248 (1833). Bir ae from typical 
M. setosa in having fewer (5-8, rarely —-10) and usually more 
scattered leaves; phiidis usually green, ibe purple-tinged, 
usually more interrup ted, the cylindric to oblong-ovoid often 
iabenis lower oenehine often remote; glumes slightly broader, 
with merely scabrous keel and awn (the panicle, therefore, not 
appearing “misty’’).—Similar habitats, Newfoundland to Al- 
berta, south to Nova Scotia, northern Massachusetts, Connec- 
ticut, New York, central Pennsylvania, Michigan, Wisconsin, 
Minnesota, Wyoming, Nevada and Oregon. PuLate 757; MAP 


Puate 749. MuUHLENBERGIA FRONDOSA (Poir.) on Fics. 1, 2 and 3, 
fragments from Type of Agrostis foliosa Roem. & Schultes, x 5, from Syn ions 
of type in Herb. U.S. Nat. Mus., kindness of Mr. Jas ae. Swallen; FIG. 4, 
two spikelets, x 4, from Bridgeport, Connecticut, Sse ae 7, 1898, Eames; 
FIG. 5, floret, x 10, fr prs Shirley, Massachusetts, October 4, 1914, J. Bt. 
as syn FIG. 4 grain, < +10, from last re men. 

IUHLENBERGIA FRONDOSA (Poir.) Fern.: ric. 1, a character- 
istic. pacha Ee — Pes , from East Jaffrey, New Hampshire, jes 
nson, no. 3 characteristic glabrous internode, open and ¢ 
pressed shea aad: ak pores vga $ panicle, X 5, from Stratford, 
New Hampshire, Pease, no. 23, ee portio n of. rhizome, x 4, to 
eusiiintearctin: scales, from no. 28 f 
Puate 751. MuHLENB BERGIA aos aga A (L.) Trin.: Ftc. 1, type of AGROSTIS 


e ‘ . 
649; Fia. 3, floret a; 10, from St. Tiahert. oo: Victorin & Rollan d, no. 
,968; Fic. 4, gr 10, from shore of re River, Sussex County, 
New Jersey, Siematec 15, 1917, E. B. Daten 
PLatTE 7 Aas UHLENBERGIA RACEMOSA (Michx.) BSP.: Fic. 1, TYPE (two 
piece) x? x Ko Michx from Herb. Michaux 
sd of Professor H ene FIG. 2, panicle, X 1, from Menard 
Connie Tikva, 1861, HE. Hall; F 3 _Panicle, from near Princeville, 
Peoria County, Illinois, V. H. Chase 940; ric. 4, spikelet, X 10, from 
Fort Snelling, Minnesota, M earns, no. 747: ; FIG. 5, floret, showing basal villi, 
x 10, from no. ; Fic. 6, grain, X 10, from near Oquawka, Illinois, Patterson. 
[UHLENBERGIA RACEMOSA (Michx.) BSP.: Fic. 1, a charac- 
teristic summit of a plant, x 1, from Fort Sn ailing. Signi bina M Le gs no. 
747; FIG. oe _sumnit of Eisbrous and peers internode and csc mm of 
sheath, from La Wisconsin, 1861, T. J. Hale, ria. 3, portion of 
_ nicle, x 2? to show the small ake gm near setae Taig Peoria County; 
llinois, V “Ae Chase, no. 940. 
E MUHLENBERGIA sETOSA (Spreng.) Torr.: Fic. 1, TYPE of 
Polypogon setosus Di eae of abe species, X ca. 14, from Herb. Berol., 
re 


fl 
& 


r7,1 Church ngs 3 panic cle, X 1, sae northwest of 
Piel Valley, Bake t County, nnsylvania, Sie at r 15, 1923, Benner; 
n 


LE: LR TT TLE 
i oepeepenenmmmmenmmnetiilineneenmenmmnsmenn tee 


1943] Fernald,—Notes on Danthonia 239 


2, 1883, Walter Deane; Fic. 6, group of Fs sassiyghal x 10, yas ieee Pond, 
West Roxbury, Massachu setts, C. &. FIG. 7, flor 10, showing 
few slender basal villi, from Mt. Bethel, Noctuinrkoe Co sake Big ennsylvania, 
October 4, 1908, Van Pelt; ria. 8, grain, x 10, from the last (Van Pelt) speci- 


m 
Piate 756. MUHLENBERGIA seTosA (Spreng.) Trin.: Fic. 1, TYPE of 


internode and node and base of sheath, x 5, from northwest of Pleasant 
Valley, Bucks County, Pennsylvania, September 15, rag ' 

LATE . MUHLENBERGIA SETOSA (Spreng. (ae NNOIDES 
(Link) Fern.: ric. 1, rypr of Dactylogramma cinnoides tisk, basis of the variety, 
x : 


portion of panicle, to show elongate anthers (as black ‘lines), x 4, fto m Fort 

‘ent, Maine, August 21, 1913, R. W. Wo pig ; FIG. oret, to show villi 
running high on lemma, X 10, ‘from no. 4531; Fig. 5, grain pA 10, m River 
Ste. Anne des Monts, Gaspé County, Quebec, pines 1905, Collins & Pani. 


II. NOTES ON DANTHONIA 


The two common species of Danthonia in eastern Canada, 
New England and New York are in general well defined, although 
the conventional differential characters of the lemma used by 
Hitchcock in Gray’s Manual, ed. 7, and by Nash in Britton & 
Brown’s Illustrated Flora are essentially valueless. As expressed 
by Hitchcock these are: 


“Teeth of the lemma bers eg, Not Aree. oo ek 1. D. spicata. 
Teeth of the lemma aristat 
Flowers not over 5 mm, hee ck MUS Va Cosh lieon eine es 2. D. compressa.” 


Under the specific treatments the lemmas of D. spicata are said 
to be ‘4-5 mm. long, sparsely clothed with stiff hairs, teeth 
triangular’’, while D. compressa has the “‘teeth of the lemma aristate, 
at least 2 mm. long.” In practice this character so frequently 
breaks that it has been abandoned by later authors, and in 
his Manual Hitchcock relies upon the contracted panicle and 
relatively short blades of D. spicata as contrasted with the more 
open panicle and longer leaves of D. compressa. 

The best key I have seen is that of Wiegand & Eames in their 
Flora of the Cayuga Lake Basin. For wholly typical D. spicata 
and D. compressa it is satisfactory; the difficulty is that so much 
material is not typical. In the series which everyone would 
accept as D. spicata in the broad sense, the lemma varies from 
2-6.5 mm. long and its back may be densely pilose, sparingly 
pilose, strigose or quite glabrous, its teeth from broadly triangular 
to lance-attenuate and awnless or awned. Such variation is 


240 Rhodora [JUNE 


largely without apparent geographic limit, though two well 
defined geographic varieties, var. pinetorum Piper and var. 
longipila Scribn. & Merr., are strongly marked; the former 
boreal and transcontinental, with lemmas often so large as to be 
mistaken for those of D. intermedia Vasey; the latter with the 
shortest lemmas of the series and a definitely southern range. 
The sheaths and blades of any of the variations may be glabrous, 
sparingly pilose or even long-villous; and to the latter tendency, 
without checking the original account, has been erroneously 
applied the name var. longipila. If forms of the common north- 
ern plant with pubescent foliage are to be distinguished, they 
have an earlier name, evidently intended for them, in D. spicata, 
var. villosa Peck (1894). 

My chief reason for now publishing this note is to direct atten- 
tion to a complex series of very large plants which in some ways 
stand between Danthonia spicata and D. compressa but which, 
in extreme development, have been mistaken for the south- 
eastern D. sericea Nutt. These are the coarse midsummer- or 
autumn-flowering plants with stout culms up to 1 m. high; long, 
stiff and mostly erect basal leaves; panicle as in D. spicata but 
with more numerous and larger spikelets; glumes up to 2.5 cm. 
long; and lemmas up to 6.5 mm. long; these and the long glumes 
often leading collectors to place New England specimens wit 
the more southern D. sericea. In this complex and mostly north- 
ern coarse series the culms may be terete to base, as in D. spicata, 
or the lower internodes may be trigonous or laterally compressed 
and with the narrower side broadly concave, as in D. compressa. 
The uppermost leaf may be short and remote from the long- 
exserted panicle, as in D. spicata, or (even from the same root) 
prolonged and reaching or overtopping the panicle as in D. 
compressa. The twisted base of the awn may be dark brown to 
purplish, as in D. spicata, or stramineous or pale, as in D. com- 
pressa. ‘The panicle is more like that of D. spicata but large and 
dense. Furthermore, whereas D. spicata (typical) flowers in 
New England from late May to early July and D. compressa in 
June and July, the coarse plant flowers chiefly from August to 
October. If culms with trigonous to compressed lower inter- 
nodes be selected they will belong to D. Alleni Austin (1872), 
while other culms (often in the same tussock), with the inter- 


TT LTT, 


1943] Fernald,—Notes on Danthonia 241 


nodes terete, will be D. Faxoni Austin (1877). It might be 
assumed that D. Alleni (including D. Faxoni) is a series of hy- 
brids, with unusual hybrid vigor; but it is found to the northeast 
of the limit of D. compressa, while to the northwest it extends 
450 miles north of the northwestern limit of the latter species. 
If sometimes a hybrid it is not always so. 

In many of its stations Danthonia Alleni occurs in recent clear- 
ings or burns where the loose litter and often the ashes from fires 
stimulate all plants; in such cases it appears like overstimulated 
specimens, in which rankness of growth, multiplication of 
spikelets and enlargement of their parts are the result. In other 
cases the autumnal coarse culms and panicles suggest the second 
flowering in many species of Carex, where autumnal inflorescences 
are larger, fuller and often more crowded than the normal 
vernal ones. Again, the panicles of D. Alleni may be greatly 
distorted and with apparent signs of fungus- or insect-attack, 
with gnarled or crumpled branches, tendencies to fasciation, 
and sometimes the gathering of spikelets into glomerules. All 
in all, D. Alleni is a heteromorphic series well worth close obser- 
vation by those situated to watch it. That it is a true species is 
very improbable. It is presumably the result of very diverse 
conditions which have resulted in parallel developments; and, 
in view of the well known development within the leaf-sheaths 
of this genus of cleistogamous florets, it is not improbable that 
every alteration of D. spicata and D. compressa, whether by 
crossing through wind-pollination or stimulation in other ways, 
may be somewhat perpetuated through the cleistogenes. Simi- 
larly the very diverse lemmas of D. spicata, whether with awnless 
or awned teeth triangular or lanceolate, the backs pilose or 
glabrous, may thus be carried on in local colonies. The genus is 
an appropriate one for close study from many viewpoints. 

I am distinguishing the two undoubted species and the wholly 
dubious Danthonia Alleni by the following characters. 


0.4-1.5 mm. long; panicle-branches ascending to erec 


242 Rhodora [Jone 


dark brown to es strongly contrasting with the paler 
and straightish su 
Culms slender, 0. ru 5 mm. thick (dried) at base, 1-6 dm. 
, with terete internodes es; panicle remote from et 
leaf, ‘mostly with 213 spikelets; toegti glume 7-11 m 
long, if longer with faint or obscure lateral nerves; hale 
Ol i a ee ee 1. D, spicata. 
Culms stout, 1.5-2.5 mm. thick at saps mostly 0.5-1 m 
high, the lower internodes terete, or triangular and with 
one concave side; panicle either hah or closely sub- 
tended by uppe r le af, dense, u wee ith 9-20 spikelets; 
longer glume batine attenuate 1-2.5 em. long, promi- 
nently 3-7-ribbed; base of awn ‘tieht to dark brown... .2. D. Alleni. 
a. valne slightly geniculate at the nodes, the summit usually 
ing; some of the lower intern odes s trigonous 
a te and often with the narrower side broadly concave; 
leaves prolonged, one half as lo i 
culm; lower cauline leaves prolonged, 1.5—4 dm. long, the 
uppermost nearly reaching or overtopping the panicle; 
e 


pressed in fruit; spiraling base of ect pale reatbont ae stra- 
Ci ekce AQ Peet ee eet eras OO AAS ee 3. D. compressa. 


. D. spicata (L.) Beauv. consists af three fairly defined and 
ee less definite variations, as follow 


a. Be art florets three fourths to quite as long as the 
eas parents with (1—) 2-13 (-15) sikeiets 
loan leaves t or involute; culms firm, 0.5-1. 
thick at bas 


Glumes lance-attenuate, tapering from near base 


cescent leaves strongly curving and twisting 

a var. typica. 

Glumes casigac ire again oe from near 
above with wi 


[ries or er ane eee r. pinetorum. 
a. Colatan ‘of florets only one half to three fifths as lo Pies 
as the thin and hyaline domi or only faintly ribbed 
shanian: spike ached 3-7, ttered; culms delicate, 
0.5-1 mm. thick a hen r taead leaves filiform- 
involute or Lesonies We, Outed 6 eS Var. longipila. 


D. spicata (L.) Beauv., var. typica. Avena spicata L. Sp. Pi. 
80 (1753). A. glumosa Michx. Fl. Bor.-Am. i. 72 (1803)- 
D. nee Seago Beauv. Ess. Agrost. 92, 153 and 160 (1812). 
D. spicata 02) Basa. oe ean & Schultes, Syst. ii. 690 (1817)- 
Mer athena spicata (L.) Raf. ex Hook. f. & Jacks. Ind. Kew. i. 


1943] Fernald,—Notes on Danthonia 243 


211 (1894) in synonymy. D. spicata, var. villosa Peck in N. Y. 
State Mus. Ann. Rep. xlvii. 168 (repr. 42) (1894), the form with 
villous blades. Pentameris spicata (L.) Nels. & Macbr. in Bot. 
Gaz. lvi. 470 (1913).—Dry to damp and peaty soils or in thin 
woodland, southern Quebec to Minnesota, south to Nova Scotia, 
New England, Long Island, northwestern Florida, Alabama, 

Tennessee and ee May-ea 
Var. PINETORUM Piper, as D. pee de pinetorum Piper in Ery- 
thea, vii. 103 (1899), deaniibad 4 in detail, although its author did 
not appreciate the characters, saying of it‘ ‘scarcely more than a 
variety of that species [D. spicata], differing mainly in character 
of pubescence.” D. thermalis (as og Seribn . Dept. 
Agric., Div. Agrost. Cire. no. 30: 5 (1901). M rerathrepta pineto- 
rum (Piper) Piper, Contrib. U.S. Nat. Herb. xi. 122 (1906). M. 
12 


ex Fedde & Schust. in Just, Bot. a Biot Xxxvii. 128 (1911). 
Pentameris thermalis (as thermale) (Seribn.) Nels. & Macbr. I. ce. 
(19138). -D. that lc (Piper) Piper in Piper & Beattie, Fl. Nw. 
Coast, 46 (1915)—Dry to moist open soil, Newfoundland and 
Céte Nord, Quebec, to British Columbia, south to Nova Scotia, 
ew Brunswick, northern New England, Bruce Peninsula, 
Ontario, northern Michigan, northern Wisconsin, Black Hills, 
South Dakota, New Mexico and Oregon.—Since this variety has 
not been heretofore recorded from east of northern Michigan a 
few characteristic eastern specimens of it may well be cited. 
NEWFOUNDLAND: Glenwood, Fernald & Wiegand, no. 4600; 
Grand Falls, Fernald, Wiegand, Bartram & Darlington, nos. 4601 
and 4602 : St. John’s Island, Fernald et al. no. 27,466; Middle 
Arm, Bonne Bay, Fernald, Long & Fogg, no. 1300; North Arm, 
Bay ‘of Island, Long & Fogg, n no. 74; French (or Tweed) Island, 
Bay of Islands, Fernald, Long & Fogg, no. 76; Blomidon (‘ ‘Blow- 
me-down’’) Mountains, "Fernald & Wiegand, nos. 2591 and 2595; 
Mt. Musgrave to Humber Mouth, Fernald & Wiegand, no. 2590. 
QueBEc: Mingan, Si. oe no. 90,140; Anticosti Island, Victorin, 
or Victorin et al. no . 4077, 20, 536 and. 20,537, 20, 542, 20, 598, 
24,344-24 "346, 27, 795, 27, 797, 27, 798, 28, 050, * 055 and 28, ‘060: 
Pereé, Fernald & ‘Collins, no. 870; River Ste. Anne des Monts 
Fernald & Collins, no. 413; Bic, Fernalds & Collins, no. 869: 
Riviére du Loup, Victorin, nos. 22 and 81; Montmorency Falls, 
Macoun, no. 69,231; Saint-Romuald, Louis-M arie, no. 20,541. 
NEw BRUNSWICK: Bass River, July 30, 1875, Fowler; Shediac 
Cape, F. T. Hubbard, no. 720. Nova Scorta: St. Paul Island, 
Perry & Roscoe, nos. 64-66; ee aaa oe Pease & Long, 
no. 19,953; Argyle, Long & Linder, no. 19,957; Abram River, 
Fernald, Bean & White, no. 19,956. Macie: at 4000-4500 ft., 
North Basin, Mt. Katahdin, July 14, 1900, Fernald; Township 


244 Rhodora |\JUNE 


ix, Range 17, Somerset County, St. John, no. 2134; Madison, 
August 21, 1894, Fernald. New HaAmpsHireE; near Half-way 
House, Thompson & Meserve Purchase, Pease, no. 12,817. 
VERMONT: Willoughly, July 11, 1896, July 18, 1898 and Septem- 
ber 14, 1898, G. G. Kennedy. 

Var. LoncipiLa Scribn. & Merr. U. S. Dept. Agric., Div. 
Agrost. Cire. no. 30: 7 (1901)—Sandy or rocky woods and 
clearings, North Carolina and Alabama to New Mexico, north to 
Connecticut, Pennsylvania, Kentucky, Missouri and eastern 
Kansas. Late May-September, The following are characteris- 
tic. Connecticut: Woodbury, July 13, 1932, Eames. PENN- 
SYLVANIA: Philadelphia, 1862, C. EZ. Smith. Vircinra: eastern 
slope of Bull Run Mountain, Prince William County, Allard, no. 
3850; south of Aldie, Fauquier County, Allard, no. 859; 4 miles 
northwest of Dixie Caverns, Roanoke County, Wood, no. 2609; 
Great Neck, Princess Anne County, Fernald, Griscom & Long, 
no. 4553; Claremont Wharf, Surry Co., Fernald & Long, no. 
9829; Carey Bridge, Southampton County, Fernald & Long, no. 
11,950; Cypress Bridge, Southampton County, Fernald & Long, 
no. 11,949. No Caroutna: Locust, Stanly County, Blom- 
quist, Correll & Garren, no. 7753; Highlands, Macon County, 
Biltmore Herb., no. 343°. Kenrucxy: Hawesville, Hancock 
County, E. J. Palmer, no. 17,804. AtaBama: Mobile, C. Mohr, 
as D. sericea. Muissourt: Indian Creek, Benton County, De- 
metrio, no. 87; Mansfield, Lansing, no. 3103. Kansas: Chero- 
kee County, Hitchcock, no. 905. OKLanoma: base of Rich Mt., 
near Page, Leflore County, Stevens, no. 2674. Texas: Dallas, 
Reverchon; Terrell, Kaufman County, May 4, 1904, F. J. Tyler. 
New Mexico: 1847, Fendler. 


When they published var. longipila, from Arkansas, Scribner & 
Merrill laid stress on the “scattered ascending hairs”, on the 
leaf-blades, a character which appears in the other varieties of 
the species and which may be absent from much material of 
otherwise good var. longipila. The distinctive features of this 
southern extreme are the very slender culms, usually filiform- 
involute leaves, and the small florets in relatively short columns, 
characters brought out in the original description: “A slender 
form . . . with small, few-flowered panicles and smaller 
spikelets than in the species. Leaf blades very narrow, involute 
: Flowering glumes 2 to 2.5 mm. long, pilose.” 

2. D. Auuent Austin in Bull. Torr. Bot. Cl. iii. 21 (1872)- 

_ D. Faxoni Austin, |. ¢. vi. 190 (1877).—Open shores, rocky °F 
arid openings, clearings and burns, Magdalen Islands to Algoma 
District, Ontario, south to Nova Scotia, Maine, Massachusetts, 


1943] Fernald,—Notes on Danthonia 245 


Delaware, mountains of North Carolina, and Ohio. Late June— 
October.—The following, selected from twice as many specimens, 
are characteristic. QuEBEc: Baie St. Paul, Charlevoix County, 
Pease, no. 27,471. MaGpaLen IsLANDs: Grindstone, Fernald, 
Bartram, Long & St. John, no. 6870. Prince Epwarp ISLAND: 
Alberton, Fernald & St. John, no. 6869. Nova Scotia: Canso, 
Guysborough County, Rousseau, no. 35,452 bis; Bridgewater, 
Lunenburg ee aon al., no. 19,951. Marne: Houlton, 
Fernald & Long, no. - Orono, September 25, 1890, and 
September 1, 1893, Foca Peaked Mt., Clifton, "August 22, 
1897, Fe rnald; Pleasant Pond, Somerset "Co ounty, August 18, 
1902, Collins & Chamberlain; Chesterville, Kate Furbish; Oxford, 
July 12, 1914, Weatherby; Pleasant Mt., Denmark, Pease, no. 
19,628; Cutler, August 19, 1902, Kate Furbish; Pembroke, 
Fernald, no. 1309; Somesville, 4 Desert I., September 22, 
1892, Fernald; Southport, August 1894, Fernald; Bear Mt., 
Livermore, July 25, 1896, Kate Furbich: South Berwick, Parlin & 
Fernald, no. 034. New Hampsuire: Gorham, Pease, no. 17,337; 
Notch of the White Mts., August 27 and September 3, 1 877, C. 
E. Faxon, TYPE collection of D. Faxoni; Crawford Notch, 
Greenman, nos. 1276 and 1277; Crawford Notch, Hart Location, 
Pease, no. 11,727; base of Mt. Will ard, August 28, 1877, Faxon; 
North Woodstock, Woodstock, F ernald, no. 11 550; Plymouth, 
Fernald, no. 14, 995; Washington, Fernald & Svenson, no. 768; 
Nashua, Fernald & Svenson, no. 769; Pelham, 1895, F. W. Batchel- 
der. MASSACHUSETTS: Amesbury, 1897, TRS Eaton; Pepperell, 
August 30, 1886, C. W. Swan; Concord, September 8, 1858, 
Thoreau; Ashland, July 3, 1884, W. Deane; Savin Hill, Dor- 
chester, September 5, 1852, Wm. Boott; Plymouth, Fernald. no. 
an Brewster, Fernald, no. 17,948; Chatham, Fernald & Long, 
8751: N antucket Island, Bicknell, no. 9854; Shrewsbury, 
Es W. Robbins, as D. sericea Nutt.; Whitcombe Summit, Berkshire 
County, H. D. House, no. 25,893. New York: Babylon, Long 
Island, T. F. Allen, isotyres of D. Alleni, also Svenson, no. 6800; 
Rockaway, Long ‘Tsland, September 20, 1869, W. H. Leggett: 
New Jersey: Manasquan, June 26, 1929, J. R. Churchill. 
PENNSYLVANIA: Chester Valley, 2 miles north of Devon, Chester 
County, October 24, 1921, H. B. Meredith. DELAWARE: “WM. 
M.C Y, WILMINGTON, DELAWARE”. NortaCaro.tna: Roan 
Mt., July, 1889, Scribner. ONTARIO: Cloche Peninsula, Manitoulin 
District, Fernald & Pease, no. 3103; cliffs by Lake Superior, ood 
cake Bay, Algoma District, Pease & Ogden, no. 24,989. OxI 
near Garrettsville, Portage County, June 19, 1910, "A. N. Rood. 
. D. compressa Austin in Peck, N. Y Y. State Mus. Ann. Rep. 
xxii, 54 (1869). D. spicata, var. compressa (Austin) Wood, Am. 
Bot. and Fl. pt. 4: 396 (1873). Merathrepta compressa (Austin) 
Heller, Muhlenbergia, v. 120 (1909). Pentameris compressa 


246 Rhodora [JUNE 


(Austin) Nels. & Macbr. in Bot. Gaz. lvi. 469 (1913).—Wood- 
lands and clearings, southern Quebec to Ohio, south to Nova 
Scotia, New England, Long Island, Virginia, and upland of 
North Carolina and Tennessee. June- ugust. 


III. ERIANTHUS BREVIBARBIS AND OTHER SPECIES 
(Plates 758-761) 


“ecin n Elana sp. ea (TaB. 758), culmis rigidis 0.75- 
1.5 m. ad basin 3-6 m penne nodis 4-6 barbatis 
barbis dees foliis saute 4 1-6, vag is glabris, laminis scabris 
e basi valde angustat Oo eablaadeolnke Rapae ibus 2- nae mm. latis 
nerviis ee ikea she utrinque 3-5; lamina superiore 
valde reducta 4-12 cm. longa; panicula fai Gsolata densa 1-1.7 
dm. uae a em. diamnetrs basi deinde exserta, racemis valde 
adpressis 2-5 cm. longis; spiculis sessilibus lanceolatis, glumis 
strigoso-hirtellis 6.5-8 mm. lon ngis, coma basilari 4-5 mm. longa; 
pedicellis strigoso-hirtellis; arista tereti <taarees 1.6—2 cm. longa. 

—Delaware, eastern Maryland and eastern Virginia. DELAWARE: 
fencerow, x mile east of Ellendale, Sussex County, October 12, 

1940, R. R. Tatnall, no. 4745. Maryuanp: roadside 5 miles 
north of Princess Anne, Somerset County, October 2, 1937, R. R. 
Tatnall, no. 3574. IRGINIA: peaty swale (cut-over cypress 
swamp), about 4 miles northwest of Homeville, Sussex County, 
September 20, 1937, Fernald & Long, no. 7301, as E. brevibarbis 
Michx. (TYPE in Herb. Gray); alluvial woods along Nottoway 
River, Green Church Bridge, northwest of Owen’s Store, Sussex 
County, October 14, 1941, Fernald & Long, no. 13,884. All but 
the last distributed as 2. brevibarbis. 

Var. Elliottianus, var. nov., planta major; oye ad 2 m. altis 
ad basin 6-10 mm. diametro; laminis 7-12 mm. latis; panicula 
laxiore majoreque 2-4.5 dm. longa 4.5-10 em. ainiiettor © racemis 
4-8 em. longis—North Carolina to Florida. Type: thicket 
bordering pond near Live Oak, Florida, October 10, 1901, A. 
Curtiss, no. 6940, as E. alopecuroides, var. brevibarbis —The plant 
beautifully described as E. brevibarbis by Elliott, Sk. i. 39 (1816) 
and very crudely illustrated by him 

Erianthus coarctatus and var. Elliottianus have passed, ever 
since Elliott, as E. brevibarbis Michx. Nash in North Am. Fl 
xvi’. 93 (1909) cites for E. brevibarbis the ‘““Typr LOCALITY: 
Tennessee” and then gives the range “Delaware to Florida, west 
of Louisiana”; while Hitchcock, Man. 723 (1935) says ‘Moist 
places, Coastal Plain, Delaware to Florida and Louisiana”, but 
on p. 854 cites the type as from “Tennessee and Carolina, 
Michaux.” Michaux, in originally describing it, Fl. Bor.-A™- i. 


1943] Fernald,—Erianthus brevibarbis and other Species 247 


55 (1803), said “H AB. in collibus Tennassée et Carolinae’”’. The 
discrepancy in Michaux’s statement (and his label) and the 
Coastal Plain range of the plant usually taken to be EZ. brevibarbis 
has often been noted; and when Hitchcock examined the Michaux 
TYPE he wrote: 

Erianthus brevibarbis. Michx. 

“In collibus desertis ab amnio Wabash ad Ostium Missouri 5 diebus 
distantibus’’. The specimen belongs to the species described i in Small’s 
Flora under thisname. The range as originally published is “in collibus 
Tennassée et Carolinae”’. The known range is from Delaware south- 
ward along the coast to Florida, and west to Louisiana. We do not 
snow of its occurrence in southern Illinois, as given on Michaux’s 
abel —Hitche. Contrib. U. 8. Nat. Herb. xii. 151 (1908). 


Michaux’s type, with the label as quoted by Hitchcock, is here 
reproduced X 1%, as PLATE 759, Fic. 1. That it is very unlike the 
Coastal Plain plant for which it has passed is evident. Its 
apparently mature and disintegrated panicle is hidden amongst 
the broad and prolonged leaves and enlargements of the spikelets, 
FIG. 2, X 1%, show them to be like those of the plant (PLATE 760) 
of Pulaski County, Arkansas, distributed by Dr. Delzie Demaree 
(by creek near old quarry, Pulaski Heights, Little Rock, Septem- 
ber 23, 1931, Demaree, no. 8228). Like that of the Michaux 
type the panicle (PLATE 760, FIG. 1, X 14) of Demaree’s no. 8228 
is disintegrating. In equivalent latitudes of the Coastal Plain 
E. coarctatus sheds its fruit from mid-October into November. 
Since the Demaree plant is so like Michaux’s type from well up 
in the Mississippi Valley we may note that it has 10, instead of 
only 4-6 nodes; the leaf-blades broader (up to 1.5 em. wide) and 
with more numerous veins (the more prominent veins 6-8 each 
side of the midrib); the mature panicle partly included at base 
and greatly exceeded by the broad upper blade (2.3 dm. long); 
the glumes (PLATE 759, Fic. 3, X 6) with glabrous and lustrous 
surfaces (Michaux said “‘valvis acutissimis, nudis’’); the more 
abundant beard up to 6 mm. long; and the awn (PLATE 760, 
FIG. 3, X 3) only 8-10 mm. long, less than twice, instead of nearly 
thrice the length of the glumes. 

The collection in Arkansas of a plant which closely matches 
the Michaux type and which is so different from the Erianthus 
“brevibarbis”’ of most authors supports Michaux in his statement 
on the label. Although the assertion (generally attributed to 


248 Rhodora [JUNE 


Richard who issued the work after the death of Michaux) in the 
original publication, that it came from hills of Tennessee and 
from Carolina does not coincide with the original label, there is 
now clear evidence that a plant like Michaux’s does occur in the 
Mississippi basin. It is not without significance that Hackel, the 
master of the Gramineae, doubted the identity of Michaux’s 
plant with that of Elliott. In his great work on the Andropo- 
goneae in DC. Mon. Phan. vi. 131 (1889), treating FE. brevibarbis, 
as E. saccharoides subsp. brevibarbis, he accurately described the 
newly recognized EF. coarctatus, var. Elliottianus, doubting if he 
had the Michaux species (transferred by Persoon to Saccharum), 


his citations reading: “Er. brevibarbis Mich. . . . ?, certe 
Elliott, Sketch . . . et aliorum auctt. amer.; Sacch. brevi- 
barbis Pers. . . . ?” His doubt seems to have been justi- 


fied; at least, E. coarctatus and E. brevibarbis seem quite as distinct 
as do most of our species in the genus. As to the range of the 
latter, it is yet to be worked out. The botanists of Indiana, 
Illinois, Tennessee and Missouri seem not to have noted it; and, 
although Demaree’s label bears the memorandum, ‘‘New to 
Ark.”, Hackel, 1. c. cited as E. brevibarbis Arkansas material at 
Berlin, received from Engelmann. Michaux’s label, giving the 
data, on dry hills 5 days distant from the River Wabash toward 
the mouth of the Missouri, means that he got it in southern 
Illinois, presumably between Jefferson County at the east and 
Randolph County at the west. On August 23, 1795, Michaux, 
with an Indian, and a horse to carry his baggage, left Vincennes 
on the Wabash, in Knox County, Indiana, and on the 28th spent 
the day drying out his water-soaked collections by a camp-fire, 
reaching Kaskaskia, in Randolph County, Illinois, on the 30th. 
From late August to October 2 he collected up and down the 
Mississippi, with Kaskaskia as a base, and then returned to the 
Ohio. Five days travel from Vincennes, allowing for the stops 
recorded in Michaux’s diary, means that he got Erianthus brevi- 
barbis in southwestern Illinois; and it is clear that his over- 
mature material was collected, at latest, in early October. The 
latter fact and the over-ripe material from Arkansas, collected 
on September 23, indicate that true E. brevibarbis, although 
little known, is a relatively early species to mature. 

In habit and in dense panicle with appressed-ascending 


EEE “ CS FF ae JF... eSslUmMEllmUlC(‘( QS! )U6UCUC 


a a Ss 


ae 


1943] Fernald,—Erianthus brevibarbis and other Species 249 


branches, typical Erianthus coarctatus strongly suggests E. 
strictus Baldwin; but the panicle of EF. strictus is more slender and 
elongate and its spikelets are naked at base or with the merest 
suggestion of a greatly abbreviated coma at the tips of some 
pedicels. Furthermore, in eastern Virginia FE. strictus is the 
earliest-flowering species of the genus, our 8 collections, from 
young anthesis to mature fruit ranging in date from July 20 to 
September 19, with a single one, from wet woods, secured on 
October 18; the Virginia collections of E. coarctatus, both im- 
mature, were made on September 20 and on October 14. 

Michaux, who established the genus Erianthus, did not realize 
the complexity of the genus. He recognized but two species: 
his E. saccharoides, ‘“‘a Carolina ad Floridam”’, with ‘‘gluma villis 
involucrantibus multo breviore’’, identical with Anthoranthum 
giganteum Walt. (1788); and E. brevibarbis. FE. strictus, E. 
coarctatus and other species which he must have encountered 
and collected, were not worked out by him or, presumably, were 
confused with those of which types are preserved. 

Related to Erianthus brevibarbis and E. coarctatus in having 
the terete awns projected forward (rather than flattened ones 
spirally twisted at base and with the straightish tip thrown some- 
what to one side) are two plants with thicker panicles and with 
coma exceeding the glumes: EH. saccharoides Michaux or Anthoz- 
anthum giganteum Walt. = E. giganteus (Walt.) F. T. Hubbard 
and sensu Hitchcock, but certainly not E. giganteus Muhl., to 
whom Hitchcock erroneously ascribes the species; and E. com- 
pactus Nash. Before considering the differences between these 
two it is necessary to consider the correct name for the plant 
which Hitchcock, Man., is calling FE. giganteus, for it is quite 
clear that his discussion (Man. p. 854) was based on confused 
ideas and inaccurate quotation of Muhlenberg’s Catalogue. 
Hitchcock’s paragraph is as follows: 

(5) Erianthus giganteus (Walt.) Muhl., Cat. Pl.4. 1813. Based o 
Anthoxanthum giganteum Walt. Late Deser. Gram. 192. 1817) Mublen- 
berg uses the name for bo th E. saccharoides ents 1803] a 
alopecuroides [L. (Ell.)] (his herbarium specimen un nder this Tne 
rite both species), but the description = twisted) applies mg 

o E. alopecuroides. Erianthus giganteus was published as new 
Hubbard (Rhodora 14: 166 (1912) based on gre tran Poa 
Walt. 


250 Rhodora [JUNE 


If, as Hitchcock definitely states, the name Erianthus giganteus 
Muhl. Cat. had been based on Anthoxanthum giganteum Walt., 
there was no need of a new combination by Hubbard; but Hubbard 
in 1912 was following the International Rules of that period and, 
as he clearly and correctly explained, HZ. giganteus Muhl. Cat. 
(1813) was Not based on Anthoxanthum giganteum Walt., but was 
a change of name by Muhlenberg of Andropogon alopecuroides L. 
Since by present-day rules Muhlenberg had no right to give the 
new specific name, instead of using the one assigned by Linnaeus, 
Muhlenberg’s quite new name, Erianthus giganteus, is illegiti- 
mate; but, by the “homonym rule’’, adopted in the International 
Rules since Hubbard wrote, there is no room for a second £. 
giganteus, based on Walter’s name, especially since Walter’s 
species is admittedly identical with E. saccharoides Michx. As 
to Hitchcock’s flat statement that Muhl. Cat. (1813) based the 
name E. giganteus upon Anthoxanthum giganteum Walt., the 
following reproductions of Muhlenberg’s text indicates that the 
statement could not have been verified. The first reproduction 
is from ed. 1 (1813), the second from ed. 2 (1818). 


[ 


mest 


] 26. ANTHOXANTHUM, SPRING GRASS, 
2. giganteum 


Walter v.erianthusS ®! ena, 
27. ERIANTHUS, ERIANTHUS, semen I. 
1. giganteus, 21 
andropogon, \ gigantic, Car. Virg. 
alopec. L. 
, 
[2] 27 ae t SPRING GRASS semen 1. 
2 giganteum, : : 
Walter. v. ucthast ———_ 
28 ERIANTHUS ERIANTHUS semen I. 
1 giganteus 2) : 
posed \ gigantic Car. Virg. 
a 


It is perfectly clear that Muhlenberg was maintaining Walter’s 
Anthoxanthum giganteum under Anthoxanthum!, though with the 
rather vague intimation (‘‘v. [vel] erianthus”) that it was perhaps 
an Erianthus. He made no combination based directly upon it; 
but his Zrianthus, the next genus, consisted of two other species, 
1. giganteus, a substitute-name for Andropogon alopec(uroides) L., 
and 2. E. brevibarbis Michx., already discussed. In his Cat. 


1943] Fernald,—Erianthus brevibarbis and other Species 251 


ed. 2: 4 (1818) Muhlenberg repeated (see quotation above) the 
treatment of ed. 1, merely making Andropogon alopecuroides 
more emphatically the nomenclatural basis of E. giganteus by 
using italics: “1 giganteus Andropogon alopec. L.” 

There is no question, apparently, about the identity of Erian- 
thus alopecuroides (L.) Ell., which was based on Andropogon 
alopecuroides L. Sp. Pl. ii. 1045 (1753), the type being Clayton, 
no. 601 from Virginia. A. photograph of the type, X %, is 
reproduced as PLATE 761, Fic. 1, with an enlargement (FIG. 2) 
of spikelets, X 3, showing the characteristic flattened and twisted 
awn and the copious long coma. FErianthus giganteus Muhl. 
Cat. (1813), based directly upon it, has nothing to do with E. 
giganteus (Walt.) F. T. Hubbard; and the latter, a valid combi- 
nation when published, must give way to E. sACCHAROIDES 
Michx. As to the definition by Muhlenberg of a plant he subse- 
quently called Erianthus giganteus, that simply confirms his 
identification of it with Andropogon alopecuroides L., for he 
emphasized the twisted awn. I cannot follow the reasoning by 
which £. giganteus was taken up by Hitchcock in his Manual as 
E. giganteus ‘(Walt.) Muhl.’’ In 1908 he was apparently right 
when he wrote of Anthoxanthum giganteum Walt. ‘The specific 
name can not be taken up because there is an Hrianthus giganteus 
Muhl., based upon Andropogon alopecuroides L.’’—Hitche. in 
Contrib. U. 8. Nat. Herb. xii®. 151 (1908). 

Although Nash in N. Am. Fl. xvii'. 94 (1909) reduced to 
Erianthus saccharoides Michx. his own E. compactus Nash in 
Bull. Torr. Bot. Cl. xxii. 419 (1895) and although Hitchcock, 
Man. also reduces it to the ill-fated F. giganteus, it seems to me 
a very well defined variety. Typical £. saccharoides, as shown 
by a photograph of the type before me, has the excessively hairy 
panicle 2-6 dm. long, the long beard of the spikelet 2 or 3 times 
as long as the blades of the glumes. It occurs from Florida to 
Texas, north to southeastern Virginia. Typical E. compactus 
has the panicle only 1-2 dm. long and the beard from slightly 
longer than to barely twice the length of the blades of the glumes. 
It occurs from the Carolinas and Alabama northward to south- 
eastern New York, New Jersey, eastern Pennsylvania, the 
District of Columbia, northern Virginia and Kentucky, in the 
southern part of its range working back to the Appalachian 


252 Rhodora [JUNE 


Mountains. Throughout the region where it is beyond the range 
of E. saccharoides it is readily recognizable and quite distinct, but 
a large proportion of specimens before me from South Carolina 
show a mixture of the two trends (panicles up to 3.5 dm. long 
but with short coma, the blades of the glumes thus very evident 
in the panicle as contrasted with the hidden blades in typical 
E. saccharoides) while some specimens from Georgia lie between 
E. saccharoides and E. compactus; so that I am forced to consider 
E. compactus an essentially northern and inland variety rather 
than a true species: 

E. saccHaropEs Michx., var. compactus (Nash), comb. nov. 
E. polepint Nash in Bull. Torr. Bot. Cl. xxii. 419 (1895). 

It is, unhappily, necessary to discuss the type of Andropogon 
divaricatum L. Sp. Pl. i. 1045 (1753). This species has been 
made by Hitchcock and by Nash identical with A. alopecuroides 
L. 1. c. (1753) and in 1908 Hitchcock took it up, apparently 
because of priority on the page, to replace E. alopecuroides (L.) 
Ell. (1816), saying 

Andropogon divaricatum L. Sp. Pl. 1045. 1753. 

The type specimen is marked “2 divaricatum’’ and is from Gro- 


novius. As pointed out ae » [> Bot. Gaz. 35: 215. 1903], this 
is the same as A. gs bir L., which is an Erianthus. It should 
be called Erian 


which is based on Clay ton no. 600. This is Sorghas m linnaeanum 
(Hack.) Nash eae ore in Contrib. qi S. Nat. Herb. ‘ai. 125 (1908). 


My faith in the acumen of Linnaeus is such that I do not expect 
to find him describing identical species twice on the same page; 
although such accidents did happen. The diagnosis in 1753 of 
Andropogon alopecuroides was “4. ANDROPOGON panicula 
laxa, aristis tortuosis’”. That was all except literary citations, 
which, since Linnaeus had a specimen (our PLATE 761, FIGS. 
and 2) matching the diagnosis, are wholly secondary. Similarly 
A. nutans L., type of Sorghastrum nutans (L.) Nash, had 4 
“panicula nutante” and our familiar Andropogon virginicum was 
described “paniculae spicis conjugatis” &c. All these accounts 
(except of Andropogon nutans L.) are borne out by photographs 
of the types before me. Andropogon divaricatum did not have 4 
panicle. Instead it was clearly defined: “2. ANDROPOGON 
spica oblonga, floribus lanatis remotis divaricatis: arista flexuos® 


1943] Fernald,—Erianthus brevibarbis and other Species 253 


nuda”; and the specimen in Linnaeus’s Herbarium (PLATE 761, 
Figs. 3 and 4) when he prepared his diagnosis, therefore the TyPE, 
coincides most accurately with the brief but clear account. 
What it is I do not know. It was misidentified by Linnaeus 
with a plant of Clayton’s from Virginia, “Lagurus humilior, 
panicula conica laxa nutante culmum terminante’’, a plant which 
Hitchcock says is the same as 3. A. nutans L. (Sorghastrum 
nutans). It would be most extraordinary if Linnaeus confused 
specimens of his nos. 2 and 3 and if Gronovius and Clayton before 
him treated as two different species from Virginia material of 
only one, A. nutans, while they did not recognize the conspicu- 
ously different Sorghastrum Elliottii (Mohr) Nash, which is fre- 
quent in eastern Virginia. 

According to Hitchcock’s statement in 1908, ‘Linnaeus also 
cites a synonym from Gronovius which is based on Clayton no. 
600. This is Sorghastrum linnaeanum (Hack.) Nash’’—Hitche. 
in Contrib. U.S. Nat. Herb. xii®. 125 (1908); and he subsequently 
(Man. 951) states that S. Linnaeanum (Hack.) Nash, going back 
to Sorghum nutans, subsp. Linnaeanum Hackel in Martius, FI. 
Bras. ii®. 276 (1883), was “misapplied” by Nash “to S. Elliottit 
(Mohr) Nash’. Most unfortunately, here, as in so many cases 
already discussed, error seems to have crept in. Clayton’s no. 
600, basis of the Gronovian reference given by Linnaeus under 
Andropogon divaricatum, is beautifully preserved material, for a 
photograph of which (our PLATE 761, Fie. 5) I am indebted to 
Dr. Ramsbottom. It is, indeed, the best sort of Sorghastrum 
Linnaeanum, i. e. S. Elliottii; and my faith in the acuteness of 
Clayton, Gronovius and Linnaeus is justified. To be sure, 
Hitcheock reduced S. Linnaeanum to S. nutans (L.) Nash; but 
it seems improbable that he could have read Hackel’s original 
diagnosis: 

Panicula laxa, 25 em. lg., nutans, oblonga, ramulis longioribus apice 
bispiculatis. Spiculae intense rufae, 6 mm. lg.; gluma prima ad medium 
parce pilosa, secunda glabra. a 23-25 mm. lg., columna subulam 
eNom OE 1. II. 1045 (non Mant. II); Ell. Sketch. 
i. aT 


Sorghum nutans Chapm. l. ¢. 
America borealis: Florida, Georgia — Texas. 


The clear description by Hackel is very close to Hitchcock’s 


254 Rhodora [JUNE 


account of Sorghastrum Elliottzi, with ‘“‘panicle loose, 15 to 30 em. 
long, nodding at apex, the filiform branchlets and_ pedicels 
flexuous . . . ; spikelets 6 to 7 mm. long, chestnut-brown 
at maturity, . . . first glume hirsute or glabresecnt on the 
back; awn 2.5-3.5 cm. long, twice-geniculate”’. This strongly 
contrasts with the account of the plant Hitchcock calls S. nutans, 


with “panicle . . . yellowish, rather dense, contracted 
ooei a et maturity «....:; ewe 1-L5 .cm~ long, ones 
geniculate’. 


Returning to the actual Andropogon divaricatum L., the 
simple fact remains that its Type does not have a panicle. The 
photograph of it, kindly sent me by Mr. 8. Savage (our 
PLATE 761, Fics. 3 and 4), shows the summit of a culm with an 
oblong spike, the spikelets lanate, remote and divergent, the 
flexuous awn naked (spica oblonga, floribus lanatis remotis 
divaricatis: arista flexuosa nuda—Linnaeus). That it is not 
Erianthus alopecuroides (our PLATE 761, Figs. 1 and 2) or any 
member of that genus is obvious. I have tried in vain to place 
it with anything Virginian or eastern American. The truncated 
pedicels of some of the spikelets suggest Andropogon, as does the 
spiraling awn; but no Andropogon which I know. It is not 
impossible that its source was far from Virginia. The photo- 
graph, poor as it is and showing the spikelets heavily impreg- 
nated with glue, may lead to its proper identification. It should 
be noted that the long-exserted peduncle is quite naked, with @ 
prolonged and divergent blade at base. All eastern American 
species of Andropogon which have to be considered have close 
sheaths with appressed-ascending tips extending nearly or quite 
to the inflorescence. It should further be noted that one spike- 
let (r1G. 4) bears 2 or 3 spiraling awns, suggestive of Danthomia, 
but not of that genus. It is hoped that those who know the 
grasses will clarify the identity. 

As to the type of Andropogon nutans L. I have no satisfactory 
information. The panicle of the wide-ranging species with short 
awns and pale spikelets is not nodding (nutans); but there is no 
doubt that the plants of Elliott, basis of S. Elliottii, and of Chap- 
man were the latter very definite species. Since the identities 
of types throughout the group have been so discouragingly 
misunderstood, it is not at all improbable that the type of 4. 


1943] Fernald,—Why not Andropogon Gerardi? 255 


nutans, when critically compared, may lead to some alterations 
of our ideas. At least, it is probable that somewhere amongst 
the many names placed by Hitchcock in the reputed synonymy 
of his S. nutans others may be found earlier than Chrysopogon 
Elliottit Mohr (1897). 

rom the situation in Erianthus and in Muhlenbergia (see pt. I) 
it is evident that the Types of our eastern North American 
grasses need much further and closer study. 

Puiate 758. ERIANTHUS COARCTATUS Fern.: Fic. 1, TYPE, X %; F 
panicle, X 1; Fic. 3, summit of rachis of raceme, X 3; FIG. 4, spikelet, x 3. 
FIG. 5, spikelet, x 6. 

PLATE a E. ap tletyemt Michx., kindness of Professor Humbert: 
FIG. 1, TYPE, X ‘er - 2, spikelets of TYPE, x 11%; Fic. 3, spikelet, x 6, 
from Demaree, no 

Pate 760. E. secaora BARBIS Michx.: dicons 1, plant, X 1%, from Pulaski 
Cointity Arkansas, Demaree, no. 8228; 2, ‘summit of rachis of raceme, 

x 3, wg: no. 8228: FIG. 3, spikelet, 3, rom no. 

oor E761. Fig. 1, TYPE of ANDROPOGON ALOPECUROIDES L. and of 
Erta al ALOPECUROIDES (L.) EIl., X %, kindness of Dr. J. _— sagen gpa 
Fig. a; spikelets of same, 9 3, to show twisted awns. Frc. YPE of AND 

N DIvaRIcaTuM L, and of ERIANTHUS DIVARICATUS (L.) ‘Hitche., va x, 

a FIG. 4, summit of peg em x Fie. 5, 

Clay 0. 600, pi pe of A. DIVARICATUM, i. ORGHASTRUM ELLIOTT 
(Mabe) N may oo %, ‘indies of Dr. pate fe 


IV. WHY NOT ANDROPOGON GERARDI? 


As early as 1700 the common plant of eastern North America, 
known either as Andropogon provincialis Lam. Encyel. i. 376 
(1785) or as A. furcatus Muhl. ex Willd. Sp. Pl. iv. 919 (1806), 
was cultivated and perhaps escaped in Provence, in southern 
France. Tournefort, Inst. i. 521 (1700) had it as his Gramen 
dactylon, villosum, ramosum, altissimum, Gallo-Provinciale; but 
it was not until 1761 that the cultivated plant of Provence was 
beautifully described and illustrated by a figure as Andropogon 
spica digitatis, flosculis alternatim geminis, hermaphrodito aristato, 
sessili; masculo mutico, pedunculato by Gérard in his Flora Gallo- 
Provincialis, 106 (fig. 4) and 107 (1761), a plant which grew in 
southern Provence (Oritur in gallopr. australi. Perenne). Gérard’s 
description was so detailed and so lucid that it is here given in 


Des. Radix numerosdé fibrarum oe implicatarum prole luxurians. 
Culmi tripedales & ultra, glabri, striati. Folia radicalia multa, lata, 
glabra, cespites constituentia; ca ant Sg quinque vel sex, admodixm vaginan- 
tia, inferne ad margines pilis raris vestita. Membrana ex apice vaginarum 


256 Rhodora [JUNE 


brevis, lacinulata, sub villosa. Spicae ex apice culmi ange an 
; ; ini, basi villosi; 


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stigmata plumosa, purpurascentia; semen ovatum. Masculus pedicellatus, 
pedicello plumoso, anguloso; glumd calycind exterior, interiort paulo 
ongior, corollae glumae inaequales, muticae, calyce breviores. Filamenta 
ut in hermaphrodito, pistillum abortiens. 

s. Variat calycibus glabris, & villosis. 


In his Encyclopédie méthodique, i. 376 (August, 1785) La- 
marck published it as Andropogon provinciale from the Royal 
Garden at Paris, merely copying the diagnoses of Gérard and of 
Tournefort, and adding, in French, a brief interpretation, ending: 
“On trouve cette plante dans la Provence, & on la cultive au 
Jardin du Roi.” Seven years later the plant of Gérard received 
another binomial, A. Gerardi Vitman, Summa PI. vi. 16 (1792), 
Vitman copying Gérard’s diagnosis and citing both that and the 
illustration, the plant occurring “In Gallo provincia”. Then in 
1806 the wide-ranging North American plant appeared as A. 
furcatus Muhl. ex Willd. Sp. Pl. iv. 919 (1806). That A. pro- 
vincialis Lam. is the same species as A. furcatus Muhl. there 
seems no doubt. Franchet in Bull. Soc. Bot. France, xxxi. 350- 
352 (1884) gave a very detailed history of the plant in France 
and he concluded the study: ‘La plante figurée et décrite par 
Gérard et cultivée au Jardin du Roi dés 1763, plante qui est 
certainement le type de |’A. provincialis Lamk, comme en font 
foi les exemplaires desséchés 4 cette époque et qui se trouvent 
dans l’herbier du Museum, est une espéce américaine, nommée 
quarante-deux ans plus tard A. furcatus Muehl. in Willd. Sp. 
IV, 919 (1805 [1806]). L’A. provincialis doit done étre rayé de la 
flore francaise.” That it has, consequently, been “struck out 
from the French flora” is indicated by Rouy, Fl. Fr. xiv. 21 
(1913), where he states in a note that “1’A. Provincialis Lamk 
. . . = A. furcatus Mihlenb. . . . qui n’a existé en 
France que naturalisée et cultivée.’’ Hackel, in his great and 
comprehensive monograph of the Andropogoneae in DC. Mon. 
Phan. vi. 441 (1889), had no doubt about the matter, definitely 
citing in the synonymy of A. provincialis both A. Gerardi and A. 
Surcatus. 


1943] Fernald,—Why not Andropogon Gerardi? 257 


As it proves, unfortunately, the name Andropogon provincialis 
Lam. (1785), the name used by Hackel, by Nash in N. Am. Fl 
and by many others, is clearly antedated by another A. pro- 
vincialis Retz. Obs. Bot. iii. 43 (1783), a low annual, with 15 or 
16 fascicled spikes, unequal subulate glumes, etc., obviously not 
Lamarck’s species.! There is, however, little support for Hitch- 
cock’s doubt as to the identity of A. provincialis nor for his 
perplexity in settling which was earlier, Retz., 1. c. or Lam. 1. ¢. 
Hitcheock’s statement is as follows: 

The name Andropogon provincialis Lam. (Eneycl. i. 376. 1783), was 
applied to this species by Hackel (in DC., Monogr. Phan. 6: 441. 1889) 
and others, but Lamarck’s species is uncertain. He states that he 
saw a plant i in the Paris Botanical Garden, but his description is taken 


from Gerard (Fl. Gall. Prov. 107. pl. 4 1761) and does not well apply 
oO our species. Furthermore, A. praddes Retz. (Obs. Bot. 3: 43. 


: term 
The part of Lamarck’s Foam containing page 376 appeared in 
August 1783.—Hitche. Man. 790 (1935) .? 
Hitchcock does not state why Gérard’s ‘‘description  . 
does not well apply to our species.”” Franchet and Hackel, both 
accurate students of grasses, felt positive of the identity; and 
perusal of Gérard’s description side by side with Hitchcock’s 
(Man.) shows nothing specifically distinctive. Any one can 
readily make the comparison. He is likely to arrive at the con- 
clusion of Franchet, Hackel, Ascherson & Graebner, Rouy and 
others. As to Hitcheock’s inability ‘‘to determine which is the 
earlier [Retz. Obs. Bot. iii. or Lam. Encyel. i. 376], he was 
apparently relying on his own conclusion that ‘The part of 
Lamarck’s Encyclopedie containing page 376 appeared in August 
1783”. It is too bad that the latter error in date confused the 
question for, as indicated by Sherborn and Woodward in Journ. 
Bot. xliv. 319 (1906), the part including p. 376 was published in 
August, 1785 (not 1783). 
1 * Hacket (1889) placed it in Chloris. 
nference that the plant, formerly cultivated in France, is ‘‘not 
es’’ may have arisen form Hackel’s oiggteo it as subvar. genuinus, 
with ‘‘ecleamis . . pedicellatae basi fascicul pilorum manifesto 1,5 mm, 
longo barbatae”’ while his subvar. furcatus was “ut cieche: sed spiculae pedicel- 
It is a very exceptional plant of eastern 
North America which lacks the bearding at the base of the pedicelled spikelet or at 
the summit of the pedicel. The distinctive point stated by Hackel is of no impor- 
tance. 


258 Rhodora [JUNE 


Since Andropogon Gerardi Vitman (1792) is the same as A. 
provincialis Lam. (1785) and A. furcatus Muhl. (1806), that 
appears to be the proper name for our common species. The 
following are the most marked varieties: 

ERARDI Vitman, var. paucipilus (Nash), comb. nov. A. 
paucipilus Nash in Britt. Man . 70 (1901). A. provincialis Lam., 
(93 perenne (Nash) Fern. "& Grisc. in RHopora, xxxvii. 147 


A. GERARDI, var. chrysocomus (Nash), comb. nov. A. chryso- 
comus Nash, 1901). . provincialis, var. chrysocomus 
(Nash) Fern. & Goes l. c. (1935). 


(To be continued) 


V. STUDIES IN NORTH AMERICAN SPECIES OF 
SCIRPUS 


(Plates 762-767) 


Tuer AMERICAN REPRESENTATIVE OF SCIRPUS PUMILUS (PLATES 
762—764).—One of the rarest or most localized sedges of North 
America is the tiny plant with solitary terminal spikelet, belong- 
ing in Scirpus, § Baeothryon and passing for nearly forty years in 
America as the Eurasian S. alpinus Schleicher (1828) and subse- 
quently as the identical S. pumilus Vahl (1806). First recorded 
as American by Britton in Trans. N. Y. Acad. Sci. xi. 75 (1892) 
as S. alpinus from the “‘Rocky Mountains’, Hall & Harbour, 
and from Moreley in the Canadian Rocky Mountains, the species 
next attracted attention from its discovery on Anticosti Island 
and the adjacent Mingan Islands of Saguenay County, Quebec, 
and was the subject of one of the interesting articles by Marie- 
Victorin, in Trans. Roy. Soc. Can., ser. 3, xxiii. pt. 2, sect. V. 25 
(1929). Somewhat later, in Ruopora, xxxiii. 23, 24 (1931), I 
pointed out that the name S. pumilus Vahl antedates S. alpinus 
Schleicher and I then recorded additional stations in the Cana- 
dian Rocky Mountains; and more recently Beetle, in Am. Journ. 
Bot. xxviii. 421 (1941), has maintained S. pumilus as American: 
‘‘Mountain meadows of Eurasia and North America’’, he citing 
American plants of Quebec (Anticosti and the Mingan Islands), 
Saskatoon, Saskatchewan, Alberta and Colorado. The habitat, 
‘‘mountain meadows”, while perhaps applying in the Rocky 
Mountains, is hardly appropriate for the Quebec area, with 
Anticosti rarely attaining a height of 175 m.' and the Mingan 

1“Jes plus hautes plateaux cités par eux ne dépassent pas 150 et 175 metres’’— 
Despécher and Combes as quoted by Schmidt, Monographie de I'Ile d’Anticosti, 10 

rent r of § Baeothryon, the flat-leaved woodland Scirpus pegareiny 
ican, ea cited 8 Beetle, ]. c. 174, from ‘‘Vermont: Willoughby, W. Boott in 
William Boott’s herbarium is incorporated in the Gray Herbarium. There . pes 
no Vermont material of S. planifolius from him, and Kennedy and others who inti- 
mately knew the Willoughby region did not know it there. Dole lists it only from 


280 Rhodora (JULY 


Islands still lower, while Saskatoon is well to the east of the 
mountains, in the flat Canadian prairie. 

Ever since Britton’s identification of the Rocky Mountain 
plant with S. alpinus Schleicher or S. pumilus Vahl of central 
and southeastern Europe, eastward into Persia and central Asia, 
we have accepted the identification without question. The 
plants of Eurasia are at least two quite different species, or 
perhaps geographic varieties, while some of the Asiatic material, 
that of alpine regions (4305-5200 m.) of Tibet may be distinct. 
This Tibetan plant may well be Isolepis oligantha C. A. Meyer, 
Mém. des Sav. Etrang. Pétersb. i. 197, t. 1 (1831); but until 
some one who has more adequate nterial and who can study 
Meyer’s Siberian type can check it, the Tibetan plant may wait. 

True Scirpus pumilus of the alpine regions of south-central 
and -eastern Europe, thence into Persia and Siberia (our PLATE 
762) is subcespitose or quite densely cespitose (as shown by 
Reichenbach and others), with short stoloniferous offsets which 
promptly send up erect tufts of culms; the lowest scale of the 
spikelet often has the green midrib prolonged (rics. 3 and 4) as 
a green and blunt mucro; the anthers (Frias. 2 and 3) have the 
connective prolonged as a distinct subulus; the achenes (FIGS. 
6-8) are rather slenderly ellipsoid-obovoid, 1.6-1.8 mm. long, 
0.4-0.5 mm. broad, subequilaterally trigonous (rics. 6 and 9) 
and usually broadly rounded to truncate at the sessile base; 
while all but the lowermost scales of the spikelet have thin 
scarious margins. 

Fully 2100 km. to the north, within the Arctic Circle, there is @ 
plant, occurring on Porsanger Fjord (south of Cape North) in 
northernmost Norway which is identified in current Scandinavian 
floras as Scirpus pumilus. This plant (pLaTE 763, FIGs. 1-3) is 
extensively creeping, with very elongate filiform rhizomes and 
with small and scattered tufts of culms. Its anthers (FIG. 2) and 
achenes (FIG. 4) are like those of S. pumilus, except that the 
achenes are longer (2-2.2 mm. long); but its scales are much 


two Vermont stations, Mt. Philo (Addison County) and North Pownal (BenningtoD 
County). It has also been collected in Arlin vac 


Willoughby, near the northeastern corner. It is probable Gus the Willoughby record 

of S. planifolius arose from S. pauciflorus Lightf., now usually treated as Eleocharis 

paucifiora (Lightf.) Link (var. —— Svenson) which abounds at Willoughby and 
was there collected by Wm. Boott 


1943] Fernald,—North American Species of Scirpus 281 


firmer and broader than in true S. pumilus and the latter have 
the margins hardly if at all scarious (Fic. 3). Furthermore the 
lowest scale (FIG. 2) is nearly like the others, without the green 
mucro strongly excurrent. Differing so markedly from typical 
S. pumilus, it was beautifully described as Trichophorum emergens 
Norman in Soc. Reg. Se. Norveg. v. 319—repr. as Spec. Loe. 
Nat. 79 (1868)—and as abundant on the sandy inundated shore 
in Porsangria. ‘‘Vaginae basilares culmi ut in Trichophoro 
caespitoso, a quo rhizomate repente, stolonifero, fasciculos pauci- 
culmeos immo culmos solitarios hue illue emittente, recedit. 
Sub accessu maris inundatur, sub recessu emergit”. So different 
is this Finmark plant of inundated tidal shores from the more 
cespitose one of the southern alpine areas that it should certainly 
be kept apart.! Even the original editors of Index Kewensis, 
who did not hesitate to reduce species they did not understand, 
were baffled by Trichophoruwm emergens, appending after the 
citation ‘‘(Quid?)—Norveg.”’ 

As stated and as shown in the plates, the anthers in the 
Eurasian Scirpus pumilus and S. emergens have prominently 
excurrent connectives and the achenes are subequilaterally 
trigonous, while in S. pumilus the lowest scale often has the mid- 
rib excurrent as a mucro; and the blunt scales of the arctic tidal- 
shore S. emergens are subcoriaceous and scarcely scarious- 
margined. When we turn to the North American plant (PLATE 
764) which has passed as S. pumilus we find a habit midway 
between those of the two European species, the culms tufted, 
sometimes without but usually with slender and elongate 
rhizomes and scaly stolons. The American plant on superficial 
examination might easily be thought to stand somewhat inter- 
mediate between those of Eurasia. In our plant the scales of the 
spikelet are as thin as in S. pumilus but the lowest (Fics. 2 and 3) 
has the midrib included, not exserted; the anthers (Frias. 2 and 4) 
merely taper to tip, with the connective not exserted; and the 
achene (rics. 5-10) is plano-convex, broad and flat on the inner 
face, gently arching to merely umbonate on the back. The 
ripe achenes (rics. 5-7) are broadly ellipsoid-oblong, only 1.2- 
1.5 mm. long but 0.8-1.2 mm. broad, and gradually rounded to 


tScrrpus emergens (Norman), comb. nov. Trichophorum emergens Norman in 
Soc. Reg. Sc. Norveg. v. 319—repr. as Spec. Loc. Nat. 79 (1868). Pare 763, figs. 1-5. 


282 Rhodora [JULY 


base or sometimes substipitate. Differing in these minute but 
morphologically important characters from the Eurasian series, 
the North American plant is clearly an endemic species. Our 
fullest representation is the abundant series from Anticosti and 
the Mingan Islands, collected by Bros. Marie-Victorin and 
Rolland-Germain. I am greatly pleased to have this opportunity 
to associate with a plant of that area the name of a modest and 
self-effacing botanist who has done much in his earnest and 
discriminating way to bring to our knowledge the rarer plants 
of Quebec, Brother Rolland-Germain. 

Scirpus (§ Baroturyon) Rollandii, sp. nov. (ras. 764). 
Planta habitu foliis culmisque ut in Scirpo pumilo sed valde 
stolonifera; culmis laxe cespitosis vel subsolitariis; spiculis 
ellipeoideo-ovoideis 3-4 mm. longis, subteretibus; squamis 
ovatis obtusis vel subacutis brunneis vel rufescentibus subcoria- 
ceis margine scariosis; antheris 1.5 mm. longis apice attenuatis; 
achaeniis nigrescentibus late oblongo-ellipticis, 1.2-1.5 mm. longis 
0.8-1 atis, plano-convexis dorso leviter convexo vel um- 
bina aoe euate Archipel de Mingan: rivages calcaires, Ie 
Sainte-Geneviéve, 9 aotit, 1925, Victorin & Rolland, no. 20, 220: 
corniches caleaires du cété du large, Ile Sainte-Genevidve 22 
juillet, 1926, Victorin & Rolland, no. 25,785 (rype in Herb. 
Gray.); rivages caleaires, Ile & Marteau, 23 uillet, 1926, Victorin 
& R Rolland, no. 25,940; parties élevées et découvertes surtout 
dans les sentiers de renard, Grande Ile & la Vache Marine, 19 
juillet, 1926; Victorin & Rolland, no. 25,782; tundra calcaire 


e 
Marine, 3 aoit, 1928, Victorin & Rolland, no. 28,374. Anti- 
costi: sur les platiéres argilo-calcaires au-dessus des gorges, 
Riviére Chicotte, 15 aofit, 1925, asia & Rolland, no. 25,783; 
sur les platiéres a plusieurs milles en aut des gorges, Riviere 
Chicotte, 24 juillet, 1927, Victorin & Rolland no. “OT, 517; sur les 


au Fusil, 20 juillet, 1927, Victorin & Rolland, no. 27 ,018; sur les 
platiéres prés de la mer, 25 juillet, 1927, Ain hig & Rolland, no. 
27,516; éboulis argilo-caleaire, le long de la mer, a l’est de la 
rividre, Riviére la Loutre, 6 aout, 1936, Victorin & Rolland, no. 


Butters & Hotes no. 50. Cotor ADO: “Rocky Mt. Alpine Flora, 
Lat. 39°-41°’, 1862, Hall & Harbour, no. 583. 


TE 762 shows details of Scrrpus pumiius Vahl: ric. 1, habit, X 1, 
fevan Pee, Semipalavitinsk, western Siberia, May 21, 1920, 0. Simonova & 


1943] Fernald,—North American Species of Scirpus 283 


T. Batueva; Fic. 2, spikelet, 10, from same collection; Fic. 3, spikelet, 
10, from Mont Cenis, Savoie, July 27, 1855, Perrier; FIG. 4, spike let, X 10, 

m Simonova & Batueva; FIG. 5, disintegrated spikelet, showing broad 
scarious margin of scale, 10, from Zermatt, July, 1882, Christ; Fic. 6, 
achene and its cross-section, after Schroeter, Pflanzenl. der Alpen, fig. 174 
(1923); ria. 7, achene, X 10, from Mont Cenis, Perrier; ric. 8, achene, X 10, 
from Zermatt, Christ; ric. 9, looking down on summit of achene, x 10, from 
or. Chrtat 

E 763, FIGS. 1-5. §. EMERGENS (Norman) Fernald: Fie. 1, habit, 

fron Poreainke Fjord, Finmark, July 9, 1898, A. L.; Fic. 2, spikelet, x XO, 


10 15 e 

2 ay from Dahl; Fic. 5, looking down on tip of achene, X 10, from Dahl. 
6, S. rufus (Huds.) Schrad.: achenes, < 10, from Skane, Sweden, July 14, 

1928, rik Asplund. Fia. 7, S. rufus, var. neogaeus: achenes, x 10, from the 


TY 
ie 764, S. Ro.tuanpi Fernald: ric. 1, habit, x 1, from the Ty 


reas some ge de Mingan n, Quebec, Victorin & Rolland, no. gy a FIG. 6, 
achene, X 10, val Riviére Chicotte, Anticosti, Victorin & R no. 
2, 317, FIG. 7, sehen x rea m Saskatoon, Saskatchewan, July 24, 1939, 
W. P.’Fras aser; FIG. 8. Joo ing down on tip of ‘achene in fig. 5, X 10; 

similar Mads, x 10, of achene in fig. 6; fig. 10, similar view, x 10, of ietins 


© Norta AMERICAN MEMBERS OF Scirpus, § LacusTRES 
Guan 765 and 766).— 

Scirpus vALipus Vahl, var. creber, var. nov. (TAB. 765, FIG. 
4-7), spiculis ovoideis 5-9 mm. longis; squamis costa margineque 
exceptis glabris lucidis achaenio maturo vix superantibus; 
antheris deinde opel wittbesd ee wage vs perianthio achaenium 
subaequante; achaentiis 1.7— . longis 1.3-1.5 mm. latis.— 
Newfoundland to British Columbia, south to Nova Scotia, New 

ngland, Long Island, Georgia, Tennessee, Missouri, Oklahom ma, 
Texas, New Mexico, northern Mexico and California. TYPE 


Forma ee f. nov. (TAB. 765, FIG. 8), spiculis 


ee ee 9-15 mm. longis; achaeniis 2.3-2.8 mm. longis 
1.4-1.8 mm. latis. —Seattered through the range of rae creber; 
the followeine are characteristic. Nova Scotta: Truro, Fernald 


oy Ns no. 2720. MAINE: Lincolnville, G. B. Rossbach, no. 
AcHusETTS: York Pond, Canton, July 8, 1894, 
Reeacdae Eaathans F. S. Collins, no. 1293; Wakeby — 
Sandwich, Sectiehe 16, 1916, Harger & Woodward. Conn 
ticuT: East Windsor, August 21, 1904, Brssell. ° dpe se was 
of Toano, Janae City a y; R. . Menzel, no. MICHIGAN: 
New Buffalo, Lansing, no. 3281. Iowa: Lost is Township, 
Clay County, Ada Tinie: no. 9195. Norra Daxota: Mandan, 
F. P. Metcalf, no. 374. Nepraska: St. Paul, July 24, 1909, "2 
M. Bates. Kansas: Joy rag Osborne County, July a1, 1894, 


284 Rhodora [JULY 
Shear, no. 239 Ske in issih Gray.); Riley Co., June 21, 1895, 


Norton. Texas: Dallas Co., July, 1877, Reverchon; El 
Paso ‘County, Cory, 83 1255. “ALBERTA: Aes (Mamawi) Creek, 
Wood Buffalo Park, Raup, 1980. Orxrcon: St. Paul, J. C 


Nelson, no. 1692; Salem, Ny aap, no. 3307. 

Scirpus validus (PLATE 765, Frias. 1-3) was described by Vahl, 
Enum. ii. 268 (1806), its ‘Habitat in Caribaeis’’, with a clear 
diagnosis and a remarkably complete description, he distinctly 
saying “‘squamis dorso villosis’”. It is the plant of eastern 
tropical America! which differs at once from the common plant 
of the United States and Canada in several points. Its inflores- 
cence (ric. 1) is usually stiffer than in the common North 
American plant, only exceptionally with arching and pendulous 
rays and pedicels, although the more northern plant may have 
the inflorescence as stiff as in S. acutus Muhl. Typical S. validus 
may, as originally described, have the scales of the spikelets 
villous or they may be glabrescent or even glabrous except for 
the keel and the fimbriate-ciliolate margin. In none of the 
tropical and subtropical American material do the achenes show 
beyond the scales; the scales strongly cover them and are nearly 
twice as long. The perianth consists of very delicate bristles 
remotely retrorse-setulose chiefly above the basal third and 
commonly overtopping the achene. The connective of the anther 
(rigs. 2 and 3) projects as a triangular-ovate sessile tip, though 
sometimes becoming elongate. North of tropical America true 
S. validus is frequent or common in Florida and there is material 
in the Gray Herbarium with the stereotyped and possibly too 
inclusive label ‘“‘Santee Canal, South Carolina, Ravenel”. 

From genuine tropical and subtropical Scirpus validus our var. 
creber differs in its often more lax inflorescence (Frias. 4 and 5), 
the backs of the scales glabrous, the scales barely covering OF 
when they are ripe (F1a. 5) not wholly covering the achenes; the 
perianth (ric. 7) of usually coarser and rather shorter bristles 
which are copiously retrorse-setose; and the anther (F1G. 6) with 
the slender tip becoming prolonged. Some material from the 
southeastern states and some from Bermuda is so transitional 

‘The citation by Beetle in Am. Journ. Bot. xxviii. 695 (1941) of Bermuda, Haiti. 
Porto Rico, Jamaica and Cuba as the “Easr Inpres” and his statement of broad 
range (p. 693), sR ae pcan North America, and bordering the a 


with the only South American specimens cited coming from Uruguay 
Argentina, suggest need of more precise knowledge of geography. 


1943] Fernald,—North American Species of Scirpus 285 


that I am treating the common plant of the United States and 
Canada as a strong geographic variety, rather than as a species. 
In its prolonged and linear-cylindric spikelets up to 1.5 em. long 
var. creber, forma megastachyus (FIG. 8), when its inflorescence 
is contracted, might be mistaken for S. acutus. In fact, by the 
recently published key to epee of this section (‘‘Spikelets 

S. 


ovoid . . . S.validus’’; “Spikelets subcylindric 
acutus’’! and the accompanying descriptions (Septicalots 5-10 
mm. long, . . . ovate’ for S. validus; ‘spikelets 0.7—2 cm. 


long . Ovate-acute to cylindrical” for S. acutus) one 
aiid handily place most specimens under S. validus; nevertheless 
several specimens, including the type, of forma megastachyus 
were cited as good S. validus. Accompanying the elongation of 
spikelet the achene of forma megastachyus is enlarged, achenes 
from the form running considerably larger than in typical var. 
creber. 

In connection with Scirpus validus, var. creber two names have 
to be considered, because they are cited by Beetle as synonyms 
under his all-inclusive S. validus. These are S. orgyalis Raf. 
Annals of Nature [not ‘‘ Amer. Nat.” as cited in the recent paper], 
16 (1820) and S. lacustris, var. condensatus Peck, N. Y. State 
Mus. Rep. no. 53: 853 (1900). Rafinesque’s S. orgyalis of 
‘creeks and rivers of New York and Pennsylvania’, had little of 
specific character in the brief description except ‘‘spikes lateral 
under the apex, glomerated, ovate, sub-sessile; scales ovate, 
mucronate, brown, arachnoidal”. What he had we do not 
know. He intended his name to apply to anything North Ameri- 
can of the lacustris group. The ‘‘glomerate . . .  sub-sessile’’ 
spikes with ‘‘arachnoidal’’ scales could as well, if not better, 
apply to S. acutus Muhl. (1842) which occurs in New York and 
western Pennsylvania and which often has villous scales, while 
those of the S. validus of that region have the scales glabrous 
except for keel and margin. 

Peck was not differentiating between the three species of 
§ Lacustris which occur in New York State. His “‘S. lacustris 
condensatus n. var.’’ was described: ‘Heads of the panicles 
sessile or on very short pedicels, forming a dense cluster about 
1 inch long and broad. Otherwise asin the common form. Lime 


1 Beetle, 1. c. 692 (1941). 


286 Rhodora (JULY 


Lake. August. F. E. Fenno.’”’ According to House, N. Y. State 
Mus. Bull. no. 254: 147 (1924) the Fenno plant is S. heterochaetus, 
a species which usually has a lax and open panicle and which 
differs from S. validus in its very pale and solitary spikelets, its 
trigonous achene, etc. Even if S. lacustris condensatus were an 
unusual form of S. validus it would be quite ridiculous to take up 
the name, intended for a trivial form, for the transcontinental 
plant with open and loosely forking panicles. Extreme literalists 
might do so, but the intent of the original author was obvious; 
the rules of nomenclature were not intended to foster absurdity, 
if they do, it is time to change them. 

S. Steinmetzii, sp. nov. (TAB. 766, Fic. 1-7). Planta statura 
habituque ut in Scirpo ee ‘spiculis ellipsoideo-ovoideis 
obtusis 5-7.5 mm. longis 4-5 mm. crassis fulvis; squamis scarioso- 

membranaceis fulvis acuntnatie atabrie s, margine apiceque 
vile ints aristo Enel antheris apice triangulari-ovatis : 


stream, Passadumkeag Stream, Passadumkeag, Penobscot 
County, Angus a 1007, #.. i. hd inmetz, no. 355 (TYPE in 
Herb. Gra yPE in Herb. Univ. of M aine), September 1, 
1938, Blecseale ay 29, 1942, Maintnets & Gashweiler, speci- 
mens originally and tentatively placed with S. heterochaetus 
Chase; shallow water near shore of river, Passadumkeag River 
at Hathaway iat Passadumkeag, August 5, 1940, Ogden & 
Wright, no. 2345 

Scirpus Saux. with which it is a great pleasure to asso- 
ciate the name of the leader of botanical exploration in Maine and 
discoverer of the plant, is like S. heterochaetus only in having 4 
prolonged basal leaf and in its lax inflorescence with no tendency 
to the formation of glomerules. S. heterochaetus has slender 
pale green to whitish-brown lanceolate to slenderly ellipsoid 
acute or subacuminate spikelets 0.75-2.3 cm. long; its pale 
scales are firm to subcoriaceous and deeply emarginate at tip; 
its styles mostly 3-cleft, its achenes trigonous. It is a species of 
calcareous or alkaline waters; and it was beautifully illustrated 
by Mrs. Chase when she published it in Ruopora, vi. 70, t. 53, 
fig. d (1904). S. Steinmetzii, on the other hand, has the reddish 
to purple-brown plump-ovoid obtuse spikelets only 5-7.5 mm. 
long; the thin and almost scarious scales tapering to the aw? 


1943] Fernald,—North American Species of Scirpus 287 


(not deeply emarginate) and heavily villous-ciliate (rather than 
slightly or hardly so) at margin; the style is 2-cleft and the 
strongly flattened achene plano-convex, merely slightly convex 
on the back. It should, therefore, be confidently watched for in 
the extensive lake-strewn area which extends from the Penobscot 
across Hancock and Washington Counties, Maine, into south- 
western New Brunswick. 

From Scirpus validus, var. creber the newly described S. 
Steinmetzii is distinguished by its non-glomerulate spikelets, the 
scales greatly overtopping the achenes (as in tropical S. validus), 
but with tapering (rather than broadly rounded and emarginate) 
tip, the anther-connective with a triangular sessile terminal ap- 
pendage, the bristles fewer and short or wanting, and the persist- 
ent old filaments very broad and ribbon-like. 

In pLaTe 765, FIGs. 1-3 are of typical Scrrpus vatipus from Cordillera 
Septantsinnal, prov. Puerto Finke, Sabaneta, in Cafio Hondo, Civ. hiya 
Domingo, Hispaniola, Ekman, . 14, 549: ric. 1, inflorescence, <1 31G. 2, 


spikelet, showing blunt anther, x ; FIG. 3, achene, showing elongate bristles 
and — eg age! x 10. Fies. 4-7 var. CREBER: FIG. 4, rego of TYPE, 


rescence of TYPE of var. CREBER, forma MEGASTACHYUS, 

In Pirate 766 Fies. 1-7 are of Scrrpus STEINMETZII, all from spat —_ 
series: fig. 1, 1 orescences, x 1; Fic. 2, axis of inflorescence, X 3; .3 
and 4, spikelets, showing long scales and blunt anthers, X 5; Fies. 5 at 6, 
achenés, with single pe rianth-bristle and broad filaments, - 10; FIG. 7, achene 
viewed from above Uncking down on mw 20, Pie 8-10, S. HETERO- 
CHAETUs, from kirk, Oswego County, New York, Pest: Wiegan & 
Eames, no. 14,192: Fia. 8, et. < 5; FIG. 9, achene, X 10; Fic. 10, achene, 
viewed from above, X 

Some AMERICAN SPECIES AND VARIETIES OF ScrRPUS (PLATES 
763, figs. 6 and 7, and 767).—ScrrRPUS RUFUS Sag Schrad., 
var. neogaeus, var. nov. (TAB. 763, FIG. Eats achaen s fusiformi- 
lanseoinee 4.5-5.5 mm. longis 1-1. ‘7 mm. latis a vaide weenie 
rostratisque.—Saline to brackish, raiely fom marshes, Ne 
foundland and shores of Gulf of and lower River St. Lawrencé: 
Quebec, south to southwestern Nova Scotia and pong ide 
New Brunswick; salt marshes from Churchill t 
River, lat. 53°, Manitoba. Type: near Hospital Point. Grind- 
stone Tsland, Magdalen Islands, Saly 18, Fernald, Bartram, 
Long and St. John, no. 6968 (in Herb. Gra 

In habit, foliage, spikes and spikelets we Avibsitin material 
of Scirpus rufus (Blysmus rufus (Huds.) Link) is quite like the 
plant of northern Europe and it shows the same diversity of 


288 Rhodora (JULY 


involucre, oftenest with it essentially obsolete or reduced to a 
short blade but occasionally with a long blade overtopping the 
compound spike. The European descriptions very generally 
define the achene as elliptic and of a yellow-gray color. There 
being no reason in European works for further detail the size is 
not often given. Holmberg, however, in his very detailed and 
unfortunately never completed Scandinaviens Flora, Hifte 2: 
304 (1926) says ‘Not spolformigt [fusiform] elliptick . . 3 
mm. ling, 1,5 mm. bred, gul-1. brungr&.”” The achenes of the 
European plant (PLATE 763, FIG. 6) range from 3-4.5 mm. long, 
with the rather definitely elliptic body usually about 3 mm. long 
and 1.5-2 mm. broad and opaque to barely sublustrous. The 
achene of the American plant is more definitely fusiform, 4.5-5.5 
mm. long, lustrous and of slightly warmer color, the body only 
1-1.7 mm. broad, the stipe and beak more prolonged than in the 
European plant. Fic. 6 shows achenes, X 5, of typical S. rufus 
from Skine, Sweden, July 14, 1928, Erik Asplund; Fie. 7, 
achenes, X 5, from the TyPE of var. neogaeus. 


S. SUBTERMINALIS Torr., forma terrestris (Paine), comb. nov. 
Var. terrestris Paine, Cat. Pl. Oneida Co., 96 (1865). 

S J ., var. Frernaupr (Bicknell) Beetle, forma 
agonus, culmis ad 1.5 m. altis et 1.3 em. crassis; foliis ad 15 mm. 
latis; spiculis 1.2-4 em. longis; achaeniis late vel subrotundo- 
obovatis ad basin sensim rotundatis plerumque 2.5-3.2 mm. latis 
plano-convexis vel lenticularibus dorso sensim rotundatis.— 


brackish marsh, Sable Island, St. John, no. 1160; near brackish 
mouth of Salmon River, Truro, Fernald & Wiegand, nos. 424 
and 4249; border of salt marsh, Jordan Falls, Shelburne County; 
September 4, 1921, Fernald & Long, no. 23,398 (ryprE in Gray 
Herb., distrib. as S. campestris, var. novae-angliae). NEW 
Brunswick: brackish marsh, Bathurst Bay, Bathurst, 8. F. 
Blake, no. 5457; marsh, Bay du Vin Island, Northumberland 
County, Blake, nos. 5706 and 5707; border of brackish pond, 


1943] Fernald,—North American Species of Scirpus 289 


Whale Cove, Grand Manan Island, C. A. & Una F. Weatherby, 
no. 5609. Marne: salt marsh toward Dennisville, Pembroke, 
Fernald, no. 1414; brackish shores, Pleasant River, Columbia 
Falls, Svenson & Fassett, no. 1005; muddy shore of Herrick’s Bay, 
Flye’s Point, Brooklin, A. F. Hill, no. 1345; fresh or slightly 
brackish border of salt marsh, South Thomaston, Bissell, Fernald 
& Chamberlain, no. 8933; wet rocky shore, Matinicus, July 19, 
1919, C. A. E. Long; sandy salt marsh, Bristol, Z. B. Chamberlain, 
no. 695, Dinsmore & Chamberlain, no. 839; salt marsh, Bath, 
August 23, 1911, Bissell; Foster's Point, West Bath, 1892, Kate 
Furbish; salt marsh, Hardin ngs’, Brunswick , September 13, 1891, 
Kate Furbish, September 27, 1898, Chamberlain, no. 936; Kenne- 
bunkport, August 7, 1888, Kennedy; mouth of York River, York, 
Bicknell, 1156, Fernald & L ong, no. 12,845. NEw Hamp- 
SHIRE: ditch near border of salt marsh, Hampton Falls, August, 
, A. A. Eaton. MASSACHUSETTS: Manchester, H. D. Tho- 
reau; salt marsh, West Manchester, F. 7. Hubbard, no. 73; 
Somerville, 1882, C.E. Perkins; Watertown, July 17, 1880, C. E£. 
Perkins; salt marsh, Scituate, ‘September , 1901 W. P. Rich, 
September 13, 1914, C. H. Kno wlton; swale, West Barnstable, 
St. John & White, no. 941. Camsucnont: salt marsh, Milford, 

. H. Eames, no. 39. 

Scirpus maritimus, var. Fernaldi, forma agonus, is the eastern 
North American plant treated by Beetle in Am. Journ. Bot. 
xxix. 84, 85 (1942) as typical S. maritimus of Europe. It is quite 
like S. maritimus, var. Fernaldi (Bicknell) Beetle, 1. c. 85, except 
in having bifid styles and thin plano-convex to lenticular achenes. 
Throughout the range of var. Fernaldz, with trigonous achenes, 
and its forma agonus the two show the same range of variation 
in spikelets and habit, with a dense glomerule or more com- 
monly with well developed rays, with the latter terminated by 
single to several and glomerulate spikelets, with the spikelets 
short and ovoid as in the type of S. Fernaldi Bickn. or slender 
and lance-cylindric, running up to 4 em. or more long, as in the 
type of S. novae-angliae Britton. Both S. Fernaldi and S. novae- 
angliae were based upon specimens with trigonous achenes; and 
in that series as well as in the one with plano-convex or lenticular 
and relatively thin achenes there is no line of demarcation evident 
by which the plants with ovoid and lance-cylindric spikelets can 
be separated. S. novae-angliae was published as a species in 
1898, S. Fernaldi in 1901. In the varietal rank they are of 
identical date, published in Ruopora, viii. 163 (1906) as S. 


290 Rhodora [Jory 


campestris, var. novae-angliae (Britton) Fern. and var. Fernaldi 
(Bicknell) Bartlett, but since the latter was transferred to S. 
maritimus as var. Fernaldi while Beetle considered S. novae- 
angliae to be a variety of S. robustus Pursh, S. robustus, var. 
novae-angliae (Britton) Beetle, 1. c. 87 (1942), they are again of 
even date. Under these cirenustances it seems less disturbing to 
maintain under S. maritimus the first of the two which was 
treated as a variety of that species. 

It is not clear to me why Beetle places Scirpus novae-angliae 
under S. robustus. The latter species is a beautifully distinct one 
of tropical America, following the Atlantic coast northward to 
Massachusetts, the Pacific to California. Its plump ellipsoid- 
ovoid to thick-cylindric blunt or bluntish spikelets are rufescent 
or fulvous, the seales (especially the outer) with very prolonged 
awns. Its leaf-sheaths have very characteristic orifices, the 
strong ribs running up the summit to the semicircular or promi- 
nently convex dark scarious ligule. S. novae-angliae, on the other 
hand, like European S. maritimus and American S. Fernaldi, has 
the ligule V-shaped, with truncate or concave (rarely low- 
convex) summit and the nerves at the summit of the sheath are 
slender and relatively inconspicuous. The ovoid to lance- 
cylindric acute to acuminate spikelets are, as in S. maritimus, 
castaneous to fuscous or blackish, and the awns of the scales are 
relatively short. 

From typical European Scirpus maritimus, our var. Fernaldi 
is distinguished merely on size, the leaves running higher on the 
culm, and upon a slight difference in shape of achene. Our 
plant is usually taller and coarser, with leaves mostly 6-15 mm. 
broad (as opposed to the “+ 4 (7)”—Hegi, of the European); 
the spikelets of ours are 1.2-4 em. long (‘‘bis 2 em. lang” in 4. 
maritimus—Hegi); and the achenes are generally more broadly 
obovate and more gradually rounded to the broad base, a8 OP- 
posed to the narrower-obovate achene of S. maritimus in which 
they taper more cuneately or even with a slight concave curve to 
the slender base. 

I do not know how Beetle arrived at the conclusion that true 
(Eurasian) Scirpus maritimus has “Style normally 2-fid” 
(Beetle, 1. c. 87), for the consensus of statements of European 
taxonomists makes the 2-fid style very exceptional: ‘a t y pic us 


1943] Fernald,—North American Species of Scirpus 291 


Narben 3’’—Ascherson & Graebner; “Stigmates 3 
tar? 2 ox )’—Rouy; phages 3, or rarely 2”—Babing- 
ton; “‘Style Sole?’ Heaton “Narben 3, seltener 2”—Hegi. 
Although true S. maritimus (with 3 he and trigonous 
achenes) is in Europe the common form, while the so-called var. 
digynus (Simonk.) Godr. is there called rare, with us typical var. 
Fernaldi (with trigonous achenes) and its forma agonus are about 
equally common. At least, in the area from the Gulf of St. 
Lawrence to Connecticut, where the two somewhat alternate 
their colonies, the representation before me shows 51 sheets of 
typical var. Fernaldi (including S. novae-angliae) and 58 of 
forma agonus. Within this area the two are about equally 
common; from New York to Virginia the representation before 
me is all of typical var. Fernaldi. 

S. paLuposus Nels., var. atlanticus, var. nov., a forma typica 
‘esate foliis caulinis plerumque 2-4, sub medio gestis, vaginae 
venis apice tenuibus vix prominulis; spiculis brunneo-castaneis 
vel ee ee antheris plerumque 2-3.5 mm. longis, 
filamentis inclusis vel subexsertis; pchaene Aohahdd obuestia 
vel suborbicularibus rariter cuneatis olivaceis vel atro-brunneis.— 
Salt marshes and saline shores, Gulf of and lower River St. 
lave Quebec, to northern New Jersey; central and western 
New York. Typu: salt marsh, Bunbury, Prince Edward Island, 
Augeat 28, 1912, Fernald, Long & St. sie. no. 6982 (in Herb. 

ray.). 

Scirpus paludosus Nels. in Bull. Torr. Bot. Cl. xxvi. 5 (1899), 
like the identical S. campestris Britton in Britt. & Brown, IIl. 
FI. i. 267, fig. 626 (1896), not Roth (1795), is the plant of western 
North America, extending eastward to Manitoba, Minnesota, 
Nebraska and Missouri. Var. atlanticus is the plant of the 
Atlantic coast, with an isolated and in some ways transitional 
area in the saline region of interior New York. True S. paludosus 
is a very pale green plant, with culms mostly 0.5-2 em. thick at 
base and 0.5-1.5 m. high; the cauline leaves mostly 3-5 (-6), 
with sheaths ascending well above the middle of the culm (but 
in starved colonies the culms lower and the leaves fewer and 
borne chiefly below the middle), their blades mostly 0.5-1.5 cm. 
broad, the veins near the orifice of the sheath prominent and 
usually thickened; spikelets whitish-brown to drab or pale 
brown; anthers 3.5-5 mm. long, standing well out of the spike- 


292 Rhodora [JULY 


lets on elongate filaments, the filaments, after falling of anthers, 
usually showing above the scales and twice to thrice the length 
of the achene; achene cuneate-obovate, rarely roundish, pale 
brown to olivaceous. 

S. paludosus, var. atlanticus is not so pale; its culms are 1.5- 
7.5 dm. high, 2-8 mm. thick at base; the cauline leaves are 
usually 2-4 and borne chiefly below the middle (though in 
exceptional plants, perhaps mixed with S. maritimus, var. 
Fernaldi, more numerous and running high on the culm), the 
blades 1.5-9 mm. broad, the veins near summit of sheath delicate 
and inconspicuous; spikelets chestnut-brown to blackish-fuscous; 
anthers 2-3.5 mm. long, mostly not exceeding scales, the old 
filaments rarely exserted and shorter than to about twice the 
length of the achene; the achene rounded-obovate to suborbicu- 
lar, only exceptionally cuneate, olivaceous to deep brown. In 
central and western New York the plant geographically some- 
what intermediate between true S. paludosus and var. atlanticus 
has the anthers up to 4 mm. long and the achene often cuneate 
at base. Along the coast of the Gulf of St. Lawrence and of the 
Atlantic occasional colonies seem like hybrids of S. paludosus, 
var. atlanticus and S. maritimus, var. Fernaldi. 

True western Scirpus paludosus has an evident tendency to 
branching of the inflorescence; var. atlanticus not. Of the 186 
inflorescences of S. paludosus before me 124 have simple or at 
least closely crowded glomerules, 62 (33%) have 1 or 2 (rarely 
3 or 4) obvious elongate rays. Of 420 inflorescences of var. 
atlanticus only 20 (less than 4%) show a single (in 1 case 2) short 
ray. On the Atlantic coast the plant with fuscous spikelets 
frequently on definite or elongate rays is S. maritimus, vat. 
Fernaldi. Of the 210 inflorescences of it before me 172 (nearly 
82%) have definite (sometimes forking) rays (26 with 1 ray, 77 
with 2, 28 with 3, 20 with 4, 13 with 5, 13 with 6, 6 with 7, 5 with 
8,1 with 11, 1 with 12 and 4 with 13). 

Although Beetle in Am. Journ. Bot. xxix. 83 (1942) cites 
Scirpus paludosus as occurring in the“ East Inpies: Ekman 1325” 
(this citation placed between New Jersey and Minnesota), 
there is grave doubt about it. I have not seen no. 1325 and can, 
consequently, not check its identity; at least, it presumably 
came from the West Indies. 


1943] Fernald,—North American Species of Scirpus 293 


S. expansus, sp. nov. (Tas. 767, Fic. 1-3), planta habitu S. 
sylvatici; culmo 0.5-1.6 m. alto ad basin 0.6-1.5 em. crasso 
superne scabro; foliis 1-2.5 em. latis, vaginis coriaceis valde 
septato-nodulosis; involueri foliis 3 8, imis panicula superantibus; 
panicula 1-3 alta radiis adscendentibus vel divaricatis; 
pedicellis valde pips, spiculis 3-5 mm. longis, plerumque 
glomerulatis; squamis valde carinatis apice subulatis; antheris 
1.3-1.6 ongis.—Spring-heads, borders of rills, springy 
meadows, swales, etc., southwestern Maine to southern Michi- 
gan, south to Georgia. The following, selected from a large 
series, are representative. Marne: South Poland, 1895, Kate 
Furbish; Typha swamp, bank, Presumpscot River, August 13, 
1903, Collins & Chamberlain, no. 614 (TYPE in Herb. Gray.) ; 
springy spot, Great Chebeague Island, Fernald, no. 1401; swale 
at border of salt marsh, Ah Fernald & Long, no. 12 851; 
swamp, York Harbor, July 1901, F. T. Hubbard. New 
HAMPSHIRE: Hanover, ge “4878, Jesup; Ore Hill, Warren, 
July 26, 1910, E. F. Williams; shore of Johnson Creek, Madbu ury, 
Hodgdon, no. 2629. Vermont: Gulf Brook swamp, Townshend, 
September 10, 1912, L. A. Wheeler. MassacHusEtTts: south e 
of Horn Pond, Woburn , August 4, 1869, Wm. Boott; South Sud- 
bury, June 17, 1902, Rich, Fernald & Williams; Westfield, July 
10, 1860, Wm. ’ Boott; brook in cold bog, Scomy ‘August 27, 1916, 
Churchill; brooksides, Uxbridge, June 23, 1876, Morong; shore 
of small pond, Granville, FOC; Ptarions no. 388; ew Salem, 
July 28, 1931, Goodale, Potshay & St. John; brookside, New 
Marlboro, August 30, 1902, Ralph Hoffmann. CONNECTICUT: 
West ane ane 13, 1901, H. J. Koehler; swamp, Southing- 


ton, Bissell, - New Hartford, August 10, 1883, Chas. 
Wright; slong Side Brook, Waterbury, Blewitt, no. 456; open 
swamp, ford, HE. H. Eames, no. 4000. New York: wet 


ditch, F en ome Washington County, Burnham, no. 46; along 
a cold stream, 4 miles south of Utica, Haberer, no. 1184; Water- 
ville, Oneida ‘County, August 18, 1917, House. New JERSEY: 
Rosenkraus Run, Sussex County, August 11, 1917, E. B. Bar- 
— PENNSYLVANIA: open rshy, springy swale, Lehigh 
Gap Station, Pretz, no. 13,253; masons ditch, 2 miles north of 
Sadsburyville, Chester County, June 29, 1924, H. E. Stone; Big 
Meadow Run, Farmington, Fayette County, June 4 4, 1931, Core. 
DELAWARE: along brooks, Centreville, ni : 1869, Commons. 
District oF COLUMBIA: Washington, 1881 (‘‘the most southern 
locality known”), Scribner. Virern1a: South Fork of Holston 
River, St. Clair’s Bottom, Smyth County, July 30, 1892, Small. 
Nortu CAROLINA: near Hickory, Catawba County, Heller, no. 
274. Gxorata: in cool shaded brook at northern base of Stone 
Mountain, Harper, no. 205. MicHiGAN: swamp, Grand Rapids, 
July 20, 1900, Emma J. Cole. 


294 Rhodora [Juny 


Scirpus expansus is the plant which regularly passes in the 
eastern United States as S. sylvaticus L. of Eurasia. The resem- 
blance is merely superficial, for in most characters there are clear 
distinctions. S. sylvaticus has thinner leaves, with scarious 
sheaths, the summit of the inner band easily friable, the blades 
6-14 mm. wide; in S. expansus the hard and thickish blades are 
1—2.5 em. broad, the coriaceous sheaths strongly septate-nodu- 
lose (in S. sylvaticus only faintly, if at all, so), the summit of the 
inner band firmer. In S. sylvaticus the relatively slender culm is 
smooth to summit; in S. expansus the usually coarser culm is 
scabrous at summit (for 1-5 em.). In S. sylvaticus the longest 
smooth and thin leaf of the involucre is 0.7-2 dm. long, only 
rarely exceeding the panicle; in S. expansus it is firm and harshly 
scabrous beneath and 1.5-3 dm. long, often overtopping the 
panicle. In S. sylvaticus the mature panicle is lax, with loosely 
spreading to recurving longer rays, the panicle 1-2 dm. high; in 
S. expansus the longer rays (Fic. 4) are more stiffly ascending to 
divergent, only the short basal ones much recurving, and the 
panicle, when well developed, is 1.5-3 dm. high. In S. sylvaticus 
the spikelets (Frias. 4 and 5) are 3-4 mm. long and in glomerules 
of 2-5, the ultimate lateral pedicels often terminated by single 
spikelets; in S. expansus the principal glomerules have 3-12 
(rarely to 40) spikelets 3-5 mm. long, only a rare ultimate pedicel 
with a solitary one. In S. sylvaticus the pedicels are minutely 
scabridulous; in S. expansus closely pilose. In S. sylvaticus the 
scales of the spikelet (ric. 5) are blunt or barely mucronate by 
the extension of the relatively weak midrib; in S. expansus they 
(FIG. 3) have subulate-acuminate tips, through the extension of 
the very prominent keel-like midrib. In S. sylvaticus the anthers 
a 5) are 0.7-1 mm. long; in S. expansus (ria. 3) 1.3-1.6 mm. 
ong. 

Forma Bissellii (Fernald), comb. nov.  S. sylvaticus, var. 
Bissellii Fernald in Ruoponra, ii. 21 (1905) ; S. microcarpus, var. 
Bissellii (Fern.) House in Bull. N. Y. State Mus. nos. 243-244: 
18 (1923). S. sylvaticus, forma Bissellii (Fern.) Carpenter in 
Dole, Fl. Vt. ed. 3: 76 (1937). 

: Forma globulosus, f. nov., glomerulis globosis 7-12 mm. 
diametro, spiculis 20-60 congestis valde imbricatis 3-4 mm. 
longis—Nrw York: Lyon’s Falls, Lewis County, August, 1882, 
J. V. Haberer (type in Herb. Gray.). 


1943] Fernald,—North American Species of Scirpus 295 


A very unusual extreme, the inflorescence, with large globose 
heads of very numerous spikelets superficially suggesting the 
most extreme Juncus canadensis J. Gay. Not to be confused with 
forma Bissellii. That has the large glomerules 1-2 cm. in diam- 
eter and composed of loosely divergent linear-cylindric spikelets 
6-14 mm. long. 

Since some botanists place Scirpus expansus under the western 
North American S. microcarpus Presl, it is well to point out that 
S. microcarpus has 2 stigmas and lenticular achenes; S. expansus 
3 stigmas and trigonous achenes. In S. microcarpus the smooth 
or nearly smooth leaf-sheaths and the leaf-blades are thinner and 
smoother than in S. expansus, the involucre only slightly if at all 
exceeding the inflorescence; as contrasted with the heavier and 
septate-nodulose sheaths of S. expansus, the broader and firmer 
blades scabrous beneath and the usually longer involucres. In 
its thin and relatively narrow leaves with smooth or nearly 
smooth sheaths S. microcarpus is more like S. sylvaticus of Eura- 
sia. In the texture of its scales and the size of its panicle (0.8-2 
dm., only exceptionally—3 dm. high) it is more like the latter, 
but the glomerules have many more and usually longer spikelets. 
Unless all members of the section are to be reduced to an all- 
inclusive S. sylvaticus, with several constant and geographically 
isolated subdivisions, some with 2 stigmas, others with 3, these 
fundamental characters of the pistil associated with other 
characters, the three North American members of the series, 
S. microcarpus Presl, S. rubrotinctus Fern. and S. expansus are 
well marked species. 


n PLATE 767, FIGS. 1-3 are of Scrrpus ExPANsuS, from the Type: Fic. 1, 


inflorescence, X 24; FIG. 2, portion of inflorescence, to show characteristic 
ascending branches with spikelets mostly glomerula te, xX 3; Fic. 3, » sit et 
with anthers, X 10. Fies. 4 and 5, 8S. syLvaTicus from Tassin, 


Boulin: r1a. 4, small portion of inflorescence to show characteristic Roceed 
pier ea with intial spikelets often solitary, < 3; Fic. 5, spikelets with 
anthe 0. 


RUBROTINCTUS heliny forma radiosus, f. nov., ores 


me nos. 12,853 (TypPp in Herb. Grey) 12,854; ditch near 
arf, Camden, August 11, 1902, G. G. Kenn nedy. 
The counterpart in Scirpus rubrotinctus of S. expansus, forma 
Bissellit. 


296 Rhodora [JULY 


. ATROVIRENS Muhl., var. GeorGiaANus (Har per) Fern., 
forma peuaiar oases nov. , glomerulis dense confertis in capitulo 
: diametro aggregatis. TYPE: A a Veazie, 
fuine. Auguat 25, 1908, Fernald (in Herb. Gra 
S. ATROVIRENS uhl., var. GEORGIANUS wer rper) Fern., 
forma oe f. nov., spiculis anguste cylindricis ad 
. TYPE: wet shore of Housatonic River, Newtown, 
ieee oan August 17, 1928, E. H. Eames, no. 10,692 (in 
Herb. Gra 
S. POLYPHYLLUS s Vahl, forma macrostachys (Boeckl. Me ee 
nov. Var. macrostachys Boeckl. in Linnaea, xxxvi. 731 (1870). 


VI. THE IDENTITY OF SCLERIA SETACEA OF 
POIRET 


One of the most definite and easily recognized species of 
Scleria is the handsome and tall cespitose but lax plant with soft 
and almost wing-angled easily compressed culms up to 1 m. high; 
broad, lax leaves up to 8 mm: wide; long, drooping, filiform, 
lateral peduncles and loosely fastigiate terminal panicles up to 
4 cm. long; the globose and reticulate achenes with spirally 
arranged pits and pubescent surfaces, the hypogynium with 3 
broad erect lobes. This relatively tall species occurs from warm- 
temperate eastern South America, the West Indies and Florida 
to eastern Texas and Mexico, north in the Atlantic States to 
Long Island, and locally in the interior to Indiana. 

This is the handsome plant beautifully described as S. laxa by 
Torrey in Ann. Lye. N. Y. iii. 376 (1836), Torrey’s appropriate 
name unfortunately preempted by the Australian S. lava R. Br. 
(1810). Somewhat earlier Muhlenberg had described the same 
plant from North Carolina as S. reticularis Muhl. Deser. Gram. 
266 (1817), he mistaking it for S. reticularis Michx. (1803). This 
confusion was soon noted, however, and there resulted two 
names: S. Muhlenbergii Steud. Nom. ed. 2, ii. 543 (1841), based 
upon “‘S. laxa. Torr. (non R. Br.) S. niinlaris Mhlbrg. Pursh. 
(non Michz.)”; and, a little later, S. Torreyana Walp. Ann. iii. 
696 (1852), based on S. lara Torr. Other but later names were 
given to the plant; S. Muhlenbergii Steud., however, being the 
earliest available one, we may in this note omit the others, 
especially since their exact identification is not now possible. 
For this tall plant with compressed-trigonous culms, broad and 


CL LS LC CLG, 


1943) Fernald,—The Identity of Scleria setacea of Poiret 297 


flat leaves and terminal lax panicles up to 4 em. high Core, in his 
American Species of Scleria in Brittonia, ii. 79 (1936), takes up 
S. setacea Poir. in Lam. Encye. vii. 4 (1806); and others, as un- 
wittingly as myself, have trustingly followed him. 

The photograph of the type of S. setacea Poir., however, 
thoroughly agrees with the original description: 

8. Sciirie A feuilles sétacées. Scleria setacea. 

Scleria culmo foliisque setaceis, glaberrimis; spicis axillaribus, minimis, 
longé pedunculatis; spiculis angustis, pedicellatis seu igre ibus re 

es seta sont courtes, fibreuses, fasciculées: i n éléve des tiges 
nombreuse haute de huit a dix pouces & grin +easanrtlee, 
fines, bres triangulaires : 

De Vorifice de chaque gaine, méme a partir de celles du bas, il sort un 
pédoncule droit, trés-fin, long d’un pouce & plus, terminé par deux, trois, 
& peine quatre épillets pedioellta, quelquefois un ou deux sessiles, petites, 
étroits, ovales-oblongs, aigues, d’un roux-clair, ee de petites ‘bractées 
courtes, sétacées, a peu prés de la longeur de iy pi 

Cette plante croit dans |’Amérique; elle e été recuaillie par M. Ledru 
& Porto-Ricco (V. s. in herb. Lam.) 

It is most difficult to imagine how Scleria Muhlenbergii could 
ever have been identified with S. setacea, described as having 
setaceous culms up to 10 inches high, setaceous leaves, small red 
spikes, ete., the photograph before me showing the terminal 
inflorescence to be compact and 6-8 mm. high. Search in West 
Indian Scleria shows nothing like it, S: Brittonii Core and S. 
georgiana Core both having elongate horizontal rhizomes and 
larger terminal inflorescences; but the slender and acuminate 
reddish spikelets and the dense non-rhizomatous base at once 
suggest Rhynchospora. Turning to that genus, the type of 
Scleria setacea Poir. is promptly matched, even to the short- 
peduncled inflorescences in the lower leaf-axils, by R. seTacra 
(Berg.) Boeckl., based upon Schoenus setaceus Bergius (1772). 
The fact that Poiret, in 1806, used the same specific name was 
merely coincidence; but the fact that his Sclerta setacea coincides 
with Schoenus setaceus, therefore with Rhynchospora setacea, 
removes that wholly misinterpreted name from the valid species 
of Scleria. The many North American, West Indian and South 
American specimens recently marked ‘‘Scleria setacea Poir.”’, 
should be changed to S. MUHLENBERGI Steud. 


298 Rhodora |JULY 


VII. WHAT IS ANGELICA TRIQUINATA? 
(Plates 768 and 769) 


In his Flora Boreali-Americana, i. 167 (1803) Michaux de- 
scribed from ‘‘Canada”’ a single species of Angelica as 

TRIQUINATA. A. petiolo tripartito; partitionibus 

pinnato-quinquefoliolatis; foliolis in- 
ciso-dentatis; terminalium eon rhom- 
beo, sessili, lateralibus decursiv 
Obs. Glabra: pedunculo caelieee minutissima 
pube subcandicantibus. 
Hab. in Canada 

Pursh (1814) took it up, literally copying Michaux’s diagnosis 
but giving the range ‘“‘In Canada and on the mountains of 
Virginia’, from which it is probable that Pursh was stretching 
the name to cover the later published Angelica Curtisii Buckl.; 
and others, Bigelow, Fl. Bost. (1814) for instance, accepted it 
for the common New England A. atropurpurea L. 

In 1818 three different authors independently considered 
Angelica triquinata to be the plant of dry woods and thickets 
southward, with stem closely tomentulose above, with thick 
lanceolate to oblong regularly and closely serrate leaflets, the 
upper or bracteal leaves (subtending inflorescences) reduced to 
linear-cylindric or lanceolate tubular sheaths with tiny blades, 
the plant which Walter, Fl. Carol. 115 (1788) had well described 
as Ferula villosa, i. e. Angelica villosa (Walt.) BSP. Muhlenberg, 
Cat. ed. 2: 30 (1818), substituted for A. triquinata Michx. his 

own A. hirsuta, saying without quibble ANGELICA 

“1 hirsuta, triquinata, Mx.” and giving the single descriptive 
word, « downy”, Muhlenberg’s plant coming from “Pens. fl. Aug: 
N. Eb.” At best A. hirsuta Muhl. is a nomen subnudum; but 
since he used the name as a substitute! for A. triquinata Michx. 
(1803) it is illegitimate. Nuttall, also in 1818, took up A. 
triquinata, obviously for Ferula peste Walt. His description of 
the plant “Common around Philadelphia” was good, except for 
the phrase, “Leaves sharply and incisely serrate’, eV ridently 
borrowed from Michaux; and, obviously not understanding 
Michaux’s plant, he commented on the plant of ‘Canada to 


‘In his Cat. ed. 1; 31 (1813) Muhlenberg had published the trinomial Angelic? 
_— triquinata as a “downy” plant of “Pens.” It was on ba 2nd edition (1818) 
e cited A. triquinata Michx. as a synonym of his A. h 


1943] Fernald,—The Identity of Scleria setacea of Poiret 299 


Carolina” as “‘Certainly a genuine species’. In the same year 
Sprengel, Species Umbelliferarum minus cognitae, 69, t. vi. fig. 
12 (1818), based his Pastinaca triquinata upon Angelica triquinata 
Michx., “‘ Habitat in Canada et Virginia” (borrowed from Pursh), 
and gave a fine description and excellent illustration of Ferula 
villosa Walt., changing the ‘‘ Foliolis inciso-dentatis; terminalium 
impari rhombeo, sessili’’ ete. of Michaux (impossible for Ferula 
villosa) to “‘foliolis oblongis”’ of F. villosa, and noting the reduced 
upper blades and sheath-like petioles of the latter, ‘‘Superiora 
minus divisa vaginis petiolaribus insidentia’’. 

Many authors followed these false leads and the name Angelica 
triquinata Michx. became general, as the first supposedly avail- 
able combination under Angelica (following the sensible, therefore 
abandoned, ‘“‘Kew Rule”) for Ferula villosa Walt.; not merely 
Nuttall and Sprengel so using it, but Elliott, Torrey, DeCandolle 
and others accepting the identification. When Sprengel took it 
up as the basis of Pastinaca triquinata (Michx.) Spreng. he did 
so in a work on Umbelliferae ‘“‘minus cognitae”, and his phrase 
very definitely described his understanding of Michaux’s plant. 
When he first studied Michaux’s herbarium Asa Gray saw the 
error, his memorandum reading ‘‘ Not the least what we call A. 
triquinata. DC. has confounded strangely if he ever saw Michx’s. 
plant . . . It is A. atropurpurea?—tho’ very poor specimen’’. 

Gray’s guess is hardly better than that of his predecessors, for 
even his ‘‘tho’ very poor specimen” does not endow the Michaux 
plant with the characters of A. atropurpurea. In the latter the 
upper leaves, which subtend inflorescences, have large, inflated 
and round-tipped stipular sheaths extending quite to the summit 
of the obscure petiole, so that the 3 divisions of the leaf are 
essentially sessile, and the leaflets are not incised-dentate. 
Furthermore, the subspherical umbel has 20-46 rays. In 1903 I 
made a photograph of the Michaux Tyre at Paris. This (PLATE 
768) shows, X %, the incised-dentate leaflets, the terminal 
unequally rhombic, the lateral decurrent, as described by 
Michaux. It also shows the stipular sheath narrow and tapering 
to the elongate and naked upper half of the petiole; and the 
axillary branch has one slender and tubular bladeless sheath 
and an immature umbel with only 7 ascending rays. I have 
been over, with Dr. Hugh M. Raup, all known eastern North 


300 Rhodora [JULY 


American Umbelliferae. Every one of them is quickly rejected 
as not Michaux’s plant, except the Alleghenian A. Curtisii 
Buckley, Am. Journ. Sci. xlv. 173 (1843) which follows the upland 
and the mountains from Pennsylvania to Georgia. Buckley’s 
species gives a good match for A. triquinata, a more modern 
specimen from the Blue Ridge of Virginia (near Luray, alt. 3600 
ft., Steele & Steele, no. 213), reproduced as PLATE 769, showing a 
portion of a plant, X 1%, with the tapering sheath (at base), the 
elongate petiole (near base), the incised leaflets, the terminal 
one rhombic, the lateral (upper) ones decurrent and the few- 
rayed umbel of the Michaux type. Buckley’s plant, from “High 
mountains of North Carolina” had “petioles large, long, and 
sheathed at the base; segments of the leaves 3-5, leaflets large 
and deeply laciniate”. It is evident that ‘‘Canada”’ of Michaux’s 
label and description was a clerical error for Carotina, Michaux 
having collected extensively in the Carolina mountains. It is 
clear, then, that the name Angelica Curtisii Buckley (1843) must 
give way to A. TRIQqUINATA Michx. (1803 

Another reason for thus reviewing the interpretations of 
Angelica triquinata is the fact that the name A. villosa (Walt.) 
BSP. Prelim. Cat. N. Y. 22 (1888) is a later homonym, invali- 
dated by the fully described A. villosa Lagasca, Gen. et Sp. Pl. 12 
(1816), a plant of Asturia. Since, furthermore, A. hirsuta Muhl. 
was an illegitimate substitute for A. triquinata Michx., the name 
hirsuta used very carelessly for a plant which is not hirsute but 
said to be “downy” (many a youth with downy face has anxiously 
waited for it to become hirsute), the name A. triquinata, long 
used for A. villosa, seemed to come into the running. That it 
cannot be taken up for A. villosa (Walt.) BSP. should be suffi- 
ciently clear. 

Only one other name is sometimes cited as synonymous with 
Angelica villosa (Walt.) BSP. This is Cicuta venenosa Greenway 
in Trans. Am. Phil. Soe. iii. 235 (1793), incorrectly cited by 
Pursh as C. venenata in his synonymy of Ferula villosa. Green- 
way, not now a well known botanist, was ‘‘ Dr. James Greenway; 
of Dinwiddie-County, in Virginia.’ His account of the plant as 
an acute poison when eaten and his very detailed descriptio® 
are conclusive. It is unnecessary to quote the whole, but Dr. 
Greenway’s solicitude for the Philosophical Society, to whom he 


1943] Fernald,—The Identity of Scleria setacea of Poiret 301 


communicated his paper on February 19, 1790, is worth quoting: 
“T will here insert the description, as it stands in my catalogue, 
first, in botanical terms, for such as are lovers of that science, and 
then in language, as plainly English as the subject will admit, 
for the sake of those to whom those terms are less familiar.” 


The detailed description, with ‘‘Caulis . . . quatuor pedes 
altus, teres, . . . superne tomentosus.—Folia _petiolata, 


petiolis semi-amplexicaulibus, . . . triternata, bipinnata, 
; foliolis sessilibus, oblongo-lanceolatis, serratis”, and the 
habitat and flowering season (in Dinwiddie County), ‘‘Locis 
campestribus et collibus apricis gaudet: mensibus Julii Augusti- 
que floret”, are wholly indicative of A. villosa. The latter should, 
therefore, be called: 

ANGELICA venenosa page ay), | comb. nov. nr 7. 
Greenway in Trans. Am. Phil. c. ili. 335 (1793). Feru 
pe ie Walt. Fl. Carol. 115 ee A. triquinata sensu Nutt. 
Gen. i. 186 (1818) and later authors, not Michx. (1803). Pasti- 
naca triquinata Spreng. Spec. Umb. 69, t. vi. fig. 12 (1818) as to 
plant described and illustrated, not as to agen of epithet, . 
triquinata Michx. Archangelica hirsuta Torr. 1, N. Am 
622 (1840) as to plant described, only in part joe source ‘of 
epithet, Angelica hirsuta Muhl. Cat. ed. 2: 30 (1 818), an ille- 
gitimate substitute (as treated by Muhlenberg) for the early A. 
triquinata os (1803). A. villosa (Walt.) BSP. Prelim. Cat. 
N. Y. 22 (1888) by inference only, not A. villosa Lagasca, Gen. et 
Sp. Pl. 12 (1816). 

MEMORANDUM REGARDING JAMES GREENWAY.—I am indebted 
to Mr. J. M. Townsend of Petersburg, Virginia, for a reference 
to Castiglioni’s Reise durch die Vereinigten Staaten von Nord- 
Amerika in den Jahren 1785, 1786 and 1787. On p. 274 of this 
German translation we read: 

‘‘Bine Meile von Petersburg liegt die Wohnung des Obristen 
Banister, der eine ausgedehnte Pflanzung besitzt. Er is ein Enkel 
des beriihmten John Banister, der seine Stelle als Professor der 
Botanik und Bibliothekar der Universitit Oxford niederlegte, 
und sich in diesem Thiele von Virginien niederliess, wo er mit 
grosser Miihe und seltenem Urtheil eine Menge der seltensten 
Pflanzen sammelte, und beschrieb. . . Am folgenden Tage 
stattete ich einen Besuch beym D. Greenway ab, der ein Eng- 
lander von Geburt, und ein Liebhaber der Botanik ist. Er hatte 
sich selbst mit den Grundsiitzen des Linneischen Systems be- 


302 Rhodora [JULY 


kannt gemacht, und wusste mehr als 600 Pflanzen zu nennen, 
unter denen einige ziemlich seltene und noch unbeschriebene 
waren.” 

Mr. Townsend also most kindly transcribes for me extracts 
about Dr. James Greenway from the recently (1942) published 
Dinwiddie County “The Country of the Apamatica’’, compiled 
by the workers of the Writer’s Program of the Works Projects 
Administration. From this account (p. 77) I quote: ‘It was 
after he had established himself in Dinwiddie as a man of wealth 
and importance—somewhat Scottish as he was—that he turned 
his whole attention to natural history and botany. Some 40 
volumes that deal with plants of Virginia and North Carolina 
flowed from his pen and won him honorary membership in 
several European societies and friendships with scholars the 
world over. Thomas Jefferson, that great patron of learning, 
frequently corresponded with Dr. Greenway and gave consistent 
encouragement to the botanical investigations of the Dinwiddie 
scientist.”’ 

I have hunted in vain, with the collaboration of Dr. Schubert, 
for the ‘‘40 volumes that deal with plants’. Pritzel (Thesaurus) 
did not know of them. Dryander, Cat. Bibl. Hist.-Nat. Banks, 
iii. 542 and 598 (1797) and the Royal Society Catalogue, iii. 5 
(1869) could muster only the two short papers in vol. iii. of the 
American Philosophical Society’s Transactions (1793), the first 
on Cassia Chamaecrista as a soil-renovator, the second the account 
of Cicuta venenosa. If, contemporary with Thomas Walter 
(1788) and prior to Michaux (1803), Pursh (1814) and Elliott 
(1816 et seq.), there were two score volumes published, describing 
in such clear diagnoses as that of Cicuta venenosa 600 plants of 
Virginia and North Carolina, their discovery would be of utmost 
importance and extremely disconcerting. 

Through the most helpful cooperation of Messrs. Jack Dalton 
of the Alderman Library at the University of Virginia and R. W. 
Church of the Virginia State Library the excitement among 
taxonomists over the ‘40 volumes that deal with plants of 
Virginia and North Carolina [which] flowed from his [Greenway’s] 
pen” is now abated. Under date of November 21, 1942, Mr. 
Dalton wrote: ‘‘The reference . . . puzzles us as much as it 
does you. I have examined our catalogues, the calendars of 


1943] Fernald,—The Identity of Scleria setacea of Poiret 303 


Jefierson’s Correspondence, the bibliographies, English and 
American, available here, and Swem’s Index, and have not been 
able to find the slightest evidence of the existence of the forty 
volumes or the voluminous correspondence of Jefferson. Dr. 
Greenway’s name is not mentioned in our calendar of Jefferson’s 
correspondence”. Mr. Church, examining the notes made by 
the Virginia Writer’s Project in connection with their history of 
Dinwiddie County, found that the statement concerning the 
forty volumes was derived from the Memoirs of Lieutenant 
General Winfield Scott, vol. 1, pp. 3-6 of the edition of 1864. 
Here is the quotation as filed by the authors of the history of 
Dinwiddie County: 

“His professional reputation brought him patients from a 
wide circumference, but, as he became rich, he gradually with- 
drew from the practice of medicine, and gave himself up to the 
culture of polite literature and natural history, particularly 
botany, and left a hortus siccus of some forty folio volumes in 
which all the more interesting plants, etc., of Virginia and North 
Carolina, were described in classical English and Latin.” 

A hortus siccus being a collection of dried plants (an herbarium) 
it is evident that the presumably very accurate descriptions of 
oo which “flowed from his pen” did not reach publica- 


‘ts publishing the genus Greenwaya Giseke, Praelect. Ord. 
Nat. Plant. 226 (1792), wrote: ‘‘in honorem Cl. . . GREEN- 
way, Med. Dris. in Virginia, cujus amicitiam & cum eo commer- 
cium debui epistolarum Cl. Drury, inter Entomologos celebri. 
Misit ille ab a. 1773-1775. plantas Virginicas siccatas, vivas a se 
collectas, ad 400. eo fine ad me, ut novam Floram Virginicam 
juncto labore concinnaremus, sed bello inter Anglos & colonias 
orto, conatus omnis profligatus est, ita ut nesciam num vivus 
adhue supersit nec-ne? = tamen labore utique de Botanica 
meritus est”’. 
(To be continued) 


f 
Fi, anh 


% 


1943] Fernald,—Notes on Hieracium 317 


CONTRIBUTIONS FROM THE GRAY HERBARIUM OF 
HARVARD UNIVERSITY—NO. CXLVIII 


M. L. FERNALD 
(Continued from page 303) 
VIII. NOTES ON HIERACIUM 

Hieracium Robinsonii (Zahn), stat. H. smolandicum, 
subsp. Robinsonii Zahn in Engler, Piasene. iv, 468 (1921). 

Although Zahn placed with doubt the plant collected by 
Robinson & Schrenk in southeastern Newfoundland under the 
Scandinavian H. smolandicum Alma. it differs in so many strong 
characters that I think it wiser to consider it an endemic eastern 
American species. So far as I have yet found there is no evidence 
that any of the European members of the complex Section 
Vulgata are indigenous with us. We have a very diverse lot of 
introduced and weedy plants of the section. H. Robinsonii, 
however, is indigenous along chiefly slaty river-banks in New- 
foundland, eastern Quebec, Cape Breton, northern Maine and 
northern New Hampshire. From the Scandinavian H. smolandi- 
cum our native plant differs in its thick and firm leaves glabrous 
beneath or merely short-pilose along the midrib, the cauline 
leaves twice as many, mostly 4-10, and more gradually decreas- 
ing in size to the summit (in H. smolandicum the thin leaves 
copiously long-villous, the cauline 3-6 and confined chiefly to 
lower half of stem), and the phyllaries caudate-attenuate (in H. 
smolandicum blunt or merely acuminate). 

It is at present quite impossible to identify with any certainty 
the diverse series of introductions belonging in the macro-species 
Hieracium vulgatum. In his inclusive H. vulgatum Zahn recog- 
nizes 144 so-called ‘‘subspecies” and many varieties, the sub- 
species sometimes grouped under such unilluminating heads as 
“a. Folia latiora’’ opposed to “b. Folia ore recone of A. 
Squamae dorso vel margine parcefloccosae’’ opposed to “b. 
Squamae efloccosae atrae nitentes’’, the latter series containing 


“subspecies” with “Involucra . . . interdum parcefloc- 
cosa”? (subsp. no. 131) or ‘“‘squamis . . .  parcefloccosis” 
(subsp. no. 132), the former series with “subspecies” with 


“squamis . . . subefloccosis’. In other cases series of 20 
or 30 “subspecies” are enumerated without even these hair- 


318 Rhodora [AuGUsT 


splitting differentiations. Surely, without a clearer under- 
standing of and a saner guide to the European variations of this 
heteromorphic group, it is better for the present to pass all the 
introduced plants of § Vulgata, with basal leaves narrowed to the 
petiole as H. vulgatum. 

Returning to Hieracium Robinsonii, Zahn’s very full descrip- 
tion of the Robinson & Schrenk plant is accompanied by the fol- 
lowing synonymy: “An H. molle Pursh Fl. Am. sept. II, (1816) 
503? H. vulgatum Gray Syn. Fl. N.-Am. I. 2 (1884) 429”. As 
to the plants of the Synoptical Flora designated by Gray (on p. 
424, not 429) as H. vulgatum from “Labrador, Kohlmeister, &c. 
Canada, on shores of the Lower St. Lawrence (Macoun), there 
perhaps introduced”, they are of two very different species. 
The Labrador element is H. groenlandicum Arv.-Touv., which is 
native in outer Labrador, northern Newfoundland, and Anticosti, 
as well as Greenland, while the Macoun material seen by Gray 
was a sheet of Macoun’s (August, 1883) with two species, the 
data reading: ‘Shores of the St Lawrence from Point Levis to 
Anticosti. Locality. Point Levis (Large specimen), River de 
Brig, Anticosti’. The plant from Levis is the ubiquitous weedy 
species which abounds in the region of Quebec city, H. vulgatum 
Fries; the Anticosti specimen is good H. groenlandicum. As for 
H. molle sensu Pursh, not Jacquin, that was evidently H. groen- 
landicum also; incidentally the date of Pursh was 1814, not 1816 
as given by Zahn. 

The type and only specimen of H. Robinsonii seen by Zahn 
came from the southeastern side of Newfoundland, more than 
550 miles southeast of the mouth of the St. Lawrence River". 
Nevertheless Zahn, with the Germanic inaccuracy regarding 
American geography to which we have become accustomed, 
concocted the citation: ‘Am unteren Lorenzstrom: Cataracts 
of Rocky River in Newfoundland”. The following are the sta- 
tions for H. Robinsonii (often distributed as H. vulgatum) repre 
sented in the Gray Herbarium and the Herbarium of the New 
England Botanical Club. 

NEWFOUNDLAND: crevices of rocks, infrequent, South Arm 
River, Bolyeiea and Cataracts of Rocky River, Robinson & 

1 Equivalent in distance to placing Berlin in England, on the western coast of 


Norway, on the Gulf of Riga, on the Dneiper River or on the Mediterranean, OF 
equivalent in distance to placing Vienna on the Seine or in Belgium or on Sardinia. 


1943] Fernald,—Notes on Hieracium 319 


Schrenk, no. 227 gibi ledges and gravel along Waterford 
River between Waterford "Bridge and St. John’s, Fernald & 
Wiegand, no. 6439. QurBrEc: Saguenay River, Roberval, July 
18, 1892, G. . Kennedy; ledges along River Ste. ‘Anne des Monts, 
Gaspé Coun nty, August, 1905, Collins & Fernald; ee below 
Fraser Falls, Riviére du Loup, July 11 and 13, 1904, Collins & 
Fernald; rocks by Lower Fall, Riviére du Loup, A. S. Pease, no. 

‘ge Ova Scotia: gravel in veces. Big Intervale, 
Cape Breton Island, Macoun, no. 16,699. MaIne: argillaceous 
ledges by Meduxnakeag River Monticello, Fernald & Long, no. 
14,923; abundant in crevices of calcareous ledges and on cliffs by 
Mattawamkeag River, at foot of the Island, Sclaoal Falls, Sep- 
tember 8, 1897, Fernald; cliffs prec slate) along Piscataquis 
River, Sangerville, June 2 26, 1895, Fernald, no. 236; July 
29, 1900, W. Ne Park; rocky soil about ledges, Carrabassett 
River, North Anson, July 4, 1885, J. F. Collins. New 
SHIRE: ledges, Gorge of Diamond River, Darancuth. College 
Grant, Coés County, Pease, no. 10,513. 

It may be serviceable to others to have my key-characters 
separating the 4 species of § Vulgata which I can recognize in 
eastern America: 


a. Stem truly scapiform or subscapiform, naked or with 1 or 2 
cauline leaves borne near base; lowest leaves with rounded 
or: cordate pases 6 ee ee urorum L. 
(locally joke bil from Europe). 
a. Stem with 2-12 roaght ok maa to near inflorescence; low 
leaves Pape tioles. ... 


peoleclnke eta with arched- or divergentascend 
ing branches and pedicels; etic few—30 or more. H. vulgatum Fries 
‘Ai inchades = of Saretactions: 
b. Mw es and pedicels glandless or only very m 
ndular, copiously long-pilose or villous with mend 
pts rs; inflorescence strict, with 1-10(-20) bigs on erect 


us. 
Larger (lower) leaves 1-5 cm. broad, membranaceous, 
villous-ciliate, their peti tioles and the a of sda 


Mee copiously villous; heads (1-)4-20; phyllarie 
nt or acuminate: 6. VG ss H. groenla nace Arv.-Touv. 
Larger (lower) leaves 0.7-2 cm. broad, ¢ coriaceous, rg nace 
or nearly so, ad petioles and the midribs of low 
surfaces only s rsely pilose or glabrescent; nédle 
1-5(-8); phy levee candate-atiens es kes H. Robinsonit. 


The stations known to me for H. Robinsonii are given above. 
Those of H. groenlandicum (outside Greenland) are as follows: 
apor: Rama, A. Stecker, no. 347; open hillside, September 


LAB 
pene lat. 56° 31’, ‘long. 61° 10’ , Bishop, no. 608. _NEWFOUND- 
LAND: thickets on slaty “hills back of Little Quirpon, Fernald & 


320 Rhodora [AUGUST 


Gilbert, no. 29,268; thicket by Big Brook, Straits of Belle Isle, 
Fernald & Long, no. 29,267. ANTICOSTI ISLAND, QUEBEC: ae 
ealeaire et barachois, Rividre au Saumon, Victorin et al., 
21,303; sur les platidres élevées et séches, R. au Saumon, Victorin 
olla nd, no. 27,583; R. de Brig [ec ?], ‘Macoun, no. 18, in part; 
R. la Loutre, Victorin & Rolland, nos. 25,356—25, 358; a plusieurs 
milles en amont, R. Jupiter, Victorin & "Rolland, no. 25,360; sur 
ial platiéres sdches, R. du Brick,  Sasonles & Rolland, no. 27, 57 9; 
sur la rocaille calcaire prés de l’embouchure, R. 4 la Patate, 
Victorin et al., no. 21,310; talus ee. R. Galiote, Victorin & 


Ro land, no. 27,587; éboulis argilo-calcaires, R. Ferrée, Vic- 
torin & Rolla nd, no. 25, 355; sur les talus d’alluvion, R. Belle, 
Victorin & Rolland, no. 27, 591. 

H. CANADENSE Michx., var. fasciculatum (Pursh), stat. nov. 
H. fasciculatum Pursh, Fl. Am. Sept. ii. 504 (1814). 

Var. fasciculatum is the common plant which passes in the 
Northern States and southern Canada as typical Hieracivwm 
canadense. It is, however, not the extreme of the species which 
Michaux had, from Lake Mistassini, and well described as H. 
canadense in his Fl. Bor.-Am. ii. 86 (1803). Michaux clearly 
described, and the beautiful photograph before me of his TYPE 
shows, the northern extreme, mostly low, with few large and 
remote thin leaves and open paniculate-corymbose inflorescence 
of few long-peduncled heads with blackish involucre of oblong- 
lanceolate phyllaries. Typical H. canadense occurs from south- 
ern Labador to northern British Columbia, south to Newfound- 
land (where nearly ubiquitous), Cape Breton, Prince Edward 
Island, northern Maine, northern New Hampshire, Ontario, 
northern Michigan, northeastern Iowa, Saskatchewan, Montana, 
Idaho and Oregon. It is the plant which in the West passes 48 
H. columbianum Rydb. or H. canadense, var. columbianum 
(Rydb.) St. John. It was also described as H. macrophyllum 
Pursh, Fl. Am. Sept. ii. 504 (1814), Pursh noting the distinctive 
characters, “panicula divaricato-corymbosa, pedunculis elon- 
gatis nudis’’, which is quite similar to Michaux’s ‘‘corym 
paucifloro: pedunculis longis’’. Pursh also added the significant 
comment: “leaves the largest of the genus’’. 

Var. fasciculatum is the common plant which occurs from 
southern Quebec to southern Ontario and Minnesota, south to 
Nova Scotia, New England, northern New Jersey, Pennsy lvania, 
Ohio, Indiana, Illinois and Iowa. It was described by Pursh 


1943] Fernald,—Notes on Hieracium 321 


“caule erecto folioso . . . ramis paniculae divaricatis brevi- 
bus, pedicellis subfasciculatis pubescentibus’”. It was also de- 
scribed by Pursh (p. 503) from western New York and Canada as 
H. virgatum. 

Torrey & Gray understood typical Hieracium canadense. and 
its var. fasciculatum but, without knowing Michaux’s type, they 
reversed the two, calling var. fasciculatum typical H. canadense 
and treating the northern typical H. canadense Si macrophyllum 
Pursh) as H. canadense, var. latifoliwm Torr. & . Am. ii 
476 (1843). As I understand the species it falls ‘siti three well 
defined varieties as follows: 


on arched-ascending peduncles mostly 2-10 cm. long 
n or divaricately corymbiform panicle... H. canaense (typical). 


— 
5 
- 
S 
i] 
fe) 
s 
@ 
jag 
@® 
2. 
. 
5 
@ 
‘pe 
2 
5 
s- 
a 
ES 
) 
a 
oO 
D 
ge 
3° 
oF 


a. Involucres 5-10 mm. high; panicle more. or less 
fastigiate to umbelliform, with slender eee erect Motu 
peduncles; stem slender, 0.5-9 dm. high, its middle and 
upper internodes and the _Peduncles copiously villous- 
hirsute with hairs 1.5-3 m oe, eaves as in typical 
var. or the lower more tate CUMNG. 350s Si es Var. hirtirameum. 


H. cANADENSE Michx. Fl. Bor.-Am. ii. 86 (1803). H. macro- 
phyllum Pursh, Fl. Am. au ii. 504 (1814). H. canadense, var. 
latifolium Torr. & Gray, Fl. N. Am. ii. 476 (1843). H. auratum 
sensu Fries, Epic. Gen. Hierac. 124 (1862) not sig Symb. 
Hist. Hierac. isl (1848). H. columbianum Rydb. i Bull. Torr. 
Bo ‘|. xxviii. 513 (1901). H. canadense nrereet potty as 
subsp. (Rydb. ) Piper in Piper & Beattie, Fl. Se. Wash. 351 (1914). 
H. canadense, var. macrophyllum (Pursh) Farwell in Rep. Mich. 
Aead. Sci. xx. 195 (1918). H. vege Maes phe dita 
i Les Zahn in Engler, et ereng : cana- 
: nse, ne gested va (Rydb .) St. John, Fl. Se. Wash. and Adj. 


date, 459 (1937).—Range given ice 


322 Rhodora [AucusT 


Var. FascicuLaTtum (Pursh) Fernald, supra. H. fasciculatum 
Pursh, Fl. Am. Sept. ii. 504 (1814). H. virgatum Pursh, 1. ¢. 503 
(1814). H. Kalmii sensu Bigel. Fl. Bost. ed. 2: 288 (1824) ; 
sensu Torr. Compend. 280 (1826); oie Spreng. Syst. iil. 646 
feel not L. Sp. Pl. ii. 804 (1753).—Range ates vee ve. 

ar. — AMEUM Fernald in Ruopora, xvii. 19 (1915).— 
Dace to dry ledges, shores, cliffs and clearings, Nowininaianl 
to Wiseorain: south t o New Brunswick, New England and 
northeastern Vecia weeny 

As originally pointed out Var. hirtirameum is more inclined 
than other variations of the species to distortions due to aphids 
or fungi. Some altered individuals show secondary involucres 
only 2 mm. high. 

As to Hieracium Kalmii, it was said by Linnaeus to have been 
collected in Pennsylvania by Kalm. The type, of which two 
different photographs are before me, looks like a meagre speci- 
men of H. canadense, var. fasciculatum, for which Bigelow, 
Torrey and Sprengl mistook it. Its involucre, however, is less 
imbricated, with the pe obtuse; and ara studying the 
specimen Gray wrote:, . . Scales of involucre narrowly 
Enea, . °°. ee sagie geriee ss OVERS. 
glabrous, not striate . . . Pappus a single series of foarte 
strongly denticulate-scabrous bristles . . . ; we are confi- 
dent that his plant is not of North American origin”.—Gray in 
Torr. & Gray, 1. c. 479 (1843). 

In describing Hieracium canadense Michaux compared it with 
the European H. sabaudum L., a species which superficially sug- 
gests typical H. canadense, but from which our native plant 
differs consistently in many characters. Since the European 
species is appearing locally as a weed with us it may here be 
abe” 


American material. MASSACHUSETTS: vacant gras 
morial Drive near Cottage Farm Bridge, Cabcaes "October 6, 


1943] Fernald,—Notes on Hieracium 323 


1928, S. F. Blake, no. 10,789, the plant now abundant and found 
elsewhere in Cambri idge: abundant in vacant lot off Pleasant 
Street, Brookline, September 29, 1913, F. F. Forbes 

H. venosum L., var. plete (Michx.) acd in Mich. 
Acad. Sci. Rep. xx. 194 (1918). H. Gronovii, var. «. nudicaule 
Michx. Fl. Bor.-Am. ii. 87 (1803). 

In making this transfer Dr. Farwell did not clearly differen- 
tiate between true Hieracium venosum and the variety. From 
the start the two Linnean species, H. venosum L. Sp. PI. ii. 800 
and H. Gronovii L. 1. e. 802 (1753), were hopelessly mixed. As 
noted by Gray in Torrey & Gray, Fl. N. Am. 478 (1843) and as is 
apparent from the Linnean diagnoses his description of H. 
venosum, drawn directly from Gronovius, calls for a thick- 
stemmed plant: “HIERACIUM foliis cuneiformibus _hirtis, 
scapo nudo crassissimo erecto’’; while the very detailed descrip- 
tion of H. Gronovii, ‘Folia radicalia obovata, obtusa, integerrima, 
tenuitate membranacea, supra pilis raris adspersa, subtus subvi- 
olacea. Caulis pedalis, filiformis, uno alterove folio lanceolato, 
paniculatus, panicula inaequali”, vividly describes the plant 
which Willdenow, Sp. PI. iii. 1570 (1804), Torrey & Gray and all 
later authors have called H. venosum. There is no type of H. 
venosum in the Linnean Herbarium but the citations by Linnaeus 
of the two Virginian plants, Hieractum . . . lapathi venis 
sanguineis inscriptis foliis of Plukenet and H. . . foliis 
punctis & venis sanguineis notatis of Banister, and the descriptive 
name H. venosum can mean only the plant we know under that 
name. Willdenow and after him Torrey & Gray so used the 
name and, in view of the sad mixup at the start, this seems the 
right interpretation. As to H. Gronovii, as badly mixed, Gray, 
who studied the specimens involved, selected the plant of 
Gronovius cited by Linnaeus to typify that confused name. 

Hieracium venosum may have strictly rosulate leaves and a 
naked scape or 1-6 cauline leaves, the latter series of selected 
specimens being H. venosum, var. @. subcaulescens Torr. & Gray. 
Experience in the field shows that it has no definite range and 
that individuals with and without cauline leaves are frequently 
associated. Taxonomically it seems wholly unimportant. So, it 
seems to me, is var. Blombergit Zahn in Engler, Pflanzenr. iv™°. 
1117 (1922). Its only character, the presence of abundant 
glands on the involucre, is altogether too evasive. Just where one 


324 Rhodora [AuGcusT 


stops in estimating whether the involucre may be ‘‘tenuiter sub- 
glandulosa” (Zahn’s characterization of typical H. venoswm) and 
“‘densissime glandulosa” I do not know. Var. Blombergii was 
based on a single collection from Massachusetts. Of the 105 
specimens from that state before me 6 show no appreciable glands 
on the involucre, 89 have them obvious but possible to count 
(from 1 to 6 or 8 on a phyllary) and could be described by 
Zahn’s “tenuiter subglandulosa”’, and 10 specimens have enough 
glands to pass as “‘densissime glandulosa’’. I give up. 

There seems to be, however, a very real geographic variation 
within Hieracium venosum. All material before me from New 
England (with the exception of a few sheets from Cape Cod, 
Martha’s Vineyard and Nantucket) to southern Ontario, south 
into Pennsylvania and Kentucky, has the upper surfaces of the 
later and abundant basal leaves quite glabrous, although the 
smaller basal ones may have setose upper surfaces. Much of the 
material from Georgia to New Jersey and a few specimens from 
southeastern Massachusetts have the upper surfaces remotely 
setose with thick-based hairs. Thus, of the relatively small but 
significant representation before me 14 sheets (including all from 
south of lat. 38°) from Virginia have setose upper surfaces, only 2 
(both from north of lat. 38°30’) have them glabrous; 8 (all but 2 
from the outer Piedmont or the Coastal Plain) from North 
Carolina have setose upper surfaces, 7 (all from the mountains) 
not; 3 from South Carolina setose, 0 glabrous; 5 from Georgia 
setose, 1 (from Pine Mt.) glabrous; and all from Missouri setose. 
The setose upper surface, as opposed to the esetose one, seems to 
mark a real geographic variation. In view of the fact that all 
our collections from the regions best known to Banister (and 
through him Plukenet) and to Clayton (and through him Grono- 
vius) have the setose upper surfaces (“foliis . . .  hirtis— 
Gronovius; “Folia tenuitate membranacea, supra pilis raris ad- 
spersa”’—Linnaeus), I am treating the common plant of south- 
eastern Virginia as typical of the species as recognized by Will- 
denow and by Torrey & Gray and later authors. 

Michaux did not even recognize Hieracium venosum. He 
lumped the two very different plants, one with elongate sub- 
cylindric inflorescences, thick green foliage and stoutish stems, 
the other with open corymbiform panicle, often purple-marked 


1943] Fernald,—Notes on Hieracium 325 


membranous foliage and slender stems as H. Gronovii. Under 
that inclusive name he had two varieties: 
Var. «. icici aes caule sri aes panicula <porrraala 
— 6. fols : caule parce folioso: panicula oblo 
HAB. 8. in vieina et Caro lin 
— a. in Canada et Pensy a ania, 

The photograph of Michaux’s material which I took in 1903 
shows the two plants: his H. Gronovit, «. nudicaule, the common 
northern extreme of H. venosum with leaves glabrous above. 
The label bears the annotation “ Etat de N. York et Pensylvanie”’ 
and the significant note: “calyce pedunculisq. hispidis’’, which 
places it perilously near “var. Blombergii’’!, so that the latter 
name must lapse with those who think the more glandular speci- 
mens worth sorting out. AH. Gronovii, var. foliosum Michx. is 
very characteristic H. Gronovii as now generally interpreted, the 
plant with 2-10 well developed and scattered cauline leaves. 
The occasional more leafy individuals, with 20-30 cauline leaves 
seem not worth separating, at least, they are not var. foliosum. 

The highly plastic series of plants known as Hieracium mari- 
anum Willd. seems to be a group of more or less perpetuating 
hybrids with H. venosum as one parent, H. Gronovii often, or 
northward H. scabrum Michx., as the other. The resultant 
maze is very complex. It is our nearest approach to the baffling 
series of apomicts and mixed progeny with which the European 
students of Hieracium are familiar. 


LQ 


i 


Plate 


of AGROSTIS 


LO. 


Pa. 
> of 
£ ~ 
i, n_ 
Oo 
L v 
&, Te 
2° 
pc 


INDOSA M. 


spikelets 


VI UHLENBERGIA FR‘ 


! 


OLIOSA, >» 


Rhodorz Plate 750 


i SOUTHERN NEW HAMPSHIRE 
. 


Photo. B. G. Schubert. 


JHLENBERGIA FROND¢ 1, characteristic decumbent branch, te 
characteristic gle abrous iiteniedes open — rep get sssed sheath and ps Rats ine Tu 


/ 


panicle, X 5; FIG. 3, portion of rhizome, 


l 


iad 


Plate 


Rhodora 


», B. G. Schubert. 


Phot« 


a] 
— 
a 
-, 
< 
*% 
= 
RQ. 
™ =, 
nm 
2X 
an 
a) 
= 2 
ay wid 
>, T 
~ . 
laxaal 
c? 
rm. 
CS & 


MUHLENBERGIA MEXICANA: FI 


portion of panicle of type, 


A OU; 


Rhodora Plate 752 


s 


Photo. B. G. Schubert. 


MUHLENBERGIA MEXICANA: FIG. 1, Jacquin’s plate of AGRosTIS MEXIC 

FIG. * summit of ai eigat: = Sa as internode and node and base of 8 hes 
5, from oem vchu setts; , floret, X 10, from Quebec; FIG. 

from New Jerse 


Mees 
< 10, 


Rhodora Plate 753 


oy ra OF ace necSa 


. Mapes! 


ae. SP aS 
MERA A fy oP 
/ CAT pect? 82 Saw 


Photo. B. G. Schubert. 


‘ 


[UHLENBERGIA RACEMOSA: FIG. 1 (2 branches), type of Agrostis egestas x 4; 
enh x 1, from Illinois; Fra. 3, panicle, X 1, from Illinois; Fra. spikelet, x 40. “aes 
Minnesota; Fria. 5, floret, X 10, from Minnesota; Fie, 6, grain, < my ‘from Illinois 


Rhodora Plate 754 


Photo. B. G. Schubert. 


aie Prey IA ARSON A: FIG. 1, characteristic summit of plant, X 1, ter 
Minnesot: a; FIG. 2, summit of glabrous ea lustrous internode and me and baseé a 
sheath, X 5, i m Wekecuceiis . 3, portion of panicle to show characteristic smi 
anthers, x 4, from Illinois. 


Rhodora Plate 755 


WF, 
Sue 


ES 
me Cat 
aul we <= 
ad 


oo 
— 


oo 
— “ 
— 


Photo. B. G. Schube 


MUHLENBERGIA SETOSA: FIG. 1, type of PoLyP0G ON SETOS , FIG. coat 


E 
from Massachusetts; FIG. 3, pars age < 1, from Poa ania: FIG. 4, Por 


of rhizome © 4. from Nova Scotia; FIG , portion of panicle to show charac wets ic 
long anthers, * 4, from Massachusetts; FIG. 6, spikele ts, X 10, from M: assac hus a 
a, eaphGhci basal villi, * 10, from Pacis ylvania; FIG. 8, grain, < 10, from Pane. 


sylv: 


Rhodore Plate 756 


¢ 


eee i 
» he tlenberyss ylemutate Pv 


155K 


Photo. B. G. Schubert. 


ae HLE ie RGIA SETOSA (M. GLOMERATA): FIG. 1, type of PoLyPOGON GLOMERATUS , 
1G. 2, characteristic mee and pube rulent internode and node and ba se oO! 
inch, x 5, from Pennsylvani 


Rhodora Plate 757 


Photo. B. G. Schubert. 


Moa SETOS ir. Sa (OID ype of DAcrYLOGRAMMA C 
NOIDES, X 2, sande 1 x 1; to Newfoundi me FIG. 3, portion of aol ie . 
show ogee anthers, x 1, fr rom Mai uine; FIG. 4, floret to show villi running high on 


, grain, X 10, from Quebec. 


lemma, X 10, 


Rhodora Plate 758 


Photo. B. G. Schubert. 


ERIANTHUS COARCTATUS: FIG 2<: rig. 2, panicle, X 1; ria. 3, summit 


. 1, type, > 
of rachis of raceme, X 3; Fic. 4, spikelet, X 3; Fic. 5, spikelet, x 6. 


~] 
o 


Plate 


Rhodora 


~ ¢ 


’ we Sf 4 re] 

Qiao Va.ba S/o 
detec t GF? a 

eudual Oe iZe ee 


Cs 


B. G. Schubert. 


Photo. 
ERIANTHUS BREVIBA sa FIG. 1, type, X 14; F1G. 2, spikelets of type, < 114; F1G. 3 


spikelet, 6, from Arkans: 


Rhodora 


Plate 760 


Photo. B. G. Schubert. 


‘THUS BREVIBARBIS: FIG. 1, plant, X 14, from Arkansas; FIG. 2, 


Ex 
ré anh of raceme, X 3; FIG. 3, spike Ag x 3. 


summit of 


Rhodora Plate 761 


Photo, B. G. Schubert. 


. so* ETA Tne Ff T : y ECU ES ike li ; FIG ?. 
ERIANTHUS ALOPECUROIDES: FIG. 1, type of ANDROPOGON ALOPECUROIDES, X 7/2; FIG. 4, 
tip of same, X 3. i — . . ea tae ~ 

ANDROPOGON DIVARICATUM: FIG. 3, type, X 36; FIG. 4, portion of same, X 2; FIG. 9, 
paratype, Clayton, no. 600, X /. 


Rhodora 


Plate 7624 


Photo. B. G. Schubert. 


US PUMILUS: FIG. 1, habit, X 1; Frias. 2-4, spikelet, < 4; FIG. 5, disin age 


SCIRE 
acikelet : a ‘ital scarious margin oe seale, < 10; ric. . birt and its 
section, afte Schroet r; Fias. 7 and 8, achenes, < 10: Fic. 9, looking down “ 
0, 


summit of cekens. x 1 


its cross- 


yon 


Rhodora Plate 76: 


Photo. B. G. Schubert. 


RPUS EME INS: FIG. 1, topotype, 1; FIG. 2, spikelet, « 10, Fria. 3, disinte 
re spikelet, gee et ar scales, X 10; FIG. 4, ache ‘ne, X 10; Fig. 5, looking down 
"pon summit of achene, X 10. 


~~ 


RUFUS: Fia. 6, : cena: K 10; 
S. RU pie var. NEOGAEUS: achenes, X 


— 


Rhodora Plate 764 


Photo. B. G. Schubert 

Scirpus RoLuanpi: Fic. 1, type, X 1; Frias. 2 and 3, epee a type, x 10; 0; 
é t, disintegr: ated spike Jet ras type, showin ng anthers, * 10; . 6-8, achenes ’ 
FIGs. 8-10, looking down upon summits of achenes, owing ens :no- cones outline, 
x 10. 


Rhodora Plate 765 


Photo. B. G. Schubert 


Scirpus VALIDUs: FIG. 1, inflorescence, X 1; FiG. 2, spikelet, X 5; Fic. 3, achene 
and elongate bristles, X 10. 

Var. CREBER: FIG. 4, inflorescence of type, X 1; Fic. 5, mature spikelets, showing 
protruding achenes, 5; FIG. 6, flowering spikelets, X 5; F1c. 7, achene and sub- 


equal bristles, < 10. 
Var. CREBER, FORMA MEGASTACHYUS: FIG. 8, inflorescence of type, X 1. 


Rhodora Plate 766 


Scirpus STEINMETZII, all “2 from Sd : F1G. 1, inflorescences, X 1; FIG. 2, axis of 
inflorescence, X 3; FIGS. id 4, kelet < 5: Fias. 5 er 6, achenes, each with 
single bristle, * 10; FI«G. 7 achene fe we d from above, x 

s. 


HETEROCHAETUS: FIG. 8. spikelets FIG. 9. is oat é 10: ric. 10, achenes 
x 10. 


vie wed Irom above, 


Rhodora Plate 767 


Photo. B. G. Schubert 


2, portion 


CIRPUS EXPANSUS, all figs. from re FIG. 1, inflorescence, xX %: FIG. 
é ) 


of infloresee ence, X 3: FIG. 3, spikelets, with ha x 10. § 
S. SYLVATICUS: FIG. 4, small portion of “eh st nce, < 3; FIG. 5, spikelets, with 
1 


anthers, X 10. 


Rhodora Plate 768 


CQugetior 
Treg Ja 
—~ 
ce in 


en. 
* : i 


Photo. M. L. Fernald. 


Type of ANGELICA TRIQUINATA Michx., X 4. 


cian “Aaa eli a NN 


Rhodora Plate 769 


Photo. B. G. Schubert. 


ANGELICA TRIQUINATA Michx: portion of plant, X 14, of A. Curtisii Buckley. 


INDEX 


New scientific names are printed in full-face type 


Agrostis hepa 228, 230, 236; 

iformis a., 230, B., 230; foliosa, 

rian 22, as Sate 238, ~ 749, 

30; ndosa, 

298-230, "235, "238, pls. 749, 750; 
lateriflora, 226, 227, var. filiformis 


240, “ 244 ; spicata ears m, 
243, var ng ype 240, , 243, 
var , 242, var. villosa, 240, 


243; p desc 243 


6; mexicana, 223-226, 228, 236, Eleocharis pauciflora, var. Fernaldii, 


238, pls. 751, "052; racemosa, 


235, 237, 238, pl. 753; setosa, 337 Erianthus, 249, 250, 255; alopecuroi- 
es, 1 


Alopecurus peepee us, 23 
American Representative of Scirpus 


pl. 7 
pl. 761; furcatus, 255, 256, 258; 
Gerardi, 256, 258, Why’ not, ?, 255, 
var. chryso ocomus, 258, var. 
p ipilis, 258; nutans, 282-255; 
re Abeatigge md Bobet 255- 
mus, 258, subvar. 


uinus 
57, var. paucipilis, 258; spica Ferula villosa, 298-301 
digitatis, etc., 255; virginicum, 25 Five common Rhizomatous Species 


of Muhlenbergia, 221-239, pls. 
749-757 


a triquinata, 298; Gramen dactylon, villosum, ete., 255 
rece 298-301, What Hod i; onary 2 
sa 


298-301; venenosa, 301; 


— .’ Memorandum regarding 


29 Jam 
Anthoxanthum, 250; giganteum, 249- + nem 304 


Archangelica hirsuta, 3 ere 325, Notes on, 317-325 
t 


01 
Avena glumosa, 242; spicata, 242 
Blysmus rufus, 287 


Carex, 241 


’ Cassia Chamaecrista, 302 


Cicuta venenata, 300; venenosa, 300— 


Cinna, 227, 234; filiformis, 236; 
glomerata, 237; lateriflora. 226, 
os 


Chloris, 2 “ 
brvtiogsa Elliottii, 255 


Dactylogramma, 234; cinnoides, 282, 
234, 239, pl. 757 

Danthonia, 239, 254, Notes on, 239- 
= Alleni, 240-242, 244, 245; 


mpressa, , 245; Faxoni, 
24, co soil glumosa. 242: inter- 
, 240; rum, 243; 


ar "340, 345. po ters 339-243, 
(327) 


3 
subsp. canadense, 321; macrophyl- 
lum, 320, ; marianum, 325; 
molle, 318; murorum, 319; Robin- 
sonii, 317-319; sabaudum, 322; 
scabrum, 325; smolandicum, 317, 
subsp. Robinsonii, 317; venosum, 


var. nudicaule, , var. 8. subeau- 
lescens, 323; virgatum, 321, 322; 


vulgatum, 317-319 


328 INDEX 


Identity of Scleria setacea of Poiret, 
e 

Isolepis oligantha, 280 

Juncus canadensis, 295 


Marsh Muhly, -_ 223, 231 
Memora — egarding James 


301. 
Merathrepta compressa, 245; 
um, 243; route 242; showed 
243, ¥ var. pinetor , 243 
Muhlenbergia, 221, 223, 226, rosa 


forma ambigua, 
glumis, 236; rds , 235, pls. 
749, 750, forma commu ta, 

glabrifolia, 227; glomerata, 231, 
ae a 237, 239, var. ramosa, 


pl. 757; 
sylvatica, 221, 222, 224, 230, 231, 
235, 237, var robusta, 236, 237, 
var. ber Sle 

Muhly, 223, Marsh, 222, 223, 231; 
oom = 

Muley 


Notes ee ae 239-246; on 
Hiss eica 317-3 


hes pti triquinata, 299, 30 
eris compressa, 245. neha, 
“243; thermalis, 243 
Podosaemum glomeratum, 237 
Polypogon glomeratus, 232, 234, 237, 
239, pil. 6; racemosus, fe 
setosus, 234, 235, 237, 238, pl. 758 


Rhizomatous Species of Muhlen- 
ergia, Five Common, 221-239, 
pls. 7 

Rhynchospora, 297; setacea, 297 


Saccharum Soll 
Schoenus seta , 297 
Scirpus § Bucothryon, 279; § Lacus- 
tres, 285 e North American 
Members of, "283, pls. 765, 766; 
acutus, 284, 285; alpinus, 279, _ 
. gt form 
, forma nh 
phalanthus, 296; beer 291, 
ar. Fernaldi, 290, va e 


reckegs globulos 
289, * et irs rte 286, 287, 
pl. +66; lacustris, var. condensatus, 
is condensatus, 285, 


2 
280; gone ita 279; polyphyllus, 
orma macro: s, 296, var. 


G Baeothyron) 
ae 764; Prod ineaes onD95, 
form Bag mci 95; rufus, , 
aaa’ 288, var. neogaeus, 283, 287, 


inm i 
pl. 766; ‘Studies in North Ameri- 
of, oe an 


1 
aI 
or) 
a 
a 
° 
“3 
B 
=~ 
me 
be 
A 
< 
~ 
ws 


 laxa ; Mu 
bergii, 296, 297; a, 2005 Mi 296; 
Poiret, The Iden- 


st 
co 
tea 
bo 
-© 
= 
2, 
oO 
bs) 
=] 
Ae 


Some American Species and Varie- 
ae of perpas, 287, ge 763, 767; 
rth America 


INDEX 329 


gp es es 253, subsp. Linnae- 
253 


Studies in North American Species of 
Scirpus, 279-296 


Trichochloa calycina, 237; glomerata, 


Trichophorum emergens, 281 


Umbelliferae, 300 
Vilfa, 2 
What is Angelica triquinata?, 298— 


30 

Why not Andropogon Gerardi?, 255- 
258 

Wire-stem Muhly, 223 


Reprinted from Ruovora, Vol. 45, October-December, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM : 
OF HARVARD UNIVERSITY 


VIRGINIAN BOTANIZING UNDER RESTRICTIONS 


ie Saad 


Reprinted from Ruopora, Vol. 45, October-December, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


CXLIX 


VIRGINIAN BOTANIZING UNDER RESTRICTIONS 


M. L. FERNALD 


DaTEs or IssuE 


eye ae = UU re oo ee sk ek 9 October, 1943 
Pag tes 753-708 TET ee cme eer a 13 November, 1943 
Paces 485-511 aa Put TOO ii ce ik 23 December, 1943 


CONTRIBUTIONS FROM THE GRAY HERBARIUM OF 
HARVARD UNIVERSITY—NO. CXLIX 


VIRGINIAN BOTANIZING UNDER RESTRICTIONS 
M. L. Fernaup 
(Plates 770-806) 
Part I. Frev.p-Stupies or 1942 anp 1943 


At the approach of spring in New England in 1942 my former 
student, Professor Ernst C. Abbe of the University of Minnesota, 
then spending a year in research at the Gray Herbarium, was 
persuaded without too much pressure to contribute the use of 
his new Ford and to join Mr. Bayard Long and me for a short 
period of botanizing in southeastern Virginia. The need to go 
at once to the region arose through the pressure for housing near 
Camp Lee, with the result that our former headquarters at 
Century House, south of Petersburg, had been wholly taken over 
by Army officers and our great stock of driers, ventilators, press- 
frames, tramping clothes, etc. had been temporarily stored in a 
shed. There was no place left near Petersburg for such unes- 
sential people as mere botanists and we would have to move. 

Abbe and I drove from Cambridge, with spring still far in the 
offing, and on the afternoon of April 15th called in at Elkins 
Park for Long and, taking a thinly settled route (via Glen 
Burnie, LaPlata and Port Conway) out of Baltimore, reached 
Fredericksburg for the night. Starting from Fredericksburg on 
the morning of the 16th we decided to drive down the north side 
of the Rappahannock to the Port Conway-Port Royal bridge, 


358 Rhodora [OcTroBER 


thence to Petersburg. We might thus add something to the 
Checklist of Plants of the Washington-Baltimore Area, which 
covers the region south to the Rappahannock. About three 
miles southeast of Falmouth we were attracted by a brook 
cutting a ravine through the woods and by the conspicuous dis- 
play of such plants now in flower or fruit as Arabis laevigata, 
Stellaria pubera and Corydalis flavula. Parking just off the road 
at the eastern end of the bridge we stepped into a colony of 
Ranunculus abortivus var. indivisus Fernald in Ruopora, xl. 418, 
pl. 518 (1938), the extreme of the species characteristic of bottom- 
land woods of the Nottoway, ninety miles to the south; and when, 
a little farther on, we were puzzled by the Cerastiwm of the road- 
side-fill and found it to be the European C. brachypetalum Des- 
portes, which has been known in America only from roadsides of 
Southampton County, Virginia, we felt that we were promptly 
getting into stride. But we were obligated to reach Petersburg 
and to move our collecting equipment to new quarters. Our 
chief concern was not the Washington-Baltimore Flora. 

In the autumn of 1941 we had found not very far down river 
from Port Royal a fine bank of naturalized trees and shrubs and 
we wanted to secure flowering material of Vinca major L., which 
abounds there. So we took a little time to drive out to the home 
of Mr. and Mrs. Snowden on the river-bluff northwest of Return. 
After a visit with our hosts we reéxamined the colony of natu- 
ralized species along the river! and among them, unnoted the 
autumn before, there were Ulmus procera Salisb., the English 
Elm, and the American Philadelphus inodorus L. Then we 
went to the woods northeast of Loretto where, the preceding 
October, Poncirus trifoliata (L.) Raf. was fruiting?, hoping to get 
good flowering material. Luck was still with us, the shrubs 
being in the prime of flowering. Since the day was rapidly 
passing we then decided to drive from Tappahannock without 
stop to Petersburg. We did pretty well, stopping first for a few 
moments to look into a wet woodland glade where we found two 
species rare in southeastern Virginia, Viola cucullata and Sym- 
plocarpus foetidus; but when we came to the broad, green, wooded 
bottomland of the Pamunkey, slightly north of Old Church, we 
had to get out for just a peep. The usual plants of such habitats 

1See Ruopora, xliv. 372 (1942). 2 KHopora, l. c. 


1943] Fernald,—Virginian Botanizing Under Restrictions 359 


were there, Clematis crispa, Cardamine bulbosa, and a perplexing 
member of the Ranunculus hispidus-series, and with them Viola 
pensylvanica Michx.!, rare on the Coastal Plain. The great 
surprise, however, was acres and acres, as if native, of splendid 
flowering clumps? of the Old World Summer Snowflake, Leucojum 
aestivum L., as thoroughly at home as any squatter and making 
a display which we shall never forget. 

While we were busily exploring this bottomland we heard 
excited voices of Negroes and the hurrying of feet on the road 
near by; and when we came up from the woods, a man, sauntering 
quietly along the road, waited for us to greet him and to explain 
our strange doings. The colored women had rushed excitedly 
to his filling-station to report three terrible monsters down on 
the bottomland across the river, one of them with an axe (my 
little hand-pick), one with a can of dynamite (vasculum), the 
third with a big pack on his back (Long’s field-press). The 
women had “runned and runned and runned”’ and were com- 
pletely out of breath and greatly frightened. Thus the three 
devotees of what Linnaeus called “our guileless science” were 
unconsciously starting the reputation which it became difficult 
to live down in succeeding days in Virginia. The proprietor of 
the gasoline-station told us of the vast extent of the bottomland- 
woods, of the fresh tidal flats near by and of his readiness to 
guide us about the region, and we promptly arranged to employ 
him and his motor-boat in midsummer and autumn, then little 
realizing that midsummers and autumns would pass before we 
should again see the Pamunkey 

The roadside-fill (soft shoulder) near where we had parked the 
car was carpeted with two wanderers from Europe, Teesdalia 
nudicaulis (L.) R. Br. which we had only once found in the state 
(on a fill in Sussex County) and Holosteum umbellatum which we 
subsequently saw in other such habitats. The large number of 
Europeans suddenly appearing on these recently made soft 
shoulders (these, Cerastium brachypetalum, C. tetrandrum Curtis, 
and other European species) make one wonder about the source 
of seed sown on the soft shoulders of new roads in eastern Vir- 
ginia. 

‘See Ruopora, xiiii, 500 soe (1941). 

? See Ruovora, xliv. 390 


360 Rhodora [OcroBER 


Six or eight miles below Petersburg we found a place to sleep— 
in cabins where they were asking and getting $4.00 a night for 
each bed over the week-ends from families visiting loyal sons and 
brothers at Camp Lee, twelve miles away. There were no 
‘ceilings’? on cabins. Only as a special concession were we 
taken in on a Friday night, for the cabins were being rented 
only Saturday and Sunday nights during the activity at Camp 
Lee. Finding next morning that the Petersburg region was no 
longer available as a home, we decided to try Waverly, twenty- 
five miles to the southeast, a village in the center of good bota- 
nizing ground and at the junction of good roads. We had for- 
merly enjoyed the hospitality there of Mrs. Carter and her 
daughter, Mrs. Fleetwood, in their large old house, and we 
decided to try there; but Mrs. Fleetwood, we promptly learned, 
was at Fort Eustis as hostess and the old house was occupied by 
the family of an Army officer, who was forced daily to make the 
trip to and from Camp Lee. Southeastern Virginia was in the 
hands of the Army. Happily, our friend, Mrs. Calvin Horn, 
who, with her husband, runs an automobile- and gasoline-station 
at Waverly, was able to suggest a new home and in the evening 
we were established in the roomy and comfortable house of Mr. 
and Mrs. James Thompson and our equipment was stacked and 
ready for use. 

Promptly after breakfast on the 17th we started eastward to 
take a cross-road north to the James River. Only a few rods 
outside the village Abbe spied a fine colony of Iris verna in 
beautiful flower. It must be photographed. For three minutes 
he tested light and distance, with his camera pointing down to 
the innocent Iris, passing cars slowing down to watch the pro- 
ceedings. Then we went on to the James River, visiting then 
and two days later areas above Claremont, at Claremont Ferry 
and at Burwell’s Bay below Rushmere, and introducing Abbe to 
the complexities of the eastern Crabapples, now in bloom, and 
to such fine natives as Cardamine Douglassii and Sedum ternatum. 
On the weedy and turfy shore at Burwell’s Bay the little Stellaria 
media var. glaberrima G. Beck! made carpets with Ranunculus 
parviflorus and we here extended the range inland from the outer 
coast of the odorless, onion-like Nothoscordum bivalve. 

1 See Ruopora, xlii. 451 (1940). 


1943] Fernald,—Virginian Botanizing Under Restrictions 361 


We were specially anxious to show Abbe the big white-flowered 
Erigeron scaturicola Fernald in Ruopora, xliii. 524 and 654, pl. 
695 (1941), which grows in the deep and viscid, dripping wet 
lime-marl under the crest of the bluff west of Burwell’s Bay. 
We found typical FE. philadelphicus on the beach, then plants 
like small E. scaturicola but with pink, instead of white, rays be- 
gan to appear and finally, high up under the dipping crest, just 
where a magnificent mass of white-rayed E. scaturicola had 
formerly grown, there was an equally vigorous clump of plants 
with deep pink ligules. It was difficult to believe and I, natural- 
ly, wanted to get rooting specimens. These, after he had slipped 
back several times on the wet marl, Abbe dug and threw down. 
White-rayed FE. scaturicola was now pink! The dry material 
was stored in a pigeon-hole to await critical study and nine 
months later, when reéxamined, its rays were all white. The 
smaller and more typical E. philadelphicus has retained, as usual, 
the bright pink of its ligules. Again, ho-hum! £. scaturicola is, 
presumably, a physiological variant of EF. philadelphicus, induced 
by the concentrated calcium solution and cold water in which it 
grows. This is the view of Mr. Arthur Cronquist who is study- 
ing the genus. Whereas Aster is reputed to be hopelessly plastic 
and inconstant, Erigeron is usually well behaved. The “strong 
waters” along the bluffs of the James have there evidently 
“gone to the head’’s of one member of the relatively conserva- 
tive genus. It would be instructive to feed ordinary and typical 
E. philadelphicus for some years on refrigerated lime-water. 

On the way back to Waverly on the following Monday, as we 
crossed Cypress Swamp north of Dendron, we stopped to collect 
typical Senecio aureus in the bottomland-woods and as we came 
back to the car there, at the upper border of the bottomland, we 
saw a row of plants combining in different degrees the habit and 
traits of S. aureus with those of the very definite and wholly 
different S. tomentosus which was flowering in the drier open 
areas. It was clear that S. aureus and S. tomentosus sometimes 
cross, and Long and I immediately remembered a single colony 
on a bottomland west of Claremont which we had hopefully 
identified with S. Crawfordii!. Was it possible that that, too, 
was a hybrid? As I shall later show, it is. We thus returned 

See Ruopora, xliii, 521 and 657 (1941). 


362 Rhodora [OcroBER 


to Waverly at dark, after happy initial days but with some pretty 
baffling problems in mind and the first evening there we im- 
mediately met the greatest problem of all. 

When we parked the car outside the café, I promptly went in 
and sat at a table, but I was soon puzzled because Long and 
Abbe did not come in. Finally Long appeared at the door and 
mysteriously beckoned. When I went out, wondering at his 
action, he met me with the query, ‘‘Have you got your creden- 
tials? We're pinched.” ‘Why?’ I asked. ‘Because Abbe 
took a photograph.” My credentials (an identification-letter 
from the President of Harvard University; a letter from the 
Secretary of the National Academy of Sciences emphasizing that 
as a member I was a scientific advisor to the federal govern- 
ment; a letter addressed to me as President of the Botanical 
Society of America; a newspaper-clipping with a picture of me 
as recipient of a medal, in part for the botanical explorations in 
Virginia; and a passport with portrait) and my statements re- 
garding the distinguished American forebears of both Long and 
Abbe promptly convinced the police that we were not, as sup- 
posed, German spies who had crossed the Atlantic and had sought 
out the small village of Waverly for destruction. In the morning 
when little Iris verna was being photographed a zealous Waverly 
citizen, unacquainted with any such interest as that, had tele- 
phoned to State Police headquarters in Richmond of the “furrin ey 
car (from Minnesota) with three “‘spies’’ who were photographing 
the Norfolk and Western railroad-tracks, the Texaco oil-tank 
and other vital landmarks of the community, for it did not occur 
to him that our backs were toward the unsightly railroad, with 
the small oil-tank farther in the background, that the camera 
was pointed exactly away from those significant objects at some- 
thing on the ground and that our small botanizing pick (see 
PLATE 770) was not the implement to use in ripping up the 
heavy steel rails of the Norfolk and Western. Telephone con- 
versation with the motor-registry in Minneapolis revealed that 
the car belonged to Professor Abbe, with the incriminating given 
name Ernst. Radio calls had gone all over southeastern Vir- 
ginia for all officers to halt our car; and all day long we had 
innocently driven on the Suffolk turnpike (U. S. Route 460), on 
the road from near Wakefield to Surry Courthouse (Route 31), 


1943] Fernald,—Virginian Botanizing Under Restrictions 363 


on the Richmond-Smithfield road (Route 10) or on the road 
from Claremont to Waverly and far beyond, across the state 
(Route 40). We did not know that we were reputed German 
spies and that the State Police were watching for us. When we 
got back to our new home Mrs. Thompson was, naturally, upset. 
Her husband, the village barber, had come home to noon dinner 
with the tale, much amplified in the barber-shop, of the terrible 
mess they had got into, “harboring German spies’; but my 
documents and the realization that Long and I had for several 
years known Mrs. Horn, Mrs. Carter and Mrs. Fleetwood, and 
had more than once been guests at the latters’ home, and had 
regularly stopped in Waverly for meals during several years of 
botanizing, cleared the atmosphere. Nevertheless, next day 
some ladies from the other end of the town came to Mrs. Thomp- 
son to commiserate. They “hadn’t slept a wink, expecting to 
be murdered in their beds”’, etc., etc. Thus our début at Waver- 
ly was even more horrifying than our appearance on the bottom- 
land of the Pamunkey, and half-jesting, half-serious questions 
through this trip and on Long’s and my June-July trip indicated 
that the first impression of us ‘‘furriners” was still in people’s 
minds. We had become used to being “‘damn Yankees’’; Long 
and I had once been arrested in Richmond as ‘‘German spies”’ 
(see Ruopora, xliii. 516); but it was a novel experience to be 
called ‘“‘furriners’’. The people of Waverly are typical Virgin- 
ians and, consequently, strongly for the “New Deal’. They 
avoided addressing any such obvious heretics as we as “ Pro- 
fessor’. Baldness having advanced on all of us, we, perhaps, 
did not fulfill their conception of “long-haired professors”’. 
Since I know of no published photograph of Iris verna in its 
native habitat the photograph which so upset the equilibrium of 
southeastern Virginia and which somewhat ‘cramped our style” 
is here presented, as they say in all criminal cases, as exhibit A 
(PLATE 770). 

Desiring to secure in flower the remarkable variations of Acer 
floridanum (Chapm.) Pax near Grove Landing, below Williams- 
burg, which were discussed and illustrated in Ruopora, xliv. pp. 
359, 360, 426-428 and plates 725-727 (1942), and hoping to 
make the field-acquaintance of Monotropsis odorata, one of the 
Williamsburg specialties, we went on Sunday to William and 


364 Rhodora [OcroBER 


Mary as the guests of my former student and Abbe’s friend of 
student days, Professor Albert L. Delisle, and in the afternoon 
we made a pretty thorough canvas of the Acer problem. That 
has already been reported upon, but when we came to a tree 
with the trunk more than 4 feet in diameter and with its great 
sheets of exfoliating whitish bark flapping like those of shagbark- 
hickory we could not help mourning our inability to get a photo- 
graph of it. The herbaceous plants of this rich forest were 
noted a year ago. To that list should be added Nothoscordum 
bivalve which the day before we had got on the opposite side of 
the James. After supper Delisle’s student, a keen searcher for 
and discoverer of rare plants, Mr. Kenneth Winfield, took us to 
his station for Monotropsis. It was necessary for him to show 
us the first clump of slightly too mature plants; after that we 
found more and more. This aromatic saprophyte really abound- 
ed. Its favorite habitat was in the leaf-mold in the shelter of 
fallen and decaying logs. We thought we had learned how to 
find it but later searches on similar slopes elsewhere were fruitless. 

In April, 1926, Dr. Paul A. Warren collected in the Great 
Dismal Swamp a dwarf Trillium which I had misidentified as the 
extreme southern T. lanceolatum Boykin but which is not satis- 
factorily referable to that species. Its sessile flowers easily lead 
to T. lanceolatum but in other characters it is near T. pusillum 
Michx., a species with peduncled flowers. Dr. Warren and a 
party of students from William and Mary had walked in on the 
old path from Wallaceton to Lake Drummond. The path now 
being obliterated by dense growth and by tangles of briers 
following forest-fire, we went in search of the Trillium via the 
Feeder Ditch in Capt. Crockett’s motor-boat. The search was 
futile but the excursion delightful and when we came back from 
Lake Drummond to the Federal dam, the engineer, Mr. Cherry, 
asked what particular plant we were seeking. When I described 
it he promptly said: “I never saw it but once, on a hillside 
covered with beech-trees where it abounded when I was a boy”. 
His explicit directions took us back to the beech-slope described, 
near Deep Creek, now a picnic-ground, trampled and scraped and 
with no Trillium left. Most fortunately, however, Mrs. Laura 
H. Lippitt of Dinwiddie has a good station for the plant and the 
beautifully prepared specimens she has sent me and others 


1943] Fernald,—Virginian Botanizing Under Restrictions 365 


loaned me by Professor Smart of the University of Richmond 
enable me to describe and illustrate (PLATE 773, FIGs. 1 and 2) 
this endemic Virginian Trillium. 

The season was too early for many novelties, though on suc- 
ceeding days we extended some local ranges and renewed our 
acquaintance with numerous vernal species, but nothing signif- 
icant enough for special note was collected. 

Unfortunately, when we could next get away Long and I had 
to go without Abbe. We reached Waverly in the afternoon of 
June 28th and, beginning early on the 29th, we had all we could 
handle until obliged to leave on the morning of July 8th. Mr. 
Horn was able to supply a pick-up truck to transport our equip- 
ment and he found a very efficient driver and helper for us, 
Lennie Watts, son of a local police-officer, young, full of activity 
and interest and as effective a driver on back roads as our best 
drivers of other years. Only three of our old stations were 
specially revisited. We did not let ourselves be lured to the 
marl-blufis of the lower James or north of that river and we did 
~~ get down to Greensville County or the pine barrens of 

and soutl tern Nansemond. Gaso- 
line was becoming scarce and must be stretched to cover all 
mileage possible, but we had a careful driver and an efficient 
truck and Mr. Horn kept it up to full effectiveness. 

Nearly west of Waverly we went as far as McKenney in Din- 
widdie County. In October of 1941 we had found just east of 
McKenney an area of low woods and clearings where typical 
Coastal Plain plants (Helianthus angustzfolius, Cirsium virgini- 
anum, and Solidago perlonga Fern.) extended well back into the 
Piedmont and mingled with eastern colonies of upland types 
such as Viburnum Rafinesquianum Schultes. There, too, were 
the second station north of northwestern Georgia of Gentiana 
cherokeensis (W. P. Lemmon) Fern. and the third station in the 
state for the characteristic species of prairies of the Interior 
(Indiana to Texas), Muhlenbergia brachyphylla Bush. Such a 
complex area demands exploration through the summer and 
autumn and on the Ist of July it was up to its October standard. 
Viburnum Rafinesquianum was more abundant than we remem- 
bered; the local Phlox carolina L., var. triflora (Michx.) Wherry 
shared a low clearing with Vicia caroliniana, Thalictrum re- 


366 Rhodora [OcroBER 


volutum, Scleria oligantha and other rather local species not seen 
in October. The ditch bordering the woods was the home of 
Polygala sanguinea (rare in southeastern Virginia) and Rhexia 
ventricosa Fern. & Griscom in RuHopora, xxxvii. 192, pl. 346 
(1935), its range extended inland, and the Oenothera looked, as 
they so often do, a little strange. A couple of pieces were taken 
for checking. Next time we will get more, for it is Oe. tetragona 
Roth, var. riparia (Nutt.) Munz in Bull. Torr. Bot. Cl. Ixiv. 302 
(1937), known to Munz only from its original region, Cape Fear 
River at Wilmington, North Carolina, and the Santee River 
region of South Carolina. We were hot and hungry. So, 
seeking a shaded spot, we sat down in a colony of the inland, 
continental Psoralea psoralioides (Walt.) Cory, var. eglandulosa 
(Ell.) F. L. Freeman in Rwopora, xxxix. 426 (1937), Miss 
Freeman’s statement of range being: ‘‘ Higher altitudes of the 
Southeastern States, in South Carolina and Georgia, ranging 
westward as far as Texas, thence northward to Ohio, Indiana, 
Illinois, Missouri and Kansas”. This Psoralea and the Oeno- 
thera were as fine a pair as Muhlenbergia brachyphylla and 
Gentiana cherokeensis, found here in October. Solidago perlonga 
we had always considered an autumnal species and it was still in 
good condition here the preceding October, but already preco- 
cious individuals were beginning to flower—the species has a 
flowering period of four months! Clematis ochroleuca was here 
very large, some plants branching at every axil and bearing 8 
fruiting heads. Since the classification of the sub-§ Integrifoliae 
of Clematis, § Viorna has recently depended too much on incon- 
stant vegetative, rather than more stable fruiting characters, I 
have tried to find the latter in the eastern American species. 
My results, with pLates 776-782, will be found in Part II. 

On the way back to Stony Creek we stopped at several inviting 
spots. Only one may here be noted. The unique and to most 
botanists very rare Carex Collinsii Nutt. (unique in having the 
teeth of the perigynium abruptly recurved as hooks, thus sug- 
gesting the very primitive subantarctic and Asiatic Uncinia 
which has the exserted rachilla abruptly hooked) inhabits cool 
spring-fed sphagnous carpets in wooded swamps. Since we first 
detected it in Virginia we had learned to expect it in such habi- 
tats. Consequently, when we saw such a deep carpet of sphag- 


1943} Fernald,—Virginian Botanizing Under Restrictions 367 


num in the woods east of Cherry Hill, we investigated. There 
was C. Collinsii, already over-ripe, and with it was true southern 
C. venusta Dewey at a new northern limit. 

Three Creek, tributary to the Nottoway, has yielded us more 
localized specialties than any other small stream in southeastern 
Virginia. Wherever we have been on its bottomlands and 
wooded banks, from the fall-line north of Emporia nearly to its 
mouth, we have got something novel. The Arringdale Sheet of 
the Topographic Survey indicated good bottomlands at Arring- 
dale and, slightly farther up stream, near Adams Grove. So, 
having never tried them, we drove from Capron by the dirt 
road toward Arringdale. Closely watching the topographic 
Sheet, we sought the crossroad to the village which lends its 
name to that invaluable guide through the country. Nothing 
but dense thickets and woods could be found. Turning the car 
we drove back to definite landmarks. Finally, seeing a colored 
girl coming toward us we waited and, as she came up, asked: 
“Will you please tell us how to find Arringdale?” “Irene 
Dale?” she promptly replied. “I’ve never hearn of her. I 
know Irene Dunn up the road a ways but I never hearn of Irene 
Dale.” Arringdale has become extinct; its fame is perpetuated 
only on the contour-sheet! 

It was a sweltering day and the thought of plunging through 
dense thicket in search of sinuous Three Creek was not inviting; 
SO we went on to Adams Grove. As we crouched in the shade 
of the small bridge there, to keep out of the blazing heat while 
we ate our lunch, we were able to note around us several exten- 
sions of range to the west, pretty closely approaching the fall- 
line: Sagittaria Weatherbiana Fern. in RHopora, xxxvii. 387, 
plates 385 and 386, fig. 1 (1935), Carex Bayardi Fern. in 
Ruopora, xliv. 71 (1942) and Micranthemum umbrosum (Walt.) 
Blake, for example; and as we lunched we were impressed with 
the somewhat “fluffy”? aspect of the meagre umbels of the Sium 
at the margin of the Creek. Its foliage, too, was unlike that of 
the common northern plant, and its slender and flexuous, terete 
stems were not like the strongly corrugated and usually coarser 
stems of the plant commonly known as S. suave Walt. or S. 
cicutaefolium. This was young flowering material of the species 
which we had previously collected in fruit on the bottomlands 


368 Rhodora [OcroBER 


of the Blackwater and of Fontaine Creek. They all belong to 
S. floridanum Small, known to Small only from a single valley in 
Florida. It sometimes pays to stop and eat lunch! 

We saved up our allotted gasoline until we had enough to 
reach the region of Northwest River, flowing from the Great 
Dismal Swamp into Curratuck Sound. There is always some- 
thing different to be found on the marshes near Northwest or 
along adjacent North Landing River; it is important to visit 
them at all seasons. On the way, a little east of Magnolia, white 
flowers on the surface of one of the big ditches along the highway 
across the northern end of the Great Dismal Swamp attracted 
us, and we got out to collect our first material in the region of 
Cabomba caroliniana. As we approached Northwest we were 
surprised to see a striking and very “different”? Physostegia 
bordering Indian Creek. We were quite familiar, on the wooded 
bottomlands of the Blackwater and the Nottoway, farther in- 
land, with the delicate and thin-leaved P. denticulata, but this 
was a much coarser plant with larger corollas, heavy and sub- 
sessile (instead of membranous and long-petioled) leaves and 
other characters which quickly distinguish it. Unfortunately, 
the colony was densely enmeshed by Cuscuta but we cleaned off 
enough material to make a good type and in Part II I shall 
describe and illustrate it (PLATE 783) and attempt to clarify the 
genus in the ‘‘ Manual range”. Days at intervals of three weeks 
in a motor-boat along Indian Creek will sometime be very pro- 
ductive. So will days at similar intervals on little Blackwater 
River near by and, likewise, on Northwest River, both up stream 
and down to the Carolina line. Furthermore, North Landing 
River, with its broad savannah-marshes needs similar explora- 
tion. There is plenty to do; we have merely glimpsed the easily 
accessible margins of these areas. On the reed-marsh of North- 
west River the always superb orange- and red-flowered A sclepias 
lanceolata, var. paupercula (Michx.) Fern. was very handsome 
and one plant had bright, pale yellow corollas, a pretty color- 
form. I took the top, Long wistfully remarking, “We'll come 
again next year for mine”. When we are again able, after the 
“duration’’, to visit Northwest there may be more than a single 
yellow-flowered plant. Here’s hoping! And if we are there at 
the right season we may secure better material of the undescribed 


1943] Fernald,—Virginian Botanizing Under Restrictions 369 


Solidago which, when we found it in October of 1941, had been 
mowed off and was, therefore, not typical. The thicket border- 
ing the marsh is also the home of a new variety of the wide- 
ranging and variable Eupatorium rotundifolium. Whereas 
typical FE. rotundifolium has the leaves gently rounded to sub- 
truncate at base, this plant of Northwest River has the unusually 
large blades deeply cordate, with the bases clasping the densely 
white-tomentose stem. We afterward found it in a marshy 
thicket in Southampton County, and Godfrey got the same 
extreme variety only a few miles to the south, in Bertie County, 
North Carolina. It will be described and discussed in Part II. 
Another plant, a very bristly Stachys, on the reed-marsh of 
Northwest River, added to a large series of different members of 
the genus accumulated during several seasons in southeastern 
Virginia, has forced me finally to tackle the group. The results, 
with many illustrations (PLATES 787-794), will be given in 
Part IT. 

July 4th, even with restricted gasoline, may be an unsafe day 
for cautious and virtuous drivers on trunk-routes. So, when 
Lennie asked if we might keep off the main roads on that day, 
to avoid drunken drivers, we remembered that there had been 
extensive lumbering operations in the pine barrens south of Zuni, 
with the consequent partial clearing of new lumber-roads and 
the opening to sunshine of formerly shaded areas. That was 
the place to go. Clearing and disturbing of the soil, while taking 
the older trees of Long-leaf Pine and many other trees, had 
certainly stimulated many species and brought others into view. 
Near one damp, new woodroad Amianthium Muscaetoxicum, at 
a new northern limit in this section of the state, and Calopogon 
pallidus Chapm., its limit of range extended 1034 miles to the 
north, stood up bravely in the mossy thicket where they had 
formerly been hidden, and some clearing-phases of Panicum and 
Tephrosia puzzled us. When the habitat has become more 
stable the Panicum may look less unfamiliar. The Fephrosia is 
the glabrescent extreme of 7. virginiana, which we had found in 
other pine barrens. It is evidently T. virginiana, var. glabra 
Nuttall, described in 1838 from Georgia. It seems to be a well 
defined variety. In the drier white sand Bulbostylis ciliatifolius 
(Ell.) Fern. had its northeastern limit also slightly extended. 


370 Rhodora [OcTroBER 


But the greatest display in the recently disturbed sand was of 
Euphorbia Ipecacuanhae. Under normal conditions this hope- 
lessly variable species flowers in early spring (late March into 
April) but here in the loosened sand it was still in flower and 
young fruit in early July. Against the suggested explanation 
that the similar but more upright HZ. marilandica Greene might 
be a hybrid of tall and erect E. corollata, flowering in Virginia 
from early June to autumn, and depressed and matted £. 
Ipecacuanhae the reply has always been that they cannot hybrid- 
ize because their flowering periods are so different. The loosened 
sands south of Zuni dispose of that argument. 

At the Blackwater River, near the bridge west of Blackwater 
School, we established new northern limits for Asimina parviflora 
(Michx.) Dunal and for Physalis monticola Mohr, and, crossing 
the river into Southampton County, we promptly parked the 
car, for the sandy flat along the river southeast of Unity was very 
alluring. Most of the plants were of species now known to us 
but always nice to collect, Paronychia riparia Chapm. and 
Vaccinium Elliottii Chapm. at new northeastern limits and some 
others equally good. The prize here, however, was a large 
colony of hundreds of uniform plants of a particularly dainty 
Ruellia which, since June of 1941, had worried us. In low, 
sandy woods at Round Gut on the Nottoway we had then found 
a colony of Ruellia, the mature stems only 2 or 3 inches high, 
with thin and blunt membranous leaves only 1.5-3 cm. long. 
The colony was struggling under a pile of brush and had no real 
chance. So we carefully cleared away the worst obstructions 
and nursed the colony along, finally to secure a few flowers. 
Now we have a real station and the plants from it will be the 
type when a study of Ruellia which I now have in progress is 
completed. Whether it can be finished in time to include in 
Part Il is very doubtful. It is a complex problem, made worse 
by past inattention to types and original descriptions, and many 
contradictory interpretations must be reconciled. But the 
sandy flat southeast of Unity has one member of the genus 
which we know from Virginia only in Southampton County. 

Starting for home, we decided that we could save mileage and 
time by cutting over to Route 312, from Courtland to Smithfield, 


1943] Fernald,—Virginian Botanizing Under Restrictions 371 


passing through Berlin! thence turning off at Ivor for Waverly. 
On the way we passed roads leading to various ponds which we 
should have liked to investigate and when we got north of Sedley 
a little landlocked pond, slightly off our route was too tempting. 
It was really a small Cephalanthus-hole, but in the thicket we 
promptly detected the new Eupatorium of Northwest River. 
There, also, was Drosera rotundifolia, the first we have seen on 
the Coastal Plain west of the Blackwater, and on a saturated 
log the very local Utricularia virgatula made a close carpet. 
Driving on to find a good turning-place we came upon Johnson’s 
Millpond. While Lennie was backing and turning the truck we 
went to the sandy shore. There was Oldenlandia Boscii (DC.) 
Chapman, its range extended 12 miles northward, but in a couple 
of days we stretched its northern limit more than 12 miles more; 
and with it Digitaria serotina (Walt.) Michx. at its second known 
station north of the Carolinas. Here too, was Panicum Wrighti- 
anum Scribn., a species subsequently found on most sandy 
pond-shores. 

Lennie, noting that we had a weakness for pond-shores and 
marshes, asked next day if we had ever been to Harold’s Millpond. 
We had not, but since in the past most millponds had been 
fruitless, as artificially dammed branches, and several which we 
had sought had been abandoned and grown up to cypress- or 
gum-swamps, we had our doubts. However, we were willing to 
try. So on the 6th we started for Harold’s (on the contour- 
Sheet as Harris) Millpond, southeast of Waverly. It was a 
dammed-up cypress- and gum-swamp but it was better than most 
and will stand exploration by boat, for drifted ashore were 
Utricularia purpurea, which we had never met in our Virginia 
work, and U. inflata var. minor Chapm. which we had seen only 
twice. Thus encouraged we decided to try Brittle’s Millpond, 
west of Wakefield. We stuck to the old road, rather than return 
to the turnpike, and slightly before reaching Brittle’s Millpond 
we investigated a piece of spring-fed sphagnum-carpeted woods. 
Here the chief interest was centered on a Xyris, just coming into 
flower and not easily matched in the herbarium. It seems to 
have some distinctive characters but mature material is needed. 

1 Berlin, Baden and Hanover were re of them named in our honor. They were 

m Virginia maps long before we visited the state. 


372 Rhodora [OcroBER 


Brittle’s Millpond at once gave us Panicum Wrightianum and 
Oldenlandia Boscii, but the shore was not easy to follow and too 
much punched by the hoofs of grazing animals, so we proceeded 
southward to see what could be found. And the next pond was 
a true find. 

Suddenly coming into view, Airfield Millpond gave us a real 
thrill. It had a broad white-sand beach, perhaps 50 feet wide 
up to the bushes, such a pond as occurs in southeastern North 
Carolina but such as we had never seen in Virginia. Here we 
settled down for the rest of the day, until driven out by a thunder- 
storm and downpour in the afternoon. The rarer pond-shore 
species of Panicum, such as P. spretum, were there, the now 
unmentionable Oldenlandia of course, and carpets of all sizes of 
Eleocharis microcarpa, some stranded, some deeply drowned, 
and so variable in size and in thinness or inflation of culms that 
we were fooled into imagining it several things. Paspalum 
dissectum gave us a new station, and Rhynchospora perplexa 
Britton lived up to its name. We kept taking it, for on different 
belts along the beach it looked very different. Dr. Cross 
(Shirley Gale) says it is all R. perplera, but she admits that the 
quite different species which abounds on one stretch of beach, 
where Drosera capillaris Poir. is exceptionally stout and fine, is 
R. filifolia, the first known between southeastern North Carolina 
and Cape May, New Jersey. We at last had found a pond with 
real sand-beach. Its shoreline is more than 4 miles around and 
we followed only one tenth of it. Rhynchospora filifolia occurred 
in but one patch a few rods across, the Drosera was more restrict- 
ed, and we saw only one lonely individual of Styrax americana. 
The remaining shore and visits at intervals up to October should 
yield great returns. Many species unknown north of the 
Carolinas delight in such a shore. 

On our last day we, inevitably, sought more ponds. Pro- 
tracted drouth had lowered the water and now was the time to 
investigate. Driving to Windsor we took a road to Collosse, 
thence to Darden’s Pond. There is a swaley and muddy shore 
but we got nothing surprising. Next we tried Womble’s (on the 
contour-sheet Wade’s) Millpond, which is a drowned cypress- 
swamp. Sometime, with the aid of a boat, it may yield good 
things, for Utricularia inflata var. minor is there; so are great 


1943] Fernald,—Virginian Botanizing Under Restrictions 373 


carpets of U. biflora. Our greatest prize, however, was Carex 
decomposita, the rare species heretofore known in the state only 
in cypress-swamps of the lower James. We then moved on to 
Whitefield’s Millpond, southwest of Corinth, and we regretted 
that we had not found it in the early morning. This was our 
last day and twilight was approaching, and we had found another 
pond with a broad sandy beach. Most of the good things of the 
other ponds were there and we quickly found a colony of Hy- 
pericum denticulatum Walt., var. ovalifolium (Britt.) Blake, one 
of the most local plants of the state; then Viola lanceolata, var. 
vittata (Greene) Weath. & Grisc. and Proserpinaca intermedia 
Mackenzie, the latter the first from south of the James. At one 
point I shouted “Look, look!” and promptly got an echo. Long 
saw it too: a little subprostrate plant with filiform stem, lance- 
oblong leaves and deep blue glomerules, an Eryngium of the 
prostratum group, new to Virginia if not to science. Its immature 
fruits and its sepals do not well match those of the more southern 
and western FE. prostratum. This was the last plant collected 
and we promised ourselves that in three weeks we should get a 
real series in ripe fruit. Whitefield’s Pond is nearly four miles 
around. We saw one twentieth of the shore and each of the 
specialties collected was highly localized. Just as with Airfield 
and with Johnson’s the possibilities are great. 

Before leaving Waverly we called on the Chairman of the local 
Rationing Board and were assured that, when we should return 
in three weeks, we would be granted the necessary gasoline for 
continuation of our work, gasoline for trucks used in scientific 
exploration, as he showed us in the regulations, being unrestricted. 
It would be necessary, however, to write a letter to the Board 
who would have to vote on the matter. The letter was promptly 
sent, with return addressed envelope and an air-mail stamp and 
I proceeded to make a directory of the more than 100 ponds in 
Tidewater Virginia, most of which we had never seen. By 
Visiting them all we should find a few as good as Whitefield’s, 
Airfield and the Cat Ponds in northern Isle of Wight. A re- 
newed letter to the local Rationing Board, then one to the State 
Board, then one from an officer of the federal Experiment 
Station brought no replies and a considerable collection of 
uncanceled air-mail stamps was being assembled. Perhaps we 


374 Rhodora [OcroBER 


were not merely “damn Yankees’’. Could it be possible that 
we were still thought of as ‘‘German spies”? In Massachusetts 
I was assured that, if I would hire and take to Virginia a Massa- 
chusetts truck, I could have unlimited gasoline. It seemed, 
however, an unjustifiable waste of gasoline, as well as tires and 
time, to travel 1200 miles to and from Waverly in order there to 
purchase gasoline for 500 miles of local scientific research. We 
are called disloyal if we do not cheerfully accept all the local and 
contradictory rulings of OPA. Scientific research is what they 
choose to dictate and the investigators are those they select. 
If our work were in Florida or some other state we should not be 
thus penalized for being scientific investigators. 

Professor Massey asked us to guide him to definite stations in 
Tidewater Virginia for the different species of Vitis, Rubus and 
other plants of economic importance and all arrangements were 
made for our joining forces at Waverly and exploring with his 
federal truck for the practically useful wild species, including 
those with edible roots, shoots and seeds and those which might 
furnish fiber and alcohol. We could also get mature material of 
the little Eryngium and other technically interesting plants. 
Everything was set for productive field-research when Massey, 
most unhappily, got himself seriously poisoned and hospitalized 
by digging and cleaning a lot of bulbs of Fly-poison, Amianthium 
Muscaetoxicum, for experimental study. 

In the spring of 1942, Mr. John B. Lewis had sent me from 
the Seward Forest of the University of Virginia at Triplett in 
Brunswick County a most extraordinary sterile plant of an 
Asarum which seemed to combine traits of true Asarum, with 
superficial, elongate rhizomes, and § Heterotropa (the genus 
Hexastylis Raf.). This was accompanied by an invitation for 
Long and me to go to the Seward Forest to study it and other 
problems. Finally, in late August, he sent me from the same 
area Hypericum setosum L., one of the most local of Coastal 
Plain species in Virginia, and perfectly typical Cynoctonum ses- 
silifolium (Walt.) J. F. Gmel., the first evidence in the state of 
this distinctive species, to quote Small’s designation, of ‘*Pine- 
lands and wet places, Coastal Plain, Fla. to La.” We already 
knew that in the Seward Forest there is a small relict stand of 
Long-leaf Pine, 47 miles west of its concentrated area in south- 


1943] Fernald,—Virginian Botanizing Under Restrictions 375 


eastern Virginia. Hypericum setosum and Cynoctonum sessili- 
folium, added to Pinus palustris, indicated a probable intrusion 
of Coastal Plain conditions farther back into the Piedmont of 
southeastern Brunswick County than the one at McKenney in 
Dinwiddle County, recorded in Ruopora, xliv. 373 (1942) and 
on a preceding page in this journal. I promptly wrote Mr. 
Lewis that, as soon as I could arrange for it, I would go to the 
Seward Forest. I still hoped (in vain) for at least the courtesy 
of a reply, pro or con, from the Ration Boards I had repeatedly 
written, since I knew of so many colleagues with all gasoline 
needed in various southern states for their scientific explorations. 

Finally, on the morning of October 10th, I reached Emporia, 
to be met there by Mr. Lewis. I had written in advance, sug- 
gesting that, before proceeding to Triplett, we drive over to 
Sedley and have a look at Whitefield’s Millpond and collect 
mature material of the puzzling Eryngium and its various com- 
panions of October, some of which would certainly be novelties. 
It seemed, however, that Professor Alfred Akerman, the Director 
of Seward Forest, wished to see forest-conditions and experi- 
mental plantings in Greensville and southern Southampton and 
had most kindly proposed that we unite errands and spend a long 
day afield with the Forest truck on the 14th, when I could take 
a night train, at Emporia, north toward Boston. That plan 
was so superior to mine that there was no question about it; 
a few days later we could take the cream from the now abnormally 
broad sand-beaches of Whitefield’s, Airfield and Johnson’s Mill- 
ponds and Long would still more regret that he had been unable 
to join us. 

Seward Forest is a wonderful experimental area for scientific 
study of forestry and, with very restricted income, Dr. Akerman 
and his loyal and underpaid associates are projecting and carrying 
on invaluable researches. It was a comfort to me to find that 
this wise and very practical experimenter in forest problems did 
not have the love of forest fires, as necessary to the stimulation 
and growth of valuable timber, which is sometimes urged. In 
fact, the adult trees of Long-leaf Pine, a species which some stu- 
dents claim to need fire if the seedlings are to succeed, have, 
without that stimulus, successfully planted seeds, and a number 
of vigorous columns of young trees are there thriving. Most 


376 Rhodora [OcroBER 


happily Dr. Akerman did not have the prejudice against botanical 
research which so frequently blocks our work and, whenever 
possible, he joined in the local exploration and everywhere found 
special trees or local tree-species which became the subjects for 
stimulating discussion. The University of Virginia and the 
state have invaluable assets in the Seward Forest and in the 
great capacity and sound common sense of its Director. 

Even on the crest, where Long-leaf Pine is successfully renew- 
ing itself as a little isolated colony, the rock is mantled by pebbles 
with rounded sides; and at lower levels the woods and fields are 
strewn with smoothed and rounded stones and pebbles. This is 
not what one finds in the ordinary fracturing and frost-heaving 
of igneous rock. It is most difficult to escape the idea that in 
this area the old stones and pebbles have been rounded under or 
by water. The creeks and branches are small and at the lowest 
levels. In view of the several typical or elsewhere exclusively 
Coastal Plain species of plants growing there one, who is not a 
geologist, inclines to the view that an arm of the Miocene Sea 
must here have intruded back into the Piedmont and that its 
waters rounded the granitic pebbles and that typical plants, now 
chiefly of the inner Coastal Plain, subsequently found there the 
soil-conditions which suit them. At any rate, Cynoctonum ses- 
silifolium thrives on a mossy and partially wooded flat; Hy- 
pericum setosum has recently been found by Mr. Lewis at a 
second station; Panicum aciculare of “Sandy woods and pine- 
lands, Coastal Plain” (Small) and P. lucidum of ‘sphagnum bogs, 
Coastal Plain” (Small) abound. In fact, the mingling of Coastal 
Plain with ordinary Piedmont and even Appalachian Upland 
plants is very striking. Panicum aciculare and the montane 
Danthonia compressa may be collected side-by-side in dry woods. 
Kuhnia eupatorioides, though found in rich woods on the Coastal 
Plain, is preeminently a plant of the Interior. Near it may be 
collected the striking Sorghastrum Elliottii (Mohr) Nash, here at 
a new inland limit. 

A mile or two up Rattlesnake Creek, west of Triplett, there is 
a swampy pond-hole in the woods, locally known as the “bog ‘ 
Here the slightly retarded creek has spread over a flat and, after 
heavy rain, may become so flooded as to form a considerable 
pond. One might be in southern Southampton County. The 


TE ee 


1943] Fernald,—Virginian Botanizing Under Restrictions 377 


pool and its margin support several Coastal Plain types, their 
known ranges here extended 15 to 30 miles inland: Paspalum 
fluitans (Ell.) Kunth (see Ruopora, xxxix. 382, pl. 474, figs. 
6-10), Cyperus densicaespitosus Mattf. & Kiikenth., Commelina 
diffusa Burm. f. (see Ruopora, xlii. 434) and Hydrolea quadri- 
valuis. In dry woods otherwise typical Andropogon scoparius 
has glabrous, instead of densely villous sheaths, and part way 
up a wooded bluff the northern and upland Pyrola rotundifolia 
var. americana has a sterile colony, a plant one would look for in 
the mountains. 

A little mossy swamp-hole, locally known as ‘Ram-hole” 
Swamp, because of the ram installed there for pumping water, 
is largely given over to retarding brambles, but in the mossy 
thicket there is an abundance of the always beautiful Lobelia 
glandulifera (Gray) Small (see Ruopora, xxxix. 345 and map 20 
on p. 343), a species with three geographic centers, (1) from the 
inner Piedmont of North Carolina across the mountains into 
Tennessee, (2) several hundred miles to the south, the Coastal 
Plain of southwestern Georgia and adjacent Florida, and (3) 
the Coastal Plain of southeastern Virginia and northeastern 
North Carolina, inland to Amelia and eastern Brunswick Coun- 
ties. Here it might have been derived either from the south- 
west or the east; but the goldenrod growing with it may give a 
clue. Resembling the northern and upland calcicolous Solidago 
patula in having almost wing-angled square stems, it yet is very 
different. Its technical characters will be defined in Part II. 
I was delighted to make its acquaintance but somewhat annoyed 
for, having closed off three different times the treatment of 
Solidago, with 75 species and many cross-references by number 
for a revised Gray’s Manual, I was obliged to interpolate another 
and to shift the numbering. Nevertheless it is nice to have in 
Virginia so definite a species as S. salicina Ell. It is a fine 
Species of acid bogs of the Coastal Plain and outer Piedmont, 
described by Stephen Elliott from western Georgia. 

One of the finest pieces of hardwood (beech, oak and hickory) 
in the Seward Forest is back of the old Chamblis place. Here 
are splendid trees and here Dr. Akerman is carrying on some 
very productive experiments in forest-reproduction. The soil is 
deep and, as in all undisturbed beech-oak-hickory forest, full of 


378 Rhodora [OcToBER 


interesting upland plants. Panicum flexile, a characteristic in- 
land grass, follows the wood-road; and toward the base of one 
slope there is a bit of the flora of the Blue Ridge or the Alle- 
ghenies: Polygala Senega var. latifolia, Ligusticum canadense, 
and, most surprising of all, the montane Zizia trifoliata (Michx.) 
Fernald in Ruopora, xlii. 298 and pl. 623 (1940), better known 
as Z. Bebbii (Coult. & Rose) Britt. 

In an old fallow field back of the Chamblis house there is a 
large colony, spread evidently by stolons, of a small Leguminous 
shrub which, Mr. Lewis assured me, never flowers and fruits. 
It proves to be Glycyrrhiza lepidota Nutt., var. glutinosa (Nutt.) 
S. Watson, a rare Licorice from the extreme western side of the 
United States. It is a large and rapidly increasing remnant of a 
colony presumably once cultivated. Having, as a boy, chewed 
licorice-root, I was happy to taste a forgotten nibble, but stu- 
dents now in the university had never even heard of it. One of 
them returned every few minutes, asking for “a little bit more”. 
I finally had the root firmly glued to the herbarium-sheet! 

Hoping for a natural pond-shore, I pushed the inquiry, but 
the best they could suggest was Simms Millpond, southwest of 
Fitzhugh, where the dam had gone out some years ago. Pro- 
ceeding there, we were amazed to see the greatest area of Poke, 
Phytolacca americana (or decandra), probably anywhere in 
existence. The entire drained pond-bottom was a solid, dense 
thicket of it much higher than our heads. We broke our way 
into it, hoping for lingering colonies of something more interesting 
but to no avail. But economically and gastronomically the Poke 
is interesting and I was surprised to find that it was not being 
used. The young shoots, as soon as they sprout in early spring, 
are greatly appreciated as a wholesome and highly nutritious 
vegetable and are brought regularly to city-markets in many 
regions, as Charleston, South Carolina, Washington, Baltimore, 
Philadelphia, etc., as a market-vegetable. The top of the gigantic 
taproot has a circle of many buds. As soon as the leading shoots 
are cut others replace them. The leaves alone, from stems up to 
3 feet high, are a valuable and delicious potherb, and nothing 
more tasty, nutritious and quickly cooked could be found than 
the new stems 2 or 3 feet high, easily rid of the tough rind. 
Phytolacca, only locally appreciated in parts of our country, Was 


1943] Fernald,—Virginian Botanizing Under Restrictions 379 


early carried to southern Europe, where it is cultivated for its 
new shoots. Simms Millpond is locally looked upon as a dead 
loss. The self-sown crop of tremendously vigorous Poke growing 
there could feed all Brunswick County for two months in the 
spring. Tops of the big roots, boxed in earth, frozen, and then 
placed in a cellar and watered, would supply a green vegetable 
all winter, and the new shoots from such a God-given truck- 
farm, sent to an appreciative market, might bring large financial 
return. The difficulty is to get conservative people to eat what 
they do not purchase at the corner-market. 

Following weeks of unbroken drouth rain came very soon after 
I reached Seward Forest. It had come to stay and we came in 
with rain-coats dripping. In the morning of my last day there 
Dr. Akerman defied the storm and joined Lewis and me on a trip 
to the Meherrin River, a few miles to the north, above Western 
Bridge. I specially wanted to see the place because Lewis had 
collected in flower on the wooded river-bluff the southern Tiarella 
with stout rhizomes, no stolons and very slender racemes. I 
wanted to secure fruiting material. Trying to dodge the worst 
downpours, we started up-river in a forest of almost pure growth 
of Acer floridanum, but I very soon worked down to the bottom- 
land-thicket; for I was attracted by several species there. Boltonia 
caroliniana (Walt.) Fern. in Ruopora, xlii. 487, pl. 642 (1940) 
was abundant and very tall, though with pink, instead of white 
ligules, its range extended inland 15 miles or more, and the 
small-headed Heleniwm was certainly the same as that of the 
Coastal Plain farther east. We have accumulated so many 
collections of this Helenium from southeastern Virginia that, in 
desperation, I have tackled the genus. My conclusions will be 
found in Part II. Vernal and aestival species of this bottom- 
land were of course unrecognizable, but one leaning and sprawl- 
ing, very brittle Muhlenbergia at once arrested attention. I had 
just “done” Muhlenbergia for the Manual and had attempted to 
point out more fundamental characters than those commonly 
used. This one, associated with the common M. frondosa (Poir.) 
Fern. in Ruopora, xlv. 235, pl. 749 and 750 (1943), was none I 
had seen from the Atlantic slope. Study of its spikelets and 
comparison in the herbarium shows it to be another prairie 
Species (southwestern Indiana and Illinois to Texas), M. glabri- 


380 Rhodora [OcroBER 


flora Scribn., isolated, like M. brachyphylla (see p. 365) in south- 
eastern Virginia. The lower Meherrin valley begins to compete 
for honors with the Nottoway. In fact, I now hesitate to make 
estimates of the relative interest of the isolated or localized 
floras of the Meherrin, the Nottoway, the Blackwater, and the 
smaller and sluggish rivers farther east. Each has its specialties. 
In the aggregate they are a remarkable and still largely unknown 
flora. 

Below the bridge, the steep bluff gave us battered fruit of 
Tiarella. It is so unlike that of the northern 7. cordifolia, with 
which it had been placed, as a variety, that in Part II I shall 
discuss what seem to me its characters and identity. With it 
were other interesting species and, of course, spring and early 
summer would yield many more. As notable as any now recog- 
nizable were typical Amsonia Tabernaemontana, heretofore 
known in the state only from two stations on the Coastal Plain, 
and Scutellaria serrata, previously unknown in the southern 
counties east of the Blue Ridge—again the juxtaposition of 
montane and Coastal Plain specialties. 

In the late afternoon, my packing being done, preliminary to 
a daylight start next morning, I consented to go with Lewis to a 
little Cephalanthus-pool in the woods, which he has dubbed 
Triplett Pond. Repeatedly, as we drove past it he had gently 
asked: “ Wouldn’t you like to look in there?” As regularly, I had 
declined the invitation. Cephalanthus- and Decodon-holes rarely 
yield much, but in this one he had caught small animals which 
belonged much farther south. I was glad to clear my conscience 
and to have a look before leaving Seward Forest. On the way, 
as we passed through dry pine woods, I was surprised to see 
most characteristic plants of Gnaphaliwm obtusifolium var. 
Helleri (Britt.) Blake, for this very definite variety (see RHODORA, 
xxxviii. 231, pl. 434, figs. 8 and 9), although originally described 
from Northwest in southeastern Norfolk County and abundant 
about Eastville on the Eastern Shore, had never been known 
inland. Here was an extension inland of 85 miles. The slight 
inland extension of var. micradenium Weath. was negligible and 
expected. Reaching the little pond-hole, I was amazed to see 
the coastal Gratiola virginiana var. aestuariorum Pennell (see 
Ruopora, xliv. 440, pl. 730, fig. 3). Surely this was no brackish, 


1943] Fernald,—Virginian Botanizing Under Restrictions 381 


tidal shore. But I was diverted by the big clumps of a coarse 
pilose Panicum, which somehow looked familiar but which I 
could not immediately place; and no wonder, for it is P. longi- 
folium var. pubescens (Vasey) Fern. in Ruopora, xxxvi. 69 
(1934), described by Vasey from Florida and never known north 
of that state. Lewis knew what he was about when he wanted 
me to “look in there’’; and again I had had rubbed in what we 
never learn, that in a country with the complex flora of south- 
eastern Virginia every natural spot, no matter how common- 
place in aspect, may harbor strongly isolated and local plants. 

It was hard to leave the hospitality and stimulating compan- 
ionship of Dr. and Mrs. Akerman and Mr. and Mrs. Lewis, but 
next morning in driving rain Akerman, Lewis and I started in 
the truck for the sand-beaches of ponds in Southampton and 
Sussex Counties. The forester made his observations and we 
had a cheerful and interesting trip but, alas, it had rained without 
letup for five days. When we got to Sedley we were told that 
we could not get at Whitefield’s Pond from the south, for the 
road was completely under water and the dam itself flooded. 
That sounded pretty bad and when we reached Whitefield via 
Corinth, there was the overflowing pond extending back into 
the woods. The farmer living near by told us that in the forty 
years he had lived there the water had never been so low as it 
was until this five-day rain came on. We could have wept. 
Locating a spot where the little Eryngium should be, I walked 
in to shoulder-depth (I was already drenched by rain), ducked 
and grabbed. Nothing but floating Utricularia and debris came 
up. The Eryngium still evades us. Stopping in Emporia at 
the home of Lewis’s sister and her husband, Dr. and Mrs. 
George C. Faville (who were sufficiently venerable to have 
studied botany under the late Professor Charles E. Bessey (1845- 
1915) in Iowa, but endowed with unlimited alertness and vi- 
vacity) and their daughter, Mrs. Wheeldon, I changed to dry 
clothes and was ready for the all-night ride to New York. 

In April Lewis located flowers and young buds on the new 
Asarum and on the 23rd I reached Seward Forest. Restrictions 
on the use of gasoline had greatly tightened. Lewis had planned 
to drive to Emporia (18 miles away) for me but the mails had 
failed to deliver to him my post-card of a week before, saying 


382 Rhodora [OcroBER 


when I would arrive. Taxicabs were allowed to cover territory 
within 10 miles of Emporia and no farther. That distance, 
fortunately, got me to the Brunswick County line, where I trans- 
ferred to the taxi of a local farmer who relayed me to Triplett. 
My home was with one of the foresters and his family, Mr. and 
Mrs. Howard Nicholson and two delightful little daughters. 
The fallow fields were carpeted with the vernal weeds and I 
introduced the family, and especially the children, to the edible 
qualities, particularly as nibbles and salads, of young Henbit, 
_Lamium, and of Peppergrass, Lepidiwm virginicum. 

The Asarum was locally abundant at the bases of wooded 
slopes along creeks and branches, great colonies with heart- 
shaped to slightly halberd-shaped, scattered, solitary firm leaves, 
but flowers were excessively scarce. In a patch with thousands 
of leaves there was often not a single bud nor flower; in another 
patch a few could be found but their peduncles were very fragile 
and it was difficult to secure a good specimen, showing the thread- 
like subterranean rhizomes and stolons, without breaking off the 
flower. Whiteoak Creek and Rattlesnake Creek, investigated at 
many places, yielded their specimens, and colonies with several 
young flower-buds were noted, that Lewis might later secure 
fruit. We here had a very distinct new species and it seemed 
evident that its rare flowering was a result of its occurrence along 
creeks and branches where it is often submerged and where its 
very efficient vegetative reproduction, by slender, subterranean 
stolons, suffices for its local spread. On the last day, one of the 
forest-crew having errands in Emporia, we went with him in the 
truck, in search of this new and very local plant outside Bruns- 
wick County. Since both Whiteoak Creek and Rattlesnake 
Creek are tributary to Fontaine (“ Fountain’) Creek, the obvious 
place to look was farther down the latter valley. At our first 
stop, at the bridge over Fontaine Creek near Barley in south- 
western Greensville County, Lewis promptly walked into it; but 
that seems to be about its eastern limit, not far from the entrance 
of Rattlesnake into Fontaine Creek. Other crossings of Fon- 
taine Creek did not yield it, although the stretch from the mouth 
of Rattlesnake Creek to Round Hill Church could not be ex- 
amined. At Round Hill Church the banks would be forbidding 
to it, acid sand with Kalmia latifolia forming a dense thicket and, 


1943] Fernald,—Virginian Botanizing Under Restrictions 383 


where the white sand is loose and dry, supporting the most 
beautiful and extensive colony of Phlox nivalis I have ever seen. 
In Ruopora, xlii. 476 (1940) I took up for this plant the un- 
equivocal name, P. Hentzii Nutt. (1834) because in the place 
where he started the name P. nivalis Loddiges had given no 
adequate diagnosis. Subsequently, however, Dr. Wherry, in 
Ruopora, xliii. 71 (1941), showed that the name given by Lod- 
diges was validated by Sweet in 1827, when a proper diagnosis 
and fuller description were published. P. nivalis, then, has 
right of way. 

The woods along Fontaine Creek lower down, in an area 
southwest of Dahlia, seemed so like those along Whiteoak and 
Rattlesnake Creeks that we hopefully went there. As Lewis 
agreed, it is just the right habitat but no Asarum could be found. 
Subsequently, by saving up his gasoline, Lewis trailed the new 
species to a small creek in Brunswick County which empties into 
Roanoke River. How extensively it occurs along Roanoke 
drainage can be determined only when more gasoline is available. 
At any rate, this remarkable new species will be described and 
illustrated (pLATES 774 and 775) in Part II. 

Just below Emporia the Meherrin is bordered by a forest com- 
posed largely of a yellow-flowered Buckeye. In 1936 Long, 
Griscom and I had got a little of it in woods along Metcalf 
Branch, which enters the Meherrin a mile farther down; and in 
Ruopora, xxxix. 352 and 435 (1937) and xl. 441 (1938) I erro- 
neously recorded it as Aesculus discolor Pursh, a more western 
species. The freshly flowering material secured from the exten- 
sive forest of the tree shows it to be Ae. neglectus Lindl., var. 
pubescens Sargent. It was a novel experience to wander in such 
a forest on the inner edge of the Coastal Plain. Whereas at 
Triplett, only 18 miles to the west and with creeks at barely 100 
feet greater elevation, spring was just emerging, here it was 
almost summer. Corydalis flavula, which we had known on the 
Coastal Plain of southeastern Virginia only along the James and 
its tributaries was scattering seed; the fruits of Nemophila micro- 
calyx and of Viola striata (also James River specialties) were ripe. 
Great carpets of Phacelia dubia had enough lingering flowers to 
be handsome and, surprise of surprises, here were thousands of 
plants, now passing out of bloom, of upland and inland Trillium 


384 Rhodora [OcTroBER 


sessile, the first member of its genus I had ever seen growing in 
southeastern Virginia. There was no time to linger. These and 
a few other species were tucked into a portfolio and I hurried to 
catch the night train northward, hoping that some “break” 
would soon get Long and me to this botanically unexplored and 
most distinctive area of the Meherrin at the inner margin of the 
Coastal Plain. At least, I could get another and longer visit there 
when, a few weeks later, I should return to Seward Forest to fol- 
low up Rubus, which there seems to be a series somewhat different 
from that of the Coastal Plain. When I regretfully said good-by 
to Lewis and with forty people boarded the north-bound train at 
Emporia, the now familiar announcement of the Conductor 
reached my ears, ‘No seats. Standing room only.’ On the 
platform of the car, however, there were two square feet of space, 
where someone had moved. Setting my suitcase on end and 
placing the portfolio of fresh specimens across it, I had a seat, 
such as it was, all the way to Washington, where the train decided 
to make over before proceeding to New York. Three hours wait 
in a milling throng outside the train-gate from 1 to 4 A. M. was 
finally relieved by announcement of a train to New York. Such, 
inevitably, is travel in war-time. There has been no opportunity 
for another trip, gasoline-rationing having still further tightened; 
but, in spite of varied and discouraging obstacles, something has 
been done to keep the flame of botanical exploration burning. 
The results, more briefly summarized in Part II, are not dis- 
creditable. Beginning the Virginia work with three days in the 
field in 1933, continuing with two to five trips a season for nine 
years and making but one brief trip, as a guest of the Seward 
Forest, in April of 1943, we are assembling much substantial 
data on the flora of the state. In the last report on this work’ 
the records of vascular plants new to Virginia, recorded in this 
series of papers, reached 751. When and if we are again able to 
renew the exploration the score of novelties to the state can begin 
the ninth century. 


1 The Seventh Century of Additions to the Flora of Virginia, Ruopora, xliv. 341-405, 
416-452, 457-478 and plates 717-744—Contrib. Gray Herb. no. CXLV (1942). 


1943] Fernald,—Virginian Botanizing Under Restrictions 385 


Part II. RANGE-EXTENSIONS, TECHNICAL NoTES 
AND REVISIONS 


In Part II, as in previous papers of this series, the notes cover- 
ing extensions of range are assembled, even though the species 
has been discussed in the more discursive journal. Several 
species collected by others and detected in the Gray Herbarium 
during studies recently prosecuted, or some sent by others, are 
included if I can find no record of their occurrence in the state. 
A number of detailed and critical studies are also included since 
they have grown out of comparisons of our Virginia material or 
since they deal with genera occurring in the state. The many 
plates were prepared by Dr. Bernice G. Scuusert, to whose 
skill and patience I am under great obligation. The cost of 
engraver’s blocks has been covered from a grant for personal 
research from the DEPARTMENT oF BrioLogy of Harvard Uni- 
versity. As so frequently in the past, Mr. Lona has most 
generously aided in meeting the expense of publication. 

As heretofore, plants thought to be previously unlisted from 
Virginia or only recently recorded in technical studies of groups 
are indicated, by an asterisk (*). In the enumerations the names 
of the collectors, when Fernald & Long, are generally omitted. 

*POTAMOGETON EPIHYDRUS Raf., var. NuTraLum (Cham. & 
Schlecht.) Fern. XX P. PULCHER "Tuckerm. IsLE oF WIGHT 

OUNTY: forming an extensive carpet, outlet of Lee’s "Millpond. 
no. 12,230. See Ogden in Ruopora, xlv. 184 (1943). The only 

nown eer as of this hybrid. See also Ruopora, xliii. 508 
and 519 (1941). 

*DANTHONIA spicata (L.) Beauv., var. LoNGIPILA Scribn. & 
Merr. See Fernald in RHopora, xlv. 244 (1943), where several 
Virginia collections are cited. 

D. compressa Aust. See Fernald, 1. c. A northern and up- 
land species occurring only locally in the southeastern counties: 

uRRY County: rich calcareous wooded slopes along James 
River, Claremont Wharf, no. 9825. Brunswick CouNTY: oak- 
hickory-beech woods back of old Chamblis Place, Seward Forest, 
near Triplett, Fernald & Lewis, no. 14,458. 6. 
LENBERGIA GLABRIFLORA Scribn. Brunswick County: 
exsiccated bottomland woods above Western Bridge, Meherrin 
River, south of Edgerton, Fernald & Lewis, no. 14,460. The 


western Indiana and Illinois to eastern Texas). See p. 379. 
Digitaria sEROTINA (Walt.) Michx. To the single recorded 


386 Rhodora [OcroBER 


Virginia station add another, also in SourHAMPTON COUNTY 
— shore of Johnson’s Millpond, 144 miles north of Sedley, 
. 14,265. See p. 371. 

Page pissectum L. To the few recorded stations add 
one in Sussex County: sandy shore, Airfield Millpond, south- 
west of Wakefield, no. 14,264. p 

P. FLUITANS (EL.) K unth. See Ryopora, XXxix. 282, t. 474, 
figs. iis (19 7 Local range extended into BRUNSWICK 
Cou wampy pond-hole in woods, Rattlesnake Creek, west 
of Triplett, Pornalt ¢ Lewis, no. 14 462. See p. 377. 

PANICUM FLEXILE (Gattinger) Scribn. BRUNSWICK County: 
clearing in oak-hickory-beech woods back of old Chamblis 
Place, Seward Forest, near Triplett, Fernald & Lewis, no. 14,471. 
ce by Hitchcock & Chase only from Alexandria County. See 

378. 


*P. LONGI aarp tt Torr., var. PUBESCENS (Vasey) hence * 
RHoporRA xxxvi. 69 (193 4). Brunswick County: sphagn 
knolls in woods he Triplett Pond, Seward Forest, near Triplett, 
Fern eo ald & Lewis, no. 14,472. First from north of Florida. See 
p- 

P. ACICULARE Desy. Local range extended inland to BruNs- 
wick County: dry border of oak-hickory-beech woods back of 
a Seward Forest, near Triplett, Fernald & Lewis, 

P. WrIGHTIANUM Scribn. Several additional stations on pond 
shores. Sussex County: Brittle’s Millpond, west of Wakefield, 
no. 14,267; Airfield Millpond, southwest of Wakefield, no. 14,268. 
SovrHampton County: Johnson’ s Millpond, 1% miles north of 
seit no. 14,266; Whitefield’s Millpond, southwest of Corinth, 

70. Istr or WIGHT County: Darden’s Pond, southeast 
of Colo, no. 14,269. See pp. 371 and 372. 

P. auBURNE Ashe. To the few recorded stations add one in 
ISLE OF Want County: dry sandy pine barrens, south of Zuni, 
no. 14,275. 

P. SPHAEROCARPON Ell., var. inFLATUM (Scribn. & Sm.) Hitch- 
cock. In disturbed soil in wet woods along Assamoosick Swamp, 
northeast of Homeville, Sussex County, the stimulated plants 
reach a height of 1 m. with athe up to 1. '6 dm. long and 1.5 cm. 
broad, the panicles 1.5 dm. | and 1 dm. broad. The extreme 
height given by et a Cine for the species is 5.5 dm., 
with panicles up to 1 dm. 

Muhl, Giveutentait County: open, gid 
pine woods, Colonial Beach, F. T. Hubbard, no. 413, a 
Be tee Not recorded from north of North Carolina. 
MUNDUM Fernald in Ruopora, xxxviii. a, t. 443, figs. 1-5 


County: sphagnous pocket, upper border of treiti reed-marsh 
and swale along Northwest River near Northwest, no. 14,272. 


1943] Fernald,—Virginian Botanizing Under Restrictions 387 


*SETARIA VERTICILLATA (L.) Beauv. RocksripGE County: 
along sidewalk, Lexington, August 23, 1924, J. R. Churchill._— 
Not indicated for Virginia on Hitchcock’s map 1567 (Man.). 

THE COMMON NORTHERN BurGRass.—The common Burgrass 
of open sands, from New Hampshire to Oregon, south to North 
Carolina, Kentucky, Missouri, Kansas and New Mexico, is 
indigenous or spread from indigenous colonies through much of 
this range but, singularly enough, it has always passed under 
specific names which technically belong to other species. Long 
identified as Cenchrus echinatus L., it was forced to resign that 
name to the quite different tropical species. Similarly the name 
C. tribuloides L. soon proved to belong to the very coarse tropical 
and southern coastal species. Then C. carolinianus Walt. was 
despairingly grasped, but Walter’s type is unknown, our plant 
is doubtfully in his territory, and Mrs. Chase, Contrib. U. 8. Nat. 
Herb. xxii. 76 (1920), believes that C. carolinianus might have 
been C. incertus M. A. Curtis, which does grow in Walter’s 
country. In her monograph of the genus she merges our plant 
with the Mexican C. pauciflorus Benth. and, until Dr. I. M. 
Johnston, working upon his Mexican and southern Texan 
material, segregated it off from the great bulk of specimens from 
the northern and Rocky Mountain region of the United States, 
it so rested. Dr. Johnston, not wishing to get involved with the 
more northern and northeastern plant, called the matter to my 
attention. There is no doubt that C. pauciflorus is the Mexican 
(and sone) species, Bentham having described ‘it from Lower 
California: “culmis suberectis, . . .~“ “Folia plerumque an- 
gustiora [quam in C. echinata], spinis dorsalibus marginalibusque 
validis basi dilatatis’’. 

Cenchrus pauciflorus, then, is the Mexican and Texan species 
with culms usually in erect or ascending tufts, the leaves 1.5-4 
mm. broad, the summit of the sheath spreading as a chartaceous 
flange; spikes 1-5 em. long, 1-1.5 cm. thick; the stramineous 
involucres 8-12 mm. broad (from tip to tip of mature spines), the 
dorsal and lateral spines compressed and broad-based. The 
characteristic fruit, from the TyPx of C. cocsiesata is shown by 
Mrs. Chase, 1. ¢. fig. 17, p. 68. 

Our plant, on the other hand, has more generally decumbent 
or rooting culms, up to 8 dm. long and strongly geniculate; 


388 Rhodora [OcToBER 


leaves 3-8 mm. broad, the enlarged sheaths constricted at sum- 
mit; spike 1.5-8 em. long and 1.3-2 cm. thick, the mature 
stramineous to bronze or purplish burs 1—1.5 cm. in diameter, 
with the coarse spines subulate-subterete. The bur is beautifully 
shown, as that of C. pauciflorus, in Chase, 1. c. 69, fig. 18, and this 
figure, instead of that of true C. pauciflorus, was copied in 
Hitcheock, Man. fig. 1594. As compared with that of real C. 
pauciflorus it is too large, with more numerous spines, the stronger 
ones more slender and without deltoid and flattened bases. 

So far as I can find the only available name for the northern 
and transcontinental species is C. echinatus, forma longispinus 
Hackel, based upon Connecticut material collected by Harger 
and distributed by Kneucker (no. 426). As a diagnosis of a form 
of C. echinatus, in the loose sense, Hackel’s brief description was 
sufficient. Treated as a species the plant demands a fuller 
account. It is fortunate that Hackel has supplied the basonym 
and type. Even so, I take no comfort in having even my name 
intimately associated with a Cenchrus and the keen taxonomist 
who called it to my attention would not care to have the species 
named for him! 

Crncurus longispinus (Hackel), stat. nov. C. echinatus, forma 
longispinus Hackel in Kneucker, Allg. Bot. Zeitschr. ix. 1 
(1903). Pl i i 


ad 8 dm. longis geniculatis basi ramosis; foliis 3-8 mm. latis, 
— distentis apice constrictis; spicis maturis 1.5-8 cm. 
longis 1.3—2 cm. crassis; involucris maturis stramineis vel aenels 
vel purpurascentibus hirsutis 1-1.5 em. diametro, spinulis majori- 
bus tereti-subulatis. Typr of C. echinatus, forma longispinus: 
Oxford, Connecticut, Z. B. Harger in Kneucker, Gram. Exsicc. 
Lief. XV. no. 426. 

In southeastern Virginia, fortunately, Cenchrus longispinus is 
rare. Our only collections are the following. James CITY 
pda sandy field about 5 miles west of Toano, R. W. Menzel, 

one OUTHAMPTON County: dry white sand of clearing in 

and pine woods bordering Assamoosick Swamp, south o 

Sebrell no. 10,941; grassy roadside about 7 miles south of 
Franklin, no. 7297 (misidentified as C. incertus). 


ERIANTHUS COARCTATUS Fernald in Ruopora, xlv. 246, t. 758 
(1943). Note that the plant of Sussex County previously 
reported as E. brevibarbis Michx. belongs here. FE. brevibarbis 
is a very different plant of the Mississippi Basin. 


1943] Fernald,—Virginian Botanizing Under Restrictions 389 


viminea (no. 12,564) from Greensville County we had not yet 
got the flowers. The later collection (no. 13,883), made October 
13, 1941, shows the plant to be var. variabilis, which is charac- 
terized by having delicate awns up to 9 mm. long. Blake’s 
plant, collected along the James River at City Point, near 
Hopewell in October, 1931 (Blake, no. 11,472) is of typical Z. 
viminea (i. e. Pollinia imberbis, var. Willdenowiana (Nees) 
Hackel), with awnless spikelets. Typical E. viminea occurs 
southward to Alabama: shaded bank of Tennessee River, north- 
east of Sheffield, Colbert Co., Oct. 7, 1934, Harper, no. 3275. 

The earliest valid generic name for the genus is Eulalia 
Kunth, Rév. Gram. i. 160, t. 93 (1829), although Hackel, like 
most Old World botanists, has maintained Pollinia Trin. in 
Mém. Acad. Pétersb. ser. VI. ii. 304 (1833). In so doing they 
may have been misled by Index Kewensis, which lists just before 
Pollinia Trin. another Pollinia Spreng. Pugill. ii. 10 (1815), 
“farrago [mixture] = seq. &c.”’ The “&c” is correct, for 
Sprengel had 10 species, none of them considered by Hackel as 
belonging in Pollinia Trin.: 6 referred by Hackel directly to 
Andropogon; 1 to Oplismenus; 1 to Ischaemum; and 2 wholly 
doubtful “species inextricabiles’”’ (Hackel, 1. ¢. 565). Pollinia 
Trin. (1833), clearly antedated by Eulalia Kunth (1829), must 
lapse. 

In looking up the origin of the generic name Eulalia the 
quickest method was to turn to the shelf of dictionaries and to 
see what Wittstein said in his Etymologisch-botanisches Hand- 
worterbuch. There it is: ‘Eulalia KNTH. (Gramineae). Zus. aus 
ev (schén, gut) und AeAte (Rede, Gericht, Ruf); ein schénes, 
guldgelbbliihendes Gras”. The type of the genus being £. 
aurea Kunth, that derivation is plausible. Had Wittstein taken 
the trouble to look up Kunth’s own explanation, however, he 
would have found a more pleasing and splendidly appropriate 
explanation of the name, for it is difficult to imagine more beauti- 
ful illustrations (all signed E. D.) than the 220 wonderful plates 


390 Rhodora [OcroBER 


by Eviauia Dewite which clarify Kunth’s remarkable volumes. 
That Kunth was fully appreciative of his artist is made clear by 
his own explanation: “Dixi hoc genus in honorem Eulaliae 
Delile, cujus perito elegantique penicillo icones hujus operis ut et 
quorumdam aliorum debet botanice”’. 

*ANDROPOGON scopaRiIus Michx., var. GENuINUS Fern. & 
Grisc. in RHopora, xxxvii. 143, 144 (1935), forma calvescens, 
f. nov., vaginis gla bris. VrrGInia: dry woods along Rattlesnake 
eae apa of Triplett, Brunswick County, October 11, 1942, 


Herb. Phil. Acad.); mois baondes 4 nd peaty pine barrens, south 
Se 


1940, Fernald & ye no. 12,567. See p. 377.— Quite like typical 
Andropogon scoparius (= var. items Keamey) but wit 
the sheaths stabrous’ instead of se 

SorGHASTRUM Exziottir (Mohr) Nash. Ketel range extended 


inland to Brunswick County: a, woods along Rattlesnake 
ak ‘abel of Triplett, Fernald & gle no. 14,473. See p. 376. 

OSTYLIS CILIATIFOLIUS (EIl.) Fern. New northeastern 
Persie IsLE or Wicut County: dry sandy pine site: 
south o : 


on Coun 
open alluvial flat by Blackwater River, southeast of Unity, no. 
14,284. Be p. 369. 


CIRPUS a BTERMINALIS Torr. Princess ANNE COUNTY: 
shallow alee northwest eae of Salt Pond, June 28, 1922, 
L. F. & Fannie R. Randolph. 

Although Britton, Ill. Fl., cites Scirpus subterminalis § as eX- 
tending south to South Carolina and Small, Man. 169, says 
“Miss.’”’, I have been unable to locate previous material from 
south of New Jersey and Pennsylvania. There is nothing in the 
Gray Herbarium nor in the Britton Herbarium from south of 
New Jersey and Pennsylvania, except the Salt Pond specimen; 
and the rather extensive collecting in eastern South Carolina 
and eastern North Carolina by Godfrey did not bring it to light. 
Beetle, however, in Am. Journ. Bot. xxx. 396 (1943) seems to 
have seen material from both South Carolina and Mississipp1. 


Tue Speciric CHARACTERS OF Scirpus OLtNEYI—One of the 
most distinctive species of Scirpus, § Schoenoplectus, is the 
coarse, soft-stemmed S. Olneyi Gray (1845), which follows saline 
and brackish marshes from eastern South America up the 
Atlantic to southern New Hampshire and Nova Scotia, the 
Pacific to Washington. In many characters the plant is so 


1943] Fernald,—Virginian Botanizing Under Restrictions 391 


distinctive that amateurs who do not distinguish between some 
technical species usually recognize this one at a glance. It is, 
therefore, a bit startling to find in the Am. Journ. Bot. xxx. 397 
(1943) Beetle merging it with and reducing it to S. chilensis 
Nees & Meyen (1843). Beetle’s detailed description of the 
inclusive species, as S. chilensis, clearly says “‘style 2-fid’’, which 
is correct for the tropical and North American S. Olneyi, and in 
his discussion he speaks of the original diagnosis of S. chilensis as 
“adequate to identify the plant’. He then quotes the diag- 
nosis of S. chilensis, including the original and easily corrobo- 
rated “‘stylo trifido”’; but he gives no explanation of how a plant 
of the warmer half of the Northern Hemisphere with “style 
2-fid”’ is necessarily, or even probably, identical with a plant of 
temperate South America with “‘stylo trifido’”’. This distinction 
is constant, apparently. So are several others. The broadly 
wing-angled soft culm of S. Olney? is so very soft that in pressing 
it becomes flat and ribbonlike; in fresh condition, however, it 
has deep reéntrant angles which caused Asa Gray, in originally 
describing it, to give the following vivid description: ‘“‘distin- 
guished .by its remarkably 3-winged stem. The reéntering 
angles are so deep that the cross section presents the appearance 
of three rays, or plates with parallel sides, joined by a common 
center.’’ The collapsing of the culm under very slight pressure 
results from its hollow or fistulous character, the pith occurring 
as scattered remnants only. If S. Olneyi is identical with S. 
chilensis it is notable that the much more slender culms of the 
latter are firm and resistant, not flattened in drying, that they 
lack the extremely broad wing-angles and deep reéntrant con- 
cavities and that they are closely filled with pith. 

In S. Olneyi the upper leaf-sheaths have a U- or V-shaped 
orifice and the brown tissue of the sheath-summit is readily 
friable; in S. chilensis the orifice is subtruncate or very shallowly 
concave and the tissue firmer. In S. chilensis the excurrent 
midribs of the scales project more prominently than in S. Olneyi. 
In S. chilensis, as already noted, there are 3 style-branches, in 
S. Olneyi 2. Although the achenes of the bipartite S. chilensis 
sensu Beetle are given absolute dimensions, ‘2.5 mm. long, 1.5 
mm. broad’’, the few achenes of S. chilensis available, from 
Pennell, no. 12,923 and Osten, no. 22,029, are definitely more 


392 Rhodora [OcroBER 


slender than in S. Olneyi. These two numbers of S. chilensis 
have achenes narrowly obovate and three fifths to three fourths 
as broad as long: 2.6 K 1.8 mm., 2.4 X 1.8 mm., 2.3 K 1.5 mm 
On the other hand, ripe fruit from material of S. Olneyi from 
the general type-region, southern New Hampshire to Virginia, 
shows much more rounded or broadly obovate achenes four fifths 
as broad to essentially as broad as long: 2.2 *K 1.8 mm., 2.2 X 2 
mm.; 2.4 X 2.2mm., 2 X 1.8mm., 2 X 1.6 mm., ete. Whether 
or not typical S. Olney? occurs in temperate South America I am 
not situated to determine. Barros & Osten described a S. 
Olneyi, forma australis from Uruguay in Anal. Mus. Hist. Nat. 
Montevid. ser. 2, iii. 204 (1931) and in vol. xxxviii. 159 and 161 
(1935) Barros illustrated it and cited much material from 
Argentina but did not suggest that it includes S. chilensis. In 
fact, in the latter detailed study Barros definitely cited (p. 156) 
S. chilensis as identical with S. americanus Pers. I am content 
to keep up Scirpus Olney as a thoroughly distinct species. 

In Virginia Scirpus Olneyi makes extensive colonies often 2 m. 
or more oe on the saline and brackish shores. 

*S. vaLipus Vahl, var. cREBER Fern., forma MEGASTACMYUS 
Fernald in Raoneee xlv. 283, t. 765, fig. 8 (1943). James CITY 

OUNTY: marsh of Chickehorniay River about 5 miles west of 
Toano, R. W. Menzel, no 

_ BTUB ERCULATUS (Steud.) Ktze. Princess ANNE COUNTY: 

shallow water, northwest branch of Salt Pond, June 29, 1922, 
L. F. & Fannie R. Randolph, no. 462. 


Scirpus etuberculatus, one of the most localized of species, has 
a long-known station near Salisbury, Maryland, one or two in 
southeastern North Carolina, two in Georgia, one in Florida, 
two in Alabama, one in Mississippi, and one in Louisiana. The 
station of Mr. and Mrs. Randolph is apparently the first between 
southeastern North Carolina and satel gaeae 

+8, rus Pursh, sien protrusus, f. nov., spiculis plus 
iniaee: ‘ceieda ad 3-5 em. lon ngis. — Von: brackish to 
fresh marsh along Back Bay 4 at eastern margin of Long Island, 
Princess Anne County, August 25, ty Fernald & Long, no. 
10,973 (type in Herb. Gray., isotype in Herb. Phil. Acad.); 
border of salt marsh, Ragged Islan a northeast of 2 oleae 
Isle of Wight County, August 20, 1940, Fernald & Lon 
12,587.—Differing from typical S. robustus (w: ith spikelets 1. 33 
em. long) by its very prolonged spikelets. 


1943] Fernald,—Virginian Botanizing Under Restrictions 393 


S. MARITIMUs L., var. FerNaup1 (Britton) Beetle. See Fern. 
in Ruopora, xlv. 289 (1943). S. novae-angliae Britton. Our 
only collections are from the James River or its tributary creeks. 

URRY County: tidal marsh at mouth of Crouch Creek, east of 
Scotland, no. 8593, previously reported as S. novae-angliae. 
JAMES Crry County: tidal shore of Back River, opposite James- 
town Island, no. 10,972, distrib. as S. robustus Pursh. 

*S. MARITIMUS, var. FERNALDI, forma aGoNus Fern. I. ¢. 288 
(1943). New KEnt County: fresh tidal ae te Lacey Creek, 
west of Walker, no. 13,559. 

HYNCHOSPORA FILIFOLIA Gray. Sussex County: upper 
ae of siliceous and page se shore, Airfield Millpond, 
southwest of Wakefield, 14,301. Fi rst station between 
i North Canola and ‘Cape May, New Jersey. See 


"Cae DECOMPOSITA Muhl. iin oo Thor Bipesar stations 
add one in SourHampron Coun cypress swamp 
bordering Womble’s (or Wade’ a ‘Dilipenit nora of Baffle, no 
14,305. See p. 373. 

Bayarpi Fern. C. virginiana Fern., not Woods. Range 
extended northwestward in SOUTHAMPTON County: sandy al- 
luvial bottomlands of Three Creek, Adams Grove, no. 14,306. 

ee p. 367. 

C. venusta Dewey. To the few recorded stations add one in 
Dinwippie County: wet. spri ngy sphagnous woods east of 
Cherry Hill, no. 14,307. See p 

C. Coutinstr Nutt. To ths a ‘recorded stations add one in 
Dinwippie County: wet springy sphagnous woods east of 
Cherry Hill, no. 14,311. See p. 366. 

CoMMELINA DIFFUSA Burm. f. Local range extended inland to 
BruNswick County: swampy pond-hole in woods, Rattlesnake 
Creek, west of Triplett, Fernald & Lewis, no. 14, 478. See p. 377. 

MIANTHIUM ve tag nivgelny (Walt.) Gray. Local range 
extended northward in IsLe or WiGHT CounTY: ag in damp 
sandy pine barrens south of Tei, no. 14,315. See p. 369. 

NOTHOSCORDUM BIVALVE (L.) Britt. Range extended inland 
from Princess Anne and Northampton Counties: James City 
County: turf back of beach ra disturbed soil in woods and 
thickets back of sand-beach of James River, Martin’s Beach, 
southeast of Grove, Fernald, eae & Abbe, nos. 14,128 and 14,129. 

SLE OF Wigut County: turf back of sand-beach of Burwell’s 
Bay, James River, below Rushmere (Fergusson’s Wha 
Fernald, Long & Abbe, no. 14,127. See pp. 360, 364. 

alge CANADENSE L., var. editorum,' var. nov. (TAB. 771, 

5), & var, typico me foliis mediis ellipticis vel oblongis 
Hore vel greed s, vix attenuatis, latitudine 4-14 partem 
' Editorum (of the uplands) from editum, upland or a height; not from editor! 


394 Rhodora |OcroBER 


longitudinis aequante; floribus rufescentibus; perianthii tubo 
elongato, tepalis supra medium sensim arcuatis; petalis (siccis) 
0.8-1.3 cm. latis—Locally from the mountains an le- 
gheny Plateau of Pennsylvania to Kentucky and the mountains 
of Alabama. PENNSYLVANIA: Waddle, Center County, July 3, 
1939, J. P. Be thickets along Conoquenessing Creek, Butler 
County, July 17, 1932, John Bright, no. 6789. Out 0: Columbus, 
1837, Lesquereux? West VIRGINIA: els near aoe re River, 
Hampshire County, July 1, 1933, Hunnewell, no. 12,771. Vir- 
GINIA: open mountain-meadow at about 4000 feet alt., top of 
Butt Mountain and at about 3500 feet alt. near Little Stony 
Creek near “Cascade Road”, Giles County, July 24, 1943, A. 
sane no. 4617 (TYPE in Her b. Gray.); Jeon gi Virginia Hot 
Springs, Bath County, July 5, 1917, Panowal, & 4824; woods 
eb 2700 fee t alt., Shenandoah Mt., Rockin gham Saunt July 14, 
1932, Hennes no. 12,481. KENTUCKY: swampy meadows, 2 
miles south of London, Laurel County, July 4, 1939, McFarland, 
no. 3545; oak-chestnut mountain-summit forest, at 4000 feet alt., 
Black Mt., Harlan County, July 22, 1937, E. Lucy Braun, distrib. 
as L. canadense or L. michi anense. ALABAMA: rich woods on 
limestone, north slope of Monte Saho, Madison County, June 22, 

32;.7T. 8. Van Al 

Freshly collected material of Lilium canadense var. editorum 
reached me just as this Contribution was starting to the printer, 
Professor Massey sending it with a query as to its real identity. 
Typical northern Lilium canadense, described from Canada and 
occurring from the Gaspé Peninsula of Quebec to northeastern 
Ohio, south to Nova Scotia, New England, Pennsylvania and 
upland to western Virginia, has the lanceolate (narrowly to 
broadly) leaves of the median whorls attenuate to acute tips, the 
blades },—% as broad as long, the usually yellow flower (red only 
in the rare forma rubrum Britt.) with relatively short and thick 
tube, the tepals strongly arching (r1c. 1) from below or near the 
middle, the petals 1.2-2 em. broad. Occasional specimens have 
the leaves of var. editorum but the flower of typical L. canadense. 
Such a transitional series is represented by MacDaniels & 
Thomas, no. 3710 from Ithaca, New York. It is noteworthy, 
then, that Ithaca is near the northern border of the Allegheny 
Plateau as mapped by Fenneman. 

The chief points of departure of var. editorwm are its relatively 
broad and only slightly if at all tapering leaves, its red flower 
with slender and elongate tube, so that the arching of the tepals 


1943] Fernald,—Virginian Botanizing Under Restrictions 395 


starts near or above the middle, and the narrow tepals, the petals 
in dried specimens being only 0.8-1.3 cm. broad. One collection 
before me, C. EH. Wood, Jr., no. 1365, from sedge-meadow, Little 
Meadows, northwest of Mountain Lake, Giles County, Virginia, 
has the leaves narrower than and as attenuate as in the most 
extreme northern plant, with flowers nearly of var. editorum. 

A number of the above cited specimens were sent to the Gray 
Herbarium with indications of doubt as to the identity and 
usually with note of the red flower. The Bright material from 
Butler County was marked in the hand of a temporary assistant 
“Lilium michiganense’’, while the McFarland specimen was 
identified with doubt as L. canadense, and the old sheet from 
Columbus, Ohio, originally identified as L. canadense, bears 
modern annotations, first as L. superbum L., later as L. michi- 
ganense Farw. Until Professor Massey sent his material the 
question of an undescribed montane variety had not been con- 
sidered. The accumulated discontent with the identification of 
the plant lead to its more critical study. 

When we know ripe fruit of Lilium canadense, var. editorum 
the seeds may show some further differences. The ripe fruit of 
typical northern L. canadense is well represented in the collec- 
tions before me, by 19 numbers in fully mature fruit, collected in 
eastern Canada and interior New England, north or inland from 
the coastwise northern limit of L. superbum and far east of the 
quite different inland L. michiganense Farw. The fully developed 
seeds of L. canadense vary from 7-11 mm. long, mostly 9-10 mm. 
It is hoped that mature fruit of the montane plant may soon be 
available. 

The frequent misidentification of Lilium canadense, var. edi- 
torum as the eastern and southern L. superbum or as the midland 
L. michiganense reflects dissatisfaction with its being called 
simply the northeastern L. canadense but it also indicates lack of 
understanding of the morphological characters which separate 
L. michiganense and L. superbum from one another and both of 
them from L. canadense. 
ste inti by the Inte Elsie Lou TEN so Echeverria loaves 
from ~_ tburn Race hire, July 30, 1924, Walter Deane. Fras. 3-5, 
var 4. flower and median leaves from TYPE; FIG. 5, 
Bones & fren Butler Ouudee Paciayivssin: John Bright, no. 6789. All figs. ML. 


396 Rhodora [OcroBER 


TRILLIUM SESSILE L. GREENSVILLE County: rich woods along 
Meherrin River, below Emporia, Fernald & Lewis, no. 14,530, 
very abundant. Our first record from the Coastal Plain. See 
p. 383. 


Tue Dwarr TRILLIUM OF SOUTHEASTERN VIRGINIA (PLATES 
772 and 773).—In southeastern Virginia the genus Trilliwm is so 
very local and scarce that in ten seasons of field-work I had 
never met any member of the genus there growing wild until Mr. 
Lewis and I came upon the extensive colony of 7’. sessile above 
noted. I had, however, reported in Ruopora, xlii. 445 (1940) 
the occurrence in the Great Dismal Swamp of a plant which I 
then misidentified as T. lanceolatum Boykin; Professor Smart 
had told me of the occurrence near the University of Richmond of 
a scarce plant thought to be 7’. pusillwm Michx., this discovered 
in May, 1931; and in the Gray Herbarium there is, under T. 
pusillum, a sheet of exactly similar material from near Powhatan 
Creek in James City County, discovered by Mrs. W. G. Guy and 
Dr. Stetson, also in early May, 1931, and sent to the Gray Her- 
barium by Professor Warren. Furthermore, Mr. Lewis told me 
of a station in Dinwiddie County which had been shown to Mrs. 
Laura H. Lippitt by the friend who had discovered it. From 
this station, rich loamy woods near a stream in company of 
Royal Fern, Sensitive Fern and Lady-fern and Medeola (a very 
frequent association), Mrs. Lippitt has sent me a beautiful series 
of freshly flowering specimens, with the note that the petals are 
white, turning pink and then dark purple. 

Altogether, the dwarf Trillium has a considerable, though 
highly localized, occurrence in the southeastern counties of 
Virginia; and, in identifying the Meherrin River 7. sessile, it 
became quite apparent that the tiny plant is all of one distinctive 
species and that it is neither 7’. lanceolatum nor typical T. pusil- 
lum. The latter, T. pusillum Michx. (pLaTE 772), was described 
as having the leaves sessile, the flower peduncled and erect, the 
sepals scarcely longer than the petals, the petals pale flesh-color; 
and it came from pinelands of South Carolina. The Michaux 
material (our FIG. 1) well agrees with this description and the 
label gives the further information: ‘35 m. de Charlest. environ 
Gaillard road”. Just such a plant, rigs. 2 and 3, is known from 
eastern South Carolina. It has the flower long-peduncled, the 


1943] Fernald,—Virginian Botanizing Under Restrictions 397 


petals about equaling or even longer than the sepals, 1.8-2.5 cm. 
long and 4-9 mm. broad; the anthers sometimes 5-6 mm. long, 
definitely longer than the filaments, or in otherwise typical 
South Carolina plants only 3 mm. long and definitely shorter 
than the filaments. The plant of southeastern Virginia (PLATE 
773, FIGs. 1 and 2) looks like T. pusillum, but its flowers are 
sessile or elevated on a peduncle up to only 5 mm. long. If the 
latter embarrassing individuals are excluded, it is one of the 
“‘sessile-flowered”’ species; if they are taken into account it is a 
species with peduncled (‘‘pediceled’’) flowers, a bit awkward in 
key-making. Its petals are mostly shorter than the sepals, 
1.2-2 em. long and 3-5 mm. wide, the anthers 3-8 mm. long and 
only slightly if at all longer than the filaments. 

In view of the inconstancy in length of anthers and the over- 
laps in other characters I am treating the plant of Virginia as a 
geographic variety of Trillium pusillum, rather than as a distinct 
species. The sessile flowers of most Virginia material were 
responsible for one collection being placed with 7. lanceolatum. 
That more southern species, however, has the sepals soon reflexed 
and the long and tapering petals (our PLATE 773, FIG. 3) with 
slender claw-like bases. The Maker plant is 

TRILLIUM PUSILLUM Michx., var. virginianum, var. nov. (TAB. 
773, Fic. 1 et 2), a var. typicum jenedit flore sessile vel subsessile ; 
petalis 1.2-2 cm. longis 3-5 mm. latis, antheris 3-8 mm. longis.— 
Southeastern Virernia: Henrico Count nty: damp woods north of 
Westwood Golf Course, May 8, 1931, R. F. Smart & Elmer C. 
Pritchard (Herb. Univ. of Richmond); woodland north of Uni- 
versity Road, Westwood, May 8, 1931, Mary E. Billings. JAMES 
Crry County: Long Hill Swam mp, Powhatan Creek on yay 
ville Road west of Williamsburg, discovered by Mrs. W. G. 
and Dr. Stetson, coll. May 3, 1931, by Paul A. Warren. Nonistx 
County: Great Dismal Swamp, west of Wallaceton, April 24, 
1926, Paul A. Warren, no. 413. Dinwipp1r County: Tich loamy 
woods near stream, 5 miles east of tee Court House, May 
9, 1943, Laura H. Lippitt (ryPE in Herb. 

In view of the evident dimorphism as : rte of anthers in 
both typical Trillium pusillum and its var. virginianum, it is 
evident that too much weight has been placed upon the length of 
anthers in the group. See P. 364. 


Pate 772 is of Tritium pusiLLuM Michx., X 1 1, Michaux’s TYPE, 
original photograph by Cintract; Fic. 2, plant from Pinopolia, South Canlinn’ 


398 Rhodora [OcroBer 


April, 1897, Maria P. Ravenel; Fic. 3, plant from Pinopolis, May, 1895, E. 
Peyre Por cher. 
In PLATE 773, Figs. 1 — 2 stg from the stair apap x 1, of T. pusm- 
LUM, Var. VIRGINIANUM ; FIG mit o ANCEOLATUM Boykin, 
X 1, from Aspalaga, Florida, ‘April 11, 1902, Biltmore Het. no. 6085. 


*NARCISSUS BIFLORUS Curtis. IsLE or WicHT County: rich 
calcareous wooded slopes by is Shes s Bay, James River, below 
Rushmere (Fergusson’s Wharf), Fernald, Long & Abbe, no. 14,130 
Originally spread from cultivation. 

CALOPOGON PALLIDUS Chapm. Range extended northward in 
IsLe oF Wicut County: clearing in damp sandy pine barrens 
south of Zuni, no. 14,320. 

*ULmus PROCERA Salisb. CaroLine County: steep wooded 
bluff by Rappahannock River, northwest of Return, Fernald, 
Long & fssiey no. 14, si See op 

*ASAR m (§ HET monuoed! 1 ewisii, sp. nov. (TaB. 774 et 
ee elabrescens seule gracili subterraneo pallido valde mt 
(ad longo), urcato ra ius stoloniferis; oles lon 


ee solitariis coriacei ae nusve macu lat s Maioidae 
vel deltoideo-ovatis vel subrenifonnibue obtusis, sini basalari 
rotundato, laminis maturis 2-8 cm. longis 2.5-8.5 cm. latis; flore 


pedunculato subnutante, ealyed campanulato perien glabro 
griseo-brunneo, intus atropurpureo villoso, 2-3 em. longo 1.3-2 
em. diametro, lobis vix patentibus. —Southea astern Brunswick 
County and southwestern Greensville County, Vrrainia: bottom- 
land-woods along Laces es Creek, west of Triplett, Bruns- 
wick Coun ty, Octobe er 1942, Fernald & Lewis, no. 14,480; 
sandy loam in woods eae Whiteoak, ‘Creek: near Triplett, April 
23, 1943, Fernald & Lewis, no. 531; mixed woods along 
Rattlesnake Creek, boli: Wri ht t’s Beds April 24, 1943, 
Fernald & Lewis, no. 14,532 (rypx in Herb. Gray.); mixed woods 
along Rattle snake Creek north of Anku um, Brunswick County, 
May 24, 1948, Lewis, no. 3825; bottomland-woods along Fontaine 
Creek west of Barley, Greensville County, May 17, 1943, Lewis; 
along Pea Hill Creek about a mile north of Gas burg, Brunswick 
County, May 29, 1943, Lewis, no. 3826. See pp. 374, 381-383. 
In some ways combining the characters of our two eastern 
American sections of the genus. It has the deltoid to ovate or 
reniform, evergreen and often mottled leaves much as in A. 
arifolium Michx. or in A. Ruthii Ashe; while the extensively 
creeping and stoloniferous stems fork as freely as in A. canadense 
L., but they are subterranean, not superficial, much more slender, 
and bearing the scattered leaves singly along the axes and at the 
tips of the widely creeping stolons. Mr. Lewis had found only 
sterile plants, always in colonies on bottomlands or just above 


1943] Fernald,—Virginian Botanizing Under Restrictions 399 


the bottomland-levels of small creeks of southeastern Brunswick 
County, Virginia, which empty into Fontaine Creek and, even- 
tually, into Meherrin River. In October, 1942, I had the great 
pleasure of visiting the Seward Forest, as the guest of the Direc- 
tor, Professor Alfred Akerman and Mr. Lewis, and when they 
showed me the strange Asarum it was, as Mr. Lewis had written, 
characteristic of the thin bottomland-woods just above creek- 
margins and where, during high water, it is regularly overflowed. 
It is colonial, making extensive colonies, with the rather small 
deltoid to ovate long-petioled leaves scattered (one-at-a-time), 
never in tufts. 

In late April of 1943 I again was a guest at the Seward Forest, 
for Mr. Lewis had written that he had discovered a few flower- 
buds on different colonies, the very young buds showing some 
weeks after A. canadense and A. virginicum had begun blooming. 
On April 23, Dr. Akerman, Mr. Lewis and I found a few fully 
expanded flowers and several very young buds, usually only one 
fertile plant amongst hundreds of sterile ones, along Whiteoak 
Creek, near Triplett, and next day about the same meagre pro- 
portion of buds and fully grown flowers at the bases of gentle 
Slopes to Rattlesnake Creek, these chiefly among fallen beech- 
leaves. And on April 26, when I had to be driven to Emporia 
to take the train back to Boston, we conceived the idea, 
since both Whiteoak and Rattlesnake Creeks empty into Fon- 
taine Creek, one of the choice habitats of Greensville County to 
the east, of trailing the new species down that valley into the 
Coastal Plain. The first crossing to the eastward in Greensville 
County reassured us, for along Fontaine Creek, not far from its 
receipt of Rattlesnake Creek, near Barley, Mr. Lewis promptly 
detected a colony. That was the last we saw. Later, however, 
Lewis trailed it to Pea Hill Creek, which empties into the Roanoke 
River, his station being 10 miles farther west than those near 
Triplett. We are, then, forced to consider the new species as a 
highly localized one of southern Brunswick County and south- 
western Greensville County. 

Although as extensively creeping as Asarum canadense, its 
resemblance stops there. In foliage A. Lewisii is, as stated, 
related to A. arifolium and to A. Ruthii of Asarum, § Heterop- 
tropa (the genus Hezastylis Raf., Small and their followers), 


400 Rhodora [OcroBER 


while its large drab-brown to drab-purple, campanulate calyx is 
nearly as large as in A. Shuttleworthii J. Britten (southwestern 
Virginia, Tennessee and Alabama), but with the outline of that 
of A. virgintcum. In all these characteristic species of Asaruwm, 
§ Heterotropa, the rhizome is stout and short, covered heavily 
with thick and very elongate roots and at summit forking into 
ascending crowns bearing tufted leaves. The new Asarum 
Lewisii, with which it is a great pleasure to associate the name of 
its discoverer, the perennially enthusiastic naturalist of the 
Seward Forest, Joun Barziuuar Lewis, is unique in the section 
in its prolonged, slender, horizontal, subterranean, cord-like 
whitish rhizomes with few slender roots, the axes and the stolons 
bearing only scattered leaves. With this very effective vegeta- 
tive reproduction and inhabiting bottoms where the plants are 
frequently submerged, A. Lewisizi seems to flower (and then very 
sparingly) only when the colonies, as in late April and May of 
1943, remain unsubmerged during the normal flowering period. 

No fruit seems to form and the fragile peduncle readily breaks, 
even when the flowers are only partly grown. 

PARONYCHIA RIPARIA Chapm. Limit of range extended north- 
eastward in SourHaAmMpron County: dry sand of open alluvial 
sgl od Blackwater River, spntheneh of Unity, no. 14,324. See 
p. 3 


> cde ein (L.) Wood. Local range extended eastward 
into IsLe or WiauHt County: rich wooded slope above bottom- 
land woods along Blackwater River, above Broadwater Bridge, 
north of Zuni, no. 14,325 
HowosteuM UMBELLATUM L. Apparently spreading; new or 
sandy roadside-fills i in Essex and Hanover Countigs. Perhaps 
pare = ries seed bet on new soft shoulders. See p. 359. 
(L.) Cyrill., var. GLABERRIMA G. Bec 
See _Soomeestl ‘ii “451 (1904). Add a station in IsLE or WIGHT 
County: turfy waste ground — of sand-beach of Burwell’s Bay, 
James River, below Rushmere (Fergusson’s Wharf), Fenald 


magn as hacer itisitet pes Desportes. Local range extended 
north to Starrorp County: Rartorigatith about 4 miles southeast 
of Paluonthe Fenalhe Long & Abbe, 14,144. 

NYMPHAEA ODORAT er a forma rRUBRA Guillon. Sussex 
County: in water at alia. of Chappell’s Millpond (Honey 
Pond), west of eran no. 14,327, growing with the common 
white-flowered for 

ABOMBA ciiereank Gray. NansemMonp County: ditch 


1943] Fernald,—Virginian Botanizing Under Restrictions 401 


along highway, northern border of Great Dismal Swamp, east 
f Magnolia, no. 14,328. Our first station in the Tidewater 
counties. See p 68. 

AROROOLER. ABORTIVUS L., var. INDIVIsUS Fernald. aoa 
FORD County: disturbed soil at border of rich woods a 


. Sarpous Crantz. Local range Bee into Sussex 
Country: siliceous me ae oni fallow field, Jarratt, Fernald, 
Long & Abbe, no. 14,151. In view of the dominance of this 
species in fields fh pastures about Franklin, likely to spread 
rapidly. 

MORPHOLOGICAL DIFFERENTIATION OF CLEMATIS OCHROLEUCA 
AND ALLIES (PLATES 776-782)—The simple-leaved and erect 
species of Clematis, of which C. ochroleuca Ait. Hort. Kew. ii. 260 
(1789) was the first American one described, has had many treat- 
ments. So far as I can find there is no clear demonstration of 
the identity of the plant described by Aiton. It was cultivated 
in England and very briefly described, the name given on account 
of the color of the sepals. It is and has been assumed to be the 
most eastern of species, found chiefly in the Piedmont and inner 
Coastal Plain region from southeastern New York to south- 
eastern North Carolina and the mountains of northern Georgia. 
Until Aiton’s material is checked, with the real morphological 
distinctions in the group clearly in mind, the established interpre- 
tation should not be disturbed. It is evident, certainly, that 
the Plukenet plant from Banister cited by Aiton ‘‘flore unico 
ochroleuca”’ is the plant generally known as C. ochroleuca. 
Plukenet’s figure is characteristic. 

My object in this note is to define on more stable and funda- 
mental lines than are relied upon in the two latest treatments, 
the eastern species of Clematis, § Viorna, sub-§ Integrifoliae. 
The latest treatment, that of Erickson in Ann. Mo. Bot. Gard. 
xxx. 13 and 35-39 (1943), keys our species in a manner quite 
reminiscent of the key of Wherry in Journ. Wash. Acad. Sci. xxi. 
195 (1931). Wherry’s key was as follows: 

ie — sparingly branched and small leaves » posed few; 

head of fruit tending to be eee. about 6 cm. in diam- 
eter; achenes nearly symme 

Under — of leaves Valeute to moderately pubescent; 

ac nang Ta es deep, or exceptionally pale, 


voll vated range chiefly at altitudes below 1000 feet, 
mostly i in PONG a te es CG. ochroleuca ovata 


402 Rhodora [OcroBER 


Under side of me moderately to densely pubescent; 
hairs of achene-appendages pale, or naapy i argh deep, 
yellow; range vchicdly at altitudes above 1000 feet, mostly 
| UPESA ET PRO or PA Ne ene crs aye meray area a ee pois io sericea 
Plant copiously cerca and small leaves relatively nu- 
eaves 
Head of fruit nearly. pears about 5 cm. in diameter; 
“orig airly symmetrical, their appendage-hairs 
PLUME ee oe Oe tun Ce mae he Me sie es C. viticaulis 
Head of fruit sagen about 4 cm. high and 6 em. broad; 
unsymmetrical, their appendage-hairs 
Oe Nl Pathe ae Gees vee cae es at os C. albicoma” 


As above stated, the latest monographer of the section gives in 
his key no more fundamental distinctions than these and leans 
primarily on the evasive and contradictory degree of branching 
(note the contradiction in his ‘‘EE” and its subordinate “FF”’, 
below) and the still more evasive degree of coloring (note ‘‘ EE” 
with ‘achene-tails usually lighter in color” as contrasted with 
“FE” in which the color is “reddish-brown”, the first species 
under the “lighter . . color” being C. ochroleuca in which 
the tails (styles) may be “‘tawny’’. Here is Erickson’s state- 

-ment of the “‘specific”’ differences: 


“KE. Plant profusely bine ge leaves lanceolate, less than 
als glabrous; achene-tails reddish- 


SG iweup ee ok ae esc eee aw Mig ig Sie ade b:« 13. C. viticaulis. 
EE. Plant I co branched; leaves ovate, at least some of them 
more than 6 cm. ‘long; sep: 3 pubescent; achene-tails 


usually lighter in pet d 
F. Plants simple or few-branched, more or less pubes- 
achene-tails light yellowish-brown to 
ae rer eer ee COLT ere 11. C. ochroleuca. 
FF. View usually much branched, often white silky- 
pubescent throughout; achene-tails whitis - 
Saks WONOW fo eves he eRe es Seo ee es 2. C. albicoma.” 
Not having the field-familiarity of Dr. Wherry with C. albicoma 
and C. viticaulis, I have been forced to my interpretation of these 
two treatments thus handicapped. By following the above keys 
I arrive at very perplexing results, for the characters depended 
upon are such as vary with extreme plasticity; and, whereas 
Erickson says in his key that the leaves of C. viticaulis are 
“lanceolate” as opposed to “ovate” in C. ochroleuca and C. 
albicoma, his fuller account of C. viticaulis says “leaves lanceolate 
to narrowly ovate’’, the other two species being assigned leaves 
which may also be “narrowly ovate’’. 
As to the degree of branching so much relied upon in the two 
treatments, I, again, have been handicapped by not having 


1943] Fernald,—Virginian Botanizing Under Restrictions 403 


before me all the material their authors studied, except that in 
the Gray Herbarium, which was borrowed by Erickson. I have, 
however, had the great advantage of studying a large series of 
specimens from the Blue Ridge in Roanoke County, Virginia, 
most kindly sent me for study by Mr. Carrol E. Wood, Jr. 
Taking into consideration only the plants which are complete 
down to the lowest node (omitting obviously broken-off branches) 
I get the following results. 
NUMBER OF BRANCHES FROM Main AXIs 

C. LEUCA, “plant sparingly branched” or “simple or few- 
branched” “a7 plants): branches 0 sn yeni l=1; 2=5; 3=4; 

4=7;5 =6;6=7:7 =3:9=1, Average 3.2 bra 

C. viricau.ts, ‘plant copiously” ~ He heanched (7 plants): 
ia aes ae 3=1-4=2: §=1;6=1;8 =]. Average oe branches. 


‘ COMA, typical glabrescent plant, ‘“copiously ”’ ayers much 
hranehad's (19 specimens): branches 0 =4; 3=1; 4=2; 5= ae 
8 =3. ye ne 4.5 beaten 

C, MA, tomentos var., Bon copiously”’ or “usually much 
branched” (36 ’plants): prion O=4; 1=4; 2=5; 38; 4=6; 5=4; 

3;7=1:8=1. Average 3.2 sede sh 

With more than half of the compiete plants of the “simple or 
few-branched”’ C. ochroleuca (PLATE 776, FIG. 1) having 4-9 
elongate axillary branches, while the “profusely” or ‘usually 
much branched”’ species show nearly half the plants with less 
than 4 branches, I am quite incapable of applying the character 
as a clear diagnostic one or even a worth-while tendency. 

As to leaf-outline and size one also has to be pretty cautious. 
The sheet of Adams & Wherry, no. 2413, from the type-locality 
of Clematis viticaulis, defined in Erickson’s key as having “leaves 
lanceolate, less than 6 cm. long”’, shows blades 6.5 em. long by 
3cm. broad. As already noted, such leaves are narrowly ovate, 
rather then “lanceolate”. Other topotypic specimens of C. 
viticaulis show primary leaves with the following dimensions: 
Killip, no. 32,484 (PLATE 778, Fic. 1), blades 7 em. long; Adams 

Wherry, no. 2418, blades 7.8 em. long by 4 cm. broad; Wherry, 
June 11, 1930, blades 7 cm. long by 4.6 cm. broad, much more 
than “narrowly ovate”. As a key-character “leaves lanceolate, 
less than 6 cm. long” is not wholly satisfactory, especially since, 
likewise, so many specimens of C. albicoma in the most restricted 
sense have leaves scarcely different in shape and size, topotypes 
(from Kate’s Mountain) showing the largest leaves 6.2 cm. long 


404 Rhodora [OcroBER 


(Gilbert, no. 511), 6-6.5 em. (Marion S. Franklin, September 4, 
1920, our PLATE 779, FIG. 1), 6.2 em. (Small, May 16, 1892), 7 
em. (Hermann & Martin in Pl. Exsice. Gray., no. 951), 7 em. 
(Core, no. 2708), 5.8 em. (Addison Brown, July 22, 1892), while 
plants from near Deerfield, Augusta County, Virginia (Wherry, 
no. 2420), have the larger leaves only 4.5 cm. long. Here, again, 
those who hope for clarification find “a distinction without a 
difference’’. 

In his key Erickson assigns Clematis viticaulis ‘‘sepals glabrous” 
as opposed to the pubescent sepals in the other plants. I hope 
they are glabrous; Steele, in describing the species, did not know 
them, and Wherry cited besides Steele’s collection only his own 
from the type-locality, collected June 11, 1930. As shown by 
the two flowers of Wherry’s plant in the Gray Herbarium, the 
sepals (PLATE 778, FIG. 2) are closely pilose. In fact, in his 
detailed description (p. 38) the “sepals glabrous” of Erickson’s 
key were allowed to be ‘‘slightly pubescent without’’. Those of 
Wherry’s topotype, however, are certainly less pubescent than 
in C. albicoma (PLATE 779, FIG. 4) or in C. ochroleuca (PLATE 776, 
FIG. 3). 

As to the fruiting head, in Wherry’s treatment that of C. 
ochroleuca is ‘‘tending to be spherical, about 6 em. in diameter,” 
that of C. viticaulis ‘nearly spherical, about 5 cm. in diameter” 
and that of C. albicoma “spheroidal, about 4 cm. high and 6 cm. 
broad’”’. The abundant series of C. ochroleuca before me shows 
that the mature fruiting heads (PLATE 776, Fic. 4) range from 
5-10 (average 7) cm. in diameter, with the long plumose styles 
loosely separated at the margin of the head, while in C. albicoma 


the mature head (pLaTE 779, Fic. 1 and 780, Fic. 6) is compact, © 


with tightly recurving plumose styles, the heads 4-7 em. thick. 
The fruiting head of C. viticaulis (PLATE 778, Fics. 1 and 3) is as 
small as in C. albicoma but with fewer and loosely spreading 
plumes (much as in C. ochroleuca). I am not sufficiently a 
mathematician fully to appreciate the differences between “tend- 
ing to be spherical”, “nearly spherical” and ‘‘spheroidal”’. 
According to the Century Dictionary a spheroid is “A geometrical 
body approaching to a sphere, but not perfectly spherical’”’. 
Since in C. ochroleuca the fruiting head is ‘tending to be spheri- 
cal” and in C. viticaulis only “nearly spherical” while in C. 


1943] Fernald,—Virginian Botanizing Under Restrictions 405 


albicoma it is “spheroidal” (i. e. “approaching to a sphere, but 
not perfectly spherical’’) this character is too erudite for me. 
Similarly, the exact shades of difference between ‘nearly 
symmetrical’’, “fairly symmetrical’? and “rather unsymmetri- 
cal”’ are difficult to visualize. Furthermore, the distinctions 
between ‘deep, or exceptionally pale, yellow”, ‘‘pale, or excep- 
tionally deep, yellow” are not wholly eitiidactoey: In charac- 
teristic Clematis ochroleuca, the easternmost plant, the mature 
plumes range from a deep, almost cinnamon-brown to whitish- 
yellow or pale buff, the range in color being comparable to that 
in the mature perianths of Eriophorum virginicum; and, although 
the plumes of C. albicoma are commonly whitish-gray or drab 
(on white paper they do not look ‘‘whitish”’), the series of its 
more pubescent and relatively eastern variety sent me for study 
by Mr. Wood shows some numbers with plumes as brown as in 
average C. ochroleuca. Exactly the same range of color occurs 
in other species of § Viorna. In C. Viorna L. Erickson rightly 
allows the ‘‘achene-tails  . . light yellow or brownish”’. 
In some recently collected (therefext not faded) material before 
me (Godfrey, no. 5004 from Wake County, North Carolina) they 
are as pale as in extreme C. albicoma, while in other material 
(Godfrey & Tryon, no. 806 from Orangeburg County, South 
Carolina) they are as dark as in the most extreme C. ochroleuca 
or as in C. viticaulis. Color of hair, like the other characters 
used in recent keys, is not sufficiently stable to demonstrate that 
C. ochroleuca, C. viticaulis and C. albicoma are clear-cut species. 
There are, however, some characters of deeper significance, 
which seem to show that Clematis albicoma and C. viticaulis are 
really separable. In fact, Erickson recorded the most significant 
characters in his descriptions of the three but these points, which 
are the strongest ones, were omitted from his key in favor of 
superficial and wholly inconstant ones. I refer to the great 
elongation of the fruiting peduncles in C. ochroleuca (PLATES 776 
and 777), as opposed to the relatively short peduncles of the 
others, and to the very different direction of the pubescence on 
carpels and achenes. These characters are constant in all the 
material I have seen, without regard to degree of pubescence of 
stem and leaf, size of leaf, amount of branching and paleness or 
deepness of color of the coma. To me they are the soundest 


406 Rhodora [OcTroBER 


differential characters, though not so easy to see without careful 
examination. 

The white-silky or -tomentose plant (PLATE 780) concentrated 
on the Blue Ridge, of which Mr. Wood sends me a splendid 
series for study (largely as a loan) is evidently what Wherry had 
primarily in mind when he published the combination C. ochro- 
leuca, var. sericea (Michx.) Wherry. Since it is not C. sericea 
Michx. (PLATE 777) and since its more stable characters are those 
of C. albicoma, to which Wherry often referred specimens and 
with which essentially glabrous and mostly more western plant 
(pLATE 779) Erickson merges this white-pubescent one of the 
Blue Ridge, I am redefining the latter. As I understand the 
three species and the variety, I treat them as follows: 


a. Carpels and achenes with appressed pubescence, that 
the summit pointing forward; lowest villi of the pasting 
style ascending or _proasting ssceniiing, fruiting head 
with styles loosel ara 
Stems either vomedlba or loosely heacatied: the branches 

only exceptionally overtopping the main axis; larger 
leaves of primary stem 6-12 cm. long, silky-pilose to 
pebente ag appa flowers 2-3.5 cm one the cinereous 
backs of the als densely silky- — mature 
fruiting peduncle lengthening to 5-19 (av. 11+-) cm 
much overtopping the su ten ing Raves mature 
fruiting head 5-10 (av. 7) cm. in diameter, the plumose 
PUN, os eee RA ea dee es 


C. ochroleuca. 


main axis; os leaves of primary axis 5-7.8 cm. long, 
i s* e 
diameter, the brownish plumose styles 2-3 cm. long... .C. viticaulis. 
e- 


consequently ign ag fruiting peduncles 3-9 cm. long; 
flowers 1.7-2. at ig central axis often or ceaadie 


c 
Main axis 2-3 igh, the y ate loosely pilose but 
pecoming glabrescent; leaves at ae sparingly pilose 
ns eath, otherwise rous, soon quite 
elabrate, the ‘larger ones 4~7.5 on. long and 1.5-4.5 : 
hs RO Ss bik 5k Sis Galle Ch es 2 os wes ORAS be C. albicoma. 
hae axis 2-4.5 d m. high, the pres densely oe E 
sistently sik Aca eh ; leaves persistent] 
tomentose beneath, the vedi bed 6-10 cm. Sim pe 
4-9 em. broad C. albicoma, var. coactilis. 


ee 


Since the descriptions, ranges and citation of specimens of 


1943] Fernald,—Virginian Botanizing Under Restrictions 407 


C. ochroleuca (our PLATE 776) and C. viticaulis (pLATE 778, 
Figs. 1-4) have been so well given by Erickson, it is here neces- 
sary only to define the restricted C. albicoma and its var. coactilis. 


*CLEMATIS ALBICOMA Wherry, var. coactilis, var. nov. (TAB. 
780), planta habitu C. albicomae typicae simillima plerumque 


centrali 2-4.5 dm. alto densissime persistenterque _villoso- 
tomentoso; foliis dilatatis axis primarii 2-3-jugis ovatis majoribus 
deinde 7.5-10 cm. longis 5.5-9 cm. latis coriaceis subtus persis- 
tenter piloso-tomentosis; foliis ramorum elongatorum 2-4-jugis 
majoribus 6-10 cm. longis 4-6 em. latis; pedunculis crassis dense 


villoso-tomentosis deinde 4-9 (av. 6.6) em. longis; sepalis anguste 
Ovatis 2.8 cm. longis 1.3 cm. latis dorso zie villoso-tomen- 
tosis; capitulo fructifero denso 4-6 (av. 4.8) em. diametro; 


achaeniis oblique subehoinbsldet-obay atis plus minusve strigosis 
dorso apiceque divergenter villosis; stylo maturo valde arcuato- 
recurvato 3-5 cm. longo, albido- vel sordido-plumoso, villis 
inferioribus horizontaliter divergentibus superioribus plus minus- 
ve adscendentibus.—Mountains of western Viretnra: Roanoke 
County: hoe southwest-facing slope of Green Ridge, about 2.25 
miles Bets nd of Hanging oe April, 1940, Carroll E. Woo 


miles northeast of Hanging Rock, June "30, 1942, Wood, nos. 


Tuly 13, 1942, Wood, nos. 3920 and 3936. J i pa eltet County: 
shale barren 1 mile northeast of Ironto, June 12, 1939, Wherry 
(distrib. as C. albicoma Mion ry or C. ochroleuca, var. sericea 
(Michx.) Wherry). Craig County: shale barren along Craig 
Creek, 0.5 mile nd estore “of Newcastle, August 13, 1937, Fogg, 
no. 13,2 255; barren along Broad Run, 3.5 miles south of New- 
castle, ‘June 12, 1939, Wherry (distrib. as C. albicoma). 

The very full series assembled by Mr. Wood and identified by 
Dr. Wherry as Clematis ochroleuca Ait., var. sericea (Michx.) 
Wherry in Journ. Wash. Acad. xxi. 197 (1931) demonstrates to 
me very effectively the distinctness of C. albicoma, var. coactilis 
from C. ochroleuca Ait. and from typical C. albicoma Wherry 
(PLATE 779). C. ochroleuca (PLATES 776 and 777), occurring 
from southeastern New York to Georgia, is a relatively taller 


408 Rhodora [OcroBER 


plant (2-6 dm. tall), either simple or loosely branched, the 
branches only exceptionally overtopping the main axis, the inter- 
nodes more elongate; the young foliage silky-pilose beneath, the 
silkiness (PLATE 776, FIG. 2) either persistent or disappearing; the 
young stems pilose, the mature ones sparsely so to glabrate; the 
fruiting peduncles slender and elongate, 5-19 (av. 11+) cm. 
long and sparsely pilose to glabrate (PLATE 777, FIG. 2); the 
fruiting head (pLaTE 776, Fic. 4) 5-10 (av. 7) em. in diameter, 
with the long plumose styles loosely separated at the margin of 
the head, either yellowish-brown or drab; the achene (Frc. 5) 
strigose-sericeous to summit, the gradually arching style 46.5 
em. long, with basal villi (r1a. 5) loosely ascending or spreading- 
ascending. 

C. albicoma (PLATE 779) was not formally described by its 
author, who said that it was ‘‘C. ovata of current manuals, not 
Pursh’’, leaving the reader to figure out the characters except 
as stated in his key: C. ochroleuca (var. ovata (Pursh) Wherry) 
and C. ochroleuca, var. sericea (Michx.) Wherry defined: ‘ Plant 
sparingly branched and small leaves relatively few; head of fruit 
tending to be spherical, about 6 cm. in diameter; achenes nearly 
symmetrical’. Opposed to this is the call embracing C. viticaulis 
Steele and the new C. albicoma: “Plant copiously branched and 
small leaves relatively numerous; leaves glabrate”’, under which 
C. albicoma is defined “Head of fruit spheroidal, about 4 cm. 
high and 6 cm. broad; achenes rather unsymmetrical, their ap- 
pendage-hairs whitish”. Since typical C. ochroleuca may have 
3 or 4 pairs of elongate branches (Staten Island material from 
T. F. Allen—ruate 776, FIG. 1—or from E. G. or N. L. Britton, 
Brooklyn specimens of John Carey, Washington material of L. F. 
Ward, and various specimens from eastern Virginia and North 
Carolina), the first differentiation, as already emphasized, is not 
clear; and as also noted, furthermore, the difference between 
heads ‘‘tending to be spherical’ and heads ‘“‘spheroidal’’ is not 
vivid. We are left with the achene of C. ochroleuca “nearly 
symmetrical”, that of C. albicoma “rather unsymmetrical”’. 
C. albicoma is typified as the plant of Kate’s Mountain, Green- 
brier County, West Virginia. Taking that as a starting point 
and the descriptions of “C. ovata of current manuals”’ the con- 
cept is somewhat clarified. It (pLaTE 779) is a usually low- 


1943] Fernald,—Virginian Botanizing Under Restrictions 409 


branching plant with the main axis 2-3 dm. high, the lateral 
branches soon strongly overtopping it; stem loosely pilose 
(FIG. 2), becoming glabrescent; leaves at first sparingly pilose on 
veins beneath, otherwise glabrous, soon quite glabrate (FIG. 3); 
well-developed leaves of primary axis 1-3 pairs, the larger ones 
5.5-7.5 (av. 6.25) em. long and 2.5-4.4 (av. 3.6) em. broad; 
elongate ascending branches with 3 or 4 pairs of well developed 
leaves, the larger blades 4-7 (av. 6) em. long and 1.5-4 (av. 3) 
cm. broad; fruiting peduncles sparsely short-pilose or glabrate, 
in fruit 3-8 em. long; sepals (ric. 4) lanceolate, 1.7—2.7 cm. long 
and 4-8 mm. broad; fruiting head (Fic. 1) depressed-globose, 
4~7 cm. broad; achenes (ria. 5) divergently long-villous on back 
and at summit; the fruiting style about 4 em. long, horizontally 
villous at base, the whitish-gray villi more ascending toward the 
apex. 

C. albicoma, typified by the plant of Kate’s Mountain, and the 
newly proposed var. coactilis (PLATE 780) are nearer to each other 
than either of them is to the more eastern C. ochroleuca (PLATES 
776 and 777). I agree with Dr. Wherry that C. ovata Pursh, 
based upon a Catesby specimen from Virginia in the Sherard 
Herbarium at Oxford, is C. ochroleuca. A tracing of it made by 
me in 1903 shows the characteristics of the latter and it may be 
noted on PLATE 776 that in 1881 Asa Gray reached the same con- 
clusion. Judging from his very brief characterization C. ochro- 
leuca, var. sericea sensu Wherry, 1. c. 195 and 197 (1931), partly 
included C, albicoma, var. coactilis. He did not note the rather 
striking difference in the pubescence of the carpels and fruits of 
C. ochroleuca and of C. viticaulis, as contrasted with those of C. 
albicoma, here discussed and illustrated, nor the marked differ- 
ence in length of peduncle, size of fruiting head and compactness 
or looseness of the latter. 

Clematis albicoma, var. coactilis is not C. sericea Michx. Fi. 
Bor.-Am. i. 319 (1803); and the combination C. ochroleuca 
sericea (Michx.) Wherry, 1. c. 197, resting upon Michaux’s name, 
nomenclaturally is the same as C. sericea. The tyre of C. 
sericea (at least the material preserved in Michaux’s herbarium) 
consists of two branches (PLATE 777) in young fruit. They both 
have the relatively sparse pubescence (rics. 2 and 3) and the 
very elongate and slender peduncles (11 em. long) of C. ochro- 


410 Rhodora [OcroBER 


leuca, also the characteristic appressed-ascending villosity (F1G. 4) 
of the fruiting style-base. In interpreting Michaux’s original 
account it must be borne in mind that he thought that his C. 
sericea was perhaps Aiton’s C. ochroleuca, that he was comparing 
it with the Eurasian C. integrifolia L. (pLaTE 778, Fia@s. 5-8) 
which has nearly glabrous or only minutely hirtellous stem (FI¢. 
5), glabrous or promptly glabrate leaves (Fic. 6), elongate- 
lanceolate sepals up to 5 cm. long (ria. 7) and nearly or quite 
glabrous on the back, and loose fruiting head (Fra. 8) much like 
that of C. ochroleuca. With C. integrifolia before him Michaux 
described his American material as 


SERICEA. ae caule erecto, pubentissimo: foliis simplicibus, 
ted junioribus calycibusque extus incano-sericeis: 
ulo terminali, solitario; flore cernuo. 
it 


Ops. Affinis C. integrifoliae; flore conspicue oad Beg! de 
angustiore. Folia sessilia, opposita, pubescen 
HAB. in Virginia et in montanis Carolinae. 

As contrasted with that of Clematis integrifolia the stem of C. 
ochroleuca is “pubentissimo’’, and the lower surfaces of the 
young leaves (PLATE 776, FIG. 2) and the outside of the calyx 
(FIG. 3) are sericeous. The calyx of C. ochroleuca is much shorter 
than in C. integrifolia, though the sepals are of about the same 
shape. In the montane C. albicoma, var. coactilis (PLATE 780, 
ric. 5) the sepals are much broader. The Michaux diagnosis 
and comparative note give a good characterization of the common 
C. ochroleuca of the Piedmont and inner Coastal Plain region 
from southeastern New York southward, a species Michaux 
would have had great difficulty in avoiding. 

In 1875 Sereno Watson received from his relative, Dr. Louis 
Watson of Ellis, Kansas, a variable series of the most western 
representative of § Viorna, sub-§ Integrifoliae from Ellis. Some 
of the specimens, with heavily coriaceous subrotund to ovate 
or oblong, obtuse leaves, Watson associated with a poor old 
specimen without stated locality collected on Fremont’s 2nd 
Expedition; and from this series of 12 plants he described 
Clematis Fremontii Watson in Proc. Am. Acad. x. 339 (1875). 
The series shows simple or branching plants, varying from 1.4 to 
4.5 and more dm. (judging from broken-off specimens up to 5.5 
dm.) high. Some have heavy oblong leaves at most 5 em. long 


1943] Fernald,—Virginian Botanizing Under Restrictions 411 


and 2 cm. broad (PLATE 781, FIG. 3), others narrowly ovate ones 
(FIG. 2), while the broadest-leaved specimens show rounded- 
ovate and very heavy blades 1.4 X 1.15 dm. (Fig. 1). These 
form a consistent but highly variable series and are typical C. 
Fremontii. They all have coriaceous foliage strongly overlap- 
ping, short internodes and stout fruiting peduncles 0.5-4 em. long. 

One of the Ellis specimens (PLATE 782, FIG. 1), however, not 
marked by Watson as his C. Fremontii, as were all the others, 
but later so labeled by Asa Gray, differs in its elongate internodes, 
relatively thin and elliptic-oblong to lance-ovate leaves and 
slender peduncle, in anthesis more than 5 cm. long. This speci- 
- men is intermediate between the most divergent material of var. 
Riehlit Erickson, 1. c. 40 (1943), a plant set off on just these 
characters and said by its author to have ‘‘ complete geographical 
isolation”. PLatE 782, ric. 1 shows the summit of the narrow- 
leaved plant from Ellis, Kansas (the specimen identified by 
Erickson as rypicaL C. Fremontii). Fic. 2 is a tip from Allen- 
ton', Missouri (coll. June 10, 1887, by G. W. Letterman) of var. 
Riehlii. Fic. 3 is a leaf and fruiting peduncle from another 
Letterman Collection, called by Erickson var. Riehliz because 
from eastern Missouri. In view of the figures given by Erickson 
of leaf-outlines from ‘‘ Mass collections” of C. Fremontii made 
in Ellis County, it is a pity that he did not get hold of the plant 
shown in our PLATE 782, ria. 1 or such plants as shown in PLATE 
781, Frias. 2 and 3, also from Ellis. They would have materially 
changed the outlines. Var. Riehlii is, it should be evident, not 
always ‘completely isolated” from typical C. Fremontii. Dr. 
Louis Watson got it along with the other variations in Ellis 
County, Kansas. Whether it is more than an ecological variant 
of more sheltered or less arid habitats is doubtful. Otherwise it 
would not in Franklin County, Missouri, have such rounded- 
ovate leaves as that shown in PLATE 782, Fria. 3 (identified and 
cited as var. Riehlii) and in Ellis County, Kansas, sometimes so 
closely imitate the eastern plant with elongate leaves. 

1 To those who know, what any good atlas will reveal, that St. Louis, Allenton and 


citations accompanying the original description of var i COUNTY NOT DE- 
birsaieesi A OR DOUBTFUL: t. a ‘ ee eons ho cte ig ONE - 
a succeeding article, ‘but not in the one where the sta ment was published, 


ieukale: 1 c. 64, it is revealed that the doubt was as to woe the cited specimens 
tiesregit were collected in St. Louis, Allenton and Eureka, not as to the county in 
which these towns are 


412 Rhodora [OcroBER 


PLATE 776 is of CLEMATIS OCHROLEUCA Ait.: FIG. 1, characteristic branching 


plant, < 2s, from Staten Island, New York, May 23, 1884, T. F. Allen, with 
memorandum made a Gray of | its identity with C. ovata Pursh; Fria. 2, 
lower surface of leaf, * 10, from Staten Island, 18 } FIG. 3, 


x 

fruiting head, < 1, from Four Mile Run, Virginia, Pollard, no. 334; ria. 5, 
anhene and base of tail, < 5, from east of McKenney, Virginia, Fernald 

ong, = 

Pua E777, TYPE of a SERICEA Michx. = C. ocHroLeuca Ait cal 
ang, x 14: ria. 2, stem of plant at lett, x 214; ria. 3, is ih x aN, 
FiG. 4, tails of fruit, « 2 5, from plant at 

, C. vITICAULIS Stecle: FIG. 1, topotype, X 1%, from 
pec no. 32,484; Fia. "2, back and margin of sep al, x 10, from topotype, 
a 


achene and base of tail, x 5, icy topotype, Adams & Wherry, no. 2418. 

Ias. 5-8, C. InTEGRIFOLIA L.: FIG. 5, stem, X 10, from Alkabek River, pits. 
southern Seal, M eve, io 766: FIG. 6, lower surface of lea 0, 
from no. 766; G. 7, flowe 1, from Hungary , F. Schultz, Herb. Norm. n 
2501; Fic. 8, fruiting i. x ar fro dion . 2501. 

Puate 779, C. ALBICOMA Wherry: Fic. 1, portion of topotype, x 1, fro 
Maison *S. Franklin, September 4, 1920; FIG. 2, stem, X 10, from eesteiery 
F, A. Gilbert, no. 511; ria. 3, lower surface of leaf, x 10, yn no. 511; Fie. 4, 
flower, < 1, from m topotype, Heller, no, 842; ria. 5, achene and base of tail, 


PuaTeE 780, C. ‘AL BICOMA, var. COACTILIS Fernald, all (unless noted) fro 


TYPE: FIG. 1, plant, X rs FIG. 2, bgp e, X 10; Fic. 3, hig! surface of let, 
<4 ‘Eta > summit of sepal, < 10; Fic. 5, flower, x 1, 0 * 
o. 127 tania head. aay "from Wood, no. 3497. mo. 7, ‘achene de 
base of tail, 


PuaTe 781, C. ‘Fremonti §. Wats., all from original series from Ellis, 

ansas: ee 1-3, leaves, X 1, from different individuals; ria. 4, achene and 
base of tail, 

Pate 782, “C. FRremontnn S. Wats.: Fria. 1, leaf mang flower, < 1, from Ellis, 


Kansas, Dr. Lou is Watson, vio labeled by Erickson as typical C. Fremontii; 
Fria. 2, leaf and ‘decade d flower, 1, of var. Riehlii Erickson, from Allenton, 
Missouri, June 10, 1 siete: Fic. 3, leaf and fruit, < 1, labeled by 


Erickson as var. Riehliz, from Franklin County, Missouri, June, 1882, Letter- 


ASIMINA PARVIFLORA (Michx.) Dunal. Northern soit of 
range extended well into IsLz or Wicnt County: sandy wooded 
slope by Blackwater River, west of Blackwater School, no. 
14,331. Be: ee p. 370. 

Co LIS FLAVULA (Raf.) DC. To the few recorded Coastal 
Plain vations add the following. Srarrorp County: rich woods 
along brook about 3 miles southeast of froma Pane. 
Long & Abbe, no. 14,157. GrrENsviLLE County woo 
along Meherrin, River, below Eiipois Fernald ‘€ edt no. 
14,536. Seep 

FUMARIA “orviersata L. To the few recorded stations add 

ne in Essex County: weed in old field near War - ’s Wharf, 
northeast of Te aeviiis Fernald, Long & Abbe, no. 14,15 

ROTUNDIFOLIA L. Local range extended ri the west 
of ree Paskeae: River (former inland limit on Coastal Plain 
in SourHampron County: mossy thicket by Cephalanthus-pool 


1943] Fernald,—Virginian Botanizing Under Restrictions 413 


near Johnson’s Millpond, north of Sedley, nos. 14,332 and 
14,333. See p. 371. 

. CAPILLARIS Poir. To the several known stations add an 
exceptionally fine one in Sussex County: upper border of sandy 
eet of Airfield Millpond, southwest of Wakefield, no. 14,334. 

ee p. 37 
(To be continued) 


VIRGINIAN BOTANIZING UNDER RESTRICTIONS 
M. L. FERNALD 
(Continued from page 413) 

TIAREL 
(1937), 
Acad. Sci. Phil. no. 42: 3 (1940) 
Wherry, 1. c. 4 (1940). To the counties already re- 
(Amelia, Mecklenburg, Henry and James City) add the 


following. Brprorp County: May 7, 1871, A. H. Curtiss. 


446 Rhodora [NoVEMBER 


Brunswick County: bluffs of Meherrin River, near Westward 
Mill, May - 1942, J. B. Lewis, no. 3410; rich wooded bluffs 
below Weste n Bridge, Meherrin River, south o gerton, 
October 12, 1949, Fernald & Lewis, no. 14 482. See pp. 379, 380. 
I get no satisfaction in trying to separate from Tvarella 
Wherryi, described from Tennessee, southwestern North Carolina 
and Alabama, the non-stoloniferous plant found from the Blue 
Ridge eastward to the Coastal Plain. According to Wherry, |. c., 
“T. Wherry? differs in flowering later, in the stems usually bearing 
one or more leaves, and in the acutish sepals rarely being over 
2 mm. long and 0.75 mm. wide”’, whereas his 7. cordifolia, var. 
collina (type from Henry County, Virginia) has the “leaves 
similar to those of var. austrina . . . in tending to be longer 
than wide, and in having the marginal teeth broad with shallow 
sinus; flowers appearing in spring; flowering stems normally 
leafless; sepals oblong, obtuse, 2.5-3.5 by 1.5 mm.” 
Stoloniferous northern Tiarella cordifolia has the leaves so 
variable in outline that it is quite easy to find specimens from 
Maine, New Hampshire and other northern states and from 
Quebec, New Brunswick and Ontario with leaves inseparable in 
outline and toothing from those of southern plants cited as var. 
austrina Lakela or as var. collina. As defined by Wherry vars. 
austrina and collina have the leaves ‘tending to be longer than 
wide”, while 7. cordifolia var. typica has, according to him, the 
leaves ‘tending to be about as long as wide”. As already stated, 
in New England, where var. typica alone is supposed to grow, 
leaves positively longer than wide are common: 5.3 cm. long, 
3.6 cm. wide; 6 em. long, 4.5 em. wide; 6.5 em. long, 5 cm. wide; 
8 cm. long, 6 cm. wide; 9 cm. long, 7 cm. wide; 9.5 cm. long, 7 cm. 
wide; 10 em. long, 9 cm. wide; 10.3 em. long, 8.3 em. wide; 11 cm. 
long, 7.3 em. wide; 14 em. long, 11 em. wide. When, further- 
more, we find that many specimens of the nonstoloniferous var. 
collina “‘foliis var. austrina similis’? have leaves nearly or quite 
as broad as long (Amelia Co.: 8 em. long by 7.4 em. wide and 11 
em. long by 9.6 em. wide; Brunswick Co: 7 em. long by 6.5 cm. 
wide, 6.5 em. long by 7 cm. wide, 7.3 em. long by 7.5 cm. wide, 
10 em. long by 9.5 em. wide; Durham County, North Carolina: 
9 em. long by 7.8 cm. wide or 9.5 em. long by 8.5 em. wide), it 
is evident that the difference in proportion is of less significance 


1943] Fernald,—Virginian Botanizing Under Restrictions 447 


than assumed. As for size, the maximum given in the original 
description of T. Wherryi is 14 em. long and 9 cm. wide; but 
plenty of New England specimens of T. cordifolia, var. typica 
have leaves 10-14 cm. long and 9-11 cm. broad, while in the 
nonstoloniferous Piedmont 7. cordifolia, var. collina full-grown 
leaves are essentially the same size; and surely the leaf of T. 
Wherryi was originally described as longer than wide, “folia 
laminis elongato-ovatis . . 7-14 em. longis 6-9 em. latis”’ 

and the type-sheet, as aowhi by the memorandum on Dr. 
Lakela’s photograph (her plate 1) of it, was originally taken by 
her to be merely a southern variety of 7. cordifolia. 

As to the bract or bracts on the scape, I am unfortunate in 
not having before me the large series of Dr. Wherry’s material of 
T. Wherry, which is said to differ from his 7. cordifolia, var. 
collina in “having the stems usually bearing one or more leaves”. 
If, as appears from the original account of 7. Wherry?, that species 
is the nonstoloniferous plant of Tennessee and Alabama, it would 
seem that the considerable series of such plants from those states 
which has accumulated in the Gray Herbarium and at the New 
York Botanical Garden should be T. Wherry. The difficulty is, 
that in this series, including plants from Tuscaloosa County and 
other areas cited in the original description ‘“caules floriferi 

. nudi vel 1-3 foliis parvis instructi’’, only 6 scapes show 
biases! leaves (in each case solitary), while 47 scapes are bract- 
less. From the station in Brunswick County, Virginia, of what 
should be naked-seaped T. cordifolia, var. collina, one of Mr. 
Lewis’s specimens has a well developed leaf on the scape, while 
a similar specimen from near Raleigh, North Carolina (Godfrey, 
no. 3424) shows bracted scapes. Incidentally, it is notable that 
Dr. Lakela’s illustration of the type of T. Wherryt shows only 
naked scapes. 

Coming to the reputed difference between the sepals of Tiarella 
cordifolia (var. collina) and T. Wherryi: the former with ‘‘sepals 
obtuse, 2.5 to 3.5 by 1 to 1.5 mm.”’, the latter with “the acutish 
sepals rarely being over 2 mm. long and 0.75 mm. wide’’, it seems 
from the material assembled in the Gray Herbarium that too 
much stress has been given these points. Material from Tusca- 
loosa County, Alabama, has blunt sepals 3 mm. long and 1.3 mm. 
broad, while 7. Wherryi from Grundy County and from Marion 


448 Rhodora [NOVEMBER 


County, Tennessee, Svenson, nos. 7653 and 10,056 respectively, 
shows the sepals quite obtuse. Desiring a check on my own 
measurements, I asked, without telling him my results, the very 
accurate Dr. Robert C. Foster to measure under his binocular 
some selected specimens. His (and my) measurements give the 
following results, the specimens all being in late anthesis. 7’. 
Wherryi from Cumberland County, Tennessee, C. A. and Una 
F. Weatherby, no. 6249, sepals 2.3-2.6 mm. long; from Marion 
County, Alabama, E. J. Palmer, no. 38,997 (as Heuchera villosa), 
sepals 2.3-2.4 mm. long. T. cordifolia, var. typica from Wilton, 
New Hampshire, May 11, 1886, M. A. C. Livermore, sepals 2-2.2 
mm. long; from Charlemont, Massachusetts, May 15, 1915, 
Churchill & Woodward, sepals 2.4-2.6 mm. long; from Becket, 
Massachusetts, Fernald, no. 15,172 (type of forma parviflora), 
sepals 1.8-2.2 mm. long. Since northern stoloniferous T. cordi- 
folia may have the sepals down to 1.8 mm. long, while southern 
nonstoloniferous 7. Wherryi may have them up to 3 mm. long 
and 1.3 mm. wide (as well as obtuse), I find myself unable to 
recognize a specific difference in the sepals. 

Dr. Wherry finds that from his Piedmont and Coastal Plain 
Tiarella cordifolia, var. collina “The likewise non-stoloniferous 
T. wherryi differs in flowering later”, and it is a very striking 
fact that the type of T. Wherry in young anthesis was collected 
in Polk County, Tennessee, on June 12. The type-station is a 
ravine of Hiwassee Beach, between the Unaka Mountains and 
Beans Mountain, consequently relatively cool and backward. 
If the non-stoloniferous plants of Tennessee and Alabama are, 
as seems to be the case, 7. Wherryi, it becomes important to note 
the flowering period of other specimens. TENNESSEE: Sevier 
County (Gatlinburg, not far from the Great Smoky Mountains), 
April 27, Sharp, in flower; Knoxville, Knox County, May 29, 
Ruth, no. 341, fruit and lingering flowers; Cumberland County, 
May 12, Weatherby & Weatherby, no. 6249, fruit and last flowers; 
Grundy County, alt. 1500 ft., April 27, Svenson, no. 7653, 
flowers; Marion County, May 13, Svenson, no. 10,056, flowers; 
Cheatham County, April 24, 2. J. Palmer, no. 35,506, flowers. 
ALABAMA: Tuscaloosa County, April 19, Harper, no. 3490, 
flowers; Marion County, April 21, EZ. J. Palmer, no. 38,997, 
flowers. When we take the flowering material of the non- 


« 


1943] Fernald,—Virginian Botanizing Under Restrictions 449 


stoloniferous plant from the Blue Ridge east to the Coastal 
Plain which has come to the Gray Herbarium we get the following 
score. Virainta: Amelia County, May 2, J. B. Lewis, no. 87, 
flowers; May 21 Lewis, no. 89, young fruit and lingering flowers; 
Brunswick County, May 4, Lewis, no. 3410, buds, flowers and 
young fruit; Henry County, May 6, Baldwin, no. 242, flowers. 
Nortu Caroutna: Durham County, April 12, Correll, no. 5049, 
flowers; May 4, Godfrey, no. 3809, young fruit with lingering 
flowers; Wake County, April 11, Godfrey, no. 3424, flowers; 
Mecklenburg County, April 26, C. H. Knowlton, buds and young 
flowers. From these facts it can only be surmised that Dr. 
Wherry’s series of later-flowering 7. Wherryi came from cooler 
habitats than much of the material collected by others. 

Tiarella Wherryi was distinguished by Dr. Lakela by the 
“fructus carpellis 5-10 mm. longis . . . abrupte rotundatis 
ad rostellum angustum’’. Here is a real morphological character 
which seems to hold, and all the mature fruit of the Piedmont 
and Coastal Plain plant before me has the short and broad 
round-tipped valves. Northern and typical 7. cordifolia, on the 
other hand, has the fruit often more elongate, 6-12 mm. long, 
the relatively narrow valves more lance-oblong and gradually 
tapering to subacuminate at tip. The stoloniferous typical T. 
cordifolia, furthermore, has the lower fruiting pedicels 7-13 mm. 
long; the lower pedicels of the nonstoloniferous southern series 
are somewhat shorter (6-10 mm. long). In typical 7. cordzfolia 
the tip of the filament (after disarticulation of the anther) is 
relatively blunt; in the nonstoloniferous southern plant the fila- 
ment is more slenderly tapering at summit. ll in all, the short 
and round-tipped fruit and the slight differences in length of 
pedicels and in tip of filament mark a southern nonstoloniferous 
plant, 7. Wherryi, as distinguished from a more northern stolon- 
iferous one, 7. cordifolia. Differentiation of the nonstoloniferous 
plants from east of the Alleghenies from a similar plant, with 
inseparable technical characters but occurring west of the Alle- 
ghenies, seems artificial. It is reminiscent of the artificial 
Separation of like plants as two species, one occurring “east of 
the Continental Divide’’, the other west of it, or the too familiar 
key-characters, “eastern’’ versus “ western”’. 


_ 450 Rhodora |NoVEMBER 


*PHILADELPHUS INODORUS L. CAROLINE County: steep 
wooded bluff by Rappahannock River, northeast of Return, 
Sie Long & Fogg, no. 14,168, spread from cultivation. 
See p. 358. 

uae VIRGINIANA L., var. Exodar: Nutt. Local 
range extended into Sournampton Count : dry woods near 
Johnson’s Millpond north of Sedley, nos. 14, 336 and 14,337, 
strongly sonttantine with Sd aaa H. virginiana growing with it. 

Pyrus Lreconter Rehde . communis L. X pyrifolia 
(Burm.) Nakai). PRINCESS Aen County: large shrubs per- 
sisting in clearing about old house, Cedar Island, no. 12,361. 
Essex County: border of dry woods northeast of Loretto, 
Fernald, Long & Abbe, no. 14,169. SourHampron County: 
border of oat 1 mile south of Branchville, no. 10,277.—Identifi- 
cations kindly supplied by Professor Re hde er, who informs me 
that this hybrid was extensively introduced by Le Conte. 

*MALUS ANGUSTIFOLIA (Ait. Michx., var. PUBERULA Rehder. 


25 feet high, with trunk 6 inches in diameter. Identifications 
kindly supplied by Professor Rehder, who states that the only 

material he had previously seen was from Louisiana and 
Mississippi. 

Although I here enter Malus angustifolia as a species of Malus 
in deference to Professor Rehder who made the identifications, 
the so called GENERIC distinctions between the subgenera or 
sections of Pyrus sensu DeCandolle, Endlicher, Bentham «& 
Hooker, Focke, Engler & Prantl, Gray, Bailey and others are no 
different now than they were when those students of worldwide 
relationships treated them as all belonging in the single genus 
Pyrus. In Rehder’s Manual, ed. 2, the key to genera gives these 
distinctions: 


Styles connate at base: fr. usually apple-shaped, without or 
few Ct INE oe ee oy a ee oe ee 26. Malus. 


Much hangs on the shape of an apple, whether the depressed- 
globose of the old fashioned Russet or the elongate ovoid of @ 
Gilliflower. Similarly, although the obovoid form of most of our 
commonly cultivated pears is familiar, one has to look no farther 
than Hedrick’s Pears of New York to see beautiful life-sized 
plates of ovoid to globose pears: globose-ovoid, suggesting 4 


1943] Fernald,—Virginian Botanizing Under Restrictions 451 


Hubbardston apple, in Pyrus serotina; globose as a cherry in P. 
betulaefolia; depressed-globose, like a conventional apple, in 
“Gansel Seckel”’ and even more depressed in “Idaho”. In fact, 
of the Pears treated in Rehder’s Manual (‘‘fr. usually pear- 
shaped’’) 13 out of the 15 species have the fruit described as 
“globose” “globular” ‘subglobose” or “ovoid’’; while such a 
species as Malus Halliana is described in Rehder’s key as having 
“fr. pyriform’”’. The terms “apple-shaped” and “ pear-shaped”’ 
thus become a bit vague. Since, furthermore, two sections of 
Malus, as accepted by Rehder, have grit-cells, we get down to 
styles connate at base in Malus, free in Pyrus, and another 
character: “Ivs. . . folded or convolute in bud” in Malus, 
“involute in bud” in Pyrus. These, if absolutely constant, are 
much sounder characters than shape of fruit. I have not suf- 
ficient knowledge of the many species to be sure whether these 
characters are absolute. Many wise taxonomists of the past 
and the present have doubted their value as full generic differ- 
ences, when there is so much variation in other characters. It 
is often said that apples and pears do not hybridize; that that is 
a sure test. Therefore, since Sorbus and Aronia do hybridize, 
keep them apart as genera and set up a new genus, X Sorbaronia 
Rehder, for their hybrid offspring, just as the genus X Sorbopyrus 
has been set up for the hybrid of Pyrus communis and Sorbus 
aria. Neither the failure of some groups to cross nor the occur- 
rence of crosses between ordinarily quite distinct genera is the 
final test. So many proven cases of incompatability within 
genera have been noted that one can hardly argue that incom- 
patibility necessarily demonstrates distinctness of genera; and 
when such utterly unrelated genera as Cyperus and Rhynchospora 
have sometimes crossed, producing X Cyperus Weatherbianus 
Fern., we do not merge the two genera. 

If Pyrus, Malus, Sorbus and Aronia are distinct genera are not 
Azalea and Therorhodion equally separable from Rhododendron; 
Polycodium and Cyanococcus from Vaccinium; Amygdalus and 
Cerasus from Prunus? These subgenera have quite as many 
and as sound morphological characters. I am inclined to follow 
DeCandolle, Endlicher, Bentham & Hooker, Focke, Engler & 
Prantl, Gray and Bailey until much stronger cases are presented 
for the generic segregation of Pyrus in its inclusive sense. Mor- 


452 Rhodora [NOVEMBER 


phologically its ee are no more discordant than those of 
Pinus and Querc 

dere aes ible Cory, var. feta woe 
(Ell.) F. L. Freeman in Ruopora, xxxix. 426 (1937). DiIn- 
WIDDIE County: ie open ahead thickets and clearings just 
east of McKenney, no. e ,346. First from northeast of western 
South Carolina. See p. 

*TEPHROSIA aE inn a @,) Pers., var. GLABRA Nutt. ex 
Torr. & Gray, Fl. N. Am 296 (1 838). HANOVER COUNTY: 
dry pine woods south of abland, no. 7460. IsLE or WIGHT 

County: clearing in damp sandy age barrens, south of Zuni, 
~ no. 14,345. finieieiuiton County: dry sa nd of pine barrens 
about 7 miles south of Franklin, “ibe. 7459 and 7461. See p. 369. 

Tephrosia virginiana, var. glabra, originally described from 
Georgia, occurs also in pine barrens or on sand-ridges of South 
Carolina (Clarendon County, Godfrey & Tryon, no. 1021) and in 
New Jersey. Whereas typical and wide-ranging T. virginiana 
has the internodes and leaf-rachises more or less villous, the villi 
either few or very numerous, the lower surfaces of the leaflets 
pilose to sericeous and the legume copiously long-villous, the 
internodes and rachises of var. glabra are at most closely short- 
hispid and often glabrate, the leaflets sparsely strigose to glabrous 
beneath and the legumes closely silky-strigose, not copiously 
villous. In eastern Virginia it seems to be the pine-barren 
extreme. 

Recently some students have indicated the occurrence in the 
Atlantic States (New England and New York to South Carolina) 
of var. holosericea (Nutt.) Torr. & Gray, 1. c., based on T’. holo- 
sericea Nutt. in Journ. Acad. Sci. Phila. vii. 105 (1834) and I find 
many specimens in the Gray Herbarium from these states marked 
over as var. holosericea. So far as I can see they are nearly if not 
quite typical 7. virginiana, the individuals with a few pin-point 
trichomes to be found on the upper surfaces of some leaflets. 
Nuttall’s plant was a more definite variation than this: ‘‘ whole 
plant sericeous, scarcely excepting the upper surface of the leaves. 
Hab. In the plains of Arkansas.’”—Nuttall. Torrey & Gray 
correctly defined it: ‘‘stem and raceme densely villous; leaflets 
very silky-pubescent on both sides”. If we keep to the original 
accounts and restrict var. holosericea to the plants with “‘leaflets 
very silky-pubescent on both sides” (i. e. above as well as be- 


1943] Fernald,—Virginian Botanizing Under Restrictions 453 


neath) that variety assumes an inland range, from Michigan and 
Wisconsin to Texas, and we do not have to oversort the plants 
of the Atlantic States. 


*GLYCYRRHIZA LEPIDOTA Nutt., var. GLuTINosA (Nutt.) S. 
Wats. Brunswick County: fallow oye yee of old Chamblis 
Place, Seward Forest, near Triplett, no 483. A large patch, 
spreading extensively by ions nase he rootstocks, doubtless 
originally cultivated. This is the western extreme w ith glutinous 
ats the stems covered with stout glands and viscid setae. 

ee p 

VICIA CAROLINIANA Walt. Local range extended northward 
to Dinwipprr County: low open pineland, thickets and clearings 
just east of McKenney, no. 

PoLyGaLa SENEGA L., var. LATIFOLIA Torr. & Gray. Bruns- 
wick County: oak- hicko woods back of old Chamblis Place, 
Seward Forest, near Triplett, Fernald & Lewis, ee 14,484; the 
first from the southeastern section of the state. 3 8. 

P. sANGUINEA L. To the few and scattered Paks in south- 
eastern Virginia add a second one in DinwippiE County: low 
open pineland, thickets and clearings just east of McKenney, 
no. 14,355. See p. 36 

*EUPHORBIA COROLLATA L., var. PANICULATA (EIl.) Boiss. 
Henrico County: open thicket, South Richmond, no. 12,711. 
Extension north from North Carolina. 

CER FLORIDANUM (Chapm.) Pax. BruNswick County: very 
abundant in rich woods near Western Bridge, Meherrin River, 
south of Edgerton, Fernald & Lewis, no. 14,487. See p. 379. 

AESCULUS NEGLECTA Lindl., var. PUBESCENS Sargent. 
GREENSVILLE County: in great t abundance in rich woods along 
reaper River, below Emporia, Fernald & Lewis, no. 14,537. 

ee p. 383. 


Previously misidentified from meagre shrubs farther down and 
to the north of the river as Ae. discolor Pursh, which should be 
dropped from the Virginia list. 

HYPERICUM DENTICULATUM Walt., var. OVALIFOLIUM (Britt.) 
Blake. To the few and remote stations add one in SOUTHAMPTON 
County: moist sandy and peaty shore of Whitefield’s Millpond, 
southwest of Corinth, no. 14,366. See p. 373. 

H. serosum L, ‘Range extended inland to BRUNSWICK 
County: wet level opening in flat woods, 114 miles southeast of 
Triplett, August 18, 1942, and swampy old field south of Seward 
Forest, Triplett, August 5, 1943, J. B. Lewis. See pp. 374-376. 

IOLA CUCULLATA Ait. Very local on or near the Coastal 
Plain néelivadnt The following are our only stations. Kine 


454 Rhodora [NovEMBER 


AND QUEEN CouNTy: magnolia swamp about 2 miles northeast 
of St. Stephen's Church, Fernald, Long & Abbe, no. 14,203. 
Surry County: low woods, Cypress eee near Sexton, 
Perel Long & “Abbe, no. 14, 204: See p 

. LANCEOLATA o the few stations end from the coast 
add one in ISLE oF Wate County: muddy and swaley shore, 
Darden’s Pond, southeast of Collosse, no. 14,367. 

V. LANCEOLATA, Var. VITTATA (Greene) Weath. & Grise. saa 
the few recorded stations add o n SOUTHAMPTON CouUNTY: 
moist sandy and peaty shore of Whitefield? s Millpond, satire est 
of Corinth, no. 14,368. See p. 373. 

. sTRIATA Ait. A second Coastal Plain station, this one 
GREENSVILLE County: rich woods along Meherrin River, bloat 
Emporia, Fernald & Lewis, no. 14,538. 3 

RHEXIA VENTRICOSA Fern. & Griscom Rang e extended in- 
land to Dinwippre County: ditch bordering er open pineland 
just east of McKenney, no. 14,369. See p. 366. 

*OQENOTHERA TETRAGO na Ro th, var. i ton (Nutt.) Munz. 
DinwippiE£ County: low open pineland, thickets and clearings 
just east of Miseatieg: no. 14,370. First from north of Wilm- 
ington, North Carolina. See 

PROSERPINACA INTERMEDIA geet To ne recorded Vir- 
ginia station (near Lee Hall) add one in Sournampron County 
moist sandy and peaty shore of Whiteficld’s Millpond, southw est 
of Corinth, no. 14,373. See p. 373. 

*ERYNGIUM PROSTRATUM Nutt. SourHampron Cou 
moist sandy and peaty shore of Whitefield’ s Millpond, eas 
of Corinth, no. 14,375. First from north of South Carolina. 
Identification tentative, the young fruit showing characters not 
“ui matched in E. prostratum. Mature material is needed. 

ee p. 373. 


as seem (Michx.) Fern. (Z. Bebbii (C. & R.) 
Britt.). Brunswick Country: oak-hickory-beach woods back 
of old Chamblis Place, Seward Forest, near Triplett, Fernald & 
Lewis, no. 14,489. First from east of the mountains. See p. 378. 

IUM FLORIDANUM Small. SourHampron County: eoade 
bottomland of Blackwater River, southeast of Ivor, no. 13,710, 
the arching or lopping stems left stranded after submergence and 
proliferating, the proliferous shoots producing simple leaves; 

sandy alluvial bottomland of Three Creek, Adams Grove, no 
id 3376. GREENSVILLE County: wooded haltaitata of Fontaine 
Creek, southeast of Taylor’s ae plants up to 7 feet high, 
no. 11. ,384, distrib. as S. suave See p. 368. 


Sium floridanum, not era when Blake made the com- 
parison, may be S. swave Walt. Whether it is a distinct species 
or a southern variety of the transcontinental northern as well as - 


1943] Fernald,— Virginian Botanizing Under Restrictions 455 


southern plant which now passes as S. suave (S. cicutaefolium) is 
not wholly clear. The common northern and wide-ranging 
plant, which we call S. swave, has (except in the submersed forma 
Carsonit (Durand) Fassett and in the wholly atypical estuarine 
forma fasciculatum Fassett) the erect or ascending stem strongly 
corrugated; S. floridanum has the usually slender and more 
flexuous stem terete, in drying only slightly ribbed. In S. suave 
the stem branches near the summit, usually above the middle; 
the corrugate-angulate branches are strongly ascending; and the 
principal cauline leaves (below the branching) are strongly as- 
cending, with 5-17 firm leaflets 4-15 cm. long. In S. floridanum 
the stem branches usually well below the middle, the terete 
branches are spreading-ascending to nearly horizontal; and the 
larger divergent to horizontally spreading leaves have 3-11 
membranaceous leaflets 2-9 em. long. In S. suave the largest 
(leading) umbel of each plant is 4-11 em. broad, with angulate 
peduncle and 10-25 definitely angled (when dry) rays; in S. 
floridanum the more open fully developed umbels are 1-7 cm. 
broad, on filiform peduncles and with 7-15 filiform rays. In S. 
suave the angled pedicels of the umbellules are stiffly ascending; 
in S. floridanum the filiform pedicels are more curving or arching 
and prolonged. In flower and fruit the two are similar, but the 
petals of S. floridanum are narrower and the filaments and styles 
longer than in S. suave. As stated, when we have fuller material 
S. floridanum may prove to be a geographic variety, rather than 
a true species. The contrasts are here stated, that others may 
watch for this southern plant. As stated also, Walter could have 
had either as the basis of his S. suave. 

Sium suave, forma Carsonii, the submersed and weak state of 
the wide-ranging transcontinental plant, has filmy foliage and 
small mostly infertile umbels. It might seem to some to include 
S. floridanum; but it is clearly demonstrated to be merely an 
ecological phase of S. suave (cicutaefolium). Its ascending 
petioles, its angulate-corrugated umbel-rays (when dry), its 
short and stiff pedicels, broadly rounded petals, short filaments 
and short style clearly show it to be S. suave. 

PyROLA ROTUNDIFOLIA L., var. AMERICANA (Sweet) Fern. 
Brunswick County: wooded bluff along Rattlesnake Creek, 
west of Triplett, Fernald & Lewis, no. 14,538, healthy colony, 


456 Rhodora [NovEMBER 


never flowering. Station at remarkably low altitude for southern 
Virginia. See p. 377 

*VACCINIUM STAMINEUM L., var. INTERIUS (Ashe) Palmer & 
Steyermark (Polycodium interius Ashe). HaNover County: 
dry sterile woods near Polegreen, no. 13,417 (fruit blue with a 
bloom). JAmEs Crry County: woods north of Jamestown Road, 
about 1 mile from Willamsburg, J. T. Baldwin, Jr. no. 206. 

Var. interius, as its name implies, is a characteristic shrub of 
the Interior. The material (except for these sheets from eastern 
Virginia) in the Gray Herbarium is from West Virginia, Ken- 
tucky, Missouri, Arkansas, northern Louisiana, Kansas and 
Oklahoma. It is of course questionable if it is a true geographic 
variety, the characters ordinarily used in this series (genus 
Polycodium) being very plastic or fickle. In eastern Virginia we 
get three of these: typical V. staminewm with the young branch- 
lets pilose, the mature leaves somewhat so on the lower surface or 
the lateral veins, a very common plant; var. interius with young 
branchlets pilose but the leaves glabrous except along the midrib 
beneath; and the common var. neglectum (Small) Deam, with 
young twigs and leaves both quite glabrous. V. caesiwm 
Greene, a low shrub (commonly only 2-5 dm. high) of sandhills 
and pine barrens closely approaches the state in North Carolina, 
and may be watched for with confidence. Its small and broad 
leaves (1.5-5 em. long) and the very large and rounded bracts 
(1-2 em. broad) greatly exceeding the subtended pedicels well 
mark it. The group is by no means properly understood, an 
the varieties (species of Greene, Ashe and Small) may prove to 
be minor forms without good geographic ranges. 

*y;. — ek SourHampron Co ta : dry white sand 


2) 
j=) 
be) 
ie] 
ie) 
° 
= 
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ee 
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a 


fae ys ANS Torr., var. CRINITUM Fern. Surry County: dry 
thicket 1 mile north of Surry Courthouse, no. 10,773, narrow- 
leaved extreme. 


1943] Fernald,—Virginian Botanizing Under Restrictions 457 


Camp has renamed Vaccinium vacillans as originally and 
beautifully described by Torrey, Fl. N. Y. 1: 444 (1843). In its 
place he publishes the very appropriate V. Torreyanum Camp in 
Am. Midl. Nat. xxiii. 177 (1940) = ‘‘ Vaccinium vacillans Torrey, 
Fl. N. Y. 1: 444. 1843 (non exemp. in Mus. Brit.)” and to V. 
pallidum Ait. he refers ‘‘ Vaccinium vacillans Kalm, apud Torrey, 
Fl. N. Y. 1: 444. 1843 (typ. in Mus. Brit. est).” 

Torrey, unfortunately, gave to the familiar shrub so clearly 
described from New York state an old herbarium-name, V. 
vacillans ‘“‘Kalm, mss. in Herb. Banks’’. The latter is worse 
than a nomen nudum. It had never been published even as a 
nomen and, surely, Torrey had not seen it in Banks’s herbarium, 
since he did not go abroad. At best he had hearsay knowledge 
of it. I am, therefore, unable to see that a mere herbarium- 
name, given by Kalm to something which Torrey had not seen 
and which, according to Camp’s note, he wholly misunderstood, 
takes precedence over the material from “the vicinity of New 
York” which Torrey so fully and accurately described. I retain 
the name V. vacillans as of Torrey. Had the name V. vacillans 
been accompanied by a description of Kalm’s material or had it 
been previously published, it would stand for that element. 
Since Torrey, who first took up the name, described only the New 
York element, which Kalm did not have, the name is typified by 
Torrey’s material, the only material described. 

STYRAX AMERICANA Lam. To the relatively few stations in 
Virginia add one in Sussex County: upper border of sandy 
ae Airfield Millpond, southwest of Wakefield, no. 14,383. 


SOT OceWae SESSILIFOLIUM (Walt.) J. F. Gmel. Bruns- 


of Triplett, August 24, 1 Lewis. See pp. 374 and 376 
PoLypREMUM PROCUMBENS It is so often stated that 
Polypremum is an annual (“‘much branched small annual”— 


Gray, Man.; ‘ anger diffuse herb’’—Small, Man.) that the 
correct description of Gray, Synopt. Fl., sho uld be emphasized: 

“much branched from an annual (sometimes almost ligneous) 
root.’’ In sandy clearings of Brunswick County the mature 
plants, late in the autu umn, send out vigorous and  eecailel new 
: sal offshoots, much as in Lechea (see Fernald & Lewis, no. 

AMSONIA TABERNAEMONTANA Walt. Local range extended to 
Brunswick County: rich wooded slope below Western Bridge, 


458 Rhodora [NovVEMBER 


Meherrin River, south of Edgerton, Fernald & Lewis, no. 14,491. 
ee p. 380. 

*ASCLEPIAS LANCEOLATA Walt., var. PAUPERCULA (Michx.) 
Fern., forma flaviflora, floribus flavi is. Norroik County: fresh 
reed-marsh and swale along Northwest River near Northwest, 
June 30, 1942, Fernald & Long, no. 14,390—a single plant asso- 
ciated with the rather svandant typical var. paupercula in which 
the hoods are deep orange to scarlet. Typr in Herb. Gray. 
See p. 368. 


Hell, south of Petersburg, very scarce, no. 12,778. Sussex 

OUNTY: open thickets and clearings near Nottoway River at 
Readjuster Bridge, south of Peanut, no. 12,438. 

Acerates viridiflora is excessively variable in foliage but all the 
material before me from southern New England and southern 
New York southward to the Carolinas, with the exception of 
five numbers from eastern Pennsylvania (Phil. Acad.) and the 
two numbers above cited, has the blunt or abruptly short-tipped 
leaves elliptic, oblong, oval or oblong-obovate and is, I take it, 
true A. viridiflora. Rafinesque’s original description called for a 
‘“‘lanceolated, obtuse’’ leaf, “leaves lanceolated, obtuse, hirsute, 
umbells axillar, bending down, corniculas, without appendices. 
I have found it in several parts of Maryland and Pennsylvania, 
mostly in fields.” In view of the oblong, elliptic or oblong- 
obovate outline of the obtuse leaves in the plant of Pennsylvania 
and Maryland, it is assumed that Rafinesque’s ‘“‘lanceolated”’ was 
used with poetic license. 

In the representation in the Gray Herbarium there is no materi- 
al of this obtuse-leaved plant from the north-central states. It 
occurs in North Carolina (Small implies Florida), Louisiana, 
Texas and New Mexico, northward in the Mississippi drainage 
to Kentucky and Kansas. From southern Ontario to Manitoba 
and Wyoming the usual plant has lanceolate to narrowly ovate 
leaves tapering from near the base to acuminate or prolonged 
acute tips. This is var. lanceolata (Ives) Gray, occurring from 
southern Ontario, Manitoba and Wyoming southward to Louisi- 
ana, Texas and New Mexico, in the South mingling with the 
obtuse-leaved plant. I am quite unable to separate the two 
numbers from Virginia and some from southeastern Pennsylvania 
from characteristic var. lanceolata of the Interior. In the East 


1943] Fernald,—Virginian Botanizing Under Restrictions 459 


they look like mere leaf-variations of one plant. In view of the 
very different broad ranges of the two, however, I am making 
this note, especially with the hope that the plants will be care- 
fully checked elsewhere in the East. 

PHLox nivauis Lodd. (P. Hentzii Nutt.) An ies 
station in GREENSVILLE County: dry sandy woods alon 
es Creek, Round Hill Church, Fernald & Lewis, ie 

4,543. See p. 383 

P. CAROLINA L., var. TRIFLORA (Michx.) Wherry. Another 
station in Dinwippre County: low open pineland, thickets and 
clearings just east of McKenney, no 14,398. See p. 365. 

NEMOPHILA MICROCALYX (Nutt.) Fisch. & Mey. This vernal 
annual, characteristic along the James, is also in gg HE 
County: rich woods along Meherrin River, below Em 
Fernald & Lewis, no. 14,542, growing with the Seeds Phacelia 
dubia and very luxuriant (up to 4.5 dm. high). See p. 383. 

HYDROLEA QUADRIVALVIS Walt. Range extended inland to 
BruNswick County: swampy pond-hole in woods, Rattlesnake 
Creek, west of Triplett, Fernald & Lewis, no. 14,493. See p. 377. 

YOSOTIS VERNA Nutt. Local range extended. James CITy 
County: rich woods and slopes by James River, Grove Landing, 
southeast of Grove, FERNALD, Lane & Adve: no. 14,227. Sussex 
County: fallow field alon Nottoway River at Readjuster 
Bridge, south of Peanut, Fernald, Long & Abbe, no. 14,228. 

SCUTELLARIA SERRATA Andr. BRUNSWICK County: rich 
wooded slope near Western Bridge, Meherrin River, south of 
Edgerton, Feriola & Lewis, no. 14,494. See p. 380. Epling, in 
his American Species of Scutellaria, Univ. Cal. Pub. Bot. xx. 
no. 1, 85 (1942) cites no Virginia specimens from the southern 
counties east of the Blue Ridge. 

*PHYSOSTEGIA aboriginorum, nov. (TAB. 783), planta 
robusta 10. dm. alta basi an surculoso; caule glabro a 
basin 0.8-1 cm. diametro; foliis firmis sessilibus vel imis late 
subpetiolatis choses obtusis vel subacutis undulato-sinuatis, 
primariis 8-11 em. longis 1.7—2.5 em. latis; inflorescentia panicu- 
lata, ramibus lateralibus prolongatis arcuato-adscendentibus; 
spicis subdensis 0.4-2 dm. longis, calycibus fructiferis imbricatis, 
rhachi puberula; calycibus bawtor campanulatis puberulis, tubo 
maturo 5-6 mm. longo, dentibus deltoideis; corollis purpureis 
2-2.5 em. longis ean ampliato 8-10 mm. diametro; filamentis 
ad apicem villosis.—Norfolk County, VirGINIA: sw ale at margin 
of Indian Creek (whence the specific name), northeast of North- 
west, June 30, 1942, Fernald & Long, no. 14, at ad PE in Herb. 
Gray.; ISOTYPE in Herb. Phil. Pes See p. 


Physostegia aboriginorum differs strikingly oe the more fre- 


460 . Rhodora [NOVEMBER 


quent species of southeastern Virginia (of the Blackwater and the 
Nottoway drainage-systems) in its broader-oblong, merely 
undulate-margined, firm sessile leaves, the lower ones narrowed 
merely to dilated bases, the upper ones but slightly reduced in 
size; the panicled spikes densely flowered, with imbricated pairs 
of fruiting calices; the corolla more ampliate; the filaments 
copiously pilose to summit. P. denticulata (Ait.) Britton (our 
PLATE 784, FIG. 1), common in the alluvium of the Nottoway or 
the Blackwater and their tributaries in Southampton, Isle of 
Wight and Nansemond Counties, has the leaves thin and mem- 
branaceous, the lower 4 to 6 pairs oblanceolate or oblong and on 
long slender petioles, the middle and upper ones sessile and long- 
attenuate at tip, the uppermost very small. Its usually solitary, 
slenderly elongate spike is very interrupted, the pairs of flowers 
and fruiting calices remote; the corolla more slender, the fila- 
ments glabrous or only sparsely pilose at tip. 

From Physostegia leptophylla Small the Norfolk County plant 
differs in its much larger leaves in more scattered and more sessile 
pairs, with merely undulate margins, and in its more slender 
corolla, P. leptophylla having the many (up to 16) pairs of more 
prominently dentate leaves mostly narrowed to base and the 
greatly distended corolla with throat 1.5 em. broad at summit. 
From P. veroniciformis Small P. aboriginorum differs in much 
greater size, the leaves not fiddle-shaped, the calyx-lobes short 
and deltoid, instead of elongate and lanceolate, the corolla larger, 
and the long, as well as the short, filaments hairy. 

This segregation of species of the Northwest River and its 
tributary, Great Dismal Swamp, and the North Landing River 
drainage, with small tributaries interlocking with those of the 
Northwest River, both emptying into the head of Currituck 
Sound, is very real. The Blackwater and Nottoway merge at 
the State Line to form the Chowan, which, a few miles farther 
south receives the Meherrin and then empties into Albemarle 
Sound. On the bottomlands or wooded slopes of the Black- 
water, Nottoway or Meherrin on the Coastal Plain of Virginia 
hundreds of species occur which we do not know from the 
bottoms, swamps and marshes of the more eastern drainage 
systems. Conversely the Northwest and North Landing drain- 
age (or that of Back Bay slightly to the east) have scores (many 


1943] Fernald,—Virginian Botanizing Under Restrictions 461 


more yet to be discovered) of plants which we do not know from 
the Blackwater, the Nottoway or the Meherrin drainage (Pinus 
serotina, Limnobium Spongia, Arundinaria gigantea, Andropogon 
virginicus var. glaucopsis, Scleria flaccida, Cladium jamaicense, 
Wisteria frutescens, Ilex vomitoria, I. coriacea, Stewartia Mala- 
chodendron, Ludwigia pilosa, L. alata, Asclepias lanceolata var. 
paupercula, Lobelia elongata, Eupatorium recurvans, E. cunet- 
folium, Aster Elliottii, ete.). Physostegia aboriginorum is the 
latest addition to this series. 

I am using the name Physostegia denticulata in its traditional 
sense. Originally published as Dracocephalum denticulatum by 
Aiton, Hort. Kew. ii. 317 (1789) from Carolina material, its 
diagnosis was good, though very brief: “D. floribus spicatis 
remotis, foliis obovato-lanceolatis superne denticulatis”. Such 
a plant abounds, as stated, on the Coastal Plain bottomlands of 
the Blackwater and the Nottoway, thence south to Florida, 
represented in the Gray Herbarium by material from five 
counties of North Carolina: Hertford, on the Chowan River, and 
Perquimans, draining into Albemarle Sound; thence south to the 
region of Wilmington, and from Georgetown County, South 
Carolina. The general geographic source of Aiton’s material is 
unequivocal. 

In Ruopora, xvii. 134 (1915) Blake, publishing the combina- 
tion Physostegia purpurea (Walt.) Blake, based on Prasium? 
purpureum Walt. Fl. Carol. 166 (1788), identified it with P. 
denticulatum. Walter’s diagnosis was as follows: ‘‘foliis oppositis 
sessilibus lanceolatis dentatis, floribus purpureis, spicis distichis 
terminalibus”; and the very sketchy sketch (our PLATE 785, 
FIG. 1) made by Blake of the type of Prasium? purpureum is 
difficult to match in any material usually placed in Physostegia 
denticulata. Blake’s description (mss. in Gray Herb.) of the 
Specimen says “leaves . . . 3mm. wide . . . Corolla 
would be just 2 em. long if straightened out . . . It agrees 
very well with Drummond 251, e. g. in Brit. Mus.” Now it so 
happens that Drummond’s no. 251 from New Orleans (our 
PLATE 785, FiGs. 2-5) has elongate-linear leaves very strongly 
Suggesting Blake’s sketch of the much crumpled Walter type, 
but it has finely sharp-serrate margins (ric. 5) and is a quite 
characteristic specimen of a wide-ranging species (our PLATE 785, 


462 Rhodora [NOVEMBER 


Fics. 6 and 7) which occurs from Tennessee to Illinois and 
Arkansas, south to Mississippi, Louisiana and Texas, the plants 
variously misidentified as P. denticulata, P. virginiana (L.) 
Benth. (‘‘very slender form’’) and P. speciosa Sweet. YFurther- 
more, a habitally quite similar species, P. intermedia (Nutt.) 
Engelm. & Gray (our PLATE 784, Figs. 2 and 3), with slender and 
interrupted spike and with leaves narrowly lanceolate to linear 
but with flowers relatively small, extends eastward into Kentucky 
and Alabama. This small-flowered species has the leaves den- 
tate, as described by Walter for Prasium? purpureum, not 
acutely serrate as in Drummond, no. 251; but, so far as we know, 
neither of these sessile- and narrow-leaved species is east of the 
mountains. With no such plant as indicated by Blake’s sketch 
(PLATE 785, FIG. 1) now known in Walter’s territory, it is unsafe 
to assert with confidence just what his Prasiwm? purpureum is. 
Until Walter’s type can be rechecked with Drummond’s no. 251 
(with linear, sharp-toothed leaves and slender, interrupted 
spikes) clearly in mind there must remain some doubt regarding 
its identity with Physostegia denticulata. Until his type can be 
reexamined and closely compared with a series including all 
known species of the Carolinas, as well as Drummond, no. 251 
and Physostegia intermedia, it is safest to let the name rest, as it 
did until Blake took it up, as still of doubtful significance. 

I am, therefore, at the risk of possibly publishing a species 
already inadequately described, Aisi 


P. angustifolia, sp. nov. (TAB. 785, G. 2-7), planta erecta 
caule glabro 3-10 dm. alto; foliis pepe pee vel pallidis line- 
aribus vel anguste lanceolatis sessilibus numerosis (16-22 jugis), 


imis Shas vel subacutis, mediis attenuatis valde angusteque 
adpresso-serratis 4-10 cm. longis 3-10 mm. latis, foliis supernis 
valde reductis; spicis solitariis vel paniculatis elongatis ad 3.5 
longis floribus distantibus, rhachi puberulo; calycibus campanu- 
latis 6-10 mm. longis pru ruinoso-puberulis glandulosisque dentibus 
Recto hia ite corollis purpureis ie albescentibus 2.5-3 
longis TENNESSEE: thickets along fence, Lavergne, 
Rutherford. County, August 12, 1922, Goeiin no. 242. MIs- 
SISSIPPI: roadside bank near Egypt, Chickasaw County, May 18, 
33, C. A. & Una F. Weatherby, no. 6318, typE in Herb. Gray. 
Iturnors: Arlington Heights, Cook County, August 25, 1934, 
Benke, no. Sete prairie, South Chicago, en ere 15, 1910, 
ansing, no. 2803 3; Champaign, September 11, 1909, Pease, no. 
12,403; Lasalle County, Huett; Sugar Crock Prairie, Richland 


1943] Fernald,—Virginian Botanizing Under Restrictions 463 


County, September 5, 1914, Robert Ridgway. ARKANSAS: moist 
roadside and ditch- bank, Hevetioale June 13, 1938, Fassett & 
s a ge no. 19,829; low ground, northwestern Arkansas, June, 

arvey, no. 116. Lpweibea Covington, May and June, 
Hale; New Orleans, 1832, Drummond, 2 EXA 
Bejar a Austin, Avril, 1828, Berlandier, no. "166: Amelia, Jefferson 
County, October 5, 1934 , Cory, no. 11 108; Kerrville, Kerr County, 
June 25, 1894, H eller, no. 1906; 8 miles south of Bandera, Bandera 
County, June 5, 1933, Cory, no. 6607. 

Most of the material (including many specimens not cited) 
has been distributed as Physostegia virginiana (L.) Benth., but 
that, as shown by a beautiful photograph of the type before me, 
is the common northeastern plant with relatively thin, lanceolate, 
often copiously and saliently serrate leaves mostly 1-2.5 cm. 
wide, the uppermost much reduced, the flowers in relatively short 
and crowded spikes (in fruit with imbricated calices and rarely 
2 dm. long), the corolla 2-3 (in the type 2.4) em. long. It has 
also been called P. virginiana var. speciosa (Sweet) Gray, but 
that is a variation of P. virginiana with broader and often more 
prominently serrate leaves, the larger blades 2-4 cm. wide and 
less reduced in size toward the summit of the stem. Sweet’s 
original plate shows the corolla 2.5 em. long. 

Relatively small plants of Physostegia virginiana from the 
lower Susquehanna in southeastern Pennsylvania have sometimes 
been called P. denticulata because their leaves have the marginal 
teeth few and short. Similar forms, with low stature, small leaves 
and reduction of teeth, abound about Lake Champlain and in 
the vicinity of Montreal on the St. Lawrence. Others from the 
Same regions and of similar proportions have well developed 
Sharp teeth. They seem to be a reduced state of P. virginiana, 
somewhat atypical but without constant enough characters to 
merit specific recognition. The designation of the individuals 
of this small extreme of tidal shores of the St. Lawrence as a dis- 
tinct species, P. granulosa Fassett in Ruopora, xli. 377 (1939), 
our PLATE 786, FIG@s. 1 and 2, because it is stoloniferous and be- 
cause its calyx is more glandular-puberulent than in some P. 
virginiana, overlooks the fact that carefully collected P. virgini- 
ana has stolons. Witness specimens of Fernald & Long, no. 
14,465 from Hampden, Maine, of Fassett, no. 14,341 (our PLATE 
786, Figs. 3 and 4) from Bingham, Maine (toward 50 miles up- 


464 Rhodora [NovEMBER 


river beyond the limit of tide and with leaves as small as and with 
fewer teeth than in isotypes of P. granulosa—in no. 14,341 some 
leaves with as few as 2-5 low teeth on each side, the type of P. 
granulosa described as with 4-8 but the isotypes in the Gray 
Herbarium showing 5~9 pairs of teeth), of the Faxons and others 
from Lake Champlain (PLATE 786, FIG. 5), of Eames from Trum- 
bull, Connecticut, or of any other botanist who takes pains 
carefully to collect the stolons. The small extreme with small 
leaves having few and low teeth and the pubescence of the calyx 
short and glandular is noteworthy but scarcely a good species. 
I am calling it 

P. viRGINIANA (L.) Benth., var. granulosa (Fassett), stat. nov. 
P. granulosa Fassett in Roopora, xli. 377 (1939). PLATE 786. 

On the St. Lawrence var. granulosa is on areas covered by high 
tide; the lower part, at least, of its range on the Susquehanna is 
tidal shore; on Lake Champlain, where the variety abounds, and 
on the middle and upper Kennebec it is in areas which have had 
no extensive tides since the Champlain Subsidence, at which 
time a fresh arm of the sea entered Lake Champlain and a tongue 
extended well up the Kennebec. Can it be that the variety 
originated on Lake Champlain and the upper Kennebec at that 
period? 

E 783. Dh pinto ABORIGINORUM Fernald: Fic. 1, TYPE, X 1; FIG. 
2, none X 2; Fic. 3, mature calices and rachis, 
PLaTE 7! : Oe ecine (Ait.) Britton: FIG. 1, characteristic plant, 


2 76 f from Monroe Bridge, Nottoway River, Southampton County, Virginia, 
& Long, no. 13, 122. P. INTERMEDIA ages ) oom gelm. & mal bh FIG. > 


FIG. 3, leaf-margin, X 2, from no. 5089 


ATE . PHYSOSTEGIA: FIG L tracing of leaf of type of Pyare 
purpureum Walt., by Blake. PHYSOSTEGIA ANGUSTIFOLIA Fern.: . 2-5, 
Drum : i, x 5/9, said by Bake to match Walter's type of "Prasium? 
perceive (FIG. 1), which is sai Physostegia denticulata (see PL 
784, Fic. 1); Fic. 5, leaf of no. 251, x 2; ly 6, TyPE of PHYsoSTEGIA ANGUSTI- 
FOLIA, x 5/12; ric. 7, portion of leaf, « 2. 

PLATE . P. virGInrana (L .) Benth., var. GRANULOSA mer Fernald: 


Figs. 1 (X 1%) and 2 (x 2), of isotype o of P. granulosa Fassett, from Cap 
Rouge, Quebec, Victorin, no ier Figs. 3 (X 4) and 4 (xX 2) from Kennebec 
Rive, Bi ngham, Maine, Fasselt, n . 14,341; ria. 5, X %, from Lake Cham- 
plain, South Has: Vermont, H ness no. 13,902. 


Stacnys Cumnemanu Small. Surry County: rich alluvial 
woods and thickets back of sand-beach of James River, below 
Sunken Meadow Beach, no. 8446; thicket back of sand-beach of 
Cobham Bay, James River, northwest of Chippokes, no. 12,788. 


194 nald,—Virginian Botanizing Under Restrictions 465 

Both numbers have been erroneously reported in previous 
records as Stachys Nuttallii Shuttlew. The latter species of 
western North Carolina and eastern Tennessee has the sides of 
the stem minutely glandular-puberulent, the sessile or subsessile 
leaves glabrous beneath, the glandular and short-pilose calyx 
with broadly deltoid teeth. Our plant, also of the mountains, 
thence westward to Illinois, is the S. Nuttallii sensu Robinson & 
Fernald in Gray, Man. ed.7. Its stems are glabrous on the sides, 
the membranous leaves slender-petioled and copiously hispid on 
the veins and veinlets beneath, the calyx glandular and minutely 
pilose, with narrow teeth long-subulate at tip. On the lower 
James it is as notable an isolation from the Appalachian Upland 
as if it were really S. Nuttallii. 

*S. TENUIFOLIA Willd. Cuaries Crry County: alluvial 
woods along James River, Harrison Point, no. 9133. SouTu- 
AMPTON County; open sandy alluvial bank of Nottoway River, 
below Cypress Bridge, and wooded bottomland, Cypress Bridge, 
nos. 6370 and 8447. GreENsvILLE County: wooded alluvial 
bottomland of Meherrin River, near Haley’s Bridge, no. 9418. 
See PLATE and map 1. 

Apparently typical Stachys tenuifolia is not definitely recorded 
as Virginian. In his Preliminary Revision of American Stachys 
in Fedde, Repert. Sp. Nov. Beih. Ixxx. 68 (1934), Epling gives 
the range ‘‘from Western New York and Southeastern Pennsyl- 
Vania westward to Michigan and Minnesota, and Southwestward 
through Tennessee and Alabama to Louisiana and Texas”. He 
could have extended it southeastward into Georgia; at least a 
characteristic specimen from northern Georgia, coll. Chas. 
Wright, correctly identified by Asa Gray and validated by Epling 
in 1931, is in the Gray Herbarium. True S. tenuifolia is, obvi- 
ously, chiefly a plant of the Interior; its occurrence on the bottom- 
lands of southeastern Virginia is another instance of a now 
familiar isolation, as is material from river-swamp of Santee 
River in Charleston County, South Carolina, Godfrey & Tryon, 
no. 728. Epling’s only suggestion that S. tenuifolia occurs in 
Virginia is his note that “Specimens collected . . . by 
Killip (no. 6286) near Great Falls, Virginia, show an apparent 
intermixture with S. hispida chiefly in hairiness of the calyces 
and the more subulate calyx teeth’. Such transitions to S. 
hispida are so frequent that I cannot feel that the two are distinct 


466 Rhodora [NOVEMBER 


species, but are rather well pronounced geographic varieties. 
The following number from southeastern Virginia combines too 
intimately the characters of typical S: tenuifolia and those of 
ypical S. hispida: SourHampron County: bushy swales and 
borders of swampy woods near Blackwater River, Cobb’s Wharf, 
no. 10,407. As I understand S. tenuifolia and S. hispida they 
are the two extremes of a rather polymorphous species, charac- 
terized by having the sides of the stem glabrous, the leaves with 
glabrous lower surfaces or with hispidity along the veins and 
veinlets but not on the intermediate tissue, and in the calyx 
being glabrous, or bristly only on the angles, its lance-attenuate 
teeth soon outwardly curving. As a polymorphous relatively 
southern species it is distinguished from the more northern 
(circumboreal) S. palustris L., in which the surfaces of the stem 
are pubescent, the leaves pilose to puberulent beneath, and the 
surfaces of the calyx pilose. My interpretation of S. tenuzfolia 
is shown in the following brief synopsis. 


a. Bracts at base of ~ grt of spike not ciliate; calyx 
——— throughou s glabrous on both faces, the 
ncipal ones w iti y heer ig petioles 0.5- "3 ¢ m. lon 
cat-Biades lanceolate to preserve bd nierohely 
ovate, one fifth to three fifths as broad as 8 long, the 
principal ones 4-10 em. long and 1. . broa 
ORR Oe ee ee - tenia, typical. 
Leat-blades paged oblong, about one sixth as bro 
as long; median ones 1—1.5 dm. long and 1. Aerie 
b Bhat OXUOUR eae oe mere ya et Var. perlonga. 
a. Bracts bristly-ciliate; calyx often bristly on the angles; 
lea on veins beneath, sessile, or the lower 
ones ral i 
me Donkey Tenaiy oblong to oon ovate, ae 
third to three fifths as broad as lon 


‘t0 
em. long, 0.8-2.4 em. broad, mostly strigose above... Var. hispida. 


S. TeENvIFoLIA Willd. Sp. PI. iii. nd (1800). S. glabra Riddell, 
Suppl. Cat. Ohio Pl. 16 (1 et 6). aspera, var. glabra Gray, 
Syn. FI. Am. ii’. 387 (1878). 5 cincinnatensis Ktze. Rev. 
Gia: ii. 531 (1891).—Rich bottomlands, shores, low woods and 
meadows, New York to Minnesota, south to southeastern South 
Carolina, northern Georgia, Tennessee, northern Louisiana and 
eastern Texas. PLATE 787 and M 

. perlonga, var. nov. (TAB. 788, ‘FIG. 1 et 2), var. typicae 
simillima; foliis anguste oblongis,. mediis 1-1.5 dm. longis 
1.6-2 em. latis; spica flexuosa. VirGINIA: bottomland sco 


S si 

se) Ns ; se 
S } 
ry S 


EXPLANATION 


Existing kc felds. 


alter DA. 
LS, 
pear 
Boe PLLI ST OC: 
Atlontic cosstel plaire & PachTe areas 


Spur oe 


exposed wring he QUATERNARY. 
Probable fara afd Wve 


since close of the TERTIARY 


Probable Jana area aoa! 


since close of Ihe 


or unknown. 


since close of the 


, 


ses § » N ( [ntesartg Ly Mi Uf r: NP a 3 | 
Asnoawmeso, Ae Tee fae 
ae. e' 
RES SI. De 
' sop ye } ‘ 
Va ere 


es aes ae as wn 
° Te ar 
‘eo ee DL igs Sy, 
YEN 
\ 4 — ey 


Ke! 


° 


; 
eC 
? 


a6 Pr 


—Virginian Botanizing Under Restrictions 


? 


i 
<< 


——— ce ® 
= ae ane. 


enero a 
~_— 
tee 
aieo 
oy. 
645 
E-F-| 
me a 
i J 
a 
Po >) 
rh 

> op 

~& 

is 

ae 

eg 

— 

p 

z, 

a} 

= 

QD 

-~o-, 

_ 
— 


TENUIFOLIA, var. PLATYPHYLLA; 
PALUSTRIS. var. HOMOTRICHA (S. ambigua sensu 


Range of (1) Sracnys TENUIFOLIA; (2) S. TENUIFOLIA, 
HYSSOPIFOLIA 


8. 


468 Rhodora [NovVEMBER 


along Meherrin River northeast of Gaskins, Greensville County, 
June 13, 1941 (foliage), August 8, 1941 (flowers and young fruit), 
nos. 13, 128 and 13, a (TYPE in Herb. Gray.; IsoTyPE in Herb. 
Phil. Acad.). Map 
*Var. eta, var. nov. (TAB. 789), caule ad angulos 
retrorse setoso; foliis primariis late oblongis vel anguste ovatis . 
cm. latis supra glabris vel sparse strigosis 
subtus ad venas hispidis Met glabris—Low woods, rich shores 
and meadows, southern Quebec to Minnesota, south to eastern 
South Carolina, wreater North Carolina, Indiana and Illinois. 
The ieee oe ted from a large series, are characteristic. 
UEBEC: sur les rivages du Fleuve St.-Laurent, St.-Francois, 
lle d’ Sige’ Be: juillet 1935, Vietorin et al., no. 43 ,661; rivages 
Fl. St.-Laur Berthier en Bas, 3 aofit 1925, Rousseau, no. 
21,610; Valleyfald, Co. Beauharnois, 6 septembre 1930, Victorin, 
Rolland, no. 33,953; rivage, Ile Plate, Longueuil, 28 aofit 1928, 
oe no. 28,272. Marne: East Livermore, July 5, 1909, 
Kate Furbish; by small pond, Portland, June 20, 1909, Fernald, 
no. 2158. New Hampsuire: bank of Connecticut River, Strat- 
ford, September 3, 1909, Pease, no. 17,705. VERMONT: ‘pebbly 
lake-shore, Highgate Springs, July 19, 1938, ed no. 26,746; 
sandy shore of Connecticut River, Norwich, July 1 , 3910, E. F. 
Williams ae in Herb. Gray.); sandy bank, Burlington, July > 
1914, C. H. Knowlton; pebbly shore, Brattlebor 1894, Gro 
MASSACHUSETTS: Sudbury, July 19, 1886, W. Raion bank oF 
Neponset River between upper and lower falls, Milton, August 
29, 1853, Wm. Boott; ea te Wellesley, July 11, 1896, Hunne- 
well; Middleboro, August nag J. Murdoch, jr.; edge of 
thicket, Northampton, Ju : 1932, Hunnewell, no. 12,496; 
bank of Connecticut River. Springfield, July 13, 1914, Luman 
Andrews. Connecticut: bank of Connecticut River, East 
Windsor, July 13, 1902, Bissell; South Meadow, Hartford, July 
7, 1908, H. 8. Clark. New York: shores of Black Lake, Morris- 
town, August 15, 1914, Phsipe no. 814; Mohawk flats east of 
Utica, July 18, 190 00, Habere r, no. 709; Sylvan Beach, Oneida 
County, July 13 , 1916, House; bank of Hudson River, near Cedar 
Hill, south of Albany, July 17, 1910, Burnham; wet gravelly soil, 
Ulysses, July 1, 1913, #. L. Palmer, n o. 1039; moist thicket near 
Cold Spring, Alleghany State Park, ‘July 30 30, 1926, Alexander & 
ouse, no. 12,783. New Jersry: marsh, ’Rosenkraus Ferry, 
Delaware River, Sussex County, July 4, 1918, E. B. Bartram; 
wet ground, Kaign’s Point, Camden, June 21, 1922, Meredith; 
border of tidal marsh along Delaware River, north of Penns 
rove, August 9, 1935, Fogg, no. 9309. PENNSYLVANIA: Sellers- 
ville, July 18, 1923, Freie: ine County, July 2, 1923, S. S 
Dickey; Presque Isle, June. 29, 1922, Dickey. VirGINIA: swampy 
woods near Nottoway River at Readjuster Bridge, south of 


1943] Fernald,—Virginian Botanizing Under Restrictions 469 


Peanut, Sussex County, June 18, 1941, Remald & Long, no. 
13,129; ‘along branch, south of Meherrin River, south of Western 
Bridge, Brunswick County, J. B. Lewis, nos. 2800 and 3538. 
SoutH Carona: Woodstock, 1833, Gibbes in Herb. N. Y. Bot. 
Gard. Ontario: Ottawa, August 8, 1894, J. Macoun, no. cen 
Wingham, July 24, 1894, Fk, Morton; Chippewa, July 5, 858, 
Wm. Boott. MurcHiGaN: near river, Flint, July 21, 1909, Sher 
Agricultural College, July 25, 1896, C. F. Wheeler. Pete Tro 
Geauga County, July 21, 1904, R. J. Webb, no. 716; Sautiusky: 
August 1, 1894, M oseley; Rocky River, near Cleveland, July 13, 
1896, Greenman, no. 1403; bares July 19, 1885, Werner, no. 
342; Sugar Grove, Fairfield Coun y, August 6, 1892, Werner, no. 
43: ; hear Cincinnati, July 9, 1888, CoG, Lloyd. WISCONSIN: 
Fort Wing, July 10, 1897, Lé8: Cheney, no. ao ie ae 
Stony Island, Cook County, June 30, 1914 H. Smith, 
5974; damp woods, Peoria, July, 1903, F.E. MeDonald: banks of 
Mississippi, near Oqu awka, July 18, 1872, Patterson. MINNE- 
SOTA: ane City, August 13, site W. H. Manning. Map 3. 


ambigua sensu Robinson & Peruild. in Bae “Man. ed. 7: 701 
(1908), not S. ambigua (Gray) Britton—Meadows, swamps, 
low woods and shores, southern New Hampshire, eastern Massa- 
chusetts and Rhode Island, eastern New York, south to eastern 


ee 12, 1896, ver. St. Joh tae pond-shore, Walpole 
Hunnewell, no. Mb Middleboro, Richard Murdoch, no. 933; 
Taunton, ymour, no. 4187. RHODE ISLAND: Providence, 


ew Jersey: along streamlet 1 mile northeast of Delanco, 
Burlington County, Long, no. 6354; fresh marshes along Wading 
River, New Gretna, Burlington County, Long, no. 10,574; rich 
bog along railroad, Cold Spring, Cape nie County, August 29, 
1922, Fogg. PENNSYLV. Anta: Tinicum, Delaware County, 
August, 1873, Chas. Schaeffer; meadow near Red Clay Gieck: 


470 Rhodora {NOVEMBER 


Kennett Square, yeas County, June 28, 1924, Mary A. 
Williams. Mary : King’s Creek, J. A. Ho Imes, no. 98. 
VIRGINIA: wet Ri incoous thickets and ditches, Rosemont, 
Princess Anne County, Fernald & Long, no. 5017: fresh reed- 
marsh eer swale along Northwest River near Northwest, Norfolk 
County, F. & L., no. 14,005; alluvial woods, along Notto oway 
River at Readjuster Bridge, south of Peanut, Sussex County, 
F. & L., no. 12,459; bushy swales and borders of swampy woods 
near Blackwater River, Cobb’s Wharf, Southampton County, 
F.& no. 10,407 (transition to typical S. tenuifolia) ; alluvial 
woods by Fontaine Creek, west of Dahlia, Greensville Coun sr 
F. & L., no. 14,399; pea aty and argillaceous clearing about 4 
miles southeast of Emporia, vr. & L., nos. 8445 and 8834; bottom- 
land woods along Me herrin River northeast of Gaskins, Greens- 
ville County, F. & L., no. 13,434. SourH CaRo.ina: wet ditch, 
thicket, 13 ‘thiles east of Walterboro, ee County, Wiegand 
& Manning, no. 2793; border of swamp, 12 miles north of 
Georgetown, Godfrey & Tryon, no. 1067 . MIcHIGAN: near river, 
uffalo, Lansing, no. 3262. Inprana: low place in oe 
about 1 mile southeast of Woodland, St. hence County, 
no. 50,459; damp prairie, Roby, Lansing, n 2542; Feankfort, 
Photon 27, 1886, H.A. Young. Wisconsin: St. Cink Falls, — 
190 og. F. Baker. I.t1nots: open field, South Chicago, H. 
Smith, no: 5684. PrLare 790 and map 4. See p. 

Stachys hispida Pursh, typified by Epling by a Clayton speci- 
men from Virginia (presumably in the eastern section), is, ac- 
cording to the description and as indicated by Epling’s note on & 
sheet in the Gray Herbarium, this narrow-leaved extreme. Var. 
hispida has, so far as material before me shows, the broken range 
of many Coastal Plain types, great concentration in eastern 
Massachusetts and adjacent Rhode Island, absence from Con- 
necticut and much, if not all, of New York, reappearance farther 
south, and with an area centering on Lake Michigan and running 
into Wisconsin. In Massachusetts, New Jersey, eastern Penn- 
sylvania and eastern Virginia it tends to grow in more acid 
habitats than var. platyphylla, the latter preferring the sweeter 
and richer soils, as does typical S. tenuzfolia. 

The confusion of Stachys hispida with S. aspera Michaux arose 
through unfamiliarity with the latter. A photograph of the 
Michaux type now before me shows it to be, as identified by 
Epling, the southern plant which Gray (1878) described from 
Georgia, Kentucky and Illinois as S. hyssopifolia Michx., var- 


1943] Fernald,—Virginian Botanizing Under Restrictions 471 


ambigua. It was soon called S. ambigua (Gray) Britton (1894) 
but, on account of the earlier European S. ambigua Sm. (1810); 
the ndme was changed to S. Grayana House (1921). S. aspera, 
described from ‘in campestribus Carolinae”, is a plant of sands 
and prairies from northern Florida to eastern Missouri, north to 
southern New Jersey, southeastern Pennsylvania, Kentucky, 
Illinois and Iowa; also adventive as a field-weed in Somerset 
County, Maine. It is, as Gray considered it, very close to S. 
hyssopifolia Michx., having the firm and scarcely veiny leaves 
almost of the latter but mostly broader, their lowest lateral nerves 
prolonged very close to the margin of the blade. From glabrous 
S. hyssopifolia it is distinguished only by the stems more or less 
retrorse-hispid on the angles, leaves sometimes broader and 
more regularly depressed-serrate, upper bracts often broadly- 
lanceolate to ovate (instead of linear or narrowly lanceolate), 
calyx often minutely pilose (instead of quite glabrous) and fre- 
quently setose on the angles. Gray’s disposition of it, as S. hys- 
sopifolia, var. ambigua, seems wholly justified. In the Carolinas 
and northeastward there are altogether too many transitions to 
Support its separation as a species. 

In his Revision Epling, 1. c. 65, 66, including with the narrow- 
leaved S. tenuifolia, var. hispida, as above defined (pLatE 790), 
the generally more inland var. platyphylla (pLATE 789) as a 
species, S. hispida, was “inclined to believe S. hispida to have 
been derived from S. ambigua, probably after intermixture with 
S. hyssopifolia”. He identified as S. ambigua Sm. Engl. Bot. 
xxx. pl. 2089 (1810) the very pilose and narrow-leaved extreme 
of S. palustris L. which I had described as S. palustris, var. 
homotricha Fernald in Ruopora, x. 85 (1918), which he also iden- 
tified with the similarly long-pilose S. velutina Schwein. (1824), 
not Willd. (1813) and S. arenicola Britton (1901). Just why S. 
ambigua sensu Epling (PLATE 793) is identified with S. ambigua 
Sm., the European plant (pLate 791), is not clear. He was 
advised of the error but did not wholly admit it, saying, p. 65, 
“Since the above was written I have had the privilege of dis- 
cussing this form with Prof. Géte Turesson who is of the 
opinion that I have mis-interpreted the European hybrid S. 
ambigua. This, according to him is a rare and sterile plant. 
Until I can review the whole matter I am accordingly at as [a] 


472 Rhodora [NOVEMBER 


loss. Suffice it to say that the form here described as S. ambigua 
is scarcely to be distinguished from plants commonly referred to 
that species of European botanists’ 

Now, Stachys palustris, var. hombieteke (PLATE 793), i. e. S. 
ambigua sensu Epling, is a stiff and normally simple-stemmed 
plant with the sides and angles of the stem and the rachis densely 
and retrorsely long-pilose or villous (F1a@s. 2 and 3), the lanceolate 
to lance-linear leaves sessile and heavily pubescent beneath 
(ric. 2), the calyx long-hirsute (Frias. 2-5), the outside of the 
corolla strongly pilose. It occurs from Maine and southern 
Quebec to Wisconsin, south to Connecticut, southern New York, 
Ohio and Illinois (map 5). S. ambigua Sm., on the other hand, 
is a generally recognized sterile hybrid of typical European S. 
palustris (the latter only naturalized in Newfoundland, eastern 
Canada, New England and New York) and the endemic European 
S. sylvatica L., a plant with long-petioled cordate-ovate leaves. 
As shown by Smith’s original plate and description and by many 
European specimens (our PLATE 791) it is a tall and loosely 
branched plant with the stem (F1G. 2) minutely strigose-pilose, 
the principal leaves definitely petioled, the divergent oblong- 
ovate blades minutely pilose beneath, the calyx short-pilose 
(ric. 3) and the corolla only finely pilose. If it is inseparable 
from the indigenous North American S. palustris, var. homo- 
tricha (PLATE 793), it is time to give up. 

As to Epling’s belief that Stachys hispida in his sense, 7. e. both 
S. tenuifolia var. platyphylla (PLATE 789 and Map 3) and var. 
hispida (PLATE 790 and Map 4), was derived from his ‘S. ambi- 
gua”’ (taking it for the American plant cited by him, our PLATE 
793, and S. hyssopifolia Michx. (PLATE 788, Fics. 3-5 and mapP 6) 

one may be pardoned a gasp of surprise. S. palustris, var. 
homotricha (S. ambigua sensu Epling) is shown in pLaTE 793, S. 
tenuifolia, vars. hispida and platyphylla (S. hispida, inclusive, of 
Epling) in pLates 790 and 789. Together the latter have 4 
wide continental dispersal (Maps 3 and 4), while S. palustris, 
var. homotricha is northern and within the northern range of the 
former. S. hyssopifolia (PLATE 788, Fics. 3-5), is primarily 4 
plant of the Atlantic Coastal Plain, but with the typical isolation 
at the head of Lake Michigan (map 6). It is strictly glabrous, 
with linear to narrowly oblong blunt to subacute, firm, pale, 


1943] Fernald,—Virginian Botanizing Under Restrictions 473 


entire leaves, so firm that, when fresh, only the midrib shows 
beneath, but, by transmitted light its leaves show the lower 
lateral veins following forward within 0.3-1 mm. from the margin, 
and its subglobose nutlets (ria. 5) are only 1.5-1.8 mm. long. S. 
hispida and S. ambigua sensu Epling are both pubescent. Their 
leaves are from narrowly lanceolate to oblong-ovate and usually 
pubescent, veiny, with the lower pair of laterals following for- 
ward from 3-10 mm. within the copiously toothed margin, and 
their tips are acuminate; the nutlets (PLATE 792, Fic. 4) of S. 
palustris are ellipsoid-obovoid and 1.8-2.2 mm. long, those of 
S. tenuifolia, including S. hispida, similar and 2.1-2.7 mm. long 
(PLATE 788, Fic. 5). Just how, by crossing the narrow-, blunt- 
and firm-leaved, chiefly Coastal Plain S. hyssopifolia (PLATE 788, 
FIGS. 3-5) with the narrow-leaved continental and northern and 
by Epling misidentified S. “ambigua” (PLATE 793), with leaves 
1-3 cm. wide, one gets the wide-ranging S. hispida sensu Epling 
(pLatEs 789 and 790), with no trace of the pubescence of S. 
“ambigua’”’ on stem, leaves and calyx and with leaves up to 6 
cm. broad, is beyond my understanding. Is this modern phy- 
logeny? 

Our VARIETIES OF STACHYS PALUSTRIS (PLATE 792-794)— 
During this study of Stachys it has been necessary to unravel our 
variations of S. palustris. This is a northern, circumboreal 
species with two pronounced subspecies, each with several geo- 
graphic varieties. Typical Eurasian S. palustris L. has the calyx 
(PLATE 792, Fic. 3) closely viscid-pilose with many gland-tipped 
hairs mixed with short glandless ones, the latter rarely more than 
1 mm. long. It and three of the varieties recognized in Europe 
are naturalized about towns and settlements, often about ports, 
in various parts of Newfoundland, Quebec, eastern Ontario, the 
Maritime Provinces, New England and New York. The native 
North American plants differ in having the calyx long-hirsute as 
well as short-pilose, the shorter pubescence largely hidden by 
long whitish and glandless setiform trichomes mostly 1.5-3 mm. 
long. This is subsp. pilosa (Nutt.) Epling, based on S. pilosa 
Nutt. in Journ. Acad. Phil. vii. 48 (1834). The native subsp. 
pilosa consists, in the Gray’s Manual range, of five rather well 
defined varieties. These I am distinguishing as follows. 


474 Rhodora [NOVEMBER 


a. Leaves ste e on very short and inconspicuous thick 
tioles.... 
b. Principal leaves oblong to ee or oval, blunt or 
merely subacute, three tenths to one half as broad as 
long, 3.5-12 cm. long, 1.5-5 cm. broad; chiefly western... Var. pilosa. 
b. Principal leaves sassy to broadly lanceolate or nar- 
oblong, acuminate, one seventh to one third as 
heaad as long, 2-13 cm. long, 0. 43. cm. eo chiefly 
eastern and northern.... 
c. Pubescence of sides of iia m many times shorter than 
that of ss or minute and with long hairs of 


r. 
Angles of stem a h few or — heen: os sides mi- 


cian tely spabeiraleihd to glabrous beneath; plant 

of feat: Lakes revion. 3)... oe a ee Var. macrocalyz. 
c. Absence fa sides of stem very dense and lon 

not readily dinktneiiahed from that of 
the an "5 Ba PE an ery eee ye ihe ec Var. homotricha. 

a. sor bh slender-petioled, the petioles of the principal ones 

1-1.4 em. long, the oblong or lance-ovate acuminate 

i gi mostly 6-12 em. long and seg em oer angles 
stem iene relatively few or no long hairs....... Var. phanero poda 


S. PALUSTRIS L., var. pilosa (Nutt. » sa. nov. S. pilosa Nutt. 
in Journ. Acad. Phil. vii. 48 (1834). Subsp. pilosa (Nutt.) 
Epling in Fedde, Repert. Beih. Ixxx. 63 (1934). S. scopulorum 

9 ns 


reene, Pittonia, iii. 342 (1901).—Shores, stream-margins, 
British Colombia to Arizona, to Ontario, 

Wisconsin, pom Nebraska, and adventive along railroads an 
in wa astward to Ne gland (several eee | 


preteens 
Var. NIPIGONENSIS Jennings in Journ. Wash. Acad. Sci. x. 459 
(1920) Saar es grounds, Quebec to Alaska, south to western New 
England, New York, Ohio, Michigan, Illinois, Iowa, New 
Mexico and Arizona 
Var. MACROCALYX Jennings, 1. c.458 (1920).—Shores, Thunder 
Bay District, Ontario, to northern Wisconsin and northern 
die age tea 
MOTRICHA Fernald in Ruopora, x. 85 (1908). 8. 


Willd. (1813). S. = erm Britt. Man. 792 (190 1). S. palus- 
tris, var. arenicola (Britt.) Farw. in Am. Mi “ aa xl. 82 (1928). 
S. Schweinitzii Rydb. in Brittonia, i. 95 (19 S. homotricha 
(Fern.) Rydb. 1. ¢. (1931). S. pustulosa Redo. l. c. (1931).— 


1943] Iernald,—Virginian Botanizing Under Restrictions 475 


Shores, meadows and fields, Maine and southern Quebec to 
Wi isconsin, south eo Connecticut, New York, Ohio and Illinois. 
PLATE 793, MAP 5 

h ropoda, var. nov., Weatherby in — — 794) 
foliis petiole petiolis (foliorum principalium) 1- m. longis, 
laminis o longis vel lanceolato-ovatis 6-12 cm. atte nae em. 
latis; setis in angulis caulis ex comparatione paucis vel carentibus. 
Onto: Port hare Ottawa County, July ys 1894, Moseley. 
Wisconsin: Spooner, July 20, 1911, J. J. Dav Iowa: river- 
Ha. cay me 31, 1925, B.O. Wolden, tr 1118, TYPE, in 

er 


PLATE ig hginr: 1 and 2, 8. Ege: var. PERLONGA Fernald: Fic. 1 


Long, no. pi ai ig and aga calyx, X 4, from south of Yorktown, 
Fernald, 0. 5016; Fic. 5, nutlet, X 10, from Sandwich, Massa- 
chusetts, rm es 23. 1929, niga nal. ; 

PuiatE 789 UI . PLATYPHYLLA Fernald: ric. 1, TYPE, X_ 3; 


x 10, from Alburg, bois ng July 16, 1939, C. H. Knowlton; ria. 3, lower 


surface of leaf, < 10, and Fic. 4, rachis, bracts and calices, X 4, from same 
place as FIG. 2; FIG. 5. chive < 10, from Chaumont, New Yor k, Fernald, 
Wiegand & Eames, Rea 14,4 29, 

LATE TENUIFOLIA, Var. HISPIDA (Pursh) Fernald: Fie. 1, plant, 
x 1%, from Rosemont, Prine e County, Virginia, a Long, no 


x 
calices, X 4, from Sharon, Massachusetts, July 12, 1896, such plants from 
Sharon specially noted in Gray Herb., by Epling, as 7 Purs sh’s type 
,. Puate 791. 8. ampicua Sm.: Fic. 1, summit of plant, X 4, from ‘Borgholm, 
Oland, Sweden, July 20, 1886, Areskug; Fie. 2, portion of stem, X 10, fro 
m 


Piate 792. §. paLusTRIS L.: FIG. 1, portion of stem, X 10, fro m Bouillon, 


X 10, and ria. 3, = whorls of flowers, X 4, from same collection; Fr. 4, 
nutlet, > 10, from ballast-filling, Vos Gaspé County, Quebec, Collins, 
Fernald & Pease (Pease, no. 506214). 

Piate 793. S. PALUSTRIS, var. HOMOTRICHA Fernald (S. ambigua sensu 
Epling, not Sm.): ne. 1, plant of type, X 34; FIG. 2, lower bracts and calyx, 

4, from TYPE; FIG. 3, stem and lower surface of leal base, < 10, from TYPE; 

bad "4, narro elena sd extreme (S. arenicola Britton), x %, from Copper 
Harbor, Michigan, Farwell, no. 281); F1G. 5, two caiiaed, x 4, from no. 281. 

Puate 794, §S. PALUSTRIS, var. PHA ee Weatherby: as ‘, TYPE, 
x 4; ain 2, stem, X 4; Fic. 3, calyx, X 1 


PYCNANTHEMUM VERTICILLATUM (Michx.) Pers. A second 
Coastal Plain mei, this in NorrotK County: damp old 
clearings and thicke ts, eastern side of Great Dismal Swamp, 
north of Wallon nos. 13,736 and 14,007. 


476 Rhodora [NOVEMBER 


P. PYCNANTHEMOIDES (Leavenw.) Fern., var. VIRIDIFOLIUM 
Fern. A second Coastal Plain station, this in SouTHAMPTON 
County: border of dry woods northeast of Adams Grove, no. 
14,398. 

VERBENA OFFICINALIS L. To the relatively few stations add 
one in SourHamMpTton County: roadside bordering Womble’s 
(Wade’s) Millpond, north of Baffle, no. 14,395. 

HYSALIS MONTICOLA C. Mohr. Range extended northeast- 
ward to Iste or Wicut County: sandy wooded slope by Black- 
water River, west of Blackwater School, no. 14,400, unusually 
large, up to 3.5 dm. high. See p. 370. 

*LINARIA CANADENSIS (L.) Dumort., forma albina, f. nov., 
corollis albidis.—Virerinta: burned spot, white sand of dry pine 
barrens, south of Lee’s Mill, Isle of Wight County, April 21, 
1942, Fernald, Long & Abbe, no. 14,230 (rypE in Herb. Gray.). 

GRATIOLA VIRGINIANA L., var. AESTUARIORUM Pennell. 
Range extended inland to Brunswick County: open mud by 
Triplett Pond, Seward Forest, near Triplett, Fernald & Lewis, 
no. 14,497. See p. 380. 

UrricuLaRIA PURPUREA Walt. Local range extended inland 
to Sussex County: Harold’s (or Harris) Millpond, southeast of 
Waverly, no. 14,408. See p. 371. 

U. myriata Walt. Local range extended inland to Sussex 
County: back-water pool by Nottoway River at Readjuster 
Bridge, south of Peanut, Fernald, Long & Abbe, no. 14,231. 

TA, var. MINOR Chapm. ‘To the very few recorded 
stations add the following. SouTHampron County: quiet pool 
in cypress swamp bordering Womble’s (Wade’s) Millpond, north 
of Baffle, no. 14,406. Sussex County: Harold’s (or Harris) 
Millpond, southeast of Waverly, no. 14,407. See pp. 371 and 372. 

U. vircatuLa Barnhart. To the few recorded stations add 
one in SOUTHAMPTON County: on a decaying log in mossy margin 
of Cephalanthus-pool near Johnson’s Millpond, north of Sedley, 
no. 14,411. See p. 371. 

OLDENLANDIA Boscit (DC.) Chapm. To the only two re- 
corded Virginia stations, in southern Southampton County, add 
the following. Sussex County: sandy-muddy shore, Brittle’s 
Millpond, west of Wakefield, no. 14,417; sandy beach, Airfield 
Millpond, southwest of Wakefield, no. 14,419. SouTHAMPTON 
County: sandy shore of Johnson’s Millpond, 114 miles north of 
Sedley, no. 14,416. Iste or Wieut Cov : muddy and swaley 
shore of Darden’s Pond, southeast of Collosse, no. 14,418. 
pp. 371 and 372. . 

OBELIA SIPHILITICA L., forma laevicalyx, f. nov., foliis 
_glabris; caulibus vix pubescentibus; calycibus corollisque gla- 
ris.—Isle of Wight County, VirGrnia: seeping argillaceous and 
calcareous bluffs along Burwell’s Bay, James River, below Rush- 


1943] Fernald,—Virginian Botanizing Under Restrictions 477 


mere (Fergusson’s Wharf), August 27 and 29, 1940, Fernald & 
Long, no. 12,834, October 10, 1941, Fernald & Long, no. 14,030 
(TYPE in Herb. Gray; 1soryPE in Herb. Phil. Acad.). 

My attention was directed to this extremely smooth form by 
Dr. Wray M. Bowden of the Central Experimental Farm at 
Ottawa, Ontario. I had supplied Dr. Bowden with seeds from 
no. 14,030 and from some other species in the autumn of 1941. 
On October 14, 1942, I received from him a letter from which 
the following passages are extracted: 


_ “Just a_note to ask if you could examine your specimen of Lobelia 
stphilitica L., Fernald and Long, no. 14,030 . . You sent seed from 
this last year and the plants have grown well. The row was quite uniform 
but not at all like any of the numerous collections of L. siphilitica which 
ave. The stems and leaves are glabrous. The upper leafy bracts in 
the flower-spike are much longer and pointed, and the calyx is not nearly 
as pubescent—in fact only the free ends of the lobes are pubescent. e 
W was quite uniform both in respect to these characters and to colour 
of corolla which was light blue. . . The corolla is quite glabrous also.” 


The original specimens have the leaves, calyx and corolla 
quite glabrous, but the stem is remotely hispid. Other collec- 
tions from calcareous bluffs of the James are more pubescent. 
The plant below Rushmere seems to me best treated as a forma, 
typical L. siphilitica having the stem and one or both faces of 
the leaves variously pubescent, the calyx always more or less so 
and the tips of the corolla-lobes bearded. Among these more 
typical plants I find some with quite as prolonged bract-tips as 
in the Rushmere plant. 

L. GLANDULIFERA (Gray) Small. Local range extended into 
Brunswick County: sphagnous thicket, ‘‘Ram Hole Swamp”, 
Pewee Forest, near Triplett, Fernald & Lewis, no. 14,500. See 
p. : 

*EUPATORIUM ROTUNDIFOLIUM L., var. cordigerum, var. nov., 
planta robusta ad 1.3 m. alta caule densissime albido-piloso- 
tomentuloso; foliis mediis late ovatis subacutis sessilibus valde 
cordatis 5-8.5 cm. longis 4-7 cm. latis subaequaliter dentatis 
subtus dense pilosis.—VirGINIA: upper border of fresh reed- 
marsh and swale along Northwest River near Northwest, Norfolk 
County, June 30, 1942, Fernald & Long, no. 14,426 (ryPxH in 
Herb. Gray.; isotype in Herb. Phil. Acad.); mossy thicket by 
Cephalanthus-pool near Johnson’s Millpond, north of Sedley, 


October 14, 1942, Fernald, no. 14,502. Nort C : 
side near Windsor, Bertie County, October 15, 1938, Godfrey, no. 
7007. See pp. 369 and 371. 


478 Rhodora [NOVEMBER 


Var. cordigerum is a very extreme variation in the polymorphic 
Eupatorium rotundifolium. In size of plant and of foliage it 
suggests the wide-ranging var. ovatum (Bigel.) Torr. (EL. pubescens 
Muhl.) but its dense pubescence is more like that of typical 
small-leaved FE. rotundifolium. In its strongly cordate, instead 
of gradually rounded, leaf-bases it is very striking, the cordate 
bases of the middle and upper leaves reaching nearly around the 
stem. 

LIATRIS GRAMINIFOLIA Willd., var. Smauui (Britt.) Fern. & 
Grise. Local range extended into Sussex County: damp san 
pine and oak woods south of Stony Creek, no. 11,455. 

BOLTONIA CAROLINIANA (Walt.) Fern. Range extended inland 
to Brunswick County: bottomland woods above Western 
Bridge, Meherrin River, south of Edgerton, Fernald & Lewis, no. 
14,512, rays pink. See p. 379 

*ERIGERON strigosus Muhl., forma piscomprus (Robbins) 
Fern. NANSEMOND CounTY: open ground along highway, 
pg orde border of Great Dismal Swamp, east of Magnolia, no. 

*SoLipaGo SALIcInA Ell. Brunswick County: open bog near 
Seward Forest Headquarters, September 14, 1940, J. B. Lewis; 
boggy thicket, ‘‘Ramhole” Swamp, Se ward Forest, Triplett, 
Fernald & Lewis, no. 14,506.—Extension northward from Wake 
County, North Carolina; most specimens (from North and South 
Carolina and west to Texas) previously confused w ith the north- 
ern and upland calcicolous S. patula. See p 

S. salicina is a very definite southern species of acid bogs of 
the inner Coastal Plain and outer Piedmont, at once distin- 
guished from S. patula by its narrower and more crenate leaves, 
the cauline ones 20-45 (in S. patula 12-23) and rapidly reduced 
in size, the more open panicle with more elongate and glabrescent 
branches floriferous at tip, the long-pedicelled heads borne strictly 
along the upper sides of the branches (in S. patula the shorter- 
pedicelled heads tending to be in short racemes subspirally borne 
along the white-pilose panicle-branches). 

GNAPHALIUM SPATHULATUM A LATER Homonym (PLATE 799, 
the Typr, X 24).—It seems to have been overlooked that (na- 
phalium spathulatum Lam. Encyel. ii. 758 (1786) is antedated by 
G. spatulatum Burm. f. Fl. Cap. Prodr. 25 (1768) which was the 
binomial based directly on the well described Elychrysum foliis 
oblongis circa caulem auritis & tomentosis of Breyn, Prodr. 29, t- 


1943] Fernald,—Virginian Botanizing Under Restrictions 479 


xvii. fig. 3 (1739), this being referred by Index Kewensis to 
Helichrysum crispum D. Don, not H. spatulatum Moench. The 
plant which generally passes as Gnaphalium spathulatum cannot, 
therefore, maintain that name. The exact identity of Lamarck’s 
type is yet to be worked out. The habit-photograph of it before 
me is good as such, but until the technical characters can be 
closely studied it would be unsafe to base a new name upon it. 
The group of annuals and biennials to which it belongs is most 
complex, a series of ruderal and campestrian plants of tropical 
and warm-temperate regions of which the characters reside in 
habit, shape and indument of leaves, indument of stem, indument 
of involucre, shape of phyllaries, etc. The plant usually passing 
as G. spathulatum is wide-spread in South America, and in North 
America from Florida to Texas, north to southeastern Pennsyl- 
vania and southern California. Superficially it strongly re- 
sembles G. platense Cabrera, Compuestas Bonaerenses, 167, fig. 
45 (1941), but that species, as clearly illustrated by Cabrera, has 
the leaves and stems copiously glandular, the elongate pluri- 
cellular glands of the stem divergent, and the inner phyllaries 
slender-tipped. Cabrera refers G. spathulatum Lam. to G. ameri- 
canum Willd., but certainly the plant of our southern states is 
not the West Indian G. americanum, which is very near G. 
purpureum LL. Until the whole complex group can be most 
thoroughly studied, with intimate knowledge of the actual types 
and their technical characters, it would be quite unwise to rename 
the poorly understood G. spathulatum Lam. (originally a garden- 
plant). Iam, therefore, at the risk of publishing a name which 
eventually may have to be discarded, proposing a new name based 
upon an unequivocal type. 
GNAPHALIUM (§ GAMOCHAETA) peregrinum, sp. nov. (TAB. 
795), bienne vel annuum; caulibus subsimplicibus vel plerumque 
ramosis 1-6 dm. altis pi Pasay var foliis caulinis anguste spa- 
thulato-obovatis vel late oblanceolatis apice rotundatis, imis 2-7 
em. longis 0.8-2.5 em. latis, subtus villoso-lanatis supra deinde 
Vicidisioen iba: capitulis stramineis | pai atis glomerulis sub- 
distantibus ‘gene interruptam 1-2 em. crassam formantibus; 
involucris 3-4 altis basi valde enn phyllariis interiori- 
bus oblongis oa “epathulatis obtusis —Roadsides, fallow fields 
and waste places, Florida to Texas ‘and southern California, 
north, locally, to southeastern asiastiania: eastern South 


480 Rhodora [NOVEMBER 


America. Type: in burnt-over pine-scrub oak sandhills, north 
ag eo Pineville, Sales Parish, oe one July 30, 1938, 
DLS. & : B. rell, oo 9937 (in Herb. Gra 

G. peered Pii ay ar. HELLERI (Britt) Blake. To the 
stations much farther east badd one in BRUNSWICK County: sandy 
pine woods, Seward Forest, near Triplett, Fernald & Lewis, no. 
14,514. 

G. OBTUSIFOLIUM, var. MICRADENIUM Weath. Range extended 
inland to Brunswick County: dry woods near Rattlesnake 
Creek, west of Triplett, Fernald & Lewis, no. 14,513. See p. 380. 


(To be continued) 


VIRGINIAN BOTANIZING UNDER RESTRICTIONS 
M. L. FERNALD 
(Continued from page 480) 


Some VARIETIES AND Species or HELENIUM (PLATES 796- 
799).—As a genus Helenium, in its present sense, was first called 
Helenia L. Hort. Cliff. 418 (1737). It consisted of a single 
species, Helenia foliis decurrentibus, based in part on the plant of 
Clifford’s garden (our PLATE 797, FIG. 1), in part upon many 
references to previously described plants of European gardens 
called Heleniastrum, Chrysanthemum or Aster, the latter not 
actually before Linnaeus. When the genus was taken up at the 
nomenclaturally critical date, 1753, and assigned a binomial it 
was as Helenium autumnale L. Sp. Pl. ii. 886 (1753). The cita- 
tions then given by Linnaeus indicate the two plants somewhat 
generally found at that time in European gardens, and, although 
in N. Am. Fl. xxxvi?, 127 (1915) Rydberg says that the type 
came from Canada, Linnaeus himself in 1753 was not so explicit, 
he then saying “Habitat in America septentrionali”’. In Hortus 
Cliffortianus he had said “in Florida & Canada”’. 

The plant of Clifford’s garden (our PLATE 797, FIG. 1) had 
narrowly lanceolate and long-acuminate, nearly entire leaves, 
few long-peduncled heads, the mature disk 1.8+ cm. broad, the 
narrow ligules strongly narrowed at base, 2 cm. long and only 
2.5-3.3 mm. broad at the sharply trifid apex. Except. that its 


486 Rhodora [DECEMBER 


leaves are rather more spreading and its branching looser, it 
closely matches the plant of tidal shores of the St. Lawrence 
(r1G. 2), there found from near Montreal to below Quebec, and 
extending westward through much of the St. Lawrence basin and 
far to the southwest. Rydberg’s inference was correct, appar- 
ently, that this plant originated in Canada. The second plant 
cultivated in Europe was the extreme which Linnaeus preserved 
in his own herbarium (presumably from the Upsala garden) and 
which, according to the late Dr. B. Daydon Jackson, was before 
him in preparing Species Plantarum, ed. 1. This (our PLATE 
796, FIG. 1) is the coarser plant of our Northeast, with more 
elliptic and broader, toothed leaves, and large heads, with the 
mature disk of the leading head nearly 2.2 cm. broad, the flat 
and cuneate ligules 2.2-2.3 em. long and 8.3-11.7 mm. broad at 
the bluntly lobed summit. Just such a plant (Fic. 2), with 
ligules up to 2.5 em. long, occurs from western New England to 
Minnesota, south to New Jersey, upland to western North 
Carolina, Kentucky and Missouri. Helenium autumnale of 1753 
consisted of both extremes and there is logic in interpreting 
either of them as the type. In a nomenclature beginning with 
1753 (not 1737 and earlier), however, the plant in the Linnean 
Herbarium, the specimen Linnaeus definitely had before him in 
preparing Species Plantarum, is here selected. ‘It is the extreme 
of the species which most students of the past have treated as 
H. autumnale, as will later be noted. 

Both of them being in European gardens, whence they were 
described by Ventenat in 1720, Miller, with no reference to 
Linnaeus, took his cues directly from Ventenat, not from the 
Linnean Herbarium. Miller called Heleniwm autumnale the 
narrow-leaved extreme, the “ Bastard Sun-flower with a longer and 
narrower leaf,” the Heleniastrum folio longiore & angustiore of 
Ventenat; and the plant with broader leaves and larger heads, the 
Heleniastrum folio breviore & latiore of Ventenat, Miller called He- 
lenium latifolium Mill. Gard. Dict. ed. 8 (1768), the ‘‘Bastard Sun- 
flower with a broader and shorter leaf’. Among botanists of the 
period, however, Miller was essentially alone in his interpretation. 
Lamarck, Smith, Schkuhr and other botanists, far more precise 
than the very impressionistic Miller, took the broad-leaved and 
large-headed plant (our PLATE 796) of the Linnean Herbarium as 


1943] Fernald,—Virginian Botanizing Under Restrictions 487 


true H. autumnale. Thus Lamarck described as a new species, 
H. canaliculatum Lam. in Journ. d’Hist. Nat. ii. 213, t. 35 (1793) 
the narrow-leaved plant of European gardens which Ventenat 
had called Heleniastrum folio longiore & angustiore and which 
Miller took up as H. autwmnale. Lamarck’s description and 
life-sized illustration (our PLATE 797, FIG. 3) indicated unbranched 
plants with solitary large heads with ligules 1.6 em. long and only 
3-5 mm. broad at the sharply toothed summit: ‘‘Semi-flosculi 
15-18, foeminei, distincti, patentes: ligulis concavo-canaliculatis, 
apice tridentatis”. Lamarck went on “L’Helenie canaliculée 
différe évidemment de l’Helenie d’automne (dict. vol. 3, pl. 81) 

par son aspect, son port, et principalement par le 
canarias de ses demi-fleurons, qui ne sont point élargis, planes, 
et réfléchis comme dans ces [referring to both H. autumnale and 
H. quadridentatum] espéces’”’; and in summary he contrasted his 
H. canaliculatum “ semi-flosculis canaliculatis’’ and H. autumnale 
“semi-flosculis planis reflexis”’. Somewhat later Lamarck, IIl. 
iv. t. 688, fig. 1 (1797), illustrated his conception of H. autumnale, 
the plant of the Linnean Herbarium. 

Helenium canaliculatum Lam. (portion of his illustration, 
shown in our PLATE 797, FIG. 3) was unquestionably the same as 
the plant (our pLaTE 797, FIG. 1) of the Clifford garden. Except 
for the more spreading leaves, a natural response to the ameliora- 
tions of cultivation, H. canaliculatum is the extreme (F1G. 2) of 
H. autumnale which characterizes the tidal shores of the St. 
Lawrence from above Montreal to below Quebec, thence west- 
ward to Minnesota and Iowa. That region was the source of 
many plants early carried to Europe and a very inexact and 
obviously conventionalized plate of it with similarly elongate 
leaves, too many heads for the wild plant, and tremendously 
multiplied but characteristic rays was described and illustrated 
as Aster luteus alatus by Cornut, Canadensium Plantarum, 62, 
63 (1635). The life-sized head and reduced leaves from Cornut’s 
plate are here reproduced as PLATE 797, rias.4 and 5. It will be 
noted that its disk is 2 cm. broad, its ligules about 1.5 em. long 
and 4-5 mm. broad at the sharply toothed summit. It is obvi- 
ously of the same variety as the plant of the Clifford garden and 
as that of the Paris garden described and illustrated by Lamarck 
as his H. canaliculatum. 


488 Rhodora [DECEMBER 


When Sir James Edward Smith purchased and eventually 
established in London the herbarium of Linnaeus he intimately 
knew the collection and in his many articles in Rees’ Cyclopedia 
he clearly designated what he understood, from studying the 
collection and literature, as the types of many Linnean species. 
Thus, in Rees, Cycl. xvii. (1819), he took as H. autuwmnale the 
plant of the Linnean Herbarium with “ Leaves elliptic-lanceolate, 

more or less deeply serrated. Flowers large’’, etc., 
wills for the oa of European gardens with ‘Leaves linear- 
lanceolate, entire’, etc., the H. autumnale sensu Miller, he pro- 
posed the new name H. longifolium Sm. In treating as typical 
Helenium autumnale the plant which Lamarck had already (1793) 
selected and which he soon (1797) illustrated I am not only 
following the very careful Lamarck and Smith but am trying to 
keep the concept which has prevailed among most botanists until 
Rydberg. Witness, besides the illustration by Lamarck and the 
treatment by Smith, the following illustrations of H. autwmnale: 
Schkuhr, Bot. Handb. ed. 2, t. 250a (1808); Barton, Fl. N. Am. 
i. t. 26 (1821); Raf. Med. FI. i. t. 47 (1828); Hook. in Bot. Mag. 
Ivii. t. 2999 (1830); Meehan, Fl. and Fern U. S. ii. t. 29 (1879); 
and so on to Britton & Brown, Ill. FI. iii. 450 (1898) and House, 
Wild Fl. N. Y. ii. t. 261 (1918). By so doing less violence is 
done well established and fully justified usage than by taking up 
for the plant of Linnaeus’s own herbarium, labeled by him H. 
autumnale, the name H. latifolium Mill., as is done by Rydberg 
in the N. Am. Fl. xxxiv?. 127 (1915). In there taking up as H. 
autumnale the H. longifolium Sm. Rydberg showed that he did 
not understand the latter. If he had understood it he would 
not have referred H. canaliculatum Lam. (clearly H. autumnale 
sensu Miller and H. longifolium Sm.) without question to the 
synonymy of H. latifolium; and if he had understood H. autumnale 
of Herb. L. and of Lamarck, Schkuhr, Smith, Barton et al. (1. 
latifolium Mill.) he would not have contented himself with 
“Jigules 10-15 mm. long”. In the type they are 2.3 cm. long. 

Besides typical Helenium autumnale (H. latifolium Mill.) and 
the rather local var. canaliculatum (Lam.) Torr. & Gray, Fl. N. 
Am. ii. 284 (1842), a combination not cited in the North American 
Flora, we have a wide-ranging southern extreme of the species 
(PLATE 798), the plant with small and narrow rays which was 


1943] Fernald,—Virginian Botanizing Under Restrictions 489 


described as a species, H. parviflorum Nutt. Although JH. 
parviflorum in most extreme variations is very different from true 
H. autumnale, their ranges overlap and it is most difficult to 
find any stable morphological character to keep them apart. 
The chief claim of H. parviflorum to recognition is its extension 
far south of the other varieties, a plant with the flowering and 
fruiting disk only 0.8-1.5 em. broad (as opposed to 1.6—2.3 cm. 
in true H. autumnale), the ligules 3-12 mm. long and only 3-7 
mm. broad (as opposed to 1.6—-2.5 em. long and 7-12 mm. broad 
in true H. autumnale). For six days (a full week) I have boiled 
out disk-florets and ligules, seeking in the H. autumnale complex 
some stable characters of disk-corollas, pappus and achenes, and, 
although there is extreme diversity in the number, shape and 
length of the pales of the pappus, sometimes short, sometimes 
much longer and more slender, sometimes in several lengths, and 
while the disk-corollas may have very short or almost obsolete 
tubes, or the tubes may be prolonged and the throat more 
campanulate, I have been quite incapable of making these di- 
vergencies fall into clear geographic patterns or associate them- 
selves with other characters. After concentrated study for a 
week I get back to the decision of Torrey & Gray who wrote: 
“Some of our varieties are possibly species; but they accord in 
every thing but the pappus, which also presents every inter- 
mediate gradation. The var. @. [the western var. grandiflorum 
(Nutt.) Torr. & Gray] is the only state we have seen from Oregon, 
Saskatchewan, &c.: but a state with a nearly similar pappus is 
common in New York; while other specimens, otherwise undis- 
tinguishable, present a reduced and merely acute pappus’’.— 
Torr. & Gray, l. ec. (1842). Torrey & Gray had a score or so of 
specimens; the ten score of specimens from which I have boiled 
(to clarify and straighten) the florets have failed to bring any 
more clarification than they could get from their hand-full of 
material. 

Excluding the plants from west of the “Manual range”’, the 
great bulk of inclusive Helenium autumnale seems to me to fall 
into three recognizable geographic varieties: 

HELENIUM a eee us Sp. Pl. ii. om (1753) in part (as to 
plant of Herb. L. bea he identification of Linnaeus); Lam 
in Journ. d’Hist. Nat. ii. i AS (1798) and tll iv. t. 688, in 1 (1797); 


490 Rhodora [DECEMBER 


Schkuhr, Bot. Handb. ed. 2, t. 250a (1808); Smith in Rees, Cycl. 
xvii (1819); Barton, Fl. N. Am. i. t. 26 (182 1); Raf. Med. Fi. i. 
t. 47 Gee Hook. in Bot. Mag. Ivii. t. 2999 (1830); oe i 
Gray, Fl. N. Am. ii. 384 (1842); Meehan, Fl. and Ferns U 
t. 29 (1879): Britton & Brown Fl. ii. 450 (1898); cies 
Wild FL NW-Y i. t. 261 tee yab sue tae Mill. Gard. Dict. 
ed. 8, Helenium no. 2 (1768); Fl. xxxiv?. 127 
(1915) as to plant, not as to aban oilatae a elenia autumnalis 
ill, Hort. Kew. 6 (1769). Heleniwm pullin Willd. Enum. 
jay se 60 (1813). H. altissimum Link ex Rydberg 1. c. 126 
(1915).—Stem 0.5-1.5 m. high, wing-angled, simple or branching, 
corymbose-branched at summit; leaves elliptic, oblong or lance- 
olate, ese membranaceous, =e larger coarsely dentate 
ones 0.5-1. long a —5.5 em. broad; heads few-many, 
peduncled, in savtiinal eprviab: fully raewctoped disk 1.6—2.3 cm. 
broad; ligules flat, cuneate, deep yellow, soon reflexed, 1.6—2.5 
cm. long, 7-12 mm. broad at the Ghani toothed or lobed ‘summit; 
pappus much shorter than to two thirds as long as disk-corolla.— 
Rich thickets, meadows and shores, western New England to 
Minnesota, south to New Jersey, western North Carolina, Ken- 
tucky and "Missouri, northward passing into var. canaliculatum, 
southward into var. ‘parviflo orum. A few fairly typical specimens 
are the following. Massacnusetts: Stockbridge, September 5, 
1912, R. Hoffmann; Sheffield, September 30, 1919, Churchill. 


— 
— 
ane 
. 


no. 
10,815; Sylvan Beach, Oneida Co., House, nos. 3843 and 8705; 
Dryden, MacDaniels & Eames, no. 1313. New Jersey: Fields- 
abe Long, no. 18, 221; southwest of Harrisonville, Long, no. 
45, 269. PENNSYLVANIA: Washington Crossing, Bucks County, 
September 25, 1923, Meredith: Wissahickon Ravine, Philadelphia 
eee September 24, 1924, He enry A. Lang; Sayre, Barbour, no. 


536. ONTARIO: teen eg Krotkov, 7908 (mixed with var. 
canaliculatum). Onto: Garrettsville, pres County, R. J. 
, no. 400. Itirors: Bloomington, August, 1886, B. L. 
Robinson. Our PLATE 796. 
Var. CANALICULATUM (Lam.) Torr. & Gray, Fl. N. Am. ii. 284 
citation H. autumnale L. Sp. Pl. ii. 886 (1753) in part, as to some 


_ Am xiv’. 
127 (19 15). H. ince Ait. Hort. Kew. iii. 227 (1789). H. 
canaliculatum Lam. in Journ. d’Hist. Nat. ii. 213, t. 35 (1793). 
Helenia decurrens Moench, Meth. 589 (1794), both as to descrip- 
tion and citation only of the Morison figure which is of most 


1943] Fernald,—Virginian Botanizing Under Restrictions 491 


characteristic H. canaliculatum, and obviously copied from Cor- 
nut. Helenium longifolium Sm. in Rees, Cyel. xvii, Heleniwm 
no. 2 (1819). ZH. phates pubescens (Ait.) Britton in Mem. 
Torr. Bot. Cl. v. 339 (1894).—Stem relatively slender, 3.7-9 dm. 
high with 1-many heads; leaves linear to lanceolate, Moe or 
shallowly ddentiontite, firm, subrigid, the sea ones 3- 

long and 3-18 mm. broad; mature disk 1 . broad, ligule 
strongly narrowed and ofte en convolute at s Hit —2 cm. lon 

2-7 mm. broad at the shallowly and sharply to bluntly ached 


and ec rst Some characteristic specimens are as follows: 
QuEBEC: Cap Pains Victorin, no. 28,177; St. Vallier, Svenson & 
Fassett, no. 3022; Berthier, Montmagny County, Fernald & 
Pease, no. 25,324; Longueuil, Victorin, no. 18,376; phe a 

n- 


074. 

cord, September 9, 1934, R. J. Eaton (adventive, leaves more 
toothed than in most material). Nerw Yorx: Canton, or 
no. 1002; Springport, Hames & Wiegand, no. 11,007. Ont 
Tobermory, Krotkov, no. 7908 (mixed ery typical H. cadumuaiies: 
Birch Island, Lake "Huron, Macoun, no. 26,508 (distributed as 
H. huronense Britton, an ‘herbar arin ines ‘unfortunately pub- 
lished in syno nymy, therefore with no status, by Rydberg in N. 
Am. Fl. xxxiv?. 127 (1915)). Outo: Windham ee Portage 
County, R. J. Webb, no. 5554. Wisconsin: St. Cro x Falls 


pete City , August 15, 1883, W. H. Manan Lowa: 


1894, Fink. NeprasKa: Beaver ‘Creek, Holt es Fred Clements, 
no. 2858: South Fork of Dismal River, Rydbero, no. 1690. Mrs- 
SOURI: Dodson, Bush, no. 4148, dist rib. as H - montanum; Colum- 
bia, Boone Co. , Lisle Jeffrey, no. 261. Kansas: Chautauqua 
Co.  Aitehchels no. 737. Texas: banks of ae about Fried- 
richsburg, Lindheimer, no. 477. Arizona: canon at Fort 

' Passing westward, from Saskatchewan and North Dakota to eastern Washington, 


For note on H. autumnale pubescens (Ait.) Britton see p. 492. It is probable that 
Aiton had the two Ghihe which were in general cultivation in Europe, not the western 
var. montanum to which Rydberg doubtfully refers var. pubescens. 

‘ Farther west, in British Columbia, Washington and Oregon, becoming the large- 
ow = 

Var. GRaNpvIFLORUM (Nutt.) Torr. & Gray, Fl. N. Am. ii. 384 (1842); Gray, 

Fl. N. eS, i?, 349 (1884); independently published by Howell, Fl. Nw. Am en 
(1900). H. grandifiorum Nutt. 1. c. (1841) 


492 Rhodora [DECEMBER 


Huachuca, Lemmon, no. 2776. PiatTe 797.—Many specimens 
transitional to the preceding so the following, and on the Plains 
of the West to var. monta 


Aiton, Hort. Kew. iti. 227 (1789), had a H. pubescens, de- 
scribed simply “H. foliis pubescentibus”. Torrey & Gray and, 
later, Gray treated it as synonymous with the inclusive H. 
autumnale, of which the typical variety is subglabrous, in both 
places wrongly citing the page as “287’’. In Mem. Torr. Bot. 
Cl. v. 339 (1894) we get the unexplained combination H. autum- 
nale pubescens Britton, based upon H. pubescens Ait. of page 
“287”. Had the author of the new trinomial looked up Aiton, 
instead of copying the error of Torrey & Gray and of Gray, he 
would have noted that H. pubescens was on p. 227. I have not 
seen Aiton’s type; neither had Britton. From the date (1789) it 
is probably var. canaliculatum. 


*Var. parviflorum (Nutt.) stat. nov. H. parviflorum Nutt. in 
Trans. Am. Phil. Soc. ser. 2, vii. 384 (1841).—Leaves mem- 
branaceous or submembranaceous, elliptic, oblong, lanceolate or 
oblanceolate, coarsely toothed to subentire, the larger oe 
0.5-1.5 dm. long and 0.7—3.5 em. broad; mature disk 0.8—-1.5 ¢ 
broad; ligules 3-12 mm. long and 3-7 mm. broad, often fiche 
late below. —Bottomlands, swamps and other low grounds, 
Florida to Arkansas, north to Connecticut, New York, Pennsyl- 
vania, Kentucky, Illinois and southeastern Iowa, thoroughly 
typical and isolated southward, passing sig oe into the two 
preceding northward. The fo lowin ng, from a very large series 
“i an e are characteristic. Gomnkancer: Bia ague, August 
oodward; Selden Cove, Lyme, August 29, 1901, 
Bissell: ‘Stratford, August 16, 1895, Eames. New York: old 
specimens without designated. locality, Torr. & are: Fl. New 
Jersey: east of Silverton, Ocean County, Fogg, no. 4901 (nar- 
rowest-leaved extreme); Mays Picea "August "28, 1910, W. 
Stone; Cold Spring, Fogg, no. 235; Treasure Island, ‘Hunterdon 
ounty, Long, no. 38,291. Palen Delaware County, 
18et Canby, called by Gray H. parviflorum, with the later com- 
t “Passes for but ??”, the material _{relatively hineiente) 
alas by Rydbeeg as H. longifolium Sm., i. e. H. autumnale 
var. canaliculatum; Dillerville Svenp. “Lancaster eat, 
September 18, 1889, Heller. DeELAWARE: Bayville, Sussex 
County, Fogg, no. 11 27 1; north of Leipsic, Kent County, Fogg, 
no. 6230. West Vrirornta: Roland P ark, Cabell County, 
Gilbert, no. 546. Virernta: Hunting Creek, southwest of Alex- 
andria, G. H. Shull, no. 188; Bedford County, September, 1873, 
A. H. Curtiss; Jamestown Island, James City County, Ferna 


1943] Fernald,—Virginian Botanizing Under Restrictions 493 


Long, no. 11,204; Kittewan Creek, Weyanoke, Charles City 
County, Fernald & Long, no. 11,483; near Creeds, Princess Anne 
County, Fernald, Long & Fogg, no. 5139; Northwest, Norfolk 
County, Fernald & Long, no. 


8: 
north of Burgaw, Pender County, Godfrey, no. 6537; near Sun- 
ford, Lee County, Godfrey, no. 6911; Winston-Salem, Godfrey, 
no. 6094; Sylva, Jackson County, September 18, 1897, E. E. 
Magee. Sour Carourna: near Georgetown, Godfrey & Tryon, 
no. 8240. Groraia: Macon, September 3, 1883, J. D. Smith; 
Warsaw, Long County, Eyles, no. 7655. Forma: St. Mark’s, 
Wakulla County, Nash, no. 2534; Apalachicola, Biltmore Herb., 
no. 515a. Kentucky: near Wasioto, Bell County, Kearney, no. 
506. TrnnessEE: Knoxville, W. A. Anderson, no. 638; Town- 
send, September 26, 1936, Luther Burns. ALABAMA: without 
stated locality, Nuttall (possible type or isotype of H. parviflorum 
—bearing Nuttall’s asterisk and ent 


Earle, no. 89. Mussissrprt: Taylorville, Tracy, no. 8529; Jack- 
son, September 6, 1885, J. D. Smith. Iniinois: Champaign, 
Pease, no. 12,389; Peoria, September, 1904, F. E. MacDonald. 
Owa: Keosauqua, Pammel & Reis, no. 465. Mussourt: Bakers- 


In the plastic group with disk globose or globose-ovoid and 
usually brown or purplish, the receptacle ovoid and the ray- 
flowers sterile or neutral, Rydberg treats both Helenium nudi- 
florum Nutt. and H. polyphyllum Small as good species. Only 
by recognizing the smallest specimens with narrowest cauline 
Wings as the former, the largest ones with broadest wings as the 
latter, ignoring the large series of transitional specimens can one 
do so. The intermediate pile is altogether too large; but in our 
Southeast there are two extremes, native of low woods and 
Swamps and evidently local endemics, which seem to be really 


494 Rhodora [DECEMBER 


well defined species. In typical campestrian and aggressively 
weedy H. nudiflorum (including H. polyphyllum) the pales of 
the pappus are narrowly lanceolate to lance-ovate and 0.5-1.6 
mm. long, the pales attenuate to narrow tips or scabrous awns, 
the firm and usually scabridulous cauline leaves linear to narrowly 
lanceolate or oblanceolate. In southeastern South Carolina, 
however, there is a plant of this series, occurring in low woods or 
in wooded bottomlands, with membranaceous and quite smooth 
cauline leaves lance- to elliptic-oblong, the pappus of blunt and 
awnless round or oval small pales. This I am calling 
dfreyi, sp. nov. (TaB. 799, Fic. 1 et 2), H. nudifloro habitu 
simillima; foliis caulinis membranaceis laevibus lanceolato-vel 
elliptico-oblongis; achaeniis glabris verrucosis, pappi paleis al- 
bidis suborbicularibus vel ovalibus muticis 0. 4-1. 4 mm. longis.— 
OUTH CAROLINA: creek-bottom through rich Caf anie wnicds, 4 
a west of Georgetown, June 27, 1989, Godfrey & Tryon, no 
124; cleared strip along logging railroad through floodplain- 
forest, Santee River, 3 miles northeast of Pineville, Berkeley 
county, July 14, 1939, Godfrey & Tryon, no. 586 (TYPE in Herb. 
ay 


This localized species is named for Ropert KENNETH GODFREY 
whose extensive field-work is greatly clarifying our knowledge 
of Carolina plants. 

Another relative of Helenitum nudiflorum is concentrated in the 
northern half of Florida. In this plant the relatively small 
heads are either rayless or with well developed ligules, but the 
ovate pappus-pales (1-1.6 mm. long) are rounded at tip and 
terminated by a very long, filiform and smoothish awn. This 
plant seems says of recognition as 


W someone River’’); waste ground in Tampa, June 1, 1886, 
Curtiss, no. 


1943] Fernald,—Virginian Botanizing Under Restrictions 495 

ATE 796 is of on cae AUTUMNA were MS. : F1@s. 1 and 2, portions of 
Bein in Linnean Herb., nal phot panne Er ough kindness of Mr. S. Savage; 
FIG. 2, summit of plant fron Garrettsville, Patan County, Ohio, R. J. Webb, 


no. 
PLATE 797, H. AUTUMNALE, var. CANALICULATUM (Lam.) Torr. & Gray, X 1, 


except Fic. 5: Fia. 1, portion of Helenia foliis decurrentibus, ete. of Hort tus 
Cliffortianus, courtesy of Dr. John Ramsbottom; F1a. 2, summit of plant from 
River nce, Lp toor sl Montmagny County, Quebec, Fernald & 


Pease, no, 25,324; ria. 3, portion of original plate of H. canaliculatum Lam.; 
Pld ‘ oo 5, portions of Taateatioe of Aster soles alatus of Cornut, Canad. 
Piate 798, H. mp ich pe var, PARVIFLORUM Ny hes ) Fern., < 1: FIG 
portion of Typ or 1soryPE of H. parviflorum Nu 2, summit of plant 
oa isa, of South Guay. Nansemond County, "viata, Fernald & Long, 


SENECIO AUREUS AND ITS GEOGRAPHIC VARIETIES AND ALLIES 
IN EASTERN Nortuo America (PLATES 800-806)—To one who 
from boyhood has known as Senecio aureus the common glabrous 
or promptly glabrate plant of peaty meadows, swales and swampy 
thickets of New England, New York and Pennsylvania, with 
suborbicular to round-ovate obtusely crenate radical leaves which 
are tufted from very slender and cord-like elongate rhizomes and 
basal offshoots, it is very disconcerting on every trip to tidewater 
Virginia to see as a dominating plant of calcareous woods and 
fertile bottomland, never in meadows and boggy places, a coarser 
plant with stout almost finger-like purple basal offshoots, very 
large heads and, when young, a copious long tomentum at the 
bases of the petioles, on the unexpanded leaf-surfaces and over 
the unexpanded corymb, and to be obliged to call it also S. aureus. 
So different is this southern plant of rich woods from the common 
plant of swamp and swale northward that it, again, comes as a 
surprise to find that it is true S. aureus L. Sp. PI. ii. 870 (1753). 
Linnaeus based the species primarily upon the Clayton collection, 
nos. 249 and 286, from Virginia, described by Gronovius, with a 
second reference to the Jacobaea virginiana of Morison and of 
Ray. The Clayton plant, which is the Type (our PLATE 800, 
FIG. 1) is very characteristic material of the woodland species of 
eastern Virginia, and phrases in the account by Gronovius but 
not quoted by Linnaeus are peculiarly significant to one who 
knows the southern plant, “leviter superne lanatis, radice parva 
atro-rubente”’, for the woodland plant of tidewater Virginia is 


496 Rhodora | DECEMBER 


very striking on account of the purple or dark red stiff rhizomes 
and basal offshoots and the lanate tomentum of the unexpanded 
corymb. ‘Madidis & umbrosis gaudet”’ of the Clayton account 
is also wholly satisfactory. So is the contradiction between the 
brief italicised diagnosis of Gronovius with “‘foliis crenatis’’ and 
the fuller account below ‘‘foliis infimis rotundis ad marginem 
serratis’’, for the type shows the margin serrate-dentate or with 
somewhat emarginate-truncate teeth as in some of the southern 
specimens, while the phrase “foliis crenatis’”’, taken over by 
Linnaeus, fits other specimens. 

Since the common plant of the northeastern states differs in 
several characters from typical southern Senecio aureus, standing 
midway between it and var. gracilis (Pursh) Wood, and since the 
more northern membranous-leaved plant (of Newfoundland, 
Anticosti, Gaspé, northern New England, etc.) which has errone- 
ously passed as the Cordilleran S. pseudaureus Rydb., needs 
clarification, I am here summarizing my interpretaion of the 
varieties of S. aureus: 

a. Radical leaves sana 5 with rounded or blunt teeth... .b. 

b. Undivide eaves suborbicular to round-ovate 
roadly ion at summit, with a well fein 
Basal teaicole at flowering time 4-10 mm. thick, 


deep red or purple; margins of basal sheaths of many 
petioles, surfaces of unexpanded leaves and unex- 


) 

tum; involucres 8-11 mm. high; disk-corollas 6-10 

mm. long; achenes 3.54 1 mm. long.......... S. aureus (typical). 
Basal offshoots at flowering ise” 1-5 mm. thick, 


bs glabrous or sparsely short-tomentose and 
glabrescent; involucres 5-8 (-9) mm. high; disk- 
m. long 


ong an ad, 
often purplish beneath, the longer patitlne 0.5-3 

dm. long; involucres 9mm. high.......... Var. intercursus. 
Stems repr io solitary, ‘2-6 dm. high; basal 
offshoots rarely = ink radical leaves mem- 
branaceous, 1-3. 1-2.8 cm. br yey 

n both sides the “longer petioles 2-10 cm ge 

lon, ne ian pe eT sa a eS " Var. gracilis. 
b. Undivided basal leaves oblong-oval, rounded to base or 
some of them are, subcordate, the abla blades 
_ - em. long and 1.5-4.3 em. b road, submembrana- 

PRC eee upiety Clk vue oe Lure. ort Var. semicordatus. 


1943] Fernald,—Virginian Botanizing Under Restrictions 497 


a. Radical leaves sharply ans Bhpa or dentate at least at base to 
cutely or subacutely lacerate, ovate or oval to rotund, 

aocane cordate, acute to founded at tip, membranaceous. 

Var. aquilonius. 

S. aurEus L. Sp. Pl. ii. 870 (1753). Var. Ashez Greenm. in 
Ann. Mo. Bot. Gard. xvi. 406, pl. 35 (1929).—Rather stout; stiff 
basal offshoots purple, 3-15 cm. long; flowering stems 3-12 dm. 
high; radical leaves rounded- to oblong-ovate, becoming cori- 
aceous, in maturity up to 2.2 dm. long and 1.8 dm. broad.—Rich 
caleareous w oods and wooded bottomlands or in upland meadows, 
Florida to Arkansas, north to Virginia, Kentucky and Missouri. 
The following are characteristic. VIRGINIA: locality not stated, 
‘““Madidis & umbrosis _ ool Clayton (ryPE, photo in Gray 
Herb., our PLATE 800, . 1); rocky bank of brook north of 


ounty, Fernald & Gri scom, no. 451 ce ee ae Cypress 
Swamp, north of Dendron, Fernald, L g & Abbe, no. 14,243; 
rich alluvial woods, east of Cabin Point, ’ Rauie & Long, no. 
7985; bottoms of rich calcareous wooded ravines west of Clare- 
mont, Fernald & Long, no. 12,893; sandy alluvial woods, bottom- 
land of Powell Creek, Garyville, Fernald & Long, no. 7984. 
Norru Carouina: wet, shady ground, Biltmore, Biltm. Herb. no. 
889". Frorrpa: banks of Little River, Chapman. TENNESSEE: 
Maryville, Blount County, May 8, 1937, Godfrey; low wooded 
slope along Bean’s Creek, siege ‘County, Svenson, no. 10,018. 
Missourt: along James R., h of Springfield, Greene Co. 
September, 1934, Richard Seth, " Lisle Jeffrey; along Shoal 
Creek, southwest of Joplin, EZ. J. Palmer, no. 29,936. ARKANSAS: 
wet places, “N. W. Ar kansas’’, April, 1888, Harvey, no. 45. See 
p. 361. Phases 800, Fra. 1, and 801 

Senecio aureus in southeastern Virginia definitely crosses with 
S. tomentosus Michaux. In April, 1942, Long, Abbe and I 
found (see p. 361) an extensive colony growing at the upper 
border of bottomland-woods of Cypress Swamp near Dendron, 
with typical S. aureus in the woods, typical S. tomentosus in the 
open. The intermediate plants, some with narrow and merely 
round- to tapering-based radical leaves suggesting those of S. 
tomentosus, others with them broader and subcordate, all more 
or less tomentulose and with tomentose to glabrescent petioles, 
has the involucres tomentose. It is clearly S. aureus X tomento- 
sus. Foliage-material, misidentified as S. Crawfordii Britton, 
from a wooded ravine west of Claremont, Fernald & Long, no. 


498 Rhodora [DECEMBER 


12,892, is very similar but smoother, and its stout and elongate 
horizontal rhizome keeps it out of S. Crawfordii (see p. 506). 
It was a single plant growing with typical S. aureus, with S. 
tomentosus abundant only a few rods away. That much is clear. 

In Ann. Mo. Bot. Gard. xiv. 406, pl. 35 (1929) Greenman 
described as S. aureus, var. Ashei a plant of western Virginia 
and adjacent northeastern Tennessee, there growing in moist 
meadows. Evidently having in mind as true S. aureus! the 
glabrescent plant of wide northern and continental range, the 
author of var. Ashet emphasized for the latter the distinctive 
characters of typical southern S. aureus: “‘foliis inferioribus 
longe petiolatis . . . crenato-serratis juventate atringe 
albo-floccoso-tomentosis supra denique plus minusve glabratis’’. 
Evidently puzzled by the presence of tomentum on the young 
and unexpanded basal leaves in his var. Ashe7z, just such pubes- 
cence as occurs on the youngest basal leaves of the plant of 
Clayton’s region (see PLATE 801, Fic. 3), and frequently on the 
involucre as well (see ric. 5), Greenman surmised that this 
plant of the mountain-region of western Virginia and eastern 

Tennessee might be a hybrid of his S. aureus (presumably our 
var. intercursus) and the coastal plain S. tomentosus: “The origin 
of the plant isnot known . . . It may beahybrid. There 
are some indications that it may be a hybrid between Senecio 
aureus L. and Senecio tomentosus Michx. The former of these 
two species occurs in the valley of the South Fork of the Holston 
River, and the latter species is relatively common in eastern 
Virginia. While the habit of the new plant is like Senecio 
aureus, yet the prevailing outline of the leaves of the offshoots 
and those of the lower parts of the upright stem, as well as their 
texture and tomentose character, suggests Senecio tomentosus.”— 
Greenm. l. c. 405. The type of S. aureus var. Ashei was poorly 
collected, showing no rhizomes, but in outline of basal leaves and 
cauline leaves (so far as the mashed and crumpled leaves show) 
the foliage looks quite like that of the rypr of S. aureus and of 
the characteristic specimens above cited and illustrated (PLATES 

1 Among the specimens taken by Dr. Greenman to Berlin for his preliminary studies 
cursus, from Rhode Island, bears the pencilled ot “This specimen of 
Thu rber’ s parla ts nds very well with ‘specimens of S. aureus L. in herb. Willdenow at 


Berlin”. The Thurber like it sé ‘Berlin (if not now destroyed) 
are very insole from the Clayton type. 


1943] Fernald,—Virginian Botanizing Under Restrictions 499 


800, Fic. 1 and 801). Although on the coastal plain true and 
characteristic S. aureus rarely crosses with S. tomentosus, pro- 
ducing a pretty mixed progeny, some plants with horizontal 
rhizomes and basal offshoots of S. aureus, others cespitose as in 
S. tomentosus, there is no reason to imagine that the cespitose, 
heavily and permanently tomentose S. tomentosus, with its 
narrow and noncordate radical leaves, its undivided or but 
shallowly toothed cauline leaves, its columnar and_ heavily 
hirtellous drab achenes (pLaTE 800, Fic. 3) with prominent 
rounded ridges—there is no reason to imagine that such a plant, 
restricted to sterile soils of the Coastal Plain and adjacent prov- 
inces, had anything to do with the origin of the continental and 
upland species of rich or calcareous areas, with horizontal elon- 
gate rhizomes and basal offshoots, oval to orbicular and deeply 
cordate glabrous or glabrescent radical leaves, glabrous and 
deeply pinnatisect cauline ones, and glabrous, slender brown to 
reddish achenes (Fic. 2) without very prominent ridges. The 
cited stations of S. aureus, var. Ashei are more than 200 miles 
west of the nearest stations known for S. tomentosus! 

Senecio aureus, var. aurantiacus Farwell in Am. Midl. Nat. 
xii. 74 (1930) is described as having red stems and “traces of 
floccose tomentum present throughout the plant at the flowering 
time”. Its distinctive character, “Ligules . . . 3-4 lines 
ong . orange-red, reflexed” is one unknown to me. 
The deacriiiae otherwise does not make clear what var. aurant?- 
acus re be, in a region where ee variations occur. 


Var. sale ea standing between the coarse, southern typical 
Senecio aureus and the smallest variety of the species, var. 


500 Rhodora [DecEMBER 


gracilis (Pursh) Wood, is the plant most generally passing as 
typical S. aureus. 


*Var. INTERCURSUS I have seen from Virginia only from 
ROANOKE County: moist woods along Back Creek, south-south- 
east of Starkey Post Office, C. E. Wood, Jr., no. 3988. It is 
presumably of broader upland range. 

Var. GRACILIS (Pursh) Wood, Class-Bk, 211 es independ- 
ently published as a new comb. by B ritton a Britt 
Ill. Fl. iii. 481 (1898). S. gracilis Pursh, Fl. Am. Sept. ii. 529 
(1814).—Meadows, swamps and bogs, iudiautetic to ‘North 
pice _ to Grodininn): south to North Carolina and 
Arkan 

ve Sains (Mackenzie & Bush) Greenm. in Ann. 
Mo. Bot. Gard. iii. 129 (1916). S. semicordatus Mackenz. «& 
Bush in Mo. Bot. Gard. Ann. Rep. xvi. 107 (1905). S. aureus 
x Balsamitae Greenm. in Ruopora, x. 69 (1908).—Calcareous 


ern Ohio, Tllinois and Missouri. 
urEvs L., var. aquilonius, var. nov. (Tas. 803), planta 
valde hisolsntonk: rhizomatibus gracilibus valde horizontaliter- 
que elongatis (ad 3 dm . longis) ; foliis basilaribus membranaceis 
ovatis vel ovalibus vel ‘rotundis serratis vel acute vel subacute 
incisis ad basin valde cordatis; foliis caulinis membranaceis basi 
pinnatifidis—Newfoundland and Céte Nord and Anticosti 
Island, Quebec, to Algoma District, Ontario, south to Cape 


vania, northern Ohio, northern Indiana and Winton Many 
specimens previously misidentified with the w en. ‘s. pseud- 
eda Rydb. ollowing, selected from a very large series, 
e characteristic. NEWFOUNDLAND: Atagrtae Grand Falls, 
Faly 3, 1911, Fonte & Wiegand, no. 6398; park-like openings in 
amp mossy woods on the Silurian hills ‘back of Birchy Cove 
(Curling), July 5, 1910, Fernald & Wiegand, no. 4174; spruce 
woods and thickets, slope of Lookout Mountain, Bonne Bay, 
ernald, Long & Fogg, n . 2145; brooksides and damp bushy 
ravines on the Piticetones ‘tableland, alt. 200-300 m., Table 
mentee Port A Port Bay, July 16 and 17, Fernald & St. John, 
10, 872: damp gia in woods near Overfall Camp, Great 
Codkey River, July 7, 1939, Pease & Edgerton. Qursec: dans 
les bois claire prés de la riviére, Riviére "de Rénard, Anticosti, 
5 aotit 1927, Victorin & Rolland, no. 27,645; dans les bois de 
Coniféres, R. Vaureal, Anticosti, 12 aoft 1927, Vuictorin & 
Rolland, nos. 27,640 and 27,641; mossy Arbor Vitae woods east 


1943] Fernald,—Virginian Botanizing Under Restrictions 501 


of Grande Coupe, Percé, August 6, 1907, Fernald & Collins, no. 
1208; springy meadows in woods at 600 m. altitude, North Fork 
of Madeleine River, Gaspé County, July 31, 1923, Fernald, 
Dodge & Smith, n 0. 26 ,096; calcareous alpine meadows, alt. 1000— 
1125 m., Tableton Mountain, August 7, 1906, Fernald & 
Collins, no. 260; coniferous forest, “Low’s Trail” from the Forks 
of the R. Ste. Anne des Monts to Table-topped roger, July 31 
and August 14, 1906, Fernald & Collins, nos. 765, 7 6 (TYPE in 
Herb. Gray), 767: wet alluvial shores, between hatde and the 
Baie des Chaleurs, Bonaventure River, August 5-8, 1904, 
Collins, Fernald & Pease; gravel-beaches and bars between the 
Forks and Brulé Brook, Little Cascapedia River, July 29 and 30, 
1904, Collins, Fernald '& Pease (Pease, no. 5022); ey: bank, 
Restigouche River, July 24-25, 1929, Rousseau & Bonin, no. 

2,211. Nova Scotta: Barrasois Thi, Cape Breton, Nichols, no. 
52 PRINCE Epwarp ISLAND: swampy Letts and Thuja woods, 
Tignish, Fernald, Long & St. John, no. 8245. Marne: wet 
thicket, Limestone, Tune 22, 1898, Fernald; Aree Opel A 
Presque Isle, July 12, 1902, Williams, Collins & Fernald; cedar- 
swamps and clearings, Blaine, June 23, 1898, Fernald 1 no. 2404; 


Roque Blufts, July 23 to ; meadow near 
Half-moon Stream, Unity, June 16, 1935, G. a Rossbach, no. 926; 
wet ground, ae roo Clinton, July 8, rid 


July 6, 1907, A. H. Moore, no. 3523, Pease, no. 10,316; wet place 
in woods, Hill, August 29, 1933, Chas. Bullard; rich woods, 
Durham, June 21, Nghe "Ho dgdon, no. 4034. New York: 
Oneida, May 15 , 1918, ouse. PENNSYLVANIA: Corry, May 7 
1896, J. R. Churchill. (eggs black-ash swamp, Sault Ste. 
Marie, July 18, 1935, Taylor és ri no. 1687. Onto: Garretts- 
ville, Po ortage County, Webb, 202. MricHiGcan: tamarack 
Swamp, south of Ann Arbor, May 8, 1898, S. H. Burnham; Port 
Huron, May 18, 1896, C. K. Do dge. InpraNa: along a brook, 
Otis, May 20, 1911, Sherff. Wisconsin: Brown County, June 
ii, 1900, Schuette. 

Senecio aureus, var. aquilonius is the northeastern plant which 
has recently been passing in my own work and that of others as 
S. pseudaureus Rydb. It certainly strongly suggests the plant 
of the Cordilleran region in the texture and toothing of its basal 
leaves and the tendency to laceration and enlargement of the 
lower cauline blades. S. pseudaureus, however, has a stout and 
relatively short rhizome which usually forks into a tuft of crowded 
ascending crowns; its basal leaves tend to be oblong-ovate as in 


502 Rhodora [DECEMBER 


S. aureus, var. semicordatus (Mackenz. & Bush) Greenm. and 
only slightly, if at all, more cordate, though sometimes broader, 
on thickish petioles 1-10 (rarely —23) em. long; its pale-flowered 
heads are on pedicels 1—5 (rarely —9) em. long. S. aureus, var. 
aquilonius stands off from it through its slender and horizontally 
creeping rhizome (up to 3 dm. long), with scattered ascending 
branches and tufts of leaves or flowering stems; it thus belongs 
with S. aureus var. intercursus and passes into it. Its basal leaves 
are ovate to oval or even orbicular, deeply cordate, on filiform 
petioles mostly 0.5-2.5 dm. long; and its deep yellow-flowered 
heads are on filiform pedicels mostly 3-12 cm. long. 

S. eae var. AQUILONIUS, forma ecoronatus (Fernald) 
comb. n S. pseudaureus, forma ecoronatus Fernald in RHopoRA, 
xx. 595, (1928 y 

. PAUPERCULUS Michx., forma inornatus, f. nov., nang basi- 
laribus panics angusto oblanceolatis laminis 1.5-2.3 cm. longis 
: m. latis; foliis caulinis valde reductis Bitoahes: es. 
em. lo tes ‘2-5 mm. lati tis; lets ed lineari-attenuatis; floribus 
discoideis, ligulis nullis. —Micn en ne pavement, Seul 
yee School, ae Coney | July 12 Eigen Pease & Ogden, 

. 24,946 (TYPE oe Herb. Gray.). Awa : Mi ller House, on 
Ste ese Ticket 115 miles nh of Pigdenks elevation 2100 
feet, July 12-18, 1940, Edith Scamman, no. 216 6. 


Not to be confused with Senecio pauperculus, forma verecundus 
Fern. in Ruopora, xxx. 225 (1928), of Anticosti, a plant which 
has all the technical characters of S. gaspensis Greenm., to which 
it must be transferred. 


S. PAUPERCULUs Michx., var. neoscoticus, var. nov. (TAB. 806), 
planta 1-4 dm. alta; foliis ‘basilaribus indivisis 1.5-6.5 cm. longis 
1-2 em. latis; foliis caulinis imis plus sae aS pinnatifidis 3.5-6.5 
em. longis 0.7-2 cm. latis; capitulis 2-25, longe pediogeb 
involucris 4-5 mm. altis basin versus persistenter tomentosis; 
achaeniis glabris 2 mm. longis—Nova Scorta: talus of gypsum 
cliffs, Five-mile River, Hants S Cae July 19, 1920, Pease & 
Long, no. 22,914 (type in Herb. Gray.), distrib, as var. Balsa- 
mitae (Muhl.) Fern.; common on gypsum, Antigon ish Harbour, 
July 11, 1941, A. E. Roland, no. 41,741, distrib. as var. Balsa- 


1 Nore on THE TeRM PHYLLARY.—The term PHYLLARY, now largely used 
of involucral bract, is given by Jackson, Gloss. Bot. Terms, as coming from a Latin 


have, naturally, made an ablative plural “ phyllaribus o & Hooker and 
other English phytographers had e term as son defined in the Century Dic- 
ry and as originally adopted in this count i aa from the New Lati 


1943] Fernald,—Virginian Botanizing Under Restrictions 503 


mitae; rock-faces and crevices of gypsum cliffs, Port Bevis, 
Victoria County, August 27, 1920, oo & Long, no. 22,915, 
also distrib. as var. Balsam mitae. QUEB Seige ee bank near 
Percé, Gaspé County, July 14, 1928, pines no. z 

Var. neoscoticus is the only variation of the a Senecio 
pauperculus (PLATE 804) known to me from Nova Scotia. It 
superficially resembles var. Balsamitae (pLATE 805) in having 
well developed lower cauline leaves, but it is unlike any of the 
other varieties of the species in having densely and permanently 
tomentose involucres, in the Type the tomentum at the base of 
the involucre or, in some plants, extending to the tips of the 
phyllaries. The conventional and not too convincing distinction 
between S. plattensis Nutt. (1841) and S. pauperculus Michx. 
(1803) is the occurrence somewhere on the former (on stem, 
petioles, pedicels or involucre or on two or more of the areas at 
the same time) of more or less persistent tomentum, while S. 
pauperculus is glabrate or merely with flocculent tufts of tomen- 
tum. Furthermore, to quote Greenman, the former species has 
“achenes usually hispidulous along the angles, sometimes 
glabrous”. In S. pauperculus we get ‘“‘achenes glabrous or 
hirtellous along the angles’. That leaves mighty little which 
seems specific. Nevertheless, of many scores of mature collec- 
tions of S. plattensis which I have studied essentially all have 
hispidulous achenes. Of more than 200 members of the inclusive 
S. pauperculus examined all but 3 numbers have shown glabrous 
achenes. I am not ready to reduce S. plattensis, without field- 
acquaintance with it. But in its densely tomentose involucre 
S. pauperculus, var. neoscoticus might be looked upon as a con- 
necting link. In all other characters, however, it is good S. 
pauperculus. 

A southeastern variety, Senecio pauperculus, var. praelongus 
(Greenm.) House, found from eastern Massachusetts to Michi- 
gan, south to Connecticut, Maryland and the upland of Virginia, 
is strikingly similar to and has often been mistaken for the ubi- 
quitous southern S. Smallii Britton. In New England, New 
York and Michigan it passes into S. pauperculus var. Balsamitae, 
while in the elongate cauline and basal leaves and the numerous 
heads it looks like S. Smallii. The latter, however, usually has 
copiously hispidulous achenes only 1.5-2.2 mm. long, the former 
glabrous achenes 2-2.5 mm. long. There seems to be a recog- 


504 Rhodora [DECEMBER 


nizable difference in the phyllaries. In S. pauperculus they are 
linear-attenuate, gradually tapering from the base; in S. Smallii 
they are more definitely linear-oblong, with parallel sides, and 
tapering only toward the summit. In typical S. pauperculus 
and its var. Balsamitae (Muhl.) Fern., furthermore, the more or 
less elongate rhizome is commonly prostrate, with decumbent 

ranches. In S. Smallii the short rhizome has crowded erect 
crowns. The photograph of the Type of S. Balsamitae Muhl. ex 
Willd. sent me by Professor Diels is of the northern few-leaved 
plant. I am clinging to these characters for what they may be 
worth. 

Far to the northeast there is a series of plants of this affinity, 
growing from western Newfoundland to Anticosti, thence to 
northern Maine, which differs from Senecio pauperculus in 
usually broader and heavier foliage, the basal leaves tending to 
be elliptic or oval but sometimes oblanceolate, the disk very 
broad. These are the two extremes which were described as S. 
gaspensis Greenm. in Ann. Mo. Bot. Gard. iii. 138 (1916) and as 
S. Balsamitae var. firmifolius Greemn. in Ruopora, vii. 244 
(1905) or S. pauperculus var. firmifolius (Greenm.) Greenm. in 
Ann. Mo. Bot. Gard. iii. 166 (1916). In their extremes the two 
are very different, but some of the larger specimens cited by 
Greenman as var. firmifolius are inseparable from smaller ones 
determined or cited by him under S. gaspensis. Typical S. 
gaspensis differs still further from S. pauperculus in having the 
undivided basal leaves broader (1.5-4.5 em. broad, the basal 
leaves of S. pauperculus ranging from 0.5-2 cm. broad). Further- 
more, in both typical S. gaspensis and in S. pauperculus var. 
firmifolius the phyllaries are linear-oblong, with parallel (instead 
of gradually converging) sides, tapering only near the tip. This 
apparently fairly definite species also includes the discoid form 
of Anticosti which, before I had spent two weeks upon the group, 
I placed under S. pawperculus. As I understand the plants I 
should now treat them as follows. 


S. GASPENSIS Greenman i in Ann. Mo. Bot. Gard. iii. 188 (1916). 


.5-8 em. long and 1.5-4.5 em. bro ad; flowering stem very leafy; 
larger cauline leaves 5-12 em. fe 1-3 em. broad, eet pin- 


1943} Fernald,—Virginian Botanizing Under Restrictions 505 


swamps and calcareous cliffs, Anticosti Island and Gaspé Pen- 
insula, Quebec, to upper St. John Ri iver, Maine (where apparently 
crossing with S. pauperculus, var. Balsamit tae). 
orma verecundus (Fern.), comb. nov. S. pauperculus, 

forma verecundus Fernald in Ruopora, xxx. 225 (1928).— 
Swamps, flats and calcareous talus, phir a Be Island. 

With the recognition of Senecio gaspensis, the discoid form 
from Anticosti, of which I now have several numbers, finds its 
at there. 


. firmifolius (Greenm.), comb. nov. S. Balsamitae, var. 


nn. 
(1916).—Stiems 0.2-2 dm. high, stiffly erect, often much tufted; 
undivided basal blades 1-8 em. lon ng, 0.7—2.5 em. broad; flowering 
stem nearly naked except at base or with greatly reduced leaves 
1-3.5 em. long and rarely 1 em. broad; pedicels 0.2-5 em. long; 
achenes 2.4-2.6 mm. long.—Dry calcareous rock, shingle and 
talus, western Newfoundland; Nits Nord (Betchouane), Anti- 
costi and Gaspé Peninsula, Quebe 
With Senecio pauperculus, S. siiaae S. gaspensis and, per- 
haps, S. plattensis showing possibly more recognizable characters 
than they have usually been accorded, there remains the local 
S. Crawfordii Britton. This local species has linear-attenuate 
phyllaries somewhat as in S. pauwperculus and its varieties, few 
large heads and the well developed basal leaves on very prolonged 
petioles, so that the typical lower blades are only one fifth to one 
third as long as the erect petioles. The elongation of petiole was 
probably induced by its often paludal habitat. A most signifi- 
cant character seems to have been overlooked. In S. pauper- 
culus and S. gaspensis there is a more or less elongate and creeping 
or decumbent rhizome, from which spring leafy tufts independent 
of the flowering stem (snatched or merely jerked-up specimens 
rarely show good rhizomes). In all the material of S. Crawfordii 
I have seen the rhizome is reduced to a short vertical or ascending 
crown which is praemorse and not creeping. When Mr. Long 
and I collected in Surry County, Virginia, a few leaf-specimens 
which we mistook for S. Crawfordii, we did not have the rhizome 
in mind. Our Surry County material (no. 12,892), reported 
under that name, has a strongly developed horizontal rhizome. 
It seems to belong with a series collected by Mr. Long, Dr. E. C. 
Abbe and me, also in Surry County (Fernald, Long & Abbe, no. 


506 Rhodora [DECEMBER 


14,245) which is an unquestioned hybrid of S. aureus L. and S. 
tomentosus Michx., the series occurring with the two parents and 
combining their characters. 

Returning to Senecio Crawfordii, the short praemorse crown- 
like rhizome, the very long petioles and the other characters 
above noted clearly mark it. Although local, the species has a 
broader range than usually assigned it. I am quite incapable of 
separating from it the typr from Indiana of S. obovatus, var. 
umbratilis Greenm. in Ann. Mo. Bot. Gard. iii. 115 (1916). It 
has the short praemorse erect rhizome, the long-petioled elliptic- 
oblong basal leaves, the cauline leaves and the heads so nearly 
identical with isotypic material from Mr. Joseph Crawford him- 
self that the only difference I can see is that the type of S. 
obovatus var. umbratilis is in young anthesis, the Crawford ma- 
terial in fruit. Similarly the Tennessee material cited under S. 
obovatus var. umbratilis (S. M. Bain, no. 421) is so similar to 
characteristic S. Crawfordii from Camden County, New Jersey 
(Witmer Stone), Bristol, Pennsylvania (Fretz) and Suitland, 
Maryland (Blake, no. 9391), that it looks as if it were collected 
with any one of them. Material from open peaty pineland near 
Middlesex, Nash County, North Carolina, Godfrey & White, no. 
7017, correctly identified as S. obovatus, var. umbratilis, is to me 
inseparable from the Stone, Fretz, Blake and Bain specimens of 
S. Crawfordii, a species which is evidently a southern one reach- 
ing a northeastern limit in eastern Pennsylvania and adjacent 
New Jersey. 

Senecio obovatus Muhl. is a stoloniferous plant, with creeping 
rhizomes derived from the elongate stolons; its obovate to sub- 
rotund radical leaf-blades are decurrent into the relatively short 
and upwardly broadened petioles; its relatively short involucre 
with few phyllaries abruptly tapering from above the middle. 
It is very different from S. Crawfordii, which is a plant of low 
grounds (meadows and bogs), whereas S. obovatus thrives on 
calcareous rocky or rich wooded banks; it would be quite unhappy 
in an acid bog. Not all plants referred in the original account 
to S. obovatus var. umbratilis are S. Crawfordii, however. Many 
of them I have not seen, but Bush’s no. 1356 from woods, Fulton, 
Arkansas, is so like a specimen from Williams’s Bluff, Knoxville, 
Tennessee, of the reputedly endemic Knoxville specialty, S- 


1943] Fernald,—Virginian Botanizing Under Restrictions 507 


obovatus var. diversifolius Greenm. that I can find no difference. 
The latter very luxuriant series is rather closely approached by 
Overgrown individuals as far away from Knoxville as New 
England. It seems like overgrowth in unusually favorable 
rane 

8. s Muhl., var. Elliottii (Torr. & Gray), stat. nov. 
S, Elliotts: 5 Set & Gray, Fl. N. Am. ii. 443 (1843). 

Typical Senecio sao and its var. rotundus Britton are 
slender and glabrous or promptly glabrescent plants. From the 
mountains of eastern West Virginia to Alabama and northwestern 
Florida var. Elliottii stands off as a stout plant, with the stems, 
petioles, often the lower leaf-surfaces and the short pedicels at 
first arachnoid-tomentose. The radical leaves are more coria- 
ceous than in typical S. obovatus, the upwardly dilated petiole 
obsolete or very short. This is S. Elliottii as described by Torrey 
& Gray, with stem “floccose-woolly when quite young. Radical 
leaves . . . 2-3 inches in diameter, frequently coriaceous 
- . . the winged petiole . . . much shorter than the 
lamina”, ete. Torrey & Gray, noosa this plant of Georgia, 
Florida sink Alabama, supposed it to be the S. obovatus of Elliott, 
Sketch, ii. 329; but Elliott, describing the plant found on the 
Santee River, mares said “Stem . . . glabrous. Leaves 

gla as ous,” ete. There is certainly doubt whether 
Elliott had the coarse and pubescent plant of the | 


County, Pease, no. 26,587; Milton, Cabell County, Louis 
Williams, no. 309. SoutH Caro.tna: slo opes of Paris Mt., 
Greenville, Mackenzie, no. 2986. FiLoripa: calcareous woods, 
near Mariana, Jackson County, Harper, nos. 53 an 
ALABAMA: chalky ravine near Epes, Sumter County, H ae 
nos. 3001, 3005 and 3159. 

*S. PLATTENSIS Nutt. S. pseudo-tomentosus Mackenz. & Bush 
in Trans. Acad. Sci. St. fouls: xii. 88, pl. xvii (1902).— Western 
Virginia: Frepericx County: limestone ledges, Cedar Creek, 
Hunnewell, no. 12,445. SHENANDOAH CovuNTY: limestone bar- 
rens, Strasbur urg, Hunnewell, no. 14,469; old field 1 mile north of 
Strasburg, Allard, no. 103, as S. tomentosus. RoaNoKE County: 
limestones or dolomites, either wooded or cleared, along Roanoke 
River, Dixie Caverns, C. E. Wo od, Jr., nos. 3387, 3392, 3670, 
3687, 3689, 3693, 3694, 5786-5797; steep shaly slope, with lime- 
stone ledges, partly wooded, along Roanoke River south-south- 
west of Wabun, Wood, no. 510 


508 Rhodora [DECEMBER 


Although Greenman cites Senecio plattensis as extending east- 
ward only to southern Ontario, Michigan, Indiana, Missouri, 
Arkansas and Louisiana, it has several times been collected in 
Ohio; and on July 2, 1907, Messrs. W. W. Eggleston, R. W. Wood- 
ward and C. A. Weatherby collected very characteristic material 
of it on limestone cliffs at North Pownal, Vermont, in a general 
region (southwestern Bennington County) notable for the isola- 
tion of more western species, such as Carex Richardsonii R. Br. 
(see Fernald in Ruopora, xxxiv. 229) and Collinsia parviflora 
Dougl. (see Weatherby in Ruopora, xxxvii. 422). Very char- 
acteristic S. plattensis (including S. pseudo-tomentosus) was col- 
lected on limestone ledges in Frederick County, Virginia, in 1932 
by Hunnewell and soon thereafter by himself and independently 
by Allard near Strasburg in Shenandoah County. These speci- 
mens are inseparable, so far as I can detect, from the Vermont 
plant and from much S. plattensis from Ontario, Ohio, Michigan, 
Indiana, Wisconsin, Illinois and Missouri, thence westward to 
the Dakotas, Nebraska, Kansas, etc. They come from more or 
less open or exposed habitats. Mr. Wood’s extensive series from 
along the Roanoke River is mostly from woods, bluffs and slopes. 
On the whole it is greener, thinner-leaved and with greater tend- 
ency to low forking of the inflorescence, but I am quite incapable 
of finding any morphological characters to separate it. It is 
smaller-headed than in some S. plattensis but not all, and its 
low-forking inflorescence and its excessive development of pin- 
natifid cauline leaves are readily matched in material from Ohio, 
Indiana, Wisconsin, Illinois, Minnesota, Missouri, Arkansas, ete. 
It seems to be a shade-state rather than a definite variety. In 
fact the numbers from less shaded habitats have thicker leaves. 


In PLATE 800, Fic. 1 shows the Type (from Clayton) of er AUREUS L.; 


FIG. 2, an achene, x 10, from Great Neck, Princess Anne County, Virginia, 
Fernald & Griscom, no. 4517. Fig. 3, achene, x 10, of S. chemateeat from 
Buckroe, Virginia, B. L. Robinson, n 0. 

PLATE . AUREUS: FIG. l, ead, < 1, from Great Neck, Princess Anne 
County, Virgi , iscom, no. 4517; 2, portion of mat 
basal leaf, x 1, from west of Claremont, Virginia, Long, no. 12,893; 

1G. 3, unexpanded basal leaf, x 2, arysville, Prince George pois 
Virginia, Ferna no. i 4, portion of unexpanded corym), 
2, from no. 7984; Fic. 5, involucre, X 5, fr oan. ore slope of Bull Run 


Mountains, Fauquier Bounty. Virginig, Allard, n 
PLaTE 802, . AUREUS, var. INTERCURSUS, 9 poe ‘all figs. from TYPE: 
ye i, hee, < 1; Fie. 2, foliage, X 1; Fic. 3, corymb, X 1; FIG. 4, involucre, 


1943] Fernald,—Virginian Botanizing Under Restrictions 509 


E 803, S. AUREUS, var. AQUILONIUS, n. var., all rics. X si Brig TYPE: 
FIG. rs qt FIG. 2, basal leaves; FIG. 3, cauline leaves; FIG. 4, co 
PLaTE 804, S. PAUPERCULUS Michx.: FIG. 1, TYPE, X 34, original ‘photog 
by the author; ries. 2 and 3, panei peg tall plank 3c 1, from wet peat, 
Port aux Basques, Newfoahnils nd, Fernald, Long & Dunbar, oN 27,160; 
FIG. 4, basal tuft, x 1, from no. 27,160. 


Fia. 1, paige ft Me plants of ‘ypesheet, Sa Ras esy of Professor Diels: 
FIGs. 2 and 3, seit a peg 1, from near ope, Lancaster County, 
Pennsylvania, June 5, 1900, Heller; ria. 4, bas a batt, > ee raat same collec- 
we Gad 5, involucre, < 5, from same collectio 

E 80 CULUS, var. NEOSCO tie n. var., all figs. oe TYPE: 
sca ve in 2, a ‘single plant, x 1; Fig. 3, involucre and pedicel, x 6 

*CIRSIUM VIRGINIANUM (L.) Michx., forma bees (Small), 
are nov. Carduus revolutus Small, Fl. Se. U. 8.1 and 1341 
(1903). Cir. —_ —— (Small) Petrak in Beihaft Bot. 
Centralbl. xxxv. Ab. 2:558 (1927). Sussex County: exsiccated 
argillaceous vrimand Ana 2 miles east of Stony Creek, no. 9223. 

Most certainly Cirsium revolutum is not a distinct species. In 
all characters it is like C. virginianum except that its lower and 
median cauline leaves are broader and deeply pinnatifid. Typi- 
cal C. virginianum (as shown by a photograph of the type as well 
as by the original description) has the very numerous cauline 
leaves lance-acuminate, the lower and median ones only 0.5-1.5 
cm. wide, entire or with remote, divergent, short, subulate 
prickles. This plant occurs in wet pineland or sphagnous or 
peaty bogs, swales and clearings, mostly on the Coastal Plain, 
from Florida to New Jersey. In forma revolutum, which has the 
Same broad range, from Florida to New Jersey, the lower and 
median cauline leaves are 1.5-4 em. broad and deeply See 
with lanceolate to narrowly triangular simple or forked o 
toothed long-pointed lobes, while the reduced upper leaves may 
be similar, though smaller, or quite like those of typical C. vir- 
ginianum. The two forms seem to occur in somewhat pure 
Stands so that all of a single station are more or less uniform. 
Forma revolutum is worthy that much recognition but surely it 
is not a distinct species. 

Cirsium virginianum is generally supposed to extend westward 
to Texas, thence northward to southern Ohio, southern Indiana 
and southern Missouri. The plant of the Interior, however, is 
C. flaccidum Small, although its author missed some of its 
most important characters and restricted it to “Low pinelands, 
Coastal Plain, Fla. to N. C.,” while the plant of the Interior 


510 Rhodora [DECEMBER 


was left in C. virginianum. True C. virginianum has tuberous- 
thickened first year’s roots (a character rarely shown in the 
herbarium), C. flaccidum slender root-fibers; C. virginianum has 
very numerous (40-70 or more) firm or coriaceous cauline leaves, 
C. flaccidum few (10-20) thin and flexible ones. In C. virgini- 
anum the peduncle-like flowering branches have several bractei- 
form leaves, in C. flaccidum the peduncles are naked or with only 
1 or 2 bracts; in C. virginianum the involucre is 1.5-3 em. high, 
in C. flaccidum only up to 2 em. high. C. virginianum, of savan- 
nahs, bogs and wet pinelands of the Coastal Plain from Florida 
to New Jersey, flowers from mid-August to October. C. flac- 
cidum of dry, acid woods, thickets and ravines, from Georgia 
(Small says North Carolina) and northern Florida to Texas, 
north to southern Ohio, southern Indiana and southern Missouri, 
flowers in May and June, sometimes in April. 

The following taken from the Carolina and Virginia sheets in 
the Gray Herbarium of Cirsium virginianum, give, in corrobora- 
tion, the habitats and flowering dates: sphagnous bog, Sept. 13; 
boggy depression, Oct. 18; low woods, Oct. 15; open pine and 
oak woods, Sept. 18; clearing, Aug. 19 (first flowers); damp pine- 
land, Sept.; bog or savannah, Sept. 10; exsiccated argillaceous 
pineland, Aug. 24; pineland, Aug. 28 (first flowers); savannah, 
Oct. 13; pineland, Oct. 13 and Aug. 29 (first flowers); savannah, 
Sept. 1; sphagnous bog, Sept. 10; bog or savannah, Sept. 10 and 
11. Average date September 15. All material in the Gray 
Herbarium of Cirsium flaccidum with sufficient data gives the 
following result: dry woods, Lookout Mountain, northern 
Georgia, June 4; woods near Chattanooga, Tennessee, May 23; 
dry woods, Talledega Co., Alabama, May 22; head of deep 
ravine, with Kalmia latifolia, Perry County, Indiana, June 4; 
upland woods, Polk County, Texas, May 23; dry bank, Houston, 
Texas, April 18. Average date May 23. Such data, associated 
with the different roots, many and firm or few and pliable leaves, 
bracted or nearly naked peduncles, etc., are decisive.! 


1 Sir John Hill in 1765 separated the boreal genus Ozyria from Rumez. S80 mewhat 
later Sir James Edward Smith, obviously with his own views of the general soundness 
of Hill's work, wrote (the reference kindly given me by Dr. Blake): “Sir John Hill, 
it seems, first separated this plant from Rumez, and gave it the above re name. 
Sometimes, as Linnaeus says, a blind hen meets with a grain of corn.” . Eng 
Bot. ii. 189 (1828). Small had a good species but he did not know its best eur 
nor its range. 


1943] Fernald,—Virginian Botanizing Under Restrictions 511 


*HIERACIUM VENOSUM L., var. NUDICAULE (Michx.) Farwell. 
For discussion see Fernald in Ruopora, xlv. 323 (1943). The 
only Virginian material I have seen is from the slopes of Bull 
Run Mountains, Prince WiLuiAmM County, Allard, no. 647 and 
3321. The plant of the Tidewater region is typical H. venosum. 


INDEX 
New scientific names are printed in full-face type 


Acer, 364; floridanum, 363, 379, 453 
Acerates Pham 458, var. lanceo- 


4 
Allies in eastern North America, 
Senecio aureus and its Geographic 
le dani and, 495-509, pls. 800- 


Amianthium Muscaetoxicum, 369, 
374, 393 
Amsonia bag Sanne corense. 380, 457 


Amygdalus 
Fee incoming ° 380; scoparius, 377 
alves 


ndinaria gigantea, 4 

Asarum, 374, 381-383, 309, § Hetero- 
tropa, 374, 399, 400; arifolium, 
398, 399; ere 398, 399; 
Lewisii, 00, 775; 
ria 8 er vee Shuttleworth, 


Acclepias fenkalade: var. paupercula, 
var. paupercula, f. 
flavifior ora, mes 
na parviflora, 370, 4 


Asim 

Antes, 361, 485; Thiceell, 461: luteus 
alatus, 487, 495, pl. 797 

Azalea, 451 


Bastard a emt er, 486 
Beech, 3 
Bokowin’, caroliniana, 379, 478 
Buckeye, 38 

Bulbostylis ciliatifolius, 369, 390 
— ss, 387, The common northern, 


ae caroliniana, 368, 400 
gon pallidus, 369, 398 
Cand amine bulbosa, 359; Douglassii, 


Cartas revolutus, 509 

Carex pn hee 367, side eager 
366, 367, 393; decomposita, 373, 
393; Richardsonii, 508; venusta, 
367, 393; virginiana, 

Cenchrus, ; carolinianus, ' 
echinatus , 387, 388, f. longispinus, 
388; idinertiie, '387, 388; longispi- 


us, 388; eo 387, 388; 

tribaloides, 3 87 

poems 371, 380 

Cerastium, 358; bra chypetalum, 358, 
350, "400; tetrandrum, 359 

Cerasus, 

Cherry, 451 

Chrysanthemum, 485 

Cirsium flaccidum, 509, 510; revo- 
lutum, 509; Mad dota 365, 509, 
510, f. revo olut m, 509 


coma, 402-409, 412, pl. 775, var. 

coactilis, 406, "407, 409, 410, 412, 
1. 780; crispa, 359; Fremontii, 

410-412, pls. 781, 782, var. Riehlii, 

411, 412, pl. 782; integrifolia, i 
2 . 


cea, 406, , 
410, 412, i aT7, a aes 405; 
viticaulis, oF ge 412, pl. 778 


Collinsia parvi 

Commelina diffusa, 377, 393 
Common northern Burgrass, The, 387 
Corydalis flavula, 358, 383, 412 
Crabapples, 3 

Cuscuta, 368 

Cyanococeus, 451 

Cynoctonum sessilifolium, 374-376, 


Cyperus, 451; densicaespitosus, 377; 
~ Weatherbianus, 4 


Danthonia compressa, 376, 385; 
spicata, var. longipila, 385 
Decodon 


Digitaria serotina, 371, 

Dracoe tine phen 461 

Drose 372; capillaris, 372, 413; 
rotundifolia, 3 _ 

Dwarf Southeastern 

Virginia, “phe, 396, pls. 772, 773 


Eleocharis eg riage ce 

Elychrysum foliis ese etc., 478 

Erigeron, 361; philadelphiews, 361; 
scaturicola, '361; strigosus, f. dis- 


(513 ) 


514 


Eriophorum virginicum, 405 
Eryngium, 373-375, 381; prostratum, 


Eulalia, 389; ea, 389; viminea, 
389, var. variabilis, 

; cuneifolium, 461; 
478; Sos Sipe 61; 
. . cor- 
7, 478, var. oneedy 


—— corollata, 370, var. panic- 
ulat 453; pecacuanhae, 370; 
pean Bra 


Fern, Royal, 396; Sensitive, 396 
Fly-poison, 374 
Fumaria officinalis, 412 


Gentiana cherokeensis, 365, 366 
Geographic Varieties 
eastern No merica, 
paca and its, 495-509, pls. 800— 


Glycyrrhiza lepidota, var. glutinosa, 
378, 453 


79, 
Homonym, 478, pl. 795; Pic 
tum, 478 
Goldenrod, BIL 
Gramineae, 38 
Gratiola virginiana, var. aestuario- 

80, 476 


Hamamelis virginiana, 450, var. 
ether 450 

rere ae autumnalis, 490; d 

; foliis decurrentibus, 

ae “485, “106, pl. 797 

Heleniastrum 485; folio breviore, 
etc., 486; folio longiore, etc., 486, 


Helenium, 379, 485, 490; Some Varie- 


90, 492, 


autu ale pubescens, “491, 92: : 
canaliculatum, 487, 488, 490, 491, 
495, pl. 797; floridanu 494, 


m, 
495, pl. 799; Godfreyi, 494, 495, 


INDEX 


pl. 799; grandiflorum, 491; hu- 
ronense, 491; latifolium, 486, 488, 
490; lo ngifolium, 488, 491, se 


493, 494; pubescens, 490, 492; 
490; quadridentatum, 


Holieathas angustifolius, 365 
a ata! sum crispum, 479; spatula- 


Henbit 382 

Heuchera villosa, 448 

Hexastylis, 374, 399 

Hickory, 377 ; 

Hieracium ss 511, var. nudi- 
eaule, 511 

Holosteum umbellatum, ‘si, 350 


Hypericum denticulat 
folium, 373, 453; coi tat 374-376, 
453 


Ilex coriacea, 461; vomitoria, 461 
Tris, we vern a, 360, 362, 363, pl. 770 
Ischaem m, 38% 

Jacobaea virginiana, 495 


Kalmia latifolia, 382, 510 
Kuhnia eupatorioides, 376 


Lady-fern, 396 


Lamium, 382 
oahek 457 
Lepidium virginicum, 382 


Leucojum aestivum, 359 
Liatris graminifolia, var. Smallii, 478 
Licorice, 
Ligusticum canadense, 378 
Lilium canadense, 394, 395, pl. 771, 
var. editorum, 93-30 5, pl. 771; 
canadense, f. ru 394; michi- 
ganense, 394, i cen, 395 
Limnobium Spo Bia, 
Linaria sea s, f. albina, 476 
Lobelia aioienta 461; : glandulifera, 
ee 477; siphilitica, 477, f. lae 


Long-leaf Pine, 369, 374-376 
Ludwigia alata, 461; pilosa, 461 


Malus, 450, 451; i aa var. 
puberula, 450; Halliana, 451 

Medeola, 396 

Micranthemum umbrosum, 367 

Monotropsis, 364; odorata, 363 

Morphological Differentiation of Cle- 


Sal ae es 


fon 


INDEX 


matis ochroleuca and Allies, 401, 
pls. 776-782 

Muhlenbergia, 379; brachyphylla, 
365, 366, 380; frondosa, 379; gla- 
briflora, '379, 385 

Myosotis verna, 459 


Narcissus biflorus, 398 
Nemophila microcalyx, 383, 459 
Note on the Term Phyllary, 501 
Nothoscordum bivalve, 360, 364, 393 
Nymphaea odorata, f. rubra, 400 
Oak, 377 
Oenothera, 366; 
paria, 366, 4 
Oldenlandia, 372; Boscii, 371, 372, 476 
Oplismenus, 389 


tetragona, var. ri- 


Panicum, 369, 372, 381; aciculare, 
376, 386; auburne, 386; ’ensifo lium, 
386; flexile, 378, 386; longifolium, 
var. pubescens, ; ; lucidum, 
376; mundum, 386; sphaerocarpon, 
var. inflatum 386; spretum, 372; 

tenue, 386; Wrightianum, 371, 372, 


eae canadensis, 400; riparia, 


Pas spalum dissectum, 386, 372; flui- 
7, 386 


"482, 463, var. 
; : 786, var. 


Polygala sanguinea, 366, 453; Senega, 
var. latifolia, 378, 453 


515 


Polypremum, 457; procumbens, 457 
Poncirus — ata, 358 
Nut- 


Prasium ?purpur 
Proserpinaea Giteieeia; 373, 454 


us, 451 
Soueala, 366; a 
eglandulosa, 366, 4 
Pycnanthemum Ped anthemoides, 
eet " -wiidifolian, 476: verticilla- 


var. 


475 
Pyle rotundifolia, var. americana, 


55 
Pyrus, 450, 451; betulaefolia, 451; 
1; 


communis, 451; communis X pyri- 
folia, 450; X Lecontei, 450; sero- 
tina 


Quercus, 452 


Ranunculus abortivus, var. indivisus 
358, 401; hispidus, 359; carvitiiets, 
r 


Rhododen 
Rivnchospera: 451; filifolia, 372, 393; 


Rumex, 


Sagittaria Weatherbiana, 367 
Scleria flaccida, 461; oligantha, 366 — 


f, 390, 
392; robustus, 392, 393, 1 protru- 
sus, tl subt erminalis, 390; vali- 


dus, Vv creber, f. megas astachyus, 


392 
Scutellaria serrata, 380, 459 
atum, 360 
Senecio aureus, 361, 495-500, 506, 


500; aureus < tomen 
aureus, var. Ashei, 497-499 
aquilonius, 497, 500-502, : 
~ — var. aquilonius , f. ecoro- 
, 502, v: aurantiacus, ssi 

por "gracilis 496, 
, 496, 498, 499, 502, 


516 INDEX 
508, pl. 802, var. semicordatus, 467, 470, 473, 475, pl. 787, 
496, 500, 502; Balsamitae, 504, var. aspera, 469, var. ‘hispida, 466, 
a; ordii, 467, 469-472, 475, pl. 790, var. 


firmifolius, 604, 
361, 497, 498, 505, 506; Elliottii, 
507; —— 502, 504, 505, v 

firmifolius, 5, gaspensis, f. 


r 

lattensis, 503, 505, 507, 508; 

pseu aureus, 4 f. ecorona- 

pseudo-tomentosus, 507, 

508; Secnlasneaties, 500; Sma li, 

503-505; tomentosus, 361, 497- 
506-508, pl. 800 


oot 
Sium, 367; cieutaefolium, 367, 455; 


floridanum, 368, 455; suave, 
367, 454, 455, f. a onii, 455, 
€ fasciculatum, bg 


Sno e, Sum 359 
Sdluines. 369, 377: ‘patula, 377, 478; 
perlonga, 365, 366; salicina, 377, 


— Varieties and Species of Heleni- 
, 485-495, pls. 796-799 

x ‘Eocbivoaia. 451 

a: opyrus, oe ak 

wa trum Elliottii, 376, 390 

Spec = Characters of Scirpus Olneyi, 


90 
a 369, 473; ; ambigua, 467, 469, 
471-473, 475, pl. 791; 


92, var. arenicola, 474, var. 
tricha, 467, 471, 472, 474, 475, pl. 
793, » 474, var. 


474; Schweinitzii, 474; scopulorum, 
474: sylvatica, 472; tenuifo lia, 465— 


perlonga, 466, 467, 475, pl. 788, 
var. platyphylla, 466-468, 470- 
472, 475, pl. 789; velutina, 471, 7A 

Stellaria media, var. glaberrima, "360, 

400; pubera, 358 
Stewartia siege appt 461 


Sun-flower, Bastard, 486 
Symplocarpus foetidus, 358 


a nudicaulis, 359 
Tephrosi gape 452; 
visgislenk, 3 452, var. glabra, 
3 var. oloations: 452 
65 


eastern Virginia, The Dw 
pls. 772, 773; Laeger 364, 396, 
397, pl. virgini 

397, 308, pl. 713; seats, 384, 396. 


besa ah 358, 398 

Uncin 6 

Uitreulia, 381; biflora, ape Ss 
var. minor, 371, 476; 

purpurea, 371, 476; vee. Bil; 

476 


Mig 451; caesium, 456; Elliot 
y aL, 456; pallidum , 487; sta- 
al eum 456, var. interius, 456, 
var. neglectum, 456; Torre x bivong 
457; vacillans, "457, var. crinitum 
3 vi um, 456 


Vicia caroliniana, 365, 4 


Vinea major, 
Viola eucullate 453; lanceolata, 
a 


358, 
454, var. vittata, 373, 4545 J 
hago 350: preety 383, 4 
Vitis, 
Wisteria frutescens, 461 
Xyris, 371 


— Bebbii, 378, 454; trifoliata, 378, 
54 


0 


I~ 
= 


Plate 


Rhodora 


(cos pus Zoe ‘ONE ‘dd 908) 
VNUGA sry SULPVUIUTLIOUT A]Os|e} pues SNOLOTTR IAL 


eS ee ee 


‘aqdV °O “A “o1ud 


Rhodor: 


Plate 771 


l 


5, 


3 


IGs. < 


EDITORUM: F 


ur. 


mid 2, ok, 2: Vi 


« 
‘ 


I 


FIGS. 


LILIUM CANADENSE: 


Schubert. 


¥ 


B. ¢ 


Photo. 


Rhodora Plate 772 


Tr Mess. pated lee 


Ee er rer eee bie 


go} wey a hawk 


Photo. B. G. Schubert. 


TRILLIUM PUSILLUM, X 1: Fic. 1, Michaux’s Type, after photo. by Cintract; FIGS. 2 and 3, 
modern specimens from ty pe-region, FIG. 2 with short anthers, FIG. 3 with long anthers 


Rhodora Plate 773 


& gy 


Photo. B. G. Schubert. 

gece PUSHLLM var, VIRGINIANUM, TYPE, X 1: FIG. 1 with long anthers, FIG. 
cs short anther 
; bauer < 1: Fic. 3, summit of flowering plant 


Rhodora Plate 774 


Photo. B. G. Schubert. 


Asarum Lewisu, from TyPr-series: Fic. 1, two plants, < 14; FIG. 2, expanded flower, 3 


Rhodora Plate 775 


Photo. B. G. Schubert. 


Asarum Lewisir: Fics. 1 and 2, portions of two flowering plants of TypE-series, X 1 


Rhodora Plate 776 


Photo. B. G. Schubert. 


LEMATIS OCHROLEUCA: FIG. 1, plant, < 24, from Staten Island, N. a3 identified Z 
i gta by Asa Gray with type of C. ovata Pursh; Fic, 2, lower leaf-surface, < 10; FIG 
flower, X 1; FIG. 4, fruiting head, 1; Fic. 5, achene and base of t ail, X r 


Rhodora Plate 777 


< ute 


Photo. B. GQ. Schubert. 


Type of CLEMATIS SERICEA Michx. = C. OCHROLEUCA Ait.: FIG. 1, two o ints, X 14; from 
photograph by Cintract; fig. 2 stem, X 214; FIG. 3, peduncle, x 2; Fa. 4, ti ails of fruit, x 


ee | 


Rhodora Plate 778 


PS ee Se ee ne eee ae ee 


Ne Se ee ee ee we rep ee See 


POE NN a ee ee ee EEN oe 


Photo. B, G. Schubert. 


coe IS VITICAULIS: FIG. l, — ype, X /4; Fig, 2, back and ae of sepal, 

x 18: . 3, fruiting head, x 1; FIG. 4, achene epi base of f tai 

xo fora ‘RIFOLIA: FIG. ee of stem - 10; mi ete leakearthcs, x 10; 
fig. 7, flower, X 1; FIa. 8, fruiting head, x 1 


Rhodore Plate 779 


Photo, B. G. Schubert. 


Ci ATIS ALBICOMA: F1G. 1, portion of mags ne, 1; 2, surface of stem, o. 10; 
FIG. € lowes leaf-surface, 10: Fic. 4, flower, X 1; Fie. 5, poate and base of tail, 


Rhodora Plate 780 


Photo. B. G. Schubert. 


- FIG. 3, 
LEMATIS ALBICOMA, var. COACTILIS: FIG. 1, TYPE, X 3/7; FIG. 2, ang x ie 
lower le rr ore:. x 10; Fic. 4, summit of sepal, < 10; Fia. 5, flower, X 1 ; Fie. 6, tru 
wead, X 1; FIG. rd achene 4 base of tail, x 10 


Rhodora Plate 781 


| 
| 
\ 


Photo. B, G. Schubert. 
CLematis FREMONTII: FIGS. ais leaves, X 1, from TY pE-series from Ellis, Kansas; 
FIG. 4, achene and base of tail, 10 


Rhodora Plate 78 


Photo. B. G. Schubert. 


CLEMATIS FREMONTU: Fic. 1, upper leaf and flower, 1, from Ellis, Kansas; FIG. 
enuded flower, < 1, of var. Riehlii from Alle nton, Missouri; FIG. 3, leaf and young at 
Franklin, Missouri, identified by its author as var. Riehlii 


2, le af $ 
iv 1, fro 


: oe eas 


Rhodora Plate 783 


Photo. B. G. Schubert. 


PHYSOSTEGIA ABORIGINORUM, all figs. from TYPE: FIG. 1, plant, X 14; Fia. 2, flower, 
X 2; nG. 3. fruiting calices, x 5 


Rhodora Plate 784 


Photo. B. G. Schubert. 


PHYSOSTEGIA DENTICULATA: FIG. 1, plant, 
P. INTERMEDIA: FIG. 2, plant, < ! Vy. ; FIG. "3; ts ees x 2 


Rhodora Plate 785 


NEW OD) LNs 
Dire wy 


Photo, B. G. Schubert, 


PRasrum? oe Walt.: rig. 1, leaf, X 1, from oe by S. F. Blake 
Pursogr TEGIA ANGUSTIFOLIA: FIG s. 2-4, Dr ummond, no. 251, X 5/9; Fia. 5, , leaf of 251, 
X 2; Fia. 6, sin of oki x 5/12: Fic. 7, portion of leaf, x 2, from TYPE 


Rhodora Plate 786 


a ee: 


ls 


Photo. B. G. Schubert. 


i leaf 
PHYSOSTEGIA Agena pa var. GRANULOSA: FIG. 1, IsoTYPE, K 14; FIG. 2, portion of 
of ISOTYPE, 3 plants, x (2s from upper Peatiiihee Hivce Maine; FIG. 4, tea, 


x 2, from the tee po , plant, 14, from Lake Champlain 


Rhodora Plate 787 


Photo. B. G. Schubert. 


STACHYS TENUIFOLIA: FIG, 1, plant, X 14; Fia. 2, surface of stem, X 10; FiG. 3, 
low er surface of leaf, < 10; FIG. 4 regu ste flower, 


Rhodora Plate 788 


Photo. B. G. Schubert. 


STACHYS TENUIFOLIA, var. PERLONGA: FIG. 1, IsoTyPE, X 24; FIG. 2, bracts and calices, 


5p 
x 
. HYSSOPIFOLIA; FIG, 3, plant, 14; Fic. 4, bract and calyx, x 4; Fria. 5, nutlet, 10 


a ee ee 


Rhodora Plate 789 


Pag Soe 
taahy fp 


LERBARIUM O§ B 


Photo. B. G. Schubert. 


STACHYS TENUIFOLIA, Var. PLATYPHYLLA: FIG. 1 x 2; Fie. 2, surface of eo. 
x Mi FIG. Ara lower po of leaf, < 10; FIG. 4, hie. bracts ace ‘calices, x 4; FIG. 
nutlet, x 1 


Rhodora Plate 790 


Photo. B. G. Schubert. 


STACHYS TENUIFOLIA, Var. HISPIDA: FIG. 1, plant, X ! 2, por wr of stem, X 10; 
FIG. 3, lower surface of leaf, « 10; Fra. 4, Re ke mn st 


i 


Rhodora Plate 791 


ee ee 
Photo. B. G. Schubert, 


. RT ACiCrS AMBIGUA: FIG, 1, summit of plant, x 14; FIG. 2, portion of stem, X 10; FIG. 3, 
alices, X 4 


> , 706 
Rhodora Plate 792 


Photo. B. G. Schubert. 


STACHYS PALUSTRIS: FIG. 


1, portion of stem, X 10; Fic. 2, lower leaf-surface, 10; 
FIG. 3, bracts and flowers, x 4; FIG. 4 x 10 


, nutlet 


Rhodora Plate 793 


Photo. B. G. Schubert. 


S of TYPE-specimens, X 
FIG. 2, lower reeset and calyx, < 4, from TYPE; FIG. 3, stem and lower sirace, of 
leaf-base, X 10, from Typx; Fic. 4, narrow-leaved plant (S. arenicola Britt.), * 72; 

- 5, calyx of his es «x 10 


J ao . 
TACHYS PALUSTRIS, var, HOMOTRICHA. FIG. L, one 


© 


— 


)4 


Rhodora Plate 


: 
: 
7 
. 


a a eT es eee ee ee 


0. +e Area 


Photo. B. G, Schubert. 


Vo; 


STACHYS PALUSTRIS, var. PHANEROPODA, % figs. from Type: FIG. 1, plant X 
FIG. 2, surface of stem, X 4; FIG. 3, calyx, X 10 


Rhodora Plate 79 


fea | 


ype 
L : es op ra 
pa feref renters Wun - 


PLANTS OF LOUISIANA | 


DS. and H.B.Correll 


poate babies. ery 


S8PUuo-Oah gard 
» north aa f Pineville: 


Collected in coaperation hetween the 
Kotanical Muscam, ard Universi 


Arnold Arboretum and the 
ity and the Department of 
Hotany, Duke University. _ 


Photo. B. G. Schubert. 


ay 
GNAPHALIUM PEREGRINUM, TYPE, X 7% 


Rhodora 


Plate 796 


Photo. B. G. Schubert. 


LENIUM AUTUMNALE: FIGS 
Linnean Society of London; Fria 


. land 2, portions of rrpr, x 1, from photograph sent em 
e : : 3 : a ( 
. 3, flowering summit, x 1, of modern specimen from UI 


Rhodors Plate 797 


‘Om M7 7 M7 ? . 
QY Ti: WZ 


ant, 


WANG INT 


Photo. B. G. Schubert. 


HELENIUM AUTUN ACULATUM, all hii CS eee eee 8): Yee. 2 
portion oF Flat of sents Cliffortianus courtesy 0 John Ramsbottom; Fic. 2, summit 
of wild specimen from Quebec; FIG. 3, portion of ceginal plate of H. canaliculatum; FIGS. 4 
and 5, portions of Cornut pla 


| 
| 
| 
| 
| 


Rhodora Plate 798 


Photo. B. G. Schubert, 


— 


HELENIUM AUTUMNALE, var. PARVIFLORUM, both figs. X 1: Fic. 1, portion of TYPE or 
IsoryPE of H. PARVIFLORUM Nutt.; rie. 2, summit of ae specimen from Virginia 


Rhodora Plate 799 


Photo, B. G. Schubert. 


HELENIUM GopFREYI: FIG, 1, Type, X 14; Fic. 2, achene, X 10 
H. FLORIDANUM: FIG. 3, portion of TyPE, X 1; Fic. 4, achene, X 10 
H, NUDIFLORUM: FIG. 5, achene, X 10 


Rhodora Plate 800 


Tod Wig Q Ay 
sa plonecee Cormfles andane™ 
= : aa le > : 2 


Photo. B. G. Schubert. 


ean 
SENECIO AUREUS: FIG. 1, ae of trPe, <1, ‘oe photograph sent from Linn 
Society of London; PIG. 2, achene, X 10, fro m Virgi 

S. TOMENTOSUS: achene, 16, from Vinuinia 


Se ae 


Rhodora Plate 801 


Photo, B. G. Schubert. 


ENECIO AUREUS, details from modern specimens — Virginia: Fic. 1, rhizome and 
i 


bs asal offshoot, x 1; FIG. 2, portion of full-grown gud eaf, FIG, 3, expanding ros 
basal leaves, showi ing archnoid pubescence, 2; Fic. 4, unexpanded heads, showing 
arachnoid pubescence, xX 2; Fira. 5, involucre, X 9 


Rhodora Plate 802 


Photo, B. G. Schubert. 


7 
SENECIO 4 var. INTERCURSUS, all figs. from TYPE: FIG. 1, rhizome and ag 
shoots, X 1; ey basal and lower cauline leaves, X 1, FIG. 3, involucre, X 95; FIG- 
inflorescence, XI 


| Plate 803 
Rhodora 


Photo. B. G. Schubert 


: ate rhizome 
SENECIO AUREUS, var. AQUILONTUS, all pit ge aol ace: opcelayr yg rescence 
and basal offshoot; Fra. 2, basal leaves; Fic. 3, cauline le¢ 


Rhodora 


Plate 804 


E> Cate e240 vas e | bem 
(ji Ay fe 
I pete it € : “ Zz 


, ad cf Dau 
oe 


Photo. B. G. Schubert. 


ECIO PAUPERCULUS: FIG. 1, TYPE, X 34, from photograph by M. L. Fernald; FIG 
d 


oe tall plants, X 1, from ’Newfoundlan 


Ss. 9H, 


ii 


Rhodora Plate 805 


[Tae 


Photo. B. G. Schubert. 


ENECIO PAUPERCULUS, var. BALSAMITAE! FIG. ype BatsaMATAE Muhl., x 1, 
kindness of ix res Ludwig Diels; ries. 2, 3 and 4 scot ‘of a modern specimen from 
type-region, ; Fic. 5, involucre, x 5, from latter specimen 


Rhodora 


Plate 806 


x 


Photo, B. G. Schubert. 


0 PAUPERCULUS, var. NEOSCOTICUS, all figs. from TYPE: FIGS. 1 and 2, base, 


SEN 
pa sit faa and inflorescence, 


X 1; FIG. 3, involucre, 


‘ — iia. 


7 

4 ; Reprinted from Ruopora, Vol. 46, January and February, 1944 

ef 

iq 

rq 

ie : CONTRIBUTIONS FROM THE GRAY HERBARIUM 
ae OF HARVARD UNIVERSITY 


OVERLOOKED SPECIES, TRANSFERS AND 
NOVELTIES IN THE FLORA OF 
EASTERN NORTH AMERICA 


M. L. FERNALD 


Dares or IssuE = 
Pages 1-21 and Plates 807-811. ./..........- fe aa Tne 22 January, 19 
Pages 32-57 and Plates 812-816... . .. ere >= .12 February, 1944 


Reprinted from Ruopora, Vol. 46, January and February, 1944 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


CL 


OVERLOOKED SPECIES, TRANSFERS AND 
NOVELTIES IN THE FLORA OF 
EASTERN NORTH AMERICA 


M. L. FERNALD 


Dates oF Issun is cay 100 
Pages 1-21 ond Plates 807-Gl1 2. 66. 
Pages 32-57 and Plates 312-816 igh ee ee os 12 Sakeuary, 1944 


SSSA E I ARs HH ATI EATS al SRE adhe a EL Sp IO ee ot 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY—NO. CL 


OVERLOOKED SPECIES, TRANSFERS AND 
NOVELTIES IN THE FLORA OF 
EASTERN NORTH AMERICA — 


M. L. FERNALD 
(Plates 807-816) 


DuRING a restudy of the Liliaceae of the Gray’s Manual area 
I was soon confronted by the many species proposed by Rafin- 
esque in one of his relatively careful and trustworthy works, 
Autikon Botanikon (1840), a book so rare, until the offset of it 
issued by the Arnold Arboretum, that only casual names from 
it (and those picked up through other sources) were caught by 
the editors of Index Kewensis, Dr. Merrill is engaged on a 
bibliographic study of great extent, dealing with this and other 
neglected works of Rafinesque. It is, consequently, not appro- 
priate here to go into further detail than to note a few species 
in Autikon Botanikon which have immediately to be taken into 
account in my studies. Very soon I found Rafinesque referring 
to species of Bartram, and on checking Bartram’s Travels (1791) 
the same situation was found: only casual names of species by 
Bartram were entered in Index Kewensis, these picked up from 
the writings of others who had cited them. Accordingly, it was 
necessary to study Bartram’s Travels, with such limited knowl- 
edge of the flora of the southeastern United States as I have. 
The present Contribution, therefore, is made up partly of con- 
sideration of some of the species of Bartram, some in Rafinesque’s 


2 Rhodora [JANUARY 


Autikon Botanikon, some based on study of photographs of 
types of Linnaeus in the Liliaceae, and the usual miscellany 
which has accumulated in recent months. The items are ar- 
ranged in systematic sequence. 

Panicum (sub-§ Cah cite Benneri, sp. nov. (TAB. 807), 
planta cespitosa 1.7-3.5 dm. alta; culmis firmis erectis basi 
1-1.5 mm. diametro, siiteenitae ad 7.5 cm. longis pilosis pilis 
adscendentibus; nodis barbatis; foliis lanceolatis firmis valde 
adscendentibus glabris vel subtus sparsissime breviterque pilosis 
ae 5-6.5 cm. longis 5-8 mm, latis 36-42-nerviis basi villoso- 

atis, vaginis subhorizontaliter hirsutis pilis 1-1.5 mm. longis 
ee bullatis, ligulis 2-3 mm. longis; paniculis primariis breviter 
exsertis ellipsoideo-ovoideis 2.5-6 cm. longis 1.5-4 cm. diametro, 
rhachi imo hirtello sparsissime villosoque, ramis patento-adscen- 
dentibus ramulis subsimplicibus, pedicellis 2-6 mm. longis 
glabris; spiculis pubescentibus ellipsoideis basi apiceque obtusis 
2.2-2.6 mm. longis 1.2—1.4 mm. latis, gluma inferiore perbrevi 
deltoideo-rotundata subapiculata 0. 5-0.8 mm. longa, superiore 
lemmateque sterile aequilongis valde costatis fructus lucidos 
subaequantibus.—New Jersey: old field along Delaware River, 
about 114 miles east of Raven Rock, Hunterdon County, June 7, 
1941, Walter yi Benner, no. 9635, TypE in Herb. Gray.; ISOTYPE 
in Herb. Phil. Acad. 

By the treatments of Hitchcock and Chase, of Hitchcock’s 
Manual and my own manuscript-treatment Panicum Benneri, 
traced through the keys, comes directly to P. villosissimum EI. 
Its very stiff habit, glabrous or essentially glabrous, short firm 
leaves, the very short and broad first glume of the spikelet, 
short-peduncled panicle, only slight (instead of usually pro- 
nounced) development of branches at the time of primary 
anthesis, and the relatively short and strongly pustular-based 
pubescence of the sheaths at once distinguish it. In P. villosis- 
simum the culms are 2-7 dm. high and the leaf-blades of the 
vernal culms 6-11 cm. long, and 5-10 mm. wide, more or less 
pubescent on both faces; in P. Benneri the stiffly erect culms are 
1.7-3.5 dm. high, the leaf-blades more lanceolate, 3.5-6.5 cm. 
long and 5-8 mm. wide, the upper surface hard and glabrous, 
the lower barely and minutely pilose, while the strongly pustular- 
based hairs of the sheaths of P. Benneri are shorter than the 
most often slender and more villous sheath-hairs of P. villosis- 
simum. As above noted, P. villosissimum, while the vernal or 
primary panicles are intact, usually has well developed axillary 


SEN Ree 


eT Bee eee Oe ee a i ES a eS a oe ee Sor 
” pa . er 


ris ie 4d ASRS a lak — "Se ne aes a a 
ed 


1944] Fernald,—Flora of Eastern North America 3 


branches; in P. Bennert only a few, and those very short, such 
branches are developed. In P. villosissimum the fully developed 
primary panicles are 4-10 cm. long and on peduncles 0.3-1.7 dm. 
long; in P. Benneri the primary panicles are 2.5-6 cm. long and 
on peduncles only 0.5—-4 em. long. The first glume of P. villosis- 
stmum is ovate, one fourth to two fifths as long as the spikelet; 
in P, Benneri broadly deltoid-rotund and one fifth to barely one 
fourth the length of the spikelet. 

In its very short and broad first glume the spikelet of Panicum 
Benneri suggests some variations of the heteromorphic P. 
lanuginosum, such as var. fasciculatum (Torr.) Fern. (P. tennes- 
seense Ashe) or var. septentrionale Fern.; but its spikelets are ~ 
altogether too large and the panicle too simply branched. In 
P. lanuginosum and its varieties the secondary branches of the 
panicle are themselves branched (whence P. implicatum Scribn.). 

The once-forked branching of the panicle of P. Benneri is 
similar to that of P. meridionale (with spikelets 1.3-1.6, instead of 
2.2-2.6 mm. long). It also suggests the panicle of the relatively 
northern P. subvillosum Ashe, but that characteristic species has 
the leaf-sheaths mostly equaling or exceeding, instead of shorter 
than the internodes, and appressed-pilose, instead of divergently 
hirsute with pustular-based hairs. Furthermore the axis of the 
panicle of P. subvillosum is appressed-pilose to glabrate, in P. 
Benneri divergently hirtellous, with some horizontally spreading 
villi; the spikelets of P. subvillosum are 1.8-2 mm. long, 0.9 mm. 
broad, and the prolonged first glume is one third to half the 
length of the spikelet. 

Panicum Benneri, with which it is a privilege to associate the 
name of the keen and always helpful student of his local flora, 
Water MackiInnett BENNER, its discoverer, seems to be a 
real species. Whether it is relatively northern, as its type- 
region suggests, we do not know. It is more likely so, and to be 
sought in northeastern Pennsylvania and southern New York, 
than an extension from farther south in New Jersey and Pennsyl- 
vania where the small army of keen explorers would long ago 
have found it. Mr. Benner has most generously defrayed the 
expense of PLATE 807. 

In Piate 807, Frias. 1 and 2 are portions of f the Type of Panicum BENNERI, 


hel yet cede and cauline sheath, X 3; Fic. 4, axis of panicle, < 10; 
5, spikelet, 


4 Rhodora [JANUARY 


XYRIS ALBIFLORA Raf. Aut. Bot. 190 (1840). XX. torta pal- 
lescens C. Mohr, Contrib. U. S. Nat. Herb. vi. (Pl. Life Al eal 
428 (1901). X. pallescens (C. Mohr) Small, Fl. Se. U. S. 234 and 
1328 (1903). 

This distinctive iitetoweted species of northwestern 
Florida and Alabama is related to X. fleruosa Muhl. and X. 
fimbriata Ell. in having exserted and fimbriate sepal-tips. It 
was well characterized by Rafinesque, who recognized its 
affinity with the latter species: 

1420, X. or Jurica? albiflora Raf. (X. cylindrica Baldw. mpt.) fol. 
strictis ensatis lato-planis striatis, scapis duplo longior teres anceps. 
suleatis, capitulis ellipt. subcyl. obt. bract. dilatatis mecca vel oe. 

—Florida, disc. by Baldw. in 1815, not yet described, unless X. fimbri 
of Elliot be very badly so, and appears rather an akin sp. leaves sale 
seape bipedal. fl. white by a note of Baldwin, while all others are yellow. 

X. FistuLosa Raf. 1. c. (1840), based upon X. juncea Baldw., 
not R. Br., is X. Batpwiniana Schultes, Mant. (1822). His 
X. rETuSA, |. c. 190 and X. sprratis, l. c., are apparently X. 
TORTA J. E. Sm. 

LuzuLA ACUMINATA Raf. Aut. Bot. 193 (1840). L. saltuensis 
Fernald in Ruopora, v. 195 (1903). Juncoides pilosum (L.) 
page= var. saltuense (Fern.) Farwell in Mich. Acad. Sci. Re 
xx. 170 (1918). LL. carolinae S. Wats., var. saltuensis (Fern.) 
ae pe Ruopora, xl. 404 (1938). 

Luzula saltwensis of woodland and thicket from Newfound- 
land to Saskatchewan, south into the Northern States and locally 
southward, was separated from the cespitose Eurasian L. pilosa 
(L.) Willd. because of its looser habit, with elongate more or 
less repent basal offsets, its paler and acuminate sepals and its 
more pointed capsules. Later (in 1938) I reduced it to varietal 
rank under the poorly typified L. carolinae S. Wats. in Proc. Am. 
Acad. xiv. 302 (1879), the latter southeastern plant being larger 
and with more forking branches to the corymb. It now seems 
that Rafinesque was nearly 40 years ahead of Watson and more 
than 60 ahead of me. There can be no doubt that his L. acumi- 
nata was L. saltuensis. His definition of it was good: 


1447, Luz. acuminata Raf. repens, gebra: fol. lanceol. acum. striatis 
nervosis _— corymbosis congestis paucifloris fuscatis, calic. acum.— 

real America, perhaps J. pilosus Mg. often blended with last, leaves 
cg and shorter, 3 uncial, stem semipedal, corymb. not exceeding the 
eaves . 


1 


os eer erem siecle ene ener " " , = ss . ere is . ‘ ers i a i i P 


Sn gt Od Ca eae eS Pe ee ee 


1944] Fernald,—Flora of Eastern North America 5 


The taking up of Luzula acuminata (1840) necessitates the 
following combination 

L. acuminata Raf., var. carolinae (S. Wats.), comb. 

L. carolinae S. Wats. in Proc. Am. Acad. xiv. 302 (1879). Funco- 
ie i Juncodes) carolinae (S. Wats.) Ktze. Rev. Gen. ii. 724 

LUZULA LABRADORICA Raf. I. c. (1840) was unquestionably L. 
PARVIFLORA (Ehrh.) Desv. (1808) or its var. melanocarpa 
(Michx.) Buchenau, which latter rests on Juncus melanocarpus 
Michx. (1803). Rafinesque’s name antedates L. labradorica 
Steud. Syn. Pl. Gram. ii. 291 (1855), which is perhaps identical 
with it. 

Rafinesque proposed nine North American species of the pro- 
phyllate Junci. Nothing sufficiently distinctive was given 
in his descriptions for us to be certain what he had. His J. 
FLORIDANUS, I. c. 194, was from “South Florida” and Engelmann 
in Trans. St. Louis Acad. Sci. ii. 451 (1866) cited as J. Gerardi 
Loisel. (1810) material from ‘North Carolina, Curtis, and 
Florida, Ware (J. Floridanus, Raf. in Hb. Durand)”. This 
would seem to dispose of J. floridanus. It also gives the clue, 
that others of Rafinesque’s Junct may have been in Durand’s 
herbarium, now in Paris. Rafinesque’s J. FuscaTus, l. c. 194, 
from “Kentucky, Tennessee, &c”, with “fl. quite peculiar’, 
was earlier than J. fuscatus Turcz. ex Ledeb. (1853), the latter 
reduced by Buchenau to J. triglumis L. J. fuscatus Raf. is not 
clearly identified. So with his other names; they may rest 
undisturbed. 


Tue IpenTITY oF YUCCA FILAMENTOSA (PLATES 808 and 809). 
—Yucca, always a baffling genus to work with from herbarium 
material, has one species, Y. filamentosa L., which is reputed to 
follow much of the Coastal Plain from Louisiana to Florida and 
northward to North Carolina, with a related plant, Y. concava 
Haw. or Y. filamentosa, subforma latifolia Engelm., on coastwise 
sands from Georgia to New Jersey. The tendency has been to 
follow the conclusions of Engelmann in Trans. St. Louis Acad. 
Sci. iii. 17-54 (1873). In that study, treating Y. filamentosa as 
& complex species, some of the varieties native, others only in 
cultivation, Engelmann defined what he considered true ar; 

lamentosa, his forma genuina, with two subforms: “a, angustz- 


6 Rhodora [JANUARY 


folia’”’, a plant with linear-lanceolate leaves gradually attenuate 
from the middle; and b, subforma latifolia, = Y. concava Haw., 
with broad and rigid lanceolate or spatulate leaves obtusely 
mucronate and cucullate at tip. He concluded that the brief 
diagnosis quoted by Linnaeus from Gronovius, ‘‘foliis lanceolatis 
acuminatis together with the Hab. Virginia, points to the narrow- 
leaved form of what I have described as the genuine plant, as the 
one he and Gronovius had in view. Of this and other forms 
numerous specimens and full notes have been obtained from Dr. 
Mellichamp, of South Carolina, on which the following descrip- 
tions are based.” Engelmann then went on to discuss the 
“narrow-leaved form” and the “broad-leaved variety’’, which 
in his Latin account were both subordinate to his ‘Forma 
genuina’”’. In 1880 Baker, Journ. Linn. Soc. Bot. xviii. 227, 
228 (1880), split Y. filamentosa into ten varieties, many of them 
cultivated only, with the broad natural range, ‘‘ Regiones littor- 
ales Americae borealis a Maryland ad Floridam’’, treating Y. 
concava Haworth as Y. filamentosa, “‘var. Y. concava’’ and not 
citing Y. filamentosa, forma genuina, subf. latifolia Engelm. 
under it. Subsequently, Trelease, Mo. Bot. Gard. 13th Ann. 
Rep. 46—49 (1902), accepting Engelmann’s and Baker’s divisions 
in the main, likewise made typical Y. filamentosa the plant with 
“Leaves 25-40 cm. wide, gradually acute, rather rigid 
Capsules rather narrowly cylindric’, this plant shown in a 
photograph (his pl. 8, fig. 1) with narrowly linear-lanceolate 
and long, attenuate leaves, while its capsule (his pl. 12, fig. 1) 
was shown as 4-5 em. long. This plant, taken by Engelmann, 
Baker, Trelease and subsequent authors as true Y. filamentosa, 
was assigned the range: ‘‘ West-central North Carolina to south- 
eastern South Carolina, Florida from Jacksonville to Tampa, 
and doubtless in the intervening country’. The plant with 
shorter, broader, more oblong-oblanceolate to spatulate and, in 
maturity, harshly scabrous heavy leaves, the Y. filamentosa, 
forma genuina, subf. latifolia Engelm., was taken up as var. 
concava (Haworth) Baker and a good portrait of a growing plant 
given (Trelease, pl. 10), showing the very characteristic and 
short, lanceolate, broad leaves, this very distinct plant given 
a range from South Carolina and Georgia to Maryland. 

It remained for Small, Man. 303 (1933) to see stronger differ- 


1944] Fernald,—Flora of Eastern North America 7 


ences. Still adhering to the plant with ‘ Leaf-blades of a linear 
type, somewhat narrowed toward both ends, attenuate to the 
slender apical spine, flat’ as Y. filamentosa, with the range 
“Fla. to Miss., Tenn. and N. C.”’, this plant with “petals broadly 
ovate, 4-5 cm. long: capsule 5-6 cm. long’’, he correctly recog- 
nized another species, as Y. concava, with ‘‘ Leaf-blades spatulate, 
abruptly narrowed or rounded and concave at the base of the 
stout apical spine’’, the species occurring from ‘Ga. to Del.” 
and having “‘sepals and petals usually broader” than in the former 
and the capsule only 4—5 cm. long, the seeds rounder. 

Ever since our first trip together to Virginia, Long and I have 
become very familiar there with the plant we have regularly 
called Yucca concava, following the treatment of Small. It 
occurs back of the outer beaches and among the dunes, and in 
sandy fields, roadsides and dry pineland back from the coast at 
least 80 miles (to the easternmost border of Dinwiddie County). 
Baker’s ‘‘Regiones littorales’’ tells only part of the story. 
Throughout all this region of eastern Virginia, thence northward, 
the plant is constant in foliage, flowers and fruit, the plant 
beautifully illustrated in Sims, Bot. Mag. xxiii. pl. 900 (1806) as 
Y. filamentosa, its habit shown by Trelease, 1. c. pl. 10. In this 
plant (our PLATE 808) the flowers are 5-7 cm. long, the petals 
2-3 cm. broad and rounded to the short acumination, the fila- 
ments spiculate-papillate in irregular lines, the style in anthesis 
about 1 cm. long, the capsule thick-cylindrie to short-ovoid, 
inclined to be constricted at or near the middle and dumbbell- 
like, 1.5-4.5 em. long; the semiorbicular seeds 6-7 mm. long by 
3-5.5 mm. broad. 

In Yucca filamentosa sensu Engelmann, Baker, Trelease and 
others, including Small, who seems first to have recognized the 
best specific characters, the southern plant (our PLATE 809) with 
linear-lanceolate long-tapering leaves, the flowers are only 3-5 
cm. long, the petals 1-2 em. broad and tapering to gradually 
acuminate tips, the filaments nearly pilose with elongate tri- 
chomes (especially at base), the style at flowering time nearly 
obsolete or up to only 5 mm. long, the more uniformly subterete 
capsule 4-6 em. long. 

When Engelmann assumed, because he had material from 
South Carolina of the latter plant, that it must, therefore, be the 


8 Rhodora [JANUARY 


Virginian plant, “In littoribus arenosis fluminum crescit’’, of 
Clayton, which is the type of Yucca filamentosa L., he was at 
least naive. Not all plants of South Carolina and Virginia are 
identical. In exploring the southeastern counties of Virginia 
Mr. Long and I have often noticed that farmers frequently set 
young plants of their native Yucca along roadways and in sandy 
clearings. Our driver for some seasons, a farmer of keen intelli- 
gence, Leonard Birdsall, explained that they harvest the hard 
leaves as ‘‘SruKGrass’’, and after macerating them and softening 
the tissue, remove the strong threads for use in tying bunched 
vegetables. This plant, Y. concava, being the abundant and, 
so far as we know, the only native species of the genus in eastern 
Virginia, it is not surprising that the specimen preserved at the 
British Museum (photograph sent by Dr. Ramssorrom) should 
show the characteristic flowers (PLATE 808, FIG. 1)—the two 
smallish ones at the tips of branches—of typical Y. concava, nor 
that Clayton’s no. 720 (miscopied by Gronovius as 270), the 
TYPE of Y. filamentosa, should have been labeled by him: “ Yucca 
flore albo, foliorum marginibus fllamentosis. Silkgrass.”’ 

The type of Yucca filamentosa L. Sp. Pl. 319 (1753) being the 
relatively northern Y. concava, it is necessary to find the proper 
specific name for Y. filamentosa sensu Small, the Y. filamentosa, 
forma genuina, subf. angustifolia of Engelmann. Study of the 
bibliographies of Engelmann, Baker and Trelease yielding none, 
I venture to name the narrow- and smoother-leaved southern 
plant for the botanist who first saw its specific differences. It 
is a pleasure so to do; it is not always that I can follow him. I 
am ces ae 

Yuc ana, sp. nov. Y. filamentosa sensu Small, Man. 
Se. Fl. 08 "Ghee not » Y. filamentosa, forma genuina, subf. 
angustifolia Engelm. i in Trans. St. Louis Acad. Sci. iii. 51 (1873), 
not Y. angustifolia a (1814). Typ: sandy soil near J ae 
ville, eee May, A. H. Curtiss, no. 2950, in Herb. Gra 

TE 

Although Small states that the smaller-flowered and more 
southern Yucca Smalliana (Y. filamentosa sensu Small, not L.) 
has “‘panicle-branches glabrous’’, the material before me shows 
them merely glabrescent. At flowering time they are pruinose- 
pilose, only in fruit becoming glabrate. True northern Y. fila- 
mentosa has the panicle glabrous from the first. In its pubescent 


1944] Fernald,—Flora of Eastern North America 9 


panicle Y. Smalliana suggests the upland Y. flaccida Haw., of 
the Blue Ridge and adjacent uplands, which may have pubescent 
or glabrescent panicle; but, as I understand it, Y. flaccida has 
very pliable and thin leaves, very broad and abruptly short- 
acuminate petals (as shown in Lindl. Bot. Reg. xx. t. 1895 
(1836) ), and the native specimens which seem to belong to. it 
have the style elongate and the broad and flat filaments coarsely 
ciliate. In Y. Smalliana (pu. 809) the flower is smaller, with 
much narrower and acuminate sepals and petals, the style 
obsolete or very short, and the filaments less flattened and finely 
pruinose-pilose. 

PLaTE 808 is of Yucca FILAMENTOSA L.: Fig. 1, terminal flowers, x 1, fro 
the TYPE Pag photograph sent by Dr. John Ramsbottom) ; F1a. 2, aatetas 
leaves, X 14, from Old Town Neck, Northampton County, Virginia, F 
& Fogg, no. 5269; Fics. 3 and 4, flowers, X 1, from Cape Henry, Virginia, 

Fernald, Griscom & Long, n o. 4707; Fia. 5, ee style and filaments, X 3, 
from no. 4707; FIG. 6, ripe poet pee from wd 4 of South Quay, Virginia, 

ernald & Long, no, 10,585. 

Piate 809, ae ig SMALLIANA Fernald: ric. 1, leaves and flowers, X 4, 
from TYPE; FIG. 2, basal rosette, greatly rediced, — lr photo. by 

. H. Curtiss, 1887 FIG¢s. 3 and 4 , flowers, xX 1, fro Gc. 5, ovary, ai be 
and filaments, X 3, from TYPE; FIG. 6, capsule, x1, ya as Weel, A. H. Curt 


Sreciric DIsTINCTIONS BETWEEN POLYGONATUM BIFLORUM 
AND P. CANALICULATUM.—More than a decade ago, while study- 
ing with me, Dr. W. A. Anderson, Jr. clearly worked out the 
proper nomenclature and the specific characters of our American 
species of Polygonatum and, although he published! his study of 
Trillium in Tennessee, he has, unfortunately, not stated in print 
his conclusions regarding Polygonatum. In the Manual range 
we have three indigenous species, P. pubescens (Willd.) Pursh, 
quickly distinguished by having superficial rhizomes, minutely 
pilose or hirtellous lower leaf-surfaces, lowest peduncle usually 
from the Ist or 2nd leaf-axil, perianth 7-13 mm. long, with 
stamens inserted high on the tube; and P. biflorum (Walt.) Ell. 
and P. canaliculatum (Muhl.) Pursh (P. commutatum (R. & 8.) 
Dietr. and P. giganteum Dietr.), formerly needlessly confused 
and, judging from recent identifications, not usually understood, 
two very different species with deep-seated rhizomes, glabrous 
lower leaf-surfaces, lowest peduncle usually from the 3d—5th 
leaf-axil, perianth 1-2 cm. long, stamens inserted near middle of 


1 'W. A. Anderson, Notes on the Flora of Tennessee: the Genus Trillium. Rxopora, 
Xxxvi. 119-128 (1934). 


10 Rhodora [JANUARY 


the tube. Dr. Anderson, with the collaboration of Mr. C. A. 
Weatherby, found so many distinctive characters and in my 
own detailed study so many others become evident that it may 
be helpful to others to have these distinctions pointed out. The 
usual failure clearly to separate the two species, P. biflorum and 
P. canaliculatum, is reflected in many recent local floras. Thus 
Wiegand & Eames in their very helpful and usually keenly dis- 
criminating Flora of the Cayuga Lake Basin give two types of 
habitats for their inclusive P. biflorum: ‘Sandy or gravelly, 
rarely clayey, banks and thickets, in subneutral soil, on dry hill- 
sides and hilltops, or in alluvial calcareous soils on river banks’’. 
Several of the collections cited by them are represented in the 
Gray Herbarium. Those from ‘Sandy or gravelly 

banks’’, ete. are characteristic P. biflorum: “dry ravine Banik 
between Renovick and McKinney’s’’, etc., these plants having 
the characteristic slender rhizome, slander stem only 6-7 dm. 
high, flat and relatively few-nerved, merely sessile leaves, lowest 
peduncle from the 3d leaf-axil and 1.2-2 em. long, with the 2 or 
3 pedicels 0.5-1.4 cm. long, and slender perianth with lobes 3.5 
mm. long. The material from “alluvial calcareous soils” is 
characteristic P. canaliculatum: “Valley of Inlet, Ithaca’, etc., 
the plants with rhizome more than 2 cm. thick, stout stem 
(1 em. thick) 2 m. high, with clasping-based corrugated and 
undulate-margined leaves with about 200 nerves, the flowering 
peduncles up to 9 cm. long, with 4~7 pedicels up to 2.5 cm. long, 
the thicker perianth with lobes 6 mm. long. Again, in that 
compendium of accurate local field-observation, Deam’s Flora 
of Indiana, the author, beautifully distinguishing the pubescent- 
leaved P. pubescens, with superficial rhizome, from the glabrous- 
leaved plants with deep-seated rhizomes, which he unwillingly 
treated as one species, P. biflorum, wrote: ‘‘My study of this 
species complex was made from 155 specimens which I have col- 
lected from all parts of the state. I am not satisfied with the 
treatment of this species but I am not able to find differences 
sufficient to separate them . . . I donot think them all the 
Same species . . . The genus has been monographed by 
three authors and my specimens have been seen by one of them 
but I cannot accept their treatment of this complex.” Such 
observations indicate the need of better statements of the char- 


1944] Fernald,—Flora of Eastern North America ll 


acters, and the misidentifications in the herbaria (including our 
recent collections from Virginia as identified by a student of the 
group) also indicate such a need. 
Very briefly my conclusions cota 
POLYGONATUM BIFLORUM (Walt.) Ell. Rhizome 0.6-1.5 cm. 
thick; stem slender, 1.5-5 mm. thick pale: lowest leaf, 2-9 dm. 
high; leaves flat, sessile or nearly so, narrowly lanceolate to 
broadly ovate, the largest ones (of each plant) with 46-120 
nerves and 5.5-15 cm. long by 1.2-6 cm. broad, the terminal 
small ones 20—66-nerved; te a 1-4 cm. long, 1-3 (-5)- 
flowered, the lowest usually borne from the 3d (l1st—5th) axil; 
pedicels ‘becoming 0.5-2 cm. long; perianth slenderly cylindric, 
1-1.7 cm. long, its lobes 3-4 mm. long; filaments commonly 
papillate or granulose, slender; enlarged terminal joint of fruiting 
pedicel cupshaped or campanulate, with the rim flaring, 0.7-1.5 
mm. long, often as broad; seeds 2.7-3.5 mm. long.—Dry to moist, 
sandy, loamy or rocky woods and thickets, Florida to Texas, 
north to Connecticut, New York, southern Ontario, southern 
Michigan, Illinois, Iowa and Nebraska. 

. CANALICULATUM (Muhl.) Pursh. Rhizome 1.5-3 em. thick; 
stem stout, 0.5-1.3 em. thick at lowest leaves, 0.6-2 m. high; 
leaves more or less corrugated and with puckered margin (not 
drying flat), mostly narrowed to broad clasping or sheathing sub- 
petiolar bases, the larger ones with 110-220 nerves and 0.9-2.5 
dm. long by 3.5-13 cm. broad, the smallest eager ones 58- 
112-nerved; peduncles becoming 1.5-9 cm. long, 2—10-flowered, 
the lowest commonly sey from the 4th or 5th (era-Si axl 
perianth thick-cylindric, 1.7-2 cm. long, its lobes 5-6.5 m 
ong; filaments broad, smooth or merely granulose; i 
terminal joint of fruiting pedicel subcylindric to slenderly 
campanulate (except at flaring summit), 1-3 mm. long, usually 
longer than thick; seeds 3-4.5 mm. long.—Rich woods, alluvial 
thickets, river-silts and other calcareous habitats, Connecticut 
Valley of New Hampshire to southern Manitoba, south to 
South Carolina, Tennessee, Missouri and Oklahom 

Many reputed species and varieties have been eet based 
upon shade of color, breadth of leaf, etc. These I am not here 
discussing. That is for a monographer who has studied them 
all. So far as I have seen they do not affect the fundamental 
specific characters of the species long ago defined. In fact, I 
have recently been challenged to point out any real characters 
distinguishing P. canaliculatum from P. biflorum. The above 
is my answer. What better characters could be found in the 
Liliaceae? 


12 Rhodora [JANUARY 


The position of the lowest peduncle, a splendid character 
pointed out by Deam, is certainly significant. I have noted its 
position in all specimens in the Gray Herbarium and that of the 
New England Botanical Club. The results follow. 

P. pusescens: of 365 plants 100 (27+ per cent.) have the lowest 
peduncle from the 1st axil, 241 (66+ per cent.) from the 2nd, 20 from the 
3rd and only 1 from the 4th. 

P. BrrLorum: of 116 plants 3 have hee lowest peduncle from the Ist 
axil, 12 (10 per cent.) Ree the 2nd, 6 ee per cent.) from the 38rd, 28 
(244 per cent.) from the 4th, and 5 rae ‘ 

P. CANA uM: of 41 ee 4 (10 per cent.) have the lowest 
bec Me from the 8rd axil, 17 (4 be per cent.) from the 4th, 13 (31+ De 


A biologically ae ae because Dee from elong- 
ate leafy axillary branches, of Polygonatum biflorum is 

P. BIFLORUM, forma ramosum (McGivney), comb. nov. 
commutatum, forma ramosum McGivney in Am. Midl. Nat. ix. 
664, fig. (1925). 


Tur INDIGENOUS ALLEGHENIAN CoNVALLARIA.—The native 
Lily-of-the-Valley, occurring in acid rocky or sandy woods, 
summits and upper ravines of the mountains from Virginia and 
eastern West Virginia to northern Georgia and eastern Tennessee, 
is much larger than the European Convallaria majalis L., the 
plant so generally cultivated and naturalized near settlements; 
and, as Professor Massey writes me, no one knowing its stations 
high on the mountains would think of calling it “of the Valley.” 
In the European species the scape is elongate, so that the flowers 
are borne opposite the middle or upper halves of the leaves; the 
leafy axis (to the base of the upper leaf) is, except in highly 
cultivated plants, 5-12 em. high, with the larger leaves 1-2 dm. 
long and 3-7.5 cm. broad, their veins and cross-partitions, as 
seen by transmitted light, relatively faint and pale; the longer 
bracts of the raceme are lanceolate, 4-10 mm. long, and much 
shorter than the pedicels; and the seeds are nearly globose. In 
the native eastern American species the scape and raceme are 
shorter than the leafy axis or barely reaching the lower half of 
the lowest leaf; the leafy axis is 1.5-2 dm. high, with the larger 
clear green (not glaucescent) leaves 1.5-3 dm. long and 4-12 cm. 
broad, strongly nerved, the dark nerves and cross-partitions 
sharply visible by transmitted light; the longest bracts of the 


1944] Fernald,—Flora of Eastern North America 13 


raceme are almost linear, 0.8—2 cm. long and nearly equaling to 
much exceeding the pedicels; and the seeds are compressed, 
either oblate or somewhat lenticular. 

In recent years the Alleghenian native has been known as 
Convallaria majuscula Greene in Fedde, Rep. Nov. Spec. v. 46 
(1908). Greene proposed two species, C. globosa, 1. c., a plant 
cultivated by the late Robert Ridgway who received it from a 
dealer as found in the mountains of North Carolina, and C. 
majuscula, ‘‘occasionally collected in the higher mountains of 
Virginia, from the Peaks of Otter northward; also in those of 
southeastern Pennsylvania. .. . it re totally from 
that [C. majalis] in its very large light-green leaves without 
trace of bloom, with excessively fibrous anatomy, insomuch that 
the surface of the leaf when growing looks to be plicate rather 
than plane and even. Both these American species 
when compared with C. bee dh flower much later, their folinge 
perishing at the end of summer.’”’ There are no evident specific 
characters separating the cultivated C. globosa and the indigenous 
plant of western Virginia, therefore the name C. majuscula has 
come into use. 

There is, however, a cence sealable name for the native 
plant, which was well described 68 years before Greene’s publica- 
tion. This is Convallaria montana Raf. Aut. Bot. 66 (1840). 
Rafinesque’s account, quite as good as Greene’s, was as follows: 


486, Conval, montana Raf. (pseudlo-majalis Bartr. in Rees cycl. Am. ed.) 
fol. binis sessilib. ovatobl. me scapo angul. fol. subeq. racemo 10-12 


floris, bract. lanc. ad ped. fl. secundis—Unaka and Cherokis Mts. 
large’ plant, leaves 6 to 8 ciebee long, ats fl. mn of C. majalis. Bartram 
says the berries are blue and ovate. C. majalis has leaves petiolate 


elliptic acute at both ends, raceme of 7-8 fl. feast shall length of pedicels. 

Rafinesque had, also, a Convallaria parviflora, 1. c. with ‘“‘seapo 
filif. fol. brevior.” ete., from ‘Appalachian and Wasioto Mts.” 
Whether this was small-flowered C. montana or the introduced 
plant I do not feel certain, but the identity of C. montana and 
C. majuscula can hardly be questioned. 

From Rafinesque’s reference to Bartram it would appear that 
the latter had still earlier given the correct name to our native 
Convallaria. In Rees Cycl. Am. ed. x. (1810 or later), after the 
original British treatment there occurs the following unsigned 
and bracketed note: 


14 Rhodora [JANUARY 


[To these we will add, from Bartram, C’. pseudo-majalis, mountain lilly 
of the valley. This charmin ing plant is indigenous to the mountainous 
parts of the. United States of America, particularly the country of the 
Cherokees, in the rich glades or shady vallies in their mountains, and in 
the like situations in Pennsylvania. It differs but little from C. majalis of 
pot only is larger every way, and the fruit blue and more oblong to 

te 


Here, of course, was where Rafinesque got his quotation from 
Bartram about the blue berries. In his Travels (1791) Bartram 
repeatedly noted Convallaria majalis from the upper slopes of the 
Cherokee country but I have been unable to find him giving in 
print a new name or noting the “blue” berries. Since C. 
montana (or C. majuscula) has, as I am assured by Professor 
Massey and others who are familiar with it in the wild, RED 
berries we can hardly accept C. pseudo-majalis as properly 
diagnosed. Bartram could quite as well have had Polygonatum! 
The synonymy of our native species is as follows: 

CONVALLARIA MONTANA Raf. Aut. Bot. 66 (1840), excluding 
the synonym C. pseudo-majalis Bartr. ex Rees Cycl. Am 
(1810 or later) as too doubtful on account of the blue aie 

°C. parviflora Raf. 1. c. (1840). C. globosa Greene in Fedde, Rep. 
Brae Spec. v. 46 (1908). C. majuscula Greene, 1. c. (1 908); 

Fernald in RHopora, xxx. 184, Hen poses (1928) and xxxix. 347, 

foot-note (1937). 


TRILLIUM CATESBAEI AND T. NERVOSUM Ell. (PLATES 810 and 
811).—Elliott, Sk. i. 429 (1817), described two new species, 
Trillium Catesbaei and T. nervosum. The first, which he took to 
be the same as the plant illustrated by Catesby, Nat. Hist. 
Carol. ete., i. t. 45, as Solanum triphyllon; flore hexapetalo, 
carneo (our PLATE 811, ric. 1) was described as follows 


: CaTESBAEI. E. 

GOP ge recurvato; petalis Peduncle recurved; petals lance- 
lan tis, calyce majoribus; foliis | olate, larger than the calyx; 
obovatis ovalibusque, acuminatis, leaves obovate and oval, acuminate, 
basi attenuatis. E. 

Catesby, Carol. p. 45, t. 45. 

T. cernuum, Miche. 1. p. 216? 

Leaves 4-6 inches long, rather obovate, 3 inches wide, gre to the 
base and not abruptly acuminate at the summit. lanceolate, 
expanding, undulate, rose coloured. Leaves of the calyx Cae narrow. 

[Then a paragraph of more or less pertinent observation, with the con- 
a ea his plant, coming from the same region as Catesby’s, must be 
i 
Pendelton county, South-Carolina; Mesrs. Baker & Perry. 


a ee ae ae Fete he eas i eo ee ee 
sees iad 


1944] Fernald,—Flora of Eastern North America 15 


The second of these new species (our PLATE 810, FIG. 2) was 


9. NERVOSUM. , 

T. pedunculo recurvato; petalis Peduncle recurved; petals oblong 
oblongo-lanceolatis, calyce majori- | lanceolate, larger than the calyx; 
bus; foliis lanceolatis ovatisque, | leaves lanceolate and ovate, acute 
utrinque acutis, membranaceis, | at each end, membranaceous, 
nervosis. E. nerved. 


Plant 6-8 inches high. Leaves generally narrower than those of the 7. 
sessile, most commonly lanceolate, membranaceous, somewhat 3 nerved. 
Peduncles about an inch long. Petals rose coloured. 

Grows in the upper and middle country of Georgia and Carolina. 
Athens; Mr. Green. The T. cernuum of Walter probably belongs to this: 
species. 


Now, if anything is clear, it is that the plants which Elliott 
was describing differed strikingly as follows: 7. Catesbaei with 
“rather obovate’ leaves 3 inches broad, 7. nervosum with them 
lanceolate and ovate (‘‘most commonly lanceolate’’) tapering 
at both ends. ; 

Most fortunately, Elliott’s Herbarium, which, in the past 
suffered serious destruction by insects, mould and the removal 
of specimens, still contains these two types in good condition. 
They were photographed in October, 1941, by Mr. and Mrs. 
Weatherby, their photographs (our PLATE 810) now in the Gray 
Herbarium. Although it is, as said, most fortunate that the 
types exist and closely agree with Elliott’s descriptions, it is 
most unfortunate that Elliott identified his plant with obovate- 
oval broad leaves with Catesby’s plate and, consequently, called 
this species 7’. Catesbaeit. The type of 7. Catesbaez has elongate, 
curving stigmas sessile at the summit of the ovary, while its 
petals are pretty broad to be called lanceolate. The type of the 
narrow- and tapering-leaved T'. nervosum has a definite style 
capping the ovary (PLATE 810, ric. 2). In other words, the type 
of 7. nervosum is identifiable with 7. stylosum Nutt. Now, if 
Catesby’s plate be examined it will be seen (our PLATE 811) that 
the leaves are those of 7. nervosum and that, in the Catesby 
drawing of the fruit (our ria. 2) there is a definite style. Elliott 
evidently ‘got the wires crossed” and identified the Catesby 
plate with the wrong plant; and Rendle, Journ. Bot. xxxix. 333 
(1901), said “In the absence of the specimens which Elliott had 
before him, Catesby’s figure (which he cites) is the only authority 
for this species.”’ ‘He, therefore, . took up T. Catesbaei for the 


16 Rhodora [JANUARY 


narrow-leaved plant with definite style as impressionistically 
shown in Catesby’s plate (impressionistically, because Catesby 
had the sepals beautifully roseate and petaloid, just like the 
petals!) and a plant of the Carolina mountains with broadly 
rhombic leaves he described and illustrated as T. Rugelit Rendle 
in Journ. Bot. xxxix. 381, t. 426 B (1901). Rendle’s illustration 
shows broadly rhombic leaves 11.5 cm. wide; material from the 
same general region, Highlands, North Carolina, April, 1903, 
Harbison, has them less broadly tapering at base, tending to 
obovate and 6-9.5 cm. broad; Harper’s no. 1891 from Randolph 
County, Georgia, generally identified with 7. Rugeliz, has them 
in outline nearly as in the type of 7. Catesbaez and 9-12 cm. 
broad; and Harper’s no. 3492 from Tuscaloosa County, Alabama, 
has them definitely rhombic-obovate and 7.5 em. broad; while 
the type of 7. Catesbaez has them 9-10 cm. broad. All these, as 
I understand the plants, show the usual range of variation in the 
leaves and are all T. Carresparr Ell. (1817). They are also T. 
Rugelii Rendle (1901), for, since ‘‘the absence of the specimens 
which Elliott had before him”’ is now rectified by looking in 
Elliott’s Herbarium at Charleston, not in the British Museum, 
the identity of 7’. Catesbaei has become clarified. 

Early Carolina botanists clearly recognized that T. nervosum 
Ell. (1817) was the same as T. stylosum Nutt. (1818) and many 
sheets from M. A. Curtis had both names (often bracketed) on 
their labels. They were right. The error occurred when T. 
nervosum was dropped (as by Small) and 7’. Catesbaez (following 
the gratuitous assumption of Rendle) was wrongly used in the 
sense of TJ’. stylosum. 

In PLATE agai jo ~ ve bey TYPE of TRILLIUM CaTESBAE!I Elliott, 1, after 
bil: rg. 2, the we ae F. Weatherby. Fics. 2 and 3, T. NERYOSUM 


FIG. 2, rd dle ih x 4, after a photograph by C. A. and Una 
Wenn: ; Fic. 3, detail of flower in central specimen of fig. 2, to show style, 


gee 811, rics. 1 and 2, portions of Catesby’s plate of Solanum triphyllon; 
flore hexape etalo, carneo, Nat. Hist. Carol. i. t. 45: FIG. 1, flowering summit, X< 1; 
FIG. 2, fruit, showing style, X11. Fia. 3, leaf, x 1, of TYPE of SMILAX Bona- 
Nox L., var. EXAURICULATA Fernald. 


TRILLIUM FLExIPES Raf. Aut. Bot. 133 (1840). 7. erectum, 
var. declinatum Gray, Man. ed. 5: 523 (1867). T. declinatum 
(Gray) Gleason in Bull. Torr. Bot. Cl. xxxiii. 389 (1906), not 
Raf. Aut. Bot. 135 (1840). 7. Gleasoni Fernald in Ruopora, 
xxxiv. 21 (1932). 


1944] Fernald,—Flora of Eastern North America 47 


Hatiperdne s description of Trillium flexipes was unusually 
good, for him 

968, Tril. A [i. e. his subgenus Anthopium, with peduncled flowers and 
sessile stigmas] fleripes Raf. caule sulcato, fol. sessil. obovatis acuminatis 
undul. 3nervis, pedunc. equante inc ato flexuoso, calix lanceol. acum. 
petalis albis eq. obl. acut. undul.—West Kentucky ret Tennessee, rare, 
pedal, leaves 3 inches long, 2 broad, flowers middle siz 

The inclined peduncle about equaling (in early anthesis) the 
sessile, acuminate, obovate leaves, the acuminate, lanceolate 
sepals and the equal, oblong white petals are all good characters 
of Trillium Gleasoni. That it occurs in western Kentucky there 
can be no question. Witness Shacklette, no. 378 from Union 
County, Kentucky. Wiegand & Eames, in their study of the 
group, cited it from Tennessee in Ruopora, xxv. 190 (1923) as 
did Small (Man.); and it occurs in eastern Missouri. There is no 
reasonable doubt about 7. fleripes. The form with maroon or 
purple petals is 

Forma Walpolei (Farw.), comb. nov. 7. cernuum, var. de- 
clinatum, forma Walpolei Farw. in Rey. Mich. Acad. Sci. xxi. 
363 (1920). 

T. pecLINATUM Raf. |. c. 135 (1840) and T. BALDUINIANUM 
Raf. 1. c. 185 (1840) are probably both forms of T. NeRVosum EII. 

T. LANCIFOLIUM Raf. |. ¢. 132 (1840) can hardly be anything 
but 7. recurvatum, var.(?) lanceolatum (Boykin) S. Wats. in 
Proc. Am. Acad. xiv. 273 (1879), based upon T. LANCEOLATUM 
Boykin in herb. in Wats. 1. c. 274 (1879). Watson treated T’. 
lanceolatum as a doubtful variety of the northern and wide- 
ranging 7. recurvatum Beck, but, as Small, Bull. Torr. Bot. Cl. 
xxiv. 171 and 174 (1897), showed, the two are well distinguished 
species. Small (Man.) gives the range of 7. lanceolatum as 
“W. Fla. to La., Tenn. and Ga.” and in Ruopora, xlv. plate 773, 
fig. 3 (1943), I showed, life-size, the summit of a characteristic 
plant, X 1, from northwestern Florida, the original specimen 11 
inches high. If Rafinesque’s description be compared with this 
figure the identity of 7. lanceolatum Boykin (1879) and T. lanci- 
folium Raf. (1840) will be apparent: 

962, Tril. S. [subgenus Sessilium] lancifolium Raf. caule elato, fol. sess 

neeol. acutis trinervis patulis planis sepe maculatis, calicib. patulis vel 
reflexis, petalis erectis longior rubris unguic. lanceol. —Florida to Alabama 
and Apalachian Mts. stem often pedal leaves 3 inches, flowers uncial. 


18 Rhodora [JANUARY 


As to “flowers uncial.’’, the petals vary from 1.5-4.5 cm. long, 
the lower measurement being less than “uncial. 

We do not know true Trillium lancifolium (T. lanceolatum) in 
the Gray’s Manual area, although Rendle, Journ. Bot. xxxix. 327 
(1901), referred to “Specimens which I have seen from Kentucky 
(Short). The specimens which are in American herbaria, ac- 
companied by Short’s label with the print, “C. W. SHORT, 
M.D. KENTUCKY, 1840” fortunately bear, in his hand, the 
written memorandum, ‘‘From Dr. Boykin, Sha ”. In other 
words, they are isotypes of Boykin’s species. 

T. cungeatum Raf. Aut. Bot. 133 (1840). 7. Hugeri Small, 
Fl. Se. U. S. 277 and 1328 (1903). 


Trillium cuneatum was thus described: 


964, Tril. S. [subgenus Sessilium] cwneatum Raf. caule elato, fol. ses- 
silib. ovatobl. acutis trinervis planis seman calicib. = ag obl. petalis 
cuneatis duplo longior acutis purpura: ns.—Unaka Mts. of Cherokis, 
pedal leaves 3 at flowers large akin't to i tlioas of Tr. maculatum, but 
leaves very differe 


It is most difficult to believe that Rafinesque’s plant from the 
Cherokee country, with ovate-oblong acute green leaves, oblong 
sepals and cuneate purplish petals twice as long as the sepals is 
not the characteristic large-flowered 7. Hugeri (type from 
Tryon Mt., North Carolina), which abounds in the Appalachian 
region from northwestern Florida and Alabama northward to 
North Carolina and Kentucky. Rafinesque compared it with 
his earlier published 7. maculatum, ‘of the coastal plain and 
piedmont regions from the Carolinas to Alabama and Missis- 
sippi’”’, with, further, to quote Dr. W. A. Anderson in RHopora, 


XXXV1. 122, 123 (1934), “leaves . . . lance-ovate, . 
strongly sngttled. mottling tending to form longitudinal he. 
sepals 2.5-5 cm. long. . . , lanceolate, acute; petals 3.5-6 


em. long”. Anderson, identifying T. macuLtatum Raf. (1830) 
with T. Underwoodii Small (1897), said “Among the numerous 
species of Trillium described by Rafinesque, this one is unmis- 
takable’’. 

By Small (Man.) his Coastal Plain Trillium Underwoodii (T. 
maculatum Raf.) is, in the key, separated from his montane T. 
Hugeri (T. cuneatum Raf.) by its narrower leaves (“‘bracts’’), in 
the former “much longer than wide, often twice as long”, in the 


1944] Fernald,—Flora of Eastern North America 19 


latter ‘‘nearly or quite as wide as long’. From the type-locality 
and the description 7. cuneatum well matches the more oblong- 
ovate-leaved extreme of the Appalachian 7. Hugeri, originally 
described by Small with “blades . . . bright green [Raf. 
said ‘concolor’] . . . , sepals oblong to oblong-lanceolate 
[Raf. said ‘obl.’], . . . petals oblong-lanceolate to spatulate, 
purple, somewhat longer than the sepals [Raf. said ‘cuneatis 
duplo longior . . . purpurascens’].”’ 

Rafinesque had no less than 34 so-called species and 67 so- 
called varieties! of Trillium. A few species, as noted above, 
were described with sufficient clarity to be recognizable. In 
these cases his names, when having priority, are automatically 
taken up. T. flexipes Raf. thus displaces T. Gleasoni Fernald, T. 
lancifolium Raf. throws out T. lanceolatum (Boykin) 8. Watson, 
T. cuneatum Raf. antedates T. Hugeri Small, and T. maculatum 
Raf. replaces 7. Underwoodii Small. Most of the others are too 
vague for recognition, others are definite renamings of already 
properly published species, often with mere undefined varietal 
names: for instance, 7. rotundifolium Raf., with a brief descrip- 
tion which is applicable to 7. erectum L., and “Var. 1. Flexicaule. 
2. Rubricaule. 3. Maculatum. 4. Orbiculatum. 5. Pallidum. 6. 
Undulatum.’”’ These six varietal names are, of course, nomina 
nuda and have no status; their publication does not indicate a 
sound mentality or genius as a phytographer. Nevertheless, 
although (his p. xxi) acknowledging what any clear thinker 
quickly sees, that ‘ Rafinesque’s genius is debatable in the 
extreme’’, that “unbalanced . . he may have been” (his 
p. 166) and (his p. 263) that ae Dr. Faustus, dangerously 
experimenting, tempting the rack or devil-ridden madness, 
would certainly correspond in a crude way to Rafinesque’’, one 
of the few monographers of Trillium, Mr. Donald Culross Peattie, 
becomes ebullient over this erratic and “unbalanced” man, 
whom, nevertheless, he considers “one of the most prodigious 
and prophetic scientists of the century 1750 to 1850” (his p 
261)—the century, to mention a few botanists only, of Haller, 
Linnaeus, Lamarck, Humboldt, Robert Brown, DeCandolle, 
Darwin, young Gray and his friend, young Joseph Hooker, and 


1 Dr. Merrill Ce hy weenie a with 67 varieties in 
Trillium, set a score not achieved by the much advertised H 's 57 varieties”’. 


20 Rhodora [JANUARY 


countless others for whom no one has the “‘nerve”’ to apologize 
and who were respected by every one for sanity, clarity and sound 
scholarship. Peattie, over-enthusiasite about the “prophetic” 
nature of Rafinesque’s unbalanced work, wrote (Green Laurels, 
p. 266): ‘If the rules of priority were strictly and justly applied, 
Rafinesque would be found to have antedated a large part of the 
work . . . in botany of Gray and DeCandolle”’. Whew! 
Rafinesque himself might well have written such a sentence; his 
introverted brain (likewise that of Marcus Jones) frequently 
brought forth just such statements. 

But, returning to Trilliwm. One of the few monographic 
studies of Trillium is that of Peattie, Trillium in North and South 
Carolina, in Journ. Elisha Mitchell Soc. xlii. 192-206 (1927) and, 
as an outgrowth of this, the treatment of the genus in Small’s 
Manual (1933). In the former treatment we read (p. 193): 
“Following Asa Gray, who eschewed anything from Rafinesque’s 
pen, Sereno Watson! . . . reduced to synonymy or varietal 
rank the greater part of Rafinesque’s . . . names of Tril- 
lium. It was not until Small showed the limited nature of T. 
sesstle L. and set off 7. Underwoodii and later T. Hugeri that our 
knowledge of the genus drifted out of the doldrums into which 
workers like Watson [whose great genius and just fame need no 
defense], with almost no field knowledge, had put us”. In view 
of Peattie’s evaluation of the work of Rafinesque it is, conse- 
quently, amazing that he so consistently ignored his 34 species 
and 67 varieties of Trillium, many of them described from the 
Carolinas or other Southern States, and that he failed to weleome 
the opportunity to displace T. Hugeri and T. Underwoodii of 
Small by Rafinesque’s names of 1840 and 1830 respectively. 


1 Peattie seems to overlook the patent fact that both Asa Gray and his friend 
Alphonse DeCandolle (dim lights, one gathers, as compared with his hero) regularly 
took up Rafinesque’s genera and species when they were definite and had priority: 
Peltandra Raf., including P. undulata and P. alba Raf. ate Raf., with the species 

. borealis Raf, ; Hesalecris Raf., with the species H. aphylla Raf.; Adlumia Raf., with 
the species A. rrhosa Raf.; and so on with scores pipet genera. This was far from 

wing’ his work, When sound. It was not the fault of either DeCandolle or Gray 
that the great majority of Rafinesque’s genera and bed 
or were sO vague as to be undecipherable. Neither is it ono the ‘‘prejudice”’ 
imagined by Peattie that earlier specific names have been taken up in Rafinesque’s 
genera Peltandra, Hezalectris, Adlumia, Cladrastis, Nemopanthus, esochins. etc., nor 
that later monographers have discarded his genera Didiplis, Steironema, Ilysanthes, 


founded statements have no place in real science; they belong in fi 


| 
? 
] 


. 
4 
F 
: 
e 


1944] Fernald,—Flora of Eastern North America 21 


If anyone has ‘‘eschewed anything from Rafinesque’’ it is the 
author of a recent monograph who has ignored all but 4 (scarcely 
4 per cent) of Rafinesque’s published names in the genus—these 
all reduced by the admirer of that “‘prophet’’ to synonymy. 

(To be continued) 


32 Rhodora [FEBRUARY 


OVERLOOKED SPECIES, TRANSFERS AND 
NOVELTIES IN THE FLORA OF 
EASTERN NORTH AMERICA 
M. L. FERNALD 
(Continued from page 21) 
Smitax Pseupo-Cuina L. Sp. Pl. 1031 eae S. tamntfolia 
Michx. Fl. Bor.-Am. ii. 238 (1803). Puat 
Smilax Pseudo-China, named from eis e the Asiatic 
Smilax China L., the China radix of Bauhin, was included, along 
with S. lanceolata L., by Linnaeus in a section of the genus with 
unarmed terete stems. The treatment was as follows: 


*Caule inermi tereti 


13. SMILAX caule i inermi tereti, foliis inermibus: “ caulinis Jame Chi- 
poatbebe —- [corrected to rameis in ed. 2] 
ova 
Smilax fraiapelated Simlax] caule tereti inermi, foliis inermibus: caulinis 
cordatis, ramorum genet ie pedunculis longissimis. Gron. virg. 193. 
Smilax caule tereti in foliis cordato-ovatis acutis rate, 
petiolis bidentatis. "Hort. “Cl lif. 495. Gron. virg. 120. 
virginiana, spinis innocuis armata, latis canellae foliis, radice 
0. f.6 


Habitat in Vina Jasna 


It is at once evident that, as usual, Linnaeus had no clear 
understanding of American species and that his citations covered 
different species. The phrase ‘‘Caule inermi’” of his major 
grouping of the primary diagnosis (of the plant he had actually 
before him in his own herbarium), of the quotations from Gro- 
novius and from Linnaeus’ Hortus Cliffortianus at once contrast 
with “‘spinis . . . armata’’ of the Plukenet reference and 
“Smilax aspera” of Sloane. Singularly enough, Alphonse De 
Candolle in his Smilacées in DC. Mon. Phan. i. 82 (1878), 
although saying ‘‘Linn. Sp. p. 1461 [ed. 2], excl. syn. post 
Gronov.”’, placed it, along with S. rotundifolia, S. glauca, etc. in 
the group with ‘Folia persistentia vel subpersistentia’’ and 
described it ‘‘aculeis in caule crebris; . . . limbis margine 
setaceo-ciliatis”. In other words the S. Pseudo-China, sensu A. 
DC. (and many followers), not L., is S. Bona-nox L., which was 


Ses er 3 suai is ere teen reo! ren Speen ia " , “ Er: 
SER Fe ce ee Pe rag ey ae ere ae = EO TT TT eS eet Renee Ne! ROR Ree ese al ala 


1944] Fernald,—Flora of Eastern North America 33 


originally described ‘‘foliis ciliato-aculeatis.”” It, therefore, 
becomes necessary to determine what Linnaeus actually had in 
his own herbarium prior to 1753. Fortunately there is a char- 
acteristic sheet of foliage marked by Linnaeus ‘‘K. Pseudo- 
China’. This, undoubtedly collected by Kalm (“K’’) in New 
Jersey or Delaware, is characteristic S. tamnifolia Michx. FI. 
Bor.-Am. ii. 238 (1803). <A portion of the Linnean TyPE is re- 
produced as our Fic. 2. The first Gronovian reference given by 
Linnaeus is supported by a fruiting branch in the herbarium of 
Gronovius at the British Museum, badly crumpled and unsightly, 
also by a very beautiful flowering plant (our Fig. 1). These are 
likewise S. tamnifolia with very long peduncles. The third 
reference, to Hortus Cliffortianus, carries back to the Plukenet 
figure cited by Linnaeus, which is of some woody species, pre- 
sumably S. Walteri Pursh, since its foliage will do for that species 
and, in the place cited, Pluketet identified it with Virginian 
specimens with ip oe coccineis’; and the second Linnean 
reference to Gronovius is supported by a characteristic lateral 
sprig of flowering S. rotundifolia! The Smilax aspera of Sloane 
need not specially concern us, since it is so clear that the primary 
material, the plant which Linnaeus had in his own herbarium and 
the two Clayton (Gronovian) specimens with smooth stems are 
so definitely the true type and syntypes of S. Pseudo-China. 
The beautiful photograph sent from Paris of the type of S. 
tamnifolia Michx. is unequivocal. It is, consequently, worth 
noting that Michaux thought it might be the S. caule terett, 
folits inermibus: caulinis cordatis, ramorum lanceolatis, pedunculis 
longissimis of Gronovius (our FIG. 1). The point which Michaux 
did not note is, that this was the best syntype of S. Pseudo- 
China L. (1753). 

Further showing the utter confusion of Linnaeus in interpreting 
North American species of Smilaz is the fact that, although he 
described in his section with Caule aculeato, tereti a new species as 
S. tamnoides L. Sp. Pl. 1030 (1753), the specimen in his own 
herbarium marked S. tamnoides is an unusually good piece of 
unarmed, herbaceous-stemmed S. Pseudo-China (S. tamnifolia 
Michx.) while the Catesby plate, from which the characters 
were obviously drawn, is of a prickly-stemmed, evergreen, 
high-climbing and broad- and eciliate-leaved extreme of an- 


34 Rhodora [FEBRUARY 


other species! In fact, the specimen preserved by Linnaeus as 
representing his S. tamnoides not only has the unarmed stem, 
the slightly panduriform leaves, the long peduncles and loosely 
globose inflorescence of S. Pseudo-China (S. tamnifolia Michx.). 
One of its inflorescences even shows with diagrammatic sharp- 
ness the elongate-clavate styles of that species. 

In my conclusion that the plants selected are the real types of 
Smilax Pseudo-China I am quite in agreement with Dr. Pennell 
who, in 1916, wrote: 


Smilax Pseudo China L. l. c. 1031. 1753. “Habitat in Virginia, 
ica.”” Species clearly aggregate, represented in the Linnean 
herbarium by Se age ens written up by Linnaeus as follows, according 
to a letter of B. D. Jackson: “three sheets pinned together; the first 
is ‘II K Pseudo China’, it is a barren branch, the leaves leathery; the 
second sheet is written’ up ‘II’ and seems quite the same plant as the 
former, but has one berry, the third sheet is of a West Indian species, 
coll. by Patrick Browne in Jamaica, probably S. celastroides.” From 
the wording of the Linnaean description none of these can be con 
sidered the type of S. Pseudo China, but this would be rather a a 
specimen of Gronovius, also studied by Linnaeus, now in the Grono- 
vian herbarium in the British Museum. The description of Linnaeus 
is word for word from Gronovius, except for the addition of the 
phase “racemis ovato-oblongis,” inappropriate for any Smilax what- 
ever. ‘Smilax caule tereti i inermi: fo =e ermibus, caulinis cordatis, 
ramorum lanceolatis unculis longissimis,’ > Gron novius, Fl. Virg. 
156. 1742, citing Plavion’s Nos. B41 561 and 630, is represented in 
the herbarium by Nos. 561 and 630 (for No: 541 see above note under 
S. herbacea). These two numbers are identified by Dr. A. B. Rendle 
as both the same as No. 541, that is, as seemed Smilax herbacea L. 
For the verification of these two Linnae types I am indebted to 
Dr. B. Dayden Jackson, of the Taianean| Saeiaty of London, bs 
Dr. gah dle, of the British Museum.—Pennell in Bull. T 
Bot. Cl. xiii. 413, ‘414 (1916). 


Unfortunately, Pennell did not himself see the specimens dis- 
cussed; had he done so he certainly would not have reduced 
Smilax Pseudo-China, correctly typified by him, to S. herbacea 
L. He depended upon identifications by the late Drs. B. Daydon 
Jackson and A. B. Rendle, neither of whom clearly understood 
the species involved. The plant generally passing as S. herbacea 
rarely if ever has any tendency to panduriform leaves. Its 
leaves, varying from oblong-ovate to cordate-subrotund, are 
glaucous and glabrous beneath, the flowers much larger than in 
S. Pseudo-China, with the styles rather broadly lingulate, the 
berries glaucous. Whether the type of S. herbacea, from Vir- 


Ie ee 6 Ce 6) CO Eee! 86 «oe ee 


1944] Fernald,—Flora of Eastern North America 35 


ginia (Clayton), is of this species (S. pedunculata Muhl.) or S. 
pulverulenta Michx. can be determined only by actual (future) 
examination of the material. The two photographs of the 
specimens (one sheet in Herb. L., the other in the Gronovian 
Herbarium) are of identical plants. The foliage is young, but it 
looks green and lustrous beneath, as in S. pulverulenta Michx. 
Until the specimens can be actually studied we may retain the 
names S. herbacea and S. pulverulenta as currently used. 

Although Pennell, 1. c. 414, placed Smilax inermis Walt. FI. 
Carol. 244 (1788), as well as S. Pseudo-China, in the synonymy 
of S. herbacea, rather than place them with S. tamnifolia, it now 
seems clear that Walter’s species was identical with S. Pseudo- 
China. Pennell said “‘Type, presumably from Berkeley County, 
South Carolina, not verified”. Walter’s diagnosis was perfectly 
good for S. Pseudo-China; Berkeley County is on the outer 
Coastal Plain and it is notable that Pennell’s only stations, in his 
early paper, for S. herbacea from Virginia (south of Fairfax Coun- 
ty) North Carolina, South Carolina and Georgia were all from 
along the Blue Ridge or the Alleghenies, while he specially 
designated the area of S. tamnifolia as “Coastal Plain; Long 
Island to South Carolina’. In the Gray Herbarium, as in those 
studied by Pennell, there is no so-called S. herbacea from the 
Coastal Plain from south of Virginia. Walter’s S. inermis, 
“presumably from Berkeley County, South Carolina’, had the 
weak (“infirmo”) stem only 3 feet high (‘“tripedali”). The 
southernmost specimens in the Gray Herbarium of S. Pseudo- 
China are three from Berkeley County, South Carolina (Santee 
Canal, Ravenel; west of Chicora, Godfrey & Tryon, no. 868; 
Moncks Corners, Godfrey & Tryon, no. 1411), one from Charles- 
ton County, South Carolina (north-northwest of McClellanville, 
Godfrey & Tryon, no. 1112) and one from eastern Georgia 
(Savannah, Mrs. Say). Incidentally there are in the Gray 
Herbarium no specimens (not even from Wilmington) from 
North Carolina and Pennell saw none from that state. Is it 
really absent from or rare in the broad area between southern 
South Carolina and southeastern Virginia? This gap is frequent 
in the known ranges of many species. 

NEMEXIS ELLIPTICA Raf. Aut. Bot. 131 (1840) with stem “6 to 
12 inches high, leaves uncial’, the stem with quadrate, elliptic 


36 Rhodora 3 [FEBRUARY 


leaves acute at each end, from: Alabama, might have been 
Smilax Hugeri (Small) J. B. Norton in Pennell in Bull. Torr. 
Bot. Cl. xliii. 420 (1915), based upon Nemexia Hugert Small 
(1903). Since the combination Smilax elliptica already exists 
for other species Rafinesque’s Nemezzs elliptica need not disturb 
us, but by those who maintain Nemexia as a genus it must be 
considered. 

Another and quite clear synonym of Smilax Hugeri is S. 
herbacea sensu Walt. Fl. Carol. 243 (1788), not L. His “‘caule 
annuo inerme stricto suberecto 2—5 pedali, simplici, foliis verticil- 
latis ovatis . . .  cirrhis obsoletis’’, etc. are unequivocal for 
a plant which is well known from southern and southeastern 
South Carolina and from Georgia. 

Smitax Bona-nox L., var. exauriculata, var. nov. (TAB. 811, 
FIG. 3), foliis oblongo-lanceolatis subacuminatis basi cordatis nec 
panduriformibus, sgt fk setoso-ciliatis—VireiInta: Norfolk, 

eed? Sa in Herb. pe 8 

NA-NOX, Var. eodsvciilia (Beyrich), oe nov. S. 
hedsraefolse Beyrich ex Kunth, Enum. v. “ae (1850). S. Bona 
oo hederaefolia (Beyrich) A. DC. in DC. Mon. i. 77 


As it extends northward into eastern Virginia (more locally 
northward) Smilax Bona-nox is nearly as variable as farther 
south. In this northeastern area of its broad dispersal it occurs 
in three (perhaps four) quite definite variations. Typical S. 
Bona-noz L. Sp. Pl. 1030 (1753) was, most exceptionally, not so 
much confused as most of Linnaeus’s North American species. 
Although he included a West Indian shrub of Bauhin, he gave 
the “Habitat in Carolina” and the species rests primarily on 
Smilax, foliis latis in margine spinosis, caroliniana,  stipite 
quadrato, Pluk. Alm. 348, t. 111, fig. 3; the Linnean diagnosis 
drawn directly from Plukenet: ‘“SMILAX caule inermi [because 
Plukenet had merely a sterile tip] angulato, foliis ciliato-aculea- 
tis.” Typical S. Bona-noz, therefore, is the slender, straggling 
and rarely climbing, freely branched shrub with deltoid-ovate 
to slightly panduriform leaves usually mottled with white, the 
rounded basal lobes short and tapering into the upper part of 
the blade, the margins bristly-ciliate. Its northern limit seems 
to be in Wicomico County, Maryland (J. J. Carter in Herb. 
Phil. Acad.). 


1944] Fernald,—Flora of Eastern North America 37 


Linnaeus picked up another of Plukenet’s figures. This was 
S. Bona-nozx @. of L. Sp. 1. c. “Smilax caroliniana, stipite quadrato 
leni, foliis angustis asperis auriculatis ad basin angulosis. Pluk. 
alm. 348, t. 111. f. 3.””. Somewhat later this was taken up as a 
species: S. hastata Willd. Sp. iv?. 782 (1806), Willdenow empha- 
sizing the point shown by Plukenet, that the very narrow leaves 
are “margine ciliato-aculeatis”. This extreme with narrowly 
lanceolate bristly-ciliate blades with narrow divergent basal 
lobes occurs occasionally in the Carolinas and southward. The 
Virginia material is slightly transitional but may be ealled S. 
Bona-nox, var. hastata (Willd.) A. DC. in DC. Mon. i. 79 (1878). 

The third variety with strongly bristly-ciliate leaves is the 
very extreme plant described above as var. evauriculata. The 
type material is an old sheet from Norfolk, collected probably by - 
Reed who got in the neighborhood of Norfolk several species 
otherwise and not recently known from Virginia. Var. exauricu- 
lata is very extreme in having elongate oblong-lanceolate, taper- 
ing, ciliate-margined leaves with nearly parallel sides and cordate 
(not hastate or subhastate) bases. It piques the curiosity! The 
Specimen was sent out as S. hastata, var. lanceolata Pursh. It 
can hardly be that, however. Pursh gave no new description; 
he was simply giving a name to the S. lanceolata Walt., perhaps 
not L.: ‘‘caule angulato spinoso; foliis longis angustis lanceolatis 
inermibus”. Var. exauriculata could not be described ‘“‘foliis 
inermibus’”’. I am tentatively placing with it a sheet from dry 
bank, Santee Canal, 5 miles west of Pineville, Berkeley County, 
South Carolina, Godfrey & Tryon, no. 630 (distrib. as S. glauca). 
Its leaves are chiefly oblong, rather than lanceolate. 

The extreme of Smilax Bona-nox with “foliis inermibus”’ is a 
coarse and usually high-climbing shrub with ovate, deltoid or 
broadly panduriform leaves usually green above and with 
eciliate or very weakly and casually ciliate leaves on the fertile 
branches. Basal sprouts may have ciliate and mottled leaves 
but the foliage of the adult branches is rarely so. This often 
high-climbing vine has the leaves up to 8 (on sprouts to 12) cm. 
broad and in good development is very different from typical S. 
Bona-nox. It is var. hederaefolia, which rests on S. hederaefolia 
“Beyrich ined.” ex Kunth (1850), treated by Alphonse 
DeCandolle as S. Bona-noz, subsp. hederaefolia. Beyrich’s 


38 Rhodora [FEBRUARY 


original material was from shores of Savannah River, Georgia. 
To this citation DeCandolle added “ Virginia (Rigel ) 
Var. hederaefolia is the commonest extreme of the species. In 
southeastern Virginia it abounds on damp shores or in low woods, 
the typical form of the species preferring open and drier habitats. 
Var. hederaefolia reaches Delaware (A. Commons in Herb. Phil. 
Acad.) and is isolated on Nantucket Island in southeastern 
Massachusetts. 

Smilax tamnoides L. Sp. Pl. 1030 (1753) was another species not 
understood by its author. His species rested upon two items: 
the first a specimen of the unarmed herbaceous plant which on 
the next page he described as S. Pseudo-China, the plant which 
has regularly passed as S. tamnifolia Michx. The second and 
sure basis of S. tamnoides was the description and plate of 
Smilax Bryoniae nigrae foliis, caule spinoso, baccis nigris of 
Catesby, Carol. i. 52, t. 52. Linnaeus took his brief diagnosis 
primarily from Catesby: ‘“SMILAX caule aculeato tereti’’ and 
the species was placed by him in the section *Caule aculeato, 
tereti. There can be no question that the type of S. tamnoides 
was the Catesby plate. 

Catesby’s description was vivid: 


This plant shoots forth co many pliant thorny stems; which, when at 


full bigness, are as big as a walking cane, and jointed; and rises to the 
height usually of ey ok, climbing upon and spreading over the adja- 
cent Trees and shrubs. In Autumn it produces clusters of black 


— berries, hanging pendant to a foot-stalk, above three inches long,” 
Cc. 

The only possible competitor is S. Bona-noz, var. hederaefolia, 
but that has square or at least 4-angled stems (Catesby’s shrub 
terete), the leaves are strongly reticulate with prominent cross- 
veins (Catesby’s not), and the fruiting peduncles are 1-3 cm. 
long, merely ascending to divergent or rarely recurved (Catesby’s 
shrub with pendulous fruiting peduncles 5-6 cm. Jong, with 
pedicels much longer than in S. Bona-noxr). Catesby’s plate is 
a beautiful match for the terete-stemmed plant, with relatively 
thin though firm, and delicately veined often panduriform leaves, 
elongate, arching and finally drooping peduncles (up to 6.5 cm. 
long) and long pedicels, which occurs on the Coastal Plain from 
Florida to southeastern Virginia (perhaps farther north). This 
plant has been considered a variety of the wide-ranging, conti- 


1944] Fernald,—Flora of Eastern North America 39 


nental S. hispida, from which it differs in the development of 
some or many panduriform leaves. It has been designated in 
the Gray Herbarium by a critical student of the group as a 
southeastern variety of S. hispida, with an unpublished name 
indicative of its southern occurrence. In the Gray Herbarium 
there is no material of S. tamnoides, var. hispida! (S. hispida), 
the continental plant with leaves regularly ovate, from the 
Coastal Plain south of Virginia, where it extends from the interior 
down the calcareous valleys. All specimens from the Coastal 
Plain of Florida, Georgia and South Carolina have some or all 
of the leaves panduriform and are typical S. tamnoides. I have 
seen no such material from North Carolina. 

Pursh, treating Smilax tamnoides as herbaceous (meaning S. 
tamnifolia Michx.) described as S. pandurata [he said us] Pursh, 
Fl. Am. Sept. i. 251 (1814), a species in his **Caule fruticosa; 
ramis teretibus. Pursh’s brief description could have applied 
only to typical S. tamnoides: 


14. S. aculeata; foliis ovato-panduraeformibus acumi- panduratus. 
te ‘S-nervibus, pedunculo communi petiolo duplo 


In sh wes New Jersey to Carolina. k. July. 
v.v. Leaves smooth and shining on both sides. 
Notre on SmMILAx LANCEOLATA.—S. lanceolata L. Sp. Pl. 1031 
(1753) was published as follows, under *Caule inermi, tereti: 


13. SMILAX caule inermi tereti, foliis inermibus lanceolatis. 


Smilax, viticulis aspe s angustis laevibus 
nullis : auriculis pret ic ee ae 349 i 110. f. 4? Raj 
supp 
Habitat in Virginia 
Differt nostra a E Fiebaakand aculeorum absentia. 


Linnaeus himself, according to the late Dr. B. Daydon Jack- 
son, had no material and my efforts to secure a photograph of the 
Hortus Cliffortianus and Gronovian plants have been unsuccess- 
ful. The Plukenet figure, cited by Linnaeus in Hortus Clifforti- 
anus without query, is available and it is significant that Linnaeus 
stated that S. lanceolata differed from Plukenet’s figure in lacking 
prickles. It should also be noted that S. lanceolata was from 


'Smitax Tamnores L., var. hispida (Muhl.), stat. nov. S, hispida Muhl. ex Torr. 
Fl. N. Y. ii, 302 (1843). 


40 Rhodora [FEBRUARY 


Virginia. During ten seasons in the field my companions and I 
have sought, in vain, for the familiar rather thin-leaved plant 
with 3 stigmas and dark red berries which regularly passes as 
S. lanceolata. This species occurs in our extreme Southern 
States, Florida to Texas, north into South Carolina and Arkansas. 
I have seen no specimens of it from north of South Carolina. 
Everywhere in southeastern Virginia the coriaceous-leaved, 
evergreen S. laurifolia L. occurs, varying tremendously, with 
leaves from elliptic-oblong to narrowly lanceolate, obtuse to 
acuminate, the blades anywhere from 0.6-2 dm. long and 1-7.5 
em. wide. But it is always S. laurifolia, with intermittent 
tendrils, flowers expanding in autumn, black fruit ripening the 
second year, stigma and seed 1, etc. In its narrowest-leaved 
extreme, such as Fernald & Long, nos. 6778 and 13,919, its leaves 
are narrowly lanceolate and very close to those of the Plukenet 
figure. Of two sheets of one number retained at the Gray Her- 
barium, one has the branches as fiercely prickly as in Plukenet’s 
illustration, while the second has them prickleless, as described 
by Linnaeus. Furthermore, the Plukenet figure is quickly 
matched by vigorous, prickly, narrow-leaved vegetative sprouts. 
It is most difficult for one who knows southeastern Virginia to 
believe that the plant inadequately described by Linnaeus as S. 
lanceolata was anything but the narrowest-leaved S. laurifolia, 
the type of which, as shown by the photograph before me, was 
the broad-leaved phase of the species. 

Of great importance in interpreting the confused ideas of 
Linnaeus is the fact, that in Species Plantarum, after citing the 
reference to his brief account in Hortus Cliffortianus (which was 
based wholly or in part on the Plukenet figure) he appended 
“Gron. virg. 120.” Now, if one looks in Gronovius he will find 
Smilax treated on pp. 120 and 193. On p. 120 two species have 
completed treatments, the third treatment was unfinished (as 
printed) and as an erratum was given on p. 193, “Ibidem linea 
penultima lege.”’ The first species on p. 120 had cordate-ovate 
leaves and was cited by Linnaeus under his S. Pseudo-China. 
Gronovius included under it “Smilax humilior, floribus dilute 
luteis, baccis rubris. Clayt. n. 82’, which is, of course, S. 
Walteri Ell. (photograph of the Clayton plant before me). The 
second species in Gronovius, on p. 120, had “caule angulato 


1944] Fernald,—Flora of Eastern North America 41 


aculeato, foliis dilatato-cordatis’ and included “Smilax late 
scandens Bryoniae nigrae foliis, caule spinoso . .' . baccis 
atro-purpureis. Clayt. n. 81”; i. e., with angulate stems, leaves 
of Bryony and purple-black barred: it was one of the variations 
of S. Bona-nox L., surely not the red-berried so-called “SS. 
lanceolata”. The third species, revised on p. 193, with the 
admonition to substitute for the account on p. 120, was 

SMILAX caule spinoso tereti: folits ovato-oblongis, trinervits, inermibus. 

Smilax laevis Lauri folio, baccis nigris. Catesb. Hist. Carol. Vol. oT: WS. 

Smilax sempervirens Lauri folio crasso: floribus parvis herbaceis; 

caule spinis rigidissimis armato; baccis nigricantibus. Clayt. n. 617. 

Smilax laevis Lauri folio of Catesby was the best kind of S. 
laurifolia L.; in fact it was cited by Linnaeus in publishing that 
species. So too was the Smilax caule spinoso tereti (with his 
own changes to “‘caule aculeato tereti’’, etc.) of Gronovius, p. 193. 
These, in fact, were the only bases of S. laurifolia L. (1753). 
Surely, there is not much left to stand for S. lanceolata L. as a 
distinct species! One sometimes doubts the wisdom of starting 
our nomenclature of American plants with Linnaeus (1753). It 
is almost an exceptional North American species about which he 
was not hopelessly confused. 

As pointed out by Morong, Bull. Torr. Bot. Cl. xxi. 434 (1894), 
Asa Gray, thinking of Smilaz lanceolata in the usually accepted 
but obviously erroneous sense, as the southern, thinnish-leaved 
plant with 3 stigmas and dark red 3-seeded berries, “examined 
in 1881 the specimens in the Herbaria of Enslen and Sherard 
upon which Pursh founded his species [S. ovata Pursh, Fl. Am. 
Sept. i. 249 (1814)], and declares positively that they are Smilax 
lanceolata’. S. lanceolata sensu Gray, Morong, Small and 
others is, then, S. ovata Pursh, whose description was good 
except for ‘berries black”. In the herbarium they always lose 
their red color. Unfortunately, however, S. ovata Pursh (1814) 
is antedated by a different S. ovata Jaume St.-Hilaire (1800). 

The first available name for Smilax ovata Pursh, not Jaume 
St.-Hilaire, seems to be S. SMaLLut Morong in Bull. Torr. Bot. 
Cl. xxi. 484 (1894). Morong described it from young flowering 
branches, with staminate flowers only, but a very full sheet of 
isotypic material before me is surely of the shrub erroneously 
passing as S. lanceolata. Small himself, in his Manual, reduces 


42 Rhodora [FEBRUARY 


to S. lanceolata sensu authors, not L., both S. Smalliz Morong 
and S. cinnamomifolia Small in Bull. Torr. Cl. xxv. 609 (1898), 
the latter described as having ‘‘bluish-black”’ berries. In his 
Manual, where he reduces S. cinnamomifolia, the berries are 
called ‘‘dull-red”. The type of S. cinnamomifolia, Heller, no. 
4109, from Arkansas, was in ripe fruit. The isotype in the Gray 
Herbarium shows the crushed berries darkened, just as are the 
crushed and dried fruits of the admittedly red-berried shrub. 

Until Smilax Smallii (S. lanceolata sensu most auth., not L.) 
is really collected in Virginia it may safely be excluded from the 
flora of the state. 


Four or RAFINESQUE’S SPECIES OF SISYRINCHIUM.—In his 
Autikon Botanikon (Cent. V), 65 and 66 (1840) Rafinesque, with 
the unusual clarity which contrasted much of his work in this 
rare volume with the slipshod technique of most of his publica- 
tions, described four species of Sisyrinchium. Since these were 
overlooked by Bicknell in his detailed study of the genus and 
find no mention in current floras, it is desirable to note their 
probable identities. The original descriptions are copied below. 


481, Sisyrincuium flecuosum Raf. caule dichot. flexuoso anceps, fol. 
brevis ensatis acutis vix nervosis, spathis lanc. diphylis ineq. fl. sube 
3-4fl. capsulis ovoideis truncatis torulosis—Arkanzas and Texas, semi- 
pedal, leaves 1 or 2 inches, specimens in fruit o 

482, Sisyr. tenuifolium Raf. caulib. cespitosis genicul. filif. 2ang. fol. 
tenuis ier? “BY mre lin. filif. sip a aoe subeq. lanceol. 


483, S: Stee Raf. Scaposum, fol. lin. lanc. acutis vix nervosis, 
scapis "fol oe lato bialatis, spathis ineq. bivalvis lanc. 2—4floris equante, 
petalis retusis sos aya caps. obovatis—Florida, found by Baldwin, 4 to 
6 inches high, annual leaves 2-4 inches, one line broad, flowers large white. 

484, Sisyr. oh Raf. scaposum fol. lin. angustis striatis nervosis 
acutis, scapis fol. longior bialatis, alis striatis, spathis sepe coloratis 
3valvis, 2int. brevis subeq. 1 ext. _longissima lane. plicata, 6-Sfloris. 
petalis retusis cuspid. capsulis globosis—Alab. Kentucky &c, 6 to 8 inches 
high, leaves 3 to 5, half line wide, alas size of S. anceps, snowy white, 
spathas more or less colored of re 


Species no. 481, Sisyrinchium fleruosum Raf., not Spreng. 
(1825), is undoubtedly S. minus Engelm. & Gray in Bost. Journ. 
Nat. Hist. v. 263 (1845). The description might well have been 
based on the type of S. minus, and the description of S. minus in 


1944] Fernald,—Flora of Eastern North America 43 


Small’s Manual emphasizes the short leaves (2-7 cm. long) and 
the capsule “corrugated”. Rafinesque said ‘capsulis 
torulosis.”” Although S. flexrwosum Raf. was the earliest name 
for this southwestern species, it was antedated by S. fleruosum 
(L.) Spreng. 8S. mrnus stands. 

Species no. 482, S. tenuifolium Raf., suffers the same fate. 
Rafinesque gave a good diagnosis of S. Langloisii Greene, Pit- 
tonia, iv. 32 (1899). Compare with Rafinesque’s diagnosis these 
phrases from Greene’s description of S. Langloisii: ‘ Densely 


tufted and very slender . . . : leaves very narrowly linear, 
2 to 4 inches long . : stems scarcely ancipital, rather 
subterete [Raf. said “filif. os ue . . peduncles 
each with a single _— fawdiduercd sntles, its bracts equal 
or nearly so . . . perianth large for so small a 


plant, blue.” Characteristic plants of S. Langloisii closely 
match the account of S. tenutfolium. Rafinesque, however, was 
again out of luck, for there was an earlier S. tenuifolium Humb. & 
Bonpl. ex Willd. (1809). S. Lanetorsi stands. 

Species 483, S. floridanum Raf., is more puzzling, on account 
of its large “white” flowers, for, so far as I can make out, no 
ee white-flowered species occurs in Florida, unless possibly 

S. albidum Raf. Atl. Journ. 17 (1832). The “scapis 
lato bialatis’’, their height and the leaves 1 line broad would do 
for S. albiiem. It may so rest for the present. S. floridanum 
Raf. (1840), however, invalidates the name S. floridanum Bickn. 
in Bull. Torr. Bot. Cl. xxvi. 222 (1899), given to a very slender 
and pale plant first collected by Nash in high pineland of Lake 
County, Florida. Various coarser and darker-drying plants 
probably referable to S. arenicola Bickn. (incl. S. fibrosa Bickn.) 
have been wrongly distributed as S. floridanum, but an ISOTYPE 
(Nash, no. 13) and such material as Blanton, no. 6952 stand 
clearly apart. Since there is already a S. Nashii Bickn. his S. 
floridanum may appropriately be called 

Sisyrincuium Bicknellianum, nom. S. floridanum Bickn. 
in Bull. Torr. Bot. Cl. xxvi. 222 (1899), not Raf. Aut. Bot. 66 
(1840). 

Rafinesque’s species no. 484, S. niveum from Alabama and 
Kentucky, can scarcely be anything but his already described 
S. albidum Raf. Atl. Journ. 17 (1832). 


44 Rhodora [FEBRUARY 


QuERCUS INCANA Bartram, Trav. 378 (1791). Q. cinerea 
Michx. FI. Bor.-Am. ii. 197 (1803). 

I fully concur in the decision of Dr. Francis Harper in Bartonia, 
no. 22: 3 (1943), that Quercus incana is Q. cinerea. Bartram’s 
description was very brief: ‘‘Q. incana, foliis ovalibus integerri- 
mis subtus incanis.” Bartram. was describing the trees and 
shrubs of high gravelly ridges of the upper Ogeechee River in 
central-eastern Georgia: ‘“‘This day’s journey was for the most 
part over high gravelly ridges, and on the most elevated hills 
appeared emerging out of the earth, rocky cliffs of a dark reddish 
colour; their composition seemed to be a coarse, sandy, ferru- 
ginous concrete . . . The trees and shrubs common on 
these gravelly ridges are as follows, Diospyros, Quercus rubra 
[meaning falcata], Q. nigra, Q. tinctoria or great Black Oak, Q. 
alba, Q. lobata, post White Oak, Q. incana, . . . Pinus 
lutea, Pinus taeda, . . . Pinus palustris’, etc., a rather 
characteristic group of oxylophytic and more or less xerophytic 
species. These are the regular associates of Quercus cinerea, 
a characteristic species of dry sands and gravels, which, according 
to R. M. Harper, Phytog. Sk. Altamaha Grit Reg. of Ga. Ann. N. 
Y. Acad. Sci. xvii. pt. I. 249 (1906), “Ranges from the fall-line 
sand-hills [near Bartram’s station] almost to the coast”. Ordi- 
narily the leaves of Q. cinerea are oblong, but they vary to oval 
or elliptic in broader-leaved individuals (Small says, ‘blades 
elliptic, varying to lanceolate or oblanceolate . . . Sand- 
ridges, dry woods, and pinelands.’’) 

Trelease, The American Oaks, Mem. Nat. Acad. Sci. xx. 113 
(1924), guessed that Q. incana Bartr. is the Live Oak, Q. vir- 
giniana Mill., but Bartram was thoroughly familiar with Live 
Oak; in fact he mentioned it more often than any other species 
and would not suddenly, in describing the characteristic covering 
of the inland Sand Hills, proceed to describe it as a novelty. 
Quercus virginiana, the true Live Oak, so familiar to Bartram, is, 
from Virginia to Georgia, an evergreen tree primarily of the 
outermost Coastal Plain. Harper, op. cit., did not know it on 
the Altamaha Grit which, at its northern boundary, approaches 
Bartram’s station of Q. incana. The only representative of 
Q. virginiana in Harper’s area is Q. geminata Small, which 
Bartram would have included under Q. virginiana and which is 


1944] Fernald,—Flora of Eastern North America 45 


found only well to the south of Bartram’s area and in a different 
soil. So far as I can find, Q. cinerea is the only species of the 
“fall-line sand-hills’” which Bartram could have meant by Q. 
ineana.! 


QUERCUS MARITIMA Bartr. Trav. 164 (1791). From the 
“endless wild desert, the upper stratum of the earth of which is 
a fine white sand, with small pebbles, and at some distance 
appears entirely covered with low trees and shrubs of various 
kinds, and of equal heighth’’, along St. John’s River north of 
Lake George, eastern Florida. From the description alone, 
“foliis obcuneiformibus obsolete trilobis minoribus”, I do not 
venture to guess its identity. Too many shrubby species of 
Florida are candidates. Bartram’s Q. maritima (1791), however, 
antedates Q. maritima (Michx.) Willd. (1805) and Raf. (1838). 
The last two were caught by Index Kewensis, the earliest of the 
series not. 


SILENE scABRA Raf. Aut. Bot. 18 (1840), from “barrens of 
West Kentucky”, is very definitely 8S. recia Sims (1814). 
Rafinesque’s name, moreover, is invalidated by S. scabra Kit. 
(1814). 


PRUNUS NEMORALIS Bartr. Trav. 408 (1791) should be added 
to the synonymy of Padus caroliniana Mill. Dict., no. 6 (1768) 
and of PrRuNUs CAROLINIANA (Mill.) Ait. Hort. Kew. ii. 163 
(1789). 

Bartram’s species, not entered in Index Kewensis, was from 
above Mobile, Alabama: “ Prunus nemoralis, floribus racemosis, 
foliis sempervirentibus, nitidis.”’ 

SEBASTIANIA fruticosa (Bartram), comb. nov. Stillingia fruti- 
cosa Bartr. Trav. 476 5 (1791), Stil. ligustrina Michx. Fl. Bor.-Am. 
ii. 213 (1803). Sebastiania ligustrina (Michx.) Muell.-Arg. in DC. 
Prodr. xv.? 1165 (1866). 

Index Kewensis cites Stillingia fruticosa as starting with 
Spreng. Syst. iii. 805 (1826), while Sprengel credited it to Mi- 
chaux (1803) who had no such name but described Stil. ligus- 

1 terpreta’ incana Bartr. (1791) 
Ng one et caph eae Rageacdieniy pista pei ene ae See | Beng. 104 Jose oars and 


Fl. Ind. iii. 642 (1832),. The Indian species should, apparently, be called Q. lanata 
Sm. in Rees Cycl. xxix. no. 27 (1819). 


46 Rhodora [FEBRUARY 


trina from “sylvarum umbrosis, ad ripas amnium Carolina et 
Georgia.” Bartram, however, describing the “very curious and 
beautiful flowering and sweet scented shrubs” northwest of Cape 
Fear, North Carolina (near Wilmington), enumerated many of 
them: “particularly Callicarpa, Aesculus pavia, . . . Sty- 
rax, Stewartia, . . . Stillingia fruticosa, foliis lanceolatis, 
utrinque glabris, fructu tricoceo”. Though brief, the diagnosis 
of this characteristic shrub of the Coastal Plain, northward to 
southeastern North Carolina, is sufficient. M. A. Curtis in his 
Catalogue of the Indigenous and Naturalized Plants of North 
Carolina (1867) cited Stil. ligustrina as occurring in the ‘‘ Low 
Dist.”’, 7. e. on the Coastal Plain, and Wood and McCarthy in 
their Wilmington Flora, covering the country drained by both 
the Northeast and the Northwest Cape Fear Rivers, also listed it. 
In both lists the species is recorded on the authority of Dr. J. 
F. McRee, who was born at Wilmington and later had a planta- 
tion some miles north of that city. 

CYRILLA RACEMIFLORA L., var. subglobosa, var. nov. (TAB. 
813, Fic. 1 et 2), foliorum venis subtus vix prominulis; sepalis 
ovatis 1 mm. longis; fructibus depressis subglobosis sulcatis; 
stigmatibus perbrevibus. —VIRGINIA: wooded swamp along Mill 
Creek, about 1 mile north of Skipper’s, October 14, 1938, Fernald 
& Long, no. 9600 (rypx in Herb. Gray.; 1isorype in Herb. Phil. 
Acad.). 

Typical Cyrilla racemiflora has a characteristic conic-ellipsoid 
or -ovoid drupe definitely longer than broad, though in excep- 
tional specimens only slightly longer. In ripe material the 
drupes (Fic. 3) show a continuous rounded surface with little or 
no sign of furrowing and the styles and stigmas are rather 
prominent, the lanceolate to lance-ovate sepals are 1—-1.8 (usually 
1.5) mm. long, their sharp tips evident beyond the half-diameter 
of the fruit. In typical C. racemiflora, furthermore, the mature 
foliage is usually prominently reticulate-veiny beneath. Var. 
subglobosa, the type in ripe fruit, collected in mid-October, has 
the reticulum of the lower, as well as the upper, leaf-surface 
rather faint; the sepals are the shortest in the species, 1 mm. 
long and ovate; and the depressed-subglobose drupes (Fras. 1 
and 2), definitely as broad as or broader than high, so that the 
sepals are relatively hidden, are broadly rounded to an almost 
emarginate summit and with a deep longitudinal furrow running 


1944] Fernald,—Flora of Eastern North America 47 


from base to apex and marking the boundaries of the 2 carpels; 
the style and stigmas are the shortest in the species. 

Although occasional specimens of the wide-ranging and more 
typical Cyrilla racemiflora approach in one character or another 
the shrub from near Skipper’s, the latter combines so many of 
these exceptional characters that I am designating it as a variety. 
Flowering material may show other differences. 

In shape of drupe Var. subglobosa suggests the fruit described 
by Rafinesque for his Cyrilla polystachya Raf. Aut. Bot. 8 (1840), 
with ‘‘capsulis subglobosis”. That shrub of Louisiana and 
Florida was described, however, as having the “fol. 
imis . . . acutis . . . racemes 6 to 8 inches”. It is 
most probably a phase of the narrow-leaved and small-fruited 
C. parviflora Raf., 1. ¢., which differs in several characters from 
C. racemiflora. 

In PLATE 818, Fics. 1 and 2 are of TyPE of CYRILLA RACEMIFLORA L., var 
SUBGLOBOSA: FIG. 1, portion of branch, X 1; Fic. 2, portion of fruiting raceme, 

. Fig. 3 is a portion of a fruiting raceme of typical C, RACEMIFLORA, X 5, 
from .west of Winfield’s Mill, Dinwiddie County, Virginia, F & Long, 
no, . 

AESCULUS SYLVATICA Bartr. Trav. 476 (1791). Ae. neglecta 
Lindl. Bot. Reg. xii. t. 1009 (1826); Sargent in Journ. Arn. Arb. 
v. 43 (1924), 


The tree known as Aesculus neglecta is characteristic of wood- 
lands along streams in central and eastern North Carolina and 
southeastern Virginia. In the latter state it reaches the inner 
border of the Coastal Plain along the Meherrin River, below 
Emporia. From North Carolina Sargent cites specimens from 
the Piedmont eastward to the fall-line sand-hills or the inner 
margin of the Coastal Plain: Alamance, Orange, Durham and 
Wake Counties, on Cape Fear drainage or (the Wake County 
station) on the Neuse. According to Lindley’s original account 
and plate the flowers are cream-colored or pale-yellowish, “and 
beautifully veined with red; the lateral petals are also pinkish”. 
The red and pink coloring may sometimes be nearly absent but 
the non-glandular pedicels and calyx, which characterize the 
species, hold. Bartram, ascending “the North West of Cape 
Fear’ River, found “Aesculus sylvatica, floribus ex albo et 
carneo eleganter variegatis, caule arboreo”. What else could it 
have been than Ae. neglecta, with “flowers . . - beautifully 


48 Rhodora [FEBRUARY 


veined with red’’, the species characteristic of eastern North 
Carolina and adjacent Virginia, South Carolina and Georgia, 
which Sargent cites from definite stations on Cape Fear River? 
The name was not recorded in Index Kewensis. 

Neither did Index Kewensis pick up Aesculus floridana Bar- 
tram, 1. c. 401 (1791) from northern Florida, described: ‘‘ Aesc. 
Florid. ramis divaricatis, thyrsis grandis, flosculis expansis 
incarnatis.”” What he had I do not know. He was definitely 
distinguishing it from Ae. Pavia which he enumerated in the 
same list. If the latter species is ever subdivided Bartram’s 
species must be taken into account. 

VITIS CAMPESTRIS Bartram, Trav. 400 (1791), not Nutt. in 
Fenses’é s Cat. (1813) nor Baker in Mart. Fl. Bras. xiv’. 200 (1871). 
V. rotundifolia Michx. F]. Bor.-Am. ii. 231 (1803). 

Vitis rotundifolia, the Muscadine, and its pale-fruited form, the 
Scuppernong, are among the most familiar and most prized wild 
grapes of the Southeastern States. When fully ripe the large 
grapes are intensely sweet and are harvested in great quantities 
for preserving. Unlike the great majority of our grapes, this 
species has few and simple tendrils, instead of many and forking 
ones. Consequently, although it may climb high when oppor- 
tunity presents, it is most often sprawling over bushes or reclin- 
ing on the sand or gravel, so that, in such habitats, its often too 
tempting and almost over-sweet berries are gathered by stooping. 
Thus, Elliott said: ‘‘This species of vine varies much in size, 
sometimes ascending the loftiest trees, more frequently humble” 
(Sk. ii. 687) and Darby said “sometimes not climbing’. As to 
the fruit, Mohr (Pl. Life Alab. 6: 3), pays it the compliment 
which he accords no other species of Alabama: “Important as a 
table and wine grape” and, after speaking of the ‘Berries large, 
plum-purple”’, he adds: “ Plants of spontaneous growth have been 
observed near Mobile bearing light amber-colored berries.” 

Bartram’s party, crossing northern Florida en route to Mobile, 
where “ Our road now for several miles led us near the Alabama”’, 
eventually left “the Alabama bearing away Southerly, and 
enter[ed] a vast open forest which continued above seventy 
miles, East and West, without any considerable variation . 
During our progress over this vast high forest, we crossed eiten- 
sive open plains, the soil gravelly, producing a few trees and 


YR EOL IE EP IY GG LT EN LI 


1944] Fernald,—Flora of Eastern North America 49 


shrubs or undergrowth, which were entangled with Grape vines 
(Vitis campestris) of a peculiar species [Vitis rotundifolia Michx. 
is so “peculiar”? in many characters that Small makes it a 
separate genus, Muscadina], the bunches (racemes) of fruit were 
very large, as were the grapes that composed them, though yet 
green and not fully grown, but when ripe are of various colours, 
and their juice sweet and rich. The Indians gather great 
quantities of them, which they prepare for keeping, by first 
sweating them on hurdles over a gentle fire, and afterwards dry 
them on their bunches in the sun and air. and store them up for 
provisions: these Grape vines do not climb into high trees, but 
creep along from one low shrub to another, extending their 
branches to a great. distance horizontally round about [Bailey, 
Gent. Herb. iii. 238, says of V. rotundifolia “climbing to 100 
feet over bushes and trees’’], and it is very pleasing to behold the 
clusters pendant from the vines, almost touching the earth, 
indeed some of them lie upon the ground.” I have many 
times gathered from near the ground or from the surface of 
sand-hills or/dunes and eaten more than my fill of the “very 
large . . . grapes [Michaux described his V. rotundifolia 
baccis magnis”’], . . . their juice sweet and rich” of V 
rotundifolia. Not only did the Indians of northwestern Florida 
and of Alabama gather and dry the large and sweet grapes “and 
store them up for provisions.’’ Their successors in occupation 
of the region continue the custom, with obvious refinements. 
In his Nature’s Garden for Victory and Peace, revised (Bull. 
Tuskegee Inst. Alab. no. 42), the late Dr. George W. Carver, 
ignoring the other 5 (and sour-fruited) species of Vitis, concen- 
trated on the Muscadine, giving explicit directions for cleaning 
and preparing the fruits, after which ‘“‘They may be dried whole 
or made intoaleather . . . Imuchprefertheleather . . 
ofafine flavor . . . I wish every housewife would try this.” 
To those who intimately know the Muscadine and the Skupper- 
nong, their growth-habit and fruit there can be no serious ques- 
tion that V. campestris Bartr. (1791) is the same as V. rotundi- 
folia Michx. (1803). The change of name would have been 
made a half-century ago if Bartram’s species had not been 
ignored by the editors of Index tires! 

AMMANNIA TERES Raf. Aut. Bot. 39 (1840). A. Koehnei 
Britton in Bull. Torr. Bot. Cl. xviii. 71 (1891). 


50 Rhodora [FEBRUARY 


Rafinesque certainly had the plant of fresh to brackish tidal 
shores of Virginia to Florida (once in the Hackensack marshes 
of New Jersey) which Britton described as Ammannia Koehnet. 

Rafinesque’s description was good: 

296, Am. teres Raf. caule teres vix ramoso fol. obl. obt. carnosis, caulinis 
basi cord. ad medio. angustatis, rameis cuneatis brevis, fl. sepe solit— 
Virg. ad Carol. pedal, leaves twice as broad as in last, broader at both ends. 

This necessitates the new combination: 

A. TERES, var. exauriculata (Fernald), comb. nov. A. Koehne 

var. exauriculata Fernald in RHopoRA, xxxviii. 437, tab. 449, = 
4 and 5 (1936). 

The “last”, referred to above by Rafinesque, was his AMMAN- 
NIA MULTICAULIS, 1. c. (1840). This was evidently fRotala 
ramosior (L.) Koehne, with the virgate stems simple, ‘‘fol. 
linearib., acutis, fl. sepe vertic—Virginia, pedal, leaves uncial’, 
while no. 294, A. LINEARIFOLIA, cited as the same as A. ramosa 
of authors, but with “fol. linearib. obt. nonnulis basi dilatatis”’ 
was ROTALA RAMOSIOR, Var. INTERIOR Fern. & Grisc. 

A. LONGIFOLIA Raf. 1. c. (1840) from its excellent description 
and the region, “Arkanzas and Louisiana,’”? was A. COCCINEA 
Rottb. (1773); and A. pirrusa Raf. 1. c. (1840) is apparently A. 
AURICULATA Willd. (1806). 

None of these species of Rafinesque are recorded in Index 
Kewensis. 

HERACLEUM MAXIMUM Bartr. Trav. 344 (1791). H. lanatum 
Michx. Fl. Bor.-Am. i. 166 (1803). 

It is with real sadness that I seem forced to relinquish so long 
established and familiar a name as Heracleum lanatum Michx. 
But the species, which in the East extends south along the 
mountains to Georgia (Small), was detected by Bartram in the 
mountains of northwestern Georgia and briefly but passably 
described: “I observed growing in great abundance in these 
mountain meadows, Sanguisorba Canadensis and Heracleum 
maximum, the latter exhibiting a fine shew, being rendered con- 
spicuous even at a great distance, by its great height and spread, 
vast pinnatified leaves and expansive umbels of snow-white 
flowers.”” We are sorry to give up the long established name; 
we should have been familiar with Bartram’s earlier one had it 
been picked up, as were several names in the same book,! in 


1 Such names as Anona pygmaea, ——- of Asimina pygmaea (Bartr.) Gray; 
Andromeda formosissima, Stewartia montana, 


Se pa Se ene EE TERE Opa ae Cage PTS PEE eee Te NOT A PwORy Mw NRTA cee sie, Se Me Se eed ee ee ee Oe eee hen Ee a eee OMe Re ee ee a Ae ne mentee le oe 


1944] Fernald,—Flora of Eastern North America ol 


Index Kewensis. Incidentally, the type of Heracleum lanatum 
consists of a portion of leaf of the Heracleum and a fruiting 
umbel of Pastinaca sativa! 


THE PLANTS PASSING AS PHACELIA HIRSUTA (PLATES 814-816). 

PuHace ia fallax, sp. nov. (Tas. 814), a P. hirsuta differt 
foliorum caulinorum lobis terminalibus cuneato-obovatis seg- 
mentis oblongo-obovatis; caulibus pedicellisque minutissime 
pilosis strigosisque; sepalis adscendentibus deinde tuberculato- 
ciliatis extus glabris vel glabratis intus glabris; seminibus 
big Nd lineato-rugosis.—Mountain-region, western Virginia 

orgia. VrirGInta: Giles County, May, 1869, Canby. 
Cusine Stone Mountain, May, 1869, Canby, May 13, 1897, 
Biltmore Herb., no. 4263, May 3, 1899, Canby & Sargent, May 
24, 1899, near summit, A. H. Curtiss, no. 6458 (TYPE si — 
Gray. i, April 16, 1932, thin soil on 'slope, E. J. Palm 
39,909; slope of Pine Mountain, 1 mile north of Tathonta, on 
8, 1934, Perry & Myers, no. 998. 

Phacelia fallax has been mistaken for P. hirsuta Nutt. in Trans. 
Am. Phil. Soc. v. 191 (1837), our PLATE 815, Fries. 1-3; and 
Brand in Engler, Pflanzenr. iv”. 65 (1913) gave under the name 
P. hirsuta a description of P. fallaz, based exclusively on the 
plant of Stone Mountain (the Biltmore and the Curtiss exsicca- 
tae), Brand specially emphasizing the tuberculate-based cilia of 
the sepals and the broad lobing of the leaves and saying: ‘‘ Von 
anderern Standorten nicht gesehen. Gray gibt die. Pflanze 
ferner an fir Missouri bis Osttexas.” The latter, the 
region from which Brand saw no material, is the real area of P. 
hirsuta Nutt. Nuttall’s species is definitely HrRsuTE; the TYPE 
or IsoTYPE (PLATE 815, Fic. 1) in the Gray Herbarium closely 
matches the original description of the species which came from 
“sylvan prairies” in Arkansas, in having the summit of the stem 
and the pedicels spreading-hirsute, the ‘pectinately-pinnatifid”’ 
leaves with linear segments, and the sepals during anthesis 
widely spreading. It is matched by a few collections from 
Arkansas, Oklahoma and eastern Texas; and Mr. Long sends me 
for checking a characteristic specimen, collected as an adventive, 
near Telford, Pennsylvania, June 14, 1901 by Dr. C.D. Frets. 
The contrasts between the two species are given below and in 
PLATES 814 and 815. 


52 Rhodora [FEBRUARY 


P. urrsuTA: principal cauline leaves deeply or poeeiely ya sor 
with linear to linear-lanceolate segments; summit of ste pe 

spreading-hirsute as well as pilose; sepals during anthesis setlde ace 
to slightly reflexed, later ascending, their margins ciliate-hirsute with 
mostly slender-based hairs, the upper surfaces strigose. PLATE 815, FIGs. 


P, FALLAX: principal cauline leaves with terminal cuneate-obovate 
lobe rarely cleft to the middle, the broader segments oblong to oblong- 
obovate; stem and pedicels minutely pilose, sometimes also strigose; 
sepals i in anthesis appressed-ascending, their margins when mature ciliate 
with tuberculate- or pustular-based hairs, their upper surfaces glabrous. 
PLATE 814. 

The habitat of Phacelia fallax is rarely given on the labels 
before me. Such as indicate it say thin soil or mountain-slopes 
(in Georgia on granitic rock). Small, familiar with P. fallaz, 
rather than P. hirsuta, says ‘“‘Dry soil, rocks and open woods’; 
and McVaugh!, who, like Brand, defines as the western P. 
hirsuta Nutt. the plant of Stone Mountain and vicinity with 
hairs of the sepals ‘“‘often enlarged at base and terminating in 
raised pustules or teeth on the sepals,” cites it for Georgia as a 
secondary member of the flora of the granite flat-rocks. Its 
habitat in Giles County, Virginia, is not recorded but since 
MecVaugh’s map (his fig. 1) of granitic outcrops of the eastern 
United States, showing them to extend northward across Vir- 
ginia, indicates none of them in or very near Giles County, it is 
safe to say that there it was not found on granite. The localities 
in Georgia are, quite clearly, thin and subarid acid soil. 

The locality for Phacelia fallax in Giles County, Virginia, 
cited by Gray, Syn. Fl. ii!. 164 (1878), by Brand, 1. c. (1913) 
and the basis for the inclusion of Virginia in the range of the 
composite P. hirsuta in Gray’s Manual and in Small’s Manual, 
if taken into account by MeVaugh, would have changed the tiny 
ellipse in northern Georgia (on his fig. 29) to an elongate tongue 
reaching western Virginia. 

The habitats of true southwestern P. hirsuta (Arkansas, Okla- 
homa and Texas), on the other hand, are given as follows on the 
labels in the Gray Herbarium and in the Bebb Herbarium of the 
University of Oklahoma, the latter material kindly sent me for 
examination by Dr. Milton Hopkins, in addition to Nuttall’s 
“sylvan prairies”: wet prairie, Arkansas, Bush, no. 258; field, 


1 aah The pg gt of the —_ Flat-Rocks of the Southeastern United 
States, Monogr. xiii., especially p. 158 and fig. 29 (map) on p. 163 (1943). 


1944] Fernald,—Flora of Eastern North America 53 


Ark., F. L. Harvey, no. 12; limestone, hillside, Ark., Damaree, no. 
14830; wet places, Oklahoma, McClary, May, 1935; meadow 
soil, Okla., DeVztt and Clark, no. 50; prairies, Okla., Robert Bebb, 
no. 3856 and Demaree, no. 12,646; on slough, Okla., Richard F, 
Mason, no. 127; shaded bank of slough, Okla., R. F. Mason, 
April 14, 1937; creek-banks and open woods, Okla., Houghton as 
Stevens, no. 2137; dense shrubbery along small stream, Okla., E. 
L. Little, Jr., no. 128; wooded grassy glade in river-valley, Okla., 
Hopkins, no. 2943; wooded hills, Okla., Goodman, no. 2137; 
granite hills, Okla., P. B. Sears, no. 1317; dry sandstone ledges 
and boulders in oak-hickory forest, Okla., Hopkins and Cross, 
no. 1487; sandy soil, Okla., V. Johnson, no. 116; arid hillside, 
Okla., M. Fielder, April 20, 1927. The southwestern P. hirsuta, 
then, is evidently more calcicolous and less xerophytic than the 
southeastern P. fallax. 

The Missouri dots for Phacelia hirsuta on McVaugh’s map 
(his fig. 29) were presumably based on material of P. gilioides 
A. Brand (our PLATE 815, Frias. 4-6). At least, all the Missouri 
specimens in the Gray Herbarium which were sent out as P. 
hirsuta are P. gilioides, a grayish or canescent plant with leaves 
cut into linear or linear-lanceolate segments, the stem and pedi- 
cels canescent-pilose without hirsute pubescence, the appressed 
calyx strigose, the corolla relatively small, its lobes undulate to 
dentate. It is P. eiz1o1pEs (misidentified as P. hirsuta) which 
Palmer & Steyermark assign in Missouri to “Rocky prairie, 
glades, ledges along bluffs, low rich woods, and alluvial ground 
along streams. Calciphile to circumneutral.”” The Oklahoma 
material in the Bebb Herbarium and the Gray Herbarium shows 
the following habitats: creek-bank, Stevens, no. 1377; prairies, 
Robert Bebb, nos. 3850 and 5090, Ezra Brainerd, April 12, 1908; 
prairies and wood-openings, Robert Bebb, no. 3884; woods, Mrs. 
W. L. Ducker, no. 39; shale, rocky mountain, April 6, 1941, H. 
Randel Griffith; rocky hills, Robert Bebb, no. 2720 and May, 
1935, J. E. McClary; vacant lots and waste lands, Auval H. 
Brown, no. 18; cottonfield, April 19, 1927, R. E. Jeffs. Like P. 
hirsuta, it is also calcicolous and scarcely a xerophyte. 

In Phacelia dubia! (PLATE 816) the sepals are unequal, the 


‘ Although the combination Phacelia dubia (L.) Trel. is here credited to Trelease, 
it is done so under strong mental protest. The combination was published by Tre- 
lease in Branner & Coville, Ann, Rep. Geol. Surv. Arkansas for 1888, iv. 205 (1891) 


54 Rhodora [FEBRUARY 


outer ones usually broader and longer than the others. As the 
plant occurs through southeastern Virginia, the region best 
known to Clayton, thence northward into Pennsylvania, south- 
ward to southeastern North Carolina and west to the Blue Ridge 
and less frequently to Ohio, West Virginia and Tennessee its 
outer sepals are lanceolate to lance-oblong or -linear (Fies. 1 and 
2), sometimes slightly spatulate, the inner ones from lanceolate 
to linear. In mature fruit the lower and middle calices of each 
raceme are 4.5—-7.5 mm. long and nearly to quite twice as long as 
the capsule; and the lower mature pedicels are 5-25 mm. long. 
About Nashville, Tennessee, and in Alleghany County, Virginia, 
there occurs an extreme (FIG. 4) of the species with the outer 
sepals ovate, the fruiting calyx only 4-5 mm. long, and the 
longer (lower) pedicels only 3-7 mm. long. Although small, it 
is not P. dubia, var. georgiana McVaugh, I. c. 158 (1943). That 
has narrow sepals and pinnate or deeply pinnatifid leaves, 
while the plant of Alleghany County and of the Nashville region 
has them shallowly lobed as in P. dubia, var. dubia McVaugh. 
I am calling the new plant 

P. pusta (L.) Trel., var. interior, var. nov. (TAB. 816, FIG. 4), 
calycibus maturis 4-5 mm. longis; sepalis exterioribus ovatis; 
pedicellis imis maturis 3-7 mm. longis.—Western Virginia and 
north-central Tennessee. VIRGINIA: dry roadside, Covington, 
Alleghany County, May 12, 1916, H Seite no. 4080. TEN- 


as follows: Phacelia ‘‘dubia (1..), (P. parviflora, Pursh); Harvey.’’ That was all. 
Disregarding the somewhat unusual punctuation and the ra that ‘Paueatia “qubia 
is own from so far wi nsa arvey’s material being P. hirsuta Nuttall 


cite the basonym of P, dubia, ~~. was Polemonium dubium L. Sp. Pl. 163 (1753)—a 
photograph of the rypr before m The combination Phacelia dubia (L.) Small in 


going good nature by which many slipshod combinations by Britton, Stern and 
Poggenberg, by Small (elsewhere) and by some in recent years, have been accepted 
as ‘‘validated”’ by the next author who correctly made them with proper bibliographic 


the errors become doubled. I yndicder more than once eraser — as ee of taking 
up such combinations as validly pu ed, 

d thousands of regularly used proccess ere of the ceo Siete were thus 
published. When we can again have International Congresses it may be a safeguard 
against such opportunist and inconsiderate ip stage to establish a dead-line after 
which no such combinations will be valid. If I am here I will work for such legisla- 
tion! Since the adoption of the more rane ai, rules of botanical nomencia- 
ture there is no excuse for slipshod bibliograph 


1944] Fernald,—Flora of Eastern North America 55 


NESSEE: gregarious in open shrubberies and pastures or in open 
grounds, Nashville, April, 1878, Gattinger (ryPE in Herb. ies ); 
barrens near Nashville, April, Gattinger as Curtiss, no. 2131 

One of Gattinger’s specimens was called Phacelia puntic 
Buckley in Am. Journ. Sci. xlv. 172 (1843). Buckley’s species 
has generally been considered identical with P. dubia, but it 
-came from Alabama and had the “sepals linear-oblong, acute’, 
which certainly is not the case with var. interior. I have not 
seen Buckley’s type but his description of the Alabama plant, 
with the pinnatifid leaves with obovate and abruptly acuminate 
segments and with linear-oblong, acute sepals 2/3 the length of 
the corolla, might well have been drawn from Harper, no. 82, 
from Elmore County, which is one of the paratype-collections 
of P. dubia, var. georgiana McVaugh. Some specimens, like 
one from Nashville, April, 1884, Mzss Cushman, and Biltmore, 
North Carolina, Biltm. Herb. no. 786°, are transitional between 
var. dubia and var. interior. 

In Phacelia dubia the seed (ria. 3) is rugose-reticulate, in P. 
fallax (pLATE 814, Fic. 4) transversely lineate-rugose. 

Although McVaugh delimits his tabulation of plants occurring 
on granite flat-rock at the North Carolina-Virginia boundary,’ 
his maps of ranges often extend farther north. In view of the 
abundance of Phacelia dubia on the Coastal Plain of Virginia 
(in Henrico, Prince George, Sussex, Southampton and Greens- 
ville Counties, and presumably the type-locality of Clayton) 
and McVaugh’s correct reference to it on his p. 158 as “abundant 
in many parts of Virginia”, it is difficult to interpret his map of 
the range of P. dubia which shows only 3 or 4 stations in the 
state, these all on the granites of the northwestern section of the 
state. The inclusion of the many stations 100-135 miles to the 
southeast would make a somewhat different picture, as would 
the inclusion of Newbern at the mouth of ears co in oe 


1 Near the fall-line in south t 
in connection with McVaugh’'s study of the flora of the granite te flat-rocks farther 
south. Cheilanthes lanosa, which ‘‘is approaching here the eastern edge of its range” 
e flat- 


@ small outlying colony on a low granite hill at the outer edge of the Piedmont in 
Brunswick County,—see RHopora, xlv. 374-376 (1943). 


56 Rhodora [FEBRUARY 


County, North Carolina (specimen from M. A. Curtis) and of 
Wilmington at the mouth of Cape Fear River in southeastern 
North Carolina (see Wilmington Flora by Thos. F. Wood and 
Gerald McCarthy in Journ. Elisha Mitch. Sci. Soc. for 1885-86, 
p. 131, as P. parviflora Pursh).. In fact, M. A. Curtis, who 
intimately knew North Carolina, did not restrict P. dubia (or 
parviflora) to the 2 stations on the granitic belt of western North 
Carolina. By him it was not known from the mountains but 
was thought to be characteristic of the ‘‘Low Distr.’’, 7. e. the 
Coastal Plain (see M. A. Curtis, Geol. and Nat. Hist. Surv. N. 
Ci Pt. A. ents 42 (1867)). 

PuaTeE 814 is of PHaceLia FALLAX, all figs. from the Typx: Fic. 1, portions 
of plant, 1; gee 2, young inflorescence, * o 3: FIG. 3, portion of mature calyx, 
xX 10; ria. 4, ‘seed, X 10. 

In pLaTE 815, Frias. 1-3 are of PHACELIA HIRSUTA: FIG. 1, TYPE Or ISOTYPE, 
X 1; rig. 2, portion of eee, X 3, from northwestern Arkansas, F. L. 
Harvey, no cet = 3, seed, X 0, from Prescott, Arkansas, Bush, no. 258. 
Fias. : LIOIDES: FIG en Tian. < 1, from Joplin, Missouri, EB. J 
Palmer, no. 1879 A (as P. dubia) : FIG. 5, young inflorescence, < 3, from same 
no.; Fic. 6, seed, X 10, from near Muskogee, Oklahom a, Robt. Beb bb, no. 3850 


FIG. ’ 

Virginia, Fernald & Long, no. 10,015; ric. 3, seed, X 10, from no. 10,015. 

Fic. 4, var. INTERIOR, portion of mature inflorescence, X 3, from TYPE. 
HYDROLEA UNIFLORA Raf. Aut. ee 34 (1840) must replace 

H. affinis Gray, Man. ed. 5: 370 (18 

Rafinesque got hold of material en axillary spines, which 
is not unusual, both this glabrous species of the Interior and 
Southwest and the pubescent H. quadrivalvis Walt. having spines 
or not, the character variable. Likewise, the glabrous species 
may have solitary or glomerulate flowers. Rafinesque’s descrip- 
tion is unequivocal: 

273, Hydr. wie | Raf. caule flexuoso inerme glabro, fol. alt. lanceol. 
acutis subsessil. pedune. axil. unifl. bract. 2 obl. ineq. cal. lin. glabris— 
Louisiana on Red - Rives, sent me by Torrey as Hydr. 4valvis, but unlike 
the last, smooth, not spinose, leaves smaller, biuncial, flowers bluish 
white, anthers blue shaped like x, caps. bivalve 

LycoPUs AMPLECTENS Raf. Aut. Bot. 115 (1840). L. euro- 

aeus, Var. sessi ba Gray, Man. ed. 5: 345 (1867). L. sesszli- 
folius Gray, Proc. Am. Acad. viii. 285 (1870). 

There can be no dake that Rafinesque had the plant known for 
three fourths of a century as Lycopus sessilifolius. His descrip- 
tion is beyond dispute, for he restricted the genus to species 
with acerose teeth: 


eee ee ee 


ee ey ee ee ee 


a A ee a ee TN ee eee | ee 


a ae a ee ee ae ee ee eee 


1944] Fernald,—Flora of Eastern North America 57 


834, Lyc. amplectens Raf. glaber, caulib. virgatis simpl. angulis obt. fol. 
arcte ‘sessil. subamplex. basi dilatatis oblongis acum. remote serratis, 


beneath, this sp. differs from all by obtuse angles and leaves not attenuate 
at base. 


Plate 807 
Rhodora 


Photo. B. G. Schubert, 


ai 4 . YPE 
de ae vices 4 Abed $; 5C 10; all trom + 
) ee Rieti ne ; 2, X 1; FIG. 3, X 3; Fias. 4 an 
PANICUM BENNERI: FIGS. 1 and “4 SS 


Rhodora Plate 808 


: and 4 
YUCCA FILAMENTOSA: FIG. 1, floy x 1, from TYPE; FIG, 2, leaves, X 14; FIGS. 3 and 4, 
bee 


Photo. B. G, Schubert. 
flowers, X 1; FIG. 5, ovary, style res Eishenits x 5; fig. 6, -apsules, X 
| 


Rhodora Plate 809 


Photo. B. G. Schubert. 


cca Sy er eat ee pe 1, leaves and flowers, X 14, from TYPE; FIC 2, basal rosette, 
: greatly reduced; FIG and 4, flowers, < 1, from TY a. FIG. 5, Ovary, ay le and filaments, 
Xx 3; Fig. 6, capsu ule, aoe 


Rhodora 


Plate 810 


Si hiun 
i. 
Tob. wh has mm 


i ies Mt Cean: 


btllin In 
AA Falmer Sup 
SF Hess Burnley 


Photo B. G. Schubert. 


TRILLIUM CaTESB AEI> FIG. 


, X 1, after photograph by C. A. and Una F. Weather 


rby 
LIUM NERVOSUM:? FIG. 2 aye -SHE ET, X 4, after photogrs ph by C. A.and Una F.W platter 
3 


4% 
FIG. 3, dete ul, to show style, 


} Rhodora Plate 811 


i 
F 
' 
K 
, 


Photo, B. G, Schubert. 


UM NERVOSUM: FIGs. 1 and 2, portions, X 1, of Catesby’s plate of Solanum 
| lr ih ition; Revi he cape talo carneo 
| ax BoNA-NOX, var. EXAURICULATA: FIG. 3, leaf, X 1, from TYPE 


Rhodora Plate 812 


-=TYPE SPECIMEN 
DEt ci7ey 
PM tactic at Giant Latte wok 
: ofa PRE Oa FA te oP or 
7 raed ALD Le OW GPE 0 Come 
gle We 218 Teh: 


M5 gf} 


"Baigce a ra » 
teas: < 


mets soca aps a8 fig ved 


Photo, H. G. Fernald. 


SmMivAx Psevupo-Caina: after photograph sent by Dr. John 
Ramsbottom; vgs 2, TYPE, after pT aa sea from Linnean Society of Linwiniis seth 
reduced; 1 leaf x 1 


i { 


Sane Nach nde ae eee ergs 


I 


thodora 


Photo, B, G. Schubert. 


CYRILLA RACEMIFLORA! FIG. 3, 


TY 


Var. SUBGLOBOSA: FIG. 1, portio 
PE 


fruit, x 5 re 
n of fruiting branch, X 1; Fic. 


Plate 813 


2, fruit, & 5; both from 


Rhodora Plate 814 


Photo, B. G. Schubert. 


Co 
PHACELIA — _ figs. from TYPE: FIG vier ie! of of apo sa - ie 2, young 
inflorescence, X 3; FIG. 3, portion of mature i ~ 10; 3. 4, seed, 


# Rhodora mes 


Je Photo. B. G. Schubert. 

PHACELIA HIRSUTA: FIG. 1, TYPE or ISOTYPE, X 1; FIG. 2, portion of inflorescence, X 3; 
FIG. 3, seed, 

P. GILIOIDES: FIG. 4, plant, X 1; Fic. 5, young inflorescence, X 3; Fic. 6, seed, XK 10 


Rhodora Plate 816 


Photo. B. G. Schubert, 


PHACELIA DUBIA: FIG, 1, young inflorescence, X 3; F1G.2, portion of mature inflorescence, 
X 3; FIG. 3, seed, 


Var. INTERIOR: FIG. 4, portion of mature inflorescence, X 3, from TYPE 


INDEX 
New scientific names are eae in full-face type 


Adlumia, 20; cirrhosa, 2 

jaime floridana, 48; i 47; 
Pavi : sylvatica 

Misgheman pomeany ie The Indige- 


12 

Ammannia a 50; Pr. 

50; di 50; Koehne 0, 
var. peaiioiat ta, 50; Hate Ie 
50; longifolia, Tat multica ulis, 50; 
ramo; - res Q, var 
exauriculata "60 

Andromeda formoxissi 50 

Anona pygmaea, 50 

Asimina ie te, 50 


Benner, Walter Mackinnett (dedica- 
tion of “ore to), 
Black Oak, 4 


Callicarp 

Chelanthes lanos, 55 

China rad 

Cladrastis, he 

Clintonia, 20; borealis, 20 

Convallaria globosa, 13, 14; majalis, 
1. —14; majuscula, 13, 14; montana, 


The Indigenous 
Alleghen 

Cyrilla seers ve ioe 47; polys br he 
47; racemiflora, 46, 47, pl. 813, v: 
subglobosa, 46, 47, pl. 813 


Didiplis, 20 


Diospyros, 44 

Distinetions betw een Polygonatum 
ifl and P. canaliculatum, 
Specific, 9 


Four of Rafinesque’s Species of Sisy- 
rinchium, 42 


pecionn lanatum, 50, 51; maxi- 


Hees, 20; aphylla, 20 
Hydrolea affinis, 56; ‘quadrivalvis, 
56; uniflora, 56 


Identity of Yucca filamentosa, The, 
» pis. ’ 
Ilysanthes 20 
Indisecans Alleghenian Convallaria, 
The, 12 


June 
Fines ee at 5; pilosum, var. 
saltue 


Juncus Tondae s, 5; fuscat — 5; 
Gerardi, 5; melanocarpus, 5 ; pilo- 
sus, 4; triglu 

Jupica ; aptcm, 4 

Liliaceae, 1, 

Lily of the Valley, 12 

Live Oak, 

Luzula acuminata, 4, 5; var. caro- 


linae, 5; carolinae, 4, 5; var. 
saltuensis, : ; iebradeeot, 5; parvi- 
elanocarpa, 5; pilo- 
sa, 4; saltue ria 
Lycopus eke ar 56, 57; 
aeus, var. sessilifolius’ 56; gra 
Folia, 56 


Museadina, 49 
Muscadine, 48, 49 


Nemexia, 36; Hugeri, 36 
Nemexis elliptica, 35, 36 
Nemopanthus, 20 

Note on Smilax lanceolata, 39 


Osmorhiza, 20 

Padus agg sone 4 

Panicum Benn , 2, 3, pl. 807; im- 
var. 


onale, 3; meridionale, subvillo- 
> ; tennesseense, 3; villosis- 


Porrsoves sativa 
Peltandra, 20; hs 20; ales ea 20 


Phacelia dubia, 53-56, 816; 
dubia, 54, 55; var. 2 rgiana, 54, 
55; var. interior, 54-56, pl. 816; 
fallax, 61-53, 55, 56, pl. 814; 
gilioides, 53, 56, 815; hirsuta, 
51-54, 56, pl. 815; Plants 
as, 51; parviflora, 54, 56; pusilla, 55 


Pinus ee a, 44; palustris, 44, 55; 
— 
gens as Phacelia hirsuta, 
51, pls. 814-816 
Polemonium dubium, 54 
Polygonatum, 9, ve ’piflorum, 9-12; 
and P. canaliculatum, Specific 


ul INDEX 


Distinctions between, 9; biflorum, 
—_ oe 2; canalicula- 
Specific Sages daa 
rere GB ote orum 
and, 9; comitutatata} Q;' forma 
ce a 12; giganteum, 9; pu- 
bescens, 2 
Prunus caroliniana, 45; nemoralis, 45 


Quercus alba, 44; cinerea, 44, 45; 
faleata, 44; geminata, 44; incana, 
44, 45; lanata, hee lobata, 44; 
maritima 45; nigra, 44; rubra, 44; 
tinctoria, 44; Sniatin, 44 

Bs eee s Species of Sisyrinchium, 
Fo 


Rotala ramosior, 50, var. interior, 50 


Sanguisorba canadensis, 50 
Scupperno ng, 

ee ruticosa, 45; ligustrina, 
Silene regia, 45; scabra, 45 

Silk 


gras 
Sisyrinchium, 42; Se ss areni- 


cola, 43; Bicknellian 43; 
fibrosa, 43; cere rr} : 
florid , 42, 48; Four of - 


m, 42, 43; penta 42, 43 
Siniles, 33, 34, 36, 40, as 
6 8d 


Cc 
811, hastata, 37, subsp. 
had-ecfolia, 36, Si; var. hederae- 
folia, 36-38; Bryoniae nigrae 
age 38; § Caule aculeato, tereti, 


38, caule aculeato tereti, ca 

pinoso ter : celastroides, 

34; Chi ; cinnamonifolia, 42; 
a2, orf 


hispida, , 36; inermis, 
35; laevis Lauri folio, 41; lanceo- 
lata, 32, 37, 39-42; Note on, 39; 
laurifolia, 40, 41; ovata, 41; pan- 
durata, 39; edunculata, 35; 
Pseudo-China, 32-35, 38, 40, pl. 
12; pulverulenta, 35, rotundi- 


, 39; tamnoides, 33, 


oa 39, var. hispida, 39; 
Walte , o3, 40 
Solanum erosion: flore, ete. 14, 16, 
811 


— Distinctions between Poly- 
pe tum biflorum and P. canalicu- 
m, 9 


a 
a, 50 
Stilingi ee 45, "46: ligustrina, 
5, 46 
A si 46 


Trillium, 9, 18-20; — 17; 
Catesba aei, 14-16, pl. ; Cates- 
baei and T. ae Bll, 14; pls. 

1 5, var. 


ac ee 17-19) maculatum, 18 
; nhervosu 14- pl, ; 
Trillium Catetbasi and, "14, 1. 810; 


15, 20; stylo- 
sum, 15, 16; subg. Anthopium, 17; 
subg. veg 17, 18; Under- 
woodii, 18— 


Vitis a 48, 49; rotundi- 
49 


folia 
White Oak, 44 


Xyris albiflora, 4; Baldwiniana, 4; 
cylindrica, 4; fimbri ata, a gir 
osa, 4; flexuosa, 4; jun i 
pallescens, 4; ua. 4; sotiaiay 2 
torta, 4; ar ens 4 


Yucea angustifolia, 8; concava, 5-8; 
filamentosa, 5-9, pk 808; The 
Identity of, 5, pea 8, 809; forma 
genuina, 5, 6, vibto rma angusti- 
folia, 5, 6, 8, Subforma latifolia, 5, 

var. concava, 6 . concava, 
6; flaccida, 9; ore alba: etc., 8; 
Smalli iana, 8, 9, pl. 809 


Reprinted from Ruopora, Vol. 46, April—July, 1944 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


CLI oa 


RHYNCHOSPORA, SECTION EURHYNCHOSPORA, 
IN CANADA, THE UNITED STATES AND THE 
3 T INDIES 


SHIRLEY GALE 


i 


Dates oF Issue 


Reprinted from Ruopora, Vol. 46, April—July, 1944 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


__ RHYNCHOSPORA, SECTION EURHYNCHOSPORA, 
IN CANADA, THE UNITED STATES AND THE 
WEST INDIES 


' 
‘ 


SHIRLEY GALE 


: 
Bs: 
a 
a 
a 
Ss 
. 
a 
a 
Bey: 
a 


Dates or IssuE 


CONTRIBUTION FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY—NO. CLI 


RHYNCHOSPORA, SECTION EURHYNCHOSPORA, IN 
CANADA, THE UNITED STATES AND THE 
WEST INDIES 


SHIRLEY GALE! 
(Plates 818-835) 


In 1806 Vahl? segregated the genus Rhynchospora from 
Schoenus as the latter had been defined in the inclusive sense of 
Linnaeus. Vahl’s brief diagnosis, with its emphasis upon the 
indurated, persistent style, was followed by descriptions of 
nineteen species. Of these original species nine are now recog- 
nized members of the Section Eurhynchospora. R. inexpansa, 
R. fascicularis, R. distans (R. fascicularis var. distans (Michx.) 
Chapm.), R. capitellata, R. sparsa (R. miliacea (Lam.) Gray) and 
R. ciliata (R. ciliaris (Michx.) Mohr) had been previously 
described under Schoenus by Michaux: R. glomerata and R. alba 
under Schoenus by Linnaeus; and R. glauca was only a new 
name given by Vahl to his own previously described Schoenus 
rugosus.> R. fusca (L.) Ait. f. was also included by Vahl as R. 
alba & fusca. Of these species R. alba has been selected to typify 
the genus.® 


' Now Mrs, Chester E. Cres. 


* Vahl, Eclog. Am. ii. 5 (1798). 
*M. L. Green, Standard-Species Nom. Conserv. no. 492 (1926), mimeographed. 


90 Rhodora [APRIL 


In 1816 Elliott published the first pages of his Sketch of the 
Botany of South Carolina and Georgia. Here he described three 
new species of Rhynchospora, R. plumosa, R. punctata, R. caduca, 
and, under Scirpus, a fourth, R. schoenoides (Ell.) Wood. He 
also made the new combination, R. rariflora, from Michaux’s 
Schoenus rariflorus. 

Unfortunately nothing but confusion has resulted from 
Muhlenberg’s treatment of the genus. He attempted to rein- 
state Schoenus as inclusive of Cladium and Rhynchospora. 
Although several new species were listed in the first edition of 
his Catalogue! in 1813, they were not validated until the publi- 
cation of his Descriptio Uberior Graminum? four years later. 
The names of his new species S. ciliaris, S. capitatus, S. cymosus 
and S. setaceus were later homonyms under Schoenus, nor are 
they available for use under Rhynchospora. Finally, he frequent- 
ly neglected to give any indication of the authorship of his 
species. 

Asa Gray’s’® early monograph of North American Rhyncho- 
spora is the first treatment of the genus to be done in the care- 
fully documented, modern style; and it forms the basis for all 
later work on the genus as it is represented in Canada and the 
United States. Gray was able to verify, by means of fragments 
from the Michaux Herbarium, all of the Michaux species. He 
had the use of John Torrey’s herbarium which contained among 
others, specimens collected by Elliott, Schweinitz, Ingalls and 
Curtis, as well as the herbarium of Baldwin. At the Philadelphia 
Academy of Natural Sciences he saw the Schweinitz herbarium 
and the Muhlenberg herbarium. Of the latter he remarks, 
“Specimens of many of these [Muhlenberg’s species], however, 
do not exist in his herbarium; and those which have a place are 
in such a state of confusion, (there being often three or four 
species with a single label) that little information is to be obtained 
by consulting it.”” Gray wisely chose to base his treatment on 
the details of the achene; and, to aid in correct identification, a 
plate figuring the achene of each species was prepared. The 
descriptions are original, detailed, and accurate, and are accom- 
panied by synonymy, the citation of specimens and careful dis- 


1Thid. 5 (1813). 
2 Ibid. 4 (1817). 4 
Ann. Lyc. N, Y. iii. 191-220 (1835). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 91 


cussions. The exigencies of the primary division into “nuts 
rugose”’ and ‘‘nuts not rugose” forced the separation of R. 
plumosa from the closely related R. oligantha; but, in good part, 
the related species were placed together. There was, however, 
no attempt to separate the thirty recognized species into formal 
groups.' R. Torreyana, R. microcarpa Baldw. ex Gray, R. 
Elliottii (renamed R. Grayii by Kunth), R. megalocarpa, R. 
Baldwini, R. oligantha, R. gracilenta, and R. cephalaniha were 
the additions to the Section Eurhynchospora. The correct com- 
bination R. miliacea was made from Lamarck’s Schoenus mili- 
aceus, and given preference over R. sparsa (Michx.) Vahl. How- 
ever, to Scirpus schoenoides of Elliott, Gray gave a new name, 
Rk. multiflora, rather than the correct combination which was 
later made by Wood. R. patula Gray was a nomen confusum 
since it was applied to sheets of R. microcarpa and R. caduca. 
Rk. dodecandra Baldw. ex Gray and R. pycnocarpa were synonyms 
of R. megalocarpa, as was R. semiplumosa of R. plumosa Ell. 
R. paniculata is true R. glomerata (L.) Vahl and the species to 
which the latter name was applied by Gray is R. capitellata 
(Michx.) Vahl. 

A year later in Torrey’s North American Cyperaceae*® Gray 
reworked his treatment of Rhynchospora, segregating R. oligan- 
tha, R. plumosa and its synonym, R. semiplumosa, as 1, ‘‘Erio- 
chaete”’ separated by the subglobose achene and plumose bristles; 
and 2, “True Rhynchosporae”’, including species with lenticular 
or compressed achenes. R. macrostachya and R. corniculata were 
transferred to the genus Ceratoschoenus. One new species, R. 
filifolia Gray, was described, and R. pycnocarpa was reduced to 
synonymy under R. megalocarpa. 

In the year previous, Nees von Esenbeck‘ published his 
Synopsis Generum Cyperacearum. This was a much needed 
attempt to give the diagnostic characters of the Cyperaceae, and, 
in some cases, of the principal subgeneric divisions. He limited 
the genus Rhynchospora to species with ‘“spiculae polygamae. 
Stylus bifidus. Perigynium setosum, setis rigidis denticulatis. 


' With the exception of a suggestion that R. corniculata and R, macrostachya might 
be referred eph us Nees. 

? Class-Book of Bot. 744 (1861). 

* Ann. Lyc. N. Y. iii. 362-372 (1836). 

* Linnaea ix. 282 (1835). 


92 Rhodora [APRIL 


Caryopsis styli basi persistente discretaque latirostrata.” 
Although this definition is practically identical with that now 
applied to the Section Eurhynchospora, it was followed by a list 
of species many of which belonged to other sections. 

Kunth’s Enumeratio, Volume ii, appeared. in 1837. The 
forty-four species of Rhynchospora were divided into the Capi- 
tatae, Longirostres and Communes. Of these only the Communes 
are considered in this paper; but it is important to notice that, 
as in Rhynchospora sensu Nees, the group Communes had the 
“Stylus bifidus’” and was, in part, a precursor of the Subgenus 
Diplostylis. Kunth’s treatment of the species consisted of a 
careful review rather than the addition of new species. His 
descriptions, like those of Gray, drew particular attention to the 
achenes. His special contribution is his elaborate synonymy, 
to which I am indebted for the clue to the identity of Schoenus 
cymosus Willd. In the Addenda and Corrigenda, he gave to R. 
Elliottii Gray its present name, R. Grayii, on account of the 
earlier R. Elliottii of Dietrich. 

The treatment in Steudel’s Cyperaceae,! despite its inclusion 
of several new species, was less original than that of Kunth. 
Steudel’s main divisions were geographical. Under “I. Species 
Americae septentrionalis’” he placed Gray’s Eriochaete and. 
Rhynchosporae verae (latinizing the latter). Under ‘II. Species 
Americae australis” he had a vague key, emphasizing, in the 
primary divisions, plumose as contrasted with scabrous bristles; 
and, in the secondary divisions, the type of inflorescence. Many 
of Steudel’s descriptions were condensed latinizations of Gray’s 
descriptions, both from the Monograph and from the North 
American Cyperaceae. He even included a full description of 
R. pyenocarpa Gray, although he explained in a note that it had 
been reduced by its author to a synonym of R. megalocarpa. 
Of Steudel’s new species, R. longiseta, R. microseta and R. foliata 
are unknown to me. R. longiseta and R. microseta are referred 
by the Index Kewensis to the synonymy of R. caduca Elliott and 
R. ciliata Vahl (R. ciliaris (Michx.) Mohr) respectively. FR. ? 
monostachya Steud. is Eleocharis tuberculosa? and R. etuberculata 
is referred again by the Index Kewensis to Scirpus leptolepis. 


1 Ibid. 139 (1855). 
venson, RHopora xxxix. 248 (1937). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 93 


Steudel also gave to R. pallida Curtis another name, R. Curtisii, 
thinking that R. pallida Nees, which had not previously been 
validly published, took precedence. 

Grisebach! in 1857 set up the Section EHurhynchospora to con- 
tain species with ‘‘Setae elongatae. Stylus bifidus”, and he 
typified it by R. glauca Vahl (R. rugosa (Vahl) Gale). This 
definition was later amplified in Part vi. of the Flora of the 
British West Indies? with the additional character “achaenium 
separated from the beak by a broad transverse joint.” Of the 
species listed by him, I am excluding R. cephalotes Vahl from the 
section on the basis of its indurated scales and coarse, peculiar 
habit. R. gracilis Vahl is of uncertain identity, but Grisebach 
was probably using it in the sense of R. globularis (Chapm.) 
Small, var. recognita Gale and R. rugosa (formerly R. glauca 
Vahl). 

Grisebach’s chief contribution to the knowledge of this genus 
came with his work on the Wright collections from Cuba. R. 
pruinosa was described in Part II. of the Plantae Wrightianae’, 
but the bulk of the new species were published in the Catalogus 
Plantarum Cubensium.* Here were added to the Series Cernuae, 
previously represented only by R. pruinosa, 4 additional species 
hk. Lindeniana, R. scabrata, R. tenuifolia and R. cernua, although 
Grisebach mistakenly placed the last named species in a new 
section, Microchaeta. R. odorata Wright ex Griseb. was also 
included. R. penniseta, however, is R. plumosa Gray; R. deflexa 
is R. cubensis Rich., and R. setacea sensu Grisebach, non Vahl, 
is R. rariflora (Michx.) Ell. R. cephalotoides, also included by 
Grisebach in the Section Eurhynchospora, is closely related to R. 
cephalotes. I have excluded both of these species from this 
section. Known from the mainland, but new to Cuba, was R. 
miliacea. 

The work begun by Grisebach was continued by Wright, 
working in collaboration with Francesco Sauvalle. The portion 
of the Flora Cubana' dealing with species of Eurhynchospora was 
published for the first time in 1871. Wright’s new species 2. 


1 Goetting. iad vii. 272 (p. 124 of repr.) (1857). 
64). 


5 Anal. Acad. Ci. Habane. viii. 83 (1871). 


94 Rhodora [APRIL 


leptorhyncha and R. brachychaeta were carefully described; 
others, already treated by Grisebach or earlier authors, were 
merely listed with the number of the Wright collection. The 
authentic specimens are at the Gray Herbarium, as are also 
duplicates of Grisebach’s species, but special care has been 
necessary in their citation, for the sheets are badly scrambled; 
the same number has often been applied to specimens belonging 
to 2 or more species and the accompanying memoranda as to 
locality and habitat often cannot be oriented with certainty to 
one of possibly three specimens on a sheet. 

In 1873 Otto Boeckeler published, under the title Die Cyper- 
aceen des Kéniglichen Herbarium zu Berlin, a treatment of the 
Tribe Rhynchosporeae. He described in all 136 species of Rhyn- 
chospora, several of which he derived from Dichromena as 
transfers. These were divided into two groups: A. Capzitatae, B. 
Corymbosae. The Eurhynchosporae appeared with species of other 
sections under the latter heading. In this treatment most of 
the new species were Brazilian, but Boeckeler’s descriptions of 
such North American species as R. alba, R. caduca, R. plumosa, 
etc. were original and among the best that have been written. 

The designation Dichostyleae appeared in the summary of the 
genus by Bentham and Hooker.! It set off from the Haplo- 
styleae, with apically bilobed styles, those species in which the 
styles were deeply cleft, forming slender stigmatic branches 
equal in length to the undivided portion of the style. 

Prior to 1892 Gray’s original treatment of species of the 
Coastal Plain had been augmented by scattered publications of 
other authors, notably Curtis, Carey, Boott, and more especially 
by Chapman’s Flora of the Southern States. In that year, 
however, Britton? published a list of North American Rhyncho- 
spora. This list covered the Mexican species, but excluded those 
from the West Indies. It gave little information outside of the 
few characters employed in a rudimentary key to the groups of 
species. Its principal emphasis was placed on the synonymy, 
which was frequently incorrect, and statements as to the range 
of the species. Britton had been supplied by Clarke with an 
abstract of the latter’s arrangement of the North American 

1 he Pl. iii. 1060 (1883 

ns. N. Y. Acad ee xi, 83-93-repr. 10-19 (1892). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 95 


species, and the Hurhynchosporae, as well as some species of other 
sections, were placed under the “Subgenus Eurhynchospora 
Clarke.” R. alba var. macra Clarke, R. glomerata var. discutiens 
Clarke, and R. fuscoides Clarke were new; and were apparently 
taken from the abstract referred to in the introductory note. 
R. fuscoides was here without description but was validly pub- 
lished two years later in Clarke’s treatment of the Cyperaceae 
for Urban. R. glomerata var. minor (R. capitellata (Michx.) 
Vahl) and R. axillaris var. microcephala (R. microcephala Britt. 
ex Small) were additions by Britton himself. R. compressa 
Chapm. was reduced to a variety under R. cymosa (R. globularis 
var. recognita Gale). 

. B. Clarke wrote the treatment of the Cyperaceae for Urban’s 
Symbolae Antillanae, Volume II. With the fifty-five species of 
Rhynchospora appears the first key to species. Presumably it is, 
in part, a natural key, since series and sections are directly 
assigned to its main divisions. Its chief faults are those of 
brevity and too great a reliance on measurements. There is no - 
mention of a subgenus, or even of a section Eurhynchospora. 

Species of that section, as it is here interpreted, occur under 
the heading of Series B. Diplostyleae, and are classified as Section 
2, Plumosae, Section 3, Albae, Section 4, Fuscae, and Section 5, 
Glaucae. Section Plumosae includes, in addition to R. plumosa 
Gray, the totally different R. lwnata which has a curious horned 
tubercle. Section Albae, as defined, is synonymous with Series 
Glomeratae Small. It is represented only by R. alba from the 
mountains of Puerto Rico. Section Fuscae includes R. fuscoides 
(which is here supplied with a description), R. leptorhyncha, R. 
fascicularis, R. distans sensu Clarke (also R. fascicularis), R. 
gracilenta sensu Clarke (R. leptorhyncha), R. Lindeniana, and R. 
pallida sensu Clarke (R. brachychaeta). This section is defined 
as having setae antrorsely scabrous, nut smooth, not transversely 
wrinkled. Clarke must have overlooked the slightly rugulose 
surface in R. Lindeniana. R. brachychaeta (wrongly identified 
by Clarke as R. pallida) I have put with true R. pallida in the 
Series Chapmaniae. Section Glaucae is merely a name for a 
jumble of species having achenes with rugulose surfaces. Clarke 
placed a strong, and I think mistaken, emphasis on the impor- 
tance of the original spelling of Rhynchospora as “‘Rynchospora.” 


96 Rhodora |APRIL 


As a result of this he took up the first specific name to appear in 
combination with the original ‘‘Rynchospora.” If a specific 
name had always appeared with the spelling, ‘thynchospora,” 
he made a new combination under the old spelling. This ex- 
plains his recognition of R. sparsa Vahl in preference to the 
earlier combination R. miliacea (Lam.) Gray. It also explains 
why Vahl, Britton and Clarke are the only authorities cited 
after names of species long in good standing. 

After the turn of the century, important treatments were 
included in the following: Small’s Flora, 1903; the revision of 
Gray’s Manual by Robinson & Fernald, 1908; and, later, Small’s 
Manual, 1933. Britton also published Rhynchospora of Cuba,’ 
another list which included new species and their descriptions. 


SpEciFic CriTeR1A.—The species of Rhynchospora which come 
within the scope of this paper are perennials. The roors are of 
little aid in classification. They are usually fibrous and richly 
- branched or occasionally thickened and spongy with only a few 
short branchlets. The BAsEs are more often distinctive. In R. 
pallida they are usually cormous and covered with short, broad, 
imbricated scales—a condition occurring to a lesser degree in R. 
megalocarpa. In the Series Cernuae a short, densely fibrous 
caudex is developed, the fibres resulting from the fraying-out of 
the fibro-vascular strands in the lower portions of the old dried 
basal leaves. This character is one of several which are probably 
correlated with the intermittently exsiccated habitats of these 
species. The bases of R. cubensis and R. stenophylloidea are 
subligneous. A few species have stolons. Those of R. fusca 
are slender, whereas short, thick stolons bud out from the bases 
of R. pallida. R. megalocarpa, which populates the sand-hills of 
the southern Coastal Plain, also spreads by means of stolons. 
The commonest growth-habit, however, is a tuft; and several 
species, R. inexpansa among them, are reported as forming dense 
stools. An exception to the caespitose habit, as its name con- 
notes, is R. solitaria. _ 

The BASAL LEAVES are usually linear-elongate, from 1 to 7 mm. 
in width, or filiform. Commonly they are flat or somewhat 
canaliculate. The under surface is short-carinate, the keel be- 

1 Mem. Soc. Cubana Hist. Nat. ii. 185 (1916). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 97 


coming more prominent toward the apex so that the tip of the 
leaf is sharply triquetrous. This character is present, but less 
obvious, in a few species with broad, short leaves, such as R. 
ciliaris and R. pruinosa, in which the leaf is abruptly narrowed 

to an obtuse tip. Rarely the leaf-tips are rounded, as in R. 
plumosa. The angles of the tip, if not the entire margin of the 

q leaf, are always more or less serrated. These microscopic teeth, 
__ apparently formed of pure silica, contribute in great part to the 
material worthlessness of the plants, for they render the leaves 
unfit for cattle-fodder. In R. ciliaris these serrations reach 
their maximum development as stiff, silvery cilia. Occasionally 
the leaves are involute, although this condition is difficult to 
determine from dried specimens. In R. pruinosa the leaves are 
not only canaliculate but also moderately revolute. However, 
the chief distinction of the leaves of this species lies in their 
peculiar upper surfaces. These are roughened and bear small = 

white inclusions of lime which are responsible for the rimy or 4 

silvery appearance emphasized by the specific name. In another 

closely related species, R. scabrata, the upper surfaces of the 
leaves are definitely exasperate. The basal leaves vary from 
stiffly erect to flat and spreading to curling. The curling habit 
reaches its climax in the circinately coiled leaves of the tiny, 
depressed R. crispa. The CAULINE LEAVES are similar, in most 
cases, to those of the base. They decrease in size upward and are 
finally reduced to the setaceous or, less often, somewhat frondose 
bracts of the inflorescence. The typically closed sheaths are 
- definitive of the Cyperaceae as a whole. 

The cuLMs are usually triquetrous and slender to robust, or 
filiform and subterete. The 2-several nodes each bear a leaf | 
which may or may not subtend a lateral cluster of spikelets. is 
The internodal surfaces are smooth or striate, and the culms - 
range from stiffly erect to procumbent. ve 

The INFLORESCENCE varies in size from a ‘solitary terminal = 
fascicle, as in R. pallida, to a terminal decompound fasciculate | 
cyme which is accompanied by 4-8 smaller lateral cymes, as in 
R. miliacea. Throughout this paper I have applied the term 
“eyme”’ to the compound, flat-topped or concave clusters of 
spikelets which characterize in particular the Series Caducae. 
Compact bundles of mainly ascending spikelets are ‘fascicles’, 


98 Rhodora [APRIL 


whether they are numerous and approximate in the cyme or are 
solitary and distant in the axils of the cauline leaves. Fascicles 
may be globose, turbinate or corymbiform, depending upon 
their shape. Compact masses of spikelets oriented in all 
directions are heads, capituli or glomerules and may also be of 
primary or secondary degree. Unfortunately I have not been 
able to study flowering material of the genus, for from the dried 
specimens it is difficult to ascertain whether the so-called cymes 
are determinate, but to all appearances the main axis and the 
successive axes in turn are terminated by a spikelet. 

The sPIKELETS are usually ovoid, but they may be subglobose, 
as in R. globularis, or fusiform as in R. capillacea. The lower 2 or 
3 of the spirally imbricated scales are comparatively small and 
empty. The succeeding 1-10 subtend perfect florets, each of 
which may produce an achene. The terminal 1-2 scales subtend 
rudimentary florets or are empty. Occasionally the spikelet 
contains only a single floret. 

The ScaLes are papery (not indurated), tightly, but more 
often loosely, imbricate, and persistent or caducous. They are 
commonly brownish and fairly constant in shade, most commonly 
castaneous or fuscous. In R. alba and several members of the 
Series Cernuae they are whitish; in R. pallida they have a reddish 
tinge. Those of R. leptorhyncha, on the other hand, are black- 
ened, as are frequently those of R. cephalantha var. typica. The 
general outline of the fertile scales is ovate, on the one hand, to 
narrowly lanceolate, and on the other to suborbicular. When 
tightly imbricate they are concave. The midrib frequently 
extends as a mucro which is often serrulate, and in R. ciliaris 
bears straggly reddish-brown cilia. 

he STAMENS are variable in number, from 1-12, but are 
usually 3. In R. odorata the filaments are conspicuously mar- 
cescent. 

The hypogynous BRISTLES are presumably the remnants of 
a perianth. They are borne at the base of the achene and are 
arranged in upper and lower series consisting of 3 bristles each. 
In R. alba, however, the bristles are from 10-12 and in R. macra 
from 18-20. The increase in number is probably due to chorisis; 
occasionally I have seen an achene of R. alba with 2-pronged, 
presumably imperfectly divided, bristles. The broad, straplike 


1944] Gale,—Rhynchospora, Section Eurhynchospora 99 


bristles of the Glomeratae are unique, not only in their shape and 
size, but in their retrorse barbs. In other series the bristles are 
capillary, or flattened only at the base, and the barbs, or better 
the minute serrulations, are directed upwards. There are a few 
forms in the Series Glomeratae which, like the species of other 
series, have bristles with antrorse barbs or with barbs failing. 
This problem and its importance has been discussed under R. 
capttellata f. discutiens. The bristles in the Series Plumosae are, 
as indicated by the name of the series, characteristically plumose. 
Several species of other series, however, have a few silky hairs 
at the bases of the bristles. Occasionally the bristles fail to 
develop. This is true in particular of R. Chapmanii and R. nuda, 
and also of occasional achenes of R. perplexa. 

The sTyLe has two stigmatic branches which are characteristic 
of the genus and which indicate the bicarpellary nature of the 
gynoecium. 

The AcHENE is the most important single character in the 
determination of a specimen, for, almost without exception, that 
of every species is distinct. It is derived from the development 
of one of the paired ovules of the gynoecium at the expense of 
the other. The shape is commonly ovate, lenticular and bicon- 
vex; usually the achene is marginate. Those of species belonging 
to the Series Glomeratae are frequently slenderly prolonged 
toward the base, forming a gynophore. Other peculiarities of 
the achenes of this series are the heavy wire-like margins and the 
prominent central umbo or boss. The umbonate condition also 
appears in achenes of other series, e. g. Rk. Harperi of the Fuscae. 
More often the central area of the achene is picked out by a 
smooth, pale disc. This is particularly true of R. filifolia the 
white disp of which has a sheen like that of glass. The achenes of 
R. compressa suggested the specific name, for they are so flattened 
that they have a shrunken appearance as if they had dried while 
immature. Some other species with strongly compressed achenes 
are R. perplexa, R. schoenoides, and R. tenuifolia. Very few of the 
series have achenes with an absolutely plain smooth surface. 
Usually the surface is alveolate or cancellate. The alveoli may, 
as in the case of the Chapmaniae, be reduced to tiny pricks; or 
they may be shallow, superficial and isodiametric as in the 
Harveyae. ‘The most common modification of the alveoli is that 


100 Rhodora [APRIL 


in which they are more or less aligned across the achene with 
the transverse walls pulled up into ridges. This produces the 
rugulose to ridged effect which is common in the Series Rari- 
florae, Cernuae, Cubenses, Harveyae, Globulares, Caducae, and 
Glaucae. Although the surface is ridged the individual alveoli 
may remain distinct, as in R. caduca, or they may be crowded so 
as to appear only as fine glistening striae between the ridges, or, 
in R. tenuifolia, over the faces of broad corrugations themselves. 
Like the scales, the achenes are brownish, but they vary in 
degree from the pale, lustreless shade of R. nuda to the dark, 
gleaming black-mahogany of R. megalocarpa. The measurements 
of the achenes are very dependable, and have been made with 
the aid of a finely divided steel rule, to tenths of a millimeter. 
The largest achenes in this section of the genus are those of R. 
megalocarpa (2.8-3.4 mm. wide, 3-4 mm. long). Among the 
smallest. are those of R. Knieskernii, R. sulcata, and several 
species in the Cernuae the measurements of which do not exceed 
a millimeter in width and length. 

Occasional trigonous achenes have — seen in the Series 
Plumosae and Glomeratae. Such anomalies are accompanied by 
tripartite styles. They probably represent the last traces of a 
tricarpellary condition. 

The TUBERCLE, consisting of the indurated and persistent base 
of the style, is triangular and compressed. The achenes of the 
Harveyae which are, for the most part, tumid above, have conical 
tubercles. These are buttressed and slightly incrusted at the 
base by the narrowed summit of the achene. The tubercles in 
the Series Chapmaniae are extremely short, often apiculate. 
The tubercle of R. compressa is strongly depressed with a pro- 
jecting basal flange. The margins of the tubercles in the Series 
Fuscae are thickly setose, and those of other series may be less so, 
or bare. 


ACKNOWLEDGMENTS.—This study has been made under the 
guidance of Professor M. L. Fernald, and for his advice and 
patience I am most sincerely grateful. I also wish to express My 
appreciation to the many friends who have so freely given both 
time and encouragement—in particular to Mr. Charles A. 
Weatherby and Miss Bernice Schubert of the Gray Herbarium, 


ata 


=a 
= 
- 
ong 
= 
ia 
£ 
a 
be 


PPE CT te Ae 
WePE Rn Teme eee R 


1944] Gale,—Rhynchospora, Section Eurhynchospora 101 


and to Dr. Chester E. Cross. I have been generously given the 
opportunity to study material from the herbaria of the following 
institutions to the curators of which I express my great obligation: 
Academy of Natural Sciences, Philadelphia (P); California 
Academy of Sciences (CA); Catholic University of America (CU); 
Duke University (D); Gray Herbarium (G); Atkins Institution 
of the Arnold Arboretum (A); Louisiana State University (La); 


Missouri Botanical Garden (Mo); New England Botanical Club 


(NE); New York Botanical Garden (NY); St. Bernard College 
(StB); United States National Arboretum (USNA); United 


States National Museum (US); University of North Carolina 


(NC); University of Pennsylvania (Penn). 

Ruy A Vahl. Seales spirally meine the lower 1 
(rarely )—2 to ait scales vacant, the following 1— 10 subtending 
perfect florets, the upper 1-2 flo rets DE he or rudimentary: 
stamens 1-— 12, usually 1-3: hypogynous bristles 0-20, when 
present usually 6: style bilobed at the apex or with 2 long, 
slender, stigmatic branches: achene more or less oo 
crowned with a conspicuous tubercle consisting of the broad, 
persistent, indurated base, or even the greater part, of the stale. — 

hiefly perennials with more or less triangular culms and axillary 
inflorescences. Species of tropical and subtropical regions of both 
hemispheres, temperate North America and Eurasia. (Name 
taken from elyyeos, a snout and oxopd, a Ree from the beaked 


_ achene.)—Enum. ii. 229 (1806); Gray, A Lye : Na Y. a 194 


(1835); Kunth, Enum. ii. 287 (1837) ; Steud. Orn 139 (1855); 


- Bentham & Hooker, Gen. PI. iii. 1058 (1883); Pax in Engler & 


bees Pfizfam. ii. 2: 116 (1887); Clarke in Urban, Symb. Ant. 

. 103 (1900); Pfeiffer in Fedde, Rep. Spec. Nov. xxxviii. 89 
(i 1935). tose L. C. Richard in Persoon, Synops. i. 60 (1805) 
in a note; nomen rejiciendum. Phaeocephalum Ehrh. Beitr. iv. 


; 146 (1789), nomen illegitimum 


Since this paper is limited to a consideration of the Section 
Eurhynchospora the generic definition given above is neither 


_ Original nor complete; nor has any attempt been made to include 


in the list of generic synonyms the numerous tropical genera of 
Nees, none of which come within the scope of this paper. — 
In 1935 the name Rhynchospora! was conserved over Triodon 


__L.C. Richard, which threw into synonymy all previous com- 


binations made by Farw in ee Triodon. House’ in 1920 at- 
‘Int. Rules Bot. Nomencl. 90 (19: 

* Rep. Mich, Acad. Sci. xv. 167 Shae § xix. 253 (1917). 

‘Am. Midland Nat. vi, 201 (1920). 


102 Rbodora [APRIL 


tempted to set up the generic name Phaeocephalum Ehrh., but 
the name is excluded by the International Rules, under Art. 67 
(3), as a unitary designation of species, not intended as a generic 
name. 

However, at present the status of the generic name is once 
again challenged, for Pfeiffer! has brought up the problem of the 
inclusion of Pleurostachys Brongn.? and Dichromena Michx.’ 
within the genus. Whether Pfeiffer’s solution, which reduces 
both genera to the status of synonyms under Rhynchospora, is 
sound, I do not know. It is a problem for the student of the 
Rhynchosporeae as a whole. However, the name Dichromena has 
priority over Rhynchospora, and the inclusion of the two under 
one generic name would necessitate either the transfer of the 
species of Rhynchospora to Dichromena, following the precedent 
of Macbride*, or the conservation of the name Rhynchospora 
over Dichromena as proposed by Pfeiffer’, with the necessary 
combinations under Rhynchospora, many of which have already 
been made by that author. 

Subgenus Disty iis Pax. Style deeply cleft; stigmatic branches 
linear, equal in length to the undivided portion of the style.— 
In Engler & Prantl, Pflafam. ii. 2: 117 (1887). Rhynchospora 3. 
Communes Kunth, Enum. ii. 295 (1837). Rhynchospora ii. 
Dichostyleae Besthann & Hooker, Gen. Pl. iii. 1060 (1883). 
Rhynchospora Series B. Di plostyleae Clarke in Urban, Symb. 
Ant. ii. 104 (1900). ree Diplostylis Pfeiffer in Fedde, Rep. 
Spec. Noy. _XXXvVili. 91 (1935). 

Section Eurny A Griseb. Seales thin and papery 
not indurate, often ‘oer ts imbricate: bristles 0 (rarely)-20, 
usually 6 and equal to or exceeding half the achene in height, if 
bristles consistently failing achenes smooth (e. g. R. Chapman, 
R. nuda): achenes commonly ovate in pablus, ienticdlar and 
gradually biconvex or umbonate, less often subglobose; surface 
smooth to strongly alveolate to striate and nde ed: tubercle 
discrete, triangular or conical, compressed. —Goetting. Abh. Vil. 
272 (1857); Bentham & Hooker, Gen. Pl. iii. 1060 (1883); 
Pfeiffer in Fedde, nei Spec. Nov. xxxviii. 91 (1935). Rhyncho- 
spora Series B. Diplost yleae, Divisio 5. Eu-Rhynchospora. Clarke 
in Kew Bull. Add. Ser. viii. 119 (1908). Rhynchospora ii. ee 

enus pa Hea ag Clarke ex Britton, Trans. N. Y. Acad. 
Sci. xi. 85 (1892). 

' Fedde, Rep. Spec. Noy. xxxviii. 88 (1935); xliii. 258 (1938). 

? Brongniar ileglen ares Voy. Coq. Bot. 172 t. 31 (1829). 

2 Fl. Bor.-Am. i. 3 

‘ Field Mus. Pub. be iv. 166 (1929); viii. 113 (1930); xi. 5 (1981); xiii. 301 (1936). 

’ Fedde, Rep. Spec. Nov. xliii. 261 (1938). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 103 


Key To SERIES IN SECTION EURHYNCHOSPORA 
a. Bristles retrorsely barbed, or if barbs antrorse or failing, 
ristles robust, straplike, pale, exceeding the ache 
eries Glomeratae. 
a. Bristles payer barbed or failing, rarely smooth, then capil- 


ge 


ary 
b. Bristles welt developed and popes orp for at least 14 


their length or reduced to six basal tufts... ... 2. age Plumosae. 
b. ivintlad upwardly arte ge Swit or without a few basal 
hairs, rarely smooth... . 


c. Surface of Aerts Siecle or pricked by small dark pits 
achene chestnut-brown or dark hove (then lentiading 
and moderately biconvex), not ne 

d, Surface of achene pricked by ree "dak pits 0 = if sur- 


face smooth, bristles 1-3 and ailing; 
lateral fascicles rare, with exception of 13 in 
bracketed 6.5 2 Fee ho he Series Chapmaniae. 


d. joan = Recarag =e ti bristles 6 and well devel- 
in R. cularis var. typica and R. 
debilis vie they are fe 5 to 6 and rudimentary ; 
sities pee present in well developed speci- 
é. nate aie usually less than 1 mm. wide, light 
brown, py yriform or nen ce ellipsoid with the | 
‘maximum width near the apex............. Series Fuscae. 
e. Achenes vieodiid 1 mm. in width (with the excep- 
tion of that of R. Fernaldii which is minute and 
blackish), broadly ovate in outline, their maximum 
idth corresponding to their midpoint. vil eee) Fasciculares. 
ans 


ee es ve Pi pa nee te spiciform 


und cory 
with ‘yelatyely, ‘ie “often _straw-coloed spikelets; 
culms relatively short to depressed. .. . 

g. Spikelets remote on phe ete pedicels; is corym- 
biform, spreading; 2 species only, both of the 
Coastal Plain, R. rariflora also in the a, Indies 

Series Rariflorae. 

q. oe approximate, if Vian peiclae one 
t di es limite es ; 
Indien os eae 7" Series Ceriuac. 

erulose, corymbiform, congeste oose cyme; ; 
culms relatively t tall re ly dep oad. ie 

h. Spikelets with 1 flor e solitary 

sp its axis; ase longitu 
if shriveled; 2 species of the Greater yates 


a 2 
PEs 
5.8 | 
15 ® 
dB 

4 


Series Cubenses. 
h. age wok airy 2-several florets, one of which may be 
rudiment; achene not longitudinally 
eB ir rg many species of the Coastal Plain 
the West Indies... .i. 
i. Achene surmounted by a grayish, conical-apiculate 
tubercle with a subterete base which is mad 
buttressed and often slightly incrusted by the 
narrowed neck of the achene; surface of achene 


104 Rhodora [APRIL 


deeply alveolate and Tidged only i in hp hea 


otherwise mahogany-brown, smoot with a 
honey eomb-pattem of shallow eadaatie 
CM ars Pie Pe et wae ie este eb 9. Series Harveyae. 


t. Achene usually surmounted by a deltoid, com- 
essed tubercle which is often attenuate, if 
eonieal-diseoid mh te at the base not 
visibly buttre 

j. Achene broadly eberrald or slenderly ellipsoid- 
eh abie Alam castaneous or catching the light 
minent, deeply etched alveoli, 

Seaualty te ed....k. 

k, Cymes stiff, erect or spreading; periee ar 
terminating in glomeruli whic y be 
dense or limited to 3-4 globose snikelots 

10. Series Globulares. 

k. Cymes usually flexuous, corymbi form, densely 

asciculate ase for the stiff, open netlike 
ies of R. miliacea), occasionally glo- 
merulose in reduced phase of R. micro- 
rd MAR hie edie Week roe i ees eas 11. Series Caducae. 


& 


which are intercepted by transverse merce 
tions or vague rugulosities........... Series Glaucae. 


Series 1. GLoMERATAE Small, emend. Plants of acid bogs of 
temperate North America and Eurasia with several species 
limited to the Coastal Plain; R. alba also in the mountains of 

uerto Rico and (?) northeastern Brazil. Habit caespitose: 
leaves filiform to flattish and up to 5 mm. wide: culms capillary 


cymose-fasciculate: spikelets 1—2-fruited: scales acute to mu- 
cronate, tightly or loosely imbricate: achene pyriform or with a 
conspicuous gynophore, smooth to granular or faintly rugulose, 
usually with a pale central disc; bristles 6, retrorsely barbellate, 
or if with barbs antrorse or failing the bristles robust, straplike 
a pale-—Rhynchospora, V. Glomeratae Small, Man. 175 (1933), 
in part. Rhynchospora, Series B. Diplostyleae Sect. 3. Albae 
Clarke in Urban, Symb. Ant. ii. 105 (1900). 


Key To Species IN SERIES GLOMERATAE 


a. Achenes with a ogre arg pale, wirelike margin, smooth, 
castaneous and unlined, usually lustrous, umbonate, with a 
ale disc and basally pr rolonged 

b. — 1-fruited, the solitary ‘achene terminating the 
c. futovensdaee of 2-6 dense, globose ee (rarely 
subhemispherical and looser on poorly devel oped 

ra © eR the crowded spikelets ing, to re 

ff x 


1944] Gale,—Rhynchospora, Section Eurhynchospora 105 


d. Achenes minute, 0.9-1.1 mm. wide, 1.4-1.6 mm. long 

1. R. microcephala. 

d. Achenes 1.1—-1.6 mm. wide, 1.8-2.5 mm. long....2. R. cephalantha. 
c. Inflorescence of 3-7 turbinate to loosely hemispherica 
sretaticed the loosely aggregated spikelets ascending to 

WIGOING fo e6 2. os a hoe ace Ee R. chalarocephala. 

b. Spikelets usually 2- or more-fruited, or if one-fruited es 


e. Achene prominently umbonate, with a aie: aise a 

pressed sides and raised wire-like margins; inflorescence 

of usually several strict, fasciculate, rarely glomerulose, 
CVINCR i Sis ea a le a cet ee ere R. glomerata. 

e. Achene gradually eis without a prominent umbo, 

surface a uniform brown, margins narrow; inflorescence 

of 2-6 compact, seeeaalneey lobed fascicles, less often 
CyMonelAsCIMURTO 23 iho. chances eee eens 5. &. capitcllata 

a. Achenes inconspicuously marginate, finely granular to slightly 

gulose, dark brown toward the margins from which short, 

dark, broken lines run in between the roughenings toward a 

more or less definite te pale, polished disc... .f. 


q. 
g. Bristles 10-12; achenes 0.9-1.2 mm. wide, 1.6-1.8 mm 
long; spikelets several-fruited or, if 1-fruited, with 
terminal rudimentary floret; ac typically whitish 6. R. alba. 
g. Bristles 18-20; achenes 1.3-1.4 mm. wide, 2—2.1 mm. 
ong; spikelets 1-fruited, without a terminal sane 


h. — chong with a ae slander ee 
R. capillacea. 
1. R. MICROCEPHALA Britt. ex Small. Se leaves 1-3 


m 

cones globose, or less frequently looser, subhemispherical 
heads 1.1—-1.8 em. wide, on included aodnohe: spikelets slenderly 
lanceolate-attenuate, sessile, ascending to reflexed, typically 
forming a solid echinate ball, 1-fruited; the solitary fertile floret 
terminal with no trace of a sterile rudim ent: scales ovate to 
lanceolate, acute, chestnut to dark brown, tightly inrolled 
; especially ‘at the apices: bristles 6, straplike, retrorsely barbellate; 
3 their apices aren about the tubercle which they slightly 
exceed: achene 0.9-1.1 mm. wide, 1.4-1.6 mm. long; its body 
suborbicular, with a short but narrow and distinct gynophore; 
surface smooth, lustrous, brown, with a prominent light umbo 
and a raised wirelike margin: febcaile subulate-attenuate, 0.7- 
1.1 mm. nigh; its base not wholly covering the summit of the 
achene. Piate 818, rigs. 3A and 3B; Map 4.—Fl. 195, 1327 
(1903) and MG. 181 (1933); Fernald, RHODORA, Xxxvii. 404, pl. 


106 Rhodora {APRIL 


391, figs. 4 and 5 (1935); Fernald & Gale, Ruopora, xlii. 428, 
figs. 3 and 4 (1940). R. azillaris var. microcephala Britton, 
Trans. N. Y. Acad. Sci. xi. 89 (1892); Britton & Brown, IIl. FI. 
i. 279 (1896); Britton, Man. 185 (1901); Robinson & Fernald in 
Gray, Man. ed. 7: 201 (1908). R. axillaris sensu Britton in 
Britton & Brown, Ill. Fl. ed. 2: i. 344 (1913), in part; Britton, 
Mem. Soc. Cubana Hist. Nat. ii. 195 (1916), not as to name- 
bringing syn. Schoenus axillaris Lamarck.—Wet peaty or sandy 
clearings, swamps and pond-margins of the Coastal Plain from 
New Jersey southward to the Florida Peninsula, thence west to 
Mississippi; also in western Cuba. Nrw JERSEY: east of Park- 
dale (2 miles), Atlantic Co., Aug. 17, 1905, Van Pelt (P); Mon- 
mouth Co., 1845, Knieskern (NY, Type of R. axillaris var. micro- 
cephala). DeLAWARE: Indian River, Millsboro, Sussex Co., 
Sept. 21, 1907, Brown (P). Wasuineton, District oF CoLum- 
BIA: Sept. 18, 1901, Steele (NY). Maryann: Salisbury, Wico- 
mico Co., Sept. 28, 1863, Commons (G, P). Virernia: sphagnous 
magnolia swamp at head of Garnett Creek, about 1 mile north- 
east of St. Stephen’s Church, King and Queen Co., Fernald & 
Long, no. 13275 (G); fresh to brackish swales along North Land- 
ing River, near Creed’s, Princess Anne Co., Fernald, Long «& 
Fogg, no. 4830 (G, P); wet peaty clearings in woods of Pinus 
serotina, south of Grassfield, Norfolk Co., Fernald & Long, no. 


mile east of Balley, Wilson Co., Qosting, no. 1670 (CU, D); 


Hartsville, Darlington Co., July 12, 1920, Norton (NC); sandy 


drainage-ditch, 2 miles west of Salters, Williamsburg Co., 


1944] Gale,—Rhynchospora, Section Eurhynchospora 107 


Godfrey & Tryon, no. 502 (D, G, NY); Sumter Co., Holdaway, 
no. 11 (D); boggy swale, 5 milés south of Columbia, Lexington 
Co., Godfrey & Tryon, no. 1263 (G). Groreta: Cypress Pond, 
Bethesda Church, Effingham Co., Eyles, no. 6398 (CU); Chase 
Prairie, Okefenokee Swamp, Charlton Co., J. S. Harper, no. 
731 ( FLoripa: South Jacksonville, Duval Co., Aug. 11, 
1909, Lang (P); low pineland bordering Lake Geneva, near 
Keystone Heights, Clay Co., Feb. 24, 1925, O’Neill (CU); 
margin of pond along west boundary of Welaka, Putnam Co., 
June 26, 1940, Laessle (CU); in a low pineland, 7 miles north of 
Ft. Christmas, east of Orlando, Orange Co., O’ Neill, no. 7679 
(CU; US, without number); wet ditch, Lake Jovita, Pasco Co., 
O’ Neill, no. 2609 (NY); flatwoods, 1 mile from Tiger Lake, Polk 
Co., McFarlin, no. 3466 (CU); ditch, Hardee Co., July 29, 1940, 
Schallert (G); margins of ponds in pine barrens, Apalachicola, 
Franklin Co., Chapman in Biltmore Herb., no. 864b (G, in part). 
ALABAMA: in a wet place, Perdue, Coffee Co., Blanion, no. 83 
US). Mississippi: Petit Bois Island, Jackson Co., Tracy, no. 
iv . 


17240 (NY). 

2. R. CEPHALANTHA Gray. Caespitose: leaves 1-4.5 mm. 
broad, flat or tending to become inrolled on drying, serrulate on 
upper margins, erect; radical leaves equalling about 14 the culm; 
cauline leaves short: culms slender to stout, erect, obtusely 
trigonous, 0.4-1.1 m. tall: inflorescence 1-5 dm. long; glomerules 
densely subglobose to loosely hemispherical, 1-2.2 em. wide, 
terminal and solitary or accompanied by 1-4 smaller lateral 
glomerules on subincluded peduncles: bracts foliaceous and con- 
spicuous: spikelets ovoid, 4-6 mm. long, compact, sessile; the 
fertile floret consistently terminal and solitary with no trace of 
a succeeding rudimentary floret: scales castaneous to blackish, 
slightly mucronate, so tightly involute at their apices as to give 
an echinate appearance to the glomerule: bristles 6, robust, 
strap-like, with the margins and upper surfaces retrorsely 
barbed, stramineous to light brown, equalling to slightly exceed- 
ing the tubercle around which they converge: achene 1.1-1.6 
mm. broad, 1.8-2.4 mm. long; its body suborbicular, usually 
with definite shoulders, lenticular and prominently umbonate, 
with a very slender gynophore; the sides depressed and rimmed 
by a raised wire-like margin; surface smooth, lustrous, castane- 
ous, paler over the umbo: tuberele compressed, subulate-attenu- 
ate, 1.4-2.4 mm. long. . 


108 Rhodora [APRIL 


Key To THE VARIETIES OF R, CEPHALANTHA 
Achenes 1.4—1.6 mm. — 2-2.5 mm. long; habit usually stiffly 
erect; culms often robus 
Glomerules subglobose o loosely hemispherical, 1-3 (rarel 
with the lowermost poorly formed) ; culms slender to ode 
a. var. typtica. 
Glomerules eee: eHepee, 4-7 in number, the lowest 
POUAOU; CUR BLOG rs eke ew CV ee var. pl ioccph:la, 
Achenes 1.1-1.2 mm. ute 1.8 mm. long; habit weakly greet; 
culms slender, attenuated in appearance............. var. attenuata. 


Var. Typica Fernald & Gale. Leaves 1.5-2.5 (-3) mm. 
ears tending to become inrolled on drying: culms slender, 0.4—1 
m. in ‘height: inflorescence 1—2.8 dm. long (rarely longer) com- 
prising about 4% the culm; glomerules subglobose to loosely 
hemispherical, 1.3-2.2 ¢ m. wide, terminal and solitary or accom- 
panied in the upper ale by 1-2 (rarely 3) smaller glomerules, 
then only the terminal glomerule subcompound to several-lobed: 
achene 1.4-1.6 mm. broad, 2-2.4 mm. long. PuaTe 818, FIGs 
4A and 4B; Map 7. —Ruovora, xlii. 423 (1940). R. cephalantha 

Ann. Lye, Ni Yodan 218, pl. 6, fig. 30 (1835) and Man. 
533 (1848), in part var. pleiocep hala, as with later authors; 
Chapman, FI. So. U. 8. 528 (1860); Fernald, RHODORA, XXXVii. 
403, pl. 391, figs. 2 and 3 (1935). R. axillaris Britton, Bull. 
Torr. Bot. Cl. xv. 104 (1888) and Trans. N. Y. Acad. Sci. xi. 89 


name-bringing syn. . Schoenus axillaris Lam.; Britton “& Brown, 
Ill. Fl. i. 279, fig. 655 (1896); Britton, Man. 185 (1901); Small, 
Fl. 195 (1903) and Man. 181 (1933); Robinson & Fernald in Gray, 
Man. ed, 7: 201, fig. 327 (1908). Phaeocephalum axillare House, 
Am. Midland Nat. vi. 201 (1920).—Sphagnous bogs of southern 
New Jersey, southward on the Coastal Plain to Georgia, thence 
ee to eastern Louisiana. New Jersey: bog southeast along 


Parkdale, Atlantic Co., Aug. 17, 1905, Van Pelt (P); Gray ?, 
1 (NY, Types; accompanied by notes in Gray’s handwriting); ; 
al bog along Mullica River about 1.5 miles southeast 
of Atsion, Burlington Co., Fogg, no. 5662 (G). DELAWARE: 
ree along Queen Anne Railroad, near Ellendale, Sussex Co., 
Aug. 17, 1899, Commons (P). MARYLAND: open white gravel 
bog, Powder Mill Bogs, near Lewiston, Prince George Co., 
Blake, no. 10670 (CA). Vrirernta: argilla aceous and siliceous 
ggy depression southeast of Petersburg, at head of Poo Run, 
Prince George Co., Fernald & Long, no. 6090 0 (G, P); depression 
in Rebar ig woods west of Winfield’s Mill, Dinwiddie Co " 
F Long, no. 13902 (G); sphagnous argillaceous boggy 
peak just north of Wakefield, Sussex Co. , Fernald & Long, 
no. 7352 (G, P); sphagnous bog about 1 mile northwest of 
Dahlia, Greensville Co., Fernald & Long, no. 8993 (G, NY, P). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 109 


Nortu CAROLINA: sphagnous bog at Method, Wake Co., God- 
frey, no. 4985 (CA, D, G); drainage-ditch at Carolina Beach, 
New Hanover Co., Godfrey, no. 4719 (G, NC); roadside-ditch 
between Beaufort and Atlantic, Carteret Co., Blomquist, no. 
11300 (D). Groreta: cypress-pond, Bethesda Church, Effing- 
ham Co., Eyles, no. 6 U); edge of cypress-pond, near 
Smithville, Lee Co., Eyles, no. 1711 (CU). Lovistana: in low 
pine barrens, St. Tammany Parish, Sept. 14, 1892, Langlois (US). 
Forma antrorsa, f. nov. Setis antrorse hispidulis.—Occa- 
sional in range of the typical form. New Jersey: Hammonton, 
Atlantic Co., Aug. 4, 1907, Bartram (P); Parkdale, Camden Co., 
Sept. 18, 1916, Pennell, no. 9032 (NY); Quaker Bridge, Burling- 
ton Co., Sept. 3, 1867, Parker (G, in part the typical form; US); 
same locality, Aug. 18, 1866, Diffenbaugh (P); border of cranberry 
bog along Little Hauken Creek, north of Jenkins, Burlington Co., 
July 28, 1937, Long, no. 50858 (P); Symmes’ Place 3 miles bagch of 
i 1 j 


Detaware: Ellendale, Sussex Co., Aug., 
US); same locality, July 9, 1908, Van Pelt (P); sandy swamps near 
Georgetown, Aug. 26, 1897, Commons (P). Nortu Caro.ina: 
savanna 5 miles east of Jacksonville, Onslow Co., Aug. 6, 1938, 
Godfrey, no. 5808 (G, type). SourH Caro.ina: drainage ditch, 3 
miles north of McClellanville, Charleston Co., July 19, 1939, 
Godfrey & Tryon, no. 677 (G); grass-sedge bog or savanna, 12 miles 
north of Georgetown, Georgetown Co., Aug. 2, 1939, Godfrey & 
Tryon, no. 752a (D, G, NY). 

2b. Var. PLEIOCEPHALA Fernald & Gale. Leaves 2.5-4.5 mm. 


Surry Co., Fernald & Long, no. 8994 (G, TYPE; NY and ig 
isoTyPES) ; dominant in flat sphagnous pinelands, Collier's Yard, 


Co., Blomquist, no. 10872 (D, G); ditches near Wilmington, New 
Hanover Co., Biltmore Herb. no. 279a (G, NC, US); moist place 


110 Rhodora [APRIL 


in savanna, Southport, Brunswick Co., Aug. 13, 1930, Blomquist 
(D); pineland at Nakina, Columbus Co. , Godfrey, no. 6347 (G). 

oUTH CAROLINA: sandy ‘drainage ditch, "2 miles west of Salters, 
Williamsburg Co., Godfrey & Tryon, no. 504 (D, G, NY, P): 
grass-sedge bog or savanna, 12 miles north of Georgetown, 
Georgetown Co., Godfrey & Tryon, no. 1061 (G, NY, P); boggy 
ditch in pine barrens, 2 miles east of Megeetts, ‘Charleston 
Co., Wiegand & Manning, no. 582 (G). Groreta: Waycross, 
Ware Co., Aug. 18, 1909, Lang (P); pine barrens, near Lem 
Griffin’s Camp, Okefenokee, Clinch Co., Eyles, no. 133 (CU); 
cypress head, Valdosta, Lowndes Co., May 27, 1940, Sargent 
(Sargent Herb.); wet meadow, Leslie, Sumter Co., Harper, no. 
413 (G, NY, US). FLorrpa: swampy places in pine barrens near 
Jacksonville, Duval Co., Curtiss, no. 5016 me Ee 
swamp, vicinity of Eustis, Lake Co., Nash, no. as (CUl GEN, 
P, US); swamp, Okefenokee region, ’ Brevard Co., Fredholm, no. 
5821 (G); bogs and shady swamps, Apalachicola, Franklin 'Co., 
Chapman in Biltmore Herb., no. 279b (G, NY). ALABAMA: 
Miflin Creek, vicinity of Elberta, Baldwin Co., Aug. 21, 1925, 
Wolf (StB); in a wet place, Perdue, Coffee Co., Blanton, no. 83 
(CA); roar and swampy thickets, Mobile, Mobile Co., June, 
1878, r (US). Mussissrept: Ocean Springs, Jackson Co., 
Pst "06 (NY); Wisdom, Harrison Co., Tracy, no. 3418 
(G, NC, NY, US); open pine woods, 2 “idles west of Bay of St. 
Louis, Hancock Co., Correll & Correll, no. 9103 (D). Lousiana: 
pine flatwoods west of Covington, St. Tammany Parish, Brown, 
no. 6649 (La); open pineland north of Abita Springs, St. Tam- 

many Parish, Pennell, no. 4137 (D, NY, P); low moist grassy 
soil of open prairie, 3 ‘miles east of Robert, Tangipahoa Parish, 
Correll & Correll, no. 9217 (D). 

Forma controversa, f. nov. Satis antrorse hispidulis.—In- 
frequent in the range of the typical form. SourH CAROLINA: 
drainage ditch, 3 miles north of McClellanville, Charleston Co., 

odfrey & Tryon, no. 675 (G); grass-sedge bog or savanna, 12 
miles north of Georgetown, Georgetown Co., Godfrey & Tryon, 
no. 752 (G, type; NY, 1sorypr); shallow peaty pond in pine 
barren, 9 miles north of Georgetown, Georgetown Co., Godfrey 
& Tryon, no. 759 (D, G). 

ar. attenuata, var. nov. Foliis 1-3 mm. latis planis: culmis 
6-8 dm. altis gracilibus attenuatis: inflorescentia 2.2-3 dm. alta, 
culmo duplo vel triplo breviore; glomerulis 3-5 laxe subglobosis 
vel subhemisphaericis, 1-1.8 cm. latis: spiculis 4-4.5 mm. longis 
castaneis: achaenio 1.1-1.2 mm. lato 1.8 mm. longo: tuberculo 
1.4-1.8 mm. alto. Map 6.—Swamp-margins and moist pinelands 
of the Coastal Plain of the Carolinas (rare), Mississippi and 
Alabama. NortH Caroutna: Wilsons Mills, Johnston Co., 
Ashe (NC). Sours Ceiin flats across from paper mill, 


1944] Gale,—Rhynchospora, Section Eurhynchospora 111 


Hartsville, Darlington Co., July 5, 1909, Coker (NC). ALABAMA: 
in a wet place, Perdue, Coffee Co., Aug. 23, 1933, Blanton, no. 
83 (G, TyPE; CU, US, ISOTYPES) ; about swamp, Elberta, Baldwin 
Co., Aug. 21, 1926, Wolf (StB); common in swamp, Sprin ng Hill, 
Mobile Co. , Aug. 6, 1897, Bush, no. 248 (NY, US); moist sandy 
eae Theodore, Mobile Co., Aug. 30, 1912, Pennell, no. 4423 
(NY, Mobile, Mobile Co. _ July 20, 1897, Baker, no. 843 (NY). 
ioe Rac Ocean Springs, ‘Jackson Co., ‘Aug 21, 1889, Tracy, 
no. 124 (NY); same locality, July 18, 1891, earls (NY); same 
locality, Aug. 27, 1891, Seymour, no. 8 (CA, G, NC); Biloxi, 
Harrison Co. , July 31, 1900, Tracy, no. 6993 (G); same locality, 
July 25, 1892, Tracy, no. 1361 (G, US). 

An examination of R. microcephala Britt. ex Small and R. 
cephalantha var. pleiocephala from the coastal areas of the 
Carolinas, Mississippi and Alabama has brought to light the 
existence of certain specimens, the achenes of which are smaller 
than those of the already known varieties of R. cephalantha, but 
larger than those of R. microcephala. At first sight these speci- 
mens appear as intermediate between the two species mentioned 
and suggest, as preferable, a restoration of Britton’s R. cephalan- 
tha var. microcephala. However, one would expect these southern 
plants, if truly intermediate, to present a series of achenes 
ranging in size from those of R. microcephala to those of R. 
cephalantha. Actually, however, the achenes of these specimens 
appear to be of a definite size larger than those of the former, 
smaller than those of the latter species. The glomerules of this 
southern variety are from 2-4, hemispherical and indistinguish- 
able from the occasionally smaller, looser, hemispherical glom- 
erules of R. cephalantha. I am, therefore, assigning it to R. 
cephalantha as var. attenuata, to be distinguished from var. 
typica and var. pleiocephala primarily by its smaller achene; 
secondarily by its attenuated appearance, consistently weak 
sting and restricted southern range. 

R. CHALAROCEPHALA Fernald & Gale. Caespitose: leaves 
Fy (rarely) 2 mm. wide, flat, crowded, erect; upper margins often 
minutely serrulate; tips triquetrous: ‘culms subterete, slender to 
rarely stout, 1.8-8.3 dm. high: fascicles 3-7, remote, the majority 
2-5 lobed; the terminal fascicle turbinate to loosely subhemi- 
spherical, 0.9-1. 8 em. in diameter; lateral fascicles consistently 
turbinate, on included peduncles: spikelets lanceolate in outline, 
loosely aggregated, ascending to divergent; the fertile floret 
solitary, abruptly terminating the axis of the spikelet: scales 


112 Rhodora [APRIL 


lanceolate, acute, tightly imbricate about the achene and tuber- 
cle: bristles 6, robust, strap-shaped, retrorsely barbellate, erect, 
equalling the tubercle to slightly exceeding it: achene obovoid, 
lenticular, 0.9-1 mm. wide, 1.4-1.7 mm. long, with more or less 
definite shoulders and a drawn-out excessively slender gyno- 
phore; surface smooth, shining, umbonate, with depressed sides 
and a raised wirelike margin: tubercle subulate-attenuate, com- 
pressed, 1-1.6 mm. long; its base much narrower than the apex 
of the achene. PLATE 818, rigs. 1A and 1B, Map 8.—Ruopora, 
xlii. 426, figs. 1 and 2 (1940)—Swamps, lake-borders and road- 
side ditches of the Coastal Plain from the New Jersey pine 
barrens to Florida. New Jersey: wet pine barrens, Sims Place, 
Burlington Co., Drushel & Svenson, no. 6860 (G); Parkdale, 
Camden Co., S. Brown, no. 56 (P); peaty and sphagnous pond- 
hole depression near Hardingville, Gloucester Co., Long, no. 
47134 (P); moist pine barrens, Egg Harbor City, Atlantic Co., 
Mackenzie, no. 5558 (NY); Maurice River flats east of Vineland, 
Cumberland Co., Aug. 12, 1923, Bassett & Long (P); wet peaty 
pond-hole about 1 mile west of Bennett, Cape May Co., Long, 
no. 23488 (P). DrLAaware: moist soil, Lewes, Sussex Co., Aug. 
15, 1895, Commons (P); burned swamp, standing water, near 
Maryland line, Beaven, no. 69 (D). Maryianp: marsh in 
abandoned mill pond near Sharptown, Wicomico Co., O’Neill, 
no. 7430 (CU, . VIRGINIA: sandy and peaty border of Cat 
Pond south of Benns Church, Isle of Wight Co., Fernald & Long, 
no. 7357 (G, TYPE; P, isoTyPE); deep peat and mud, southeastern 
shore of Lake Drummond, Great Dismal Swamp, west of 
Wallaceton. Norfolk Co., Fernald & Long, no. 13570 (G). 
Nortu Caro.ina: pineland, Sampson Co., Blomquist, no. 5657 
(D); 4 miles north of Beaufort, Carteret Co., July 18, 1939, 
Engels (NC); savanna, 8 miles southwest of Jacksonville, Onslow 
Co., Godfrey, no. 6469 (G, NC); Pender Co., Hyams, no. 4979 


. GLOMERATA (L.) Vahl. Caespitose: leaves flat, 2.5-5 mm. 
wide, linear-attenuate, carinate toward the apex with finely ser- 
rulate margins: culms triquetrous, smooth, ascending, 0.6-1.1 


1944] Gale,—Rhynchospora, Section Eurhynchospora 113 


m. high, the upper 144-4 bearing the inflorescence, flexuous to 
arching: inflorescence composed of 3-6 narrow, flexuous, fascicu- 
late cymes or large loose glomerules: entire ovoid, 4.56.5 mm. 
long, subsessile, ascending to sprea 2-3-fruited (or if I- 
fruited the spikelet terminated by a ierile floret): scales acute 
to obtuse, loosely imbricated except when the spikelet is 1- 
fruited, originally apiculate but usually soon erose, castaneous 
to dark brown: bristles 6, straplike; the margins retrorsely and 
heavily echinate; the tips connivent about the apex of the tu- 
bercle: achene 1.4 mm. wide, 1.5-1.7 mm. long; its body sub- 
orbicular, with definite shoulders, basally prolonged; surface 
glossy with pronounced pale umbo and heavy wirelike margin: 
tubercle compressed-subulate, 1.3-1.8 mm. long. 

a. Var. typica. Inflorescence composed of 4-6 narrow 
cymes, the small dense ultimate fascicles of which are borne on 
3-4 slender approximate, ascending to arching branchlets which 
exceed one another so as to goal a continuous, wandlike 
effect: achene 1.2-1.4 mm. wide, 1.5-1.7 mm. long; gynophore 
of the achene thick and short: tubercle subulate, compressed; its 
base usually nearly apr iee! the summit of the achene. PLaTE 
819, ras. 1A and 1B; Mar 2.—R. glomerata Vahl, Enum. ii. 234 
(1806) ; Blake, RHODORA x 25, fig. 101918); Fernald, RHODORA 
Xxxvil. 401 (1935); Small, Mai. 180 (1933). Schoenus ile: Foes 
Linnaeus, Sp. Pl. i. 44 (1753). R. paniculata Gray, Ann. Lye. 
Ni ¥en: O11, pl. 6, fig. 21 (1835), non Presl (18 28). R. glomerata 
6 robustior Kunth, pass il. ate a. ar fe: glomerata var. 


(1920). Trion ‘glomerats [-a] Rael: Rep. Mich. Acad. Sci. xv. 
167 (1913).— Soa or sandy soil, New Jersey, Delaware 


achians of the Gavslenaat southward; Sentinal in Tennessee and 
northern een fairly frequent along the drainage of the 
Arkansas, Red, and Sabine Rivers. The species is so definite 
that only rootanasaaie specimens from western Louisiana and 
eastern Texas, where var. typica might be confused with the 
smaller var. angusta, are here cited. LovursiaNa: stream-bottom 
6 miles south of Franklinton, Washington Parish, Brown, no. 
6643 (La); open pineland, 1-2 miles north of Abita Springs, St. 
Tammany Parish, Pennell, no. 4133 (P); banks of Stoke Creek, 
4 miles southeast of Harrisonburg, Catahoula Parish, Brown, no. 
7383 (La); low wet soil along ditch, 1 mile south of Derry, 


114 Rhodora [APRIL 
Natchitoches Parish, Correll & Correll, no. 9988 (D); Sodus, 


miles east of Ruston, Lincoln Parish, Brown, no. 60 La); 
margin of pond near Minden, Webster Parish, Brown, no. 5354 
(La); meadow in pine hills near Mansfield, DeSoto Parish, 
Brown, no. 6093 (La); stream-bottom in long-leaf pine hills near 
Flatwoods, Rapides Parish, Brown, no. 6125 (La); in low prairies, 
Pointe aux Loups, Acadia Parish, Sept. 17, 1894, Langlozs (CU); 
mixed pine-hardwoods west of Sulfur, Calcasieu Parish, Brown, 
Nyland & Rogers, no. 8593 (La). Texas: swamps east of 
Mineola, Wood Co., Aug. 13, 1902, Reverchon (Mo); 10 miles 
northeast of Gilmer, ‘Upshur Co., Cory, no. 25659 (CU); swamps, 
Swan, Smith Co. , Aug. a 1902, Reverchon (Mo); sandy bogs, 
Grapeland, Houston Co., Palmer, no. 14436 (Mo, US); % mile 
east of Keechi, Leon Co. Cory, no. 25256 (CU); near Huntsville, 
Walker Co., duly, 1913, Young (Mo); = eae Polk Co., 
Palmer, no. 1 (Mo); Orange, Orange Co., 8, 1880, 
Letterman (Mo): Houston, Harris Co., July 20, 1919, e risher, no. 
45 (US). 

4b. Var. angusta, var. nov. Cymis 3-6, subglomeratis, re- 
motis: achaenio 1-1.1 mm. lato, 1.5-1.6 mm. longo, subgloboso, 
basi angusto, attenuato: tuberculo anguste subulato, basi quam 
achaenii apice valde angustiore, 1.3-1.8 mm. longo. —Low 
prairie, Arkansas (one collection), western Louisiana, aie eastern 
Texas. ARKANSAS: Grand Prairie, Pulaski Co., Har no. 7 
(G) and same locality, July, Harvey in Redfield H erb., ie. 15190 
(Mo). Lovistana: ditch in Holloway Prairie, Holloway, Rapides 
Parish, June 24, 1936, Brown, no. 6442 (La); common, low 
prairie, vicinity ‘of Lake Charles, Caleasieu Parish, Aug. 25, 
1898, Mackenzie, no. 442 (Mo, NC, NY): Lake Charles, Caleasieu 
Parish, Aug. 7, 1897, Tracy (G, U S); 1839, ex herb. Torrey (G). 
Texas: 21 miles north of Deweyville, Newton Co., Oct. 4, 1934, 
Cory, no. 10866 (G); 2.6 miles east Bie Camp Jackson, ante Co., 
Sept. 13, 1936, Cory, no. 19711 (CU, G); Cypress City, Harris Co., 
pee no. 886 (G); brook banks, Hempstead, Waller Co., June 10, 

1872, Hall, no. 718 (G, TyPr; Mo, US, ISoTYPEs). 

R. glomerata var. angusta possesses in common with var. 
typica a several-flowered spikelet maturing 1-3 achenes and 
terminated by a rudimentary floret. Its habit, however, is 
generally more robust; the leaves are 3-5 mm. wide, and the 
ultimate fascicles are densely aggregated to glomerulose. It is, 
however, on a basis of the achene that the varietal distinction 
chiefly rests. That of R. glomerata var. typica is broadly ovoid 
with a short, thick gynophore. The achene of var. angusta, 
however, is, as the name implies, narrower, somewhat shorter, 


— 73 im reeitt . rs ra eri x shine - aio a ai i ii basis nae a i c eee " sins ee i pe = i ee a i aa aT . bi as = eee wy eaee Lee pom if bill ae en aah tae oT ee > bi ror ee 


1944] Gale,—Rhynchospora, Section Eurhynchospora 115 


with a slender accentuated gynophore as in R. microcephala. 
Both varieties have in common the heavy wirelike margin, the 
prominent pale umbo and straplike retrorsely echinate setae. 

The geographic range of the new variety impinges upon that 
of var. typica in Arkansas and Louisiana, but continues westward 
into eastern Texas. 


5. R. CAPITELLATA (Michx.) Vahl. Caespitose: leaves 1.5-3.5 
mm. wide, flat, short, smooth, slightly carinate, becoming 
minutely serrulate on the upper margins: culms erect, slender, 
obtusely triangular, smooth, 1.9—9.2 dm. tall: the terminal cyme 
composed of 1-several ultimate, turbinate (rarely globose) 
fascicles on short included branchlets; lateral fascicles 1-5 on 
subincluded peduncles: spikelets ovoid, 3.4-5 mm. long, sub- 
sessile, 2—5-fruited (rarely 1-fruited and then terminated by a 
sterile floret): scales obtuse to acute, short-mucronulate, casta- 
neous, swiftly caducous, often forced apart by the maturing 
achene: bristles 6, straplike, but weaker than those of R. glo- 
merata, with retrorse barbules dwindling toward the base; tips 
convergent around the tubercle which they fail to equal or but 
slightly exceed: achene pyriform, 0.9-1.2 mm. wide, 1.3-1.8 mm. 
long, lenticular, plump, without a prominent umbo, edged by a 
narrow wirelike margin; the brown surface at maturity entirely 
or nearly uniform in coloring, smooth, lustrous: tubercle com- 
pressed-subulate, pale, 0.9-1.6 mm. long; its base widening 
nearly to cover the summit of the achene. PLATE 819, FIGs. 


and New England (excepting northernmost Maine and Upper 
Vermont), thence southward on the Coastal Plain to North 


116 Rhodora [APRIL 


Carolina; inland to the vicinity of the Great Lakes and along the 
Blue Ridge and Alleghanies from Maryland to Georgia; less 
common throughout western Tennessee and Missouri, with 
scattered stations in lower Georgia, northwestern Florida, Ala- 
bama, Mississippi, Arkansas, and eastern Texas; re-occurring in 
the coastal ranges of northern California and southern Oregon. 
The citation of specimens of this common, well-known species is 
limited to those from the extremities of its range. NEw Bruns- 
wick: ledges by Miramichi River, Northumberland Co., Fernald 
& Weatherby, no. 2408 (G); damp shores below Oakiok, York Co., 
Brittain, no. 11 (G). Marne: St. Francis, Aroostook Co., 1881, 
Furbish (NE). Carouina: 1794, Michaux Herb. (G, TYPE- 
PHOTO of Schoenus capitellatus). Grorara: low wet woods along 
small stream, near Thomasville, Thomas Co., Correll, no. 6484A 
(D); wet woods near Whigham, Decatur Co., Harper, no. 1185 
(NY, US); in the canyon at Tallulah "Falls, Rabun Co., alt. 
1600 ft., Aug. 3, 1893, Small (NY); densely ‘shaded sphagnum 
pockets i in aswamp, Sylvester, Worth Co., Svenson, no. 6928 (G). 
FLoripa: Quincy, Gadsden Co. (G, no collector designated but 
handwriting that of John C arey); ee near De Funiak 
Springs, Walton Co., Curtiss, no. 5926 (G, NC, NY). ALABAMA: 
moist sandy soil, DeSoto Falls, Jefferson Co. | Ruth, no. 127:(N Y}- 
ONTARIO: marshy places, Moon River, Muskoka, July 1882, 
Burgess (G); au bord dun petit lac, Timagami Park, Victorin, 
Germain & Meilleur, no. 45386 (G); Sandwich, Macoun, no. 
25338 (G, NY, US). Kenrucxy: near Harlan Court House, 
Seg? on » Kearney, no. 24 (G); wet flats of Red River, Logan 


Co. Me Short (P); wet flat on Ky. 98, Marshall Co., 
oar io. 3 (G). TENNESSEE: gravelly oak pea 6 miles 
east. of Crossville, Cumberland Co., Svenson, no. 4171 (G, P); 


wet roadside depression, Jamestown, Fentress Co., Svenson, 
de 4104 (G); bog, South Indian Creek, Unicoi Co., Price, no. 

oe ion ponds at Thompson’s, Williamson Co., Ruth, no. 
ee (N ; dry oak woods, Lawrenceburg, Lawrence Co., 
oa fe 4298 (G); along road in swamp, about 1.5 miles 
southeast of Hollow Rock Junction, Carroll Co., Svenson, no. 
425 (G, US); ditches, Henderson, Chester Co. , Bain no. 245 (G). 
Mississippt: Saratoga, Simpson Co. , Tracy, no. 8616 (G, NY). 
WISCONSIN: damp sandy shore of Crooked Lake, Siren, Burnett 
Co., Fassett, no. 7457 (G, f. discutiens); moist meadow, Mar- 
quette, Green Lake Co., Hotchkiss & Martin, no. 4407 (US); 


OKLAHOMA: wet open Deciea Antlers, Pushtamaha Co., Palmer, 
no. 9004 (CA, Mo, P). Trxas: Swan, Smith Co., Reverchon, no. 


Pe ee TON eS ee ee pee eS eee 


1944] Gale,—Rhynchospora, Section Eurhynchospora 117 


2917 (Mo). Catirornia: near Trinity Center, Trinity Co., 
Howell, no. 12845 (CA, G); Pitkin Marsh, 5 miles north of 
Sebastopol, Sonoma Co., Howell, no. 12677 (CA, in part f. 
discutiens; G, entirely f. discutiens). OrnGon: sphagnum bog, 
Brookings, Curry Co., Peck, no. 8793 (NY). 

In a recent paper Pfeiffer! appropriates the name R. capitellata 
(Michx.) Vahl for the common tropical and subtropical species 
of the Western Hemisphere which has heretofore been known as 
R. glauca Vahl. Since the name-bringing synonym, Schoenus 
capitellatus Michx., has been generally listed by American 
botanists, following the precedent of Torrey and Gray, in the 
synonymy of a widely spread and common species of the Series 
Albae, it is necessary to review the available information con- 
cerning the authentic material of S. capitellatus in the Michaux 
Herbarium. 

According to notes made by M. Gadaceau of the Paris Her- 
barium and quoted by Blake? in 1918, the Michaux Herbarium 
contains two sheets with the label, Schoenus capitellatus. On 
one of these are mounted specimens of R. Grayii Kunth (R. 
Elliettii Gray non Dietr.) and of the species mentioned above as 
belonging to the Series Albae. The Gray Herbarium has a 
photograph of this sheet, and the two specimens of R. Grayzi, 
designated as A and B, can be plainly seen, one on either side of 


_ the centrally placed subcapitate plant with inflorescence labeled 


The original label with the name, “Schoenus capitellatus” 
and the data “Hab. in Carolina”’ is affixed to the righthand side 
of the sheet and two slips of paper bearing Gray’s annotations 
“R. Elliottii Gray” and “R. glomerata” (the name wrongly 
applied by Gray and later botanists up to 1918 to the species 
here recognized as R. capitellata (Michx.) Vahl) are mounted in 
the opposite corner. Of the second sheet M. Gadaceau writes, 
“T’autre offre quatre beaux echantillons du R. glauca Vahl.” 
He then continues with remarks on the various Michaux speci- 
mens from the Richard Herbarium. To this herbarium belongs 
a sheet of Schoenus capitellatus, as labeled by Michaux, which 
has been referred by A. Richard to R. glauca Vahl. 

Pfeiffer apparently has not seen either the paper by Blake or 
the decisive specimens in the Michaux Herbarium; for neither are 


1 Fedde, Rep. Spec. So xlix, 75 (1940). 
* RHoporA, xx. 26 (1918). 


118 Rhodora [APRIL 


mentioned in his article. He does state, however, that authors 
prior to Boeckeler (especially Kunth) were cognizant of the 
relationship of Schoenus capitellatus to R. glauca. This state- 
ment, although incorrect in so far as it includes Torrey and 
Gray, both of whom identified S. capitellatus with plant C 
mentioned above, does indicate the probable source of Pfeiffer’s 
synonymy. He is apparently only taking over the synonymy of 
R. glauca as given by the older European botanists and inter- 
preting it in the light of the present International Rules of 
Nomenclature. Such an explanation also accounts for Pfeiffer’s 
inclusion of the basonym, Schoenus fascicularis Michx., of another 
well-known American species, R. fascicularis (Michx.) Vahl, in 
the synonymy of R. capitellata; for it is so placed by Kunth!. 
Undoubtedly the treatment by the earlier European botanists 
of R. glauca as a synonym of R. capitellata was the result of the 
inclusion of the specimens which were later identified as R. 
glauca Vahl under the name of Schoenus capitellatus in the 
Michaux Herbarium. However, viewed in conjunction with 
the original description of S. capztellatus in Michaux’s Flora 
Boreali-Americana”, there can be within the mixture of R. 
Grayii Kunth, R. glauca Vahl (?) and plant C of the Series 
Albae only one possible correct application of the name Schoenus 
capitellatus; for Michaux states that the specimen under con- 
sideration has “Capitula breviter pedunculata, interdum gemi- 
nata: semen compresso-obovatum” and ‘setulae retrorsum 
muricatulae.”” The habitat he gives as Carolina. Of the three 
species mentioned above R. glauca is eliminated at once by its 
range; for it is unknown in the United States. Nor does it have 
retrorsely barbed bristles. It is possible that the specimens of 
R. Grayit had a part in the more generalized portions of Michaux’s 
descriptions, but that they could not have been of sole considera- 
tion is evidenced in the portions of the description quoted above. 
The peduncles of the axillary “capitula” of R. Grayii are ex- 
serted and attenuated, the achene is conspicuously swollen 
above, and, as in R. glauca, the bristles are upwardly hispidulous. 
Following the precedent established by Blake, I am, therefore, 
applying the name R. capitellata (Michx.) Vahl to the species 


Bre ii. 297 (1837). 
Bor.-Am. i. 36 (1803). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 119 


aforementioned as of the Series Albae which is typified by speci- 
men C of the sheets labeled Schoenus capitellatus in the Michaux 
Herbarium. 

Forma controversa (Blake), comb. nov. Bristles geseany! 
serrulate.— Var. controversa Blake, Ruopora, xx. 28, fig. 3 (1918). 
R. glomerata fe minor f. controversa (Blake) Fernald, TinaDeeL 
xxxvil. 402 (1935). R. Smallii Britton ex Small, Fl. 1321, 1327 
(1903) and Man. 182 (1933).—Occurs infrequently throughout 
the range of the typical R. capitellata, with the exception of the 
southern states. 

Forma discutiens (Clarke), comb. nov. Bristles smooth.— 
ms glomerata var. discutiens Clarke ex Britton, Trans. N. Y. 

Acad. Sci. xi. 89 (1892); Britton, Man. 185 (1901); Small, Fl. 
195 (1903); Robinson & Fernald in Gray, Man. ed. 7: 201 (1 908). 
R. capitellata var. discutiens (Clarke) Blake, Ruopora, xx. 28, 
fig. 4 (1918). R. glomerata Meg minor f{. discutiens (Clarke) 
Fernald, Ruopora, xxxvii. 402 (1935).—Occurs sporadically 
throughout the range of ie “pied R. capitellata with the 
exception of the southern state 

In 1933 Britton elevated the specimens of R. capitellata with 
antrorsely barbed bristles to specific rank under the name, R&. 
Smallii. However, the antrorsely barbed phase of R. cephalan- 
tha, which has been generally known since the publication of 
Asa Gray’s Monograph in 1835, has never received nomencla- 
torial recognition. Smooth-bristled specimens have been found 
in R. capitellata and R. capillacea, and have been generally treated 
as varieties. The first has its var. discutiens, made by Clarke in 
1892 under R. glomerata, and transferred by Blake in 1918 to R. 
capitellata; the second, its var. leviseta E. J. Hill, 1876. 

In 1935, with a view toward establishing uniformity in the 
treatment of these similar cases, Fernald discussed the signifi- 
cance of both the variation in direction, and the failure alto- 
gether, of barbing in the bristles. He points out that in R. 
capillacea the smooth-bristled state occurs sporadically through- 
out the wide range of that species. In R. capzitellata (glomerata 
var. minor), as in Eleocharis and Scirpus, plants from the same 
locality have retrorsely or antrorsely barbed or even smooth 
bristles. Consequently he concludes that the atypical specimens 
are most adequately and satisfactorily covered as forms. I am 
accepting R. capillacea, f. leviseta (Hill) Fernald and following 
his precedent as to category in making the new combinations for 


120 Rhodora [APRIL 


hte * 
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i H 
: I “ 
f r 
an = é 
N ~ Bd ry Lr ped 
<oL ie 
Mh iP amt oo ° la 
> A { AN 
~ Rae Ae Sir Be se, | 
rh Pe ae | is L Baie e 
i Le f n 
\, z bad 
if rs z 
ey Oe oem on al : a Y/ 
> LYS i- eae 
H I mr 
Pande Peed x i| 3 
sat? a cd ek ee ; 
he , 
mec NN +\4 Pee 
1 ee Bae ees “Kage 
\! a v. “its mo , hae a git fa “Ss 
~~ afer rs a ais 3 


Range of 1, RHYNCHOSPORA CAPITELLATA; 2, R. GLOMERATA, var. TYPICA; 

, R. GLOMERATA, var. ANGUSTA; 4, R. MIcROCEPHALA; 5, R. CEPHALANTHA, 
var. PLEIOCEPHALA; 6, R, CEPHALANTHA, Var. ATTENUATA; 7, R. CEPHALANTHA, 
' var. TYPICA; 8, R. CHALAROCEPHALA; 9, R. ALBA 


19441 Gale,—Rhynchospora, Section Eurhynchospora 121 


both f. discutiens and f. controversa under R. capitellata. R. 
cephalantha var. typica f. antrorsa and var. pleiocephala f. con- 
troversa bring into line the hitherto neglected phases of that 
species. 

6. R. aupa (L.) Vahl. Caespitose, often densely so: leaves 
slenderly linear, 0.5-2.5 mm. wide, flat, becoming slightly carinate 
and then obscurely setaceous on keel and margins: culms slender, 
erect, ee 0.7-7 dm. high: fascicles 1-3, turbinate, 0.7— 
re vide; the smaller lateral fascicles exserted on slender, 
erect si ap spikelets ovoid, 3.5-5 mm. long, 2- (rarely) 2. 
flowered, often maturing 2 achenes, but if 1-fruited the spikelet 
terminated by an immature floret: fertile scales characteristi- 
cally whitish ‘be pale rufous, mucronate: bristles 10-12, stiffly 
connivent, obviously arranged i in 2:series on the elongated stipe, 
retrorsely "barbed, sparingly villous at the base, falling short of 
to exceeding the tubercle: achene pyriform, lenticular, biconvex, 
obscurely margined, with a prominent pale disc, irregularly 
lined, then generally darker toward the margins and faintly 
rugulose, 0.9-1.2 mm. wide, 1.6—-1.8 ee) mm. long: tubercle 
attenuate-subulate, compressed, 0.6-1. . long; the narrow 
base not equalling the breadth of Ao clan of the achene. 
PLatE 819, rigs. 4A and 4B; Map 9.—Enum. i. 236 (1806) ; 


Cyp. i. no. 92 (1834) and Ann. Lye. N. Y. iii. 213, pl. 6, fig. 24 
(1835) ; Boeckeler, ee XXXVil. 570 (1873); Britton & Brown, 
Ill. Fl. i. 277, fig. 651 (1896), i in part k. macra (Clarke) Small; 
Clarke in Urban , Symb. Ant. ii. 124 (1900); Britton, Man. 185 
(1901), R. macra (Britt,) ect - to specimens from Florida; 
Robinson & Fernald in Gray ed. 7: 200, fig. 323 (1908) ; 
Small, Fl. 194 (1903) and Reet 180 (1933); M. L. Green, List 
of Standard Species of Nom. Conserv. 9, no. 492 (1926), mimeo- 
graphed; Victorin, Fl. Laurent. 689, fig. 248 (1935); Fernald, 
Ruopora, xliv. 371 (1942). Schoenus albus Linnaeus, Sp. Pl. 1 
44 (1753); Fl. Danica ii. 5, pl. 320 Sree Michaux, Fl. Bor. nes 
i. 34 (1803). R. alba var. macra sensu Robinson & Fernald in 
Gray, Man. ed. 7: 201 ee non Disko R. luquillensis Brit- 
ton, Bull. Torr. Bot. Cl. 1. 56 (1923); Britton & Wilson, Sci. 
Surv. Porto Rico and Virgin Isl. v. 103 (1928). Triodon albus 
Farwell, Rep. Mich. Acad. Sci. xix. 253 (1917). Phaeocephalum 
album House, Am. Midland Nat. vi. 201 (1920). Dichromena 
alba Macbride, Field Mus. Pub. Bot. iv. 166 (1929).—Common 
in open sphagnum bogs from Newfoundland to Maryland, = 
in Virginia, southward only in scattered mountain bogs of Wes 
Virginia, North Carolina and Puerto Rico; inland to the area at 
the Great Lakes and Saskatchewan (one collection), reappearing 
to the west in northern Idaho and along the Pacific coast from 


122 Rhodora [APRIL 


northern California to southern Alaska; also in northern Eurasia. 
The citation of specimens of this well known and clearly defined 
species is limited to those from the outlying portions of its range. 
QUEBEC: openings in marly Arbor Vitae swamp, mouth of Bona- 
venture River, Bonaventure Co., Collins, Fernald & Pease, no. 
5819 (G); open peaty spots in larch swamp, Grindstone, Grind- 
stone Island, Fernald, Long, & St. John, no. 7015 (G, P); dans 
les tourbiéres, Isle 4 la Proie, Archipel de Mingan, V2ctorin, no. 
20225 (G); Father Point, Williamson, no. 1090 (P); in peat bogs, 
St. Hubert, Chambly Co., July, 1910, Victorin (US); tourbiére 
flottante, 93 miles au nord de Mont-Laurier, route Mont- 

aurier, Senneterre, Marie-Victorin, Rolland-Germain & Blain, 
no. 309 (G). DrLaware: New Castle, New Castle Co., Tatnall 
(G); swamps near Laurel, Sussex Co., Aug. 19, 1880, Commons 
(P). Maryuanp: 4 miles north of Salisbury, Wicomico Co., 
Shreve & Jones, no. 1278 (US); peat bog, Glenburnie, Anne 
Arundel Co., Aug. 19, 1905, Chrysler (G); Suitland Bog, Prince 
George Co., C. P. Smith, no. 3192 (CA); open white gravel bog, 
Powder Mill Bogs, near Lewiston, Prince George Co., Blake, no. 
10671 (G); sphagnous bog, northwest of Mountain Lake Park 
and vicinity, Garrett Co., on the Alleghany Plateau, alt. 720 m., 
Steele, no. 77 (US). Vireinta: wooded swamp of North Landing 
River, west of Pungo Ferry, Princess Anne Co., Fernald & Long, 
no. 13899 (G). Wersr Viraginta: Aurora and vicinity, Preston 
Co., alt. about 3000 ft., Aug. 15-Sept., Steele & Steele (NY, US). 

ORTH CAROLINA: roadside between Sparta and Roaring Gap, 
Alleghany Co., Blomquist, no. 5612 (D). SasKaTCHEWAN: bog, 
Dahlton, Aug. 9, 1936, Breitung (NY). Ipauo: Priest Lake, 
Bonner Co., Piper, no. 3756 (G, US); Minard’s Bay, Bonner Co., 
Priest Lake, alt. 660 m., MacDougal, no. 294 (NY). CALIFORNIA: 
Inglenook Swamp, Mendocino Co., Congdon, no. 67092 (G, US). 
Orecon: Hall, no. 568 (G); bog near Florence, Roosevelt High- 
way, Lane Co., Henderson, no. 13978 (P). WASHINGTON: in 
quaking sphagnum bog, Fazon Lake, Whatcom Co., Muenscher, 
no. 10145 (G); in floating bogs in Samish Lake, Whatcom Co., 
Suksdorf, no. 1014 (G, NY, US); bogs, rare, Seattle, King Co., 
Piper, no. 1121 (G, NY); in sphagnum bog, 28 miles south of 

acoma on Mt. Rainier Road, Pierce Co., Abrams, no. 9232 
(NY); Wreck Creek Prairie near Granville, Chehalis Co., 
Howell, no. 374 (NY, P, US); bog, edge of ditch, Onslow Station, 
N. P: -, 20 miles northwest of Hoquiam, Grays Harbor Co., 
Foster, no. 870 (US); bogs, Moclips, Grays Harbor Co., Cowles, 
no. 619 (G, Mo); in damp meadows of Baker Prairies, Grays 
Harbor Co., McGee, no. 556 (CA). British Cotumsta: Fort 
Rupert, Vancouver Island, 1904, Hunt (NY); Vancouver, Aug. 
28, 1893, Macoun (US); meadows, boggy and rocky, of northwest 
part of Calvert Island, south of Kwatshua, McCabe, no. 3083 


ia i i a a al 
diets i + a ten ene per aia ores ae 


1944] Gale,—Rhynchospora, Section Eurhynchospora 123 


(G); muskegs in sphagnum in wet places, borders of ponds, 
Smyth Island, Bardswell Group, McCabe, no. 3184 (G). ALASKA: 
sphagnum bog, Wrangell, Walker & Walker, no. 728 (G); Sani- 
tarium, Anderson, no. 290 (US, immature); bog, Ketchikan, 
Cowles, no. 1408 (US); vicinity of Loring, summer, 1903, Cham- 
berlain (US, immature); in marshes near Yes Bay, Howell, no. 
1683 (NY, US, immature); upland meadows, Back Bay, Gorman, 
no. 122 (NY, US, immature); marshy margin of lake, Prince of 
Wales Island, Walker & Walker, no. 904 (CU, G, NY, US). 
Puerto Rico: rocks, summit of El Yunque, alt. 1050 m., Gleason 
& Cook, no. X-86 (NY) and Britton & Bruner, no. 7626 (NY); 
Sierra Luquillo, Hioram, no. 364 (NY, Type of R. luquillensis; 
US, IsoTYPE). 

Britton’s R. luquillensis from the eastern mountains of Puerto 
Rico is morphologically inseparable from phases of R. alba. In 
general appearance it closely resembles the short, thickly caespi- 
tose specimens of R. alba collected on the Newfoundland table- 
lands. Gleason & Cook, no. X-86 is especially similar to Fernald 
& Wiegand, no. 2753. Also the spikelets of the Puerto Rican 
material, like those of the continental R. alba, are 1—3-fruited 
and usually (if 1-fruited, invariably) terminated by a sterile 
floret; and the achenes are identical with those of A. alba in all 
particulars. 

Pfeiffer! has published R. alba var. meridianus, based on speci- 
mens collected by Liitzelburg in northeastern Brazil. He 
states in the description that the bristles are 6-8, which seems to 
indicate that his plants vary, at least in this respect, from the 
typical. Unfortunately, I have not seen any of his material, for 
it would be interesting to compare it with the specimens from 
Puerto Rico. 


and borders of rills on peaty slopes, Great Barachois (or Barasway 
Bay), District of Burgeo and La Poile, Sept. 11, 1926, Fernald, 
Long & Fogg, no. 119 (G); bare spots on peaty and gravelly 
slopes, French (or Tweed) Island, Bay of Islands, Sept. 2, 
1926, Fernald, Long & Fogg, no. 118 (G, Type); wet bog-barrens, 
Trepassey, Avalon Peninsula, Aug. 16, 1924, Fernald, Long & 
Dunbar, no. 26344 (G); shallow pond-holes in tundra west of 


1 Fedde, Rep. Spec. Nov. xxxiii. 210 (1933). 


124 Rhodora [APRIL 


Rushy Pond, Valley of Exploits River, Aug. 28, 1911, Fernald & 

Wiegand, no. 4718 (G); heath on diorite tableland, Lookout 

Mt., Bonne Bay, alt. about 380 m., Aug. 26, 1910, Fernald & 
5 


Harry’s Brook, Aug. 9, 1929, R. B. Kennedy, no. 71072 (G); 
patches of damp peat with Utricularia, Middle Ridge, July 26, 
1937, Moir, no. 41 (G). MrqurLon: maricages, tourbiére prés du 
pont de Miquelon, July 31, 1901, Arséne, no. 98 (G). QUEBEC: 
dans la grande tourbiére, Saint Charles de Bellechasse, Aug. 11, 
1925, Rousseau, no. 20224 (G); Nah eee swamp, Natashquan, 
Saguenay Co., Sept. 4, 1915, St. John, no. 90201 (G). Nova 
Scotia: cg tibey pockets i in sandy plains, Middleton, Annap- 
olis Co., July 20, 1920, Bean & White, no. 20267 (G). PENNSYL- 
VANIA: ‘Vicinity ie lento: Lehigh Co., Pretz, no. 7793 (P); 
open (calcareous) marshy meadow, vicinity of Allentown, Lehigh 
o., Preiz, no. 10361 (P). Wisconsin: Oneida Reservation, July 
ricereere ee (G). British Cotumsta: Lulu Island, Aug. 11, 
BEA niger! (Clarke) Small. Caespitose: leaves 1.5-3.5 mm. 
wide, flat, ascending; upper margins serrulate: culms peal 


smaller, on slender exserted peduncles: a ikalote ovoid, 4-5 mm 

long, invariably 1-flowered, with the achene terminating the axis: 
seales mucronulate, rusty: bristles 18-20, obviously arranged in 
at least two series, connivent, exceeding the tubercle; the barbs 
of the distal portion retrorse, Abana their direction in the 
proximal half, and lengthened into a few hairs at the base: 
achene pyriform, lenticular, ra: 4 mm. wide, 2-2.1 mm. long; 
the base somewhat attenuate as in R, alba; the surface obscurely 
rugulose, pale over the umbonal region, irregularly lined, then 
generally darkened toward the margins: tubercle narrowly 
subulate, compressed, 1 mm. long; the base not verry the 
summit of the achene. PLate 820, ng 3A and 3B; Map 12.— 
pe 180 (1933). R. alba var. macra Clarke ex tae Than 


1944] Gale,—Rhynchospora, Section Eurhynchospora 125 


Texas: sandy bogs, Grapeland, Houston Co., Palmer, no. 14404 
(Mo) and no. 12844 (Mo); 2 miles south of Grap eland, Houston 
, Cory, no. 26080 (CU); Drummond, no. O81 (NY, ISOTYPE, 
srehateney 
This species, known only from Georgia, Florida, Mississippi and 
Texas,! was set off from R. alba Vahl by Clarke. He described 
it as having “clusters larger, sometimes 3 mm. broad, bristles 
more numerous 15-20.”’ In 1933 Small elevated R. alba var. 
macra Clarke to specific rank without adding anything of note to 
the original description. On examination, however, the spikelets 
of R. macra prove to differ from those of R. alba in several 
characters. In the former species each spikelet is invariably 
1-flowered so that the single maturing achene terminates the 
axis. In R. alba, on the contrary, each spikelet.is 2- (more 
rarely) 3-flowered. Commonly two achenes mature, the upper- 
most of which may or may not terminate the axis, depending 
upon the presence or absence of a third almost invariably sterile 
floret. If the spikelet is 1-fruited, however, the achene is always 
succeeded by a sterile floret. It is the presence in R. macra of a 
consistently 1-flowered spikelet, coupled with its generally grosser 
habit, its increased number of bristles, and larger achene that 
leads me to agree with Small in thinking it to be a good species. 
8. R. KNigskERNII Carey. Caespitose: leaves ap somone 
ceous to 1.8 mm. wide, involute when dry, smooth, 
serrulate on margins and keel: culm slender to filiform, jeraius 
1.5-5 dm. high: terminal fascicle 0.4-1 em. wide; 2-3 lateral 
fascicles remote at intervals along the entire length of the culm, 
with peduncles included: spikelets ovoid, 2—2.8 mm. long, su 
sessile, 2-3-fruited, terminated by a sterile Hoxets ohne encucous, 
castaneous to dark brown; lower scales apiculate, upper on 
slightly so: bristles 6, stiffly erect, retrorsely ie, falling 
short of to pyhad exceeding the body of the achene: achene 
obovoid, 0.6—-0.8 mm. wide, 1.1-1.3 mm. long, lenticular, bicon- 
vex, nearly reads rugulose, a shining yellow-brown in 
the center, becoming fragmentarily lined, then generally darker 
toward the margins: plat deltoid-subulate, sme lea 
0.4-0.6 mm. high. Pua ead ries. 2A and 2B; 
Am. Journ. Sci. Ser. 2: a 5 847); Gray, Man. hers 11848); 
Britton & Beswis Ill. Fl. i. O78, fig. 653 (1896); Britton, Man. 
185 (1901); Robinson & Fernald in Gray, Man. ed. 7: 201, 
fig. 325 (1908). R. Grayana Knieskern ex Carey, Am. Journ. 
s of R. macra from New England are erroneous, being founded on specimens 
of R. prog exhibiting gigantism. 


126 Rhodora [APRIL 


Sci. Ser. 2: iv. 25 (1847), as synonym, non R. Gray nap ere 
ines petty Knieskernii (misspelled Kneiskernit) House, 

and Nat. vi. 202 (1920).—Moist places in pine barrens ‘of 
Nee epee and Delaware. New JersEY: Point Hollow, 1843, 
Knieskern (NY, TYPE, annotated by poe Nt pe woods, Sh ark 
River, Monmouth Co., Mackenzie, no. 8003 (NY, P); (0 n bog 
iron ore?) sandy bogs southeast of Bamber, Ocean Co., Aug. 25, 
1909, Long (G, P); dominant on iron ore banks near Atsion, 
matey Co., Aug. 26, 1867, Parker, (G, P); moist humus, 
Parkdale, Camden Co., Pennell, no. 9030 (NY); pinelands, Egg 
Harbor City, Atlantic Co., Mackenzie, no. 8050 (NY). DeELa- 
WARE: swamps, Baltimore Hundred, Sussex Co., Sept. 10, 1875, 
Commons (P); swamp, near Gumboro, Sussex Co., Aug. 5, 1874, 
Commons (P). 


First distributed under the herbarium-name, R. Grayana 
Knieskern, this rare little species of the New Jersey and Dela- 
ware pine barrens was later described and published by Carey 
in 1847 as R. Knieskernii, the change in name being made 
because of R. Grayii of Kunth, 1839. : 


9. R. cAPILLACEA Torr. Caespitose: leaves filiform-setaceous, 
involute, at least on drying, becoming slightly carinate then 
serrulate on keel and margins: culms flexuous-erect, capillary, 
0.9-4 dm. high: fascicles ovoid, erect with 1—-10-spikelets, 3-8 
mm. wide; the single lateral fascicle remote on a subincluded 
peduncle: spikelets lanceolate to fusiform, 5-6 mm. long, erect, 
sessile or nearly so, 1—5-fruited: fertile scales eer RS to dark 
brown with the prominent midrib prolonged into a short mucro: 
bristles 6, erect to ascending, retrorsely G  iallnte, falling short 
of to exceeding the tubercle: achene 0.8-1 mm. wide 2mm 


rigs. 2A and 2B; 10. Mid. St. Py ae sae 
and Ann. Lye. N. Yi iii. 366 6 (1836); ney ae 
o bier: and Ann. | ye. N. Y. iii. 214, os a 835) ‘and 


: Man 

ed 7: 201, fig. 324 (1908). R. setacea crate MacMillan, 
Metasp. Minn. Valley, 104 (1892), non Vahl. Schoenus setaceus 
Muhlenberg, Descrip. Gram. 6 (1817), non Vahl. Triodon capil- 
laceus [a] Farwell, Rep. Mich. Acad. Sci. xv. 167 (1913). Phaeo- 
recom capillaceum Farwell, Rep. Mich. Acad. Sci. xxi. 361 
(1920).—Sporadic in marly bogs and on calcareous ledges from 
the western coast of Newioundland through southern Pennsyl- 


1944] Gale,—Rhynchospora, Section Eurhynchospora 127 


vania; more common inland in the region of the Great Lakes 
(excepting Lake Superior), south through Ohio, with scattered 
stations in western Virginia, Tennessee, northern Iowa, northern 
North Dakota, and Saskatchewan, and a concentration in 
southeastern Missouri. Cited specimens of this clear-cut species 
are limited to those from the extremities of the range. NeEw- 
FOUNDLAND: in bogs, Bonne Bay, July 26, 1930, Jansson (G, 
f. leviseta); Harry’s Brook near Dump Pool, R. B. Kennedy, 
no. 920 (G, f. leviseta). QUEBEC: open spots, marly Arbor 
Vitae swamps, mouth of Bonaventure River, Bonaventure Co., 
Collins, Fernald & Pease, no.4774 (G). New Brunswick: 
crevices of wet calcareous ledges, Gorge of Aroostook River, 
Victoria Co., Robinson & Fernald in Pl. Exsic. Gray., no. 44 
(CA, CU, G, NE, NY, P, US). Marne: calcareous ledges, Ft. 
Fairfield (above mouth of Aroostook River), Aroostook Co., 
July 11, 1893, Fernald (NE); ledgy river bank, Winslow, Kenne- 
bec Co., Fernald, no. 2785 (NE, f. leviseta). Vermont: 4th of 
July Slide, Mt. Willoughby, Orleans Co., Aug. 15, 1896, Faxon 
(NE, NY, US). Connecticut: border of pond, Salisbury, Aug. 
28, 1910, Phelps (G, NE). New Yorx: Watertown, Jefferson 
Co., 1834, Gray (NY, cited by Gray). New Jersey: limestone 
sink, White Pond, Sussex Co., Mackenzie, no. 4766 (NY); marl 
beach, White Pond, Warren Co., Griscom, no. 12120 (G). PENN- 
SYLVANIA: one mile east of Johnsonville, Northampton Co., 
Sept. 2, 1907, Van Pelt (G, P); in limestone, Dillerville Swamp, 
Lancaster Co., July 16, 1901, Heller (G; US, no. 4839). Vir- 
GINIA: boggy meadow fed by springs in limestone area, vicinity 
of Watauga, Washington Co., Carr, no. 585 (Penn.) Onto: 
cedar swamp, Champaign Co., Werner, no. 170% (NY); rather 
abundant in bog, Cedar Swamp, vicinity of Tremont City, Clark 
Co., Leonard, no. 2090 (US); Paxton, Ross Co., Aug. 1933, 
Pontius & Bartley (US). TeNNeEssEE: dolomitic limestone, 
Cedar Creek, Campbell Co., Underwood, no. 163 (CU). Iowa: 
bog, Estherville, Emmet Co., Walden, no. 1113 (G); marshy 
zone around springs emerging from a knoll 5 miles east of Ruth- 
ven south of the viaduct over Highway 18, Highland, Palo Alto 
Co., A. Hayden, no. 742 (G, P); abundant, forming a zone around 
a hanging bog, southwest of Silver Lake, Silver Lake, Dickinson 
Co., A. Hayden, no. 10886 (P); marshy plain in a hanging bog, 
Logan, Clay Co., A. Hayden, no. 8040 (NY). Norra Dakota: 
Turtle Mts., Bottineau, Bottineau Co., alt. 2000 ft. July 25, 
1896, Barber (US). SASKATCHEWAN: bog, rare, 4 miles southwest 
of Wallwort, Breitung, no. 13 ‘ : 

Forma teviseta (KE. J. Hill) Fernald. Bristles smooth, other- 
wise identical with R. capillacea..—Ruopora, xxxvii. 252 (1935). 


1 For discussion see treatment of R. capitellata f. discutiens. 


128 Rhodora [APRIL 


Var. leviseta E. J. Hill ex Gray, Am. Nat. x. 370 (1876); Britton 
& Brown, Ill. Fl. 278 (1896); een erst 185 (1901); Robinson 
& Fernald in Gray, Man. ed. 1 (1908). Phaeocephalum 
capillaceum var. levisetum Farwell een Mich. Acad. Sci. xxi. 
361 (1920).—Occasional in the range of the typical form in 
Newfoundland, Maine and the area of the Great Lakes. With 
the exception of the type, representative specimens of f. leviseta 
have been cited with those of the typical form. InpIANA: wet 
pine barrens, Pine Station, head of Lake Michigan, July 28, 
1875, Hill (G, Type of var. leviseta). 

Asa Gray, in reporting var. leviseta for Hill, mentions another 
possible variety of R. capillacea, characterized by 12 setae and a 
short ‘stipe’? which had been collected in Herkimer County, 
New York by J. A. Paine, 1864. The sheet in the Gray Her- 
barium labeled Litchfield, Hidden Lake, Herkimer County, 
New York, John A. Paine, 1864, is mixed. The two specimens 
with achenes bearing as many as 12 bristles are poorly developed 
R. alba (L.) Vahl. 

Series 2. PLumMosak (Clarke) Small, emend. Plants of damp 
or exsiccated pine barrens on the Coastal Plain, the West Indies 
and Central America. Caespitose: leaves filiform to 3 mm. wide: 
culms capillary to slender: inflorescence reduced to a few spike- 
ets or 1-2 spiciform or corymbiform fascicles: spikelets usually 
1— (rarely 2-3)-fruited, pedicellate to sessile: scales castaneous 
to pale brown, tightly imbricated: bristles heavily plumose in at 


Rhynchospora, Series B. Peden Sect. 2, Plumosae Clarke i in 

rban, Symb. Ant. ii. 105 (1900), in part. Rhynchospora § 
Eriochaete Gray in Torr. Ann. Lye. N. Y. iii. 363 (1836); Steud. 
Cyp. 139 (1855). 


Key TO Species IN SeRrES PLUMOSAE 


Spikelets 1-5, remote on 1-2 gene branchlets, the lowermost of 
which is abruptly CVE ao ee 10. R. oligantha. 


Fascicles ovoid he irre: eek rymbose; achene 1.4—1.7 mm 

wide, 22.2 mm. long; leaves 3-3 mim. wide: 6.0, <<<. aah 'R. intermedia. 
Fenition spieitorn: achene 1.1-1.4 mm. aides 1.4-1.8 m 

long; leaves filiform to 1 TU ee 19. R. plumosa. 


10. R. origaAnTHA Gray. Densely caespitose: leaves filiform- 
setaceous, canaliculate, smooth, blunt-tipped, ascending: culms 
capillary, erect or often weak, leafless, 1.6-3.7 dm. tall: inflores- 
cence reduced to 1-2 elongate capillary branchlets, the one erect 
and with 1—2-spikelets, the other abruptly divergent, with 1-3- 


a “ 
3 
q 
i 
9 
| 
3 
: 
; 
E. 
4 


1944] Gale,—Rhynchospora, Section Eurhynchospora 129 


spikelets, subtended by a long filiform erect often circinately- 
tipped bract which appears as a continuation of the culm; rarely 
a single remote lateral spikelet present: spikelets ovate to lanceo- 
late in outline, with 2-4 florets, 1-3-fruited, pale cinnamon- 
brown, 4-7 mm. long; scales ovate, som metimes mucronulate, 
tightly imbricate: bristles 6, partially plumose, slightly shorter 
than the achene to exceeding the tubercle or reduced a pth 
achene broadly elliptic or ovate, subterete, 1. 
2-2.6 mm. long, ener’ rugulose with faint longitudinal 
striae: tubercle conical, 0.3—-0.6 mm. in hei 

10a. Var. typica. Bristles slightly shorter than the achene 
to exceeding the Gowtee the lower Leh covered by dense, 
reddish, silvery-tipped hairs; the upper portion upwardly his- 
pidulous: achene broadly elliptic, Taba. As mm. wide, 
2.3-2.6 mm. long, occasionally having a trace of a bluish bloom 
over the castaneous to dark brown surface; the summit noticeably 
constricted under the wide basal flange of a conical-attenuate 
eee (0.4—0.6 mm. in height). sgeae 821, Frias. 2A ae 2B; 
Map 15.—R. oligantha Gray, Ann. Lye. N. Y. iii, 212. - ph 6 , fig. 
ya) (1885): Chapman, FI. So. U. s” ‘52 4 (1860), in ’ part ‘var. 
breviseta Gale: Gray, Man. ed. 6: 585 (1890); Britton & Brown, 
Ill. Fl. i. 277, fig. 650 (1896); Britton, Man. 184 (1901), ee 
breviseta as to specimens from Florida; Small, Fl. 194 (1903) 
and Man. 180 hoe Robinson & Fernald in Gray, Man. ed. 7: 


ae bog near th Sussex on ene 15, 1895, RTE 
(G, P). Norra CAROLINA: Schweinitz oe nyPE); i — 


clumps of wire grass, Sylvester, Worth oe Svenson, no. 7 
G). Frorma: west Florida, Chapman (P). ALABAMA: about 
swamp, Elberta, Baldwin Co., Aug. 24, 1924, Wolf (StB); low 
sandy soil, Mobile, Mobile Co., May 1, 1940, Sargent (Sargent 
Herb.). Texas: swamps, Swan, Smith Co., Reverchon, no. 2919 
(G, Mo, US); bogs, Hempstead, Waller Co., Hall, no. 715 (G, 
US); Drummond, no. 282 (G). 

10b. Var. br eviseta, var. nov. Setis plerumque ad 6 cristas 
villorum argenteorum reductis; saepe 1-2 prolongatis apicem 
versus serrulatis achaenio duplo brevioribus: achaenio ovoideo 
1.6 mm. lato 2 mm. longo, caeruleo-pruinoso; tuberculo breviter 


130 Rhodora [APRIL 


conico 0.3-0.4 mm. longo. PLATE 821, rig. 2C; Map 14.—R. 
oligantha sensu Kiikenthal, Fedde Rep. Spec. Nov. xxiii. 207 
(1926) and xxxii. 76 (1933); non Gray.—Low areas in pine 
barrens and savannas, Florida Peninsula, western Cuba, Jamaica 
and Hispaniola. FLortpa: damp pine barren, Duval Co., 
Fredholm, no. 5179 (G); moist pine barrens near Jacksonville, 
Duval Co., June, Curtiss, no. 3165 (P, US); damp pine barrens 
near Jacksonville, Duval Co., June 19, 1896, Curtiss, no. 5687 
(G, TypE; NC, US, IsoTyPEs); near Jacksonville, Duval Co., 
May 26, 1893, mise pe: 4116 (US); low pine barrens, June, 
1884, Curtiss (US); Winter Park, Orange Co., Apr. 1919, Francis 
(US); rather dry pine piece about 1144 miles south of Starke, 
Bradford Co., May 18, 1909, Harper, no. 39 (US); cut-over 
flatwoods, west of Lake Reedy, Frostproof, Polk Co., May 7, 
1931, McFarlin, no. 5138 (CU); Tampa, Hillsborough Co., May, 
1876, Garber (G). CuBA: in alg moist places (between 
Aleatrez Grande and Ale. Ch hico) at Laguna “ges Pinar del 
Rio, Nov. 18, 1923, Ekman, no. 18129 (NY, US); Laguna 
Restinga, between Palmarejo and Las Set Pinar del Rio, 
Nov., 1923, Ekman (NY). Hispaniona: savanna in Rhexia 
association, not rare, El Valle, Sabana de la Mar, prov. de 

amana, Cordillera ee Santo Domingo, July 11, 1930, 
Ekman, no. 15652 (G, NY, US). Jamarca: in small patches 
amongst shrubs and | grasses, Hollis’s Savanna, Upper Clarendon, 
alt. 2400 ft., Jan. 12, 1915, Harris, no. 12249 (NY, US). 

11. R. INTERMEDIA (Chap m.) Britt. Caespitose: leaves 
canaliculate, carinate, 2-3 mm. wide, margins mostly upwardly 
serrulate: culm s terete, erect, lancer 2.5-6.7 dm. tall: fascicles 
1, rarely 2, congested, irregularly corymbiform or broadly ovate 
in outline, 0.9-2.2 em. wide; the lateral fascicle when present 
smaller and eres deve! ovoid, Sasa sessile, with 2 


browned, 1.4—-1. mm. wide, 2-2.2 mm. long: tubercle conical- 
apiculate, depressed, 0.6 mm. P aye PLATE 820, rics. 4A and 
4B; Map 13.—Trans. N. Y. Acad. Sci. xi. 87 (1892), non Bey- 
rich; Small Fl. 194, 1397 (1903) and Man. 180 (1933). R. plu- 
mosa var. intermedia Chapman, Fl. So. U. 8. 524 (1860). BR. 
pineticola Clarke, Kew Bull. Add. Ser. viii. 40 (1908). Phaeo- 
cephalum intermedium House, Am. Midland Nat. vi. 202 (1920). 
Dry sandy pine barrens, peninsula of Florida. FLortpa: Hiber- 
nia, Clay Co., March, 1869, Canby (NY); Gainesville, Alachua 
Co., March, 1876, Garbe r (NY): dry pine barrens, oy 
Inlet, Volusia Co., Vartan no. 3173 (CA, G, NC, NY US); 
Lake Butler, Orange Co., "Beckwith, no. 558 (US): Packie’ Lake 


i 
PN ae eee ae Ne RE cere Neve es 


1944] Gale,—Rhynchospora, Section Eurhynchospora 131 


Co., Nash, no. 2020 (G, NY, P, US); dry pine barrens, Eau 
rallie, Indian River, Brevard Co., Curtiss, no. 5703 (D, G, NY, 

US); hammock near St. Cloud, Osceola Co., Small, DeWinkler 
& Mosier, no. 11166 (NY); Tampa, Hillsborough Co., Britton & 
Wilson, no. 20 (NY); flatwoods, Lee Co., Hitchcock, no. 427 (G, 

Y, US); Palma Sola, Manatee Co., Tracy, no. 6998 (G, NY, 
US); Miami, Dade Co., June, 1877, Garber (G, P, US); dry 
sandy ridges near coast, Apalachicola, Franklin Co., Chapman 
in Biltmore Herb., no. 4481 (G, NY, US). 

R. intermedia can be arbitrarily distinguished from R. plumosa 
on a basis of size. Its leaves are wider, the spikelets larger, and 
the achene 2—2.2 mm. long in contrast to 1.6—-1.8 mm. long for 
that of R. plumosa. There is a pronounced tendency also for the 
fascicles of R. intermedia to be irregularly globose rather than 
elongated-spiciform. 


cylindric spike, occasionally 3 em. long, 1 cm. wide; a single 
remote lateral ‘spike’? may also occur on a slender exserted 


i. 58 (1816); Gray, Ann. Lye. N. Y. iii. 203, pl. 6, fig. 10 (1835); 
Chapman, FI. So. U. 8. 524 (1 . Ant. 
ii. 123 (1900); Small, Fl. 194 (1903) and Man. 180 (1933); 
Britton, Mem. Soc. Cubana Hist. Nat. ii. 195 (1916). BR. semi- 


pma : me 8S 
Man. 180 (1933). R. penniseta Grisebach, Cat. Fi. Cub. | 
(1866); C. Wright in Sauvalle, Anal. Acad. Ci. Habana, viii. 84 


Plain from North Carolina south to the tip of Florida, and west 
to Texas; also in the western provinces of Cuba, the Isle of Pines 


a 


132 Rhodora [APRIL 


Range of 10, Ruyncno 
MACRA; 13, R. INTERMEDIA: 


CuapManil; 20, R. BRAcHYCHAETA; 21, R. soua; 22, R. souirarta; 23, R. 
upA; 24, R. rusca; 25, R. pueranrua; 26, R. Curtissi; 27, R. HARPERI; 28, 

R. crinipes; 29, R. ruscorpeEs; 30, R. Leproruyncna; 31, KR. Gacert; 32, R. 

JOVEROENSIS; 33, R. BaLpwinu: 34, R. FerNaupu; 35, R. FrLIFOLIA 


TT ee ee re eee eee 


Ee ee an nee ey eh Tee 


1944] Gale,—Rhynchospora, Section Eurhynchospora 133 


and Central America. NortH Carouina: dry savanna, 15 miles 
southeast of Greenville, Pitt Co., Blomquist, no. 11252 (D); 
savanna at Chocowinity, Beaufort Co., Godfrey, no. 5403 (G); 
sandy soil, marsh, Harker’s Island, Carteret Co., Randolph & 
Randolph, no. 843 (G); moist black soil, low ground, cen, 
Onslow Co., Randolph & Randolph, no. 946 (G); savanna at 
Burgow, Pender Co., Godfrey, no. 4736 (G, NC); sandy pine 
barrens, west of Leland, Brunswick Co., W tegand as Manning, 
no. 619 (G); dry sandy soil, Fayetteville, Cumbe 
Biltmore Herb., no. 243b (CU, G, P). Sours Yap peaty 
excavated area in savanna at side of road, 12 miles north of 
Georgetown, Georgetown Co., Godfrey & Tryon, no. 7: (CU, DB, 
oN ; 4 mile south of Manning, Clarendon Co., Stone, 
no. 713 (P); open white sandy, oak-pine woods, 1 mile east of 
Eutawville, Orangeburg Co., Godfrey & Tryon, no. 833 (G). 
EORGIA: in pinetis aridis, Geor. Car Elliott (G, TyPE-PHOTO); 
moist pine barrens, Bullock Co., Harper, no. 893 (G, NY); low 
pine barrens southeast of Dublin, Laurens Co., Harper, no. 1372 
(G, NY, US); areas rocks near Ohoopee River, Tattnall Co., 
Harper, no. 2156 (G, US); pine barren, 2 miles south of Savannah, 
Chatham Co., Eyles, no. 6087 (CU ); pine barrens a few miles east 
of Eulonia, McIntosh Co., Eyles, no. 6437 (CU); ir mney at. in 
piney woods, Billy Island, Charleton Co., Bradley, no. P); 
moist pine barrens, Coffee Co., Harper, no. 706 (NY, US): 
forming wiry clumps i in moist pine barrens south of Sylvester, 
Worth Co., Svenson, no. 6936 (CA, G); pine barren, north of 
Hahira, Cook Co. , Eyles, no. 3796 (CU); damp margin of cypress 
pond, about 1 mile north of Jakin, Early Co., Harper, no. 3629b 
(NY). Frioripa: dry oe e barrens near Jacksonvil e, Duval Co. is 
Curtiss, no. 4867 (G, NY, US); north of Macelenny, Baker Co., 
re no. 5864 (C U); ieee Clay Co., March, 1869, Canby 
P, US); Cypress-Ilex-Myrt. swamp, west of Lawtey y, Brad- 
ford Co., West & Arnold (CU); in a wet ditch in pineland, 
Gainesville, Alachua Co., O’ Neill, no. 621 (CU); moist flatwoods, 
Welaka, Putnam Co., June 29, 1939, Laessle, no. 17 (CU); low 
pine barren, Okeechobee Region, Brevard Co., Fredholm, no. 
740 (G); in a low pineland, Aripeka, Pasco Co., Leonard, no. 
1414 (CU); Tampa and vicinity, Hillsborough Co., May, 1897, 
Berg (NY); in dry sandy soil, about 5 miles north of Parrish, 
Manatee Co., Moldenke, no. 1052 (D, NY); Punta Gorda, De- 
Soto Co., Eaton, no. 1284 (G); in moist nog at at edge of 
ditch, Fort Myers, Lee Co., Moldenke, no. 675 (D, NY); Miami, 
Dade Co., Mar., 1877, Garber (G, P); in low pineland, Otter 
Creek, Leon Co., O’N eill, no. 725 (CU); Slgmagater Pe 
Franklin Co., Small, DeWinkler & Mosier, no. 11248 (NY); 
boggy ee: sandy meado ow, 7 miles west of Sneads, Jackson 
Co., Wiegand & Manning, no. 678 (G); moist pine barrens, 
Argyle, Walton Co., BS hae: no. 6488 (G, NC, NY, US); small 


134 Rhodora [APRIL 


savanna near coastal highway, about 12 miles east of Pensacola, 
Santa Rosa Co., Harper, no. 3745 (G, NY). ALABAMA: Gates- 
wood, Tracy, no. 8617 (G, NY, US); Mobile, Mobile Co., Sulli- 
vant (G). Mississippi: Ocean Springs, Jackson Co., Skehan, no. 
22617 (G); Biloxi, Harrison Co., Tracy, no. 4861 (G, US). 
LOUISIANA: open pineland, 1 mile north of Abita Springs, St. 

ammany Parish, Pennell, no. 4154 (NY); New Orleans, 1834, 
Dr. Ingalls (NY, type of R. semiplumosa, very immature); wet 
prairies, Welsh, Jefferson Davis Parish, Palmer, no. 7669 (CA, 

o, P, US); low prairies, vicinity of Lake Charles, Calsasieu 
Parish, Mackenzie, no. 444 (Mo, NC, NY). Texas: San An- 
tonio, Bexar Co., Sept. 20, 1891, Plank (NY); E. Tex., Wright 
(G). Cusa: in sandy pinelands close to the laguna, Laguna 
Santa Maria, Pinar del Rio, man, no. 17250 (G); moist 
places, Mateo Sanchez, Pinar del Rio City, Pinar del Rio, Ekman, 
no. 17941 (US); pine woods, Herradura, Pinar del Rio, Earle, 
no. 756 (NY, US); Vinales, Pinar del Rio, Leon, no. 18905 (CU); 
Pinales, Los Almacigos Pinar del Rio ? July 29, and high pinales, 
Vinales, Pinar del Rio, Mar. 1, Wright, no. 3398 (G; NY, US, 
same number without locality); savanna, San Pedro and vicinity, 
Isle of Pines, Britton & Wilson, no. 14296 (G, NY, US); vicinity 
of Sopopo, Buenos Aires, Trinidad Mt., Santa Clara, Smith, 
Hodgdon & Gonzalez, no. 3341 (CU, G, US 

Series 3. Chapmaniae, ser. nov. Inflorescentia saepe ©¢ 
fasciculo solitario terminali composita: spiculis sterilibus fusi- 
formibus saepe numerosis: setis antrorse serrulatis achaenio 


short, often apiculate.—Rhynchospora III. Pusillae Small, Man. 

175 (1933), in part. Rhynchospora, Series B. Diplostyleae, 

Divisio 4. Psilocarya, Sect. ii. Pauci-Nucigerae Clarke, Kew 
Bull. Add. Ser. viii. 119 (1908), in part. 
(To be continued) 


Soe . 
pO EE ne ee ee ee eee ee 


ee Se ee Oe ee ee ae 


the wees (ated 


Nene tins 


1944] Gale,—Rhynchospora, Section Eurhynchospora 159 


RHYNCHOSPORA, SECTION EURHYNCHOSPORA, IN 
CANADA, THE UNITED STATES AND THE 
WEST INDIES 
SHIRLEY GALE 
(Continued from page 134) 


Ky TO SPECIES IN SERIES CHAPMANIAE 


a. apres ¢ Pepe seigestc by tiny dark pits with the exception 
h oth region of the disc, o oe 2 the p shes are obscured, 
the 34 ‘bristles equalling the sath els. 2% 
b. Basal leaves 4-6 m: wide, obtusely tipped d, short, forming 
a foouthe: distal pont n of the midribs of at least the 
Circe floral scales conspicuously see settee ciliate : 
. RR. ciliaris. 
b. Basal leaves 4 5-3 me: wide, acute, elongate, erect; floral ; 
scales not ciliate... . 
Se ae se 3-4, ecuailing fh Giberele 2 ye ae 14. R. solitaria. 
3 2-3, rudinentaty 6.4.) 95. 52 Se ea ees 15. R. sola. 
a. heres or aioe a0 ae S camneks striate as the result of 
e crowding of pape inconspicuous oblong alveoli; bristles 
absent or rudimentary. ... 
d. Fascicles 1-3, italy ovoid; tubercle triangular-subulate; 
scales acute a OrisuINte os or Sees SA ee ee 16. R. brachychaeta. 
d. Fascicles solita Ponies rm; tubercle triangular to tri- 
pega alate Scales with aristate tips at least 0.3 
¢. Bases typically bulbous, sheathed by short fibrous acute 
te scales; floral scales silvery to-redlieh. 3... 24. 17. R. pallida. 
é. Duonn ellowish-brown to chestnut. oh 
sf personne homogeneously pale, not lustrous, 1.2-1.3 m 
ide, ath m. long; (Sena exceeded by 1-2 ak 
Btall racte hicccas 4 0 CRG a he Ne ee ae 18. R. nuda. 
f. Achenes ale except for a conspicuous dark patch at 
the base and apex, glossy, 0.8-1 mm. wide, 1-1.2 
mm. racy fascicles panel St by. long. setaceous 
bracts which are often somewhat circinate....19. R. Chapmanat. 


13. R. crurarts (Michx.) Mohr. Coarsely tufted: — 
leaves short, suggesting a rosette, flat, 4-6 mm. wide, their m 
gins and narrow keels silvery-ciliate ; median and costal veins 


chee a elliptic, strongly erp 2 L 5-1.6 mm. wi e, 
1.8 mm. long; the surface pricked by minute pits and dark 


160 Rhodora [May 


brown except for the rege pale central disc: tubercle broadly 
deltoid, compressed, 0.4-0.6 mm. high, usually wrinkled an 
slightly encrusted by the ae of the achene. PLATE 822, 
rigs. 2A and 2B; Mar 17.—Contrib. U. 8. Nat. Herb. vi. 408 
(1901) ; Small, Man. 183 (1933). Schoenus ciliaris Michaux, Fl. 
Bor.-Am. i. B ei — R. ciliata Vahl, Enum. ii. 235 (1806): 
Gray, Ann. L Y. iii. 209, pl. 6, fig. 19! (1835); Chapman, 
Fi-Se.. UU. 8: mie! (1860); Small, FI. 196 (1903); Britton, Trans. 
cc ¢ CRG, et: XL. 90 (1 892). Rk. Rappiana Small, Man. 179, 
1503 (1933). Phaeocephalum ciliatum House, Am. Midland Nat. 
vi. 201 (1920).—Boggy savannas and low pinelands of the Coastal 
Plain from North Carolina southward throughout the peninsula 
of Florida and west to eastern Louisiana. NortH CAROLINA: 
savanna at Newport, Carteret Co., Godfrey, no. 5772 (G); moist 
black soil, low ground, Dixon, Onslow Co. , Randolph & Randolph, 
no. 954 (G); pineland near Carolina Beach, New Hanover Co., 
Godfrey, no. 4699 (G); Supply Road, Southport, Brunswick Co., 
Blomquist, no. 5653 (D); pineland at Hallsboro, Columbus Co. 
Godfrey, no. 6252 (G, NC). SoutrH Caro ina: grass-sedge bog 
or savanna, 12 miles northwest of Georgetown, Georgetown Co., 
Godfrey & Tryon, no. 747 (D, G, NY, P); grass-sedge bog or 
savanna, 3 miles southwest of Manning, Clarendon Co., Godfrey 
& Tryon, no. 924 (G, NY); Sumter, Sumter Co., Bartram, 
no. 3288 (P). Grorgia: rather dry pine barrens, Coffee Co., 
Harper, no. 701 (G, US); intermediate pine barrens, Suwannee 
ake, Ware Co., J. S. Harper, no. 84 (D, P); rather dry pine 
barrens south of Empress, Brooks Co., Harper, no. 1630 (G, Fs 
; dry soil, Valdosta, Lowndes Co., May 27, 1940, Sargent 
(Sargent Herb.): moist pine barrens, Bullock Co., Harper, no. 
887 (G, NY, US); pine barrens, Bethesda Church, Effingham 
Co., Eyles, no. 6106 (CU); in meadow, New England Camp, 8 
miles north of Brunswick, Glynne Co., M oldenke, no. 5207 (NY); 
piney woods, Billy Island, Charleton Co., July 7, 1912, Bradley 
(P). FLORIDA: moist ~ barrens near Jacksonville, Duval Co., 
Curtiss, no. 3146 (CU, D, G, P, US); Hibernia, Clay Co., March, 
1869, Canby (G, US); left side Palatka Road ‘about 1 mile from 
florist’ s, Gainesville, Alachua Co., West & Arnold, no. 7 (CU); 
low pineland near swamp on ro ad betwee ae Piskan d and New 
Smyrna, Volusia Co., Sept. 11, 1926, Dr. Ball? (NY, type of R. 
Rappiana); ine Tomoko Creek, Volusia Co. , Michaux Herb. (G, 
TYPE-PHOTO of Schoenus ciliaris; NY, fragment from Michx. 
Herb.); low pineland, vicinity of Eustis, Lake Co., Nash, no. 
' Figures 18 and 19 were apparently reversed in the printing. R. ciliata is fig. 19, 
not 18 as listed. 
* The type of R. Rappiana, although undoubtedly sent to Small by Rapp, 
year later, followed up by a collection made by Rapp himself, was actually eit 
. Ball as stated on the memorandum attached to the sheet. 


RNa tO Oe ee S 


1944] Gale,—Rhynchospora, Section Eurhynchospora 161 


534 (CU, G, NY, P, US); in sandy field about 3 miles west of 
Bithlo, Orange Co., Moldenke, no. 201 (D, NY); low pine barren, 
Okeechobee region, Brevard Co., Fredholm, no. 5177 (G, US); 
on the prairie, Kissimmee Park, Osceola Co., O’Neill, no. 6234 
(CU); flatwood east of Loughman, Polk Co., McFarlin, no. 4282 
(CU); Tampa, Hillsborough Co., Oct., 1877, Garber (G, P); in 
water’s edge, near St. Petersburg, Pinellas Co., Deam, no. 
2904 (G); Braidentown (? Bradenton), Manatee Co., Tracy, no. 
7126 (G, NY, US); Istokpoga Prairie between Lake Istokpoga 
and Kissimmee River, Highlands Co., Small & DeWinkler, no. 
9056 (NY); in a low pineland 11 miles east of Okeechobee City, 
Okeechobee Co., O’ Neill, no. 7678 (CU); Dade Co., Nov—Dee., 
1903, Eaton (NY); wet pine barrens, Apalachicola, Franklin Co., 
Chapman in Biltmore Herb., no. 862a (G, NY, US). ALaBaMa: 
swamp, Elberta, Baldwin Co., Aug. 21, 1926, Wolf (StB); com- 

1 h 295 (NC, NY 


’ 


Small compared his species, R. Rappiana, with R. pallida 
Curtis, but did not mention any possible relationship with R. 
ciliaris. However, Small’s species has the short, blunt-tipped 
tuft of basal leaves with their silvery cilia, the ciliate bracts and 
scales, and the pitted, strongly lenticular achene of R. ciliaris. 
I cannot help but think that Small neglected to compare his 
material with the older species, as the two are unquestionably 
identical. 


4. R. sonrraria Harper. Sparingly caespitose or solitary: 
leaves linear, erect, flat, 2.5-3 mm. wide, smooth with blunt tips: 
culms terete or nearly so, slender, erect, smooth, 5.2-6.4 dm. 
high: inflorescence a single turbinate to subhemispherical fascicle, 
1.5 cm. wide: spikelets lanceolate to fusiform, 6-7 mm. long, 
acuminate, sessile, 1-flowered, split open by the maturing achene: 
scales chestnut, with an aristate tip 0.6-0.8 mm. long: stamens 
2: bristles 3-4, extremely fragile, upwardly serrulate, equalling 
the tubercle: achene obovate, lenticular, 1.3 mm. wide, 1.4 mm. 
long; the surface, with the exception of a pale smooth disc, 
brown and pitted: tubercle triangular-apiculate, compressed, 
0.6 mm. long. Prats 822, rigs 4A and 4B; Map 22.—Bull. 
Torr. Bot. Cl. xxviii. 468 (1901); Small, Fl. 193 (1903) and Man. 
182 (1933). Phaeocephalum solitare House, Am. Midland Nat. 
vi. 202 (1920).—Southern Georgia. Gxroreta: moist pine bar- 


162 Rhodora [May 


rens, Tifton, Berrien Co., Harper, no. 668 (NY, TYPE; G, ISOTYPE) 
and no. 1677 (G, US) 


At first glance R. solitaria appears very similar to R. pallida 
M. A. Curtis, but its details—the simple non-tuberous bases, 
blunt-tipped leaves, terete culm, chestnut spikelets, aristate 
scales, well developed bristles and pricked surface of the achene— 
indicate that it is clearly a distinct species which, so far as I am 
aware, is known only from collections made in the type-locality. 


15. R. sola, sp. nov. Planta laxe caespitosa: foliis erectis 
valde involutis saltem siccatis 1.0-1.5 mm. latis; apicibu 
obtusis: culmis subteretibus, tenuibus, erectis, saepe flexilibus, 
3.2—5.7 dm. altis; inflorescentia ex fascicu o uno parvo compacto 
turbinato vel hemisphaerico 0.8-1 em. lato constata; bracteis 
obscuris fasciculum non aapbrenkiius? spiculis lanceolatis, con- 
fertis, sessilibus 2-floris, monocarpis, 5 mm. longis: squamis 
mucronatis, dense imbricatis castaneis: setis 2-3, rudimentariis, 
antrorse serrulatis: achaenio valde lenticulari-obovoideo 1.1-1.2 
mm. lato 1.2-1.6 mm. longo foveolato fusco; disco medio pallido: 
oe compresso-deltoideo. PLATE 820, rigs. 1A and 1B; 
Map 21.—R. fascicularis sensu C. Wright in Sauvalle, Anal. 
head “Gk Habana, viii. 84 (1871) and FL. Cub, 180 See m 
part; non (Michx.) Vahl. R. distans sensu Grisebach, Cat. 
Cub. 243 (1866), non (Michx.) Vahl.—Low pinelands of saute 
Cuba. Cusa: pinales, Hato Quemado, Pinar del Rio, Nov. 20, 
1862?,! and San Juan-Guanes, Wright, no. 3397, sheet ‘labeled B 
(G, in part); Wright, no. 3397, ‘sheet labeled A (G, Type); Wright, 
no. 3399 sheet labeled A (NY, in part)’; Wright, no. 3397, in 
Herb. Canby, no. 396 (US, in part); ae savannas, Chirigota, 
Pinar del Rio, Oct. 26, 1863?,° Wright, no. 3399 (US, in part); 
Sabana de la Maguina, south of Pinar del Rio City, Pinar del 
Rio, November 28, 1940, Léon & Alain, no. 19422 (G). 


With the exception of the Léon & Alain collection, no. 19422, 
Rk. sola is known only from specimens collected by Charles 
Wright and distributed as R. distans no. 3397 and R. deflexa 
Gris. no. 3399. I have seen 3 sheets of no. 3397; of the two located 
at the Gray Herbarium, one contains specimens of R. sola 


exclusively ; this I am designating as the rypr. The other sheet — 


is mixed, containing on the left a specimen of R. fascicularis 
(Michx.) Vahl, var. typica. Sheet no. 3397 from the Canby 

t See Underwood, ae Torr. Bot. Cl. xxxii. 297 (1905). Underwood misread “Que- 
mado” as “Quenia 


2 See footnote to ee no. 3399 under R. fascicularis (Michx ) Vahl. 
3 See Underwood. Bull. Torr. Bot. Cl. xxxii. 297 ( 1905) 


ee ee ee ee See, ee ee ee ee 


Pe Day he Ce 


Se eS eee eT ae 


« 


1944] Gale,—Rhynchospora, Section Eurhynchospora 163 


Herbarium, now at the National Museum, is even more con- 
fusing. The specimen on the left is R. sola, that in the center, 
accompanied by two inflorescences, is R. fascicularis and that on 
the far right, R. Wrightiana Boeckl. A single sheet, no. 3399, 
from the Herbarium of the New York Botanie Garden contains 
a specimen of R. sola, located centrally and accompanied, right 
and left, by specimens of R. fascicularis to which the label R. 
deflexa pertains. 

Although it is evident that the original distributor failed to 
distinguish between R. sola and R. fascicularis, these two species 
are not of the same series, and, once several important details 
have been observed, can be rather easily distinguished. As 
stated in the description, the leaves of R. sola end in a relatively 
abrupt blunt tip; and the slender culm bears, without exception, 
a single terminal fascicle which is subtended by inconspicuous 
bracts not exceeding the fascicle in height. The leaf-tips of R. 
fascicularis are triquetrous and attenuated; the culm usually 
bears one or more distant lateral fascicles, and the terminal 
fascicle is exceeded by a bract approximating twice the height of 
the fascicle. Occasionally young or reduced plants of R. fascicu- 
laris bear but a solitary terminal fascicle, but in all cases the 
spikelets remain divergent and ovoid in shape, not mainly 
ascending and ovate-lanceolate as those of R. sola. 

Less obvious differences between the species are supplied by 
the details of the achenes. That of R. sola is obovoid, extremely 
lenticular, with a puncticulate surface, and accompanied by 2-3 
rudimentary bristles. The achene of R. fascicularis, on the 
contrary, is subelliptie to suborbicular, markedly biconvex, 
with a smooth surface, and accompanied by six bristles varying 
in height from 14 the achene to exceeding the tubercle. 

The natural affinities of R. sola lie, not with R. fascicularis, but 
with the very rare continental species, R. solitaria Harper, 
which is known only from its type-locality in Berrien County, 
Georgia. Although larger in all its parts than R. sola, this species 
also has obtusely tipped (but wider) leaves, a slender culm, and 
a single terminal fascicle. However, the lanceolate-acuminate 
to fusiform spikelets of R. solitaria, measuring 6-7 mm. long, are 
easily distinguished from the ovate-lanceolate (4-5 mm. long) 
spikelets of R. sola. The achenes again emphasize the close 


164 Rhodora [May 


relationship existing between the two species, for they are identi- 
cal in both shape and surface-sculpturing, differing only in size, 
color, and the relative development of bristles. 

The name, R. sola, has been selected because this species is 
closely related to R. solitaria and shares its characteristic feature 
of bearing only one fascicle. 

16. R. pRacHYCHAETA (by error appearing as brachychata) C. 
Wright. Caespitose: leaves filiform, promptly involute, ascend- 
ing, smooth except for the sparingly serrulate upper margins: 
culm terete or nearly so, filiform, wiry, flexuous, smooth, 3.5-5.2 
dm. high: inflorescence of 1-3 remote fascicles, 0.61.2 em. wide, 
usually oblong in outline to approaching corymbiform: spikelets 
fusiform and sterile, or ovoid, acute and fertile, 1-flowered, 3 mm. 
long: scales acute to aristulate, drab-chestnut: bristles 1-2, 


rudimentary: achene broadly obovoid, with a pale central disc, * 


smooth or faintly cancellate, 0.9-1.1 mm. wide, 1.2 mm. long: 
tubercle subulate, 0.4 mm. long, with a broad base. PLATE 822, 
Figs. 1A and 1B; Map 20.—C. Wright in Sauvalle, Anal. Acad. 
i. Habana, viii. 85 (1871) and FI. Cub. 180 aie as “ brachy- 
hata” ; Br ritton, Trans. N. Y. Acad. xi. 90 (1892), in part. & 
pallida sensu Clarke in Urban, Symb. Ant. ii. 126 (1900), in 
part, non M. A. Curtis; sensu Kiikenthal, Fedde, Rep. Spec. 
Nov. xxiii. 209 (1926), in part, non M. A. Curtis. R. Chapmanit 
sensu Britton, Mem. Soc. Cubana Hist. Nat. ii. 194 Soe non 
M. A. Curtis. R. Blawneri Britton, Bull. Torr. Bot. 56 
(1923). Phaeocephalum brachychaetum House, Am. Sadland 
Nat. _vi. 201 (1920).—Fresh-water shallows of western Cuba, 
Dominican Republic and eastern Puerto Rico. CuBa: in occa- 
sionally flooded places in pinelands south of railroad, Herradura, 
Pinar del Rio, Ekman, no. 17737 (G); “en sabanas bajas y a 
arillas de lagunas, jurisdiccion de Pinar del Rio,’! Wright, no. 
3 (G, type; NY, US, probable isotypes). H1spANIOLA: 
hard soil, shallow water, Laguna Ahoga-los-perros, Sabana 
Guabatico, prov. Santo Domingo, Llano Costero, Dominican 
Republic, Ekman, no. 13309 (US). Purrto Rico: Sierra 
Luguillo, Blauner, no. 247 (NY, type cf R. Blawneri Britton). 
In his list of North American Rhynchospora, published in 
1892, Britton recognized R. brachychaeta C. Wright, attributing 
it to Sauvalle. However, that Britton’s conception of this 
species was at best uncertain is seen in his synonyms “R. gra- 
cillima Sauv.” and “R. fascicularis var. stenophylla Chapm. mss.” 
R. gracillima Wright in Sauvalle, I am recognizing under its 
1 Wright in Sauvalle, Fl. Cub. 181 (1873). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 165 


legitimate name, R. Wrightiana Boeckl.; R. fascicularis var. 
stenophylla was later authentically published by Chapman as 
R. stenophylla and has been maintained as such. Keeping this 
confusion of 3 species in mind, one is not surprised to find that 
Britton, in 1916, revised his earlier opinion, this time referring 
hk. brachychaeta to R. Chapmanti Curtis. 

Superficially R. brachychaeta and R. Chapmanii have much in 
common. The former is readily distinguished, however, by its 
acute, not long-aristate, spikelets, and its inflorescence consis- 
ting of 1-3 remote narrowly elliptic fascicles in contrast to the 
single terminal corymbs of R. Chapmanit. Close observation by 
means of a lens enables one to distinguish the two species on a 
basis of their achenes as well. The achene of R. brachychaeta is 
obovate, dull, with a pale central disc, with a subulate tubercle 
and with rudimentary bristles. The achene of R. Chapmanii, 
on the contrary, is subelliptic, the dise so enlarged as to cover 
the entire surface with the exception of a small dark patch at 
the base and the apex. The surface is glossy, the tubercle del- 
toid-apiculate, and the bristles are lacking. Clarke, on the other 
hand, lists R. brachychaeta in the synonymy of R. pallida M. A. 
Curtis; but Blauner no. 247 bears Clark’s annotation “ Ryncho- 
spora divergens Curtis!” 

In 1922, Britton, studying the same specimen, Blauner no. 
247, realized that it was a good species wrongly annotated by 
Clarke; but, failing to see its connection with Wright’s R. 
brachychaeta, set it up as a new species, R. Blauneri. 

The confusion of R. brachychaeta with R. pallida, initiated by 
Clarke, was furthered by Kiikenthal who, in 1926, assigned 
Wright, nos. 3782 and 3397 to the latter species. No. 3397 at 
the Gray Herbarium is a mixed sheet, containing on the left a 
specimen of R. fascicularis (Michx.) Vahl, and it suggests the 
possibility that the sheets seen by Kiikenthal are also mixed. 
However, even if this were so, it is quite improbable that the 
specimens of either sheet are R. pallida, since this species, un- 
mistakable in its larger achene, pale corymb and tuberous bases, 
has not been reported south of the Carolinas. 

I have corrected the original spelling of this name, following 
the precedent of authors since Wright. 


166 Rhodora [May 


17. R. pauura M. A. Curtis. Caespitose with short stolons; 
base bulbous, sheathed by fibrous, acute, ovate scales: leaves long, 
narrowly linear, 1-3 mm. wide, erect, flat with the exception of 
the triquetrous tip; upper margins upwardly scabrous: culms 
acutely trigonous, slender, flexuous, 4.4-9.4 dm. tall, upper 
angles sparingly serrulate: inflorescence a single turbinate to 
hemispherical fascicle, 1.4-2.6 em. wide: spikelets ovoid-atten- 
uate, 4.5-5 mm. long, sessile, 1-flowered: scales forced apart by 
the developing achene, with aristate tips 0.3-0.4 mm. long, 
silvery, pale to reddish: stamens 2: bristles obsolete or 1-3 
rudimentary stubs 0.2 mm. long: achene obovate to broadly 
elliptic in outline, strongly lenticular, the surface finely striate, 
the umbo light and the marginal regions chestnut-brown, 1.2—1.5 
mm. wide, 1.4-1.8 mm. long: tubercle compressed, ‘deltoid- 
apiculate, 0.2-0.4 mm. high. PLare 821, ries. 4A an d 4B; ce 
18.—Am. Journ. Sci. ser. 2. vii. 409 (1849), non ape Steud 
Chapman, Fl. es U.S. 527 (1860); Gray, Man. ed. 5: 568 (1867): 

: ‘ (1 ; 


£. . 
184 (1901); Small’ FI. 194 (1903) and Man. 179 (1933); Robinson 
& Fernald in Gray, Man. ed. 7: 200, fig. 322 (1908); Fernald, 
Ruopora, xlii. 378, map 14, and 381 (1940). R. Curtisii Steudel, 
Cyp. 141 (1855); Boeckeler, Linnaea, xxxvii. 564 (1873); non 
Britton ex “yee Phaeocephalum pallidum House, Am. Midlan 
Nat. vi. 202 (1920)—Open bogs and wet depressions, often in 
pineland, Long Island, New York, south through North Carolina. 
New York: pine-barren swamp, Central Islip, ee Co. 
Ferguson, no. 515 (G, NY). New Jersey: near P. R R. 1% 
Sure northwest of Allaire, Monmouth Co., Van Pelt & Brown, no. 
4 (P); Lawrence Station, Mercer Co. , Aug. 6, 1885, Peters (P); 
poe ib in dense tufts, Tom’s River, ” Ocean Co., Parker (G); 


Long, no. 4835 (Py; boggy meadow, Swain, Cape May Co., Aug. 
8, 1925, Stone (G). DELAWARE: pine-barren bogs near Laurel, 
Sussex Co., Aug. 19, 1880, Commons (NY, P). MARYLAND: 
swamps, Eastern Shore, near Salisbury, Wicomico Co., Sept., 
Canby (US). Virernta: sphagnous savanna-like swale "east of 
herry Grove, south of South Quay, Nansemond Co., Fernald 
& Long, no. 10550 (CU, G, P, NY). Norra CaroLina: wet 
depression, pineland, beside railroad, 1 mile east of Bailey, 
Nash Co., Oosting, no. 1677 (CU); damp or peaty sandy soil, 3 
miles west of Sims, Wilson Co., Wiegand & Manning, no. 
(G); savanna 8 miles Kidthwcat of Washington, Beaufort Co., 


1944] Gale,—Rhynchospora, Section Eurhynchospora 167 


Godfrey, no. 4393 (G); wet soil, open pinelands, Newport, Car- 
teret tev} Randolph & Randolph, no. 925 (G); moist sandy soil, 


Aug. 1879, "Hyams (ug), low cera near Wilmington, Nee 
Hanover Co., ere Herb., no. 4472 (NC). 

18. a, sp. nov. Planta caespitosa: foliis filiformibus vel 
1.5 mm. - latis, planis, saepissime laevibus, laxe ascendentibus: 
culmis subtriquetris gracilibus ascenden tibus, apicem versus 
flexilibus vel laxis, 3—4.3 dm. altis: fasciculo solitario, compacto 
subhemisphaerico vel turbinato, 0.5-1 em. lato: bracteis setaceis 
quam fasciculo paullo longioribus: spiculis fertilibus lanceo- 
ovoideis 4 mm. longis: spiculis sterilibus fusiformibus et numero- 
sis, confertis ascendentibus pn ba squamis dense imbricatis, 

allidis; mucrone promi inente 0.4 mm. longo: setis nullis vel 
a 1: ‘ia antrorse polar achaenio obovoideo 
lenticulari leviter tiGhaedat 1.2-1.3 mm. lato 1.5 mm. longo, 
laevi pallido opaco: tuberculo ere arts sea eg apiculato 
0.4 mm. alto. Puare 821, rics. 1A a AP 23.—Moist 
sand, Isle of Pines. CuBA: moist ar eri ao eine of Los 
Indios, Isle of Pines, Feb. 13, 1916, Britton, Britton & Wilson, no. 
_ 15809 (N Y, TYPE) and no. 14177 (NY)3 i in a wet palm grove of 

Sess Wrightii, between Nueva Gerona and McKinley, 
Isle of Pines, Feb. 23, 1939, Leén, Victorin & Carabia, no. 18770 
(CU); sandy savanna between Nueva Gerona and McKinley, 
Isle of Pines, Feb. 23, 1939, Leén, Victorin & Carabia, no. 
18757 (CU, in part). 

This West Indian species is known to me only from three 
specimens collected on the Isle of Pines. It is similar to R. 
Chapmanii M. A. Curtis both in the slender habit and details of 
the inflorescence—the numerous fusiform sterile spikelets, 
strongly awned scales, ovoid, lenticular, bristleless achenes. It 
differs from the latter in having stiffly erect, not circinate 
bracts and larger fertile spikelets which are not forced open by 
the growing achene and which are few in number so that the 
resultant fascicles are smaller and less corymbiform than those of 
R. Chapmanii. 

R. nuda also differs from R. Chapmanii in details of the 
achene. That of the latter measures 0.8-1 mm. wide, 1-1.2 mm. 
long, is typically lustrous, pale, with dark brown patch at the 
base and apex. The achene of the Cuban species, on the other 
hand, is 1.2-1.3 mm. wide, 1.5 mm. long, with a uniformly pale 
dull surface. 


168 Rhodora [May 


The only close relative of R. nuda on the islands is R. brachy- 
chaeta Wright. However, the terminal fascicles of the latter, 
although similarly characterized by sterile fusiform spikelets, 
are occasionally accompanied by 1 or 2 smaller lateral fascicles. 
Moreover, the scales of R. brachychaeta are acute, or, at the most, 
aristulate, in contrast with the strongly aristate scales of R. 
nuda; and the achenes of the former are smaller (0.9-1.1 mm. 
wide, 1.2 mm. long), dark brown relieved by a large pale dise and 
usually accompanied by 1—2 rudimentary bristles. 

The specific name has been chosen with reference to the almost 
complete failure of the bristles, an uncommon condition in the 
Section Hurhynchospora. 


19. R. Cuapmanit M. A, Curtis. Densely caespitose: leaves 
capillary to 1 mm. wide, flat, becoming involute on drying; upper 
margins finely serrulate: culms obtusely trigonous, slender, wiry, 
erect, smooth, 3-5.1 dm. high: inflorescence a single terminal 
corymbiform fascicle, 0.5-1.7 em. wide, closely compacted, less 
often slightly exceeded by a smaller secondary fascicle: bracts 
several, filiform, exceeding the fascicles, erect or slightly circin- 
ate: fertile spikelets slenderly ovoid-aristate, mostly ascending, 
closely approximate, 1-flowered, split apart by the maturing . 
achene, 2.5-3 mm. long; sterile spikelets fusiform: scales with 
aristate tips 0.4-0.6 mm. long, pale chestnut-brown: stamens 1-2: 
bristles none: achene subelliptic in outline, strongly lenticular, 
0.8-1 mm. wide, 1—-1.2 mm. long, pale except for the dark brown 
patch at base and apex; surface obscurely rugulose to smooth, 
glossy: tubercle deltoid-apiculate with a broad base, compressed, 
0.2-0.3 mm. high. Puars 821, rigs. 3A and 3B; Map 19.—Am. 
Journ. Sci. ser. 2. vii. 409 (1849); Chapman, Fl. So. U. 8S. 528 
(1860); Small, Fl. 194 (1903) and Man. 179 (1933). R. conferta 
Chapman ex M. A. Curtis, Am. Journ. Sci. ser. 2. vii. 409 (1849), 
in syn. of R. Chapmanii. R. Grayana Chapman ex M. A. Curtis, 
Am. Journ. Sci. ser. 2. vii. 409 (1849), pub. in syn. of R. Chap- 
manit. Phaeocephalum Chapmanii House, Am. Midland Nat. 
vi. 201 (1920).—Low, sandy pineland of the Coastal Plain of 
North Carolina to Florida and west to eastern Louisiana. 
NortH Caro.ina: wet sandy soil, waste ground, Beaufort, 
Carteret Co., Randolph & Randolph, no. 795 (G); savanna, 8 


m 
quest, no. 5617 (D); pineland at Delway, Sampson Co., Godfrey, 
no. 6170 (D, G); low pineland at Dunn, Harnett Co., Godfrey, 


i il 


1944] Gale,—Rhynchospora, Section Eurhynchospora 169 


no. 6139 (D, G). Sourn Caro.ina: grass-sedge bog or savanna, 
18 miles north of Georgetown, Georgetown Co., Godfrey 
Tryon, no. 1609 (D, G, NY); damp ome levee, Santee Canal, 
Berkeley Co., Ravenel, no. 25 (G); grass-sedge bog or savanna 
14 miles south of Monks Corner, Berkeley Co., Godfrey & 
Tryon, no. 1432 (G, NY); pineland pool, 5 miles south of 
Hardeeville, Beaufort Co. , Lyles, no. 4378 (CU). Geroreta: 
moist pine barren near Monteith, Chatham Co., Kyles, no. 6455 
SUE sandy borders of pine-barren stream, F ‘itzgerald, Ben Hill 
is Harper, De 1420 (G, NY, US); moist. pine barrens, Sweet- 
nite Creek, Clinch Co., Eyles, no. 244 (D). FLORIDA: moist 
pine barrens near Jacksonville, Duval Co., Curtiss, no. 5015 
(G, NY, US); Toeai, St. John s Co., Palmer, no. 605 (G); Tampa, 
Hillsborough Co., Oct., 1877, Garber (G, P): prairie, 18 miles east 
of Okeechobee City, St. Lucie Co., Small et al., no. 9300 (NY 
edge of dried-up pool in low pineland, vicinity of Eustis, fake 
Co., Nash, no. 1396 (NY); turfy, boggy, sandy meadow, 7 miles 
Ww est of Sneads, Jackson Co., Wiegand & Manning, no. 589 (G); 
sloping moist pine barrens about 5 miles south of Bristol, Liberty 
Co., Harper, no. 47 (G, NY, P); flat pine barrens, Apalachicola, 
Franklin Co. , Chapman in Biltmore Herb., no. 20 la (G, NY, US); 
low open places, Lynn Haven, Bay Co., Oct. 12, 1921, Billington 
(US). ALaBama: low pi ineland about Miflin Creek, Elberta, Bald- 
win Co., July 9, 1926, Wolf (StB); sandy pineland, Theodore, 
Mobile Co. yy eels no. 4446 (NY). Mississippi: Ocean Springs, 
Jackson Ca. Seymour, no. 15 (CU, D, NY, US); Biloxi, Harrison 
Co., Tracy, no. 4888 (US). LOUISIANA: open ‘pineland, 1 mile 
north of Abita Springs, St. Tammany Parish, Pennell, no. 4167 


” 


Series 4. Fuscae (Clarke), stat. nov. et emend. Represented 
in the peat bogs of the northeast by the common R. fusca, also 
infrequent species in low places and ‘pond-margins of the Coastal 
Plain and Cuba. Habit solitary to caespitose: leaves filiform to 

mm. wide: culms filiform to slender: inflorescence 2-4 turbinate 
to hemispherical fascicles: spikelets maturing several achenes: 
scales loosely imbricate, castaneous to fuscous: bristles upwardly 
serrulate, well developed: achenes ddd slenderly elliptic, 
often strongly biconvex, smooth to glossy (minutely granular in 
R. fuscoides): tubercle triangular, compressed, thickly setose.— 
Rhynchospora Series Sg Diplostyleae Sect. 4. Fuscae Clarke in 
Urban, Symb. Ant. 105 (1900), in part. enees V. 
Glomeratae Small, Maa: 175 (1933), in small part. 


Key To Spectres IN SERIES FuscaE 


a. Fascicles 1-3, turbinate or ovoid, the eg eee few — 
strongly ascending; leaves filiform to 1.5 mm. wi ial De 
b. Slenderly sealoedtctoiens fascicles exieeded by 2 ime 
bracts; bristles naked at their bases.........--------- 20. R. fusca. 


170 Rhodora [May 


b. Caespitose; bracts shorter than or barely one the 
fascicles; bristles sparsely hairy at their bases... . 
e a peel ovoid, homogeneously een: A 
PO SA ees pee Oe ee be ee rene 21. R. pleiantha. 
R. Curtissii. 


De eee Ole Bie se Bee a he ee eee Se Oe 8 s-(8 eae) wi IF 


peeing en 4-5 in number, with sant spikelets sat 
the leaves 2 mm. wide or wider and languidly Ea edie. eels 
d. Corymbs 2, rarely 3, the terminal internode strongly 
myohind © bracts fOleCOUR 68 ss eis ote ee 23. kk. Harperi 
d. Corymbs 2-5, the sna internode erect or flexuous; 
racts setaceous.... 
6. oe borne on a pessicteht basal stipe which is 6 m 
n length and covered with a tangle of white haiti 
habit weak; leaves 2 mm. wide or wider; corymbs 4— 
24. R&R. crinipes. 
e. Achene without a conspicuous basal stipe; stipe, if, 
present, ae hairy; habit wiry, erect; leaves filiform to 
mm e; corymbs ie 
7. Surface of fF ashenas glassy, with ‘a white lustrous disc; 
Pome filiform, rarely 2 mm. wide; Coastal Plain, 
mand contrat Cuba; 2. oo. oi cess 25. R. filifolia. 
‘. Sines of achene tending to become granular or some- 
what polished, cinnamon-brown with a paler central 
disc; leaves 1-1.5 mm. wide; range limited to 
Ww a 26. 


R. fuscoides. 


20. R. rusca (L.) Ait. f. Slenderly stoloniferous: leaves 
promptly involute, filiform to 1.5 mm. wide, ascending: culms 
slender, erect, terete, 0.8-4.6 dm. high: terminal fascicle often 
compounded of 2-3 closely approximated secondary fascicles, 
turbinate or ovoid, 0.3-2 em, wide, exceeded by the 1 or 2 long 
circinate or erect bracts: lateral fascicles 1-2, on exserted 


spicuously serrul ate. PLATE 824 |. FIGS. 2A and 2B; Map 24.— 
Hort. Kew. ed. 2: i. 127 (1810); Gray, Ann. Lye. N. Y. iii. 215, pl. 
6, fig. 26 (1835); Torrey, Ann. Lye. N. Y. iii. 366 (1836); Britton 
& Brown, Ill. Fl! i. 279, fig. 656 (1896); Britton, Man. 185 (1901); 
Shaye Fl. 1321 (1903) and Man. 181 (193 33): Robinson & Fernald 
ray, Man. ed. 7: 200, fig. 319 (1908); Victorin, Fl. Laurent. 

089, fig. 248 (1935). Schoenus fuseus Linnaeus, Sp. 1. ed. 2: 
1664 (1763). R. alba 6. fusca Vahl, Enum. ii. 236 (1806). 
Phacomephales fuscum House, Am. Midland Nat. vi. 202 (1920). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 171 


—Peat bogs and sandy or peaty pond-shores, Newfoundland, 
eastern New Brunswick, Nova Scotia and southern portions of 
Maine, New Hampshire and Vermont; general over the other 
New England states; southward along the coast to Maryland; 
inland in central New York State with perm: stations on the 
shores of Lakes Superior, Michigan and Huron; also in Europe. 
The characters of R. fusca are sufficiently stne to make the 
citation of representative specimens unnecess 
21. R. pleiantha (Kiik.), stat. nov. ice ae. leaves filiform 
to 1 mm. wide, flat, setaceous, ascending: culms obtusely trigo- 
nous, filiform to slender, erect, exceeding the radical leaves, 2.5-4 
m. tall: terminal fascicle corymbiform, composed of not more 
than 15 spikelets; lateral fascicles 1- 2, smaller, on exserted erect 
peduncles: spikelets oblong-ovate, sessile, erect to spreading, 
approximately 6-flowered, 2-fruited, 6 mm. in length: fertile scales 
lanceolate-aristate, loosely imbricate, greatly exceeding the 
achene in length: bristles 6, variable, the tallest barely exceeding 
to twice the height of the tubercle, upwardly serrulate, with a 
few short white hairs at their bases: achene id lenticular, — 
convex, with depressed margins, 0.7-0.8 mm. , 1-1. 
long; surface smooth, dark shining brown at aneraiay Sith an 
indefinite paler disc: tubercle triangular, attenuate, flat, pale; 
margins sparingly serrulate ee smooth, 0.6—0.8 mm. long. PLATE 
823, Frias. 4A and 4B; Map 25.—R. filifolia Torrey var. pleiantha 
Kikenthal, Fedde Rep. Nov. xxiii. 208 (1926). R. fusca 
sensu Harper, Bull. Torr. Bot. Cl. xxx. 324 (1903), non (L.) Ait. 
f.—Infrequent, shores of ponds and lakes in southeastern North 
Carolina, southwestern Georgia, central Florida, and western 
Cuba. Nort H CAROLINA: shore of Silver Lake near Wilmington, 
New Mokouer Co., July 5, a) Godfrey, no. 4846 (G); shallow 
pond near Carolina Beach, New Hanover Co., July 2, 1938, 


31, 1894, Nash, no. 1321 (G, NY, 'P, US}. CuBA: on moist 
white sand on shore of the laguna, Laguna Santa Maria, Pinar 
del Rio, Aug. 22, 1923, Ekman, no. 17242a (NY, US, immature. 
This number cited by ‘Kiikenthal in original deseription of var. 
pleiantha) and no. 17242b (G. This number cited by Kiken- 
thal in original description of var. pleiantha.). 

Specimens of R. pleiantha have generally been identified as 
either R. Curtissit Britt. ex Small or R. fusca (L.) Ait. f. The 
confusion with R. Curtissii obviously results from a similarity 
in habit, but the most superficial examination of the achenes 
enables one to separate these species. Those of R. pleiantha are 


172 Rhodora [May 


pyriform, dark brown, not exceeding 1 mm. in length; those of 
R. Curtissii, on the other hand, are narrowly ellipsoidal, 1.3-1.5 
mm. long and light brown, with a pale conspicuous oval disc. 

The confusion of R. pleiantha with R. fusca, however, is due 
to a likeness which extends to details, not only of the habit and 
spikelet, but to the achene as well. Nevertheless, specimens of 
R. plecantha can rather easily be separated from those of R. 
fusca, for the fascicles of the latter are exceeded by a long seta- 
ceous bract which is often circinate at its tip; whereas those of 
the southern species are subtended by a short stiff bract which 
is at the most only twice the height of its facsicle. 

The specific distinction of the one from the other species rests, 
however, on the more technical differences of the achene. That 
of R. pleiantha is 0.7-0.8 mm. wide and 1 mm. long; in color it 
is a dark mahogany-brown, and the bristles are sparsely plumose 
at their bases. The achene of R. fusca, by contrast, is larger, 
1-1.1 mm. wide and 1.2-1.3 mm. long, consistently light brown 
in color, with the bases of the bristles naked. 


23. R. Curtissu Britt. ex Small. Caespitose: leaves filiform 
to 1 mm. wide, wiry, involute with the exception of the flattened 
tip 0.6-1.4 mm. long: culms filiform, loosely ascending, 1.5—3 dm. 
high: fascicles 1-2, turbinate, 4-8 mm. wide, composed of less 
than 10 spikelets; lateral fascicle borne on an included peduncle: 
spikelets ovoid-elliptic to fusiform, similar to those of R. fusca, 
2-3-flowered, 2-3-fruited, strongly ascending, 4-6 mm. long: 
scales mucronulate, ovate-oblong, rather loose, castaneous: 
bristles 6, erect, delicate, tenuous, scarcely equalling to well 
exceeding the tubercle, antrorsely hispidulous, sparingly plumose 
at the base: achene narrowly ellipsoid, lenticular, smooth, often 
lustrous, light brown, with a pale elliptic disc, marginate, 0.6-0.7 
mm. wide, 1.3-1.5 mm. long: tubercle deltoid, sometimes subu- 
late, compressed, with conspicuous upward serrulations, 0.8-1.2 
mm. long. PLATE 823, rigs. 3A and 3B, Map 26.—FI. 195, 1327 
(1903) and Man. 181 (1933). R. fusca sensu Fernald, Bot. Gaz. 
xxlv. 433 (1897), non (L.) Ait. f. Phaeocephalum Curtissii House, 
Am. Midland Nat. vi. 201 (1920).—Low places, coastal counties 
of Northwestern Florida, Alabama, and Mississippi. FLORIDA: 
moist roadside, Milton, Santa Rosa Co., Curtiss, no. 5929 (NY, 
TyPE; G, NC, US, tsorypss); moist pine barrens about 114 miles 
northeast of Milton, Santa Rosa Co., Harper, no. 46 (G, NY, US). 
ALABAMA: swamp, Elberta, Baldwin Co., Aug. 21, 1926, Wolf 
(StB, in part R. filifolia Gray in Torr.). Murssisstppr: Ocean 
Springs, Jackson Co., Tracy, no. 4891 (G, NC, NY, US). 


i a 


1944] Gale,—Rhynchospora, Section Eurhynchospora 173 


23. R. Harpreri Small. Solitary: radical leaves filiform, few, 
short; cauline leaves 1-1.5 mm. wide, long-attenuate, erect, 
canaliculate, smooth: culm obtusely trigonous, slender, 6—7.3 dm. 
tall; the terminal internode strongly arched: fascicles 1-2, rarely 
3, corymbiform, 1.1-1.5 cm. wide, separated by the arching 
internode; occasionally a third fascicle distant and smaller: 
bracts foliaceous: spikelets ovoid, loosely ee 5-flowered, 
4-fruited, 5-5.6 mm. long:. seales aristulate, margins free, cas- 
taneous: bristles 6, slender, upwardly hispidulous, falling ‘short 
of the tubercle: achene obovate to pyriform, strongly pee: 
brown, with a slightly paler disc, smooth, 1-1.1 mm. wide, 1.3- 
. long: tubercle subulate, pale, 0.6-0.9 mm. eae 

margins thickly hispidulous. PLare 823, rigs. 1A and 1 
27.—Man. 182, 1503 (1933). R&R. leptorhyncha sensu Small FI. 
195 (1903), non R. leptorhyncha C. Wright. —Infrequent on 
borders of ponds and in low places in the pine barrens of on 
oastal Plain of Georgia, Florida, and Alabama. GroRG 
edge of pond, just north of Lu dowici, Long Co., Eyles, no. 6527 
(CU); wet. pine barrens about 3 miles east of Hawkinsville, 
Pulaski Co., Harper, no. 1877 (NY, TypE; US, IsoTyPE); wet 
pine barrens, Sumter Co., Harper, no. 467 G, US). Froripa: 
Tampa, Hillsborough Co., May, 1876, Garber (NY, U US). Ata- 
BAMA: ae of pond, Elberta, Baldwin Co., July 15, 1926, 
Wolf (StB). 

Prior to 1933, R. Harperi was identified as R. leptorhyncha 
Wright, the type-specimen of R. Harperi being originally so 
determined by Britton. However, the two species can be readily 
distinguished from one another. Of the two, R. leptorhyncha 
has the much stiffer habit, an inflorescence exceeded by a tall 
upright bract, and tightly involute floral scales. &. Harperi, 
on the contrary, has the culm arched to nodding, a short bract 
subtending the fascicle, and loose floral scales. Nor is there 
any close resemblance in the achenes. Those of R. leptorhyncha 
are of the fascicularis-type—large, dark, dull, and broadly ovate. 
The achenes of R. Harperi, on the other hand, are pyriform, 
light brown, resembling, but surpassing in size, those of R. fili- 
folia Gray. 

24. R. ecrinipes, sp. nov. Planta caespitosa: foliis 2 mm. 
latis planis lineari-elongatis, debilibus, apicem versus triquetris 
et sparse serrulatis: culmis gracilibus subtriquetris, 6.9-7.4 dm 
altis, laxe ascendentibus: fasciculis 4-5 compactis 1-2 cm. latis 
turbinatis lobatisque vel corymbosis; pedunculis fasciculorum 
lateralium subexsertis: bracteis setaceis parvis ornatis: spiculis 
lanceolatis 5 mm. longis confertis, ascendentibus vel patentibus 


174 Rhodora [May 


3-floris sed quarto terminali rudimentario, 2-carpis: squamis 
lanceolatis, aristulatis, laxe imbricatis, mox caducis chartaceis 
pallide castaneis: setis 6 rigide erectis, tuberculo approximate 
aequalibus: achaenio pyriformi 1.4 mm. longo 1 mm. lato, bicon- 
vexo laevi, marginem versus leviter depresso; umbone candido, 
conspicuo; stipa tereti persistente 6 mm. longa villis longis albis 
irregularibus ascendentibus vestita: tuberculo compresso-triang- 

; m. alto, margine hispido-scabrato. PLATE 823, FIGs. 
2A and 2B; Map 28.—Coastal Alabama. ALABAMA: dry places, 
roadsides (exsiccated), Mobile, Mobile Co., June, 1868, Mohr 
(US); ditches, border of ponds, Mobile, Mobile Co., June 18, 
1868, Mohr (US, TyPE). 

This species, represented by two specimens collected by Mohr 
in the vicinity of Mobile, Alabama, is closely related to R. 
filifolia Gray. It differs from the latter in its generally larger 
habit with wider leaves, coarser culms, and in its languid habit 
and looser, more numerous and more irregular fascicles. Both 
species have, however, the pyriform achene, with its glossy sur- 
face picked out by a prominent pale disc, the six stiffly erect 
bristles and the deltoid, compressed and marginally hispid 
tubercle. The signal character which distinguishes R. crinzpes 
from R. filifolia is the unique basal stipe of the former, which is 
0.6 mm. long, remains attached to the achene, and is clothed 
with a tangle of white, ascending hairs. 


3A and 3B; Map 35.—Gray in Torrey, Ann. Lyc. N. Y. iii. 366 
(1836); Chapman, FI. So. U. S. 527 (1860); Britton, Mem. Soc. 


vi. 202 (1920).—Margins of ponds or damp pockets in pinelands 
on the Coastal Plain of New Jersey, south to Florida, and west 


1944 | Gale,—Rhynchospora, Section Eurhynchospora 175 


to eastern Texas; also in central and eastern Cuba. New Jersey: 
aes and peaty, pond-hole ca. 1 mile west of Bennett, Cape May 
Co., Long, no. 13625 (G). DrLaware: Queen Anne Road, east 
of Ellendale, Sussex Co. , Aug. 17, 1899, Commons (P). VIRGINIA: 
upper border of siliceous and argillaceous shore, Airfield Mill- 
ere southwest of Wakefield, Sussex Co., Fernald & Long, no. 
14301 (G). Nortx Carona: Mr. Curtis, in Gray’s handwriting 
(NY, TYPE); moist ee between Newport and New Bern 
Highway, no. 70, Craven Co., Blomquist, no. 11241 (D, G); 
moist sandy soil between "les ehead City and Newport, Highway 
no. 70, Carteret Co., Blomquist, no. 11238 (D, G); wet sand, 7 
miles southwest of Mai De Brunswick Co., Godfrey & 
Shunk, no. 4117 (G, NC). SourH Carona: grass-sedge on 
or savanna, 12 m iles north of Georgetown, Georgetown Co. 
Godfrey & Tryon, no. 749 (CU, D, G, NY, P); damp soils, Oct.., 
Ravenel (G). GrorGiA: margin of cy press pond, north of 
Douglas, Coffee Co., Harper, no. 1434 te US); pineland pool 
just east of the Clinch Co. line on U. 8. Route 84, Ware Co., 
Eyles, no. 6328 (CU). FLorima: moist pine barrens near Jackson- 
ville, Duval Co., Curtiss, no. 3153 (CU, G, P, US); margin of 
flatwoods pond, Welaka, Putnam Co., Laessle, no. 13 (CU); 
Indian Mound near Citrus Center, DeSoto Co., Small, no. 9918 
(NY); cypress swamp, vicinity of Ft. Myers, Lee Co., Standley, 
no. 12865 (US); hammock north of Eagle Bay near Kissimmee 
River, Okeechobee Co., Smaill, no. 9188 (G); margins of ponds in 
pine barrens, ta NY Ue Franklin Co. , Chapman in Biltmore 


Baldwin Co., Aug. | 24, 1926, Wolf (StB): i in woods, Spring Hill, 


Tracy, no. 2926 (NY). epee a saat of PGapanven, $ 
Tammany Parish, Arséne, no. 11869 AO Foote low prairies, 
vicinity of Lake Charles, Caleasieu Parish, Mackenzie, no. 443 
(Mo, NC). Texas: ponds, Hempstead, Waller as Hall, no. 
717 (Mo, US). Cusa: in lagoon near El Paynes, between 
Guane and Remates near sea level, Pinar del Rio, Killip, no. 
32373 (CU, US); Laguna Los Indios and vicinity, Pinar del Rio, 
Shafer, no. 10819 peer Herradura, Pinar del Rio, Baker & 
Abarca, no. 4195 (NY); s euler ivigeis, Isle of Pines, 
Britton, Britton & Sb itei no. 15018 (NY); in wet sand, ai of 
Laguna Yaiti, Mordazo, ‘Santa Clara, Ekman, no. 170 G); 
near lagoon, Asiento Viejo, Sabana de Manacas, Souk Hones 
Leoén, no. 9288 (N 

26. iced be Clarke. Caespitose: leaves 1-1.5 mm. wide, 
involute o n drying, stiffly erect: culms obtusely triquetrous, 
slender, ee 4.7-7.8 dm. high: fascicles 2-4, the terminal one 
turbinate to hemispherical, barely if at all exceeded by the sub- 


176 Rhodora [May 


tending bract, 0.9-2 cm. wide; peduncles of the lateral fascicles 
included or nearly so: spikelets lanceolate in outline, closely ap- 
proximate, 3—6-flowered, 1—5-fruited, 3.5-5 mm. long: scales 
aristulate, loosely imbricate, somewhat caducous: bristles 6, 
approximating the height of the tubercle, stiffly erect, upwardly 
serrulate: achene pyriform, lenticular, conspicuously marginate, 
0.7 wide, 1.2-1.38 m m. long: the surface tending to 
become granular, drab to Satelit polished cinnamon-brown, 
with a paler central disc: tubercle deltoid, subulate, compressed, 
with the margin thickly setose. PLATE 822, Figs. 5A and 5B; 
Map 29.—Clarke in Britton, Trans. N. Y. Acad. Sci. xi. 89 
(1892), in part and excl. syn. R. fascicularis, var. distans Chapm 

nomen nudum; Clarke in Urban, Symb. Ant. ii. 124 (1900), excl. 
syn. R. fascicularis, var. distans Chapm. and R. fusca b. Chapm. 

first valid publication; Britton, Mem. Soc. Cubana Hist. Nat. 
ii. 195 (1916). R&R. filifolia sensu Wright in Sauvalle, Anal. Acad. 
Ci. Habana, viii. 84 (187 e and Fl. Cub. 180 ea non eis in 


Cusa: Wright, no. 3783 (G, NY, us ISOTYPES, this number 
cited by Clarke as type of R. fuscoides and by Boeckeler as type 
of R. distans var. microcarpa); wet grassy places, pinelands, at 
12 km. of highway to La Coloma, Pinar del Rio City, Pinar del 
Rio, Ekman, no. 17807 (NY); swale in pinelands, Laguna Santa 
Maria, Pinar So Rio, Britton, Britton & Gager, no. 7137 (NY); 
border of lagoon, Laguna Santa Maria, Pinar del Rio, Britton, 
Britton & foe o. 17179 (NY, US); wet pine woods, Pinar del 
Rio, Sept., 18632! “Wright, (NY). 

hk. fuscoides is closely related to R. filifolia Gray, a species of 
the Coastal Plain which also occurs in Cuba. R. fuscoides can 
be distinguished in the field, however, by its coarser, stiff un- 
bending habit, its lack of filiform basal leaves, its culms which 
frequently bear 4 fascicles, and its long spikelets. R. filifolia has 
a delicate habit, with the culms normally flexuous, the basal, 
and often the cauline leaves as well, filiform and arching, and 
the fascicles limited to 3. In R. fuscoides the terminal fascicle is 
subtended by a short bract which may slightly exceed the fas- 
cicle; in R. filifolia, however, the bract is prominent, undulant, 
and at least twice the height of the fascicle. 

Under a lens the tiny achenes of R. filifolia, with their glassy 
surfaces picked out by the lustrous white discs, are unmistakable. 
Those of R. fuscoides are, by contrast, less pyriform in outline, 


1 See Underwood, Bull. Torr. Bot. Cl. xxxii. 297 (1905). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 177 


longer, with a dull and often granulose drab-brown surface, 
somewhat relieved by lighter discs. 

Boeckeler in 1880 was the first to recognize that the specimens 
of Wright’s no. 3783! were not R. filifolia as labeled. He set 
them off, therefore, as var. microcarpa of R. distans (Michx.) 
Vahl but added, “. . . Torrey’s Pflanze wird durch eine 
andere schmachtige Form, Rh. gracilenta A. Gray, mit der 
typischen Form der R. distans genau verbunden.”’ However, so 
utterly different in all but the most superficial details are the 
Wright specimens from R. distans and R. gracilenta that it is 
difficult to imagine wherein lay Boeckeler’s basis for the sug- 
gested relationship. 

By 1892 Clarke had evidently studied the Wright material 
and come to the decision that its status was that of a new species; 
for in that year Britton included in his list of North American 
Rhynchospora, R. fuscoides Clarke, based on Wright no. 3783. 
The description of the new species was not published, however, 
until 1900, when it appeared in Clarke’s treatment of the West 
Indian Cyperaceae for Urban’s Symbolae Antillanae ii. Clarke, 
in giving the synonymy of R. fuscoides, lists both R. distans var. 
microcarpa Boeckl. and R. fascicularis var. distans (Michx.) 
Chapm., probably taking his cue for the inclusion of the latter 
from Boeckeler’s original misalliance. 

Series 5. Fasciculares, ser. nov. Inflorescentia fasciculata vel 
cymoso-fasciculata (R. Gageri excepta) rigida: squamis castaneis 
vel furvis: setis antrorse serrulatis rudimentariis vel bene evolutis: 
achaenio late ovato vel elliptico laevi castaneo vel fusco; saepe 
disco pallido. 

Plants of usually moist areas in pineland and savannas of the 
Coastal Plain, West Indies and Central America. Habit caespi- 
tose: leaves filiform to 4 mm. wide: culms stout and erect to 
capillary, then occasionally procumbent: cymes usually fascicu- 
late, simple or compound (spiciform in R. Gageri), stiff: scales 
castaneous to blackened: bristles rudimentary to well lormed, 
antrorsely serrulate: achene broadly ovate to elliptic, smooth, 
castaneous to blackish-brown, often with a prominent pale disc, 
usually dull: tubercle triangular, compressed, often prolonged.— 
Rhynchospora V. Glomeratae Small, Man. 175 (1933), in part. 

1 One of the sheets of Wright’s no. 3783 at the New York Herbarium is mixed. 
The other species is R. leptorhyncha Wright in Sauvalle. According to C. B. Clarke 
(Urban, Symb. Ant. ij. 125 (1900) ). this is also true of a sheet of the same number 
the herbarium at Kew. 


178 Rhodora [May 


Rhynchospora Series B. Diplostyleae Sect. 4. Fuscae Clarke in 
Urban, Symb. Ant. ii. 105 (1900), in part. 


Key To SPECIES IN SERIES FASCICULARES 


a. Terminal cyme ovoid, spiciform, or consisting of 2 corymbi- 
form fascicles aligned one above the er setaceous bracts 
long; species endemic to the West Indies... .b. 

b. Terminal cyme composed of 2 se hepsisaiy corymbiform 
fascicles aligned one above the other; scales dark brown 

to blackish; achenes 1.6-1.8 mm, wide, 1,6-1.8 mm. long 

27. R. leptorhyncha. 

b. Terminal cyme spiciform to ovoid, the division into fas- 

cicles not distinct; scales castaneous; achenes 1.2-1.6 
—1 ong.... 
c, Cymes spiciform; surface of achene polished ; bristles 
ring a ba sal tuft of long white hairs; tubercles 
BLAU MARINER 6.5 7 0. oA oa Oe ey re 28. R. Gageri. 
c. Cymes red cangeatod: surface of achene mostly dull; 
od onc basally hispidulous or with a few short incon- 
us hairs; tubercle deltoid................ R. joveroensis. 
a. Terminal sheet fasciculate, corym mbi form when well devel- 
rt a setaceous bracts short, slightly if at all exceeding the 
; species continental, with a few also represented in 
the Wes t Indies... .d. 


d. Bristles 12: nenene 2-2.0 mm. long. eo. i ek. 30. R. Baldwiniit. 
d. Bristles 6; achene 1.8 mm. long or less. ... 
e. Achene minute, 0.8 mm. wide, 0.9-1 m aay ig ea 31. R. Fernaldiit 
e. Achene exceeding 1 mm. in width and 1-2 mm. in length... .f. 


lace with as prominent sink or beak-like pro- 
on ee 
g. Bristles ‘Gmaltte to falling short of the tubercles; 
tubercle broad-based, gradually tapering toward 
apex Me POUete eo 32. AR. gracilenta. 
g. gepren ately aren the achene; tubercle basally 
t abruptly contracted into a broad 
hin Chek BONS Ti ME Re Sage ee ee 33. R. Wrightiana. 
Ko, oe Tubercle deltoid to deltoid-subulate, not prolonged. . . .h. 
h. Bristles rudimen ay to exceeding the achene; 


culms erect, often coarse................ 34. R. fascicularis. 
h. Bristles rudimentary culms filiform, loosely ascend- 
ati Uo prOeimabenGs ys Se, ok eS 35. R. debilis 


_ Scales prominently aristate, dark brown, often blackened, tightly 
imbricate: stamens 4: bristles 8, upwardly hispidulous; the tips 


1944] Gale,—Rhynchospora, Section Eurhynchospora 179 


connivent around and exceeding the tubercle, plumose at their 
bases: achene elliptic to suborbicular in outline, strongly bi- 
convex, 1.6-1.8 mm. wide, 1.6-1.8 mm. long; surface smooth, 
dull brown: tubercle deltoid-subulate, broad-based, nearly 
smooth to setose, usually pale, 1.4-1.8 mm. high. PLATE 824, 
rics. 1A and 1B; Map 30.—C. Wright in Sauvvalle Anal. Acad. 
Ci. Habana, viii. 84 (1871) and FI, SCab. 180 (1 873); Clarke in 
Urban, Symb. Ant. ii. 124 (1900); Britton, Mem. Soc. Cubana 
Hist. Nat. ii. 195 (1916); Kiikenthal, Fedde Rep. Spec. Nov. 
xxiii. 209 (1926); usually as “leptorrhyncha. ”  R. leptorhyncha 
var. laevirostris Kiikenthal, Fedde Rep. Spec. Nov. xxiii. 209 
(1926). R. gracilenta sensu Clarke in Urban, Symb. Ant. ii. 126 
(1900), non Gray. R. tetrandra C. Wright msc. ex Clarke in 
Urban, Symb. Ant. ii. 127 (1900), pub. in syn. of R. gracilenta 
sensu Clarke, non Gray. Phaeocephalum leptorhynchum House, 
Am. Midland Nat. vi. 202 (1920).—Near or in shallow water, 
western Cuba and the Isle of Pines. Cusa: Wright, no. 3784 (G, 
TYPE; NY, US, probable 1sorypsEs)!; Wright, no. 3787 (G. This 
sheet of the number listed by Wright for R. odorata is R. lepto- 
rhyncha); in ponds, pinales, jurisdiccion Pinar del Rio, Oct., 
62 } 


and Coloma at 11 geese eae no. “18252 NY, US. 
Cited by Kiikenthal with other Ekman numbers in ty pe-descrip- 
tion of R. leptorhyncha var. pace lave in pebbly pinelands, La 
Siguanea, Isla de Pinos, Ekman, no. 12187 (NY). 

Specimens in the Wright oi without number but 
labeled R. tetrandra are undoubted R. leptorhyncha. The fact 
that Wright never published a species tetrandra suggests that 
he also realized this fact. 

C. B. Clarke in Urban’s Symbolae Antillanae, ii. page 127, 
erroneously refers R. tetrandra to R. gracilenta Gray. On a basis 
of this misidentification, Britton‘, in his publication on Cuban 
Rhynchospora provisionally lists R. gracilenta, but states that 
the Wright material of R. tetrandra in the Herbarium of the New 
York Botanic Garden is R. leptorhyncha. 

I am not keeping up Kiikenthal’s var. laevirostris. As the 
name indicates this variety is based on specimens, the achenes 

1 No. 3784 is also listed by Wright under R. gracilis. There is one sheet of this no 

? See Underwood, Bull. Torr. Bot. ak "297 (1 (1905). 

* Also one Gray Herbarium sheet, C. Wright, no. 3787, labeled R. odorata is R. 


gente A 
m. Soc. Cubana Hist. Nat. ii. 196 (1916). 


180 Rhodora [May 


of which have a smooth, rather than setose, tubercle. However, 
even in the specimens of sheets cited by Kiikenthal, I find this 
character to be inconstant; for individual achenes from the 
same inflorescence vary considerably, some having nearly smooth, 
others rather densely setose tubercles. 


28. R. Gacerti Britt. Densely caespitose with thick, often 
fibrous bases: leaves setaceous, canaliculate, firm, arched-ascend- 
ing, 1.8-3.3 dm. high: culms filiform, terete, ascending or some- 
what arched, varying within a tuft from 0.5-2.2 dm. in height: 
terminal cyme fasciculate, compact, ovoid, small (0.9-1.2 mm. 
long, 0.6—-1 cm. wide) rarely containing more than 15 spikelets, 
exceeded by a setaceous, recurving bract 0.4—1.3 dm. long which 
has the appearance of continuation of the culm: spikelets ovoid, 
crowded, 1-flowered with the achene terminating the axis, sessile, 
mostly ascending, closely approximate, 3-4 mm. long: scales 
aristulate, tightly imbricate, castaneous: stamens 3-4: bristles 
6-8, delicate, upwardly serrulate, connivent around the tubercle 
which they fall short of to slightly exceed, bearing at their bases 
a few silky hairs which are 0.6 mm. in length: achene broadly 
ovoid, strongly biconvex, not umbonate, surface evenly browned, 
smooth, occasionally lustrous, 1.3-1.6 mm. wide, 1.4—-1.7 mm. 
long: tubercle subulate-attenuate, compressed, smooth or rarely 
slightly setulose at the base, whitish, 0.9-1.1 mm. long. PLATE 
824, ries. 3A and 3B; Map 31.—Mem. Soc. Cubana Hist. Nat. 
li. 196 (1916). R. longifrons var. @. reducta Kiikenthal, Fedde 

ep. Spec. Nov. xxiii. 209 (1926).—-Moist savannas, Isle of 
Pines; western and central Cuba. Cusa: in white sand of 
Sabana de los Indios, Isle of Pines, Leén, no. 17501 (G); Hacienda 
San Julian, south of Mendoza, Pinar del Rio, Leén & Roca, no. 
6955 (NY); in moist places, Mateo Sanchez, Pinar del Rio City, 
Pinar del Rio, Ekman, no. 17939 (G, US. This number cited by 
Kikenthal as R. longifrons Kuk. var. reducta); Colpothrinax 
savanna, vicinity of Herradura, Pinar del Rio, Britton, Britton, 
Earle & Gager, no. 6618 (NY, TypE; US, 1soryprr); savannas at 
La Ciega, Coabilla, Camaguey, Acufa, no. 4396 (NY). 

29. R. Joverornsis Britt. Densely caespitose with stiff 
quill-like bases: leaves 1-2 mm. wide, 3.9-6 dm. long, canalicu- 
late, stiffly ascending, with arched filiform tips; upper margins 
setiferous: culms terete, rigidly erect, 2.6-4.5 dm. tall: cymes 
congested-spiciform; the terminal one 0.5-1 em. wide, 2.5-4 em. 
long, exceeded by a setaceous recurving bract 1.8-2.5 dm. long; 
the lateral smaller, on an included peduncle: spikelets ovoid, 
3.5-4 mm. long, extremely compact, sessile, 1-flowered, with the 
achene terminating the axis: scales aristulate, tightly involute, 
castaneous: stamens 3-4: bristles 6-8, delicate, upwardly serru- 


1944] Gale,—Rhynchospora, Section Eurhynchospora 181 


late, connivent at their tips, approximating the tubercle in 
length: bases without prominent hairs: achene obovoid, lenticu- 


wide, 1.4-1.5 mm. long: tubercle deltoid-compressed, with 
heavily ta margins, 0.6—-0.8 mm. long. PLarer 824, Frias. 4A 
and 4B; Pp 32.—Mem. Soc. Cubana Hist. Nat. ii. 195 (1916). 
R. reieme ‘Kakenthal, Fedde Rep. Spec. Nov. xxiii. 209 (1926). 
—Lake margins, western Pinar del Rio, Cuba. CuBA: wet sand, 
Laguna Jovero to Laguna del Bufeo, Pinar del Rio, Shafer, no. 
10992 (NY, TtypE; G, US, isorypEs); Hacienda San Julian, south 
of Guane, Pinar del ‘Rio, Leon & Roca, no. 6935 (NY); shore . 
Laguna El Punto, Pinar ‘del Rio City, Pinar del Rio, Ekman, 
18263 (NY. One of several Ekman numbers cited by Kikenthal 
after type-description of R. longifrons); banks of south Lagoon, 
anta Maria, near San Luis, Pinar del Rio, Leén, no. 19627 (G). 
30. ALDWINII Gray. Forming a coarse clump: radical 
leaves commonly short, flat becoming carinate, 3-4 mm. broad, 
tips acutely triquetrous due partially to development of the keel: 
upper margins and keel finely serrulate: culms triquetrous, erect, 
becoming flexuous, 0.5—1 m. high: terminal fascicle corymbiform, 
cm. wide, rarely accompanied by a small fascicle on a short 
stiff erect branchlet: lateral fascicles smaller, simple, exsertly 
pedunculate: spikelets ovoid, 1—3-flowered, 1—2-fruited, sessile, 
bursting, 5-5.5 mm. long: scales aristate, caducous, castaneous 
to dark brown: ppatles 12, slender, typically convergent, up- 


subrotund, lenticular-compressed, emarginate, dark brown, with 
a pale disc, smooth to minutely pitted, dull, 1.8-2 mm. wide, 
2—2.5 mm. long: tubercle deltoid-compressed, whitish, 0.8-1 mm. 
long. PuaTE 825, rics. 4A and 4B; Map 33.—Ann. Lyc. 

iil. 210, pl. 6, fig. 18! (1835); Cha apman Fl. So. U. S. 526 (1860); 
Small, Fl, 196 (1903) and Man. 182 (1933). Phaeocephalum 
Baldwinii House, Am. Midland Nat. vi. 201 (1920).—Peaty 
savannas and low pine barrens of the Coastal Plain from North 
Carolina southward through the peninsula of Florida and west 
to Mississippi. NorrH CaroLina: savanna near Jacksonville, 
Onslow Co. Wadia. ~ 5822 (G); savanna near Burgaw, Pender 
Co., Godfrey, no. 4737 (G, NC); long-leaf pine and wire grass 
savanna, Carolina Beach, ‘New Hanover Co., Godfrey, no. 4685 
(G, NC). SourH CAROLINA: peaty excavated area in savanna at 
side of road, 12 miles north of Georgetown, Georgetown Co., 
Godfrey & Tryon, no. 8 (G, NY); in damp stiff soils, Santee 
Canal, "hosel (G). Geroreta: rather dry pine barrens between 


1 Figures 19 and 18 were apparently reversed in the printing; R. Baldwinii is fig. 18, 
not 19 as listed. 


182 Rhodora [May 


Guyton and Springfield, Effingham Co., Harper, no. 934 (G, NY, 

S); sandy bog, Bullock Co., Harper, no. 852 (NY); moist pine 
barren 2 miles west of Glennville, Tatnall Co., Eyles, no. 6435 
(CU); Dr. Baldwin, fragment from Herb. Schweinitz (NY, 
TYPE). FLORIDA: moist pine lees near Jacksonville, Duval 
Co., Curtiss, no. 4868 (G, NY, US); Hastings, St. Johns Co., 
Tracy, no. 9286 (iS, NY) US): Tampa, Hillsborough Co., May, 
1876, Garber (US); prairie, ve miles east of Okeechobee City, St. 

ucie Co., Small et al., no. 3 (NY); wet places, Bear Creek, 

Gadsden Co., June 1841, Hse (G, in part) ; wet pine barrens, 
Apalachicola, Franklin Co., Chapman in Biltmore H erb., 
256a (G, NY, US); pine barrens 8 miles west of Apalashiaets 
Gulf Co., Eyles, no. 5783 (CU); swamps, Walton Co., 1885, 
rou (NY ). AtaBaMa: low pine barrens, Mobile, Mobile Co., 

ne, 1870, Mohr (NY); Sartwell (G).  Mussissrprr1: Biloxi, 
Fickich Co., Tracy, no. 4894 (NC, NY, US). Lovrstana: New 
Orleans, Drummond (G) 

31. R. Fernaldii, sp. nov. Planta caespitosa: foliis basilaribus 
1—1.5 mm. latis planis marginibus laevibus vel subtiliter serru- 
latis, apicem versus triquetris: culmis teretibus tenuibus rigide 
erectis 2.5-5.2 dm. longis: fasciculis 1-2 congestis corymbiformi- 
bus; lateralibus one pedunculatis; pedunculis erectis: spiculis 
ovoideis 2-2.5 m m. longis sessilibus confertis erectis vel patenti- 
bus 3—4-floris 2-3-carpis: squamis aristulatis laxe imbricatis 
fuscis: setis 6 antrorse hispidulis rigide erectis, achaenio fere 

ry eepeatres achaenio lenticulari-obovoideo, biconvexo, parvo 
mm. lato 0.9-1 mm. longo) laevi furvo; disco leviter pallid- 
iore: tuberculo compresso-deltoideo, 0.2—0. 3 mm. alto. PLATE 
825, Frias. 3A and 3B; Map 34.—Pine barrens of southern Georgia, 
northern Florida and coastal Alabama. GroraiA: rather dry 
sandy road in pine barrens near Camp Cornelia, Charlton Co., 
Aug. 8, 1902, Harper, no. 1487 (G, TypE; NY, ‘US, IsoTYPES) ; 
near Lem Griffin’s Camp, Okefenokee Swamp , Clinch Co., Oct. 
1938, Eyles, no. 164 (CU); piney woods, Billy Island, Charlton 
Co. July 7, 1912, Bradley, no. 4 (P). FLORIDA: in a low pineland, 
7 miles southwest of St. Augustine, St. Johns Co., Aug. 8, 1929, 
O'Neill (CU); Sanford, Orange Co., Aug. 23, 1899, Pieters, no. 
301 (US); in low pinelands, Alva, Lee Co., Oct. 30, 1917, Francis, 
no. 41 (US); low pine barrens near the seashore, Apalachicola, 
Franklin Co., Oct. 27, 1895, Mohr (US); pine barrens, Apalachi- 
cola, Franklin Co., July, 1870, Chapman (US); low pine barrens, 
Apalachicola, Franklin Co., Chapman in Biltmore H erb., no. 9 
(G, NY; US, in part) ; Chapman (P, no locality given), ALABAMA: 
Buckley, no. 35 (NY). 


In habit, R. Fernaldii suggests a dwarfed state of R. fascicu- 
laris var. distans. The plant is tufted, the slender culm is stiffly 


a ; 


1944] Gale,—Rhynchospora, Section Eurhynchospora 183 


erect, and the small terminal fascicle is corymbiform and fre- 
quently accompanied by a lateral fascicle on a slender peduncle. 
However, among the members of this series the spikelets and 
achenes of this species are unique in their minuteness. The 
former measure only 2—2.5 mm. in length and the tiny blackish 
achene varies from 0.9-1 mm. in length. I suspect that this 
character was in part responsible for the long neglect of this 
species, for at first glance the inflorescence does suggest an un- 
developed state of var. distans. This species has been named in 
honor of Professor M. L. FERNALD. 


32. R. GRACILENTA Gray. Caespitose: radical leaves filiform, 
about 10 mm. high, cauline leaves filiform to usually 1-2.5 mm. 
wide, flat; upper margins serrulate; tips triquetrous: culms terete, 
slender, erect to flexuous, 0.3—1 m. in height: terminal cyme a 
single fascicle, irregular in outline to corymbiform, with few 
spikelets, 0. 8-1. 2 cm. wide; or less often several fascicles on stiff 
branchlets: lateral fascicles 1-2 (rarely 3) on exserted peduncles: 
spikelets ovate, split open by the developing achenes, bearing 
2-3 florets, the lowermost of which is often abortive, 1—2-fruited, 
sessile, 3-3.5 mm. long: scales aristate, castaneous: bristles 6, 
delicate, upw atdly serrulate, falling short of to slightly exceeding 
the tubercle: achene oval to suborbicular in outline, 1.1—1.7 m 
wide, 1.3-1.8 mm. long, compressed, umbonate, ‘smooth, dull 
dark brown at maturity, often with a conspicuous light disc: 
tubercle broad-based, with a long slender strap-like terminal 
extension, strongly compressed, 1-2.6 mm. long. Pxate 826, 
rics. 6A and 6B; Mar 36.—Ann. Lye. N. Y. iii. 216, pl. 6, fig. 27 
(1835) ; Chapman, FI. "So. U8. ‘507 (1860); Britton & Brown, Il. 
Fl, i. 279, fig. 657 (1896); Britton, Man. 186 (1901); Small, Fl. 
195 (1903) and Man. 182 rere Robinson & Fernald in Gray, 
Man. ed. 7: 200, fig. a (19 08); Britton, Mem. Soc. Cubana 
Hist. Nat. ii. 196 (i9 16); Kikenthal, Fedde Rep. Spec. Nov. 
xxi. 208 (1926). R. gracilenta var. diversifolia Fernald, Ruo- 

DORA, XXXVii. — ( coe R. fusca sensu Gray, Gram. et Cyp. i. 
no. 93 (1834 ), in part.1 R. Drummondiana Boeckeler, Fl. xli. 
644 (1858). R. ieichomhe Fernald, Ruopora, xxxix. 389 (1937). 
Phaeocephalum gracilentum House, Am. Midland Nat. vi. 202 
(1920).—Bogs and moist areas in swales, common along the 
Coastal Plain from New Jersey to southeastern Virginia; ap- 
parently more scattered southward in the Carolinas Hed Georgia; 


0. 93 was apparently made up from a mixed collection. In the volume belonging 
to the. library of the New York Botanic Garden, no. 93 is R. gracilenta, as sta 
Gray in an appended correction; but in the volume seem, to the Gray Herbarium, 
no. 93 is R. fusca (L.) Ait. f., as stated on the original I 


184 Rhodora [May 


westward along the coast to eastern Louisiana; infrequent inland 
stations in the mountains of Virginia, North Carolina and 
Tennessee, and in central Arkansas and eastern Texas. The 
citation of the bulk of specimens from New Jersey and states 
south to Virginia has been omitted. Nrw Jersey: ‘pine barrens 
of N. J., Aug.—Sept.”’ in Gray’s handwriting (NY, Type). Vir- 
GINIA: cranberry-meadow in wet spots, Stuart’s Draft, Augusta 
Co., Carr, no. 303 (G). Norra Caro.ina: wet grassy pineland, 
12 miles west of Swan Quarter, Hyde Co., Oosting, no. 22 : 
low pineland at Dunn, Harnett Co., Godfrey, no. 6140 (D, G); 
sphagnum swamp, vicinity of Black Mt., Buncombe Co., 
Standley & Bollman, no. 10464 (US); dry sandy soil near Supply, 
Brunswick Co., Blomquist, no. 11266 (D, G). SourH CAROLINA: 
grass-sedge bog or savanna, 12 miles north of Georgetown, 
Georgetown Co., Godfrey & Tryon, no. 60 (D, G, NY); grass- 
sedge bog or savanna, 1 mile west of Chicora, Berkeley Co., 
July 24, 1939, Godfrey & Tryon (G); Sumter, Sumter Co., 

. 3740 (P); Aiken, Aiken Co., July—Aug., 1866, 
Ravenel (G). Grorata: pine barren, Pooler Road near Ogeechee 
Canal and Little Ogeechee River, Chatham Co., Eyles, no. 6239 
(CU); moist pine barrens, Rushing Pond, near Statesboro, 
Bullock Co., Eyles, no. 6185 (CU); moist meadow, Leslie, Sumter 
Co., Harper, no. 414 (G, NY, US). FiLoria: sphagnous swamps 
near Jacksonville, Duval Co., Curtiss, no. 5072 (CA, NC, NY, 
US); Marianna, Jackson Co., Tracy, no. 4887 (US); wet pine 
barrens, Apalachicola, Franklin Co., Chapman in Biltmore Herb., 
no. 4478a (G, NY, US). Atapama: Lee Co., June 29, 1897, 
Earle & Baker (NY); in a wet place, Perdue, Coffee Co., Blanton, 
no. 84 (G, US). TENNESSEE: permanently moist meadow with 
ox-bows of Abrams Creek, west end of Cades Cove, Blount Co., 
Great Smoky National Park, alt. 1700 ft., Camp, no. 1993 (US); 
grassy borders of small pond south of Altamont, Grundy Co., 
Svenson, no. 9181 (G); bogs between Tracy City and Coalmont, 
Grundy Co., Svenson, no. 9568 (G). Muisstssrpp1: Ocean Springs, 
Jackson Co., Pollard, no. 1005 (G, NY, US); Biloxi, Harrison Co., 
Tracy, no. 7003 (G, NY); Bay of St. Louis, Hancock Co., Sept. 
13, 1883, Langlois (G, NY). ArKANnsas: sandy bogs, Malvern, 
Hot Spring Co., Palmer, no. 8095 (CA, Mo, P, US). Louisiana: 
vicinity of Covington, St. Tammany Parish, Arséne, no. 11735 
(Mo); dampish soil, 2 miles west of Hammond, Tangipahoa 
Parish, Trotter & Chilton, no. 153 (La); New Orleans, Orleans 
Parish, Drummond, no. 397 (G, 1soryrE of R. Drummondiana 
Boeckeler). Trxas: swamps, Swan, Smith Co., Reverchon, no. 
2921 (Mo) and no. 2918 (Mo); ponds, Hempstead, Waller Co., 
Hall, no. 717 (G). 


A study of specimens of R. gracilenta selected to represent its 


—— 


1944] Gale,—Rhynchospora, Section Eurhynchospora 185 


entire range tends to minimize the significance of var. diversifolia 
Fernald. This variety was defined on a basis of its “stiffer habit, 
broadish cauline leaves, and longer spikelets, fruits and tuber- 
cles.”’ However, a stiffer habit and broader cauline leaves are not 
always accompanied by spikelets with larger achenes, and vice 
versa; nor would it be practical to re-define var. diversifolia 
solely upon the size of its achene, for the common occurrence 
of achenes of intermediate sizes would make the choice of defini- 
tive measurements purely arbitrary. 

R. trichophylla Fernald was based on a peculiar colony in 
Sussex Co., Virginia. Identical in habit with the more robust 
specimens of FR. gracilenta, this material bears achenes dis- 
tinguished by bristles which barely equal the slightly prolonged 
deltoid tubercles. Typical achenes of R. gracilenta, on the other 
hand, have long bristles often exceeding the long strap-like 
tubercle. However, a close study of the material of R. gracilenta 
reveals that short bristles as well as the deltoid-attenuate 
tubercle of R. trichophylla fall within the range of achenial 
variability of the former species. 


33. R. WricgutiaAna Boeckl. Densely tufted: leaves usually 
filiform to rarely 1.3 mm. wide, ascending to loosely spreading: 
culms filiform or very slender, wiry, stiffly erect to weakly ascend- 
ing, leafy, 1-4.6 dm. high: fascicles 1-2, rarely 3, with 1-few 
spikelets, not ue 1 cm. in width; lateral fascicles on subex- 
serted peduncles: bracts setaceous, several well exceeding the 
fascicle: spikelet ovoid, usually split, revealing the tubercle and 
upper portion of the achene, 1—2-flowered, with a rudimentary 
floret terminating the axis, Ma 2-fruited, sessile, 3-3.5 mm. long: 
scales mucronate, castaneous, caducous: bristles 6, variable in 
height but rarely exceeding the achene, upwardly serrulate: 
achene elliptic in outline, biconvex, 1.2- 1.3 mm. wide, 1.3—-1.5 
mm. long, smooth, dark brown; a somewhat paler dise indistinct 
or lacking: tubercle triangular-compressed, 0.6-0.8 mm. long, 
the tip prolonged as a broad, blunt beak. PLaTE 825, FIGS. 5A 
and 5B; Map 37. —Flora, lxiv. 78 (1881); Britton, Mem. Soc. 
Cubana Hist. Nat. ii. 197 (1916); Britton & Wilson, Sci. Surv. 
Porto Rico and Virgin Isl. v. 104 (1923); Small, Man: 183 (1933), 
excl. syn. R. brachychaeta Sauv. R. graci illiima C. Wright in 
Sauvalle, Anal. Acad. Ci. Habana, viii. 85 (1871) and Fl. Cub. 
181 (1873). R. tenuis Baldwin ex Gray, Ann. pode ee yo. 
217 (1835), publ. in syn. of R. distans var. $., non Link. R. 
distans var. 8. Gray, Ann. Lye. N. Y. iii. 217 (i835). i distans 


186 Rhodora [May 


ANA, Var. T 


yrica; 53, R, LInDENIANA, var. BAHAMENSIS; 54, R. Grayu; 55, 
R. cunrxa; 56, R. cupensis; 57, R. STENOPHYLLOIDEA; 58, R, MEGALOCARPA, 
59, R. Harveyi. 


2 


1944] Gale,—Rhynchospora, Section Eurhynchospora 187 


var. tenuis (Baldwin) Britton, Trans. N. Y. Aead. Sci. xi. 
(1892). R. brachychaeta sensu Small, Fl. bat (1903), non C. 
Wright in Sauvalle. &. pallida sensu Clarke in Urban, Symb. 
Ant. 11. 126 (1900), in part, non M. A. Curtis. R. distans (Michx.) 
Vahl var. y. gracillima (C. Wright) Kiikenthal, Fedde Rep. Spec. 
Nov. xxiii. 208 (1926) and xxxii. 77 (198 3).— Wet sand or peat 
of ditches, swamps, ponds or pockets in pineland of the Coastal 
Plain from southern North Carolina south to the Florida Penin- 
sula and westward in coastal Alabama; also ponds of Cuba and 
mountains of Puerto Rico. Norra Carouina: wet sand, Fort 
Caswell, Brunswick Co., Godfrey & Shunk, no. 4140 (G, imma- 
ture). SourH CAROLINA: peaty pocket in pine barren, 5144 
miles south of Georgetown, Georgetown Co., Godfrey & Tryon, 
no. 201 (G, NY, P); peaty pocket in pine barren, 5 miles north- 
west of McClellanville, Charleston Co., Godfrey & Tryon, no. 
1120 (G, NY); gum-pond i in pine barren, 4 miles west of Bonneau, 
Berkeley Co., Godfrey & Tryon, no. 1625 (G, NY); margin 
of barrow-pit near Savannah Rives Refuge, Jasper Co., Eyles, 
no. 6123 (CU). Groreta: drainage-ditch, Wilmington ‘Island, 
Chatham Co., Eyles, no. 4380 (D); pine barrens, Bethesda 
Church, Effingha am Co., Eyles, no. 6104 (CU); moist sandy place, 
Bullock Co., Harper, no. 897 (NY, US); margin of pine-barren 
pond about 2 miles northeast of Hawkinsville, Pulaski Co., 
Harper, no. 1376 (G, NY); sandy — along railroad near Douglas, 
Coffee Co., Harper, no. 684 (NY, US). Fuioripa: moist pine 
buries heat a ri Nees, Duval Co. Curtiss, no. 4946 (G, NY, 
US) and 3152 (CU, D!, G, NY, P, U 8); low pineland, vicinity of 
Eustis, Lake Co., Nash, no. 662 (G, NY, P, US) and no. 642 
(G, NY, US); grassy swamp, Okeechobee ‘Region, Brevard Co., 
Fredholm, no. 5750 (G, NY, immature); prairie, ft ae east of 
Okeechobee City, St. ee Co., Small et al., no. 9304 (NY); wet 
place, Hardin Co., July 2 , 1940, Schallert (G, ae nes ALA- 
BAMA: Mobile, Mobile Rae ” July 16, 1897, Mohr (CU). CuBa: 
swampy place, pinelands at km. 12 of the highway 4 La Coloma, 
Pinar del Rio City, Pinar del Rio, Ekman, no. 17824 (NY); shore 
f Laguna de Junco, Pinar del Rio City, Pinar del Rio, Ekman, 
no. 17861 (G, US); “sobre tembladeras en medio de lagunas, en la 
Vuelta de Abajo”, Wright, no. 3781 (G, rrpg; NY, US, probable 
ide Bees Puerto Rico: plants of the Luquillo Mt., Wilson, 
7 (NY); in monte Jimenes, S* de Luquillo, Sintenis, no. 
i381 (NY, US). 
This species was originally described by Wright in 1873 under 
the name of R. gracillima. Boeckeler, however, in 1881, noting 


' One sheet of this number _ Duke University is R. fascicularis (Michx.) Vahl 


~ var. distans (Michx.) Chapma: 


2C. Wright in Sauvalle, FI. Gab. 181 (1873). 


188 Rhodora [May 


an earlier application of this name by Thwaites' in 1864 to a 
species from Ceylon, renamed the West Indian species R. 
Wrightiana, in honor of its collector. Subsequently its status 
was confused by C. B. Clarke who mistakenly placed R. Wrighti- 
ana in the synonymy of the Atlantic coastal species R. pallida 
M. A. Curtis. With R. brachychaeta it formed the basis for 
Clarke’s report of R. pallida from the West Indies. The plant 
which extends northward into southeastern Virginia and which 
ey fe passed as R. Wrightiana is R. debilis. 


. R. rascrcunLaris (Michx.) Vahl. Caespitose, often coarse- 

Ms so: eae leaves 1 (rarely less) to 4 mm. wide, curling to erect, 
oe: margins and keel finely serrulate: culms subterete and slender 
stout and obtusely 3-angled, stiffly erect, becoming flexuous, 

0. 4 (rarely)—1.3 m. high: terminal fascicle simple or corymbosely 
compound, 1—5.5 cm. wide, the ultimate densely fasciculate 
clusters borne on stiff erect peduncles, lateral fascicles 1—3 or none, 
simple or less commonly compound, distant, exserted to subex- 
serted on short erect peduncles: bracts several, foliaceous to 
setaceous, exceeding the fascicles: spikelets lance-ovoid to ovoid, 
3-4.5 mm. long, erect, sessile, 2—4-flowered, with a terminal 
rudimentary floret, 1-3-fru ited: scales ovate to lanceolate, 
prominently mucronate to aeistate, caducous, 3-4 mm. long: 
bristles 5—6, rudimentary and short to well developed and exceed- 
ing the achene, stifly erect, antrorsely serrulate: achene ovate 
to orbicular or elliptic, smooth, chestnut- to blackish-brown, 
evenly biconvex to oe with . without a pale disc, ob- 
scurely marginate, 1.1-1.5 mm. wide, 1.3-1.5 mm. long: tubercle 
ese but sagas deltoid to Hci eabnilate. 0.4-0.7 mm. 
34a, Var. typica. Habit usually very robust: spikelets 
usually with a conspicuous recurved mucro: achene ovate to 
saianaare rarely elliptic, dark to blackish brown, umbonate, 
usually relieve by a prominent pale disc, 1.2-1. 5 mm. wide, 
1.4-1.5 mm. long: tubercle variable but essentially deltoid to 
deltoid-subulate. PLatE 825, Frias. 1A and 1B; Map cer 


Britton, Mem. Soe . Cubana Hist. Nat. ii. 197 (1916). Schoenus 
fascicularis Michaud , Fl. Bor.-Am. i. 37 (1803). R. ona var. 
6. fascicularis Kikenthal, Fedde Rep. Spec. Nov. xxi . 208 


1 Thwaites, Enum. Pl. Zeyl. 435 (1864). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 189 


(1926). Phaeocephalum fasciculare House, Am. Midland Nat. 
vi. 202 (1920).—Roadside-ditches, peaty. savannas, low pine 
barrens and lake-margins, Princess Anne County in southeastern 
Virginia, and common southward on the Coastal Plain from 
North Carolina to the Peninsula of Florida; less frequent in the 
coastal states west to eastern Texas; also in western Cuba, 
amaica, Hispaniola, Puerto Rico and Central America. Vir- 
GINIA: wet peaty depressions in sandy pineland, The Desert, 
Cape Henry, Princess Anne Co., eee & Long, no. 3790 
(G, P). Norra Caro.uina: New B ern, Craven Co., T. H. 
Kear rney Jr., no. 1974 ee Soi at sea-level, Carteret Co., 
Codkrey, no. 6408 (G, N Y); pineland, White Lake, Bladen 
Co., Godfrey, no. 5982 (D a): drainage-ditch at Carolina Beach, 
New Hanover Co., Godfrey, no. 4724 (G); savanna, 7 miles 
southwest of Wilmington, Brunswick Co., Godfrey. & Shunk, no. 
4113 (G, NC). SourH Caro.ina: sandy drainage-ditch, 2 miles 
west of Salters, Williamsburg Co, Go dfrey & Tryon, no. 512 (CU, 
DG, SY, P); shallow peaty pond in pine barrens, 9 ee 
north of Fea each Georgetown Co., Godfrey & Tryon, n 

758 (D, NY); drainage-ditch, 3 miles north of MeClellan. 
ville, Siicun Co., Godfrey & Tryon, no. 683 (G); grass- 
sedge bog or savanna, 3 miles southwest of Manning, Clarendon 
Co., Godfrey & Tryon, no. 940 (G, NY); grass-sedge bog or 
savanna, 1 mile west of Chicora, Berkeley Co., Godfrey & Tryon, 
no. 842 (G, NY); wet sand pockets, 5 miles north of Ridge- 
land, Jasper Co., Wiegand & Manning, no. 599 (G). CaroLina, 
uin, Michaux Herb. (G, TyYPE-PHOTO.; NY, TYPE-FRAGMENT 
from Michx. Herb.). GroroGia: moist sandy roadside in pine 
barrens, Bullock Co., Harper, no. 878 (G); Wilmington agri 
Chatham Co., Eyles, no. 4891 (CU); Wayne Co., Baldwi i 
green sphagnum swamp, 9 miles north of Darien on U.S. Rou te 
17, MacIntosh Co., Hylan, no. 6439 (CU); pine woods, Kingsland, 
Camden Co., Small & D eWinkler, no. 9682 (NY); pineland-pool 
just east of the Clinch Co. line on U. §. Route 84, Ware Co., 
Eyles, no. 6327 (CU); hammock at edge of swamp on Billy’s 
Island, Okefenokee Swamp, pene Co., Eyles, no. 635644 
CU}; m margin of pees en pond n r Downing, Coffee Co., 
Harper, no. 1440 (G, NY, US); moist ne barrens, Thomas Co., 
Harper, no. 1173 (G, U8). FLoripa: moist pine barrens near 
Jacksonville, Duval Co., Curtis, no. 4945 (G); low pineland, Lake 
City, Columbia Co., O'Neil no. 7675 (CU); Hibernia, Clay Co., 
Mar., 1869, Canby (G US); ditch near Clyatt’s Station, 
Gainesville, Alachua Ce: me 13, 1940, Arnold (CU); wet flat- 
woods, Welaka, Putnam Co., Laessle, oun 15 (CU); dry sand, 
high pineland, vicinity of Eustis, Lake Co., Nash, no. 641 (G, 
NY, US); near Sanford, Seminole Co., Aug. 1931, Rapp (NY): 
in a low pineland, Bithlo, Orange Co., O'Neill, no. 5268 (CU); 


190 Rhodora [May 


prairies near Lake Washington, coke ard Co., Small e he — 
no. 9741 (NY); Kissimmee, Osceola Co. . May 10, , Mearn 

(US); ditch, Polk Co., July 27, “ner Schallert ne in as ditch, 
Lake Jovita, Pasco Co., Britton, no. 2609 (CU); near water, near 
St. Petersburg, Pinellas Co.,: Mre.-C 4. Deam, no. 2899 (G); 
pinelands near Fellsmere, Indian River Co., Small, no, 8868 
(NY); Okeechobee Prairie north of Okeechobee City, Okeecho- 
bee Co., Small et al., no. 9238 (NY); Palma Sola, Manatee Co., 
Tracy, no. 6996 (G, NY, US); ditch, Sarasota Co., July 29, 1940, 
Schallert (G); pinelands, east of Punta Gorda, Charlotte Co., 
Small, Mosier & DeWinkler, no. 10928 (NY); around ponds, 
Myers, Lee Co., Hitchcock, no. 423 (G, NY, US): low pineland, 
5 miles south of Stuart on east coast, Martin Co., O’ Neill, no. 
5265 (CU); St. Vincent Island, Franklin Co., M cAtee, no. 1806 
(US); low pine barrens, Apalachicola, Franklin Co., "Chapman 
in Biltmore Herb., no. 4469 (G, NY, US); Lake Gentry, Santa 
Rosa Co., Howell, no. 1092 (US). Atasama: low pineland about 
Miflin Creek, Elberta, Baldwin Co., Aug. 25, 2 Wolf (StB); 
low swampy pine barrens, Mobile, Mobile C July, 1872, 
Mohr (US). Tetons Ocean Springs, ae Co., Pollard, 
no. 1016 (D, G, NY, US); Long Beach, Harrison Co., ’ Aug. 25, 
1896, Joor (Mo): Cat Island, Hancock Co. , Lloyd & Tracy, no. 
359 (NY). LOUISIANA: In pine wood clearings, Slidell, St. 
Tammany Parish, Oct. 5, 1891, Langlois (NY); near Indian 
Village, east of New Orleans, Orleans Parish, Killip, no. 13968 
(US). Trxas: Jefferson Co., Tharp, no. 3055 (U S). CupBa: in 
savannas, towards Bibijaguas, Nueva Gerona, Isla de Pinos, 
Ekman, no. 12535 (G, US); dry ground, Laguna Jovero and 
vicinity, Pinar del Rio, Shafer, no. 10752 (G, NY); sabanas, El 
Sabalo, Finca Sabanalamar, Pinar del Rio, near sea level, Killip, 
no. 32260 (CU); palm barrens west of Guane, Pinar del Rio, 


aguna Santa Maria, Pinar del Rio, Ekman, no. 17274 (US): 
sandy pine woods (damp) Pinar del Rio, Oct. 1863?! C. Wright, 
no. 3392? (NY); on edge of laguna, Herradura, Pinar del Rio, 
Ekman, no. 11585 (G); low savannas, Chirigo ota, Pinar del Rio, 
Oct. 26, 1863? Wright, no. 339922 (US); low wood at 420 m., 
north slope of Loma Pelada de Buenavista, Cayajabos, Pinar del 
Rio, Leén, no. 13561 (NY). JaMArIca: summit of Bull Head, 
Chapelton to Bull Head, Middlesex, Underwood, no. 33646 (NY). 
Hispantoxa: laterite soil at Dutreuil, Corail, Western Group, 
Massif de la Hotte, Dept. du Sud, Haiti, c. alt. 250 m., Ekman, 


1 See Underwood, Bull. Torr. Bot. Cl. xxxii. 297 (1905). 
2 The penciled number 3399 appears on 3 Wright sheets, the specimens of which 
are for the most part R. fascicularis. These are undoubtedly no part of the original 
‘Cuba or. (Wr. 3399)"’ on which Grisebach based in part his description of R. deflera. 
’ See Underwood, Bull. Torr. Bot. Cl. xxxii. 297 (1905). 


1944) Gale,—Rhynchospora, Section Eurhynchospora 191 


no. 10741 (US); in savannas, El Valle, Sabana de la Mar, prov. 
de Samana, Cordillera Central, Dominican Republic, Ekman, no. 
15667 (NY); forming colonies, in Sabana de Ponton, toward 
Cotul, Rinc6n, prov. de la Vega, ae del Cibao, Dominican 
Republic, Ekman, no. 14623 (G, US). Purrrto Rico: in sphag- 
num, western end of Laguna Tortuguero, Britton & Britton, no. 
7872 (NY, US); wet white sand, vicinity of Vega Baja, Britton, 
Britton & Brown, no. 5788 (NY); moist sandy sojl, Santurce, 
Heller & Heller, no. 583 (NY, US). 

. Var. DISTANS (Michx.) Chapm. Habit more slender 
than that of var. typica: spikelets mucronulate: achene elliptic 
in outline, gradually biconvex, not umbonate, chestnut to dark 
brown, without a well-defined central disc, 1.1-1.8 mm. wide, 
1.3-1.5 mm. long: bristles 6, always exceeding the achene: 
tubercle compressed, triangular-subulate with a narrow base. 
PLATE 825, Frias. 2A and 2B; Map 40.—FI. So. U.S. 527 (1860). 
Schoenus distans Michaux, Fl. Bor. ea 1.36 (1803). R. vcore 
Vahl, Enum. ii. 235 (1806) ; Elliott, Sk. me fer Car eee Ga. 

59 (1816): Gray, Ann. Lyc. N. Y. iii. 216, 6, fig. 28 (1 835): 
Clarke in Urban, Symb. hat. ii. 125 (1900); Fal FL. oe (1903) 
and Man. 182 (1933): Britton, Fl. Bermuda 53, fig. 81 (1918); 
Britton & Wilson, Sci. Surv. Porto Rico and Virgin Isl. v. 103 
(1923) ; Kiikenthal, Fedde Rep. Spec. Nov. xxiii. 208 (1926). 
R. dommucensis A. H. Moore, List of Pl. Coll. in Bermuda, 1905: 
6, pl. 1 and 2 (1906). Phaeocephalum distans House, Am. Mid- 
land Nat. vi. 202 (1920). Dichromena distans Macbride, Field 
Mus. Pub. Bot. xi. 5 (1931).—Moist. sandy or peaty pineland 
from southeastern Virginia southward along the Coastal Plain 
to the tip of Florida and west to Mississippi; also Bermuda. 
VirRGINIA: moist sandy and peaty pine barrens, south of Lee’s 
Mill, Isle of Wight Co., Fernald & Long, no. 12273 (G, P); wet 
peaty pine barrens, east of Cox Landing, south of ete Quay, 
Nansemond Co., Fernald & Long, no. 10551 (CU, G, NY, P) 
Nortu CAROLINA: savanna, Chocowinity, Beaufort Co. , Godfrey, 
no. 5417 (G); ee 5 miles west of Clinton, Sampson n Co., 
Godfrey, no. 4502 (D, G); moist boggy places, north side of White 

ake, Bladen Co., Blomquist, no. 10863 (D); pineland fons 


Carolina Beach, New Hanover Co. , Godfrey, no. 4708 (G, NC). 
Sourn CaRroLina : peaty pocket in pine apa 5 miles northwest 
of McClellanville, Charleston Co., Godfrey & Tryon, no. 1122 


(DG, PR, NY), Canora: Michaux G, TYPE-PHOTO; NY, 
TYPE-FRAGMENT from Michaux Herb.). Groreta: shady woods 
Leas Sry ies and Springfield, Effingham Co., Harper, no. 

6 (G, NY, US); border of pond near Middleground Road, 
ie Co. , Eyles, no. 6228 (CU); moist sandy roadside in pine 
barrens, Bullock Co., Harper, no. 878 (C, NY, US); margin of 
shallow ‘pond i in sandhills of Satilla River, Coffee Co., Harper, no. 


192 Rhodora [May 


1447 (G, NY, US); sphagnous bogs near Bugaboo Island, 
Okefenokee Swamp, Charlton Co., Harper, no. 1477 (G, N 
US); pine barren near Lem Griffin’s, Okefenokee Swamp, Clinch 
Co., Eyles, no. 6373 (CU). FLorIpA: damp shady woods near 
Jacksonville, Duval Co., Curtiss, no. 5072¢e (G, NY); low moist 
ground, Welaka, Putnam Co., Laessle, no. 20 (CU); flatwoods, 
Lake Helen, Volusia Co., Apr. 29, 1911, Hood, (G): marsh, 
Bithlo, Orange Co., O’ Neill, no. 5117 (CU); “Bayhead” i in wet 
soil, vicinity of Eustis, Lake Co., Nash, no. 802 (CU, G, NY, 
US); cypress swamp, northeast of Labelle, Lee Co., April 24, 
1921, Small & DeWinkler (NY); eerie: east of Sebring, 
DeSoto Co., Small & DeWinkler, no. 9074 (NY); ancient sand 
dunes about Lake Jackson, Leon Co., Small Mosier & De- 

inkler, no. 10874 (NY). ALABAMA: low pineland about 
Miflin Creek, Elberta, Baldwin Co., Aug. 25, 1926, Wolf (StB). 
BERMUDA: Pembroke “Marsh, Brown, Britton & Bisset, no. 2122 
(NY); Devonshire Marsh, Brown & Britton, no. 170 (NY); 
Devonshire Marsh, A. H. Moore, no. 3004 (G, type of R. dom- 
mucensis A. H. Moore; NY, 1soryPe). 

The variability of this species, both in habit and appearance 
of the achene, was undoubtedly responsible for its original 
sepsration by Michaux into two distinct species, Schoenus fas- 
cicularis and S. distans. In 1805, Vahl transferred both species 
to Rhynchospora; and later Gray, following established precedent, 
also recognized both R. fascicularis and R. distans. However, in 
1860 Chapman reduced R. distans to varietal rank under R. fas- 
cicularis, and in the third edition of his Flora dismissed it as 
“R. distans Nutt. the form with longer bristles.’ In 1926, 
Kiikenthal, although apparently unfamiliar with the Flora of 
the Southern United States, reached a conclusion similar to that 
held by Chapman. The former, however, preferred to keep R. 
distans as the species and subordinated R. fascicularis as R. 
distans var. fascicularis. Obviously this was an unfortunate 
choice since, by International Rules, if the varietal rank is to be 
maintained the earlier decision of Chapman to preserve R. fas- 
cicularis and reduce R. distans must be followed. 

Along the coastal plain from Virginia to Florida and west to 
Louisiana, R. fascicularis has been identified as a caespitose 
plant with a shock of coarse, often curling basal leaves 2-4 mm. 
wide, much shorter than the stiffly erect, subtriquetrous culms. 
The terminal cyme is nearly always irregularly compound, the 
spikelets 3-4.5 mm. long and composed of prominently aristate 


a ee ee ee ee eee 


SL eee ee eee ee 


1944] Gale,—Rhynchospora, Section Eurhynchospora 1938 


scales 3-4 mm. in length. The achene is usually ovate to orbicu- 
lar in outline, less frequently elliptic. Its surface is consistently 
smooth, blackish-brown, with a prominent pale disc. On 
smaller achenes, however, the disc may be indistinct or absent. 
Specimens from Virginia and Carolina have short rudimentary 
bristles not exceeding 144 the achene in height; but in Georgia 
and Florida occasional specimens occur in which the bristles 
may equal or exceed the achene.! 

R. fascicularis in Cuba, Jamaica, Hispaniola and Puerto Rico 
is usually represented by comparatively poorly developed speci- 
mens with reduced inflorescences. The achenes of these speci- 
mens are, however, both long- and short-bristled, and are indis- 
tinguishable from those of the mainland. 

All specimens with long-bristled achenes were formerly as- 
signed to R. distans. According to previous definition, in addi- 
tion to the increased length of its bristles, R. distans was separable 
from &. fascicularis by its consistently more slender habit. 
However, typically robust specimens of R. fascicularis from 
Florida and Georgia bear achenes with the long bristles of R. 
distans; and, vice versa, many of the more slender plants of the 
West Indies have rudimentary bristles, indicating R. fascicularis. 
To discriminate between R. fascicularis and R. distans on the 
basis of bristle-length or comparative size is to set up an arbitrary 
distinction. 

On the other hand, slender specimens from Virginia, Bermuda 
and Carolina south to the tip of Florida and west to Mississippi, 
bear achenes which are consistently long-bristled, smaller in 
size, characteristically elliptic in outline, with evenly browned 
surfaces, and narrow-based, triangular, subulate tubercles. 
To this category belong the fragments of S. distans from the 
Michaux Herbarium secured by Dr. Torrey and now in the 
Herbarium of the New York Botanical Garden, so that the name, 
Rk. fascicularis var. distans (Michx.) Chapm. is properly applied 
to them. The coarse plants with strongly mucronate scales but 
dark ovate achenes and well developed bristles may be con- 

' The original description of S. fascicularis was possibly based on a ‘ge Rages 
individual, for Michaux states ‘‘setulis semine laevi pain longioribus.’’ Asa Gra, 
however, reports that achenes from the Michaux type have ¢ short radixieantary bristles, 


which is true of the fragment from the Michaux Herbarium now in the Herbarium of 
the New York Botanical Garden 


194 Rhodora [May 


sidered as possibly transitional between this variety and the 
di ate short-bristled, widely-spread R. fascicularis var. typica. 

R. debilis, sp. nov. oo. foliis vel filiformibus 
debilibus ascendentibusque vel 1 mm. latis erectis planis; mar- 
ginibus apicem versus subtiliter serrulatis: culmis filiformibus 
rigidisque subteretibus, laxe ascendentibus vel procumbentibus, 
2.5-4.4 dm. altis: fasciculis 1— (rare)2; fasciculo terminali con- 
gesto acim! parvo (0.5-1.1 em. lato) : spiculis late ovoideis 
2.5-3.2 m Sear sessilibus, confertis, 2-floris, tertio rudi- 
mentario ree pis, saepe a achaenio maturante apertis: 
squamis slic "fissilibus, quam achaenio brevioribus, valde 
caducis, castaneis vel fuscis, costis pallidis; setis 5-6 quam 
achaenio duplo Secnoubad aut minus, antrorse  serrulatis: 
achaenio late ovato vel orbiculari, lenticulari biconvexo 1.4-1.6 
mm. lato 1.3-1.5 mm. longo laevi castaneo; disco pallidiore 
saepe obscuro; tuberculo compresso-deltoideo saepe apiculato 
0.3-0.4 mm. alto ; basi lato. PLatr 826, rigs. 5A and 5B; Map 
38.—? R. fascicularis var. trichoides Chapman, Fl. So. U. S. 527 
(1860), non R. trichodes Clarke-—Damp sandy or peaty clearings 
of the Coastal Plain from southeastern Virginia south to northern 
Florida and west to Alabama, with an inland station in the 
Alleghanies of Aa a VIRGINIA: exsiccated argillaceous 


Y, P); wet pineland south of. Petersburg, Prince George Co., 

Tul 13, 1938, Fernald & Long, no. 8602 (G, P); wet peaty clear- 
ing about 1 mile northeast of Burgess, Dinwiddie Co., Sept. 13, 
1937, Fernald & Long, no. 7355 (G, P); sandy and peaty bor der 
of Cat Pond, south of Benns Church, Isle of Wight Co., June 15, 
1938, Fernald & Long, no. 8118 (G, P, immature); damp sandy 
and peaty depressions back of the dunes, Rifle Range, south of 
Rudy Inlet, Princess Anne Co., July 30-31 and Aug. 4, 1934 
Fernald & Long, no. 3797 (G, P): peaty and argillaceous clearing 
in pineland east of Courtland, Southampton Co., June 10, 1938, 
Fernald & Long, no. 8119 (CU, a. immature); depressions in 
argillaceous field north of Littleton, ‘Sussex ae July 22, 1936, 
Ferna Long, no. 6085 (G, TYPE: NY, US, ISOTYPES); 
argillaceous and sphagnous meadow Beat ‘of Taylor’ s Mill- 
He! Greensville Co., June 12, 1939, Fernald & Long, no. 10145 
(G, P). Norra CAROLINA: Curtis (P, no locality given); damp 
open sand 5 miles west of Swanquarter, Hyde Co., June 22, 1927, 
Wiegand & Manning, no. 598 (G); moist rich soil, low eround 
along main highway gee Carteret Co., July 17, 1922, Ran- 
dolph & Randolph’ no. 8 (G). Sourn CAaRoLINa: cartroad 
through slg. 5 miles south of Andrews, George- 
town Co., Aug. 11, 1939, Godfrey & Tryon, no. 1376 (D, G, NY); 


PEE ey ee) aS See ae ae SSPE See ee nae ae Fee ae 


1944] Gale,—Rhynchospora, Section Eurhynchospora 195 


ney Mag aoe cr 2 miles west of Salters, Williamsburg 
Co., July 14, 1939, Godfrey & Tryon, no. 513 (G, NY); damp 
soils, Aiken, Aiken Co., Aug., 1866, Ravenel (G). Grorata: 
roadside, 5 miles east of Statenville, Echols Co., Dec. 31, 1937, 
Eyles, no. 2682 (CU). Fiorina: Chapman (US, no. 61756). 
ALABAMA: wet situation, low woods, St. Bernard, Cullman Co., 
July 17, 1909, Wolf, no. 479 (StB); Mobile, Mobile Co., May, 
1845, Sullivant (G); May, 1841, Buckley (G). 

R. debilis, R. Wrightiana and the two varieties of R. fascicularis 
probably constitute the most difficult group of species within 
the Section Eurhynchospora. Both R. debilis and R. Wrightiana, 
for which R. debilis has been mistaken, are characterized by 
their wiry, filiform to slender culms which tend to become pro- 
cumbent. The basal leaves of R. Wrightiana are typically fili- 
form and sinuously ascending, often equalling the culm in height; 
rarely they are 1.3 mm. wide and stiff. The reverse is true of R. 
debilis. Its leaves are more often 1 mm. wide, flat, stiffly as- 
cending and shorter than the culms. Occasionally, however, they 
are filiform, and then indistinguishable from those of R. Wrighti- 
ana. Similarly the fascicles of both species may be identical in 
appearance but, whereas those of R. Wrightiana are often 
reduced to 1 or 2 spikelets, those of R. debilis are typically 
corymbose and congested. A comparison of the achenes gives 
the most certain means of discrimination. That of R. debilis 
is round-ovate to orbicular, slightly umbonate, normally pale in 
the center, with dark brown somewhat depressed sides and blunt 
margins. It is surmounted by a deltoid broad-based tubercle 
which may be apiculate. The 5-6 subtending bristles are rudi- 
mentary, uneven in length, and rarely equalling 44 the achene. 
The achene of R. Wrightiana, on the contrary, is elliptic to sub- 
orbicular in outline, evenly biconvex, not umbonate, homo- 
geneously brown or nearly so, and immarginate. The tubercle 
is basally deltoid but, unlike that of R. debilis, is apically pro- 
longed into a broad blunt’bill. The six bristles, although uneven 
in length, are well developed, often equalling the achene. 

R. fascicularis is, by comparison with R. debilis and R. Wrighti- 
ana, a coarse, robust species. Of its two varieties only the weaker 
specimens of var. distans, with their narrow leaves, attenuated 
culms and reduced corymbs, might be confused with the more 
robust specimens of R. debilis or even R. Wrightiana. However, 


196 Rhodora [May 


the culms of normal specimens of var. distans are rarely, if ever, 
procumbent, the basal leaves are coarse, often spreading and 
curling, never filiform, with some of them at least 2 mm. wide, 
and in no case approximating the culm in height. The achene 
of R. distans most closely resembles that of R. Wrightiana. It is 
elliptic in outline, 1.1-1.3 mm. wide, 1.3-1.5 mm. long, evenly 
browned, and surrounded by six well developed bristles which 
frequently exceed the achene in height. However, the tubercle is 
merely triangular-attenuate, without the broad bill-like exten- 
sion characteristic of R. Wrightiana. It is this difference in the 
character of the tubercle, coupled with the difference in habit, 
which leads one to believe that R. Wrightiana is a good species, 
and not merely a delicate state of R. fascicularis var. distans. 

The filiform (rarely 1 mm. wide) leaves and culms, both of 
which are characteristically lax to procumbent, the small spike- 
lets with their round-ovate caducous scales, and its suborbicular 
achenes distinguish FR. debilis from the much coarser R. fascicu- 
laris var. typica. In habit it more nearly approaches R. fascicu- 
laris var. distans; but, unlike the latter, does not possess the 
broad radical leaves, the stiffly erect culms, and the bristles 
which in var. distans equal the elliptic achene and occasionally 
exceed the tubercle. 

k. debilis has, however, been most commonly confused with 
R. Wrightiana Boeckl. The latter species, first collected in Cuba 
by Wright, has since been found on the Coastal Plain from Flori- 
da to North Carolina. Like R. debilis, it has a weak, lax habit 
coupled with strictly filiform leaves and culms. The fascicles of 
h. Wrightiana, however, contain even fewer spikelets than those 
of R. debilis, and rarely form the small corymbiform glomerules 
characteristic of the latter. The essential distinction appears 
upon a comparison of the achenes. The achene of R. Wrightiana 
is commonly elliptic, exceeded by the six bristles, and surmounted 
by a deltoid-attenuated tubercle 0.7-0:8 mm. high. That of R. 
debilis is suborbicular, accompanied by 5-6 abortive bristles 
which do not exceed Y the achene in height, and is topped by a 
short deltoid tubercle only 0.3-0.4 mm. high. 

This entity was probably singled out by Chapman under the 
name f. fascicularis var. trichoides; but since Chapman’s types 
are obscure, and the name frichoides is not available for transfer 


. 


1944] Gale,— Rhynchospora, Section Eurhynchospora 197 


to the species, being already preémpted by Clarke for a West 
Indian species, R. trichodes, it seems best to treat the plant as a 
new species, and to designate a type. I have selected as descrip- 
tive of the lax to procumbent habit the epithet, ‘‘debilis”” which 
is taken from a memorandum written in Chapman’s hand and 
attached to a sheet of this species in the Gray Herbarium. 

(To be continued) 


oe 
of 
De ate 
te one 


- 


ay 
ae 


ae 


1944] aale,—Rhynchospora, Section Eurhynchospora 207 


RHYNCHOSPORA, SECTION EURHYNCHOSPORA, IN 
CANADA, THE UNITED STATES AND THE 
WEST INDIES 


SHIRLEY GALE 
(Continued from page 197) 


Series 6. Rariflorae, ser. nov. Foliis culmisque filiformibus 
erectis vel debilibus: cymis pluribus parvis patentibus: spiculis 
paucis remotis saepissime longe pedicellatis: setis antrorse ser- 
rulatis: achaenio ovoideo valde biconvexo castaneo vel pallido 
cancellato vel striato rugoso; rugis acutis. 

xrowing in bogs and on pond-margins of the Coastal Plain— 
R. rariflora infrequent inland and also occurring in the West 
Indies and Central America. Habit caespitose: leaves and 
culms filiform, erect to reclining: inflorescence of several small, 
open cymes: spikelets few, remote, mostly long-pedicellate: 
bristles upwardly serrulate, fragile: achenes ovoid, strongly 
biconvex, abruptly ridged, cancellate to striate, castaneous or 
pale, with 2 small whitish tongues of spongy tissue pushed out 
on either side at the base: tubercle deltoid, compressed.—Rhyn- 
chospora V. Glomeratae Small, Man. 175 (1933), in part. hyn- 
chospora, Series B. Diplostyleae, Sect. 4. Fuscae Clarke in Urban, 
Symb. Ant. ii. 105 (1900), in part. 


Key To SPECIES OF SERIES RARIFLORAE 


Spikelet 2—4-flowered, the achene when solitary accompanied 
by a sterile floret; bristles shorter than the achene; tubercle ; 
0.4-6.6 sm. Nigh. fo. or we a aes ae tas 36. &. rariflora. 


36. R. rartrtora (Michx.) Ell. Densely tufted: leaves fili- 
form to 1 mm. wide, involute at least on drying, loosely erect; 
upper margins finely serrulate: culms filiform to very slender, 
wiry, flexuous to reclining, 2.4-5.6 dm. high: eymes 1-3, small, 
lax, open, with few spikelets, corymbiform, the capillary branch- 
lets ascending to spreading; spikelets broadly ovoid, rarely 
sterile, remote, long-pedicelled, 2—4-flowered, 1—3-fruited, 3-4 
mm. long: scales round-ovate, obtuse, castaneous, often pushed 
apart at maturity: bristles 6, upwardly serrulate, unequal in 
length, the longest shorter than the achene: achene obovoid to 
rotund, strongly biconvex, 1.1—1.4 mm. wide, 1.3-1.4 mm. long, 
castaneous, traversed by abrupt ridges, striate to oblong- 
cancellate between the ridges; two small whitish ascending 
tongues of spongy tissue pushed out on either side at the base: 
tubercle compressed, deltoid, 0.3-0.6 mm. high. Pare 828, 


208 Rhodora [JuNu 


rics. 4A and 4B; Map 49.—Sk. Bot. 8. Car. and Ga. i. 58 (1816) ; 
Gray, Ann. Lye. N. Y. iii. 197, pl. 6, fig. 3 (1835); Chapman, Fl. 
So. U. S. 524 (1860); Britton, Trans. N. Y. Acad. Sci. xi. 92 
(1892); Clarke in Urban, Symb. Ant. ii. 130 (1900); Small, Fl. 
196 (1903) and Man. 183 (1933); Britton, Mem. Soc. Cubana 
Hist. Nat. ii. 197 (1916); Kiikenthal, Fedde Rep. Spee. Nov. 
xxxii. 78 (1933). Schoenus rariflorus Michaux, Fl. Bor.-Am. 1. 
35 (1803); Muhlenberg, Descript. Gram. 10 (1817). &. setacea 
sensu Grisebach, Cat. Pl. Cub. 243 (1866); C. Wright in Sauvalle, 
Anal. Acad. Ci. Habana, viii. 84 (1871) and Fl. Cub. 180 (1873); 
non Vahl. Phaeocephalum rariflorum House, Am. Midland Nat. 
vi. 202 (1920).—Open peaty depressions, bogs, or pond-margins 
of the Coastal Plain from New Jersey and southeastern Virginia 
south to the Florida Peninsula and west to Texas, with isolated 
inland stations in the mountains of Georgia, in central Tennessee 
and northern Texas; also in western Cuba, the Isle of Pines, 
Jamaica, the Dominican Republic, and Central America. NEw 
JeRSEY: clay-bottomed bog, Cold Spring, Cape May Co., 
Gershoy, no. 168 (G). VirGcinia: boggy swale, Otterburn, about 
1 mile west of Amelia Court House, Amelia Co., Fernald & Long, 
no. 8986 (G, P); sphagnous bog about 1 mile northeast of Burgess, 
Dinwiddie Co., Fernald & Long, no. 7354 (G, P); argillaceous and 
siliceous boggy depressions about 3 miles southeast of Peters- 
burg, at head of Poo Run, Prince George Co., Fernald, Long & 
Smart, no. 5656 (G); wet peaty margin of pine woods about 3 
miles southeast of Zuni, Isle of Wight Co., Fernald & Long, no. 
6083 (G, P); damp peaty meadows behind dunes, Rifle Range, 
south of Rudy Inlet, Princess Anne Co., Smith & Hodgdon in Pi. 
Exsic. Gray. no. 624 (CU, G, NY, P, US); peaty openings 
bordering wooded swamp along Mill Creek, about 1 mile north 
of Skippers, Greensville Co., Fernald & Long, no. 8601 (G, P). 

ORTH CAROLINA: open pine forest, used soil south of Bennett 
Memorial, west of Durham, Durham Co., Blomquist, no. 9803 
(D); edge of swamp, 14 mile within west boundary of county, 
highway 264, Nash Co., Blomquist, no. 7686 (D); rich moist low 
soil near Chocowinity, Beaufort Co., Correll, no. 1589 (D); 


no. 11 ouTH CarRoLINA: Aiken, Aiken Co., July, 
Ravenel (G); sunny banks near pond, Lotus Pond, Savannah 
River Refuge, Jasper Co., Eyles, no. 60 GEORGIA: 


woods about 2 miles west of Wrightsville, Johnson Co., Harper, 
no. 1345 (NY, US); low grounds near depot, Millen, Jenkins Co., 


Se Sg 


1944] Gale,—Rhynchospora, Section Eurhynchospora 209 


Harper, no. 781 (G, NY, US); moist sandy roadside, Bullock 
Co., Harper, no. 879 (G, N ¥, US); pine barren, 13 miles north 
of Waycross, Ware Co., Eyles, no. 7027 (G); low wet edge o 
woods between Folkston and Mattox, Charlton Co., Correll, no. 
5520a (D); moist pine barrens, near Huntington, Sumter Co., 
Harper, no. 13896 (G, NY, US); damp margin of cypress-pond 
about a mile north of Jakin, Karly Co., Harper, no. 3629 (P, US); 
Michaux Herb. (G, TypE-PHOTO; NY, TYPE-FRAGMENT from 
Michaux Herb.). FLorrma: moist pine barrens near Jackson- 
ville, Duval Co., Curtiss, no. 4870 (G, NY, US); Eustis, Lake Co., 
Nash, no. 438 (CU, G, NY, US); in a wet ditch in low pineland 
at the crossroads east of Fivay, Paseo Co., O'Neill, no. 2613 
(CA, CU, US); Tampa, Hillsborough Co., Oct., 1877, Garber 
(G, P, US); in moist ditch along roadside, about 5 miles north of 
Parish, Manatee Co., Moldenke, no. 1053 (D, NY); cut-over 
flatwoods 5 miles east of Lake Rudy, vicinity of Polk Co., 
McFarlin, no. 5145 (CU); Istokpoga Prairie, between Lake 
Istokpoga and Kissimmee River, Okeechobee Co., Small & 
DeWinkler, no. 9058 (NY): Alapittah Flats, St. Lucie Slough to 
Lake Okeechobee, St. Lucie Co., Small &- ‘DeW inkler, no. 9513 
(NY); low grassy pine barrens, Apalachicola, Franklin Co., 
Chapman in Biltmore Herb., no. 259a (NY, US). ALABAMA: 
Auburn, Lee Co., July 3, 1897, Earle & Baker (NY); boggy ser? 
slope near Jasmine (on M. an nd O. R. R.), Chilton Co., Apr. 

1921, Harper (NY; US with collector’s no. 171); Mobile, Probile 
Co., May, 1845, Sullivant (G). TENNESSEE: swamps between 


Spencer and Cagle, Van Buren Co., son, no. 9644 (G) 
Mississtppr: Ocean Springs, Jackson Co., Tracy, no. 4863 (NY, 
US); Biloxi, Harrison Co., Tracy, no. 489 NC, NY, US) 


LovtstaNna: in low pine barrens near Mandeville, St. Tammany 
Parish, May 1, 1893, Langlois (G, Mo, US); in cut-over pines 
four miles west of Hamm ond, Tangipahoa Parish, Trotter & 
Chilton, no. 141 (La); New Orleans, Drummond, no. 414 (G); 
in low prairies, Pointe aux Loups, Acadia Parish, Sept. 17, 
1894, Langlois (CU, US); wet prairies, Sulphur, Caleasieu 
Parish, Palmer, no. 7712 (CA, P, US). Texas: along road 
on dry sandy soil, West Orlando, Orange Co., Apr., 1936, 
Shee! (CU); bogs, Hempstead, Waller Co., Hall, no. 716 (G, 

y NE; US); San Antonio, Bexar Co. | Sept. 20, 1891, Plank 
(NY); e. Texas, Wright (G); Hookley Co., Thurow (US). Cua: 
white sand, vicinity of Los Indios, Isle of Pines, Britton, Britton 
& Wilson, no. 14175 (NY); Isle of Pines, Taylor, no. 59 (G, 
Ny, US); Colpothrinax savanna, between Pinar del Rio 


no. 3400 (G); Hacienda San Julian, south of Guane, Pinar 
el Rio, Leon & & Roca, no. 6937 (NY): eee places, Mateo 


210 Rhodora [JUNE 


Sanchez, Pinar del Rio City, Pinar del Rio, Ekman, no. 17935 
(NY); common, moist places near rivulets in pinelands, Herra- 
dura, Pinar del Rio, Ekman, no. 17719 (US). Jamatca: in the 
morass, Pedro Morass, Upper Clarendon, alt. 3000 ft., Harris, 
no. 11223 (NY, US). HIsPaNIoLa: slightly moist places, Pimen- 
tal, prov. Duarte, Valle del Cibao, Santo Domingo, Ekman, no. 
13328 (G, US); in savanna, El Valle, Sabana de la Mar, prov. de 
Samana, Cordillera Central, Santo Domingo, Ekman, no. 15700 
(NY, US 
37. R. STENOPHYLLA Chapm. Tufted: leaves filiform, strongly 
involute, flexuous, ascending, upper margins minutely serrulate: 
culms filiform, wiry, flexuous, 3.1-9 dm. tall: cymes 1—2, small, 
weak, lax, corymbiform, bearing only 1-5 fertile spikelets on 
capillary ascending to divaricate branchlets: fertile spikelets 
eng to lanceolate in outline, usually 1-flowered, 1-fruited, 3-5 
long, remote; sterile spikelets frequent, smaller, fusiform: 
scales lanceolate, pale, tightly imbricate: bristles 6, delicate, 
darker than the achene, with few antrorse serrulations, unequal 
in height, shorter than or exceeding the tubercle: achene obovoid, 
strongly biconvex, with surface transversely ridged, cancellate, 
pale, 1-1.3 mm. w ‘ide, 1.3-1.5 mm. long, with two small whitish 
ascending tongues of spongy tissue 0.4 mm. in height, pushed 
out on either side at the base: tubercle See aoa deltoid- 
pare ‘Gee 4 mm. long. Puarr 828, rics. 5A and 5B; 
Map 41.—F ie S. 525 (1860); Small Fl. 198 (1903) and 
Man. 186 eae Britton, Trans. N. Y. Acad. Sci. xi. 92 (1892), 
excl. syn. R. tenuifolia Griseb. Mal-ecuent 3 in swamps or boggy 
ground, often in pine woods, Coastal Plain of the Carolinas, 
northwest Florida and Alabama. NortH CAROLINA: boggy 
ground ca. 3 miles north of Hampstead, Pender Co., Blake, no. 
11906 (G, US). Sour Carouina: Aiken, Aiken. Co., July, 
1866, Ravenel (G, NY). Fuoripa: Apalachicola, Franklin Co., 
Chapm man (NY); low grassy pine barrens, Apalachicola, F as 
Co., Chapman in Biltmore Herb., no. 4479 (US); swamp nea 
Argyle, Walton Co., Curtiss, no. 5945 (G, NC, NY, US); fer rile 
pine woods, Walton Oo. Curtiss, no. 3174 Cu, ’P, US) 5, Warring- 
ton, Escambia Co., Trac y, no. 8613 (G, Y, US). ALABAMA: 
Great Bog, Mobile Co., July, 1889, Mohr “8; Bigelow (G). 
In contrast to R. rariflora (Michx.) Vahl, which it closely 
resembles, R. stenophylla has pale lanceolate scales with acute 
tips which, in the fertile spikelets, are closely imbricated about 
the solitary achene. A number of spikelets are characteristically 
erect, sterile, and fusiform. The scales of R. rariflora, on the 
other hand, are castaneous, broadly ovate, rounded or obtuse 
at the tip. Frequently the 2-several-flowered spikelet of the 


pS or ae eee ees 


1944] Gale,—Rhynchospora, Section Eurhynchospora 211 


latter matures 2-3 achenes which, during growth, cause the 
scales to separate. Unlike the condition in R. stenophylla, the 
spikelets of R. rarzflora are usually fertile and distant on wiry 
capillary pedicels. 

Two sheets of R. stenophylla from Chapman’s Herbarium 
(both of which are now at the National Herbarium, no. 968501 
and Biltmore Herbarium, no. 4479) have specimens the spikelets 
of which are comparatively numerous and more than 1-flowered. 
However, none of these spikelets have mature achenes, and, 
although they may have been merely immature when collected, 
there is also the possibility that a are pathological. 


Series 7. Cernuae, ser. nov. Plantae valde caespitosae saepe 
veep basibus saepe densis abrillosegue foliis setaceis saepe 


sis: setis antrorse serrulatis vel laevibus: achaenio parvo sub- 
tiliter ruguloso vel rugoso nitido et fusco vel castaneo ab valde 
corrugato et pallido. 

Small plants (with exception of R, nipensis) growing in ex- 
siccated locations, such as rocky crevices of stream-beds and 
dry pine barrens, ‘mostly restricted to the Selb of nan 

uba. Strongly. caespitose, often depressed: bases often dense, 
occasionally fibrillose, due to the fraying out and the persistence 

of the fibro-vascular strands of the old cauline leaves: leaves 

filiform, setaceous, often curling, or broad, flat and subcoriaceous 
with the upper surfaces exasperate or pruinose: cymes small, 
ovoid-capitate, spiciform or weakly corymbiform: spikelets : 
fruited: scales often pale, with obtuse apices, frequently short, 
leaving the upper portion of the achene and tubercle exposed: 
bristles 6, extremely fine, weakly ascending; antrorse serrulations 
sometimes imperceptible: achenes small, their surfaces finely 
rugulose, glossy dark brown to castaneous and rugose, or pee 
and heavily corrugate: tubercle compressed, deltoid.—Rhyncho- 
spora, Series B. Diplostyleae, Sect. 5. Glaucae Clarke in Urban, 
Symb. Ant. ii. 106 (1900), in part. 


Key To Species in Serres CERNUAE 
a. ee of 1-3 loosely genre cymes or a solitary 


spreading; upper s vid ige pruinose..........-..+-. a pruinosa, 
b. Radheal I leaves capill if 1 mm. wide flat, attenuate, 
approximating the culm in length, ascending; upper 
surfaces smooth. .. . 


212 Rhodora [JUNE 


c. Inflorescence loosely ae spikelets erect to ascend- 
ing; leaves capillary... . 

d. Plants Orne depressed tufts; bases hard, fibrous; 
leaves short, 0.6 dm. in he ight o r less, circinately 
ones reais 0.2-0.9 im. tall; achene obovate, 1 

ele ca eS ee ek ne ee ne 39. RK. crispa. 
d. Plants rea erect tufts; bases not fibrous; aie 
ed 


n he ight, sinuously ascending to erect; 
cule 2-3. 3 dm. high; achene slenderly er ser 
m. m. long PED A ne aia et ares «et 40. R. Shaferi. 


.3-1.4 m 
ce. Inflorescence of 1-2 dense, ovoid capituli; spikelets erect, 
ivergent and reflexed, the lowermost nodding in 
cernua; sth filiform to 1 mm. wide... .e. 
e. Leaves 1 m phe ascending, attenuate, approxi- 
fated 3.5 Bai length, nearly equalling ‘to exceed- 


to divaricate, occasionally reflexed; achene elliptic- 

ovoid, 0.8-0.9 mm. wide, | Sian mm. long... 2. 41. R. nipensis. 
e. Leaves filiform, curling, short, 0.15-0.8 dm. in length, 

less than V the height of the culm; capituli 1— 

(rarely)2; spikelets erect to nodding on _ short 

curved pedicels; achene obovoid, 0.8 mm. wide, 


BAUME oe Sl fee Ag Cha eee ae 42. R. cernua. 
a. incre consisting of 2 terminal, small, spreading cyme 
eral cymes; or, if cymes are poorly developed, 
of ae Spe echicallate: bases of the caespitose 
tufts not ae 
f. Mature Eanes oe ‘bristles Pag the former flattened, 
corrugate and stipitate. ... 
g- Gre forming Geared sed tufts: (nl 1-1.5 dm. or 
eak; scales whitish; achen e 0.7 m wide, 1.2 m 
tong; bri stles shorter than the BHNENE (ee ew 43. R. depressa 


g. Plants forming erect tufts, not genoa ed; culms 2.4—4.4 
dm. enderly erect; “petllen light brown; achenes 
aii wide, 1.3-1.4 mm. long; bristles exceed- : 
Sail oe 44. R. tenutfolra. 
f. Mature achenes and bristles castaneous to dark brown, e 
ormer strongly biconvex to plano-convex, ly t 
obscurely rugulose, not Pl a stipitate. cucu 
h. Radical leaves usually exasperate on the upper ony 
5-2 mm. wide; the sho oad, 


achene dark ply biconvex, with the 
bristles equally iatibited On Doth ides ss 45. R. scabrata. 
h. Radical leaves smooth-surfaced, 1 mm. wide, usually 


R. Lindeniana. 


38. R. pruinosa Griseb. Caespitose, often Ges basal 
leaves commonly short, 1.5 (rarely)—3 mm. wide, canaliculate, 
marginally revolute, with the ha Eee upper surface and the 
small white inclusions in many of the epidermal cells producing 4 
silvery to rimy effect; culms triquetrous, slender, erect, 3 

m. high, seldom bearing more than two short divaricate leaves: 
inflorescence of 1-2 (rarely 3) small glomerules which are oblong- 


\ 


Sa at 


— 


1944] Gale,—Rhynchospora, Section Eurhynchospora 213 


ovate in outline, 3-6 mm. broad; the smaller axillary glomerule 
subexserted to exserted on a slender erect peduncle: the bract 
subtending the terminal glomerule occasionally setaceous and 
inconspicuous, but more often short and leafy: spikelets ovoid, 
2—2.5 mm. long, compact, turgid, 1-fruited with the tubercle of 
the achene protuberant: scales suborbicular, tightly imbricate, 
pale: bristles 6, delicate and weakly ascending, upwardly serru- 
late: achene broadly obovoid, biconvex, 0.7—0.9 mm. wide, 1—1.4 
mm. long, gleaming dark brown, with barely discernible rugu- 
losities: tubercle subulate-triangular, 0.4-0.6 mm. long. PLATE 
826, rics. 3A and 3B; Map 42.—PI. Wright. pars 2: 535 (1862), 
preprint of Mem. Am. Acad. Sci. new ser. vill. 535 (1863): 
Grisebach, Cat. Pl. Cub. 243 (1866); C. Wright i ~ Sad alle, Anal. 
Acad. Ci. "Habana, vill. 84 (1871) and FI. Cub. 9 (1873), excl. 
syn. FR. scabrata Griseb. : Clarke in Urban, ae Ant. ii. 132 
(1900), exel. syn. R. scabrata Griseb. ; Britton, Mem. Soc. Cubana 
Hist. Nat. ii. 198 (1916), excl. syn. R. scabrata Griseb. —Damp 
woods or thickets often bordering brooks, mountains of eastern 
Cuba and northern Dominican Republic. CuBa: in woods, Gato 
Mts., Cobre Range of Sierra Maestra, alt. 1000 m. , Leon, Clement 
& Roca, no. 10424 (NY); in umbrosis, Loma Mensura, in cacum- 
ine montis, Sierra de Nipe, Oriente, alt. 1000 m., Ekman, no. 
9936 (NY): road to pinal, Mayari Abajo, Aug. 6, 1860," Wright, 
no. 1532 (G, type-number cited without jakalicy ‘by Grisebach); 
damp banks of stream in thickets, Sierra Nipe near Woodfred, 
Oriente, alt. 450-550 m., Shafer, no. 3488 (NY, US); Camp La 
Gloria south of Sierra Moa, Oriente, Shafer, no. 8084 and no. 
8085 (NY); banks of rivulets, pe villam Monte Verde dictam, 
Jan—Jul., 1859, Wright, no. 1532 (G, NY; type-number cited 
without locality by Grisebach) and no. 729 (G); Wright, ne 3391 

, US; also the number of the type-collection of R. scabrata). 
HISPANIOLA: rare, in forest, Loma Quita Espuela, prov. Duarte 
(formerly Pacificador), Cordillera Septentrional, Santo Domingo, 
alt. 800 m. Ekman, no. 12269 (NY). 

39. R. crispa, sp. nov. Caespites parvos valde depressos 
formans; basibus saepe fibrillosis: foliis basilaribus brevibus 
cireinnatis: culmis filiformibus flexilibus ascendentibus 0.2-0.9 
dm. altis, fasciculis 1-2 rare 3 parvis laxe spiciformibus; spiculis 
paucis; fasciculis axillaribus minoribus gracillime pedunculatis: 
bracteis et squamis imis setose prolongatis: setis 6, subtilissimis 
inaequalibus quam achaenio saepe longioribus fere laevibus: 
achaenio obovoideo biconvexo transverse ruguloso pallide cas- 
taneo 0.8 mm. lato, 1 mm. longo: tuberculo anne conico 0.6 
mm. longo. PLATE 827, rics. 4A and 4B; Map 43.—Wet rocks 
of the mountain streams, Sierra Nipe, eastern Cuba. CuBA: in 
wet crevices of rock subject to overflow, Arroyo del Medio above 
: ‘See Underwood, Bull. Torr. Bot. Cl. xxxii. 295 (1905). 


214 Rhodora [JUNE 


the falls, Oriente, Dec. 22, 1909, Shafer, no. 3266 (NY, TYPE); 
in pinetis (et carrascales) Sierra de Nipe, Oriente, July 24, 1914, 
Ekman, no. 2171 (NY); banks of upper Sojo River ~ 600 m. alt. 
Sierra de Nipe, Oriente, May, 1940, Carabia, no. 3731 (G); Pinal 
Mayari, 700 m. alt. Sierra de Nipe, Oriente, April 7, 1941, Leon 
& Victorin, no. 19906 (G, in part). 

The Ekman sheet, no. 2171, in the Herbarium of the New York 
Botanical Garden bears the copied annotation “R. cernua Gris. 
det. Kiikenthal, 1926.’ Considering the evident similarity of 
R. crispa and R. cernua, it is not strange that Kiikenthal failed 
to distinguish the one from the other. Both form small depressed 
tufts of curling leaves (although the leaves of R. crispa are 
apparently more tightly and usually inwardly rolled), from which 
short, capillary, wiry culms arise. Both have characteristically 
pale spikelets and the stubby ovate achenes with an identical 
pattern of surface-elaboration. The cymes of R. crispa, however, 
are strictly elongate, and are composed of more or less distant 
and erect—never crowded and pendulous—spikelets. The long 
setose subtending bracts pass by gradations into the shorter, 
less strongly setose, basal scales of the spikelets proper. The 
upper scales of R. crispa appear to be acute with aristulate tips, 
although from the over-ripe condition of the only available 
specimens neither this fact, nor the degree of protuberance of 
the achene from the spikelet can be satisfactorily determined. 

The small ovoid congested capituli of R. cernua, on the other 
hand, bear erect to pendulous spikelets, and are exceeded by only 
1-2 setaceous bracts at the most, which are sharply differentiated 
from the poorly developed lower scales. The uppermost scales, 
beyond which the tubercle and the upper end of the achene 
protrude, are characteristically emarginate with a short central 
mucro. 

Although the achene of R. crispa is of the same shape and size 
as that of R. cernua, it is. exceeded by several of the slender 
bristles and surmounted by an elongate flattened tubercle; 
whereas the achene of R. cernua is short-bristled, with a low 
conic tubercle. 

Shafer’s sheet, no. 3266, which I am designating as the TYPE 
of R. crispa, was identified by Britton as R. Berterii Clarke 
(R. pusilla Griseb.); but the two species are so utterly different 
in all but their general size and the bleached appearance of their 


1944] Gale,—Rhynchospora, Section Eurhynchospora 215 
spikelets that it is improbable that such a misidentification 
should reoccur. FR. pusilla is a lax little plant with flat, soft- 
textured, spreading leaves which are rarely filiform to 2 mm. 
wide—very different in appearance from the thick hard-based 
tufts of R. crispa with their strictly filiform subcoriaceous and 
inwardly coiling leaves. | 


40. R. SHarert Britt. Forming dense, upright tufts: basal 
leaves capillary, wiry, attenuate, flexuous, ascending; cauline 
eaves few, attenuate, approximating the height of the culms: 
culms many, capillary, wiry, flexuous-ascending, 2—3.3 dm. high: 
cymes solitary, small, loosely spiciform, 3-4 mm. wide, each 
exceeded by an upright, setaceous bract: spikelets ovoid, sessile, 
erect or ascending, 1—2-fruited, 2.4-2.6 mm. long; the tubercle 
and 14—\% the achene exposed: scales broadly ovate, pale; the 
apex rounded, with the midrib slightly, or not at all projecting: 
bristles 6, not equalling the achene in height, upwardly serrulate: 
achene slenderly ovoid, lenticular, compressed, transversely rugu- 
lose, castaneous, 0.8 mm. wide, 1.3-1.4 mm. long: tubercle conic- 
subulate, compressed, 0.5 mm. high. PLaTe 827, Frias. 2A and 
2B; Mar 44.—Mem. Soc. Cubana Hist. Nat. ii. 197 (1916). R. 
lingulata Kiikenthal, Fedde Rep. Spee. Nov. xxiii. 211 (1926).— 

hin soil, rocks on banks of mountain streams, Sierra Nipe, 
eastern Cuba. Cua: thin soil near base of mountain, Loma 
Mensura, Oriente, alt. 680 m., Shafer, no. 3797 (NY, Type; US, 
ISOTYPE); on the edge of Arroyo Machete, Sierra de Nipe, 
Oriente, Ekman, no. 15121 (NY, this number cited by Kiikenthal 
in type-description of R. lingulata); in carrascales-pinetis, Sierra 
de Nipe, Oriente, Ekman, no. 2172 (NY, US; this number cited 
by Kiikenthal in type-description of R. lingulata); rocks, bank of 
stream, Arroyo del Medio above the falls, Sierra Nipe, Oriente, 
alt. 250-500 m., Shafer, no. 4477 (NY); carrascales at Rio Pedra, 
Sierra, Sierra de Nipe, Oriente, alt. 200-300 m., Ekman, no. 
19107 (G, US), and same locality, Ekman, no. 10010 (G, this 
number cited by Kiikenthal in type-description of R. lingulata). 

41. R. nrpensis Britt. Densely caespitose with thick often 
heavily fibrous bases: roots coarse, sparingly branched, spongy: 
radical leaves 1 mm. wide or less, subcoriaceous, flexuous, 
ascending, approximating the culms in height; the tips flat and 
blunt, the margins finely serrulate: culms few, slender, erect, 
3-3.5 dm. high, bearing 1-2 short, blunt-tipped leaves and ter- 
minated by a small ovoid dense capitulum of spikelets: bracts 
few, short, filiform, divergent: fertile spikelets ovoid, 1-fruited, 
sessile, ascending to divergent, 2.5 mm. long; the tubercle and 
upper portion of the achene protruding; sterile spikelets numer- 
ous, small, fusiform: fertile scales ovate, short, with rounded 


216 Rhodora [JUNE 


apices: bristles 6, extremely tenuous, loosely ascending, antrorsely 
serrulate: achene elliptic-ovoid, compressed, often slightly asym- 
metric, 0.8-0.9 mm. wide, 1.3-1.4 mm. long; mutates dull pale 
brown, transversely rugulose, the rugulosities conspicuously 
striolate: tubercle seemingly confluent with the apex of the 
achene, conical-acuminate, 0.6 mm. long. PLarr 826, rias. 4A 
and 4B; Mar 45.—Mem. Soc. Cubana Hist. Nat. ii. 197 (1916), 
non Kikenthal, R. fibrillosa Kiikenthal, Fedde Rep. Spec. Nov. 
xxill. 212 (1926).—Rocks bordering mountain streams of the 
Sierra Nipe, eastern Cuba. Cusa: rock ravine, dry but subject 
to overflow after heavy rain, Sierra Nipe, along ‘trail from Piedra 
Gorda to Woodfred, Oriente, alt. 400-500 m., Shafer, no. 3103 
(NY, TYPE); ad ond sve fluvii Rio Piloto, Sierra de Nipe, 
Oriente, Ekman, no. 3244 (NY, cited by Kiikenthal in type- 
description of R. fibrillosa); on rocks of the high cascades of 
Rio Piloto, Sierra Nipe, Oriente, ca. alt. 700 m., Ekman, no. 
15173 (NY, cited by Kiikenthal in type-description of R. fibril- 
losa); in rock fissures, carrascales at Rio Piloto, Sierra de Nipe, 
ca. alt. 375 m., Ekman, no. aa (G, US; cited by Kiikenthal i in 
type-description of R. fibrillos 
42. CERNUA Griseb. Depressed, the basal leaves forming 
dense curly tufts with more or less fibrous bases: leaves filiform, 
setaceous, wiry, sparingly serrulate, much shorter than the culms: 
culms filiform, erect, 1—2-leaved, 0.4-2.5 dm. high: inflorescence 
1 (rarely 2) small, ovoid capituli 3-4 mm. wide; lateral capituli 
exserted on long filiform ascending peduncles: ‘spikelets ovate, 
mm. long, subsessile, erect to pendant, 1-fruited; the achene 
and tubercle protruding: scales obovate, notched at the apex, 
with the midrib slightly prolonged, stramineous to whitish: 
bristles 6, upwardly serrulate, height variable but falling short 
of the achene: achene obovoid, biconvex, transversely rugulose, 
Leieegion 0.8 mm. wide, 1 mm. long: tubercle a stubby cone 
0.3-0.4 mm. high. PLaTE 826, Figs. 2A and 2B; Map 46,—Cat. 
Pl. Gib. 248 (1866); C. Wright in Sauvalle, Anal. Acad. Ci. 
Habana, viii. 87 (1871) and Fl. Cub. 182 (1873); Clarke in 
Urban, By mb. Ant. ii. 133 (1900), excl. syn. R. brevirostris 
Griseb.; Britton, Mem. Soc. Cubana Hist. Nat. ii. 197 (1916).— 
Open places in pineland, mountains of eastern Cuba. CUBA: 
pinal near Baracoa, Oriente, June 15, 1861?,2 Wright, no. 3413 
¥, fragment. from Gray Herb. This number cited by 
Grisebach in type-description); 1860-64, Wright, no. 3365 (G. 
This number cited by Grisebach in type-description) ; forming 
small clumps in open places, Sierra Nipe near Woodfred, Oriente, 


BS B. Clarke lists R. brevirostris in the synonymy of R. cernua, due to a misprint 
in Griseb. which gives the Wright pole no. of R. brerirostris as 3414 (instead of 3410), 
making it the same as coll. no. of R. ¢ 

2 See Underwood, Bull. Torr. Bot. os atm 295 (1905). 


Se SEE ET ER er Te ee ee ee ee ST ee ON I ee ne ee 


Pe nae oa 
Fae Pe Re Te Serer ee ee ee 


Se ore 


ee ee 


1944] Gale,—Rhynchospora, Section Eurhynchospora 217 


alt. 500-650 m. NYG no. 2995 (NY, in part); same locality, 

pans no. 3048 '(N US); on bank of stream, Rene ae a Gloria, 
uth of Sierra Mek ’ Oriente, Shafer, no. 8218 (NY); baike of 

Smal stream, Aserrio de Moa, Oriente, Leén, no. 20254 (G). 

On a basis of his new species, R. cernua, Grisebach! drew up 
the section Microchaeta to contain those species which possessed 
tripartite styles. However, the specimens of R. cernua which I 
have examined have the typically bifid styles. Grisebach’s 
description was probably based upon an anomalous specimen 
with a tricarpellary gynoecium, a condition which occurs spo- 
ae throughout the Section Hurhynchospora. 


R. depressa (Kiik.), stat. nov. Growing in depressed 
Gite leaves filiform, canaliculate and wiry, erect or somewhat 
arching with the apices rounded and sparingly serrulate: culms 
wiry-filiform, subterete or flattened, 1-1.5 dm. high, with 1-2 
nodes towards their apices; cymes ix 2, strictly acelin the 
terminal cyme limited to 1-2 branchlets each of which bears 
either a pair of spikelets, the one subsessile, the other slenderly 
pedicellate, or a central sessile and two lateral pedicellate spike- 
lets; lateral cymes smaller, exsertly pedunculate, erect: bracts 
attenuate, setaceous, paralleling the culm, serrulate: spikelets 
2.2m m. long, erect, approximate, sessile or slenderly pedicellate, 
1 atk with the tubercle of the achene slightly protuberant: 
scales ovate, with the midrib extended slightly from a rounded, 
emarginate apex, papery, whitish, flecked with cinnamon toward 
the apex: bristles 6, shorter than the achene, with the fine up- 
ward serrulations somewhat prolonged and silvery toward cons 
bases: achene slenderly obovoid, lenticular, biconvex, 0.7 m 
wide, 1.2 mm. long, corrugate with fine | ongitudinal Oxi 
yellowish- brown: tubercle eed -based, pike-like, 0.4 mm. long, 
pale, scurfy. PiaTE 826, -lAa nd 1B; Map 48.—R. Lin- 


UBA: ad marginem rivuli, Minas de Iberia (a Taco Bay), 
Oriente, Ekman, no. 3808 (NY, US. This number cited by 
Kiikenthal in type-description of R. Lindeniana var. depressa). 

R. depressa was described by Kiikenthal in 1926 as a provi- 
sional variety of R. Lindeniana Griseb. Following the brief 
description the author stated that the specimens at hand were 
over-ripe and did not permit of a more reliable diagnosis. I 
have examined two sheets of the original collection, and, after 
careful comparison of these with specimens of R. Lindeniana, 


'Cat. Pl. Cub. 248 (1866). 


“ 


218 Rhodora [JUNE 


believe that Kiikenthal’s variety is not conspecific with Grise- 
bach’s plant. The matted, depressed habit, emphasized by the 
name, and the dense wiry filiform leaves distinguish the former 
from both R. Lindeniana var. typica and var. bahamensis (Britt.) 
Gale. 

Most fundamental, however, are the differences of inflores- 
cences and achenes. In R. depressa the cymes are of few spikelets, 
delicate, strictly ascending, and the scales are silvery white, 
flecked with cinnamon, emarginate, with short mucronulate 
tips. The cymes of R. Lindeniana, however, are congested, often 
irregularly corymbiform, and the scales, although often margin- 
ally bleached, are definitely castaneous to dark brown, with 
rounded or acute apices. The achene of R. Lindeniana is ovate 
to oblong-ovate in outline, plano-convex, indistinctly to finely 
rugulose, and a glossy dark brown at maturity. The base of 
the tubercle includes the summit of the achene. The achene of 
R. depressa, however, is obovate and so flattened as to be only 
slightly biconvex; its surface is banded by a few pale yellowish- 
brown corrugations. The narrow base of the tubercle of R. 
depressa does not include the summit of the achene, and extends 
to form a small pike-like projection. 

The type-locality, Taco Bay, Oriente, tends to strengthen the 
argument for the specificity of R. depressa, for the high rate of 
endemism in this province is well-known. 

44. R. renvurrouia Griseb. Tufted: basal leaves filiform to 1 
mm. wide, short, often curling, triquetrous and sparingly serru- 
late at the tips: culms filiform, loosely ascending, leafy, 2.4-4.4 
dm. high: cymes 2-3, 6 mm. wide or less; the filiform branches 
erect, approximate, with few spikelets; lateral cymes distant on 
subexserted peduncles, the lowermost not far from the base of 
the culm: spikelets lanceolate-acuminate in outline, 3.5-4.5 mm. 
long, 3-flowered, 1-fruited, erect on slender pedicels: scales 
slenderly ovate, acute, light brown, papery, loosely imbricate, 
promptly caducous: bristles 6, pale, extremely delicate, ascend- 
ing, heavily and upwardly serrulate, exceeding the tubercle: 
achene slenderly oblong-ovate, strongly flattened, broadly and 
heavily corrugate with fine longitudinal striae, pale, frequently 
dar brown in the narrow interstices between the corrugations, 
conspicuously stipitate, 0.7-0.9 mm. wide, 1.3-1.4 mm. long: 
tubercle attenuate-subulate, pale, 0.7—0.9 mm. high. Pate 827, 
rigs. 1A and 1B; Map 47.—Cat. Pl. Cub. 244 (1866); non R. 
tenuifolia Benth. (1878); C. Wright in Sauvalle, Anal. Acad. Ci. 


1944] Gale,—Rhynchospora, Section Eurhynchospora 219 


Habana, viii. 86 (1871) and Fl. Cub. 181 (1873). R. pee g 
sensu Boeckeler, Flora, lxiv. 78 (1881), non Torrey. R. 
phylla sensu Britton, Trans. N. Y. Acad. Sci. xi. 92 (1802), in 
part, non Chapman; Britton, Mem. Soc. Cubana Hist. Nat. ii. 
200 (1916), non Chapman. R. stenophylla sensu Clarke in 
Urban, Symb. Ant. ii. 130 (1900), exel. syn., non Chapman. 
R. stenophylla Chapm. var. albescens Kiikenthal, Fedde Rep. 
Spec. Nov. xxiii. 211 (1926).—River-banks and moist open wood- 
lands of northwestern Cuba. UBA: in dense tufts, banks of 
rivulets San Marcos, jurisdiccion Bahia Honda, Pinar del Rio, 
Apr. 17, 1863?! Wright, no. 3393 (NY: US, without locality. This 
number cited ae eee in type-description.)?; 1865, Wright, 
no. 313 ane no. 315 (NY); in cuabales, Minas, Habana, Ekman, 
no. 1313 8 (NY); Con Hill, Campo Florido, Havana, Leén, no. 
4733 (NY); in a clearing ‘of a wood, Loma de la Pita, San 
Miguel de Casanova, Havana, Leén, no. 11547 (NY); very 
sterile locality, in cuabales towards Canasi, Ceiba Mocha, Matan- 

Ekman, no. 18597 (US); Guabal del Espinal south of San 
aca station, Matanzas, Leén & Roig, no. 4142 (NY). 

Grisebach followed the original description of R. tenuifolia 
with the citation of the Wright numbers 3392 and 3393. The 
only sheet of number 3392 which I have seen is at the Gray 
Herbarium. On it are mounted three specimens; the central and 
largest one is R. setacea which Grisebach mentions as growing 
with R. tenuifolia. The other two specimens, which have fili- 
form leaves and culms, are R. Lindeniana, easily distinguished 
from R. tenuifolia by the finely rugulose rather than deeply 
rugose achene. 

According to Grisebach, R. tenuzfolia occurs in both eastern 
and western Cuba. However, the specimens which I have seen 
come exclusively from the western half of the island, that is, if 
the memorandum on Wright’s no. 3393 in the Herbarium of the 
New York Botanical Garden can be taken to apply also to no. 
3393 in the National Herbarium. However, the misidentified 
no. 3392 at the Gray Herbarium is from the vicinity of Baracoa, 
Oriente, suggesting the possibility that, if the sheet of no. 3392 
which Grisebach saw was similarly mixed, the inclusion of 
eastern Cuba in the range of R. tenuifolia is erroneous. 

R. tenuifolia Griseb. is the basis of the reports of R. stenophylla 
Chapm. from the West Indies. soee R. tenuifolia is very 

‘See Underwood, Bull. Torr. Bot. Cl. xxxii 297 (190 


*The sheet bearing numbers 3393 and 3392 at pee Gray Herbarium consists of 
two specimens of R. Lindeniana Griseb. and one of R. setacea Vahl. 


220 Rhodora [JUNE 


like Chapman’s species in appearance, both having the caespitose 
habit, filiform leaves ‘and culms, and small, weakly corymbose 
cymes of pale pedicellate spikelets, the organization of the 
spikelets and the appearance of the achenes are strikingly dissimi- 
lar. The spikelet of PR. tenwzfolia is 3-flowered, the lowest floret 
not maturing and the upper remaining rudimentary. The soli- 
tary achene, during its growth, forces apart the rather loosely 
imbricated, promptly caducous seales. The achene itself is 
slenderly oblong-ovate in outline, strongly flattened, pale, and 
broadly and heavily corrugate with the cancelli reduced to very 
fine longitudinal striae. Frequently the narrow interstices 
between the corrugations are dark brown, the tubercle narrowly 
attenuate-subulate, and the bristles extremely delicate (as is 
true in general of the Series Cernuae) pale and heavily serrulate. 

The spikelet of R. stenophylla, however, is usually 1-flowered 
(rarely 2-3). The solitary achene is tightly enclosed by the per- 
sistent scales. The achene, unlike that of R. tenuzfolia, is ovoid, 
prominently biconvex, and traversed by abrupt ridges between 
which occur the oblong cancelli. The surface is uniformly pale, 
except for two small tongues of whitish spongy tissue which 
occur, one on either side, at the base. The tubercle is triangular, 
and the bristles, which are heavier than those of R. tenwifolia, are 
only sparsely serrulate. 


45. R.scaBrata Griseb. Caespitose: basal leaves tufted, 1.5-2 
mm. wide, short, with obtuse to acute tips; margins serrulate; 
upper surfaces smooth to exasperate; cauline leaves few, short- 
bladed, ascending to divaricate: culms obscurely triquetrous, 
slender, flexuous, erect, 2.3-3.8 cm. high: cymes 2-4, corymbi- 
form, small (0.8-1 cm. wide), congested; more loosely organized 
cymes occurring on later-formed shorter culms; lateral cymes on 
included peduncles: spikelets ovoid, 3-3.5 mm. long: scales ovate 
to lanceolate, tightly imbricate, pale to castaneous, papery: 
bristles 6, extremely delicate, weakly ascending, upwardly 
serrulate, falling short of the tubercle: achene ovoid-ellipsoid, 
biconvex, gleaming dark brown, transversely rugulose, 0.9-1 mm. 
wide, 1.4 mm. long: tubercle attenuate-subulate, whitish, 0.9-1 
mm. long. 

45a. Var. typica. Basal leaves obtusely tipped, 1.5-2 mm. 
_ wide; upper surfaces exasperate: scales pale, concealing the 
tubercle. Puare 827, rics. 3A and 3B; Map 50.—R. scabrata 
Griseb. Cat. Pl. Cub. 243 (1866). R. Randii Britt. Mem. Soc. 
Cubana Hist. Nat. ii. 198 (1916).—Moist, shaded spots, moun- 


) 


a a ee Smet ee ae ne eteeee emer’ na ‘ = 
TIS OM Re eae oe RL Te NN Ne Se, Le: ea eae ea ee ogee ee ene eA eee et Pay ae 


ES a ee Sele ee in he ee 


1944] Gale,—Rhynchospora, Section Eurhynchospora 221 


tains of eastern Cuba. CuBa: pinal of Monte Libano, Oriente, 
Oct. 7, 1861," and pinal of Mayari, July 24, 1860,2 Wright, no. 
3391 (G; NY, US, without locality, this typE-NUMBER cited 
without locality by Griseb.); shaded bank of small ye Sierra 
Nipe near Woodfred, Oriente, Schafer, no. 3077 (NY, e of R. 
Randi Britt.) ; Sierra de Nipe, Oriente, Carabia, no. S08 (G); 

inal Mayari, East of ae Mensura, Sierra de Nipe, Oriente, 
Leén & Victorin, no. 19934 (G); open pineland, La Casimba, 
Sierra de Nipe, Oriente, 700) m. alt. Leén & Alain, no. 19241 (G). 

Since the publication of Sauvalle’s Flora Cubana in 1871, R. 
scabrata has been considered synonymous with the earlier R. 
pruinosa Griseb. This confusion may have had its origin in an 
unfortunate mixture of both species within the type-materia! 
(Wright, no. 3391) of R. scabrata; for one of Wright’s specimens 
of this number, at the National Herbarium, is undoubted R. 
pruinosa, label and citations to the contrary notwithstanding. 
And possibly the perpetuation of the confusion may have been 
due to the superficial similarity existing between the two species. 
Both are caespitose, often depressed, with a tuft of coarse, flat, 
basal leaves which are subject to degrees of the same peculiar 
roughness, and from which arise the slender, triquetrous culms. 

The similarity ends, however, with the inflorescence. The 
cyme of R. scabrata, although small, is spreading and corymbi- 
form, and the scales cover the tubercle or, in the case of var. 
laevifolia, the tubercle protrudes but does not extend beyond the 
lanceolate scales. The cyme of R. pruinosa, on the other hand, 
is glomerulate, ovoid-oblong in shape, and so congested as to 
simulate a small spike; while the characteristic suborbicular 
scales of the spikelet are exceeded by the wholly exposed tubercle. 
Then, too, the pronounced transverse rugulosities on the achene 
of R. scabrata are not at all like the fine almost indistinct sculp- 
turing on the achene of R. pruinosa. 

Britton apparently followed previous opinion in reducing #. 
scabrata to the synonymy of R. pruinosa and, accordingly, the 
Wright sheet, no. 3391, at New York bears his annotation “ R. 
pruinosa.” However, for identical material, also collected in 
Oriente, Britton published the name R. Randii which, now that 
R. scabrata is given rightful recognition, becomes a synonym of 
that species. 


1 See Underwood, Bull. Torr. Bot. Cl. xxxii. 296 (1905). 
?See Underwood, loc. cit. 295 


222 Rhodora [JUNE 


45b. Var. laevifolia, var. nov. Foliis basilaribus acuminatis, 
1.5 mm. latis, supra laevibus vel minute granularibus; squamis 
castaneis; tuberculis squamas superantibus. AP 51.—Moun- 
tains of eastern Cuba. Cusa: moist places, side of trail, Sierra 
Nipe, along trail Piedra Gorda to Woodfred, Oriente, Dec. 8, 
1909, Shafer, no. 3098 (NY, TYPE). 

The leaves of var. laevifolia and var. typica are short, approxi- 
mately 1.5 mm. wide, and basally tufted—identical except in one 
_ detail: whereas those of the latter have the peculiar exasperate 
upper surface, from which the specific name scabrata derives, 
those of the former are smooth or only minutely granular. 

It is notable that the achenes of both var. laevifolia and var. 
typica, although inseparable, are, at best, very similar to those 
of R. Lindeniana. Possibly the one specimen by which var. 
laevifolia is typified is the result of hybridization between the 
two very closely related species. 

46. R. Linpeniana Griseb. Densely caespitose: leaves lax, 
setaceous, 1 mm. wide or less, flat, marginally scabrous toward 
the often rounded apices; cauline leaves long-attenuate, erect: 
culms obtusely trigonous to subterete, slender to filiform, flexu- 
ously ascending, sometimes weak, 1.5 (depressed)—7 dm. tall: 
cymes 2-5, remote, with few spikelets; uppermost cyme small, 
congested, corymbiform, with many of the spikelets undeveloped 
and sterile; lowermost cyme not far from the base of the culm, 
loosely ascending; axillary bracts slender, equalling or exceeding 
the height of the cymes: spikelets ovoid to rotund, 2-3.2 mm. 
long, 1-3-flowered, 1-fruited; the tubercle protruding: lower 
scales mucronate; upper scales acute to obtuse, papery, castane- 
ous to ferruginous: bristles 6, extremely fragile, weakly ascend- 
ing, upwardly serrulate, assembled on the flat side of the achene: 
achene ovate to oblong-ovate in outline, plano-convex, 0.7—0.9 
mm. wide, 1.1-1.4 mm. long; surface indistinctly rugulose, 
glossy, castaneous: tubercle triangular-acuminate or sometimes 
prolonged and subulate, compressed, whitish, 0.8-1 mm. high. 

46a. Var. typica. Spikelets ovoid, 3-3.2 mm. long, with the 
tubercle protruding: scales castaneous, acute: achene ovate in 
outline, 0.7-0.9 mm. wide, 1.2 (rarely)-1.4 mm. long; tubercle 
prolonged, subulate, 0.8-1 mm. high. PLatTe 827, rics. 5A an 
5B; Map 52.—R. Lindeniana Griseb. Cat. Pl. Cub. 244 (1866); 
Clarke in Urban, Symb. Ant. ii. 126 (1900), in part var. baha- 
mensis; Britton, Mem. Soc. Cubana Hist. Nat. ii. 196 (1916); 
Kiikenthal, Fedde Rep. Spec. Nov. xxiii. 211 (1926).—Moist 
woodland in western and eastern Cuba. Cuba: low woods 
bordering manglares, Toscano, Pinar del Rio, July 29, 18637, 

1 See Underwood, Bull. Torr. Bot. Cl. xxxii. 297 (1905). 


1944] rale,—Rhynchospora, Section Eurhynehospora 223 


Wright, no. 3393! (G, in pert 1843-44, Linden, no. 1945 (G, 
IsoTyPE; NY, drawing of no. 1945 from Mus. Bot. Berol. with 
data ‘‘ Mt. Lebanon sur les soaee endroits couverts alt. 600 f. v. 
1844 leg. Linden’’)?; moist bank of trail, Sabanilla to Yamuri 
Arriba, Oriente, Shafer, no. 8428 (NY); Yamuri Arriba to Berme- 
jal, Oriente, Shafer, no. 8455 (NY, US); Camp la Gloria, south 
of Sierra Moa, Oriente, Shafer, no. 8086 Ne); Baracoa to Flori- 
da, he Shafer, no. 4328 (G, NY, 

Var. bahamensis (Britt.), stat. nov. Similar to var. 
typica in habit, but the spikelets more nearly rotund, 2-2.4 mm. 
long, with tubercle and achene protruding: scales often ferru-. 
ginous, usually white-margined; the uppermost blunt: achene 
oblong-ovate in outline, 0.8 mm. wide, 1.1—1.2 mm. long: tubercle 
triangular-acuminate, 0.5-0.7 mm. long. PLATE 827, FIG. 5C; 
Mar 53.—R. bahamensis Britton, Torreya, xiii. 217 (1913); 
Britton, Mem. Soc. Cubana Hist. Nat. ii. 198 (1916); Britton & 
Millspaugh, Bahama Fl. 55 (1920); Britton & Wilson, Sei. Surv. 
Porto Rico and Virgin Isl. v. 105 (1923).—Open to shady brook- 
margins and damp slopes of the Bahamas, eastern and western 
Cuba, eastern Hispaniola and Puerto Rico. BAHAMAS: along 
path in coppice, Soldiers Road, New Providence, Britton & 
Brace, no. 588 (NY, tTyPE of R. bahamensis); vicinity of Blue 
Hills, “New Providence, Wilson, no, 8241 (NY, US); mudholes of 
mangrove swamp, Deep Creek Andros, Brace, no. 5195 (NY). 
Cusa: in cuabales, Loma de Cajalbana, Pinar del Rio, Ekman, 
no. 12710 (G); dry open place, Loma Pelada de Buena Vista, 
Cayajabos, Pinar del Rio, alt. 400 m., Leén, no. 13797 (N Y): 
bushy savanna near Loma de la Pita, San Miguel de Casanora, 
Havana, Dec. 6, 1923, Leén (NY); in humidis, secus Rio Piedra, 
Sierra de Nipe, “Gusie Ekman, no. 1796 (G, NY). HISPANIOLA: 
very steep open mountainside, M. Bonpere, Gros-Marne, Massif 
du Nord, Haiti, Ekman, no. 4959 (G, NY, US). Purrro Rico: 
rocky slopes, Maricao to Monte Alegrillo, alt. 650-750 m., 
Britton, Stevens & Hess, no. 2552 (NY); Rio de Maricao, Hess, 
no. 559 (NY). 

The specimens of R. Lindeniana which I have studied fall into 
two groups. That to which the Gray Herbarium isotype, Linden, 
no. 1945, belongs, I am designating as var. typica. The other 
group, var. bahamensis (Britt.), includes Britton’s species R. 
bahamensis. 

Extremely diverse in habit, both varieties range from subde- 
pressed, filiform tufts to robust caespitose clumps. The spikelets 

‘This is the type-number of R. tenuifolia C. Wright and as such was obviously 
misapplied to the specimens on this sheet. 
seg scopulosis montis Liban.” as cited by Clarke in Urban, Symb. Ant. ii. 126 


224 Rhodora [JUNE 


of var. typica, however, are ovoid, acute, 3-3.2 mm. long, and 
1-fruited. The solitary achene is oblong-ovoid and slender, 
1.2 (rarely)—1.4 mm. long, completely enclosed by the castaneous 
scales, although the tubercle may protrude. 

The spikelets of var. bahamensis, by contrast, are nearly 
rotund, 2-2.4 mm. long, and often 2-fruited. Their scales are 
characteristically stubby and usually white-margined. Beyond 
them protrudes the upper portion of the ovoid achene which is 
1.1—-1.2 mm. long. 


Series 8. Cubenses, ser. nov. Spiculis ovoideis apicem versus 
tenuiter prolongatis pedicellatis; pedicellis longis, tenuibus, 
divaricatis vel reflexis: squamis dense imbricatis: achaenio fusco 
longitudinaliter irregulariter rugoso sive contracto et transverse 
laevissime ruguloso: tuberculo prolongato. 

rowing on damp, usually shaded hillsides, endemic in the 
West Indies. Coarsely caespitose: bases hard: leaves 2-4 mm. 


to reflexed, forming a globose network 2.5 em. wide; lateral 
cymes on included peduncles: spikelets basally ovoid with a 
prolonged, acute tip, borne on long, slender, divaricate to reflexed 
peduncles: scales tightly imbricate: achenes dark brown_ to 
black, longitudinally wrinkled as if shrunken, transversely but 
slightly rugulose: tubercle prolonged.—Rhynchospora, Series B. 
Diplostyleae, Sect. 5. Glaucae Clarke in Urban, Symb. Ant. il. 
106 (1900), in small part. 


Key TO SPECIES IN SERIES CUBENSES 


; ; 1.4-1.6 mm. long; 
bristles equalling to exceeding the tubercle; species limited to : 
Gat ek eC, Me wee Ae Gogoi e ox Seta Oke 7. BR. cubensis. 
Achene rhombic-elliptic in outline, conspicuously flattened, 
-1. . wide, 1.6-1.8 mm. long; bristles usually shorter : 
than the tubercle; species limited to Hispaniola. ...48. R. stenophylloidea. 


47. R. cupensis A. Rich. Coarsely caespitose with hard 
bases: leaves 2-3 mm. wide, flat, attenuate, loosely ascending, 
with acutely triquetrous scabrous tips: culms 3-angled, slender, 
leafy, loosely ascending: cymes 2-4, compound or decompouna, 
the wiry filiform, stiffly divaricate to mainly reflexed branchlets 
forming a globose network, 2-5 em. wide; lateral panicles on 
_ included peduncles: bracts foliaceous: spikelets basally ovoid with 
a prolonged acute apex, often split open by the maturing achene, 

1-fruited, 5-6 mm. long, solitary on slender divaricate to reflexed 
pedicels: lower fertile scale ovate-aristate, upper scale lanceolate- 
acuminate and tightly convolute about the prolonged tubercle, 


1944] Gale,—Rhynchospora, Section Eurhynchospora 225 


papery: bristles 6, extremely tenuous, irregularly ascending, 
upwardly serrulate, equalling to exceeding the tubercle: achene 
ovoid, swollen, 1.2-1.4 mm. wide, 1.4-1.6 mm. long, dark brown 
to black, longitudinally wrinkled as if shrunken, especially 
toward the apex, faintly rugulose from side to side: tubercle 1.1- 
1.8 mm. long, whitish, encrusted at the base, abruptly narrowed, 
then stoutly prolonged to a blunt apex. PLarr 828, Frias. 1A 
and 1B; Mar 56.—FI. Cub. Fanerog. ii. 294 (1853); Clarke in 
Urban, Symb. Ant. ii. 131 (1900), in part R. stenophylloidea 
(Kuk.) Gale; Britton, Mem. Soc. Cubana Hist. Nat. ii. 199 
(1916), in part R. stenophylloidea (Kiik.) Gale. R. deflexa Grise- 
bach, Cat. Pl. Cub. 243 (1866); C. Wright in Sauvalle, Anel. 
Acad. Ci. Habana, viii. 84 (1871) and Fl. Cub. 179 (1873).— 
Damp usually shaded hillsides of eastern and western Cuba. 
Cusa: in a low wood, north slope of Loma Pelada de Buenavista, 
Cayajabos, Pinar del Rio, alt. 420 m., Ledén, no. 13565 (NY); 
La Magdalena, Cayamos, Havana, Boker, no. 4644 (NY); Bafios 
de Casanova, Loma de la Pita, San Miguel de Casanova, Havana, 
Leon, no. 12480 (NY); Loma de la Coca, near Campo Florido, 
Havana, Leén, no. 2939 (NY); along brook, vicinity of Madruga, 
Havana, Britton, Britton & Shafer, no. 709 (NY); edge of creek- 
bank, dry open serpentine savannah, 10 kilometers west o 
Santa Clara, Santa Clara, Howard, no. 5089 (G, A); deep woods, 
limestone outcrop, Loma Ventana, Trinidad Mt., Santa Clara, 
Howard, no. 6533 (G, A); grassy shaded hillsides, Jucaral, Cien- 
fuegos Bay, Santa Clara, Britton & Wilson, no. 5742a (NY); not 
uncommon in wet shaded grassy lands, Cieneguita, southwest 
district of Cienfuegos, Santa Clara, Combs, no. 419 (G, NY); 
Monte Verde, Oriente, Aug. 28, 1859?! and Mayari-Abajo, 
Oriente, Wright, no. 3399 (G. This number cited without locality 
by Grisebach in type-description of R. deflexa); in tall clumps, 
gorge of the Rio Yamuri, Oriente, Shafer, no. 7809 (NY); dry, 
rocky hillsides, Sierra Nipe along trail Piedra Gorda to Woodfred, 
Oriente, Shafer, no. 3315 (NY); hills about Tabajo, base of El] 
Yunque, Oriente, Shafer, no. 8370 (NY); prope litus, prope 
Baracoa ad Navas, Oriente, Ekman, no. 3853 (NY). 

Grisebach apparently described R. deflexa in ignorance of the 
already existing R. cubensis A. Rich. He cited as the type, 
Wright no. 3399. The Wright sheet of this number at the Gray 
Herbarium bears two specimens, both of which are R. deflexa 
Griseb. (R. cubensis A. Rich.) as labeled; but no 3399 at the 
National Herbarium, although similarly labeled, bears by some 
mischance several specimens of the utterly different R. fascicularis 
(Michx.) Vahl, var. typica. 

See Underwood, Bull. Torr. Bot, Cl. xxxii, 294 (1905). 


226 Rhodora [JUNE 


48. R. stenophylloidea (Kiik.), stat. nov. In habit identical 
with &. cubensis, caespitose with hard bases: leaves 1.5-2.5 mm. 
wide, flat, long, slender, flexuous-ascending; the tips triquetrous 
with serrated angles: culms obtusely 3-angled, 0.3-1 m. high, 
lax, wiry, often extremely tenuous, with the upper of the long 
internodes filiform: cymes 2-3, compound to decompound, 1-3 
em. wide, loosely globose, subtended by a foliaceous bract; the 
wiry filiform branchlets divaricate to reflexed; lateral panicles on 
subexserted peduncles: spikelet 4-5 mm. long, basally ovoid 
with a prolonged acute apex, 1-fruited, with no trace of a suc- 
ceeding rudimentary floret, solitary on slender divaricate to 
reflexed pedicels: lower fertile scale ovate-aristate; upper scale 
lance-acuminate and tightly convolute about the prolonged 
tubercle, papery: bristles 6, extremely fragile and tenuous, 
irregularly ascending, upwardly serrulate, variable in length, 
the tallest approximating the tubercle: achene rhombic-elliptic 
in outline, conspicuously flattened, longitudinally wrinkled as 
if shrunken, faintly rugulose from side to side, 1-1.2 mm. wide, 
1.6-1.8 mm. long: tubercle slenderly conical or slightly com- 
pressed, somewhat sunken at the base, whitish, 1—-1.6 mm. long. 
PLATE 828, ric. 2A; Map 57.—R. cubensis A. Rich., var. steno- 
phylloidea Kiikenthal, Fedde Rep. Spec. Nov. xxxii. 78 (1933) .— 
Mountainsides on northern coast of Hispaniola. HispANioLa: 
edge of the Estére, Le Borgue, Massif du Nord, Haiti, Sept. 18, 
1925, Ekman, no. 4853 (G, US); in sylvestribus ad J yet Santo 
Domingo, alt. 150 m. , June 23, 1887, Eggers, no. 2600 (NY, US); 
mountainside, M. Bo onpere, Gros-Morn ne, Massif du Nord, 
Haiti, c. alt. 800 m. , Sept. 30, 1925, Ekman, no. 4958 (NY); in 
fruticetis ad Isabel de Torres, Apr. 23, 1887, Eggers, no. 1658 

Kiikenthal described R. cubensis var. stenophylloidea of His- 
paniola thus: ‘‘Corymbi perparce spiculosi, nux oblonga (haud 
late ovalis sicut in forma typica) enervis (non longitudinaliter 
striata), setae hypogynae 3 breves nucem superantes (nec 6 cum 
nuce aequilongae).” He concluded with “ Vielleicht eine gute 
Art, aber das vorhandene Material reicht nicht aus, um diese 
Frage zur entscheiden.”’ 

According to this description Kiikenthal’s specimens differed 
from the material which I am designating as R. stenophylloidea 
with regard both to the bristles and the longitudinal wrinklings 
on the nut. It is possible, therefore, that Kiikenthal’s type, 
Ekman, no. 14873, which is not available at the present time, is @ 
different entity. But the fact that this specimen came, as it did, 
from Hispaniola, to which island my species is apparently 


1944 Gale,—Rhynchospora, Section Eurhynchospora 227 


Rsond 


limited, and had a “‘nux oblonga,” described above as rhombic- 
elliptic but which is in any case larger than that of the typical 
R. cubensis, makes me suspect that the two are conspecific. 
Possibly three of the six extremely fragile bristles may have 
broken off at their bases prior to the time of Kiikenthal’s exam- 
ination; for when this occurs it is nearly impossible to ascertain 
their place of previous attachment. Kiikenthal’s statement as 
to the comparative length of the bristles is inaccurate, for those 
of R. cubensis equal or even exceed the tubercle in length and 
are longer than the bristles of the new species. The achenes of 
the specimens I have examined are, in all cases, more or less 
longitudinally wrinkled as if shriveled. I cannot suggest why 
this feature should be absent on the achenes seen by Kiikenthal 
unless they were in this respect atypical. 


Series 9. Harveyae, ser. nov. Culmis rigide erectis: cymis 
rigidis vel diffuse fasciculatis vel congestis ragga pe 
squamis pallidis vel castaneis: spiculis rity monocarpis: 
setis antrorse serrulatis: achaenio saepissime eta a tumido, 
infra compresso fusco leviter alveolato (ies lenticulari ruguso): 
tuberculo conico; basi haud discoideo sed saepe in achaenii 
apicem intruso 

Often growin on white pera - the Coastal Plain from 
North Carolina to Florida and west to Texas; inland in the 
southern and irwshers states; vate in Cuba. Stoloniferous or 
caespitose: leaves 1.5-7 mm. wide, not filiform: culms sti 
erect: cymes stiff, diffusely { fasciculate or congested and eles 
late: scales pale or castaneous: spikelets turgid, 1-fruited: 
bristles iiiandig serrulate: achenes usually tumid above and 
compressed below, dark mahogany in color, lightly pitted or 
cancellate in a ones te mk: pattern, rarely lenticular and ridge 
With oblong alveoli: tubercle always grayish or whitish, conical, 
not projecting at the base but slightly rina — overgrown 

y the summit of the achene.—Rhynchospor Glomeratae 
Small, Man. 175 (1933), in part. Paka Series 
Diplostyleae, Divisio 5. Eu- rr ogen satinge Sect. iii. Fuscae 


Key To Species IN Serres HARVEYAE 


Spikelets 6-7 mm. long, remote, some if not all slenderly pedicel- 

ate; achenes large, 2. cai mee wide, 3-4 mm. long. .49. i megalocarpa. 
Spikelets not exceeding 5.5 mm, n length, sessile in small c 

pact glomerules; a eons not exicedtl ng 2.2 mm. in width a 

2.4 mm. in len gth. 


228 Rhodora [JUNE 


Achene suborbicular to broadly ovate in outline, tumid above, 
ape ae sed below, the entire surface mahogany-brown at 


Spikelets 4-5.5 mm. long; bristles caning 1% the achene 
- ie the tubercle; achene 1 2 mm. wide, 2—2.4 

ep re link wg EOE ne ep es 50. R. Grayit. 
Spikelets 2 paca ve m. long; bristles equalling or shorter than 
1% the achene; aed 1.3-1.6 mm. wide, 1.5-1.8 mm. long 
51. R. Harvey. 

Achene obovate, slenderly lenticular, not tumid, surface 
crossed by pale irregular transverse ridges.............. 52... R. culiza, 


R. MEGALOcARPA Gray. Stoloniferous, the bases swollen 
and covered by short rusty imbricated scales: leaves rigid, sub- 
coriaceous, flat, 4-7 mm. wide, becoming conspicuously chan- 


ices on slender tory lateral cymes on long exserted 
slender ascending peduncles: spikelets ovoid, plump 5 bursting, 
some if not all pedicellate and remote, mainly ascending, 1- 
flowered, 1-fruited, 6-7 mm. long: scales mucronulate, chestnut- 
rown: stamens 10-12: bristles 6-8, very tenuous and scarcely 
thickened at the base, brittle, upwardly hispidulous, falling 
short of to barely exceeding the achene: achene round-ovoid, 
turgid in the upper portion, compressed below, 2.8-3.4 mm. wide, 
3-4 mm. long, emarginate, mahogany-brown.to blackish, glossy 
Bisentaiie where covered with a honey-combed surface-pattern 
of minute, extremely shallow pits: tubercle buttressed and some- 
what encrusted by the summit of the achene, conic-apiculate, 
grayish or often white, 0.7-1 mm. high. PLaTE 829, FIGs 
and 1B; Map 58.—Ann. Lye. N. Y. iii. 208, pl. 6, fig. 16 (1835); 
Torrey, "Ann . Lyc. N. Y. iii. 368 (1836) ; Chapman, Fl. So. U 
526 (1860); ‘Bacek alas Linnaea, xxxvii. 606 (1873). AR. wie 
candra Baldwin ex Gray, Ann. Lye. N. Y. iii. 207, pl. 6, fig. 15 
(1835); Britton, Trans. N. Y. Acad. Sci. xi. 90 (1892); Small, 
Bl. 196 (1903) and Man. 183 (1933). R. pycnocarpa Gray, Ann. 
Lye. N. Y. iii. 208, pl. 6, fig. 17 (1835). eaten dode- 
candrum House, Am. Midland Nat. vi. 202 20).—Deep 
Ww ners sand-ridges of the Coastal Plain from North Carolina 
poclahage: becoming frequent on the Florida Peninsula; less 
soataien est to Mississippi and inland along the Mississippi 
River reais collecting). Nortu Carona: sand-ridge at Carolina 
Beach, New Hanover Co., Godfrey, no. 4675 (G, NC); Wilming- 
ton, New Hanover Co., Mr. Curtis (NY, type of R. pycnocarpa) ; 
near White Lake, Bladen Co., Oosting, no. 33080 (D); dry open 
sand-barrens, 14 miles southeast of Lumberton, Robeson Co.. 
Wiegand & Manning, no. 597 (G); sandy ee tasiok Oak Island 


| Oe eS eee en 


SN SS de “ 


_ im Biltmore Herb., uo 


1944] Gale,—Rhynchospora, Section Eurhynchospora 229 


off Southport, Brunswick Co., Blomquist, no. 5613 (D). SourH 
CAROLINA: excavated area in coarse white sandy pine barren, 5 
miles “ges of pave Williamsburg Co., Godfrey & Tryon, 
no: 676 (CU; DG, NY, P): rarer: very dry sand-hills along 
Fifteen Mile Creek, Emanue 1 Co., Harper, no. 977 (G, NY); 
common, sand- scrub, Floyd’s [sland, Okefinokee Swamp, 
Charlton Co., J. S. Harper, no. 700 (G); very dry white sand 
south of open pond, Decatur Co., Harper, no. 1216 (G, NY, 
US). FuLoripa: Fort George, Duval Co., Dr. Baldwin (NY, 
TYPE); Baldwin (NY, type of R. dodecandra, immature; anno- 
tated by Gray); dry sandy soil near Jacksonville, Duval Co. ‘ 
Curtiss, no. 83161 (CU, D, G, P, US); in a serub, 18 miles north 
of St. Augustine, St. Johns Co., O’ Neill, no. 7688 (CU); Palatka, 
Putnam Co., April, 1869, Canby (G, NY); ; Gainesville, Alachua 
Co., April 12, 1897, Crawford ( He dry er in scrub, vicinity of 
Eustis, Lake Co., Na sh, no 2 (G, NX, US); Lake Butler, 
Orange Co., Beckwith, no. 556 (US); Ms pine barrens, Okeecho- 
bee Region, Brevard Co., Fredholm, no. 5734 (G, US); in a low 
pineland, Kissimmee, Osceola Co., O’Neill, no. 5121 (CU); 
scrub near Gadsen Hammock, vicinity of Winter Hav en, Polk 
Co., McFarlin, no. 4741 (CA); sand-barrens, Ballast Point, 
Tampa, Hillsborough Co., March 28, is Churchill (G); 
Dunedin, Pinellas Co., Tracy, no. 6995 (G, Y, US); scrub near 
Sebastian, Indian River Co. Small, DeWenklor & Mosier, no. 
11120 (NY): ; in ascrub, 2 miles north of Ft. Pierce, St. Lucie Co., 
O'Neill, no. 7689 (CU); in a scrub, 8 miles east of Lake Placid, 
Highlands Co., O'Neill, no. 7687 (CU); sandy ridge beside 
stream in pineland, 6 miles east of Manatee, Manatee Co., 
Oosting, no. 170 (D); Indian Mound near Citrus Center, DeSoto 
Co., Small, no. 9914 (NY); flatwoods, Alva, Lee Co., Hitchcock, 
no. 417 (G, NY, DB): sterile pine woods, Lantana, Lake Worth, 
Palm Beach Co., Curtiss, no. 5389 (G, NY, US); in serub-land 
9 miles north of Miami, Dade Cu., 0’ Weill, ‘no. 7691 (CA, CU); 
dry sand along the coast, ere Franklin Co., Chapman 
60 (G, NY, US); Port St. Joe, Gulf Co., 
Eyles, no. 3723 (CU): i in a high ee Pensacola, Escambia 
Co., O'Neill, no. 6091 (CU, US). ALABAMA: Fort Morgan, 
Baldwin Co., Tracy, no. 7684 (G, NY, US). Mi£ssissrppi:. 
Avondale, Bolivar Co., Tracy, no. 4840 (G, NY); moist sandy 
soil west of bay, Biloxi, Harrison Co., Pennell, no. 4388 (NY, P); 
Cat Island, Hancock Co., Lloyd & Tracy, no. 361 (NY). 
R. dodecandra Baldwin, R. pycnocarpa and R. megalocarpa 
Were simultaneously described by Gray in 1835. The type- 
specimen of the first was admittedly immature; that of the 


- second only somewhat so. A year later, in his revision of the 


North American Rhynchospora for Dr. Torrey’s' North Ameri- 
‘Ann. Lye. N. Y. iii, 368 (1836). 


230 Rhodora [JUNE 


can Cyperaceae, Gray placed R. pycnocarpa in the synonymy of 
R. megalocarpa with the remark: ‘‘ More perfect specimens com- 
municated by Mr. Curtis and Dr. Chapman have enabled us 
satisfactorily to ascertain that R. megalocarpa and R. pycno- 
carpas.. are different states of the seme species.”” He 
also added that: “It (R. megalocarpa) approaches R. dodecandra 
with which it also agrees in the prevalent number of its stamens.”’ 

Twenty-four years later Chapman, publishing his first edition 
of the Flora of Southern United States, reduced R. dodecandra 
to the status of a synonym under R. megalocarpa; and, in so 
doing, established a precedent for the use of the specific name, 
megalocarpa. However, in 1892, Britton!, in his list of North 
American Scirpus and Rhynchospora, gave preference to the 
specific name, dodecandra, and that name has recently come into 
general use. I have examined the types of R. pycnocarpa and R. 
dodecandra, as well as that of R. megalocarpa. With the mass of 
material now available for comparison, they are unquestionably 
referable to one species. Applying the rules of priority, the name 
R. megalocarpa, as selected by Chapman in 1860, must be rein- 
stated. 


50. R. Graym Kunth. Coarsely tufted: leaves arching, flat, 
2-4 mm. wide, smooth, becoming carinate; the upper margins 
and keel minutely serrulate; the midrib prominent: culms ob- 
scurely triquetrous, smooth, stifly erect, 0.4-7.5 dm. tall, with 
elongated internodes: the terminal cyme 1-1.5 cm. wide, com- 
pounded of 1-3 glomerules of few spikelets on short, erect to 
spreading peduncles; lateral cymes 1—4, consisting of a single 
small glomerule on an exserted peduncle: spikelets plumply 
ovoid, compact, 2-3-flowered, 1-fruited, 4-5.5 mm. long: scales 
mucronate, sandy to castaneous, tightly imbricate and entire at 
maturity: stamens 3— (rarely)6: bristles 6, upwardly hispidulous, ~ 
brittle, varying in length from equalling 14 the achene to exceed- 
ing the tubercle: achene suborbicular in outline, tumid toward 
‘the summit, with a more or less compressed base, the surface 
honeycombed with minute shallow pits, mahogany-brown, 
1.8-2.2 mm. wide, 2-2.4 mm. long: tubercle conic-apiculate, 
0.4-0.6 mm. high, buttressed and partially encrusted by the 
narrowed apex of the achene. Piare 829, Frias. 3A and 3B; 
Map 54.—Enum. ii. 539 (1837); Chapman, Fl. So. U. S. 526 
(1860); Small, Fl. 196 (1903) and Man. 183 (1933); Britton, 
Mem. Soc. Cubana Hist. Nat. ii. 197 ( 1916). R. distans Elliott, — 


1 Trans. N. Y. Acad. Sci. xi, 90 (1892), 


1944] Gale,—Rhynchospora, Section Eurhynchospora 231 


Sk. Bot. 8. Car. and Ga. i. 59 (1816), non Vahl. R. Elliottii Gray, 
Ann. Lye. N. Y. iii. 204, pl. 6, fig. 12 (1835), non Dietrich. 
Schoenus distans Muhlenberg, Descrip. Gram. 11 (1817). 
Schoenus fuscus Muhlenberg, Descrip. Gram. 6 (1817). Phaeo- 
cephalum Grayt House, Am. Midland Nat. vi. 202 (1920). Rare 
in Virginia (one collection); common southward on sandy pine- 
lands of the Coastal Plain from North Carolina to the tip of 
Florida and west to eastern Texas; also western Cuba. VIRGINIA: 
Norfolk, Norfolk Co., Read (P). Norru Caroxiina: Minnesat 
Beach, near Arapahoe, Pamlico Co., Oosting, no. 33208 (D); 
dry sandy soil near Wilmington, New Hanover Co., Biltmore 
Herb., no. 239g (CU); dry sandy pine woods, 2 miles southeast 
of Fair Bluff, Columbus Co., Wiegand & Manning, no. 606 (G); 
pineland at Roseboro, Sampson Co., Godfrey, no. 5723 (G); 
sandhill, 12 miles north of Laurenburg, Scotland Co., Godfrey, 
no. 5044 (D, G); Southern Pines, Moore Co., May 18, 1895, 
Blankinship (G). SoutH CaroLina: swampy spots in sandy 
land along stream, sandhills north of Hartsville, Darlington 
Co., Mar. 20, 1921, Norton (NC); Florence, Florence Co., May 
18, 1912, Bartram (P); Columbia, Richland Co., May 16, 1912, 
oe (P); 10 miles east of Paxville, Clarendon Co., Godfrey 


? 
Yard, Charleston, Charleston Co., Robinson, no. 259 (G). 
GrorGia: about Augusta, Richmond Co., June 27-July 1, 1895, 
Small (NY); sandy pinelands at Magnolia Springs, Jenkins Co., 
Eyles, no. 6255 (CU); dry pine barrens near Graymont, Emanuel 
Co., Harper, no. 806 (G, NY, US); dry sand-hills along Big 
Lott’s Creek, Bullock Co., Harper, no. 916 (G, NY, US); 
sand-hill west of Altamaha River on State Route 38, west of 


pine barren, just north of Ashim, Turner Co., Eyles, no. 5569 
(CU). Fioripa: dry pine barrens near Jacksonville, Duval Co., 
Curtiss, no. 4801 (G, NY, US); sand-hills, Welaka, Putnam Co., 


Ie Rhodora [JUNE 


Dade Co., Moldenke, no. 607 (D, NY); near Tallahassee, Leon 
Co., summer, Berg (NY); dry pine barrens, Apalachicola, Frank- 
lin Co., Chapman in Biltmore Herb., no. 239a (G, in part; NY, 
US); DeFuniak, Walton Co., Tracy, no. 9010 (G, NY, US). 
ALABAMA: Tensaw, Baldwin Co., Tracy no. 8028 (G, NY, US); 
common in woods, Spring Hill, Mobile Co., Bush, no. 335 (NY, 
US). Musstssrppi: Biloxi. Harrison Co., Baker, no. 1129 (NY). 
Louisiana: Hale (G). Texas: Liberty, Liberty Co., Mar. 25, 
1892, Plank (NY). Cusa: Laguna Los Indios and vicinity, 
Pinar del Rio, Shafer, no. 10820 (NY). 

Gray in his monograph pointed out that Elliott’s R. distans, 
as described in the Sketch of the Botany of South Carolina and 
Georgia, is not Schoenus distans Michx. on which it was nomen- 
claturally based. Accordingly Gray designated the former species 
R. Elliotizi in commemoration of its discoverer. He then placed 
in its synonymy two of Muhlenberg’s species, Schoenus distans 
and S. fuscus, with the explanation that, as the Muhlenberg 
Herbarium contains no specimen labeled S. distans, Muhlenberg 
probably derived his material from Elliott. 3S. fuscus Muhl., on 
the other hand, is represented in the herbarium by a specimen 
labeled “S. fuscus Elliott.” This, Gray states, is definitely P. 
Elliottii Gray. That Muhlenberg himself apparently suspected 
S. distans and S. fuscus to be conspecific (and incidentally that 
he also perpetuated Elliott’s mistaken reference to Michaux’s 
species) is evident from his suggestion, appended to the descrip- 
tion of S. fuscus: ‘An S. distans Michaux?” 

The legitimate name, R. Grayii, came from Kunth who, in 
1837, on the basis of the priority of Dietrich’s R. Elliotti 1833, 
renamed the species in honor of Asa Gray. 


51. R. Harveyrt Wm. Boott. Caespitose: leaves flat, obscurely 
carinate, with upper margins finely serrulate, ascending to curly, 
1.5-3 mm. wide: culms obtusely triquetrous, stiffly erect, smooth, 
0.3-1.1 m. high: terminal cyme 0.8-2.2 em. wide, usually com- 
pounded of 1-4 small glomerules on stiff ascending to spreading 
peduncles; lateral glomerules 1-2, usually solitary on wiry erect 
peduncles: spikelets ovoid, turgid, castaneous, 2-flowered, 1- 
fruited, 2.5-3 mm. long: scales with midribs continuing into 
conspicuously recurved mucros: stamens 3: bristles 6, delicate, 
equalling to falling short of 14 the achene, upwardly hispidulous: 
achene broadly ovate to suborbicular in outline, tumid above, 
somewhat compressed below, 1.3-1.6 mm. wide, 1.5-1.8 mm. 
long, rich mahogany-brown when mature; the surface honey- 


1944] Gale,—Rhynchospora, Section Eurhynchospora 233 


combed by small shallow isodiametric pits which may be ob- 
scured, appearing as faint rugulosities: tubercle squat, conic- 
apiculate, 0.4-0.5 mm. long, grayish, buttressed by the nar- 
rowed apex of the achene. PLatTEe 829, Fias. 2A and 2B; 
Map 59.—Bot. Gaz. ix. 85 (1884); Small, Man. 183 (1933). 
R. Earlei Britton ex Small, Fl. 197, 1328 (1903) and Man. 184 
(1933); Fernald, Ruopora, xxxix, 338 (1937) and xl. 398 (1938). 
R. Plankii Britton ex Small, Fl. 196, 1328 (1903); Small, Man. 
183 (1933), in syn. of R. Harveyi Wm. Boott. Phaeocephalum 
Plankit House, Am. Midland Nat. vi. 202 (1920). Phaeocephalum 
Karlet House, 1. e.—Low or frequently dry open areas in pine- 
lands of the Coastal Plain from southeastern Virginia to Florida 
and west to Texas; scattered inland stations in the coastal states, 
in western Tennessee, and on the lower drainage of both the 
Missouri and Arkansas Rivers, and on the Sabine River. Vir- 
GINIA: argillaceous and siliceous boggy depressions about 3 miles 
southeast of Petersburg, at head of Poo Run, Prince George Co., 
Fernald, Long & Smart, no. 5647 (G, NY, P); pinelands at western 
side of Wilcox Lake, Petersburg, Dinwiddie Co., Fernald & Long, 
no. 8596 (G, in part, P); very local, open pineland near Mason’s 
Siding, about 1 mile north of Henry, Sussex Co., Fernald & Long, 
no. 13274 (G); depression in dry open sandy pine and oak thickets 
near County Line, north of Emporia, Greensville Co., Fernald & 
Long, no. 8114 (G, ORTH CAROLINA: moist humus soil, 
open woodland 4 miles southeast of Wilson, Wilson Co., Randolph 
& Randolph, no. 723 (G); savanna, 1 mile east of Middlesex, 
Nash Co., Blomquist, no. 6358 (D); sandy soil, Cumberland Co., 
Biltmore Herb., no. 239f (CU); damp sand near brook, Pinehurst, 
Moore Co., Wiegand & Manning, no. 595 (G); Winston-Salem, 
Forsyth Co., Denke in Botanical Science Series, no. 5003 (D). 
SourH Carouina: cart-road through pineland-clearing, 5 miles 
south of Andrews, Georgetown Co., Godfrey & Tryon, no. 
1371la (G, NY); pine barren, 2 miles west of Pineville, 
Berkeley Co., Godfrey & Tryon, no. 619 (G, NY, P). Grorata: 
Marshallville, Macon Co., Earle, no. 2977 (NY, type of R. 
Earle?) and same locality, no. 2976 (NY, annotated as isotype 
of R. Earlei); rather dry pine barrens near Cobb, Sumter Co., 
Harper, no. 2217 (G, NY, US). Forma: 1839, Torrey (G). 
ALABAMA: low pineland, Elberta, Baldwin Co., Aug. 13, 1926, 
Wolf (StB); Mobile, Mobile Co., May, 1845, Sullivant (G). 
TENNESSEE: dry hill, Henderson, Chester Co., Bain, no. 244 (G). 
Missourt: ferruginous sandstone glade south of Birdsong, St. 
Clair Co., Steyermark; no, 13416 (Mo). ARKANSAS: Benton Co., 
1899, Plank, no. 29 (NY, type of R. Plankii); Grand_Prairie, 
eastern Arkansas, Harvey, no. 2 (G, TYPE). OKLAHOMA: Sapulpa, 
Bush, no. 656 (G, Mo, NY). Lovrstana: vicinity of Covington, 
St. Tammany Parish, Arséne, no. 11879 (US); along drain in 


234 Rhodora [JUNE 


Seymour Prairie, north of Bastrop, Morehouse Parish, Brown 
no. 6510 (La); Tong-leaf pine-hills, north of Po lock, Grant 
Parish, Brown, no. 6458 (La). Trxas: sandy woods, Mineola, 
Wood Co., Reverchon, no. 2278 (Mo, NY); damp sands, Will’s 
Point, Van Zandt Co., Reverchon, no. 2277 (Mo); Angelina Co., 
Tharp, no. 3056 (US); 144% miles northwest of White Ranch, 

hambers Co., Cory, no. 22402 (CU, G); 2 miles northwest of 
Benchley, Robertson Co., Cory, no. 21682 (CU); moist sandy 
soil, Kurten, Brazos Co., Walece. no. 13486 (Mo); low prairie, 
Hempstead, Waller Co., Hall, no. 714 (Mo, NY, US); Alvin, 
Brazoria Co., Apr. 20, 1894, Plank (NY); in wet ditch in red 
sand on clay subsoil; in railroad right-of-way cut through oak 
woods just east of Elgin, Bastrop Co., Innes, no. 874 (G); % 
mile south of Kicaster School, Wilson Co., Parks, no. 18795 (G); 
Indianola, Calhoun Co., Ravenel, no. 139 (NY). 


William Boott published R. Harveyi in the Botanical Gazette 
of June, 1884, and cited ‘‘Grand Prairie, E. Arkansas, F. L. 
Harvey, 1883.” At the Gray Herbarium are three Harvey 
collections of this species from this locality. One of these is 
dated July, 1884, so could not have been the original material 
seen by Boott. The two remaining specimens are not dated; 
one is unnamed. The other, however, is labeled R. Harveyi in 
Watson’s hand. This I take to be the -type, presuming the 
material collected by Harvey to have been sent to Watson and 
finally turned over to Boott for study. 

The achene of R. Earlei Britton ex Small has a low conic 
tubercle which is uplifted by the narrowed summit of the mahog- 
any-brown achene. These characters, seen in conjunction with 
the general size of the achene, indicate at once the close relation- 
ship of this plant to R. Harvey’. Unfortunately R. Earlei was 
published in Small’s Manual, and thus appeared without a word 
of discussion. Nor does its position between R. Torreyana and 
R. Edisoniana (R. microcarpa Baldw. ex Gray) in the text of 
that work give an indication that its relationship with R. Harveyt 
was appreciated. In the key, R. Earlei is separated from the 
former by the following character: “achenes transversely 
wrinkled” as against ‘“achenes cancellate.”’ However, in R. 
Harveyi the shape of the alveoli, on which the degree of wrink- 
ling depends, is subject to considerable variation. Typically, 
the alveoli are shallow and isodiametrie without any accentua- 
tion of the transverse walls, but occasionally they are crowded 


1944] xale,—Rhynchospora, Section Eurhynchospora 235 


into irregular transverse rows and take on narrowly oblong out- 
lines, the shorter horizontal walls of which are then accentuated 
and appear as fine ridges. More rarely the alveoli are nearly 
obliterated and the surface becomes as smooth and glossy as 
that of the achene of R. megalocarpa Gray. LEarle’s specimen 
from Marshallville, Georgia, was one with a finely ridged achene, 
and I presume that Britton, in recognizing it as a new species, 
was unacquainted with the amount of variation in the achenes 
of R. Harvey. 

However, the decompound, somewhat lax cyme of R. Earlei 
does vary from the smaller, stiffly upright, capitate cyme of 
typical R. Harveyi. This larger, looser inflorescence appears to 
be the only possible basis for segregation. Further collections 
from Georgia and Florida will be necessary in order to determine 
whether or not plants which were the basis of R. Earle: should 
receive recognition as a variety of R. Harveyt. 

52. R. culixa, sp. nov. ?Caespitosa: foliis basilaribus 2 mm. 
latis planis interdum brevibus et crispis; apicem versus leviter 
carinatis et subtiliter serrulatis; foliis caulinis brevibus: culmis 
triquetris gracilibus attenuatis crectis, cirea 3-foliatis; internodis 
ongis: glomerulo terminali circa 1 em. lato; glomerulo laterali 
minore pedunculato; pedunculo erecto: bracteis setaceis brevibus: 
spiculis late ovoideis tumidis 2-floris l-carpis 3 mm. longis 
dense aggregatis: squamis late ovatis, pallide castaneis dense 
imbricatis; apicibus fissilibus, mucronulatis: setis 6 antrorse 
serrulatis, achaenio duplo brevioribus aut minus; achaenio 
lenticulari-obovoideo biconvexo 1.2 mm. lato, 1.4 mm. longo, 
alveolato fusco; inter alveolas rugis transversis prominentibus 
pallidis: tuberculo breviter eonico, interdum apiculato, 0.3 mm. 
alto. PLaTE 828, rics. 3A and 3B; Map 55.—Rare in southern 
Georgia and northern Florida. GEorGIA: Irby, Tift Co., Aug. 
28, 1890, Tracy, no. 1498 (US, TYPE). FLorrpa: Chapman 

in part). 


The two specimens which I have designated as R. culixa 
differ from those of R. Harveyi Wm. Boott and R. globularis 
(Chapm.) Small, var. recognita Gale in the attenuate appearance 
of their slender culms, in the abbreviated cauline leaves and the 
compact, unbranched, small, terminal glomerules. Specimens 
of R. Harveyi and R. globularis, var. recognita are, when well 
developed, robust, with stiffly erect culms, linear-attenuate 
cauline leaves and usually compound, stiffly branched ultimately 
glomerulate cymes. 


236 Rhodora [JUNE 


The achene of R. culixa combines characters of both R. Har- 
veyi and R. globularis, var. recognita. It has the unmistakable 
grayish, squat, conical tubercle of R. Harvey: but the achene, 
like that of R. globularis, var. recognita, is obovate, lenticular, 
biconvex, not swollen above, and the transverse walls of the 
prominent alveoli are accentuated as ridges. However, unlike 
the ridges on the achene of var. recognita, those of R. culixa are 
pale, broader, and developed at the expense of the alveoli which, 
in the central area, have been rounded and thrown out of the 
orderly parallel alignment so conspicuous in the achene of var. 
recognita. 

Series 10. Globulares, ser. nov. Culmis rigide erectis vel 
rare tenuibus et laxe ascendentibus: cymis saepissime rigidis, 
glomerulosis vel fasciculatis; cymis lateralibus pluribus: spiculis 
turgidis, 1~3-carpis: squamis dense imbricatis: setis antrorse 
serrulatis saepe achaenio duplo brevioribus: achaenio late ovato 
vel suborbiculato ruguloso vel valde rugoso, cancellato vel 
striato, emarginat Oo. 

Growing in low peaty areas, often in pineland, of the Coastal 
Plain; inland in the midwestern states; also in the West Indies 
and Central America. Habit often coarse: leaves never filiform, 
1.5-5 mm. broad, often forming a coarse basal tuft: culms usually 
stiffly erect, rarely slender and loosely ascending: cymes glomer- 
ulate or fasciculate, usually stiff: lateral cymes several: spikelets 
turgid, 1—-3-fru ited: scales tightly imbricate: bristles upwardly 
serrulate, often poorly developed: achenes broadly ovate to 
suborbicular, rugulose to ridged (with the exception of R. globu- 
laris var. pinetorum), cancellate to striate, emarginate, com- 
pressed or swollen in the upper portion: tubercle wee basally 
conical.—Rhynchospora V. Glomeratae Small, Man. 175 (1933), 
in part. Rhynchospora, Series B. Diplostyleae, Sect. S Glaucae 
Clarke in Urban, Symb. Ant. ii. 106 (1900), in part. 


Key To Species 1n Series GLOBULARES 

a. ar wap nw flat or acon sunken; habit coarse; the basal leaves 
b. Tubercle with a subulate apex rising abruptly a a pro- 
nounced basal flange; achene dark reddish-brown. .53. R. compressa. 

d. bd conten deltoid-apiculate, the base somewhat esha 
pores: s achene castaneous 54. 
a. eae icon somewhat plano-convex, castaneous; 
habit slender or divides the bas. idea 2h 4 mm. wide or less... .¢. 

c. Bristles exceeding the achene i 8 height. ... 

d. Achene 1.2-1.3 mm. wide EGE oa tone the trans- 
verse ridges numerous: tubercle depressed, discoid- 
55. 


R. punctata. 


OU. oi es a Uh tee eos R. saxicola. 
d. Achene 0.8 mm. wide, 0.9 mm. long, the transverse 
ridges less than 6; tubercle Saab comicesmuent See es 56. R. sulcata. 


a a a a al aA a 


1944] Gale,—Rhynchospora, Section Eurhynchospora 237 


c. Bristles not exceeding 24 the achene.... 
é. ae imited to western Cuba; pertieh 0.8 mm. wide, 
MAT. FONG aces = rw Nee eee oa ge 7. R. Brittonii. 


F3 a vansverse Bak widening, becoming lighter and ob- 
scuring the alveoli over the central portion of the 
eae scikeleth ascending, not more than 6 in 
mall ultimate fascicles... .2..0) 2. eos eae n se 58. R. obliterata. 
2 Ticncas ridges pate adokiia r becoming light- 
colored, the alveoli not less aeenet over the orca 
area of the achene; spikelets ascending to diver 
and numerous in afennhte fascicles or glomerules 59. ar 4 globularis. 
53. R. compressa Carey ex Chapman. Caespitose: basal 
leaves flat, smooth or with margins Snel serrulate, 4-5 mm 
wide, forming a coarse rigid tuft: culms obtusely trigonous, 
stiffly erect, leafy, 6.6-9.6 dm. high: cymes 2-4, 1.8-3 cm. wide, 
densely bracteate, consisting of several glomerules on slender 
erect to spreading branchlets; lateral cymes remote, exserted on 
slender erect peduncles: spikelets ovoid, with a slightly irregular 
contour, 2—3-flowered, 1—2-fruited, castaneous, 3.5-4 mm. long: 
scales papery; the lowest mucronulate, the others acute: bristles 
6, upwardly serrulate, stiff, rarely equalling, never exceeding, 
the achene: achene obovate to nearly orbicular, 1.4-1.6 mm. wide, 
1.4-1.7 mm. long, characteristically flattened, often centrally 
depressed; the dark reddish-brown surface cancellate and trans- 
versely ridged: tubercle compressed-subulate, 0 0.6-0.8 mm. high, 
abruptly rising from a conspicuous ee collar. PLaTE 830, 
Fics. LA and 1B; Mar 64.—FI. So. U. S. 525 (1860); Small, Fl. 
197 (1903) and Man. 184 (1933); Robinson & Fernald in Gray, 
Man. ed. 7: 199, fig. 316 (1908). R. cymosa var. compressa 
toes tear Clarke ex Britton, Trans. N. Y. Acad. Sci. xi. 91 
(1892). Phaeocephalum compressum House, Am. Midland Nat. 
vi. 201 (1920).—Low pinelands and swamps, southern Georgia, 
Florida and west to eastern Louisiana. Gworata: flat pine 
barrens east of Ocilla, Irwin Co., Harper, no. 1414 (G, NY, Us). 
FiLoripa: Carey (G); low pine ‘barrens, Apalachicola, Franklin 
Co. 1 So (Pp). Knalachieahe Franklin Co., Chapma ). 
ae ama: Wilcox Co., May, ie Buckley oe brackish 


. Lovur 
Arséne, no. 11750 NY US). 
Old records from Missouri were based on misidentifications. 


238 Rhodora [JUNE 


54. R. puncrata, Ell. Caespitose, with a large clump of 
coarse basal leaves: leaves 5 mm. wide, carinate, becoming 
triquetrous at the tip, with setaceous margins: cauline leaves 
short, erect: culms stiffly erect, triquetrous, smooth, approxi- 
mately 7.6 dm. high: cymes 4, decompound, the stiff wiry 
ascending branchlets of varying lengths and terminating in 
small glomerules; terminal cymes 4 cm. wide; lateral cymes 
smaller, on long slender ascending peduncles: spikelets ovoid, 
5 mm. long, approximately 4-flowered, 1—2-fruited: scales pale 
chestnut, frayed; the midribs of the lower scales free at their tips 
and slightly projecting: bristles 6, equalling 14 the tubercle in 
length, upwardly hispidulous, ascending: achene obovate to 
suborbicular in outline, extremely compressed, cancellate, with 
fine transverse rugulosities, pale chestnut, 1.8 mm. wide, 2.2 
mm. long: tubercle deltoid-apiculate, compressed, pale, 0.9 mm. 
high. Puate 830, Frias. 2A and 2B; Map 65.—Sk. Bot. 8. Car. 
and Ga. i. 60 (1816); Gray, Ann. Lyc. N.Y. iii. 203, pl. 6, fig. 11 
(1835); Chapman, Fl. So. U. S. 526 (1860); Small, Fl. 198 (1903) 


about 1 mile southeast of Douglas, Coffee Co., Harper, no. 2200 
(G, NY, US). Fuorma: St. Mary’s, Baldwin (NY); “St. 
Mary’s & Savan,”’ Baldwin (P). 

55. R. saxicota Small. Caespitose: basal leaves forming a 
coarse curly tuft; cauline leaves 1.5-3 mm. wide, harsh, mostly 
erect, flat, becoming trigonous near the summit; margins finely 
serrulate: culms triquetrous, slender, erect, short, 2.6-3 dm. 
high: inflorescence of 2-4 cymes; the terminal one 1.3-2 cm 
wide, with a few slender erect branchlets bearing small clusters 
of sessile spikelets: bracts short, inconspicuous: scales tightly 
imbricate, castaneous; lowermost mucronate and usually split: 
spikelets ovoid, turgid, 3-flowered, 1-2-fruited, sessile, 3.5-4 mm. 
long: bristles 6, fragile, well exceeding the tubercle, upwardly 
serrulate: achene obovoid, compressed except for the slightly 
swollen region of the umbo, finely cancellate, ridged, castaneous, 
1.2-1.3 mm. wide, 1.3-1.5 mm. long: tubercle depressed, conic- 
apiculate, with a discoid base, 0.2-0.3 mm. high. Pxiate 830, 
rics. 3A and 3B; Map 66.—Man. 185, 1503 (1933).—Granite 
outcrops in the Piedmont of Georgia. Groreia: boggy slope on 
south side of Little Stone Mt., DeKalb Co., Harper, no. 2308 
(NY, type; US, 1soryre), and no. 2309 (NY); dry thickets on 
flat granite rocks, 1 mile east of Logansville, Walton Co., Pyron 
& McVaugh, no. 549 (US); shallow soil about granite outcrops, 
9 miles southeast of Greensboro, Greene Co., McVaugh, no. 
53828 (USNA); shallow soil about granite outcrops, 4 miles 
southeast of Sparta, Hancock Co., McVaugh, no. 5345a (USNA). 


1944] Gale,—Rhynchospora, Section Kurhynchospora 239 


a8 
ca 
Mee 


ts ee 


oe 


a 
< 


Range of 60, Ruyncnospora cALIFoRrNIcA; 61, R. Mariscuius; 62, R. 
CACUMINICOLA; 63, R. RuGosA; 64, R. compressa; 65, R, puncrata; 66, 
R. saxicona; 67, R. Brrrront ; 68, R. opniTeRATA; 69, R. suLCATA; 10, te. 
GLOBULARIS, var, TYPICA; 71, R. GLOBULARIS, var. PINETORUM; 72, R. GLOBU- 
LARIS, var. RECOGNITA; 73, R. oporaTA; 74, R. mi~tacea; 75, R. INEXPANSA; 
76, R. capuca 


240 Rhodora [JUNE 


The habit of R. saxicola has little to distinguish it from that 
of R. globularis (Chapm.) Small, var. typica. However, its 
spikelets are usually a millimeter longer and tend to be erect in 
clusters, rather than erect to divergent in glomerules or fascicles. 
The most obvious character by which to separate the achene of 
R. saxicola from that of R. globularis, var. typica is, as empha- 
sized in the key, the comparative length of the bristles. Those 
of R. saxicola are 6 in number, and, although extremely frail and 
apt to be broken off, do, when entire, exceed the achene and often 
the tubercle. Those of the former, however, rarely equal 1% the 
achene in height. The alveoli of R. saxicola also differ from the 
usually dull alveoli of R. globularis, for the former are so narrowly 
oblong as to appear merely as fine glistening striae running 
between the transverse wrinkles. The ‘‘depressed conic or 
disc-like tubercle’’, as described by Small, is not too dependable 
a character; for, although the average tubercle of R. globularis 
is short and conical, it may less frequently be depressed and even 
basally discoid. 

R. saxicola is peculiar in being strictly limited, so far as is 
known, to margins of shallow ephemeral pools on the granite 
outcrops in the Piedmont of Georgia. 


56. Re sulcata, sp. nov. Planta caespitosa: foliis 1-2.5 mm. 
latis laxis planis; apicibus trigonis: culmis 3.6-6.5 dm. altis 


turgidis 1-6-carpis 3 mm. longis: squamis fere orbicularibus 
castaneis deciduis: setis 6 fragilibus, achaenio aequalibus, 
antrorse et minute serrulatis: achaenio parvo (0.8 mm. lato 0.9 
mm. longo) subgloboso nitido castaneo profunde transverse 
sulcato; sulcis valde longitudinaliterque striolatis: tuberculo 
compresso-deltoideo albescente 0.2 mm. alto. PLaTE 831, FIGS. 
3A and 3B; Map 69.—Infrequent along streams, on pond- 
shores, and in low places of the Coastal Plain from Southern 
South Carolina to northern Florida. Sourn CaroLina: damp 
soils, Santee Canal, Berkeley Co., Sept., Ravenel (G); sedge- 


1939, Eyles, no. 6465 (CU). Gxorata: pond near Luciene Bay, 
Effingham Co., July 25, 1939, Eyles, no. 6378 (G, TYPE; CU, 
ISOTYPE); sandy shore of Open Pond, Decatur Co., Aug. 12, 


1944] Gale,—Rhynchospora, Section Eurhynchospora 241 


1901, Harper, no. 1204 (G, NY, US). Fiorrpa: Quincy, Gadsden 

0., Chapman (NY, in part): Tologee Creek, in John Carey’s 
handwriting (G). 

R. sulcata is closely related to both R. Brittonit Gale of Cuba 
and R. microcarpa Baldw. ex Gray of the southern United States 
and the Greater Antilles. The danger of confusion with R. 
Brittonvi is small due to the geographical separation of the two 
species, but the similarity in habit is so striking as to deserve a 


short discussion. Both species have in common slender flexuous, 


_culms which customarily bear small cymes at every node includ- 
ing the first. The lax branchlets in R. sulcata are, however, 
spreading to divergent so that, as a result, its panicles do not 
have the somewhat congested appearance of the inflorescence, 
with spikelets borne on the mainly ascending branchlets, of R. 
Brittonii. The achenes of R. Brittonzi are slightly larger than 
those of R. sulcata. Both are ridged, but those of the latter more 
abruptly so and with the concomitant grooves deeper. Lastly, 
the 5-6 bristles surrounding the achene of R. Brittonii are short, 
not exceeding 14 its height; whereas the achene of R. sulcata is 
surrounded by 6 bristles which equal it in height. 

R. microcarpa, on the other hand, is found within the range of 
R. sulcata, and in its smaller attenuated specimens simulates the 
typical habit of the latter. In R. sulcata, however, as mentioned 
above, the second, if not the first, node of the culm bears the 
first lateral panicle, so that the inflorescence occupies 24-34 the 
length of the culm. In R. microcarpa the first, second, usually 
the third, and often the fourth nodes are barren, and the inflores- 
cence occupies only the upper 14, rarely 4% the culm. A com- 
parison of the sculpturing on the surface of the achenes of the 
two species is even more conclusive, for that of the new species is 
emphatically ridged, whereas that of R. microcarpa is typically 
pitted, with the transverse walls only rarely accentuated so as to 
produce a rugulose effect. 

57. R. Brittonii, sp. nov. Planta caespitosa: foliis 1 mm. 
latis aut minus, planis, apicem versus trigonis et minute serru- 
latis: culmis vel obtuse trigonis gracilibus vel filiformibus tereti- 
busque, flexilibus, 2.8-5 dm. altis: paniculo terminali decomposi- 
to, leviter congesto, 0.8-1 em. lato; ramulis ascendentibus vel 
divergentibus: paniculis lateralibus exsertis pedunculatis: “igunon 
rotunde ovoideis, turgidis, 3—5-floris, 2—4-carpis, 2-2.56 mm 


aes MD i a Ni aaa 


242 Rhodora [JUNE 


longis: squamis fere orbiculatis dense imbricatis fuscis: setis 
5-6 achaenio duplo brevioribus, antrorse et minutissime serru- 


latis: achaenio subhemisphaerico, parvo (1 mm. lato, 1 mm. 
longo) inter rugas paucas profundas transversas hares tem oni 
striolato: tuberculo compresso-deltoideo, 0.2 mm. alto. PLATE 


831, rias. 2A and 2B; Map 67.—Borders of lata Isle of Pines 
and western Cuba. CusBa: savanna, San Pedro and vicinity, 
Isle of Pines, Feb. 12—Mar. 22, 1916, Britton & Wilson, no. 14301 
(NY, US); shore of Laguna de Junco, Pinar del Rio City, Pinar 
del Rio, Oct. 31, 1923, Ekman, no. 17862 (US); lagoon in savanna, 
vicinity of Pinar del Rio City, Pinar del Rio, Mar. 8-15, 1911, 
icin no. 10023 (NY, Typx; US, 1soryPs); dryish sand, 
aguna ero and vicinity, Pinar del Rio, Dec. 12 --3O1t, 
Shater, ioe 1090 (NY); border of lagoon, vicinity of Pinar del 
Rio City, Pinar del Rio, Sept. 5-12, 1910, Britton, Britton & 
Gager, no. 6946 (NY); Laguna de la Maguina, south of Pinar del 
(a City, Pinar del Rio, Nov. 28, 1940, Leén & Alain, no. 19410 


This species has been named in honor of Dr. NATHANIEL 
Lorp Brirron. It is most closely related to R. sulcata Gale, and 
the discussion of the two species follows the description of the 
latter. 


58. R. obliterata, sp. nov. Planta caespitosa: foliis 1.5-2 
mm. latis planis setaceis ascendentibus; marginibus sparse 
serratis: culmis trigonis gracilibus, 5.9—9. 2 dm. altis, apicem 
versus flexuosis: fasciculis 2-3 decompositis corymbiformibus 
2-3 mm. latis; lateralibus compositis minoribus erectis peduncu- 
latis exsertis: spiculis ovoideis, 1—3-floris, 1-2-carpis, 3.5-4 mm. 

ongis: squamis aristulatis, laxe imbricatis: setis 6, achaenio 
duplo brevioribus vel saepe rudimentariis, antrorse ’ serrulatis: 
rapist late feast lasicobovaiden biconvexo 1.2-1.3 mm. lato 

1.2-1.3 mm. longo nitido castaneo; alveolis in medio a rugis 
validis pallidis regained obscuratis: tuberculo hones apicu- 
lato 0.3 mm. alto. Pare 830, rigs. 4A and 4B; Map 68.— 
Borders of ponds in pe ae and southeastern tae ARKAN- 
sas: borders of ponds, BE. Ark.! J 1884, Harvey, no. 12 (G). 
Texas: Cypress City, Harris Co., in 1877, Boll, no. 793 (Mo); 
pende: Rates, Waller Co., April 16, 1872, Hall, no. 709 
(G, eae oe S, ISOTY YPES); prairie near Indianola, 
Calhoaa zoe 1869, Ravenel, no. 144 (NY, in part R. 
globularis ‘Ghaaesn ‘Saal var. recognita Gale). 

The specimens of R. obliterata cited above were segregated 
from collections of R. globularis (Chapm.) Small, var. recognita 
Gale (R. cymosa). They differ in general from the latter species 

‘ Located on the map as in the vicinity of Grand Prairie. 


1944] Gale,— Rhynchospora, Section Eurhynchospora 243 


both in their cauline leaves, which are narrower and setaceous, 
and in their lack of a coarse, curling tuft of basal leaves. The 
inflorescence of R. obliterata is loosely fasciculate. The numerous 
small clusters of spikelets, each borne on a flexuously ascending 
branchlet, are inconspicuously bracteate. The inflorescence of 
R. globularis var. recognita, on the other hand, has comparatively 
short, stiffly erect to divergent branchlets bearing tight ultimate 
corymbs or glomerules which are penetrated by short. stiff 
bracts. The lenticular achene of R. obliterata has a precise, 
broad-obovate outline. The transverse walls of the alveoli are 
aligned and pushed up into small ridges which, over the central 
portion of the achene, become conspicuously wider and paler, 
obscuring the alveoli. The achene of R. globularis var. recognita 
is by contrast irregularly obovate to suborbicular in outline, 
_ and tends to be more tumid above. Its small ridges are evenly 
continued across the face of the achene, and the alveoli of the 
central portion remain undiminished in size. The name of the 
new species derives from the characteristic blotting out of the 
central alveoli. 


compressed below, cancellate, transversely ridged to rugulose, 
castaneous, 1-1. . wide, 1.2-1.6 mm. long: tubercle conical, 
with or without a compressed apex, 0.3-0.6 mm. high. 
Key to Varreries oF R, GLOBULARIS 
Achenes transversely ridged or rugulose, their cancelli oblong. 
Habit frequently depressed; branchlets of the cymes termina- 
ting in pias 4 i I : 
spikelets 2.5-3 mm. long; achenes 1-1.2 mm. wide, 1.2-1. 


var. typica. 


m. long, 
coarsely cancellate to striate, transversely ridged. .59b. _ var. recognita, 


a0 es 6 Be eo a ace 0 eee be be bn ae we ee eee Le ae 


244 Rhodora (JUNE 


59a. Var. typica. Leaves 1.5-2 mm. wide: culms often short, 
but ranging from 1.4-6.8 dm. in height, slender, obtusely tri- 
gonous to subterete, often attenuate, wiry and flexuous: branch- 
lets of the cymes terminating in small knobby capituli of 3-8 
spikelets: bracts inconspicuous: spikelets 2.5-3 mm. long, 1-2- 
fruited: bristles 5 (rarely 6), usually less than 14 the achene in 
height: achene finely cancellate, transversely ridged to rugulose, 
1-1.2 mm. wide, 1.2-1.3 mm. long: tubercle short, conical. 
Puate 831, rics. 1A and 1B; Map 70.—R. globularis Small, 
Man. 184 (1933). R. cymosa var. globularis Chapman, FI. So. 
U. 8. 525 (1860); Britton, Trans. N. Y. Acad. Sci. xi. 91 (1892); 
Small, Fl. 197 (1903); Fernald, Ruopora, xxxvii. 380, 405 (1935) 
and xxxix. 391, 480 (1937).—Sandy or peaty depressions from 
Delaware south along the Coastal Plain to the tip of Florida and 
west to eastern Texas; inland to the upper Sabine and the Red 
Rivers; also in swamps of the coastal ranges of northern Califor- 
nia. DELAWARE: swamps, Newport, New Castle Co., July 12, 
1863, Commons (NY). Viratnra: ledges along Potomac River, 
Great Falls, Fairfax Co., Blake, no. 5277 (US); low woodroad 
north of Savedge, Surry Co., Fernald & Long, no. 8121 (G, P); 
sandy and peaty depression (exsiccated shallow pond) about 4 
miles northwest of Homeville, Sussex Co., Fernald & Long, no. 
6071 (G, P); wet argillaceous depressions south of Petersburg, 
Dinwiddie Co., Fernald & Long, no. 8120 (G, P); peaty and 
argillaceous clearing about 4 miles southeast of Emporia, Greens- 
ville Co., Fernald & Long, no. 8122 (G, P); near Northwest, 
Norfolk Co., Kearney, no. 1536 (US); sandy barrens, Rifle 
Range, south of Rudy Inlet, Princess Anne Co., Smith & H odgdon 
in Pl. Exsic. Gray., no. 625 (CU, G, NY, P, US). Nort Caro- 
LINA: argillaceous-siliceous clearing, 2 miles east of Conway, 
Northampton Co., June 14, 1939, Godfrey, (CU, D, G, P); open 
pine woods, acid soil, south of Bennett Memorial, Durham Co., 
Blomquist, no. 9799 (CU, D); damp thickety school yard, 7 
miles east of Lumberton, Robeson Co., Wiegand & Manning, 
no. 614 (G). Sourn Caroxina: sand pit, Combahee River, 
south of Hendersonville, Colleton Co., Wiegand & Manning, no. 
617 (G). Gxorera: Smithville, Lee Co., Earle, no. 2979 (NY); 
low grounds between Millen and Ogeechee River, Burke Co., 
Harper, no. 792 (G, US); rather dry sandy roadside in pine 
barrens, Bullock Co., Harper, no. 880 (G, NY, US); margins, 
Bouhin’s Pond, Chatham Co., Eyles, no. 6093 (CU); ditch along 
U.S. Route 17 south of Ways, Bryan Co., Eyles, no. 6286 (CU); 
dry open sink between Newton and Elmodel, Baker Co., Eyles, 
no. 7067 (Hermann Herb.). Fiorrpa: near Jacksonville, Duval 
Co., Curtiss, no. 4105 (US); Manavista, Manatee Co., Tracy, n0- 
6991 (US); in moist sandy ditch along roadside, Rialto, Lee Co., 
Moldenke, no. 1011 (US); in hammocks and pine lands, Black 


ia ; 
| 0 SLES ee er 


1944] Gale,—Rhynchospora, Section Eurhynchospora 245 


Point, below Cutler, Dade Co., Small & Carter, no. 880 (NY); 
Chipola River swamp, Jackson Co., Curtiss, no. 3 (G); swampy 
thicket, Washington Co., Curtiss, no. 3149 (CU, P, US). Mis- 
SISSIPPI: Picayune, Pearl River o., May 24, 1940, Sargent 
satpent Herb.). LovistaNna: in grassy soil, ‘Covington, St. 
mmany Parish, July 17, 1884, Langlois (CU); low prairies, 

jeaniaee Jefferson Davis Parish, ’ Palmer, no. 7624 (CA); infre- 
quent, low prairies, vicinity of Lake Charles, Caleasieu Parish, 
Mackenzie, no. 400 pe a EXAS: common in swamp, 
Big Sandy, Upshur Co., Reverchon, no. 2460 (Mo, NY); damp 
places, Dallas, Dallas Go, Reverchon, no. 3603 (G, US); wet sandy 
ground, Jacksonville, Cherokee Co., Palmer, no. 7905 (CA, Mo, 
P); springy places north of Palestine, Anderson Co., June 8, 1899, 

Eggert (Mo); low prairie, Hempstead, Waller Co., ’ Hall, no. 71 12 
(NY, US). Cauirornia: Pitkin Marsh, 5 miles north of Sebas- 
topol, Sonoma Co., Howell, no. 126 (G). 

59b. Var. recognita, nom. nov. Leaves 2-4 mm. wide: culms 
1.5 (rarely)—9.2 dm. in height, robust, trigonous and erect to 
attenuate, obscurely trigonous and flexuous: terminal capituli 
usually heavily crowded with spikelets: the pale bracts usually 
conspicuous: spikelets consistently larger than those of var. 
typica, 3-4 mm. long, 1-3-fruited: bristles 5-6, 44-14 height 
the achene in length: achene sootian ds cancellate to striate, trans- 
versely ridged, 1.2-1.5 mm. wide, 1.3-1.6 mm. long: tubercle 

0.6 mm. high, conic to triangular with the apex compressed. 

PLATE 831, rics. 4A and 4B; Map 72.—R. cymosa sensu Torrey, 
Fl. N. and Mid. St. no. 1: 56 (1823) and later authors to the 
present; Gray, Ann. Lye. N. Y. iii. 196, pl. 6, fig. 1 (1835); Tor- 
rey, Ann. Lye. N. Y. iii. 364 (1836); ‘Gray, Man. 532 (184 8) 
. Chapman, FI. So. U. 8. 524 (1860); Britton, Trans. N. Y. Aca 
‘ Sci. xi. 91 (1892), excl. syn. R. Harveyi Wm. Boott (R. Aes. 
q in error); Britton & Brown, Ill. Fl. * 280, fig. 658 (1896); Clarke 
7 in Urban, Symb. Ant. ii. 129 (1900), excl. es Britton, n. 
4 186 (1901): Small, Fl. 197 (1903) and Man. 184 (1933); Robinson 
& Fernald in Gray, Man. ed. 7: 199, fig. 315 (1908); Britton, 
Mem. Soc. Cubana Hist. Nat. ii. i98 (1916); Britton & Wilson, 
Sci. Surv. Porto Rico and Virgin Isl. v. 104 (1923); non Elliott. 
R. gracilis sensu Grisebach, Cat. PI. Cub, 243 (1866), in part; 
C. Wright in Sauvalle, Anal. Acad. Ci. Habana, viii. 84 (1871) 
and Fl. Cub. 179 (1873), i in part; non (Swartz) Vahl. R. Torrey- 
ana sensu Grisebach, Cat. Pl. Cub. 243 (1866); C. Wright in 
Sauvalle, Anal. Acad. Ci. Habana, viii. 84 (1871) and Fl. Cub. 
179 (1873); non Gray. Schoenus cymosus Muhlenberg, Cat. 
(1818), nomen nudum, and Descrip. Gram. 8 (1817); non Willde- 
now. Phaeoc ephalum cymosum ae se, Am. Midland Nat. vi. 
201 (1920). Dichromena cymosa Mcbride: Field Mus. Pub. Bot. 
xi. 5 (1931).—Low open places on the Coastal Plain from New 


246 Rhodora [JUNE 


Jersey to northern South Carolina and inland to the mountains 
of the Carolinas, Georgia and Alabama; westward through 
Tennessee, with a few stations on the southern shores of Lake 
Michigan and Lake Ontario; also in coastal ranges in northern 
California; distributed along the Mississippi southward on the 
lower drainage of its western tributaries, the Missouri, Arkansas, 
and Red Rivers; scattered in eastern Texas, Louisiana and the 
coastal counties ‘of Mississippi and Alabama; also on the penin- 
sula of Florida, the Greater Antilles, the island of Dominica and 
in Central America. The citation of representative specimens is 
omitted because this common and widely spread variety has been 
so well known under the misapplied name of “ R. cymosa Ell.” 
However, since the nomenclatural history of the plant has been 
So confused it seems best to designate a type, and that is cited 
below. Vrrcrnia: dry pinelands about 4 miles northwest of 
Waverly, Sussex Co., July 26, 1936, Fernald & Long, no. 6070 
(G, TYPE; P, 1soTy PE). 

In 1798 Willdenow published in his revision of the Cyperaceae 
for the Species Plantarum a new species, Schoenus cymosus. 
The abbreviation v. s. (vidi siceas)' appended to his description 
indicates that at that time a dried specimen of the same existed 
in his herbarium. 

Forty-one years later Kunth relegated S. cymosus Willd. to 
the synonymy of Scirpus polyphyllus Vahl. In so doing he added 
to his citation of Willdenow’s Species Plantarum, i. page 265 the 
herbarium number 1247 of the Willdenow type. 

Recently Kikenthal,? in his treatment of Schoenus as part of 
his projected monograph of the Rhynchosporoideae, lists Schoenus 
cymosus Willd. as equivalent to Rhynchospora cymosa Ell. 

The original description of Willdenow does not mention either 
style or achene. Due to the present international situation, it is 
impossible to confirm or refute by an examination of these details 
in the type-specimen itself either the diagnosis of Kunth or the 
more commonly held opinion of Kiikenthal. However, not only 
was Kunth personally in a position to study the Willdenow type, 
but his citation of the not otherwise published herbarium number 
1247 seems to indicate that he availed himself of this opportunity. 
Also, as will be indicated in the following paragraph, Kunth was 
well aware of the confusion surrounding Elliott’s application of 
the specific name a to Rhynchospora. Kiikenthal, on the 

1 Willdenow, Sp. Pl. (1797). 

* Fedde, Rep. Spec. a xliv. 187 (1938). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 247 


contrary, gives no indication that he has as yet either personally 
examined the Willdenow type or studied in detail Elliott’s 
description of R. cymosa. I am, therefore, assuming S. cymosus 
Willd. to be correctly placed by Kunth in the synonymy of 
Scirpus polyphyllus. 

In 1813 the name Schoenus cymosus appears again, this time 
in the list which constitutes Muhlenberg’s Catalogue. Muhlen- 
berg’s description of his new species, a necessity to the valid 
publication of every new species, did not follow until the publi- 
cation of his Descriptio Uberior Graminum, 1817. When the 
description appeared, however, it unquestionably applied S. 
cymosus Muhl. to the plant which now passes under the name of 
Rk. cymosa. Technically, since the Willdenow species and the 
Muhlenberg species are not identical, Muhlenberg’s name be- 
comes a later homonym of the earlier, and, as such, is illegitimate 
under Schoenus according to the International Rules. 

In 1816, while the status of S. cymosus Muhl. was that of a 
nomen nudum, Elliott published the first pages of his Sketch of 
the Botany of South Carolina and Georgia. Here he described 
in detail a Rhynchospora cymosa which he supposed to be S. 
cymosus of Muhlenberg, but, in reality, Muhlenberg’s species 
was not validly published until a year later, when it was accom- 


panied by its description. Elliott made, therefore, not a new 


combination, but a new species. Consequently it is imperative 
to determine the identity of his specimen which is obviously the 
type of Rhynchospora cymosa. 

Kunth in his Enumeratio Plantarum assigns R. cymosa 
Elliott to R. distans (Michx.) Vahl, basing his determination upon 
a specimen ‘a Beyrichio misso.”” Both Torrey and Gray had 
previously suspected that Elliott’s description did not ayply to 
the plant which they knew as R. cymosa and which they errone- 
ously attributed to Willdenow. 

To come at the bottom of this confusion, Prof. M. L. F penile 
during the spring of 1938, kindly examined the Elliott tyre for 
me, and he states that, although immature, it is undoubtedly 
R. glomerata (Michx.) Vahl. The name R. cymosa, then, since 
it has no connection with S. cymosus Willd., properly dates 
from Elliott, belongs in the synonymy of R. glomerata, and cannot 
be rightfully used to designate any other entity. 


ba] Gee ee, a eer 


248 Rhodora [JUNE 


This conclusion necessitates for the plant which was the basis 
of Muhlenberg’s S. cymosus and which has until the present 
passed as ?. cymosa, another name. Chapman in 1860 set apart 
and described the smaller R. cymosa, var. globularis. Small in 
1933 raised this variety to specific rank. Since in my judgment 
no boundary exists which specifically delimits the larger from 
the smaller plant, I am accepting Chapman’s interpretation 
rather than that of Small. R. globularis, therefore, becomes the 
correct specific name. Var. typica is the small rlant originally 
set apart by Chapman as var. globularis, and to the plant for- 
merly treated as R. cymosa, I am giving the name R. globularis 
var. recognita. 

After this discussion of R. cymosa was yut into final form, 
Professor Fernald received, through the kindness of Professor 
Diels, Director of the Botanical Museum at Berlin, a photo- 
graph of Schoenus cymosus, no. 1247, in the Willdenow Herbari- 
um. The specimen! was clearly a young state of Scirpus poly- 
phyllus, as stated in Kunth’s annotation on the right-hand side 
of the sheet. The specimen-cover bore in the left-hand corner 
the no. 1247, and on the right, in Willdenow’s hand, ‘‘Schoenus 
cymosus Sp. Pl. Habitat in America boreali.’”? This was accom- 
panied by a diagnosis of Schoenus umbellatus copied from Walter,’ 
in which the name S. umbellatus has been replaced by that of 
Scirpus polyphyllus. 

_59c. Var. pinetorum (Small), stat. nov. Leaves 1.5-2 mm. 
wide: clums obtusely trigonous, slender, erect, becoming flexu- 
ous, 2.7-6.9 dm. in height: inflorescence 1-4 weak glomerulose 
cymes: bracts inconspicuous: spikelets nearly globose, 2-flowered, 
1-fruited, compact, 2.5-3 mm. long: scales mucronulate or obtuse: 
bristles 6, not exceeding 24 the height of the achene: achene 1.4 
mm. wide, 1.3~-1.4 mm. long; the pale flat surface covered with 
a dark nearly isodiametric reticulation: tubercle triangular, 0.4— 
0.5 mm. high. Puatre 831, rig. 5A; Map 71.—R. pinetorum 
Britton & Small, in Small, Man. 183, 1503 (1933).—Moist areas 
in pinelands and savannas, the Peninsula of Florida, west along 
the coast to Louisiana; also in western Cuba and Jamaica. 
FLormA: moist pine barrens near Jacksonville, Duval Co. 
Curtiss, no. 4871 (G); prairie near St. Johns River, Osceola Co., 
McFarlin, no. 4908 (CU); in a low pineland, Aripeka, Pasco Co., 
O'Neill, no. 2611 (CU, NY); Manavista, Manatee Co., Tracy, 


' Now, with the complete herbarium at Berlin, presumably lost. 
2 Walter, FI. Car. 70 (1788). 


1944) Gale,—Rhynchospora, Section Eurhynchospora 249 


no. 6991 (G); pinelands west of Punta Gorda, DeSoto Co., 
Small, Mosier & DeWinkler, no. 10912 (NY, Type of R. pine- 
torum Britton & Small); in moist sandy ditch along seadiade: 
‘ Rialto, Lee Co., Moldenke, no. 1011 (D, NY); glade, Cutter & 
Black Pt., Dade Co., Small & Carter, no. 885 (P); swampy 
thicket, Washington Co., May, Curtiss, no. 3149 (NY). Lovtst- 
awa: low prairies, Jennings, Jefferson Davis Parish, Palmer, no. 
( 


Pinar del Rio and Caloma, Pinar del Rio, Britton, Britton & 
Cowell, no. 10084 (NY). JAMAICA! along rivulets at ’Moneague, 
1850, Alexander (G). 

R. pinetorum Small is superficially identical with the weaker 
state of R. globularis, var. recognita found in the western States 
and the Greater Antilles. It has the same attenuated appear- 
ance, the same weak cymes, turgid spikelets, inconspicuous 
bracts and nearly orbicular short scales. Even its achene is 
similar in shape, size and tubercle. Its only claim to specific rank 
depends upon the surface sculpturing of the achene. This con- 
sists of a brown nearly isodiametric reticulation over a flat, 
usually pale background. R. globularis and the var. recognita 
have, by contrast, an elongate reticulation with the shorter 
sides of the alveoli accentuated so as to form transverse ridges on 
the surface of the achene. A study of the whole globularis- 
complex in Florida and Cuba, however, reveals that in this 
region transitional stages between isodiametric and elongate 
pitting with a ridged surface occur. Such are the achenes from 
the collection of Tracy, no. 7001, from Florida, and of Curtiss, 
December, 1903, from the Isle of Pines. It seems best, therefore, 
to reduce Small’s entity to the status of a variety. 


(To be continued) 


1944] Gale,—Rhynchospora, Section Eurhynchospora 255 


RHYNCHOSPORA, SECTION EURHYNCHOSPORA, IN 
CANADA, THE UNITED STATES AND THE 
WEST INDIES 


SHIRLEY GALE 
(Continued from page 249) 


Series 11. Caducae, ser. nov. Cymis decompositis saepissime 
valde fasciculatis patentibus corymbiformibus: spiculis 1-10 
(semper fere 4—5)-carpis: squamis laxe imbricatis: setis 1-6 
antrorse serrulatis saepissime validis (R. perplera et R. micro- 
carpa exceptis): achaenio elliptico-obovato vel late obovato vel 
suborbiculari saepissime rugoso cancellato vel striato, obscure 
marginato. 

Growing in damp depressions, swamps, pond-margins and 
shallow water of the pine barrens, savannas and open woodland 
of the Coastal Plain; infrequent inland; also in the West Indies. 
Caespitose: leaves 1 ‘(rarely)-7. 5 mm. wide: culms of well devel- 
oped specimens approximately 1 m. high, usually erect, becoming 
flexuous toward the summit, less often weakly ascending: cymes 
decompound, those of most species corymbiform, densely 
fasciculate, less often weakly spreading and with few spikelets 
or forming a globose net in R. miliacea: spikelets oe 
1—mostly 4-5 (less often 6-10) achenes: scales loosely imbric 
bristles "udually well developed (1-3 and rudimentary in wt 
perplexa), antrorsely serrulate: achenes ellivinababnte to broad- 
ly obovate or suborbicular in outline, usually ridged, always 
cancellate or striate, obscurely marginate: t tubercle basally 
deltoid.— Rhynchospora V. Glomeratae Small, Man. 175 (1933), 
in part. ah faiier snk Series B. Diplo styleae, Sect. 5. Glaucae 


Key To Species IN SERIES CADUCAE 


a. Bristles equalling or exceeding the achene... .b. 

b. Achene slenderly ellipsoid-obovoid, 2-2.2 mm. in length 
R. inexpansa. 
b. Achene bersasceaiied to broadly obovoid or subglobose, less 

than 1.8 . in length.... 
c. Branchlets of the 6-9 cymes stiffly divaricate or reflexed, 
giving the ais nesta of a se web; achene pale se 
Fellow heowi yk Pia as a eee 1. R. miliacea. 
c. Ecanckiats of the 2-5 cymes ascending to spreading, if 
divaricate never stiffly so; achene castan pee & 
d, y anton broadly cheeate to eabbetiedlar = ee 
n above, comp ae 2 
wide or wider, 1.4 mm. long or rang ee 
e. Achenes striate, castaneous, 1.4-1.6 mm. wide, i r= 
.7 mm, long, with a evi, groies terete stipe 
which is 0.3-0.4 mm. in length; stamens conspicu- 
ously anpeceeenk eat Cue eels beer reas 62. R. odorata 


256 Rhodora : [JULY 


e. Achenes Lard cancellate, 1.2-1.6 mm. wide, 1.4— 
m. long, with a short thick inconspicuous 
ent....63. FR. caduca. 
d. Achenes slenderly ronan or, if dante to suborbicu- 
ar in outline, small, gradually biconvex and not 
ee reey swollen “above, 0.8-1.2 mm. wide, 
1-1.4 


ong.... 

f. Cymes loose, spre eadin ng, the lax mango branchlets 
widely divergent; spikelets few... . 

g. Bristles well exceeding the “wuberele; tubercle 


deltoid-attenuate with setose margins........ 53, dt. mnecla. 
g. Bristles equalling the achene taherdis deltoid, 
hat decurrent, the eres bare : se decurrens. 


ridges: bristles irregularly spreading... .66. R. schoenoides. 
h. ——— moderately biconvex, the alveoli subiso- 
diametric to broadly oblong, rugulose to only 
Slightly ridged; bristles amially aired erect and 
6 


closely connivent around the achene.. . R. microcarpa. 
a. Bristles shorter than the achene or failing... .7 ; 
z. Achene strongly flattened, prominently ced, aa ede Ne 


j. Bristles 6, not exceeding the achene; transverse ridges 
of the achene approximately 12, closely spaced; tuber- 


: cle decurrent; spi genes 4mm. long As FOE Bee 68. Rk. Torreyana. 

j. Bristles none or 1-3 ru c pinks transverse ridges of the 
achene less than 8, wid ery separated. os ey 69. R. perplexa. 

zt, Achene biconvex, slightly if at all ieee. the alveoli sub- 
isodiametric to broadly ODIONG ss ssh os ee ea e058 67. R. microcarpa. 
60. R. 1nexpansa (Michx.) Vahl. Fg any often forming 
stools of several—15 culms: leaves flat, 2-3.5 mm. wide; margins 
finely serrulate; tips triquetrous: culms shelisel triangular, 
becoming slender, flexuous, 0.3-1.2 m. high: inflorescence 3- 


strictly elongated decompound fasciculate cymes, 0.6-1 dm. in 
length, the filiform arching branches approximate; lateral cymes 
on exserted peduncles: spikelets fusiform, sessile or setae 
strongly ascending, 2—5-flowered, 1—4-fruited, 4.5-6 . long: 
scales castaneous, acute or aristulate, caducous: bristles 6, up- 
wardly hispidulous, capillary, erect, once again as high as the 
tubercle: achene slenderly Slipaotdoboveid, narrowing toward 
the base, extremely flattened, transversely ridged, castaneous, 
0.8-1 mm. wide, 2-2.2 mm. long: tubercle ee, compressed, 
0.9-1.2 mm. long, pale, with thickly setose margin LATE 

Frias. 1A and 1B; Mar 75.—Enum. ii. 232 (1806): Elliott, Sk. 
Bot. S. Car. and Ga. i. 61 (1816); Gray, Ann. Lye. N. Y. iii. 200, 
pl. 6, fig. 6 (1835) and Man. ed. 2, 505 (1856) ; Pear aty Fl. So. 
U.S. 525 5 (1 ~~, ; Britton & Brown , Ll. FI. i. 280, fig. 660 (1896) ; 
Britton, Man. 186 (1901); Small, Fl. 198 (1903) and Man. 186 
(1933); eatin & Fernald in Gray, Man. ed. 7: 200, fig. 318 
(1908). Schoenus inexpansus abies FI. Bor. -Am, i. 35 (1803) ; 


1944] Gale,—Rhynchospora, Section Eurhynchospora 257 


Muhlenberg, Descrip. Gram. 9 (1817).  Phaeocephalum inex- 
pansum House, Am. Midland Nat. vi. 202 (1920)——Common in 
low or exsiccated ground in open areas of pinelands on the Coastal 
Plain from southeastern Virginia to Georgia and west to eastern 
Texas; inland localities in northern Alabama, northwestern 
Arkansas and along the Red River. Citation of specimens of 
this unique and well known species is not considered necessary in 
this paper. 

61. R. mrtracea (Lam.) Gray. Stoloniferous: leaves narrowly 
linear, 6-7.5 mm. wide, erect, flat, carinate; the keel and margins 
scabrous to smooth; tip triquetrous: culms 3-angled, leafy, 
slightly arching or more commonly stiffly erect, 0.9-1.4 m. high: 
cymes 6-9, decompound, the long wiry capillary branchlets 
stiffly divaricate or often reflexed, giving to the cyme a loose, 
web-like appearance, 0.7—1 dm. wide: lateral cymes on subex- 
serted peduncles: spikelets ovoid, turg id, 3-4 mm. long, distant 
on elongated slender pedicels, rag etl 3-10-fruited: scales 
aristulate, loosely imbricate, promptly caducous exposing a 
‘‘spikelet’’ of naked achenes: bristles 6, fragile, spreading, up- 
wardly serrulate, exceeding the tubercle: achene broadly obovoid, 
biconvex, ae ridged to rugulose, longitudinally striate, 
pale, 0.9-1. mm. wide, 1-1.38 mm. long: tubercle depressed- 
conic, often ares 0.2-0.4 mm. long. Pair 833, FIGS. 2A 

and OB: Map 74.—Ann. Lye. N. Y. iii. 198, pl. 6, fg. 4 pees 
Cheon Fl. So. U.S. 526 (1860); Grisebach, Cat. Pl. Cub. 243 
(1866); C. Wright in Saye Anal. Acad. Ci. eae vill. 84 
(1871) and FI. Cub. 1 179 (1873); Small, Fl. 198 (1903) and Man. 
186 (1933); Britton, Mem. Soe. Cubana Hist. Nat. ii. 199 (1916); 
Kiikenthal, Fedde Rep. Spec. Nov. xxiii. 211 (1926). Schoenus 
miliaceus Lamarck, Ill. Gen. i. 187 (1791). &. sparsa Vahl, 


sparsus Michaux, Fl. Bor. EO o 35 Maa. Muhlenberg, 
Descrip. Gram. 7 (1817). Phaeocephalum miliaceum (mispelled 
mailaceum) House, Am. Midland Nat. vi. 202 (1920).—Often 
standing in water of swamps of the Coastal Plain from south- 
eastern Virginia and North Carolina (rarely) southward to the 
Florida Peninsula, and west to Louisiana; also in the western 
provinces of Cuba and in eastern Hispaniola. Virainta: Read 

, as ‘‘Schoenus sparsus”’), presumably from Norfolk Co. 
Nortu Carona: swamp at Newport, Carteret Co., Godfrey, 
no. 4412 (D, G); peaty grass-sedge savanna at Carolina Beach, 
New Hanover €o.. Godfrey, no. 4684 (G, NC); edge of swamp, 
Southport, Brunswick Co., Jan. 28, 1922, Bartram (G, P). 

ouTH CAROLINA: swamp, 1% miles west of Andrews, George- 
town, Georgetown Co., Godfrey & Tryon, no. 541 (G, NY); 


258 Rhodora [JuLy 


wet places along Santee Canal, Berkeley Co., Ravenel (G). 
CAROLINA: in umbrosis Carolinae, Michaux Herb. (G, type- 
photo of Schoenus sparsus); east Carolina, D. Fraser in Herb. 
Leonard (G, TypE-pHoto of Schoenus miliaceus). GEORGIA: 
shaded pool of clear cold water, Leslie, Sumter Co., Harper, no. 
1017 (G, NY, US); semi-calcareous swamp of small creek about 
Y mile east of Cedar Springs, Early Co., Harper, no. 3635 (G, 
P, US). FLoripA: swamps aes Jacksonville, Duval Co., 
Curtiss, no dre (CU; DG, NY, PF, US); Devils Mill Hopper, 
lime-sink near Gainesville, Alachua Co., O’ Neill, no. 586 (US); 
rooting under water, Silver Springs, Marion Co., O’ Neill, no. 
2606 (CU); damp places, Ormond, Volusia Co., Fuller (G); 
Eustis, Lake Co., Nash, no. 1845 (G, NY, P, US); in mud of bog 
in open woods near Oviedo, Seminole Co., Correll, no. 6361 (CU, 
D, G); in moist woodland, south of Aripeka, Hernando Co. ra 
Moldenke, no. 1065 (D, NY): rooting under water in a spring, 
pasture hammock, Lake Jovita, Paseo Co., O'Neill, no. 1060 
U, US); swamp, between Shingle and Bonnet Creek, Kissim- 
mee, Osceola Co., Mar. 13, 1938, Singletary (D); in a swamp, 
Polk City, Polk Co., O'Neill, no. 7682 (CU); sand-barren swamps, 
Tampa, Hillsborough Co., Apr. 11, 1923, Churchill (G); oe 
Okaloacoochee Slough, Big Cypress, Lee Co., Small, no. 8311 
(NY); in everglades near Camp Long View, Dade Co., Small & 
Wilson, no. 1644 (NY); bogs and deep miry ‘places, Apalachicola, 
Franklin Co. , Chapman in Biltmore Herb., no. 231a (G, NY, US). 
ALABAMA: deep loa springy places, " eastern shore, Mobile 
Bay, Point Clear, Baldwin Co., June 10, 1879, Mohr (US). 
LovisiANA: abundant in cypress-tupelo swamp, Rone Ferry on 
Tickjaw River, Livingston Parish, Brown, no. 6573 (La); New 
Orleans, 1832, Drummond (G); sandy ditch near swamp in 


Zapata, Santa Clara, Acufia, no. 4305 (NY); in coastal swamps, 
in a wood of Peralta Estate, "Batabano, Havana, Leén, no. 14195 
Sher Wright, no. 3788 (G, US). Hispanroua: Macary in 

tang Soe Marigot, Massif de la Selle, Haiti, Ekman, no. 
5978 (NY, U; 

2. Fes see ee C. Wright ex Griseb. With short stolons: 
leaves 3.5-6 mm. wide, arching, carinate, smooth, becoming 
triquetrous at the tip, with finely serrulate angles: culms stocky, 
becoming flexuous at the summit, 3-angled, smooth, 0.7—1.8 m. 
high: cymes 3-4, decompound, "densely fasciculate, 3-4 cm. 
wide, 3-7.5 cm. long; branchlets mainly arched-ascending: 
spikelets ovoid, 3-11-flowered, 1-7-fruited, 6-9 mm. long: 
scales acute-aristate, loosely imbricate, ferruginous, caducous: 


1944] Gale,—Rhynchospora, Section Eurhynchospora 259 


Range of 77, RHyNCHOSPORA MIXTA; 78, R. DECURRENS; 79, R. SCHOENOIDES; 
0, RR: TORREYANA; 81, R. PERPLEXA; 82, He MICROCARPA. 


stamens conspicuously marcescent: bristles 6, irregularly ascend- 
ing to spreading, slenderly attenuate, usually well exceeding the 
tubercle, upwardly serrulate, 3.2-4.2 mm. long: achene orbicular 
in outline, strongly biconvex, transversely ridged and strongly 
marked by longitudinal striae, castaneous, 1.4-1.6 mm. wide, 
14-4, A mm. long, with a conspicuous persistent stipe, 0. 3-0.4 
mm. long: tubercle deltoid-compressed, somewhat peace 
usually dark with setose margins. PLaTe 833, Fries. 3A and 

Map 73.—Cat. Pl. Cub. 242 (1866); C. Wri ght in Sauvalle 
Anal. Acad. Ci. Habana, viii. 83 (1871) and FI. Cub. 179 (1873). 
R. stipitata Chapman Fi. So. U. S. ed. 2: 660 (1883); Hemsley, 
Rep. Sci. Res. Voy. Challenger, Bot. i. 76, pl. 10 (1885); Clarke 
in Urban, Symb. Ant. ii. 129 (1900); Sinall Fl. 198 (1903) and 
Man. 186 (1933); Britton, FI. Bermuda, 53, fig. 82 (1918). 
Rk. Marisculus sensu Clarke in Urban, Symb. Ant. ii. 132 (1900), 
in part. &. Marisculus sensu Britton, Mem. Soe. Cubana Hist. 
Nat. ii. 200 (1916), in part, excl. syn. R. jubata Liebm. R. Maris- 
culus sensu Britton & Millspaugh, Bahama Fl. 56 (1920); non 
Lindl. et Nees in Mart. R. caduca sensu Kiikenthal, Fedde Rep. 
pec. Nov. xxiii. 210 (1926) and xxxii. 77 (1933); non Ell. 
Phaeocephalum stipitatum House, Am. Midland Nat. vi. 202 
(1920). —Swamps of the Coastal Plain a North Carolina 
(one collection from Carteret Co.) to the Florida Peninsula, 
Bermuda, New Providence of the Bahamas and the Greater 
Antilles. Norra Caro.ina: sand banks near Beaufort, Carteret 
Co., Lewis, no. 76 (NY). Gerorara: Baldwin (P). Forma: 
low rich places near Jacksonville, Duval Co., Curtiss, no. 3141 
(P); swamp near Mosquito Inlet, Volusia Co., Curtiss, no. 3174 


260 Rhodora [JoLy 


Guane, Pinar del Rio, Dec. 23, 1863?!, Wright, no. 3394 (G); in 


Clara, Cuesta, no. 764 (NY); al norte de la Bahia de Cochinos, 
Santa Clara, Leén & Loustalot, no. 9506 (NY). Jamaica: 
marsh, 1 mile west of Black River, Cornwall, Britton, no. 1356 
(NY); border of Great Morass, Negril and vicinity, Cornwall, 
Britton & Hollick, no. 2116 (NY). H1spanroua: coastal swamp 
at Carbarete, Tosua, prov. Puerto Plata, Cordillera Septen- 
trional, Santo Domingo, Ekman, no. 14535 (G, NY, US). PuERTo 
Rico: Point Congrejos, Stevenson, no. 1706 (NY, US). 

63. R. capuca Ell. Spreading by means of short stolons, 
occasionally forming dense stools: leaves 4-7 mm. wide, flat, 
slightly carinate, mainly ascending; tips triquetrous, with minute- 
ly scabrous angles: culms robust, acutely 3-angled, erect, becom- 
ing flexuous at the summit, 0.7-1.2 m. high: cymes 3-5, decom- 
pound, strict, 0.5-1.2 dm. long, usually densely fasciculate and 
often arching, but attenuated specimens occurring with scattered 
spikelets on weak, spreading branchlets: spikelets ovoid, fas- 
ciculate and sessile or single and slenderly pedicellate, 3-6- 
flowered, 2-5-fruited, 4-4.5 mm. long: scales acute to aristulate, 
caducous, dark brown, loosely imbricate: bristles 6, brittle, well 
exceeding the tubercle, upwardly hispidulous: achene obovate to 
rotund in outline, strongly umbonate, 1.2-1.6 mm. wide, 1.4-1.6 
mm. long; entire surface cancellate with the horizontal walls 
of the oblong alveoli contiguous, accentuated and forming 
horizontal ridges: tubercle deltoid, compressed and slightly 

epressed, usually pale, setose, 0.6-0.8 mm. in height. PLATE 

‘ See Underwood, Bull. Torr. Bot. Cl. xxxii, 297 (1905). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 261 


833, rics. 4A and 4B; nage 76. woe es . a and ie i. 

(1816); melts Ann. Lye 6. fig. 5 (iss 
Chapman, FI. So. U. S sn (i860); Small Pi 197 Ane and 
Man. 185 Fess R. patula Gray, Ann. Lye. N. Y. iii. 201, pl. 6 

fig. 8 (1835), for the most part. Phaeocephalum pn House, 
Am. Midland Nat. vi. 201 (1920).—Frequent in meadows and 
marshes on the Coastal Plain from southeastern Virginia south 
to Florida and west to eastern Texas; known inland only from a 
few localities in the mountains of Georgia and Alabama and in 
northwestern Arkansas. Virainia: border of damp sandy pine 
woods east of Gloucester, Gloucester Co., Fernald & Long, no. 

61 


border of brackish to fresh marsh along Back Bay at eastern 
margin of Long Island, Princess Anne Co., Fernald & Long, no. 
10555 (G); boggy swale by Northwest River, near Northwest, 
Norfolk k Co., Fernald, Griscom & Long, no. “4578 (G); swaley 


t 
Beach), Southampton Co., Fernald & Long, no. 10148 (G, P); 
exsiccated argillaceous pineland, about 2 miles east of Stony 
Creek, Sussex Co., Fernald & Long, no. 8999 (G, P). Norra 
Carouina: in bogey soil near Elizabeth City, Pasquotank Co., 
soley, no. 2110 (D); marsh at Sligo, Currituck Co., Godfrey, no. 
283 (G dfrey, no. 5340 


262 Rhodora [JULY 


Johns Co., June 7, 1940, West & Arnold (CU); St. Vincent Island, 
Franklin Co., McAtee, no. 1820a (US); in low pineland, Lake 
City, Columbus Co., O'Neill, no. 7674 (CU); wet hammock, 
vicinity of Eustis, Lake Co., Nash, no. 139 (G); in edge of low 
woods near Mabel, Sumter Co., Curtiss, no. 6631 (G, P); swamps, 
Myers, Lee Co., Hitchcock, no. 422 (G); moist open ground, 1 
mile west of Arcadia, DeSoto Co., Blomquist, no. 8958 (G, D); 
near Tallahassee, Leon Co., Berg (NY); Herb. Baldw. (NY, 
annotated as R. patula by Gray). ALAaBamMa: Auburn, Lee Co., 
Earle, no. 859 (NY); pine woods, Ball Play, Etowah Co., Mohr 
(US). Musstsstpr1: Saratoga Co., Tracy, no. 8464 (G). ARKAN- 
sas: Benton Co., Plank, no. 17 (NY). Lovistana: in prairies, 


Rouge Parish, Trotter & Chilton, no. 42 (La); low prairies, Jen- 
nings, Jefferson Davis Parish, Palmer, no. 7625 (CA, Mo, P). 
Texas: Palestine, Anderson Co., Apr. 19, 1895, Plank (NY); 


Chambers Co., Cory, no. 22404 (G); Cypress City, Harris Co., 
Boll, no. 884 (G); Caney Post Office, Brazoria Co., Bechdolt, no. 
4 (G); banks, Hempstead, Waller Co., Hall, no. 711 (G, Mo); 
Burnet, Burnet Co., Sept. 14, 1892, Plank (NY); prairie near 
Indianola, Calhoun Co., Ravenel, no. 159 (NY); coastal prairie, 
Midfields, Matagorda Co., Tharp, no. 2139 (US). 

64. R. mixta Britt. ex Small. Stoloniferous: leaves 3-4 mm. 
wide, flat, subcarinate, becoming triquetrous at the tip; margins 
minutely scabrous: culms triangular, slender, becoming filiform 
and arched toward the summit, approximately 1 m. in height: 
cymes 4-6, decompound, fasciculate, 2.5-5.5 cm. wide, often 
extremely lax and flexuous in appearance due to the attenuated 
pedicels which may be strongly divergent; axillary cymes on 
exserted peduncles: spikelets ovoid, 2 (rarely)—10-flowered, 1 
(rarely)—10-fruited, 4-6 mm. in length, commonly distant: 
scales mucronulate or acute, loosely imbricate, characteristically 
caducous, castaneous: bristles 6-8, extremely slender, brittle, 
well exceeding the tubercle, upwardly hispidulous, 2.2-2.8 mm 
long: achene slightly compressed, otherwise slenderly obovoid, 
0.8-0.9 mm. wide, 1.3-1.4 mm. long; surface pale and dull but 
covered with a brown isodiametric reticulation, the transverse 
walls of which are barely if at all elevated: tubercle deltoid- 
attenuate, 0.4-0.9 mm. long, compressed, with a conspicuously 
setose margin. PLATE 832, rics. 4A and 4B; Map 77.—F. 197, 
1328 (1903) and Man. 186 (1933). R. prolifera Small, Fl. 198, 
1328 (1903) and Man. 186 (1933). Phaeocephalum mixtum House, 
Am. Midland Nat. vi. 202 (1920). Phaeocephalum proliferum 
House, Am. Midland Nat. vi. 202 (1920).—Swampy woodlands 


1944] Gale,—Rhynchospora, Section Eurhynchospora 263 


of Coastal Plain from North Carolina to northern Florida and 
west to eastern Texas. Norra Carouina: Elizabeth City, 
Pasquotank Co., Kearney, no. 1993 (US): ditch, Beaufort Co., 
Blomquist, no. 5614 (D); stream-margin, Holden Beach, Bruns- 
wick Co., Blomquist, no. 5614 (P). Sours CAROLINA: wet 
swampy woodland, 12 miles north a Georgetown, Georgetown 
Co., Godfrey & Tryon, no. 111 (D , P). Grorata: moist 
shaded ar place in Ogeechee River swamp, Burke Co., Harper, 
no. 768 (G, NY, US); bank of Ocmulgee River, Hawkins- 
ville, aon Co. , Harper ssh 1385 (G, NY, US); Marshallville, 
acon Co., Earle, no. 3 (NY, TYpPe); rich woods i in south- 
eastern part of Sumter git Harper, no. 490 (N ae moist 
sandy places, Leslie, Sumter Co., Harper, no. 408 (N Y, type o 
Rk. prolifera Small; US, isotype) ; swamp of Chickasawhatchee 
Creek at Johnson Bridge Ne) Elmodel, Baker Co., Eyles, no. 
7059 (Hermann Herb.). FLORIDA: wet sandy soil near Perry, 
Taylor Co., Palmer, no. 27285 (G); low ground, Marianna, 
Jackson Co., May 26, 1940, Sargent (Sargent Herb.); river 
swamps, Apalachicola, Franklin Co. ., Chapman in Biltmore 
Herb., no. 4471 (G, NY, US); swamps, Walton Co., 1885, 
Curtiss (NY); swampy woods along Yellow River near Milligan, 
Santa Rosa Co., Curtiss, no. 6854 (G; US, in part). ALABAMA: 
palmetto sw ramp, vicinity of Auburn, Lee Co:, Pollard & Mazon, 
no. 42 (G, NY, US). Mrsstssterr: Ocean Springs, Jackson Co., 
Tracy, no. 114 (NY). Lovrstana: vicinity of Covington, St. 
Dee Parish, Arsene, no. 12307 (Ue) mixed a and pal- 


Baton Rouge Parish, May 19, 1874, Joor (us Texas: in bed 
of brook in high mixed forest, 5 miles south of Nacogdoches on 
Lufkin Road, Nacogdoches Co., Lundell & Lundell, no. 9643 
(CU); 1883, N Serle no. 28 (NY US); East Texas, Wright (G). 

f. mixta has a culm which, in common with so many members 
of this series, varies in degree of flexuosity from upright, with a 
curving distal internode, to weak, attenuated and nearly pro- 
cumbent. Specimens of the latter extreme have open fascicles 
with long filiform divergent branchlets; those with the greatest 
stiffness, however, have flexuous but not open fascicles, the 
branchlets of which are often shorter, usually ascending, causing 
a closer approximation of the spikelets. As in the closely related 
fk. miliacea, the number of achenes in a spikelet is very variable. 
I consider that Small, in designating those specimens which have 
spikelets bearing from 8-10 achenes as R. prolifera, set up an 
artificial boundary which alone delimited that species from R. 
mixta of Britton. 


264 Rhodora [JULY 


R. DECURRENS Chapman. Caespitose: leaves flat, 3 m 
wide, smooth, soft, with 3- angled setaceous tips: culms obtuisely 
trigonous, smooth, slender, becoming flexuous, leafy, approxi- 
mately 0.8-1.1 m. high: cymes 4-5, decompoun d, loose, some- 
what okie. 2-5 em. wide, the filiform branchlets lax or 
drooping, bearing a few scattered or somewhat congested spike- 
lets; lateral cymes distant, on exserted peduncles: spikelets 
rotundly ovoid, 3 mm. long, 3—4-flowered, 2—3-fruited, scattered 
or somewhat congested on capillary pedicels: scales mucronulate 
or muticous, chestnut-brown, caducous: bristles 6, extremely 
tenuous, fragile, upwardly hispidulous, equalling the achene in 
height: ‘achene obovoid, slightly biconvex, bright chestnut, 0.9 
mm. wide, 1.3 mm long; the surface pitted, tending to become 
transversely rugulose: tubercle deltoid, compressed, somewhat 
decurrent, not setose. PLatTEe 832, rigs. 1A and 1B; Mar 78.— 
Fl. So. U. 8. 525 (1860); Small, FI. 198 (1903) and Man 185 
(1933). Phaeocephalum decurrens House, Am. Midland Nat. vi. 
201 (1920).—Swamps and marshy stream-banks, Columbus Co., 

orth Carolina (one collection) and Florida. Norra CAROLINA: 
edge of swamp, 1 mile east of Old Dock, Columbus Co., Blom- 
quist & Correll, no. 9444 (D). FiLorripa: in swamp, Callahan, 
Nassau Co., O'Neill, no. 6017 (CU); south branch, St. Mary’s 
River, summer, 1885, Curtiss (NY); marshy banks of streams, 
Dead ‘Lakes, Calhoun Co., Mohr. no. 68 (US) 
lakes and rivers, Wewahitchka, Calhoun Co., Chapman in Bilt- 


mor ; . 
Heinen Rie damp thickets, bank of Chittahachee River, 

onro , June 25, 1880, Mohr (US, in part); swamps, western 
Florida, fie 20, 1880, Mohr (NY). 

R. decurrens is most closely related to R. mixta Britt. This is 
at once apparent in the habit which is indistinguishable from 
that of R. mixta, for the plant has the same lax and drooping 
aspect with the typical flat soft leaves, and the loose cymes with 
their spreading or drooping filiform branchlets. The distinction 
between the two species depends upon achenial characters. The 
six bristles surrounding the customarily pale chestnut achene of 
R. mixta exceed the deltoid-attenuate tubercle which is 0.4 
(rarely)—0.9 mm. high and marginally setose. The achene of 
R. decurrens, however, while similar to that of R. mixta in pitting 
and general outline, is a brilliant chestnut, surrounded by six 
bristles in height not equalling the tubercle, which is short (not 
exceeding 3 mm. long), broadly deltoid, and without a trace of 
marginal setae. 


1944] Gale,—Rhynchospora, Section Eurhynchospora 265 


66. R. scHorNnorpEs (Ell.) Wood. Coarsely caespitose: 
leaves 4.5-5 mm. wide, flat, toward the apex becoming carinate 
and finely serrulate along the margins, triquetrous at the apex: 
culms 3-angled, becoming slender and flexuous, 0.8-1.5 m. high: 
cymes 2-5, decompound, densely fasciculate, irregularly corym- 
biform, 2-6 em. wide; the branchlets approximate, arching, 
terminating in glomerulate clusters of spikelets; lateral panicles 
subexserted on slender peduncles: spikelets rotundly ovoid, 
turgid, 3-6-flowered, 2-—4-fruited, 2.5-3. . long: scales 
mucronulate, loose, caducous, dark brown: bristles 6, exceeding 
the tubercle, upwardly hispidulous, tharacteristically divergent: 
achene pyriform, extremely flattened, heavily striate between 
the few well-spaced transverse ridges, often pale to castaneous, 

m. wide, 1.2—-1.3 mm. long: tubercle deltoid, sometimes 
apiculate, compressed, 0.3-0.4 mm. high. Pate 834, Frias. 2A 
and 2B; Map 79.—Class-Book of Bot. 744 (1861); Britton, Trans. 
N. Y. Acad. Sci. xi. 92 (1892); Small, Fl. 198 (1903) and Man. 
185 (1933). Scirpus schoenoides Elliott, Sk. Bot. 8. Car. and 
Ga. i. 89 (1816). R. Elliottii Dietrich, Sp. Pl. ed. 6: ii. 69 (1833), 
non Gray; Chapman, FI. So. U. 8. 525 (1860). R. multiflora 
Gray, Ann. Lye. N. Y. iii. 200, pl. 6, fig. 7 (1835). Phaeocepha- 
lum schoenoides House, Am. Midland Nat. vi. 202 (1920).— 
Roadside ditches and low ground in pine barrens and prairies of 
the Coastal Plain from North Carolina south to Georgia and 
west to eastern Texas. Norto Caro.ina: sandy grassy pine- 


., Wiegan Manning, no. 627 (G); Leroy, 
Washington Co., Bequaert, no. 11521 (G); Drummond, no. 251 


266 Rhodora [JuLy 


(G). Muississiprr: Ocean Springs, Jackson Co., Pollard, no. 1124 
(G, NY, US); Pass Christian, Harrrison Co., Dec. 18, 1919, 
Bartram (P). Lovisiana: stream-bottom south of Franklinton, 
Washington Parish, Brown, no. 6446 (La); vicinity of Covi ngton, 
St. Tammany Parish, Arsene, no. 11066 ; same locality, 
Arséne, no. 12220 (Mo, US); plentiful i in pools along road in pine 
flats 2 miles west of Hammond, angipahoa Parish, Hester, 
no. 808 (La); New Orleans, SDeiiacni in Herb. Hook., no. 367 
Y); common in Dasara west of Crowley, Acadia Parish, 
Brown, no. 581 ; common in low prairies, vicinity of Lake- 
Charles, Caleasieu Parish, Mackenzie, no. 44 (Mo, NC). TrExas: 
grows in a pond 3 miles below Town Bluff, Tyler Co., Wright 
(G); 2.6 miles east of Camp Jackson, Hardin Co. , Cory, no. 19714 
(CU); near Liberty, Liberty Co. . Bequaert, no. 11500 (G). 

7. R. microcarpa Baldw. ex Gray. Caespitose or solitary: 
aces 1-3 mm. wide, rarely narrower, flat; upper margins ser- 
rated; tips 3-angled: culms obtusely triangular, leafy, ascending, 
becoming tenuous and flexuous above, 5-8 dm. high: cymes 1-4, 
decompound, fasciculate, often densely so, eamatly corymbi- 
form, 1-6 cm. wide; lateral fascicles short-pedunculate: bracts 
leafy: spikelets ovoid, sometimes rotundly so, 3-4-flowered, 
2-3-fruited, congested, sessile, 2.5-3 mm. long: scales aristulate 
to acute, dark brown, more or less loose: bristles 6, varying in 
length from Y% the achene to equalling the tubercle, stiffly erect: 
achene obovoid, slightly biconvex, glistening bright brown, 
deeply alveolate, with the pits subisodiametric to broadly ob- 
long, only slightly ridged, 0.8-1.2 mm. wide, 1—1.2 mm. long: 


. ie ye. N. Y. iti. 202, pl. 

fig. : (1835); Chapman, Fl. So. U. 8. 525 (1860) ; Britton, ‘Trans. 

. Y. Acad. Sci. xi. 92 (1892); Clarke in Urban, Symb. Ant. il. 
sy boo: Britton, Mem. Soc. Cubana Hist. Nat. i. 199 (1916); 
Small, Fl. 198 (1903) and Man. 185 (1933); Britton & Mills- 
paugh, Bahama Fl. 55 (1920); Kiikenthal, Medde Rep. Spec. 
Nov. xxiii. 210 sate in part. &. patula Gray, Ann. Lye. N. Y. 
iii. 201, pl. 6, fig. 8 (1835), in small part. R. Torreyana Gray, 
var. hack: Grisebach, Cat. PI. Cub. 243 (1866). &. 
gracilis sensu C. Wright in Sauvaile, Anal. Acad, Ci. Habana, 
vili. 84 (1871) and FL Cub. 179 9 (1873), in part; non (Swartz) 
Vahl. 2. involuta C. Wright in herb. ex Clarke in Urban, Symb. 
Ant. ii. 130 (1900). R. perplexa sensu Britton & Millspaugh, 
Bahama FI. 55 (1920), not as to type, R. perplera Britton ex 
Small. &. Edisoniana Small, Man. 184, 1503 (1933). Phaeoce- 
phalum microcarpum House, Am. Midland Nat. vi. 202 (192 : 
Phaeocephalum patulum (misspelled palulum) House, Am. Mid- 

nd Nat. vi. 202 (1920), in small part——Swamps and 
margins of North Canin one saleeaan: southern Georgia 


1944] Gale,—Rhynchospora, Section Eurhynchospora 267 


and Florida westward along the coast to Louisiana; also on the 
Bahamas, and in the western provinces of Cuba and Puerto Rico. 
Nortu Carouina: Wilmington, Curtis (NY, cited by Gray in 
type-description). Grorara: in shallow pools along Aucilla 
Creek near Boston, Thomas Co., Harper, no. 1636 (G, NY, US). 
FLORIDA: swampy pine barrens near Jacksonville, Duval Co., 
Curtiss, no. 5415 (G, NC, US); vicinity of Eustis, Lake Co., 
Nash, no. 437 (G, NY, P, US); moist pine barrens, Merritt’s 
Island, Indian River, Brevard Co., Curtiss, no. 3149 (CU, G, 
NY, P, US); ‘‘wet weather pond” near Oakland, Orange Co., 
Curtiss, no. 6625 (G, NY, US); hammock, north of Eagle Bay 
near Kissimmee River, Osceola Co., Small, no. 9911 (CU); 
hammock, 12 miles east*of Okeechobee City, Saint Lucie Co., 
Small et al., no. 9292 (NY); everglades along the Palm Beach 
Canal, Palm Beach Co., Small, no. 8270 (G, NY); everglades 
along Tamiami Trail west of Miami, Dade Co., Small, no. 8835 
(NY, type of R. Edisoniana); swamp, Big Pine Key, Monroe 
Co., Killip, no. 32075 (CU, G, NY); in everglades south of 
Tamiami Trail, Collier Co., Moldenke, no. 879 (D, ; in 
pineland, vicinity of Fort Myers, Lee Co., Standley, no. 227 (G, 
NY); Indian Mound near Citrus Center, DeSoto Co., Small, no. 
9911 (NY); in a pine barren pond, north of Palma Sola Bay, 
Manatee Co., Simpson, no. 122 (G); swamp, Polk Co., Fredholm, 
no. 6229 (US); Tampa, Hillsborough Co., May, 1876, Garber 
US); in wet soil near Lake Butler, Pinellas Co., O'Neill, no. 
2615 (CU); in a low pineland, Pasco, Pasco Co., O'Neill, no. 
2616 (CU); margins of ponds in the pine barrens, Apalachicola, 
Franklin Co., Chapman in Biltmore Herb., no. 21la (G, US); 
open moist soil on St. Vincent’s Island, Franklin Co., Correll, 
no. 5604 (D). Mussissrppr: Horn Island, Jackson Co., Tracy, 
no. 2329 (NY); same locality, Tracy, no. 7682 (G); Ship Island, 
Harrison Co., June, 1886, Underwood Herb. (NY). Louisiana: 
in pine barrens, Mandeville, St. Tammany Parish, May, 1893, 
Langlois (Mo); west end of Grand Morais near New Iberia, 
Iberia Parish, Brown, no. 6383 (La). Banamas: edge of fresh- 
water marsh, Southwest Bay, New Providence, Britton & Brace, 
no. 510 (NY); grows in tufts 5 miles west of Fresh Creek settle- 
ment, in a pothole a few rods from shore of a salt creek, Andros, 
Wight, no. 241 (G); pineland swamps, Eight Mile Bay, Abaco, 
Brace, no. 1863 (NY); water holes, West End, Great Bahama, 
Brace, no. 3533 (NY, US); marsh, Glass Window to Harbor 
Island, Eleuthera, Britton & Millspaugh, no. 5395 (NY); water 
hole, Orange Creek and vicinity, Cat Island, Britton «& Mills- 
paugh, no. 5781 (NY); Hog Island, Wilson, no. 8429 (NY); 
water hole, Stopper Hill, Crooked Island, Brace, no. 4832 (NY). 
Cusa: edge of coastal swamps, La Coloma, Pinar del Rio, E kman, 
no. 17835 (NY); pineland ditch, Pinar del Rio City, Pinar del 


268 Rhodora [JULY 


Rio, Britton, Britton & Gager, no. 7250 (NY); in coastal swamps, 
Playa de Guanimar, Habana, Ekman, no. 18314 (G, 

muddy soil, west coastal savanna, or aban no, Havana, Britton, 
Cowell & De La Torre, no. 138349 (NY, US); in shrubbery, Cienaga 
de Zapata, Santa Clara, Ekman, no. 18372 (US); Wright, no. 230 
(NY, labeled by Gray ‘“R. Torreyana Gray var.” and annotated 
by Britton as “R. Torreyana var. microrhyncha A. Gray co- 
type’’); voes ae 3785 (G, NY, US). Purrro Rico: marsh, 


The more stnisk re aap of R. aiivecenee eta be ioe 
ficially distinguished from specimens of R. schoenoides. The 
differences between the two species are in their achenes. That 
of R. microcarpa is surrounded by stiffly erect bristles which are 
shorter than the achene or equal to the tubercle in height. The 
achene itself is commonly biconvex, castaneous to dark brown, 
strongly and irregularly alveolate. The achenial bristles of R. 
schoenoides are usually divergent, exceeding the tubercle in 
height. Unlike the common condition in R. microcarpa, the 
achene is strongly flattened, and pale to castaneous. The alveoli 
are longitudinally sdtdneedeed and appear as nearly regular rows 
of striae between a few prominent transverse ridges. 

On the continent, R. microcarpa occurs most commonly in 
Florida with a few collections from North Carolina, southern 
Georgia, and the coastal areas of Mississippi and Louisiana. 

R. schoenoides, on the other hand, occurs on the Coastal Plain 
from North Carolina to eastern Texas exclusive of the Florida 
Peninsula. 

68. R. TorreyaNna Gray. Caespitose: leaves slenderly linear, 
2-3 | mm. wide, flat or involute on drying, ane a stiff basal 


decompound, Becca tees corymbiform, 1-3.5 em. wide; the 
filiform branchlets somewhat pear spikelets bea 3-6- 
flowered, 1-5-fruited, pedicellate, Seek ascending, 4 on 


1-1.2 mm. wide, 1.7 mm. long: tubercle deltoid, siete 
compressed, decurent, ssa 0.3 eng in height. Pate 834, 
FIGS. BA an 0.—Ann. a. 197, a 6, ae 2 


Pe ene ee eet eS eS ee 


1944] Gale,—Rhynchospora, Section Eurhynchospora 269 


ete ree & Brown, Ill. Fl. i. 280, fig. 659 (1896); Britton, 
6 (1901); Small, Fl. 197 (190 eB Mr 184 (1933); 
Robinson & Fernald i in Gray, Man. ed. 7: , fig. 317 (1908); 
‘Fernald, Roopora, xxxix. 327 (19387). R. arid sensu Gray, 
Gram. et Cyp. i. no. 96 (1834), in part! and exel. syn. Phaeoce- 
phalum Torreyanum House, Am. Midland Nat. vi. 202 (1920).— 
Sandy pond-margins and sandy and peaty depressions in pine- 
lands and savannas of the Coastal Plain from Cape Cod, Massa- 
chusetts, to Georgia. MassacHusrETts: damp sandy and peaty 
border of Israel Pond, Barnstable, Barnstable Co., Fernald, no. 
8958 (G, NE, NY, P); Sconset, Nantucket Island, Nantucket 
Co., Bicknell, no. 1051 (NE, NY). Ruopr IsLtanp: sandy and 
gravelly shore of small double pond east of Long Pond, South 
Kingstown, Washington Co., Collins & Fernald, no. 11317 
(G, NE). New York: damp meadow, Amityville, Suffolk Co., 
Ferguson, no. 7873 (G, NY). New JersEY: White Horse, 
Mercer Co., Stone, no. 6756 (P); Quaker Bridge, Burlington Co., 
Aug. 1833, Gray ee TYPE); wet sandy clearing in barrens along 
tracks of C. R. R. of New Jersey, 114 miles northeast of Atsion, 
Burlington Co., Hermann, no. 3556 (G); boggy swale near head- 
waters of Cooper Creek, Lindenwold, Camden Co., Long, no. 
26371 (P); Egg Harbor, "Atlantic Co. , July 20, 1897, Lippincott 
(G); sandy clayey border of pond-hole, 114 miles n. n. e. of 
Olivet, Salem Co., Long, no. 47085 (P); moist pine barrens, 
Swain, Cape May Co., Mackenzie, no. 7193 (NY). DELAWARE: 
Felton, Kent Ge , Canby (G, P); wet een barrens near Laurel, 
Sussex Co. , Aug. 5, 1874, Commons (P). RYLAND: wet thickets 
along Snow Hill Road, 714% miles Sania of Salisbury, Wico- 
mico Co., Tatnall, no. 4371 (G); meadows, Ocean City, Worces- 
ter Co. , July, 1893, Canby (US). VIRGINIA: wet pineland south 
of Petersburg, Prince George Co., Fernald & Long, no. 8609 
(G, P); argillaceous and sphagnous bog (Reams Bog) south of 
Burgess, Dinwiddie Co., Fernald & Long, no. 8998 (CU, G, P); 
sandy and peaty depression (exsiccated shallow pond) about 4 
miles northwest of Homeville, Sussex Co., Fernald & Long, no. 
6076 (G, P); damp clearing in woods along Wakefield Road 
northeast of Sebrell, Southampton Co., Fernald & Long, no. 
10547 (G, NY, P); sphagnous bog about 1 mile northwest of 
Dahlia, Greensville Co. - ahalevon & Long, no. 8609 (G, 
Norra Carouina: dry savanna, 15 miles southeast of Green- 
ville, Pitt Co., Blomguist, no. 11229 (D); ov pine woodland at 
Nashville, Nash Co., Godfrey, no. 5155 (D, G); mossy floor of 
pine woodland, Raleigh, Wake Co., Godfrey, no. 4924 (G, NC); 


' No. 96 was apparently made up from a mixed collection. In the volume belong- 
ing to the library of the New York Botanic Garden, no 96 is R. Torreyana, as stated 

Gray in an appended correction; but in the volume at the Gray Herbarium, no. 96 
is R. gracilenta Gray. 


270 Rhodora [JuLy 


wet open sand pits, 5 miles west of Swanquarter, Hyde Co., 
Wiegand & Manning, no. 629 (G); savanna at Chocowinity, 
Beaufort Co., Godfrey, no. 5400 (G); pineland at Roseboro, 
Sampson Co., Godfrey, no. 5712 (G); edge of savanna, low 
ground, Big Savannah, west of Burgaw, Pender Co., Blomquist, 
no. 10067 (CU, D); Wilmington, New Hanover Co., Aug. 19, 
1930, Blomquist (G); dry sandy soil, Fayetteville, Cumberland 
Co., Biltmore Herb., no. 5055b (US); damp open sand-pocket, 
4 miles north of Hoffmann, Richmond Co., Wiegand & Manning, 
no. 630 (G). SourH CaroLina: low pine woods, common, 
Society Hill, Darlington Co., Curtis (G); Camden, Kershaw Co., 
House, no. 2681 (US); cart-road through pineland clearing, 5 
miles south of Andrews, Georgetown Co., Godfrey & Tryon, 
no. 1329 (G, NY); grass-sedge bog or savanna, 1 mile west of 
Chicora, Berkeley Co., Godfrey & Tryon, no. 851 (CU, D, G, 
NY, P); grass-sedge bog or savanna, 3 miles southwest of Man- 
ning, Clarendon Co., Godfrey & Tryon, no. 939 (G). GEORGIA: 
rather dry pine barrens near Pulaski, Bullock Co., Harper, no. 
941 (G, NY, US); pine barren at Ways, Bryan Co., Eyles, no. 
6419 (CU); moist pine barrens west of Ochmalkee, Montgomery 
Co., Harper, no. 1868 (G, NY, U 

6 . PERPLEXA Britt. ex Small. Caespitose: leaves flat, 


.3 mm. high. 
oaay ee and Man. 184 (1933); Fernald, Ruopora, xl. 399 


in central Tennessee; also in western Cuba and southern Domini- 
can Republic. VirGINIA: exsiccated argillaceous pond-hole in 
Chure 


woo: 
Fernald & Long, no. 8989 (G, P); wet peaty depression in pine- 
lands, 3-4 miles northwest of Waverly, Sussex Co., Fernald & 


1944] Gale,—Rhynchospora, Section Eurhynchospora 271 


Long, no. 8115 (G, P); pond-hole in pine and oak woods near 
Three Creek, north of Emporia, Greensville Co., Fernald & 
Long, no. 9282 (G, P); in shallow water of flat pineland, Collier’s 
Yard, 214 miles south of oe Dinwiddie Co., Smith & 
Hodgdon in Pl. Exsic. Gray., no. 924 (CU, G, NY, P). Norrn 
CaROLINA: drained hast Bee ‘i nas east of Deleo, Columbus 
Co., Wiegand & Man no. 615 (G). SoutH CARo.LINa: 
common in savannas, ‘Bukiee Co., Aug. 4, 1884, J. D. Smith, 
(G, US); exsiceated pine-barren pond, 7 miles southeast from 
Columbia, Richland Co., Sept. 26, 1883, J. D. Smith (G, US). 
GEORGIA: swamp, Augusta, Richmond Co., Ruthbert, no. 343 
(NY); gum-swamp along U. 8. Route 17, south of Ways, Bryan 
Co., Eyles, no. 6285 (CU); margin of pond, north of Ludowici, 
Long Co., Eyles, no. 6526 (CU); dry soil, Valdosta, Lowndes Co., 
May 27, 1940, Sargent (Sargent Herb. ); wet clay holes, 5 miles 
west of Tifton, Tift Co., Svenson, no. 7087 (G); shallow (now 
dry) spine-barren pond near Cobb, ‘Sumter Co., Harper, no. 1403 
(G, NY, US); margin of pond in pine barrens, 7 miles north of 
Cordele, Dooly Co., Eyles, no. 2303 (CU). FLorIDA: Chapman, 
2 (NY, TYPE); swam py places in pine barrens near Jackson- 
ville, Duval Co., Curtiss, no. 5178 (G, US); pinelands, Starke, 
Bradford Co., May, 1923, Wyman (N Y): prairies near Ft. 
Shackleford, Big Cypress, Lee Co., Small, Ho. Nig (NY); nite 
de Leon, Holmes Co., Curtiss, no. 6482 (G, Y, US). AaB 
pond, Elberta, Baldwin Co:, July 15, 1926, ey (StB); oe wet 
places, Mobile, Mobile Co., Mohr, no. 142 (US). MIssIssIPrt: 
Mississippi City, Harrison Co., Llo yd & Tracy, no. 379 (NY). 
TENNESSEE: wet open woods bordaaie a bog north of Man- 
chester, Coffee Co., Svenson, no. 9139 (CU, D, G, NY). Lovisi- 
ANA: in pine barrens near Mandeville, St. "Tammany Parish, 
May 1, 1893, Langlois (CU, Mo, P, US); Lake Charles, Caleasieu 
Parish, Tracy, no. 4882 (Mo, NY, US). TEXAS: in a pond three 
miles below Town Bluff, Tyler Co., Wright, no. 102 (G); ponds, 
Hempstead, Waller Co., Hall, no. 710 (G, type of R. microcarpa 
Baldw. ?var. achaeta Hall, nomen nudum; Mo, US, isotypes). 
Cusa: border of lagoon, vicinity of Pinar del Rio City, Pinar del 
Rio, Britton, Britton & Gager, no. 6962 (NY); banks of lagoon, El 
Punto, east km. 11 of road to La Coloma, south of Pinar del 
Rio, Ekman, no. 18262 (NY. This number cited by Kiikenthal 
as R. microcarpa A. Gray). Hispanroua: in the Rhexia belt, 
near Laguna de los Derramaderos, Sabana Guabatico, prov. 
Santo Domingo, Llana Costero, Santo Domingo, Ekman, no. 
13320 (NY. Siac tip by Kiikenthal as R. microcarpa A. 
ray.). 


The description above differs in two important details from 
the original description which appeared in Small’s Flora of the 


272 Rhodora [JULY 


Southeastern United States. The original reads: ‘‘perianth 
bristles 6” and “achenes biconvex.”’ In his Manual of the 
Southeastern Flora neither of these statements reoccur. I am 
inclined to believe that they were erroneously included in the 
original, probably due to a mixture in the material under study. 
The type-specimen is clearly R. perplexa as described above, the 
achene being extremely flat, not biconvex, and the bristles 1-3 
and rudimentary. 


Series 12. Glaucae (Clarke), stat nov. Growing in marshy 
areas, grassy slopes, and open woodland of the West Indies 
(represented by &. californica in the coast ranges of northern 
California) ; also in Central and South America and warmer parts 
of the Old World. Caespitose, rarely depressed: leaves 1.5-3 mm. 
wide: culms slender to stout, stiffly erect or becoming flexuous: 
cymes compound or decompound, fasciculate; the branchlets 
rigidly erect and spreading or flexuous: spikelets sessile, 1-3- 
fruited: scales loosely imbricated: bristles equalling the achene 
or exceeding the tubercle, antrorsely serrulate: achenes broadly 
or slenderly pyriform, a shining yellow-brown or light brown; 
the surface alveoli reduced to longitudinal striae which are inter- 
cepted by transverse corrugations or vague and blurred rugulosi- 
ties: tubercle deltoid, compressed, often attenuated.—Rhyncho- 
spora, Series B. Diplostyleae, Sect. 5. Glaucae Clarke in Urban, 
Symb. Ant. ii. 106 (1900), in part. 


Key To Species 1n Serres GLAUCAE 


Tubercle deltoid-attenuate, well exceeded by the bristles: achene 
with the transverse rugulosities and fine striae blurred to often 


th. 

ene 1.4 mm. wide, 2 mm. long; scales caducous: species 

known only from the coast ranges of northern California. 

: 70. R. californica. 

Achene 1.2 mm. wide or less, 1.6 mm. long or less: scales per- 
t Indies. 


Habit coarsely erect: culms 0.6-1.4 m. high: fasciculate 
1.6 mm. long : 
Habit depressed: culms 0.3-1.2 dm. high: fasciculate cymes 
decompound, small and congested, 1-1.5 em. wide wit 
achene corrugate ie te Oo yatinwre is ca Fo eLAK ES OS ak 73. R. rugosa. 
californica, sp. nov. Planta caespitosa: foliis 2-3 


m. latis planis, apice triquetris; marginibus superioribus 
minutissime dentatis: culmis fere 1 m. longis trigonis tenuibus 


1944] Gale,—Rhynchospora, Section Eurhynchospora 273 


2 cm. lato aut minus; ultimis glomerulis in ramulis brevibus 
tenuibus rigidisque gestis; fasciculis lateralibus minoribus; 
pedunculis longis ascendentibus: spiculis late ovatis, floribus 
tribus sed cum flore quarto terminali abortivo, dicarpis 4 mm. 
longis: squamis aristulatis, valde caducis: setis 6-7, antrorse 
hispidulis tuberculo longioribus basi latioribus et sparse plumo- 
sis: achaenio obovato lenticulari biconvexo obscure cing it 
lucido pallido obscure ruguloso 1.4 mm. lato 2 mm. longo: 
tuberculo attenuato-deltoideo 1 mm. alto. PLaTE 834, Frias. 1A 
and 1B; Map 60.— Marsh in the foothills of the coastal ranges of 
northern California. CALIFORNIA: Pitkin Marsh, 5 miles north 
a Peers he Sonoma Co., July 26, 1936, J. T. Howell, no. 
9 (G, TrpE; CA, ISOTYPE), 


This fine new species was collected and sent to me for study 
by Mr. John Thomas Howell of the California Academy of 
Science. Its large pale achene rendered conspicuous by the 
caducous scales is unmistakable and unique among the species 
of the United States. The light yellowish surface with the faint 
blurred rugulosities recalls that of R. Marisculus Lindl. et Nees 
in Mart.; whereas the habit is slighter but not unlike that of R. 
rugosa (Vahl) Gale. 

In all probability, if the status of specimens which have been 
collected in Central and South America as well as in the warmer 
areas of the Old World and which have been identified as R. 
glauca (R. rugosa (Vahl) Gale), be given a much needed revision 
the relationship of R. californica to the Series Glaucae could be 
more clearly defined. 


71. R. Mariscutus Lindl. et Nees. Coarsely caespitose: 
leaves 2-3 em. wide, stiffly ascending with attenuated, rae tear 
ee tips: culms stoutly ascending, 3-angled, leafy, 0. aes 

m. high: cymes fasciculate, 1-3, loosely decompound, 
wide, flexuous; axillary cymes on pet seorcth = Seabee 
spikelets lanceolate in outline, loosely organized, 1—4-flowered, 
1—2-fruited, 5-6 mm. long: scales Siete aaetiinte: loosely 
overlapping, dark brown: bristles 6, upwardly serrulate, finely 
attenuated, irregularly ascending to spreading and contorted, 
well exceeding the tubercle in length: achene obovoid, sometimes 
slenderly so, biconvex with a depressed margin, light yellow- 
brown, 1-1. 2 mm. wide, 1.4-1.6 mm. long; the minute longi- 
tudinal striae and transverse rugulosities blurred: tubercle 
triangular-attenuate, compressed, 1.2-1.8 mm. long. Parte 835, 
Figs. 3A and 3B; Map 61.—Nees, Linnaea, ix. 297 (1835), 


bd 


nomen nudum; Kunth, ee ii. 303 (1837), cited as ‘Sp. mihi 


274 Rhodora [JULY 


nonnisi nomini notae.’’; Nees in Mart., Fl. Bras. ii. pars 1: 142 
(1842), first valid publication; Boeckeler, Linnaea, xxxvii. 590 
(1873); C. B. Clarke in Urban, Symb. Ant. ii. 132 (1900), excl. 
syn. R. odorata C. Wright and R. jubata Liebm.; Britton & Wil- 
son, Sci. Surv. Porto Rico and Virgin Isl. v. 105 (1923); Kiken- 
thal, Fedde Rep. Spec. Nov. xxxii. 77 (1933). R. tenwiseta 
Wright in Sauvalle, Anal. Acad. Ci. Habana, viii. 83 (1871) and 
Fl. Cub. 179 (1873). R. borinquensis Britton, Bull. Torr. Bot. 
Cl. xlii. 387 (1915); Britton, Mem. Soc. Cubana Hist. Nat. 11. 200 
(1916); Britton & Wilson, Sci. Surv. Porto Rico and Virgin Isl. 
v. 104 (1923). Dichromena Marisculus Macbride, Field Mus. 


valle; : , isotypes); wet soil among rocks in stream, 
Arroya del Medio above the falls, Oriente, alt. 450-550 m., 
Shafer, no. 3231 (NY); damp woods near sphagnum, Sierra Nipe 
near Woodfred, Oriente, alt. 450-550 m., Shafer, no. 3453 (NY); 
in thickets on path from Woodfred to Piedra Gorda, Sierra de 
Nipe, Oriente, Ekman, no. 15247 (US); La Prenda, Oriente, 
Hioram & Manuel, no. 4671 (NY). Hispanroua: silt flats, moist 
ground, Sabana Nueva, Lomas de la Mediania, San Juan, prov. 
de Agua, Cordillera Central, Santo Domingo, Ekman, no. 13601 
(G, NY, US); on tembladera, in a laguna, Cuenca, Santo Do- 
mingo, Llano Costero, Santo Domingo, Ekman, no. 10982 (US). 
Puerto Rico: open wet places in forest, Sierra de Naguabo, 
Rio Icaco and adjacent hills, alt. 465-720 m., Shafer, no. 3515 
(NY, type of R. borinquensis Britt.). 

R. jubata Liebm. was confused by C. Wright with his R. 
tenuiseta, and so continued on in the synonymy of R. Maris- 
culus by C. B. Clarke. I have examined a specimen of Liebman’s 
from 8. Antonio, Huatusco, which is the type-locality as given 


in Mexicos Halvgraes,' and believe it to represent a distinct 


species. It differs from R. Marisculus both in its pronounced 
lax, more tenuous habit, and in its achene, which is distinctly 
cencellate and rugulose, surrounded by six stiffly erect bristles 
that equal but do not exceed the tubercle in length. Proof of 
this opinion rests, however, on the examination of more Mexican 
material. 

An examination of the New York collections of R. Marisculus 
leads me to believe that R. borinquensis Britt. owes its existence 

1 Ibid. 67 (1850). 


1944] Gale,—Rhynchospora, Section Eurhynchospora 275 


to Britton’s mistaken application of the name R. Marisculus 
Lindl. et Nees to specimens of R. odorata Wright. 

72. R. cacuminicola, sp. nov. Planta humilis caespitosa: 
foliis 1-2 mm. latis subcoriaceis leviter carinatis; apicem versus 
serratis triquetris, basi confertis, culmo brevioribus: culmis 
obtuse triangularibus 0.3—1.2 dm. ‘altis: cyma terminali decom- 
posita; ramulis brevibus rigide erectis vel ascendentibus fasciculis 
terminalibus parvis confertis; fasciculo laterali approximato; 
pedunculo ascendenti: spiculis lanceolatis 4-4.5 mm. longis 
2-3-floris 1—-2-carpis sessilibus congestis: squamis acutis vel fere 
aristulatis ferrugineis margine erosis: setis 6-7, quam tubercu- 
lum multo longioribus antrorse hispidulis: achaenio pyriformi 
0.9 mm. lato 1.1 mm. longo lenticulari biconvexo marginato 
luteo-castaneo, obsolete ee. pune compresso-subulato, 
attenuato, 1-1.2 mm. long PuaTE 835, Fics. 2A and 2B; 
Map 62.—Mar shy areas in ‘Cordillera Conteal of the Dominican 
Recublic Dominican Repusuic: in pratis paludosis in valle 
nuevo, prope Constanza, a 2200 m., June, 1910, Tiirckherm, 
no. 3417 (NY, Type). 

Von Tiirckheim’s specimen is accompanied by an annotation 
which implies that this species is only an alpine variety of R. 
glauca Vahl (R. rugosa (Vahl) Gale). That it is not a depressed 
form of R. rugosa, however, is attested by the achene which is 
smaller than that of the latter (0.9 mm. wide, 1-1.1 mm. long 
as opposed to 1.2-1.3 mm. wide, 1.5-1.7 mm. long), and only 
obscurely rugulose, with definite but short inconspicuous basal 
stipe. Also the tubercle of R. cacuminicola, unlike the short 
triangular tubercle of R. rugosa, is prolonged, with a subulate tip, 
and well exceeded by the bristles. 

Despite the depressed habit, the reduced stiffened inflores- 
cence and its original reference by von Tiirckheim to R. rugosa, 
R. cacuminicola seems more closely allied to R. Marisculus; for 
in the surface sculpturing of the achene, the comparative length 
of the bristles and the shape of the tubercle, its achene is to that 
of the latter species a faithful although much smaller copy. 

I have selected for this plant the epithet, cacuminicola, in 
order to signify its mountainous habitat, originally emphasized 
by von Tiirckheim. 

73. R. rugosa (Vahl), comb. nov. Coarsely ay niga 
leaves 1.5-3 mm. wide, coarse, ascending to reflexed, ca 
inate, with trigonous serrated tips: culms eae i Poear 
erect, somewhat flexuous at the summit, leafy, 0.5-1.3 m. high: 


276 Rhodora [Joy 


fasciculate cymes 1-3, strict, with stiffly erect or somewhat 
spreading branchlets, 1-2 cm. hig h: spikelets acutely ovoid, 4.5— 
5 mm. long, sessile or subscale: scales ovate-aristate, rather 
loosely imbricate, ferruginous: bristles 6, upwardly serrulate, 
ascending or often divergent, equal to or slightly exceeding the 
tubercle: achene broadly pyriform, 1.2-1.3 mm. wide, 1.5~-1.7 
mm. long, lenticular, biconvex, with depressed margins, glisten- 
ing yellow-brown; the longitudinal surface-striae obscured or 
nearly obliterated by the transverse broad, flattened, often pale 
corrugations; the thick persistent stipe 0. 3 mm. long: tubercle 
triangular-subulate, compressed, smooth or somewhat setulose 
at the base, 0.6-0.8 mm. long. PLaTe 835, rias. 1A and 1B; 
Map 63.—Schoenus rugosus Vahl, Eclog. Am. ii. 5 (1798). R. 
glauca Vahl, Enum. ii. 233 (1806); Boeckeler, Linnaea, xxxvil. 
585 (1873); Britton, Trans. N. Y. Acad. Sci. xi. 91 (1892), excl. 
syn. R. pungens Liebm.; Clarke in Urban, Symb. Ant. ii. 128 
(1900), excel. syn.; Britton, Mem. Soc. Cubana Hist. Nat. ii. 199 
See Britton & Wilso on, Sci. Surv. Porto Rico and Virgin Isl. 
v. 104 (1923); Kiikenthal, Fedde Rep. Spee. Nov. xxiii. Behr 
(1926). R. glauca var. altior Kiikenthal, Fedde Rep. Spec. N 
xxxil. 77 (1933). R. gracilis pinks Grisebach, Fl. Brit. W. I. 37 74 
(1864), in part, and Cat. Pl. Cub 3 (1866), i in sea C. Wright 
in Sauvalle, Anal. Aead. Ci. Fahoke, vili. 84 (1871) and Fl. Cub. 
179 (1873); non (Swartz) Vahl. Dichvomena glauca Macbride, 
Field Mus. Pub. Bot. iv. 166 (1929).—Mostly on damp open 
grassy slopes of the Greater Antilles, and of the Island of Domin- 
ica; also in South and Central America and Lower Mexico. 
Cusa: banks of small streams, Loma Ratones, Rio San Sebas- 
tian, La Cumbre, ges of San Juan y Martinez, Pinar del Rio, 
Ekman, no. 18062 (NY US). Jamaica: along the trail, Mt. 
ry, Mazon, no. 852 (US); in the morass, Pedro Morass, Upper 
Checadine alt. 3000 1; Harris, no. 11169 (G, NY, US); on wet 
eres Peckham, Upper Clarendon, alt. 2000 ft., Harris, no. 
810 (G, N' , US); amongst grasses in wet hillside pastures, 
Trov, alt. 2000 ft, Harris, no. 12590 (G, NY, US). HispaNioLa: 
eruptive ee Haut-Piton, Port-de-Paix, Massif du Nord, 
Haiti, alt. 800 m., Ekman, no. 4638 (G, NY, US); Petit Borgne 
to Mt. Casse, Haiti, alt. 200 ft., Nash, no. 558 (NY); grassy 
mountain trail to Citadel, vicinity of Dondon, Depart. du Nord, 
Haiti, alt. 400 m., Leonard, no. 8643 (US); grassy summit of 
mountain east of willage, vicinity of Marmelade, Depart. du 
Nord, tee alt. 800 m., Leonard, no. 8241a (US); pineland, 
gunas de Cenobi, Moncion, Monte Cristi, Cordillera Central, 
Santo Domingo, c. alt. 1100 m., Ekman, no. 12762 (NY, US). 
gr te Rico: in re Las Mesas, near Mayaguez, alt. 350 
, Holm, no. 64 (CU), Aguada, Sintenis, no. 5742 (US); in 
euininia. Lares, Sintenis, no. 5901 (G, US): low ground bac 


1944] Gale,—Rhynchospora, Section Eurhynchospora 277 


of Catano, Heller, no. 6410 (CU, NY, US); road from Rio Piedras 
to Trugillo alto, Hioram, no. 806 (NY, US); Cidra, Feb. 28, 1928, 
Perkins (NY); on rock by falls in river, Rio Teaco and adjacent 
hills, Sierra de Nasal, Shafer, no. 3534 (NY, US). Dominica: 
common in cleared pasturelands on deep rich soils of Milton 
Estate, ca. alt. 530 m., Hodge & Hodge, no. 2608 (G).. Sourn 
AMERICA: America Meridionali,! Rohr in Vahl Herb, (G, TyPE- 
PHOTO). 

The species, Schoenus rugosus, was published with full descrip- 
tion by Vahl in his Eclogae Americanae, ii. 5 (1798). However, 
when, in the Enumeratio Plantarum of 1806, he transferred the 
species to Rhynchospora, he unfortunately chose to give it a new 
name, and it appeared as R. glauca. The old name, Schoenus 
rugosus, was then given in synonymy, accompanied by the 
original description and citation, ‘‘ Habitat in America meridi- 
onali. von Rohr’’. Since the specific name, rugosa, is available 
for use under Rhynchospora, the new combination R. rugosa 
(Vahl) Gale takes precedence over R. glauca Vahl. 

The range of this species has been generally given as warmer 
regions of both hemispheres. However, the specimens from the 
East Indies and adjacent continental lands which have been 
commonly identified as R. glauca, are not, I believe, conspecific 
with the authentic R. glauca Vahl (R. rugosa (Vahl) Gale) of the 
West Indies, Central and South America. Although these 
Asiatic plants are closely related to and have the same general 
aspect as R. rugosa—the coarse growth and strict, compound 
fascicles with mainly ascending, sessile spikelets in small ultimate 
clusters—their achenes are larger, more nearly orbicular rather 
than pyriform, and the surface of the achene is generally casta- 
neous, pitted, and finely ridged, without the prominent yellowish 
band-like corrugations which characterize the achenes of R. 
rugosa. 

The Gray Herbarium isotype of Liebman’s? R. pungens, Mira- 
dor, July, bears the annotation R. glauca Vahl followed by Brit- 
ton’s initials. The specimen itself is fragmentary and the surface 
details of the mature achenes are obscured by what appears to 
be an unnatural whitish coating. However, careful examination 
seems to indicate that the achenes are normally cancellate to 

1 Probably along the coast of sper or of French or Dutch Guiana. See La- 


marck, Eneyel. Meth, Bot. viii, 754 (180 
? Mexicos Halvgraes, 65 (1850 


278 Rhodora [JULY 


nearly smooth, with only a suggestion of transverse rugosity. 
In addition, the tubercle is plainly spongy and tongue-shaped 
rather than strongly compressed and triangular-subulate. If 
these observations can be confirmed through better material, 
R. pungens, although closely related to R. glauca, will retain its 
specific status. 


INDEX 


New scientific names are printed in full-face type 


Cephaloschoenus, 91 
Ceratoschoenus, 91 

Cladium, 90 

Cyperaceae, 91, 92, 95, 97, 246 


ca elisa 94; 102; alba, 121; 
, 245; distans, 191; gla auca, 
276: "Miceiclen: @ 


Eleocharis, 119; tuberculosa, 92 


ees 101, 102; album, 
121, var. macrum, 124; axillare, 108, 
Baldwinii, 181; brachychae tum, 
; caducum, 261; capillaceum, 


8, dod um, 228; 
Earlei, 233; fasciculare, 189; fili- 
folium, 174; fuscum, 170; glomera- 

“ , var. minus, 115; gracilen- 
tum, 183 yi, 231; inexpansum, 

; intermedium, 1 Knie- 
skernii, 126; leptorhynchum, 179; 
microcarpum, 266; miliaceum, 257; 
mixtum, 262; fe hy 166; pa- 
tulum, 266; perplexum, 270; 


Plankii, 233; * Seorereeh: 131; pro- 
liferum, 262; punctatum, 238; 
rariflorum, 08; schoenoides, 265; 
sed Ste stipitatum, 9; 
m, 269 
Pleargiath ya. 2 
Beal Sect. ii. Pauci-Nucigerae, 


Rhynchospora, subg. Diplostylis, 92, 
102; subg. Distylis, 102; subg. 
1a 7 


munes, 102; ii. Dichostvlewe 102; 
V. Glomeratae, 104, 169, 177, 207, 

36, 255; I. Pusillae, 134; 
§ Euriochaete, 128; sect. Eurhyn- 
Hg ink 89, 91 1-93, 95, 101, 102, 


sp 


4, > 
Eu-Rhynchospora, 102, Divi 


Eu-Rhynchospora, Sect. ill. andl 


227; ser. B. Diplostyleae, sect. 3. 


104, 255; ser. Cernuae, 93, se 2 98, 
100, 103, 211, 220; ser. Ghapm 
ae, 100, 103, , 159; ser 
Cubenses s, 100, 103 at se 
Fasciculares, 1(3, 177, 178; ser. 
] 1 

( 

( 


ser. 
Glomeratae, 95, 99, 100, 103° 104; 
ser. Harveyae, 99, 100, 104, 227: 
ser. Plumosae, 99, 100, 103, 128; 
ser. Rariflorae, 100, 103, 207; 
group Capitatae, 92, 94; group 
Communes, 92; group Corymbosae, 
94; group Longiros ostres, 92; alba a, 


170, f. laeviseta, 123, var. macra, 
95, 121, 124, 125, var. —— 
12: : axillaris, 1¢ i , var. 
cephala, 95, 1 06; bahaimensis, 23; 
i 178, 181, pl. 


25: Berterii, re aoe” 164, 
165; Pa us ie i brachy- 


leptocarp ar. mino 5; 
yi aah a 91, 105, 107-109, 111, 
119, 120, var. attenuata, 108, 110, 


120, f. controversa, 110, 121 
_ 98, 108, 111, 120, pl. 818, 


2 
oF 


INDEX 


f. antrorsa, 109, 121; cephalotes, 


y 212, 214, 2 
chalarocephala, 105, 111, 130, ay 
18; Chapmanii, * 102, 132, 159, 
. 821; ciliaris, 


00, oe 237, 239, 


of oe nies, 
215, pl. 827; seg 93, 96, 1 
4-227 8 st 


5, 236, 
reg tl pl. ope Sytem, 95, 


; de 
l. 826; distans, 89, 95, 162, 
177, Be 193, , 230, 232, 247, 
at r. B. fascicularis, 188, 
pected 192, var. y. 
gracillima, » var. microcarpa, 
176, 177, var. tenuis, 187; di 
gens, 165; 


phyla, hoides, 
194, 196, var. typic 03, 162, 
186, . ae, oes 196, oo 8 
825; 103, 132, 178, 182, 


ae 620; fibrillosa, 216; filifolia, 91, 


4; fuscoides, 95, 132, 169, 
170, 175-177, p « O22; 


» Var. Po asia 239, ’ ’ 
244, 245, 248, pl. 831; glomerata, 
89, 91, 105, 112. 113, is, 116, 119, 


247, var. angusta, 113, 114, i 
var. discutiens, Bs "1 9, 


i dosed oy 115, var. minor, 95, 
f controversa, 119, 
diseutien, 119, paniculata, 


113; B: robustior, 113, var. typica, 
113-115, 120, pl. 819; agracilenta, 
91, 95, 177- 179, 183-186, Oe 

$26, var. diversifolia, rea 
gracillima, 164, 185, 187; o aailis 
3, 179, 245, 266, 276; Gra ayana, 
125, 126, 168; Grayii, 91, 92, 117, 
118, 126, 186, 228, 230, 232, pl. 829; 
‘1738, pl. 823; 


824, en ar. evict "179; rh yee 
ona 93, 95, 2 2, 27 219, 222, 223, 
5.156, 218 oe, 


neler 
223, 224, pl. 827, 


269; microcarpa, 91, 104, 2 
255, 256, 259, 266, 268, 271, pl. 832, 


91, 93, 96, 97, 104, 239, 255, 257, 
3; mixta, 256, 259, 262- 
, 2; ?monostachya, 92; 
multiflora, 91, 265; nipensis, 186, 
211, 212, "O15, pl. 826; uda, 99, 
7 , 132, 159, mT 168, pl 


penniseta, 93, 131; 
255, 256, 259, 266, 270, 272, pl. 832; 


a ai 


INDEX ill 


pineticola, 130; pinetorum, 248, 
249; Plankii, 233; pleiantha, 132, 
170, 171, 172, pl. 823: plumosa, 90, 
91, 93- 95, 97, 128, 131, 132,: pl. 
820, var. intermedia, 130; : prolifera, 
62, 263; pruinosa, 93, 97, 186, 211, 
212, 221, pl. 826; punctata, 90, 236, 
238, 330.) pl. 530; pungens, 276- 
215; pyenocarpa, 
91, 92, 228-230; ’ Randii, 220, 221; 
Rappiana, 160, 161; rari riflora, 90, 
93, 103, 186, 207, 210, 211, pl. 828; 
rugosa, 93, 239, 272, 273, 276, 
277, pl. 835; saxiools, 236, 238-240, 
pl. 830; scabrat ata, 9 

0, var 


a, 132, 159, 162. 
20; solitaria, 96, 132, 
134, 159, 161-164; sparsa, 89, 91, 
96, 257; stenophylla, 165, 186, 207, 
210, 211, 219, 220, var. albescens, 
219; stenophylloidea, 96, 186, 
224-226, pl. 828; stipitata, 259; 
sulcata, 100, 236, 239, 240-242, 


1. 831; ithe, 93, 99, 100, 186, 


268, 269, pl. 834, var., 268, v. 
microthyncha, 266, 268; ahader: 
194, 196; tha phylla, 183, 185; 
Wrightiana, 163, 165, 178, 185-188, 
195, 196, 825 
Rhynchosporeae, 94, 102 
Rhynchosporoideae, 246 


Sire 89, 90, 246, sts 2 albus, 
21; axillaris, 108; tatus, 90; 
caitlin 115-1195 ater 90, 


90, ’ 
245-248; donates 191-193, 231, 
32; fascicularis, 118, 188, 192, 193; 


sp 

Scirpus, 90, 119; leptolepis, 92; poly- 
phyllus, 246-248; schoenoides, 91, 
265 


Triodon, 101: albus, 121, var. macer, 
124; eapillaceus, 126; glomeratus, 
1138 


ae Rhodora 


S.G del, 


RuYNC ees CHALAROCEPHALA! FIG 
achene, « 20, 


. 1A, portion of inflorescence, 2; Fic. 1B 
R. c APILLA ACEA: FIG. 2A, portion of ger gees ance, . 2; FIG er ac hene, x 20 
R. MICROCEPHALA: FIG. £ 2 of i x 2: ria. 3B, a¢ rg ene, X 20. 
R. CEPHAL ANTHA var. TYPIC A: a 9, Shaina ot eoreatons; xX 2: 1B, achene, 
aD. 
x 20. 


Plate 818 


Plate Plate 819 


S, G, del, 


tHYNCHOSPORA GLOMERATA Var. TYPICA: FIG. 1A, portion of inflorescence, X 2; FIG. 

LB, achene, 20. 

R. KNIeSKERNII: FIG. 2A, portion of rangers gsi X 2: FIG. 2B, achene, X 20. 
R. CAPITELLATA: FIG. 3A, inflorescence, X 2; 3B, ac he ane, X 20. 
R. ALBA: FIG. 4A, portion of nab strated x 2: pity 4B, achene, X 20. 


Pl Rhodora Plate 820 


Pe 


S. G. del. 


tHYNCHOSPORA SOLA! FIG, 1A, inflorescence, X 2: FIG. 1B, achene, * 20 
t. PLUMOSA: FIG. 2A, inflorescence, X 2: FIG. $ . 

R. MAcCRA: FIG. 3A, portion of inflorescence, < 2; FIG. 3B, ac shene, x 20. 
R. sivas i. FIG. 4A, inflorescence 2; FIG. x 20. 


Rhodora Plate 821 


‘ 


i) \ 
3 Wye we 
wi 
TS w 
NAY ye 


i 
hia Mu” 
iil big il ve 


S. G. del, 


RHYNCHOSPORA NUDA: FIG, LA. inflorescence. X 2; ric. 1B, achene. X 20. 
R. OLIGANTHA, var. TYPICA: FIG, 2A, inflorescence, 2; FIG. 2B, achene, X 20. 
) 


R. OLIGANTHA. Var. BREVISETA: FIG, 2C. achene, & 20. 
R. CHAPMANTI: FIG. 3A, inflorescence, 2; FIG, 3B, achene., x 20 
R. PALLIDA: FIG. 4A, inflorescence, x 2; FIG. 4B, achene, 20. 


Rhodora Plate 822 


S.G. del. 


RHYNCHOSPORA BRACHYCHAETA: FIG. 1A, portion of inflorescence, X 2; F1G. 1B, achene, 

20 

R. crurarts: ria. 2A, inflorescence, X 2; FIG. 2B, achene, X oe 

R. FILIFOLIA: FIG. 3A, inflorescence, X 2; FIG. 3B, achene, x 20. 

R. sourrarta: F1G. 4A, inflorescence, X 2; FIG. 4B, achene, X 20. 
~ ” ee 

R. Fuscorpss: Fa. 5A, inflorescence, X 2; FIG. 5B, achene, X 20. 


Rhodora Plate 


S.G. del, 


RuyYNCHOSPORA HarPERtI: FIG. 1A, inflorescence, aa 2: ric. 1B, achene, X 20. 
R. CRINIPES: FIG. 2A, portion of —— scence, X 2; r1G. 2B, achene, X 20. 

R. Curtissi: ric. 3A, inflorescence, * 2; FIG. 3B, ac chene, en 

R. PLEIANTHA: FIG. 4A, portion of indloncodente, x 2; PER, achene, X 20. 


823 


= 


Rhodora Plate 824 


S.G. del. 
> > 9) 
RHYNCHOSPORA LEPTORHY NCHA: FIG. 1A, coos ny . 1B, achene, X 20. 
R. rusca: Fig. 2A, portion of oie reste: X 2; Fic. 2B, “achene, X 20. 
R. GaGERI: FIG. 3A, inflorescence, X 2; 3B, ac rahene, 20. 


R. JOVEROENSIS: FIG. 4A, portion of sot Selecta < 2: Fic. “4B, achene, X 20. 


Rhodora Plate 825 


S.G. del. 
. ae Sé )- FIG. 
RHYNCHOSPORA FASCICULARIS, Var. TYPICA: FIG. 1 A, portion of inflorescence, X 2; FI 
1B, —— <x 20. mos 
R. FASCICULARIS, Var. DISTANS: FIG. 2A, portion of inflorescence, < 2; FIG. 2B, achen 


R. FERNALDI: FIG. 3A, portion of inflorescence, X 2: FIG. 3B, achene, X 20. 
R. Baupwintt: FIG. 4A, portion of inflorescence, . 2; FIG. ri schunes: x 20. Pa 
R. WRIGHTIANA: FIG, 5A. portion of inflorescence, « 2: 5B, achene, X 20. 


Pla: 


Plate 826 


Rhodora 


ngs NCHOSPORA DEPRESSA? FIG. 
2 oe 


. NIPENSIS: FIG. 4A, inflo rescence, 
. DEBILIS: FIG. 5A, inflorescence, X 2; 
 GRACILENTA: FIG. 6A, portion of inflorescence, X 2; FIG. 


S.G. del. 


ie 1B, achene, 20. 


—_ 


1A, inflorescence, 2: 

CERNUA: FIG, 2A, it orescence, X 2; FIG. 2B, ac ene x2 

: FIG. 3A. portion of inflorescence, X 2 3B, achene, x 20. 
< 2: FIG. 4B, a x2 ys 
ria. 5B, achene, X 20. 
6B, achene, X 20. 


Rhodora Plate 827 


— 


} i} 
yp 
OL 


Ya 
A 


Sis 


‘i 


= sks 


S. G. del. 


. : 9). ane 
RHYNCHOSPORA TENUIFOLIA: FIG, 1A, portion of inflorescence, X 2; FIG. 1B, achene, 
x 20. 


* 


R. SHAFERI: FIG. 2A, inflorescence, X 2; F1c. 2B, ac x 
R 


hene, 20. 
SCABR: orescence, X 2; FIG. 3B, achene, 20. 
R. cRISPA: FIG. 4A, inflorescence, X 2; FIG. 4B, acher hene 
R. LINDENIANA, var. TYPICA: FIG. 5A, ‘portion of er vas Meat X 2; ria. 5B, achene, 


: 2. LINDENIANA, var. BAHAMENSIS: FIG. 5C, achene, * 20. 


Me Rhodora Plate 828 


Ss. G. del, 


RAYNCHOSPORA CUBENSIS: FIG. 1A, portion of inflorescence, < 2; FIG, 1B, achene 


| R. STENOPHYLLOIDEA:! FIG. 2A, achene, X 20. 
R. cuLrxa: rig. 3A, portion of inflorescence, X 2; FIG. 3B, achene, X 20. 

R. RARIFLORA: FIG. 4A, portion of inflorescence, X 2; FIG. 4B, achene, X 20. 
R. sreNopHYLLA: FIG. 5A, inflorescence, X 2; FIG. 5B, achene, X 20. 


‘4 829 
Rhodora Plate 


© gt 
ta 
Vitti 
bie pe, 
1‘ Oety hy Hi 
Ad y 


‘i 
i,! 


S, G, del. 


RuHYNC HOSPORA MEGALOCARPA: FIG, 1A, portion of inflorescence, X 2; FIG. 1B, 
achene, X 4 
R. Harvevi: FIG. 2A, portion of inflorescence, x fs _ 2B, achene, 20. 
R. Grayit: Fic. 3A, portion of inflorescence, X 3B, achene, X 20. 


Rhodora Plate 830 


S G. del. 


‘HYNCHOSPORA COMPRESSA: FIG. 1A, portion of inflorescence, X 2; FIG. 1B, achene, 
x 20. 


) 


R. puNcrATA: FIG, 2A. portion of inflorescence, 2; FIG. 2B, achene, X 20. 
> P . ‘eS 7 . . ~ . € ‘ . > » 4 = 

R. Saxicoa: Fig. 3A, portion of inflorescence, X 2; FIG. 3B, achene, x 20. 
R. OBLITERATA: FIG, 4A, portion of inflorescence, X 2; ric. 4B, achene, 20. 


Rhodora Plate 831 


ri 2 @ 

SS232: oe 
Seces, 

SS e=4 


es 

SS 

<a 
SS 


~ 


~'e 


re aa SED 
ashauanis 
hs ii Re, * 


WWwNwoU 


RA GLOBULARIS, var. TYPICA: FIG. 1A, portion of inflorescence, * 2; FIG. 
1B, cid sei 
R. Brrrroni: Fic. 2A, portion of -oneeN x: 2: 2B, achene, x 20. 
R. sutcata: Fic. 3A, portion of inflorescence, X 2; vie. 3B, ache ne, X 20. 4B 
R. GLOBULARIS, var. RECOGNITA: FIG. “4A, portion of inflorescence, xX 2; Fie. 2 
achene, X 
R, GLoBu LARIS, var. PINBTORUM: FIG, 5A, achene, X 20. 


Rhodora 


Plate 832 


S. G. dei, 


RHYNCHOSPORA DECURRENS: FIG. 1A, portion of inflorescence, X 2; Fic. 1B, achene, 
X 20. 
R, MICROCARPA: FIG. 2A, portion of inflorescence, X 2; FIG. 2B, achene, X 20. 
R. PERPLEXA: FIG, 3A, portion of inflorescence, X 2; FIG. 3B, achene, < 20. 
R. Mixa: ria. 1A, portion of inflorescence, X 2; Fic. 4B, achene, X 20 


Rhodora Plate 833 


! 


y 
f 


KS | 
JN \ 


rere baby 


p< oa ESS) 
i AYU 
ail 


S. G. del. 


Pp 


SPORA INEXPANSA: FIG. 1A, portion of inflorescence, X 2; Fic. 1B, achene, 


AE. 
{. MILIACEA: FIG. 2A, portion of inflorescence, X 2; F1G. 2B, achene, X 20. 
R. ODORATA: FIG. 3A, portion of inflorescence, X 2: F1G. 3B. achene, X 20. 
R. cApucA: FIG. 4A, portion of inflorescence, X 2; F1G. 4B, achene, X 20. 


Rhodora Plate 834 


S. G. del. 
RHYNCHOSPORA CALIFORNICA: FIG. LA, portion of inflorescence, a =: FIG. LB, achene, 
X 20. 


R. SCHOENOIDEs: FIG. 2A, portion of inflorescence, 2; Frc. 2B, achene, io 
R. Torreyana: Fic, 3A. portion of inflorescence, X 2; F1G. 3B, achene, X 20. 


Plate 835 


Rhodora 


S. G. del. 


RHYNCHOSPORA RUGOSA: FIG. 1A, inflorescence, X 2: FIG. 1B, achene, X 20. 
R. CACUMINICOLA: FIG. 2A, inflorescence, X 2; Fria. 2B, achene, X 20. 20 
R. Mariscuuus: ric. 3A, portion of inflorescence, X 2; F1G. 3B, achene, K wv. 


Reprinted from Ruovora, Vol. 46, 


CONTRIBUTIONS FROM THE GRAY HEI 
OF HARVARD UNIVERSITY | 


OLD WORLD ALLIES 


me ge Ya 2% ~ oe en, Ree ee eT Cana a Ry Sa eT SR re ae ra yo ser 
- oo F 4 wae ras nig iL ern a epee % LE ee ea ey OE 
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Reprinted from Ruopora, Vol. 46, October-December, 1944 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


CLIT 


AMERICAN THALICTRA AND THEIR 
OLD WORLD ALLIES 


BERNARD BolrviIN 


Dares or IssuE 
WOME ORT-O7 0 ee ee ee ek 11 October, 1944 


Petes DOL4OR 6 Re ee ee es 
Paces Dey Rae ed eg a Te Fee LS came seg 9 Me 16 December, 1944 


CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD 
UNIVERSITY—NO. CLII 


AMERICAN THALICTRA AND THEIR 
OLD WORLD ALLIES 


BERNARD BOIVIN 


A WORLD-MONOGRAPH of this genus was published in 1885 by 
J. C. Lecoyer in the Bulletin de la Société Royale de Botanique 
de Belgique (24: 78-324. 1885) and reprinted under the title 
Monographie du genre Thalictrum. It contains a history of the 
genus from Dioscorides to 1885. This need not be repeated 
here; I will, however, sketch the highlights of the history of our 
knowledge of the American species. 


History OF THE GENUS 


Cornut described in 1637 a Thalictrum canadense, which, as 
will be shown later, was not an American species. Morrison in 
1715 reported the first three species from the Eastern United 
States and Canada, but the names he proposed were not validated 
by later authors and it is not surely known what species he had. 
He also clearly described a European species, 7. alpinum L., 
which was to be first found in America on Newfoundland by 
Banks in 1766. Again, in 1739, Gronovius in his Flora Virginica 
proposed two American species and, in the first edition of the 
Species Plantarum, Linnaeus validated Cornut’s species under 
the name Thalictrum Cornuti and described T. alpinum, T. 
dioicum and T. purpurascens. Aiton in 1789 described from 


338 Rhodora [OcToBER 


cultivated specimens a Thalictrum rugosum, purportedly an 
American species, but Lecoyer (I. ¢. 311), who saw the type of it 
along with material cultivated in many botanical gardens, made 
it a synonym of Thalictrum glaucum Desf., a European species. 
About 1790, Muhlenberg wrote a Flora Lancastriensis which was 
never published but which contained descriptions of two new 
species, 7’. polygamum and another species, the first of which was 
validated in 1825. The manuscript of this flora is at the Gray 
Herbarium. Michaux (Fl. Bor.-Am. 1: 322) described T’. laevi- 
gatum in 1803. De Candolle’s Systema in 1817 recognized eight 
species and four varieties for North America, one species for 
Mexico and four for South America. To these was added T. 
peltatum DC. in the Prodromus, vol. 1, published 7 years later. 

From 1824 on there was to be no synoptical treatment of the 
South and Central American species of Thalictrum, though 
there was a large amount of work done upon the North American 
species. Hooker in 1829 accepted 4 species in his Flora Boreali- 
Americana, Torrey & Gray’s Flora of North America included 6 
species and Sereno Watson listed thirteen species and three 
varieties in his Bibliographical Index (1878). The same year 
Lecoyer published a preliminary paper and seven years later in 
his world-monograph of the genus he attributed twelve species to 
North America, seven to Central America and three more to 
South America, in all twenty species and one variety for the 
Americas. A year later, in 1886, Trelease, working at the time 
at the Gray Herbarium, published a monograph of the North 
American Thalictra, recognizing twelve species and three va- 
rieties. 

For the next 25 years the history of this genus was to be 
dominated by E. L. Greene, who described as new forty-five 
species and seven varieties of Thalictrum. He also left many 
unpublished names on herbarium sheets in various herbaria. 
By the time this was over, there existed such a confusion in the 
taxonomy of this genus that most local floras cautiously followed 
and still follow more or less the posthumously published treat- 
ment by Gray in his Synoptical Flora of North America in 1895, 
in which thirteen species and four varieties were accepted. J.N. 
Rose started in 1899 to monograph the Central American and 
Mexican Thalictra but only part of this work was carried through 


Te ee 


1944] Boivin,—American Thalictra and their Allies 339 


and published. By the end of 1942, about 165 specific names 
had been proposed to designate American’ Thalictra, 40 of them 
for species restricted to areas south of the United States-Mexican 
boundary. 

VALUE OF DESCRIPTIVE CHARACTERS 


HEIGHT OF THE PLANT. Quite variable in tall species, more 
constant in smaller ones. 7’. rhynchocarpum Dill. & Rich. of 
the southern Cameroon Mountains and Fernando Po Island may 
reach a height of 4 meters. 

PuBEscENCE. Extremely useful characters are those based 
on pubescence; for no matter whether the plant is male or female, 
whether it is in flower or in fruit, accurate descriptions of the 
pubescence always apply and can easily be checked no matter 
how fragmentary the specimen may be. Many normally 
pubescent species do, however, occasionally present glabrous 
specimens and mature foliage often loses its pubescence, but the 
reduced leaves in the upper part of the inflorescence will retain it, 
at least up to the full maturity of the fruits. Hairs in this genus 
are either unicellular or multicellular. Furthermore, multi- 
cellular hairs may be uniseriate or multiseriate. Unicellular 
and uniseriate hairs are normally translucent. Capitate hairs 
are either unicellular or multicellular; all other types are multi- 
cellular. Capitate hairs are always very small so that a good 
binocular may be pretty useful to anybody attempting to identify 
a Thalictrum, but this type of hair usually exudes a fetid and 
sticky substance in vivo. Multiseriate hairs are usually opaque, 
often whitish and sometimes diversely colored. They may 
either be stiff, short and blunt or long (1 mm. or more), undulate 
and attenuate. Short blunt multicellular hairs always render 
the plant scabrous. In one species, 7’. inuncans nostrum, small 
hooked prickles are present all over the plant. In another, T. 
Standleyi Steyermark, the hairs are stellate, stipitate, and of a 
brownish color. This species is also the most fetid of all. 

Thus there are six types of hair, all easily recognizable either 
by the naked eye, the hand-lens or the binocular, e. g.: capitate 
hairs, uniseriate flecuous hairs, short multiseriate hairs, long 
attenuate multiseriate hairs, small prickles and stellate hairs. 
Within the same species hairs of only one of those types are 
usually present, or none at all; rarely the same plant will have 


340 Rhodora [OcroBER 


hairs of two or three different types. Such a good character has 
usually been overlooked in the past, thus rendering absolutely 
impossible the exact interpretation of many a name. 

Lear. An arbitrary distinction has been made between the 
leaves according to whether they are attached close to or at the 
base of the stem, whether they are scattered along the lower 
-unbranched half of the stem or whether they subtend a primary 
branch of the inflorescence. They are respectively termed folia 
radicalia, folia caulinaria and folia inflorescentiae. 

LEAF-pIvIsion. Contrary to pubescence, the type and degree 
of division of the leaf is usually widely variable and was usually 
stressed by describers of new species. Only rarely does the leaf 
furnish any specific character, although each species, when well 
known, may usually be recognized by the contour of the leaflets. 
But these differences are not constant and not very easily put 
into words. In most cases leaves vary from 3- to 5-ternate; 
basal leaves are usually more divided than cauline leaves. Lower 
branches of the inflorescence are in many species subtended by 
1-3-ternate leaves, sometimes by simple reduced ones. Only 
rarely will the secondary branches of the inflorescence, or even 
the pedicels, be subtended by leaves, as in 7. sparsi elk Turcez. 
and 7. Hultenii nostrum. 

PETIOLE AND Petiotutes. Their length is extremely variable 
and of little taxonomic value. However, some species, e. g. 7. 
dioicum L., are characterized by having the lower branch of the 
inflorescence subtended by a long-petioled leaf. The base of 
the petiole is always more or less dilated. Even when the leaf is 
said to be sessile it is not truly sessile, for this dilated part of the 
petiole is always present between the stem and the leaf. Not 
uncommonly this base of the petiole surrounds the stem nearly 
completely. 

STIPULES AND STIPELLULES. The dilated base of the petiole 
is sometimes so much enlarged that its margins may properly be 
termed stipules. Such stipules are not constantly present nor 
is their size or contour of any taxonomic value in most American 
species. Stipellules are always absent in some groups, while in 
many a species, e. g. 7. Fendleri Gray, their form and size are 
inconstant. Stipellules of the first degree are most common 
but those of the second or third or even the fourth degree may be 


1944] Boivin,—American Thalictra and their Allies 341 


encountered. In most North American species stipellules are 
regularly lacking. 

Root. Roots are commonly fibrous, but in a few species they 
tend to be more or less tuberous, e. g. T. texanum (Gray) Small, 
T. debile Buckley and T. pinnatum Watson. _ A few species, e. g. 
T. confine Fern., may have horizontal underground stolons 
which will at their end produce a stem the following year, for all 
species of Thalictrum are perennial. The general condition is a 
stem simply thickened at the base, and from that thickened base 
all the roots depart and there also is produced the bud for the 
stem of the following year. 

INFLORESCENCE. The inflorescence is usually paniculate, 
sometimes subcorymbose, e. g. 7. polygamum Muhl. var. hebe- 
carpum Fern., or racemose, e. g. 7’. alpinum L. Except for the 
latter all Thalictra have a more or less branched inflorescence but 
are otherwise simple, there being only one inflorescence to a plant. 
An exception to this is 7. madrense Rose which has no definite 
inflorescence but is branched from the base, with solitary flowers 
or short panicles at the end of each branch. T. occidentale Gray 
var. palouense St. John sometimes has a simple raceme of mostly 
geminate peduncles. A few species are pauciflorous or uniflorous. 

PEepUNCLES. In most species peduncles are quite variable but 
in a few they will present very constant characters. They are 
shortest in 7’. venulosum Trel., longest in § CrycineriA, arching 
in 7. alpinum L., or strongly recurved below the receptacle in 
T. sparsiflorum Turez., T. grandifolium Watson and others. 

PERIANTH. In the subgenus Lecoyerium the perianth is 
typically of 4 sepals. These are usually small and fugacious, 
more or less oval in contour, and a few millimeters in length; 
they are of little taxonomic importance. However, in the 
§ Heterogamia they are dimorphous, those of the pistillate flowers 
being much smaller and narrower than those of the staminate 
flowers. As to their color, they are usually greenish at first, then 
more or less purplish, and generally withered by the time they 
drop off. 

Sex or Ptanrs anp Fiowers. All species of Thalictrum have 
perfect flowers except for those of the subgenus Lecoyeriwm in 
which the flowers may be dioecious, polygamous or perfect, 
according to the different sections of that subgenus. 


342 : Rhodora {OcTroBER 


Stamens. The color and shape of the filament also charac- 
terize the different sections. The exact color of the filament is 
unknown for many species from Mexico and southward. In 
some groups the filament is upwardly dilated or clavate, e. g. 
§ Leucocoma and § Physocarpum. The length of the filament 
and the anther varies within definite limits for each species; 
thus it is helpful in identification. The anthers also have a 
definite shape, from linear to globose, and may be obtuse at the 
apex or with a more or less well developed acumen. ‘Anthers 
are usually yellow and give its mass color to the staminate 
inflorescence. In the § Leucocoma the filaments may be respon- 
sible for this mass effect. Asa rule anthers and sepals will com- 
pletely hide the ovaries, this being the source of many an errone- 
ous statement on herbarium labels to the effect that the plant 
is a staminate one when a dissection reveals only or mainly 
perfect flowers. This error has not infrequently crept into the 
literature, even in descriptions of new species. Anthers may 
also be purplish, e. g. 7. polycarpum 8. Watson, or whitish, e. g. 
T. clavatum DC., or even greenish, e. g. T. hypoglaucum Rydb. 
The length of the anther given in the following descriptions is 
always the length of the anther exclusive of the acumen and 
immediately after dehiscence, for mature anthers are slightly 
longer before dehiscence than after, and as they wither and 
twist, they become still shorter. 

Ovary. At flowering time the ovary is generally too small to 
be characteristic but the stigma varies within definite limits for 
each species. As the ovary matures into the fruit the stigma 
usually undergoes a slight elongation, but very often drops off 
altogether. In quite a number of species there is no sharp 
demarcation between the style and the stigma. In such cases 
the length given for the stigma also includes the base of the style. 

Fruit. Nearly all species are based mainly on characters 
drawn from the fruits, for in most species mature fruits vary but 
little, while, on the other hand, they vary from once species to 
another. Characteristic features are: shape; length and breadth; 
thickness; length of stipe; number and design of nerves and 
ridges; number, direction and pabeuterie = the fruits; thickness 
of the fruit-wall.  ~ 

Nerves of the fruit when raised on sitelk’ often may not be 


1944] - Boivin,x—American Thalictra and their Allies 343 


set off ‘from the ridge itself, but they are usually conspicuous. 
Nerves are typically eight in number, one dorsal, one ventral, 
and three on each side. They usually branch off at the base 
and join at the summit of the fruit; they are then described as 
being simple. They may run straight from the base or arch 
over the central cavity or they may be simply curved, as in the 
case of the lateral nerves of a strongly flattened fruit. Nerves 
may also be sinuous, undulating, branching or anastomosed. 


Renation BETWEEN. STaMINATE AND PIsTILLATE, FLOWERING 
AND FRUITING MATERIAL 


This is a problem which is not yet thoroughly worked out and 
is far from being so. In many species this relation has been 
established on uncertain grounds, mainly on herbarium specimens 
bearing two or more fragments representing different phases of, 
presumably, the same species. Sometimes intermediate stages 
help indicate relation or an unusual specimen will bear both 
flowers and fruits. Of course, in this problem geographic dis- 
tribution is not altogether a negligible character. But many 
species are still known only in fruit or in flower and it is not 
impossible that the other stage has been collected and is passing 
under another name. 


INTERPRETATION. OF DESCRIPTIONS 


Most describers, when proposing new species of Thalictrum, 
have emphasized the division of the leaf, the contour, size and 
figure of the leaflets, the length of petioles and petiolules, all 
characters of little, if any, diagnostic value; while the type of 
pubescence, the length of style, filament and anther, the type of 
nervation and size of the fruit, were too often omitted. This 
unfortunate neglect renders many a description unintelligible 
unless one has access to the type-specimen. Consequently an 
effort was made to study all types that were available in America, 
for European herbaria are inaccessible for the time being. In so 
doing I was fortunate enough to have access to nearly all types 
of Thalictrum preserved in America. The United States Na- 
tional Herbarium had stored away a large number of its types 
but, except in two cases, isotypes were available. Photographs 
of some 10 types or syntypes along with numerous fragments 


344 Rhodora [OcTroBER 


and isotypes of species described in Europe have enabled me to 
understand a fairly large majority of these, while Lecoyer’s 
opinions and discussions greatly helped in interpreting the 
others. Unfortunately, a few names are still of dubious applica- 
tion and might, when the types are studied, antedate some of 
the names here accepted. 


TERMINOLOGY OF TYPES 


There is much variation as to the use and meaning of the 
terms applied to typical material. There follow the definitions 
of those terms needed and used in this paper to characterize 
such material.! 

Type: an herbarium specimen upon which the description of a 
new plant is based. If more than one specimen was used, the 
type is the specimen designated as such by the describer or by a 
later student of the group. 

SyntyPe: an herbarium specimen upon which the description 
of a new plant is based, when there are more than one, and no 
type has been selected. 

PARATYPE: any specimen, other than the type and the iso- 
types, upon which the description of a new plant is based. 
Duplicates of PARATYPES may be termed PARAISOTYPES, but we 
have preferred to extend the meaning of PARATYPE to include 
such specimens in all cases when the homogeneity of the collec- 
tion was not under suspicion. 

Isotype: a duplicate of a Typ, usually bearing the same 
collection-number. 

The best discussions of type-nomenclature I have found in the 
literature are: D. L. Frizzell, Terminology of Types, Am. Mid. 
Nat. 14: 637-668. 1933, and C. X. Furtado, The Nomenclature 
of Types, Gardens’ Bull. Straits-Sett., 9: 285, 308, both with 
bibliography. Frizzell’s paper is written by a zoologist from 4 
zoologist’s point of view; it is in the form of a lexicon, with 233 
terms, 10 of which he more strongly recommends. The second 
paper, the author of which is a botanist, proposes and defines 22 
terms and 21 compounds with the possibility of a great many 
more. 


' Since this study was completed before the clarification of the subject by Blake in 
Ruopora, 45: 481, 1943, no changes have been made in the text.—Ebs. 


1944] Boivin,—American Thalictra and their Allies 345 


Such terms as LECTOTYPE, LOGOAPOTYPE, NEOTYPE, PROTO- 
MEROTYPE, and many others have been proposed to express an 
historical relation between the selection of the type and the date 
of publication of the new taxonomic group. When it comes to 
the studying of a species, it is irrelevant whether you work with 
& HOLOTYPE or a LEcToTyPE. No term expressing an historical 
relationship has been found necessary in this paper for the proper 
understanding and concise treatment of a taxonomic problem. 
Furtado also distinguishes between IsoTyPEs and HAPTOTYPES, 
according to whether they were, or were not, collected from the 
same plant and at the same time as the type-specimen. Truly 
this is a very pertinent distinction and in many cases these 
terms, no doubt, may be useful. But, unfortunately, the average 
herbarium duplicate bears little or no evidence which might help 
in making this very important distinction. I have thus felt 
compelled to use ISOTYPE in a more general sense. 

Both Frizzell and Furtado recommend the use of HOLOTYPE 
for TyPE. This proposal is in accordance with the general use 
amongst the zoologists and would promote uniformity. But we 
have not departed in that respect from what seems to be at 
present the general use amongst botanists. To other terms such 
AS CLASTOTYPE, FOTOTYPE, and others, we have preferred the 
corresponding English expressions: FRAGMENT OF A TYPE, 
PHOTOGRAPH OF A TYPE, etc., for reasons of clarity and simplicity. 


Keys 


Characters in the artificial keys have been selected according 
to their usefulness, while those used in the general keys to sec- 
tions and species are either artificial or such as to place related 
species together. The numbering of species, however, is intended 
to follow natural relationships whenever possible. 


ACKNOWLEDGMENTS 


I wish to thank many of the people at Harvard University who 
have been of great help, and especially so Dr. M. L. PeRNALp 
under whose direction this work was carried out and who also 
supplied photographs of type-specimens of three of the species 
described in Europe, and Mr. C. A. WeatHErsy and Dr. R. H. 
Wermore for their numerous and helpful suggestions. This 


346 ‘Rhodora [OcroBEeR 


research would not have been ‘possible without the help of the 
late Brother Marie-Victor1n and a fellowship from the Govern- 
ment of the Province of Quebec. The following institutions 
have offered facilities for studying the herbarium material pre- 
served in their collections: Academy of Natural Sciences of 
Philadelphia (ANS); Brooklyn Botanical Garden (BG); Na- 
tional Herbarium of Canada (C); California Academy of Sciences 
(CA); Field Museum of Natural History (F) which also supplied 
eight photographs of types in European herbaria; Gray Her- 
barium (G); Missouri Botanical Garden (M); University of 
North Carolina (NC); New England Botanical Club (NE); 
‘Notre-Dame University (N-ND for the Nieuwland Herbarium 
and G-ND for the Greene Herbarium); New York Botanical 
Garden (NY); University of California (UC), including the 
Clokey Herbarium on deposit at the University of California 
(C-UC); the United States National Herbarium (US). The 
letters between parentheses are the abbreviations used through- 
out this paper to designate each of these institutions. 


THALICTRUM 


Thalictrum L., Gen. Pl., ed. 5, p. 242, 1754. Physocarpum 
Bercht. & Presl, O Priroz. Rostl. 1: 14, 1823. nec Physacarpon 
Necker, Elem. 2: 164, 1790, nec Physocarpus Maxim. Acta Horti 
Petrop. 6: 219, 1879, nec Physocarpos Cambess. Ann. Sc. Nat. 
1: 239 et 385, 1824, nee Physocarpa Raf. New Fl. Amer. 3: 
73, 1836. Ruprechtia Opiz, Sez. Rost. Kve. 86, 1852, nec 
Ruprechtia C. A. Meyer, Mém. Acad. St-Péters. 6 sér. 4: 148, 
1840, nec Ruprechtia. Reichb. Nom. 166, 1841. Leucocoma 
(Greene) Nieuwl. Amer. Mid]. Nat. 3: 254, 1914, nec Leuco- 
coma Rydb. Fl. Rock. Mts. 108, 1917. Sumnera Nieuw. 
Amer. Mid]. Nat. 3: 254, 1914. Plantae perennes, caule annuo 
herbaceo vel suffruticoso plus minusve sulecato, caudice perenni, 
vel interdum stoloniferae. Radices numerosae fibrosae, interdum 
tuberosae. Folia alterna, vel raro subopposita, vel opposita 
etiam, plerumque ternata et amplius composita. Floralia 
saepius desunt. JInflorescentia paniculata saepius racemosa vel 
subcorymbosa, nuda vel parum foliosa, ramis pedicellisque al- 
ternis vel subaggregatis. Petioli basi dilatati, saepius vel am- 
plexicaules vel subamplexicaules, plus minusve dilatati duarum 
modo stipularum vel nonnunquam etiam stipulati et. interdum 
stipellulati quoque. Flores numerosi, perfecti vel dioici vel 
polygami. Calyx sepalis hinc quatuor illine quinque nonnun- 
quam sex vel septem, fugax, viridis vel frequentius petaloideus. 


iets deed nin a ene ee Oe |! Parke REPL STE 


1944] Boivin,—American Thalictra and their Allies 347 


Corolla deest. Stamina libera plura vel numerosa, sepala fere 
semper per anthesim excedentia, numero variabili. Antherae 
adnatae comissura filamentis basilari nec articulatae. Ovaria 


unilocularia soluta, numero in diversis diverso et variabili, 
semine solitari apicali inverso. Stylus ie stigma vel 
similiter elongatum vel triangulare yal punctatum ferens per 


longitudinem fere totam. Carpella pauca vel numerosn, uni- 
seminata indehiscentia, sicca nervosa, plus minusve inflata, 

nquam stipitata, frequenter costata. Species typica: 
Thalictrum foetidum L. ut imprimis Britton & Brown proposuer- 
unt in Ill. Fl. 2: 118. 1913. 

The proposed standard species of the Linnaean generic names 
published in the International Rules of Botanical Nomenclature, 
p. 142, 1935, suggested that 7’. aquilegifolium L. should be taken 
as a type-species; but, in view of the fact that the latter species 
has been many times separated from Thalictrum to form a 
separate monospecific genus, it seems preferable to adhere to the 
older proposal. And, after all, T. foetidwm L. is much more 
characteristic of the genus as a whole than is T. aquilegifolium L. 


SUBDIVISIONS OF THE GENUS 


Two genera, Anemonella and Trautvetteria have been separated 
from Thalictrum and are usually accepted as distinct genera. 
Many more generic segregations have been proposed at one time 
or other but are not here considered worth recognition. Those 
synonyms are listed under the subdivisions which were thus 
raised to a generic rank. 

De Candolle in his Systema (vol. 1, 1817) proposed five sec- 
tions and four more subdivisions; Lecoyer (Bull. Soc. Bot. Belg. 
24: 111-113, 1885) was the first to recognize the fact that a 
natural classification of the genus sets the American species apart 
from the Eurasian ones but, unfortunately, the names he gave 
to his sections and subsections were not in Latin; Prantl (Nat. 
Pfl. 3, 2: 66) in 1888 proposed nine new subdivisions for the genus 
but he grouped the species in such a way that it is impossible to 
ascribe to each of his new names a definite position in the scheme 
actually adopted in this paper; E. L. Greene (Leaflets, 2: 49) in 
1910 proposed the family Thalictraceae to include Thalictrum, 
Anemone, Coptis, Aquilegia and Anemonella; and in 1914 Nieuw- 
land proposed to recognize Sumnera and Leucocoma as distinct 
genera. 


348 Rhodora [OcroBER 


KEY TO THE SUBDIVISIONS OF THE GENUS 


a. Owens —— — sub anthesim a ei, are persaepi 
Poe Se Gy es ea eyes Subgenus Thanicoon pea 8 
b. Stigma ‘bialatum deltoideum ventrale sessile. Filam 
if a et saepius ad junctionem cum ce Maer paul 
ulum in a rassata. Carpella aang ra costata sessilia... . 
c. Carpella matura art a ee as Bectio HoMornauicrnox. 
é. Carpella miahira compremea 2 0) ee Sectio UINA 
b. Stigma haud sian nec eatin vel, si anguste alatum, 


d. Filaments filiformia et saepius ad junctionem cum 
antheris paullulum incrassata. Stigma nonnumquam 
circinatum. arpella ata, 
nervis tamen valde rugosis, saepius breviter sapuate 
Sectio LErrostieMa. 
d. Filamenta staminum plus minusve clavata, ad junc- 
tionem cum antheris paullulum constricta, alba vel 
fescentia, raro purpurascentia; variant antherae 
globosae usque ad oblongo-lanceolatas; stigma nun- 
e. 


é. nso inflatus vel Goa Aiea apterus. .. .f. 
ff —_ us costatus sessilis vel subsessilis nec compres- 
sed nervis singulis saepius in costas elevatis. 
fhaae OUR Gas o6 ck cis ks aes cas Sectio ERYTHRANDRA. 
2 chico stipitatns plus minusve compressus nec 
tatus, nervis tamen rugosis. . . .g. 
g. sng "plus minusve compressus nervo dorsali 


convexiore vel curvato modum ventralis; 
stigma ventrale oe Bias nome pedicelli recti; 
foha caulinaria saepius desunt........ Sect: o PHysocaRPUo. 


g. Fructus valde pe a nervo ventrali pd 
charg folia eS is — et 3-5-ternata 
igma cylindraceum............ tio OMALOPHYSA. 
é vrata: ye pi ceatrtias stipite longissimo; 
WU WO a SS — io TRIPETRIUM. 
a. Flores dioici vel ee raro perfecti, stylus sub anthes 
sepala quaterna We i ss ek es Subge a is LmcovEnrow. 
h. Flores polygamo-monoie licet perfecti, stigma filiform 
elongatum, saepius cylin ee fe aap angu 
i tie. ” Sepala head dimorpha. .. . 
t. Ovaria pauca, solitaria, vel pacts licet raro 3-4. Pedi- 
celli longissimi tempore fructuum maturitatis. Flores 
: Pe a ieee sree ee sce sas ene Sectio CrNCINNERIA. 
i, Ovaria numerosa. Pedicelli ad 6 cm. longi. Flores 
polygamo-monoici, raro a Stigma, nisi in T. 
Johnstonvi, eylin — 


j. Foliolae aud PONS: oo oS Sectio CAMPTOGASTRUM. .. -k. 
k. Ovaria 10 vel pauciora. Carpella matura compressa. .. ./. 
U vide nervo ventrali nullo modo gibboso . Subsectio SiImPLICIA 
i. Carpella nervo ven WIG ioe ces ais o GIBBOSA 
k. Ovaria 18-36. wae Meco anc ong ventre globoso. 
Stigma “baseaparae clavatum.......... Subsectio VENTURIANA. 
Je Foliolae pelt a AS NS ee ae ta Dem gee iar .-.M. 


. Foli Speer petiolis prope marginem inse 
Subsectio SuBPELTATA. 
m. Foliolae ampliores petiolis circa tertias insertis. 
Subsectio EUPELTATA- 


1944] Boivin,—American Thalictra and their Allies 349 


h. Flores — vel de cee Macon stigma breve plus sores 
bialat Sepala ge rpha, floris foeminei minora. . . .n 
n. Hamer piconet filiform ia, flores dioici, rarissime 
polygami. Poliola apice trilobata, lobis persaepius 
CrONAGS ro. ee ee Sect tio HETEROGAMIA... .0. 
o. Carpella recta ventre symmetrico, nervis crassiusculis, 
nec compressa nec curvata. Radices fibrosae vel 
tuberosae, nec orgs igs Plantae glabrae. ...p. 
p. Radices tuberosae. ... 
gq. Carpella nervis  simplicibus nec sinuosis sed — 
4 cies weedea ig ls de Subsectio DeBILIA. 


C &@ nervis bee Priel ony eee Subsectio Sinvosa. 
p. ‘Radices bong es Petioluli articulati. Nervi car- 
i hau MAO kg ees a Subsectio Drorca. 


0. peel cena curvata vel compressa. Plantae chor ee 
s, plus minusve stoloniferae. .. . 
¢. spells ote crasso et firmo, sion Ls haud 
mpressa, costata, nervis haud distinctis : 
satin ee dorsali convexiore quam ventra 
Su bsecto INCURVATA. 
r, ae ee Slice minusve compressa, recta, vel bas 
minusve recurvata, vel valde Gidinewiad 
haves conspicuis, pariete nonnunquam mem- 
branaceo. .. .s. 
s. Carpella parum compressa, recta et nes Po 
stipitata, nervo ventrali convexiore 
QOPIREE SF is i oe on cee Su bsectio CLAVOCARPA. 
8. oe subsessilia vel, si stipitata, stipite n 
rvato vel nervo ventral Genie eva quam 
Ee dota. vel valde comp my 
12 t. Crassitudo carpelli denies latitudinis partem 
1% vel attingit vel superat........... age CoMPRESSA. 
ig t. Crassitudo carpelli a dimidiis latitudinis ree 
@ Su ne LAMINARIA. 
n. Filamenta alba saepius plus minusve clavata, flores 
ol a oici. Foliola integra, vel, si apice trilobata 
fobys IIS acs ie Soils Vs Mie 8 cei eee Sectio Leucocoma. 


The following treatment takes into account all species attri- 
buted to America, plus, when a subdivision of the group is not 
confined to America, a brief world-treatment of that subdivision. 
However, the § Genutna is not fully treated, for only two, or 
perhaps three, of its numerous members have been collected in 
America. To add a full treatment of this section to this paper 
would have transformed it into a world-monograph, a study not 
possible under the present conditions. Similarly § OMALoPHYSA 
is incompletely treated for lack of sufficient herbarium material. 

Subgenus THaticrrum (DC.) Reich. Sect. Huthalictrum DC. 
Prod. 1: 12, 1824. Sect. Thalictrum DC. Syst. 1: 172, 1817. 
Subg. Thalictrum (DC.) Reich. Consp. Roe. Veg. 192, 1828. 
Plantae radicibus persaepius fibrosis, caule plus minusve foliosa, 
foliis stipellulatis vel estipellulatis. Inflorescentia ampla, saepius 
paniculata, raro subcorymbosa vel racemosa. Flores perfecti. 


350 Rhodora [OcroBER 


Sepala saepius 5in flore. Stigmata in eons a sepalis recedentia. 
Species typica Thalictrum foetidum 
ectio Homothalictrum. HH omothalictrum Fries, Sum. Veg. 
Sean. 27, 1846, nomen nudum. Sec t. Camptogastrum f. Flexuosa 
Prantl, Nat fl., 3: 2: 66, 1888. co Pileiuants filiformia nonnun- 
quam ‘Purpurascentia. Stigma bialatum deltoideum ventrale 
rpella matura costata sessilia nec compressa. Species 
typica soto T. minus L. sit. 

As created by Fries, his genus Homothalictrum included only 
T. alpinum L.; and I was also at first under the impression that 
the simple raceme was a character sufficient by itself to set off 
this species from the rest of the genus. But this character must 
be considered rather as an extreme, especially if one compares 
this species with its nearest relatives, 7. Esquirolii Lévl. & Vant. 
and 7’. isopyroides Meyer. Amongst the species to be referred 
here besides T. minus L., there are: T. alpinum L., T. angusti- 
folium Jacq., T. Esquirolii H. Lévl. & Vant., T. flavum L., T. 
galioides Nestl., T. isopyroides Meyer, T. kemense Fries, T. 
punduanum Wall., T. rugosum Ait., T. simplex L., T. squarrosum 
Steph. 

1. THaLicrRuM MINUs L, 

To this species can be referred a specimen from the herbarium 
of John Ball and preserved in the Gray Herbarium. It was 
collected in Greenland by Giesecké, so says the label. C. L. 
Giesecké was a geologist who, between 1806 and 1813, made 
collections of plants on Greenland. His report was published in 
Brewster’s Edinburgh Encyclopaedia, 10: 100-102, 1832, and 
the only Thalictrum listed is T. alpinum. Thus it is not at all 
improbable that the label was pasted to the wrong specimen. 
That label, however, reads: Thalictrum Greenland Giesecke. 

T. R. Jones, reviewing the botanical literature on Greenland 
in the Man. Nat. Hist. Gr., p. 256, 18751, casts a doubt on the 
accuracy of Giesecké’s list. ‘His list’’, he writes, ‘ ‘compre- 
hends a large number of species, but he is manifestly wrong in 
regard to many of them. Some, which may possibly be per 
bers of the Greenland flora, have never been found since his day.” 

Giesecké having spent eight years in Greenland studying the 
mineralogy of the country, it is not impossible that he might 
have chanced to botanize on peculiar spots where the local 

1 See also Trans, Bot. Soc. Edin., 9, 2: 430-465, 1868. 


| SSRI eet EN ee ee aR ae secs er re or eae ane eT 


a ST ee ee HG Ere ze. lg BR in 


1944] Boivin,—American Thalictra and their Allies 


H 
I2e 1Sb 


Deraits or Ftowers or Fruits or THaAticrrvum, all X 6 


(For explanation see end of paper. Notealso that the “c’’ of “14c” has been 
omitted by error.) 
geological formations supported an endemic flora. Greenland is 
reputedly well known botanically, but as recently as 1937, 
Polunin reported seven new species from the Julianehaab sector 
alone and confirmed six doubtful previous reports. Although 
the flora of that ‘subcontinent’? shows more affinities with 
America than with Europe, the presence of T. minus sensu 
amplo can not be rejected a priori, for it is a species ranging over 
Eurasia and Africa from the Bering Straits to Ireland and from 
Arctic Norway to central Africa. For the time being it is, 
however, preferable to wait until some other field-botanist 
confirms Giesecké’s specimen by collecting the plant anew. 

Lecoyer in 1885 recorded over 200 synonyms of 7. minus and 
no doubt many others have been coined since that time. In 
view of the types of these being all in Eurasian herbaria and also 
because that species is rather scantily represented in American 


352 Rhodora [OcroBER 


herbaria, it is preferable to leave to some other worker, who 
might have access to the types, the task of disentangling, 7f 
possible, the status of this species, for, as Prantl characterizes it, 
very accurately indeed, it is a P iornien buon’: 


2. T. Hultenii sp. nov. Planta 35-70 em.1, omnino glabra nis 
in cauli. Radix ignotaest. Caulis arenoso-pr uinosus sub tempus 
florendi. Folia caulinaria 2-4-ternata, internodos subaequantia, 
superiora et intermedia in apice dilatationis petiolaris sessilia, 
inferiora plus minusve petiolata. Stipulae auriculatae, plus 
minusve laceratae. Stzpellulae desunt. Foliolae variabiles, 
saepius subquadratae, apice trilobatae, ca. 1.5 cm. long. Inflo- 
rescentia paniculata 5-10 cm. long., parum ramosa. Pedunculi in 
axilla saepe folii ternati, saepius folii simplicis, rarissime bracteae. 
Flores omnes perfecti, pauci in planta. Sepala paullulum cucul- 
lata, elliptica, ca. 3.0 mm. long., supera erosa, albicantia, plus 
minusve luteo-tincta. Stamina 8-15 in flore. Filamenta pallida 
flavescentia ca. 3.5 mm. Antherae oblongae vel oblongo-lanceo- 
latae, flavae, ca. 2.0 mm., mucrone brevi ca. 0.1 mm Ovaria 
3-6 in flore, stigmate 0. 4-0.7 mm. Carpella matura ignota, fere 
matura sessilia ventre ovoideo vel subgloboso. Floret Julio et 
Augusto. AtasKa: Hultén 6753, Unalaska, July 20, 1932 (NY, 
TYPE); Van Dyke 38, Un alaska, Makushin Bay, moist grassy 
places on hillsides, July 16, 1905 (G); J. M. Macoun 115, Una- 
laska, July 25, 1891 (G, NY); Hall, Unalaska, Tliuliuk, Oct. 1, 
1871 (G); Harrington, Unalaska, Iliuliuk, Oct. 12, 1871 (G); 
Evermann 120, Amakuak Island, Aug. 1, 1892 (F); Eyerdam 4 

nalaska, Aug, 22, 1932 (NY): Kincaid, Unalaska, Sept. 1 
1897 LUC); A E.& R. T. Porsild 641, Inana River, Hot Srna 
64° 20’ W., recent clearings in the Nesp of recently 
cick tose July 8, 1926 (G). Fie. 

This species has hitherto passed as either 7. ae Fries or 
T. majus L. or T. minus var. kemense (Fries) Trel. It differs 
mainly in the size of the fruit and by its contracted leafy inflor- 
escence. Dedicated to Ertc Huurén, the author of “Outline of 
the History of Arctic and Boreal Biota during the Quaternary 
Period’, a fundamental book on the phytogeography of the 
Arctie-boreal floras, and of detailed works on the floras of 
Kamtchatka, the Aleutian Islands and Alaska. The habitat of 
the last cited specimen suggests that the plant might have been 
somewhat overlooked on continental Alaska. 

3. T. aLpinum L. Planta plus minusve stolonifera, erecta 


1 Where not otherwise indica all measurements in descriptions apply to the 
height of the plant, or the moe of: the organs concerned. 


ST AW RS aC I Sin | a a TAN alt ny any wicca Saag Sle UL daa Me aMia ae 


1944} Boivin,—American Thalictra and their Allies 353 


rigida, omnino glabra (0.5)-1.5-(38. 0)}-decimetralis. Radices 
fibrosae. Folia bi-quadriternata, basilaria caulinarium 
deest vel raro solitarium, folia inferiora hanveneiias interdum 
lanceolata integra, raro trilobata, saepius desunt, superiora 
semper desunt. Foliolae flabellatae, coriaceae, inferne plus 
minusve glaucae, 1 cm. longae vel brevi lores, apice 3-7-lobatae, 
margine plus minusve revoluto, nervis rugosis, inferne rugosiori- 
bus. Inflorescentia nuda racemosa, e pedunculis alternis ante 
anthesim adpressis, postea recurvatis. Sepala 1.5-2.3 mm. 
longa, ovata vel elliptica. Stamina 8-15 in floribus singulis. 
Filamenta ca. 3.5 mm. filiformia, ante anthesim aliquo modo 
flavescentia, postea tamen purpurascentia. Antherae 1.2-2.3 
mm. longae, oblongae vel lineares, apice acutae, etsi mucronatae 
breviter etiam, one flavescentes deinde ee oe 


pedicellis singulis, reflexa vel et a, 2.8-3.5 cm. longa, sub- 
globosa vel oblanceolata basi saepius attenuata vel stipitata, 
apice rotundata, nervata, nervis ca. 12, rostro ad tergum valde 
eflexo. Floret a Junio ad Septembrem. 
The three following varieties seem to be worth recognition, 
although the characters of each are not absolutely spon 


3a. T, ALPINUM var. typicum. T'. alpinum L., Sp. PIl., 1: 545, 
1753. T. alpinum var. pallidum Norman, Arch. Math. aia, 
8: 4, 1883. T. alpinum var. gaspense Greene, Ott. Nat. 23: 18, 
1909. T. alpinum var. sour ables ne Greene, I. c. 1909. T. alpi- 
num var. nesioticum Greene, l. c. 19, 1909. T. alpinum var. pudi- 
cum Greene, |. c. 1909. Planta ‘stolonibus plus minusve elongatis 
horizontalibus, raro caespitosa. Radices paucae saepius sparsae. 
Foliolae supernae laete virides saepius valde lucidae nec glaucae. 
Ovaria saepius obovata vel subglobosa, sessilia vel subsessilia. 
Carpella matura stipite 0.5 mm. vel breviore vel deente.—GREEN- 
LAND: Vahl. f.(G); A. E. Porsild 257, south coast of Disko Island 
near Godhavn, 69° 14’, fertile slopes near the warm springs, July 
27-28, 1937 (G); Seidenfaden & Ostenfeld 296, East — 
northern side of the head of Husk-ax Fiord, 73° 38’, Aug. 1 
1929 (G); M. P. Porsild & R. T. Porsild, Vest-Gréal. ‘Ubekendt 
Eijland, Igdlorssuit, 71° 14’, 21 juli, 1929 (G); Eugenius, Groenl. 
occid., ca. Neria, 61° 33’, 26 ‘juli, 1935 (G). Lasrapor: Bell 874, 
Cape ‘Chudleigh (C): Fernald & Wiegand 2430, Blane eer 
limestone and calcareous sandstone terraces, Aug. 6, 1910 (G). 
NEWFOUNDLAND: Robinson & Schrenk 26, marsh near confluence 
of Exploits River and Badger Brook, Aug. 13, 1894 ear G, 
isotypes of T. alpinum var. microspermum) ; Fernald & Long 

8,272, Quirpon Island, wet boggy tundra in dius Valley, 
Ke 7, 1925 (G); Fernald, Long & of 1708, Old Port au Choix, 


354 Rhodora [OcroBER 


turfy and peaty margins of dry gravelly limestone barrens, July 
22, 1929 (G); Fernald & Wiegand 5443, Millerton Junction, dry 
bog, July 7, 1911 (G); Fernald, Long & Dunbar 26,690, Brig Bay, 
peaty and ‘turfy knolls or slopes on limestone barrens, Aug. 6 
1924 (G). Qu&Berc: Victorin & Rolland 25,441, Minganie, tle 
Saint-Charles, prés de la tourbiére du lac Salé, 20 juillet, 1926 
(G); Victorin & Rolland 21,156, Minganie, Grande Ile, talus 
humide sur le rivage, avec Anemone parviflora, 20 juillet, 1925 
(G); Pursh, Anticosti (G); Macoun 3, Anticosti, valley of Jupiter 
River, Aug. 20, 1883 (C, type of T. alpinum var. nestoticum; G, 
isotype) ; Victorin, Rolland & Louis-Marie 21,156, Anticosti, 
riviére Vauréal, sur les talus humides le long de la riviére, 27 
juillet, 1925 (G); Fernald, Gaspé Co., banks of the Grand River, 
damp ledges, June 30-July 3, 1904 (C, type of T. alpinum var. 
gaspense, G, isotype); Fernald & Collins 1046, Gaspé Co., Bona- 
venture Island, Bonaventure conglomerate (calcareous) sea- 
_ Aug. 7 & 8, 1905 (G); Pease & Smith 25,761, Matane Co., 


Juneau, eee alpine, June 30, 1 40 (C 7A: E & R. T. Porsild 
789, Norton Sound, Pastolik, bor 22’ N., 163° W., coast and low 
swampy tundra, growing on the sides of large “niggerheads”’ , hot 
common, July 20, 1926 (G); J. P. Anderson 452, Juneau, July 
4, 1917 (NY). SrBeria, PRIMORSK: Wright, Behring Straits, 
Arakamtchetchene Island, 1853-1856 (G, US); Tiling, Ajan 
(G). yaxutsK: Nilson, Balaganach, June 27, 1898 (US). 
TRANSBAICALIA: NV. & V. Ikonnikov-Galitzky 288, montes Kentei 
orientales and fontes fluvii Kerulen, July 18, 1928 (NY). YEN- 
ISSEISK: Kusnezow & Reverdatto 973, distr. Turuchansk, 1914 
pr sesaaboay GY). Rear Krylov, Altai, Margalinsky 
prope Nishnij Uimon, June 11, 1901 (NY); 
Sehisciin, Chilikina € Sumnevicz, Altai, i in jugum Terk-Tynsky 
fluminis, Kostachta, July 4, 1931 (NY). TURKESTAN: Saposhon- 
kov, in jugo Tian-Schan, in alpinis, Ju ne 28, pie Modif: Russia, 
ARCHANGEL: Steffen 567, Vaygach Is., Aug. 16, 1926 (C). VoLe- 
pa: Soczava, montes uralenses inter 62° et 64°, ‘Tuly 20, 1928 (C). 
Nova ZEMBLA: Weber, 1891 (F); Tolmatschew 416, insula sep- 
tentrionalis, in decliviis siccis ad flumen Notschujew in vic. 
eho, ia Matotschkin, July 23, 1922 v. (G). SprrzBERGEN: 
King’s Bay, July 6-8, 1935 (G). Lavrpronra: Anderson 

84, 1869. SweEvEN: Ortendal, Ostersund, Jmelland, 1890 (G); 
Asplund, Tornetriisk-omridet, Nuoljas nordsida, July 19, 1917 
(F); Sjorgren, in alpe Areskutan, 1846 (G); Alm 6523, 'Torne 


1 Abbreviated elsewhere throughout this paper as v. or g. 


TE 


pla cle ER i oe ie Ct DG EL tt apnea ea aha ae eee eta a NR cats Rie en Ate ess ante ace nai, ae ale 


1944] Boivin,—American Thalictra and their Allies 355 


Lappmark, Jukkasjirvi s:n, in monte Kaisepakte, alt. ca. 750 m., 
Toy 18, 1926 (G): Tedenius, Herjedal bor. occid. a Malmagen, 
julio ( ). ORWAY: aylor, Ruostavand, Tromsé, July—Aug., 
1907 (G); Fidtz, Opdal i Sér-Tréndelag, Kalvellafossen, Aug. 11, 
1910 (G); Bocck, ee feed. Hid, Voss: Finnesleigen, i fjéra ved 
Vangevalnet, 46 m “si Faro Isuanps: Taylor, Videré 
Island, Videreite, "1903-19 6 (G); Hartz & ees insula 
Suderé, Kvalbéfjaed, Tay. 21, 1897 (G). take 
Jack, Perthshire, Ben Lin, July , 1922 (G) ; Clova, 1832 32 (CG). ENG- 
LAND, DURHAM: Edmondson 4691, upper Teesdale, Widdybank, 
wet places, June 27, 1909 (G). WESTMORELAND: Beanland, 
Tarn Craggs, Helvellyn, June 29, 1909 (G). YorRKSHIRE: Lees, 
alt. 1600 ft., June 1872 (F). ROSSSHIRE: Stables, moor behind 
Rosebough, growing with Si ge among cowfields, June 1, 
1843 (G). Wats: Ball, North Wales, mountains near Llan- 
beris, May 30 & 31, 1859 (ry: Ball, North Wales, Carnarvon- 
shire, rocks on Snowdon, 1850 (G). IceLanpb: Seidenfaden 969, 
eastern Iceland, Seydisfjord, 65° 17° Lert heath, June 27, 1929 
( . Sérensen, Skagafjord, July i, 0 (G); Paulsen, SW- 
Iceland, Haon efjord, Hvanu, July 4, 1904 (G); ‘Scamman 1871, 
Lakjamot, a farm in the north on the road between Stadur and 


2-19, 1938 (G); Taylor, N. Iceland, Sula mountains nw. of 
Akuseyvi, June 27, 1895 (G). Type, not seen, from Scandinavia, 
presumably preserved at the Linnaean Society of London. 
Fies. 3a, b and e 


Var. pallidum Norman, is the early stage of the plant when the 
leaflets are strongly glaucous beneath. Type not seen. 

Greene’s four varieties are rather descriptions of individuals 
than of taxonomic units. Types in the National Herbarium of 
Canada seen, except of var. pudicum for which Greene did not 
authenticate nor indicate any typical collection nor any definite 
distribution. The characters stressed by Greene to separate his 
varieties are mainly the following: height of the plant; average 
number of leaflets per leaf; curvature of the pedicels; number of 
carpels per flower; insertion of the carpel. All these characters 
exhibit such a degree of variation that the isotype of var. micro- 
spermum at the Gray Herbarium does not correspond to the 
description of the type as to the height of the plant (10 to 14 
inches), the number of leaflets per leaf (13 to 23), as to form of 
leaflets (often broader than long and as often 4—5-lobed as 3- 
lobed), the description of veinlets (either prominent or not), 
curvature of pedicels (straight to strongly recurved), number of 


356 Rhodora [OcTOBER 


carpels (one or two per flower, the specimens being overmature 
and most of the fruits gone), and insertion of the carpels (defi- 
nitely short-stipitate). And so on. 

The following year', Greene proposed seven new species, all 
based on material from the western United States. Plus the 
characters already discussed, he emphasized the following: 
leaflets shining or dull, pale or dark green; sex of the plant; size 
and form of the stamen; length of petiole; size and form of the 
leaflets. Of these seven species we can say the same as has 
already been said about the varieties. Two characters, however, 
prove to be rather constant and characteristic of the western 
American specimens. In most cases the leaves are of a pale green 
and always dull on the upper surface, the latter character being 
the most constant of the two. On the other hand, all the 
material from Gaspé Peninsula to Greenland and from Alaska 
has leaflets of a rather dark green color and definitely shining on 
the upper surface. A few, very few, intermediates were en- 
countered. Such an intermediate was Hall & Harbour 10, from 
‘ Colorado, which happens to be the type-collection of Thalictrum 
scopulorum Greene. Some specimens from the Pyrénées are 
equally more or less shining on the upper surface of the leaflets. 

4. T. atpinum var. hebetum, var. nov.—T. cheilanthoides 
Greene, Leaflets, 2: 89,1910. 7. duriusculum GREENE, l. c. 92, 
Ol. T elegantulum ‘GREENE, 1, -¢, 03, 1910. 7. leiophyllum 
GREENE, |. c. 90, 1910. T. monoense GREENE, l. c. 93, 1910. 
da scopulorum GREENE, l. c. 91, 1910. T. suspensum GREENE, 
L¢@,.: O02, 1910. —Planta parum stolonifera, raro caespitosa, 
stolonibus elongatis vel ascendentibus. Radices paucae aggre- 
gatae. Foliolae superne hebetae vel raro parum lucidae, saepius 
plus minusve glaucae, interdum viridulae. Ovaria obovata vel 
oblanceolata, sessilia vel subsessilia. Carpella agen stipite 
0.5 mm., vel breviore vel deente——Wvyomine: EF. Nelson 648, 


y Co., Fox. Park, on hummocks in bogs, th 8, 1907 


1 Leaflets, 2: 89-93, 1910. 


EE a lll hl llr SS lS lc Sel = 


1944] Boivin,—American Thalictra and their Allies 357 


& E. S. Clements 319, Seven Lakes, alt. 3500 m., 4 julii, 1901 
(G); Hermann 5544, Garfield Co., 134 miles sw. of Trappers’ 
Lake, trail to Flat Top, low moist meadow at edge of spruce 
woods, alt. 9500-10500 ft., July 31, 1933 (G); Clokey 3533, 

affee Co., Grizzly Peak, moist meadow, alt. 10,500 ft., May 7, 
1919 (G); Wolf & Rothrock 94, 1873 (G); Parry 75, from the head- 
waters of Clear Creek and the alpine ridges lying east of Middle 
Park, 1851 (G); Crandalle & Cowen 30, vicinity of Como, moist 
soil along stream, alt. 9775 ft., Aug. 3, 1895 (G); Osterhout, 
Laramie Co., North Park, July 25, 1898 (G); Baker, Earle & 
Tracy 637 {and not 437], above Durango, Hamor’s Lake, July 
24, 1898 (G-ND, type of T. suspensum; G, photograph of the 
type; NY, US, isotypes; Brandegee 890, Sierra Sangre de Cristo, 
ae 1879 (UC)). Uran: Porter, Uinta Mountains, July 25, 1873 
ANS 


358 Rhodora [OcroBER 


basses Alpes, Larche, Arvet Touvet, Guiguet, ep e du Var 4 
Esteine, dans les prairies tourbeuses, 16 juin, 1875 (G); Alpes 
maritimes, Lac de Vens, 29 juin, 1898 (F); ae Alpes mari- 
“_ St. Etienne, Lac de Rabuons, gazons humides, alt. 2410 

, 7 sept., 1919 (F). PYRENEES: "Rugel, Pyrénées-Orientales, 
val d’Eynes (G, NY); Suzzet, val d’Eynes, 11 juillet, 1890 (F); 
Trat 51a, Pyrénées Orientales, vallée de Carol, in pascuis editiori- 
bus prope Porta, 1 juillet, 1846 (NY); Bordére, Hautes-Pyrénées, 
pic du Campvieil, 1891 (F); Bordére, Hautes-Pyrénées, Tré- 
mouse, aotit 1864 (IF); Irat, Hautes-Pyrénées; Vignemale, ad 
nives, 3 sept., 1846 (F); Pyrénées, Port de Pinéde (F). Spatn: 
Bourgeau, Castanesa, Bassibes, aotit, 1847 (G). 

5. T. ALPINUM var. STIPITATUM Yabe, Bot. Mag. Tokyo, 17: 
20, 19038. T. marginatum Royle, Ill. Bot. Himal. 1: 51, 1839. | 
} ig aN we Royle, |. c. 1839, nec 7. microphyllum Anders. 

c. Bot. Belg. 24: 292, 1885, nomen nudum, ex synonymis 
Thalictri minoris L. var. elati Lec. T. brackeifilum Bertol. Mem. 
oa oe Bolog. II, 3: 422, 1848. ? 7. acaule Cambess. ex Hacque- 
t, Voy. nd. 4, Botanique, 3, 1844. ? TJ. alpinum var. 
sei Ulbrich, Not. Bot. Gart. Mus. Ber. 10: 877, 1929. T. 
alpinum var. microphyllum Handel-Mazzetti, Symb. Sin. 7, 2: 
311, 1931. T. nippono-alpinuwm Honda, Bot. Mag. Tokyo, 53: 
334, 1939.—Planta stolonifera et caespitosa, stolonibus paucis 
elongatis horizontalibus, pluribus ascendentibus. Radices num- 
erosae fasciculatae. Foliolae supernae saepius viridulae, non- 
nunquam paululum glaucae, interdum parum lucidae. Ovaria 
stipitata oblanceolata. Carpella matura stipite 0.5-2. : mm. 
ongo.—Nippon, HonbDo: Takemoto 591, Sinano, Aug. 9, 1923 
(G). Cuina, szEcHWAN: Rock 17,865, northeast of Kula, Mount 
Siga, on alpine meadow, alt. 4600 m. , June, 1929 (F, N NY, US); 
Rock 16,253, west of Muli Gomba, Mount Mitzuga, high alpine 
screes, alt. 4780 m., June, 1928 (US); Rock 5554, Muli Kingdom, 
alt. 16, 000 ft., June, 1922 (US); Rock 16,405b, Kondaling, Rison- 
quemba, Mount Konka, on alpine meadow, alt. 4770 m., June- 
hug. ., 1928 (US). yunnan: Handel-Mazzetti 4696, ad austro- 
orient. pagi Dschungdien (Chungtien) in lateris occid. montis 
Piepun, regione alpina, in glarea mobili, substr. calceo, alt. 4300- 
4650 m., 11 aug., 1914 (US); Rock 4951, Yangtze watershed, distr. 
of Likian ng, eastern slopes of Likiang Snow Range, on limestone 
gravel, alt. 16,500 ft., July 8, 1922 (US); Rock 9444, Yangtze 
watershed, eastern slope of Likiang Snow Range, alt. 14,000 ft., 
jay 1923 (US). Inp1a, stxxm: J. D. Hooker, regio alp. alt. 
000 ped. (G). KumMaon: Strachey & Winterbottom 1, Barjikang 
vee, alt. 15,000 ft., (G). Garuwat: Strachey & Winterbottom 
2, Bompras, alt. 14 500 ft. (G); Duthie 3795, above steeping ae 
12, 000—13,000 ft., "Sept. 16, 1885 (US). punsaB: Koelz 7 
Lahul, Khaksar, alpine meadow, alt. 15,000 ft., July 29, i936 


| UF et a Se Oe 


——— ee 
rr Fa a 


1944] — Boivin,—American Thalictra and their Allies 359 


(NY); Koelz 116, Kulu, Chanduakani, ~~ Phos ae ft., _— (NY); 
Koelz 8610, Chamba, Kukti Pass, alt. ft., Jul y 3, 1936 
(NY); Koelz 5039, Kangra, Rotang La, ee 12, 000 {t., "tin 30- 
July 1, 1933 (NY, US); Koelz 6851, Kangra, Lahul, oars 
Gongma, alt. 15,000 ft., Aug. 29-30 (US). KASHMIR: Thoms 
Kishtwar, regio ‘alpina, "alt. 11 ,000 ped. (G); Thomson, Ladakh 
regio alp. alt. 15-16,000 ped. (G, NY); Falconer 22 (G); R. 
Stewart 9776a, Sonamarg, alt. 12,000 ft., July 23, 1928 NY), B 

R. Stewart 19,873, ae ee Burzi Bass, alt. ca. 14 ,000 ft., 
July 27, 1940 (N TY): 9,997, Deosai, near Mir 
Panzil Pass, alt. ca. 14 060 ti san 28, 1940 (N Y); R. R. Stewart 
18,275, to Gadsai from Gangabal, alt. ca. 12, 000 ft., Aug. 11, 
1939 (NY); T. T. Stewart 7814, Tulion, above. Pahlgam, bree - 
13,000 ft., July 13, 14, 1925 (NY); Koelz, Rupshu, 

among dry rocks along stream, alt. 15,000 ft., June 30, 1931 
(NY, US). Fias. 5a and b. 

No type material has been available of any of the taxonomic 
units concerned. Var. elatum Ulbrich was described from speci- 
mens having the flowers of T. alpinum L. but the vegetative 
characters of 7. Esquirolit Lévl. & Vant. The flowers of those 
two-species are not very different. One flowering specimen from 
the Burma-Thibet frontier, Ward 9804, can be referred here only 
doubtfully until the fruits are known. 


6. T. rucosum Aiton, Hort. Kew. 2: 263, 1789. T. flavum L. 
& speciosum Reichard, ‘Syst. Pl. 2: 648, 1779. T. speciosum 
(Reichard) Poir. Eneye. Bot. 5: 315, 1804, nec TJ. speciosum 
Miller, Gard. Dict. ed. 8, no. 2, 1768. T. glaucum Desf. Tabl. 
de l’Ecole de Bot" 2: 146, 1815, nomen nudum. T. crenatum 


ae 
DC. Syst. 1: 184. 1817. 7. glaucum Desf. ex DC. 1. ce. 1817. 
T. rugosum Ait. @ umbelliferum DC. 1. ec. 185, 1817. T. discolor 
Willd. ex Lec. Bull. Soc. Bot. Belg. 24: 367, 1885, nomen nudum. 
T. purpurascens L. var. rugosum (Ait.) Farwell, Pap. Mich. Ac. 
Sci. 26: 11, 1941. 

T. rugosum Ait. was described from material cultivated in the 
Botanical Garden at Kew and was said to have been introduced 
from America. But Lecoyer, who has seen the type specimens, 
states that it is T. glaucum Desf. Apparently the Kew Gardens 
distributed seeds from the type-colony of T. rugosum Ait., for 
John Bartram cultivated it and so did Purdue University as late 
as 1924 and the Missouri Botanical Garden in 1902. Schweinitz 
got the plant from Bartram for his own harbarium and Muhlen- 
berg also had it as one of his four American species. There 


360 Rhodora [OcToBER 


seems to be no doubt that the plant was never native in America 
and that Aiton’s surmise was erroneous. A. P. De Candolle also 
saw the plant cultivated in a European garden under the name T. 
rugosum Ait. Apparently this plant was cultivated in America 
during the eighteenth century, at least in the United States, 
Mexico and Peru. A photograph of the type of 7’. densiflorum 
HBK. was lent to me by the Field Museum. It shows a plant 
in all points similar to 7. rugosum. The name 7’. rugosum Ait. 
antedates both T. glaucum Desf. and T. speciosum Poir. and 
must supersede them as the oldest name of the plant of southern 
Europe and northern Africa. 

Sectio Genuina (DC.), stat. nov.—Sect. Thalictrum, subsect. 
Genuina DC., Syst. 1: 169 & 175, 1817. Species typica 7’. 
foetidum L. , typus generis, sit (p. 347). 

Sectio Leptostigma, sect. nov. Stigmatibus Sea SEEN 
nec alatis. Species typica 7. saniculaeforme DC. s 

Among other species belonging to this group are: 7. Dalzelli 
Hook., 7. Falconeri Lec., T. rotundifolium DC., T. rufum Lec., 
T. Rochebrunianum Franch., T. tuberosum L., T. virgatum Hook. 
f. & Thom. None of them grow wild in America. 

Sectio Erythrandra, sect. nov. Filamenta antherarum apice 
clavata, alba vel rufescentia. Carpella sessilia vel breviter 
stipitata, costata, inflata. Stigma nunquam alatum nec sagit- 
tatum. Species typica T. petaloideum L. sit. 

A section including 7. calabricum Spreng., T. Fortunei Moore, 
T. javanicum Blume, T. podolicum Lec., T. sachalinense Lec., T. 
thibeticum Franch. and others. This section is not represented 
in ne on 

T. LEUCOSTEMON Koch & Bouch., Append. Ind. Sem. Hort. 
Berol 13, 1854. 

This species, erroneously attributed to America, probably 
belongs here. According to Lecoyer there does not seem to 
exist any type for this species. 

Sectio Paysocarpum DC.—Syst. 1: 171, 1817. Physocarpum 
Bercht. and Presl, O Priroz. Rostl. 1: 14, 1823, ut genus. 
Subg. a ook regs 8 Cons. Regn. vee 192, aN 


ut genus. Clavata Small, Man. South. Fl. 524, 1933, divisio 


1944] Boivin,—American Thalictra and their Allies 361 


ordo cujus indeterminatus. —Plantae saepius glabrae, foliis 
eatenaliniatee: caulinariis paucis nisi desunt, radicibus nonnun- 
quam tuberosis. Pubescentia, si adest, e pilis uniseriatis trans- 
lucidis. Inflorescentia corymbosa, nonnunquam pauciflora, raro 
elongata. lies perfecti. Sepala spathulata vel obovata 
interdum unguiculata. Filamenta alba, clavata ad apicem. 
Antherae athe sal pallido-luteae. Carpella matura pauca stipi- 
tata nervosa, plus minusve compressa, nec costata, nervis tamen 
saepius rugosis et fere simplicibus. Stigmata brevissima vel 


dentia, marcescentia. Species typica 7. clavatum DC. sit 

This subdivision of the genus was first created by De Candolle 
as a section and all five synonyms given were based on sect. 
Physocarpum DC. It included at first four species, three of 
which are now in other sections. Many subsequent authors 
have recognized this unit, always retaining 7. clavatum and 
dropping the other three species. This procedure is followed 
here, for the characters given by De Candolle clearly show that 
he regarded that species as typical of his newly proposed section. 

As most of the types of species belonging to this section are 
not accessible, it has not been possible to give a complete synony- 
my except for the American species. Furthermore, the material 
7 at hand is not quite abundant enough for a thorough treatment 
, : of this section. Since Lecoyer in his monograph treated only 

three species of this group, it is thought desirable to give a key 
to the sixteen species I recognize, cite at least some collections of 
each species and give a full treatment of the American species 
) only, leaving out of the picture all species for which there is no 
: herbarium material available to me. 
a. Folia caulinaria os 2-6, stipulis laceratis. Carpella 


14 matura lignosa, ven re globoso, nervis obscuris. .. . 
6. Stigma ovatum vel pd, contig semimillimetrale vel brevius 


. T. baicalense. 
. b. Stigma recurvatum, circinatum etiam, millimetrale lineare. 
ia 9, T. baicalense, ~ megalostigma. 
/ 4 a. Folia caulinaria 0-2, stipulis pesim ee matura pariete 
: = 


ranaceo, nervis saep’ a 
c. Stigma lineare roiltenensaies Plant ae 15-40 em.. 10. 7. philippinense. 
a c. Stigma ovatum vel discoideum vel globosum semimilli- 


. Foliola tae, basi in e, apice acutae 
olae saepius wi tegra p oe cole " 

e. Foliolae —— orbiculares, - marginem totam 
tae 24. T. coreanum. 


bt a Re era aes © gh eee ao Ole 8 be eee, ee ee 


362 Rhodora [OcroBeR 


f. Foliolae oblongo-lineares integrae, nisi we a 
SUA ine pal amtnieai pine cree ace. pcetr cates ead T. integrilobum. 
f. Foliolae dentatae. . 
. Folia caulinaria Pils opposita orbicularia simplicia. 
Radix stolonibus longis................ 11. T. filamentosum. 
g. Folia siege vel desunt, vel alternantia, vel 1-2 
ternata sunt... .h. 
h, Foliolae eteshas Semele atl 
7. Carpella ventre lanceolato. ... 
§: oo faleata. Foliclae iedatne sernet 
hy pe FO EA cone aca) ae ee . TT. declinatum. 
4 Cucpaln parum asymetrica. Foliolae basi 
RS aE Se SY ghee mere te 14. T. acutifoliwm. 
i. Carpella ventre obovoideo vel rhomboideo 
16. 7. unguiculatum. 
h. Plantae omnino glabrae... .k. 
k. Carpella matura vette faleato oblanceolato 
5 mm. longo, nervo ventrali concavo. .12. T'. clavatum. 
k. os ae nervo ventrali convexo vel 
ere ea 
l. Carpella matura jonah Seg 7 a m 
a; inflorescentia sub- 
corymbosa he ‘longats, foliolis fere 


ovatis 2-8 ¢ ongis..... 
arpella faatin ventre 3 
ol sta 9 mm.; antherae 
ROR 55s ca hes eer 4. T. acutifolium 


n, oe matura Mii 4.0-5.38 mm. 
ongo; — . 3 mm.; antherae 


reer a te ke . Chiaonis. 
Planta 1-2 din. alta; eo ed) foliolae 
ina 4-5 mm..... 
0. Polioiae ee orbiculares. Banks ca. 
Stigma discoideum i 
Carpella matura stipite ca. 3.5 ing - 
17. T. Guegueni. 


o. Foliolae rhomboideae, Stamina ca. 4 
mm. Stigma elongatum insertum a 
styli ventrum. Carpella ee 
Be ae 3S mith ss 20. . Watanabei. 


gma sessile hee ate ere sy 5. 7. microgynum. 
Dp. Canaie, nervo dorsali multo eeubec tad 
ventrali. tigma Psi sa 
insertum ad —) val styli brevis, 
Stamm entri a oo: oe 

q- ina mm.; carpella matura 1.0— 
La ee ee 13° T- mirabile. 

q. o semimetralis; stamina 5-6 mm.; 

matura ventre 3-4 

longo, 1 A-1.8 mm. lato....... 19. 7. tuberiferum. 


T. BAICALENSE Turez. ex Ledeb. Fl. Ross. 1: 7, 1841. T. 

bizalens too Paes Soc. Imp. Figs Mose. 2: 85, 1838, ie gaa 
MUR: Korshinsky, Amur mediu us, 1891 

( US): J Rorhaae Peeuaiiaa, 18 gate 1891 g. (G); Ririidasky, 


1944] Boivin,—American Thalictra and their Allies 363 


Khabarovka (na Amurye), 22 Maya, 1891 g. (G); Maak, ad 
fluvium Amur (G). Nippon, Honpo: Masamune, Nikko, July 3, 
1926 (NY). Manpssurra: Schrenk 173, ad fluv. Amur, 1855 
(G). Cura, KANsu: Ching 1003, vicinity of Choni, alt. 3100— 
3300 m., Sept. 7-15, 1923 (US); Ching 360, Hsia Mo K’ou, near 
Lichen, alt. 2000 to 2300 m., July 7-8, 1923 (US); Rock 12,862, 
T’ao River basin, banks of T’ao river, opposite Choni, flowers 
greenish, alt. 8200 ft., July, 1925 (G). Fie. 8. 

9. T. BAICALENSE Turez. var. megalostigma, var. nov.— 
Stigma lineare recurvatum, circinatum etiam, millimetrale.— 
CHINA, szECHWAN: Fang 3619, Kangtin Hsien, Tachienlu, in 
thickets, herb 3 ft., alt. 9000-9500 ft., Sept. 27, 1928 (G, TYPE; 
US, isotype); Wang 21,377, ascending to Fu-pien Hsien, road- 
side, thicket-floor, alt. 3400 m., June 19, 1930 (G), herb 3 ft. 
F 


1G. 9. 

10. T. paiuiprrnense C. B. Robinson, Bull. Torr. Bot. Club, 
35: 65, 1908.—Puriprines, Luzon: R. S. Williams 957, prov. 
Benguet, Baguio, June 22, 1904 (NY, paratypes); idem 1137, 
June 22, 1904 (NY, vypex; G, 1soryrr); Merrill 7767, Benguet 
subprovince, May, 1911 (US). Fie. 10. 

11. T. FILAMENTOSUM Maximovicz, Prim. Fl. Amur. 13, 1859. 
—SrBeriA, PRIMoRSK: Palczewsky, Vladivostok, Mt. Orlinoe 
Gnezdo (Eagle Nest), May 5, 1905 v. (NY); Saverkina 99, June 
(or July) 15, 1929 v. (NY); Topping 2060, Vladivostok and 
vicinity, May—Oct., 1919 (US). amur: Maximovicz (G, iso- 
type?); Komarov 725, vallis fluminis Sutar, 12 junii, 1895 v. 
(NY, US); Korshinsky, 1891 v. (US); Korshinsky, Bidzhanskii, 
8 June, 1891 (G). Fia. 11. 

12. T. cuavatum DC. Syst. 1: 171, 1817.—Physocarpum 
clavatum (DC.) Bercht. & Presl, O Priroz. Rostl. 1: 14, 1823. 
T. filipes Torr. & Gray, Fl. N. Amer. 1: 38, 1838. T. nudicaule 
Schweinitz ex Torr. & Gray, Fl. N. Amer. 1: 39, 1838. Sumnera 
clavata (DC.) Nieuwl. Amer. Midl. Nat. 3: 254, 1914.—Planta 
omnino glabra. Caulis luteolus glaber 1.5-7.0 dm. altus, per- 
saepius solitarius et semimetralis. Radices paucae fibrosae 
nigricantes, aliquantulum ad originem tuberosae. Folia cau- 
linaria biternata petiolata, saepius tamen desunt. Foliolae 1.0- 
5.0 em. long., 0.8-6.0 cm. lat., forma variabiles, subalbicantes 
inferne. Inflorescentia subcorymbosa. Sepala spathulata alba, 
2 . longa. Stamina 4-5 mm., Ca. 0. Filamen 
3.5-4.5 mm., saepe crassiora quam antherae. Antherae ovoid- 


pauciora per receptaculum, faleato-oblanceolata, 7-8 mm. long., 
stipite 2.2-3.0 mm., ventro 4-5 mm. long., nervo ventrali concavo. 
Stigma ca. 0.3 mm., di : 
ejusdem longitudinis. Floret ab Aprili ad Junium.— VIRGINIA: 


364 Rhodora [OctroBER 


Schweinitz, Patrick Co. (ANS, type of 7. nudicaule; NY, frag- 
ment). Nort Carouina: Radford & Stewart 1787, Wilkes Co., 
growing at spring in a shady bend of the road up Pores Knob Mt., 
north exposure, July 2, 1940 (NC); Magee, Lickstone Mt., June 
10, 1897 (G); Curtiss 28, Roan Mt., moist rocky ravines, alt. 
6100 ft., July (G, NY); Churchill, Polk Co., Melrose, May 19, 
1899 (M); Churchill, Watauga Co., Blowing Rock, Glen-Bernie, 
water-fall, June 16, 1899 (M); Biltmore Herbarium 1015b, Bilt- 
more, wet places, May 6th and June 18th, 1897 (G, M, NC); 
M. A. Curtis, mountains of North Carolina (NY, type of T. 

lipes). SoutnH Carona: Peattie, Greenville Co., Hogback 
Mts. (G); J. D. Smith, Caesar’s Head, wet rocks, Aug. 1, 1881 
(G). Grorera: Underwood, Taccoa Falls, Apr. 20, 1891 (NY); 
Leeds 2011, Union Co., on branch of Helton Creek, 1 mile n. of 
Frogtown, dripping rocks, alt. 3850 ft., June 1, 1934 (ANS). 
TENNESSEE: Ruth, Thunder-Head Mt., border of streams, elev. 
6663 ft., July, 1894 (M); Jennison & Sharp, Fentress Co., James- 
town, Buffalo Cave, wet sand, rock, cliffs, May 17, 1931 (US); 
Svenson 8901, Marion Co., Cumberland Plateau, about 5 miles 
se. of Sewanee, dripping sandstone rocks, alt. 1800 ft., June 22, 
1938 (G); Svenson 10,158, Grundy Co., east of Tracy City, Deer 
Lick, in spray of waterfall, May 15, 1939 (ANS, G); Ruth 1799, 
near Wolf Creek, along a mountain brook, June, 1896 (NC, NY); 
Eggert, Franklin Co., no. of Sherwood, shady rocks, June 8, 1897 
(M). Fries. 12, a-e. 


The type is in the Paris Museum. It has not been studied, 
but the description is clear and Delessert’s plate (Icones Sel., 1: 
tab. 6, 1820), presumably drawn from the type, represents this 
species beyond any doubt. 


13. T. declinatum, sp. nov. Planta semimetralis, in petiolis 
petiolulis foliolisque infernis pubescens. Radices fibrosae nigri- 
cantes, paululum ad originem incrassatae. Folia inferiora 
biternata, caulinaria desunt, inflorescentiae ternata. Foliolae 
ovatae asymetricae dentatae, basi cordatae, apice acutae, ca. 
em. long. et ca. 3 em. lat. Stamina ca. 3.5 mm. Filamenta ca. 
3 mm., latitudinis antherarum. Antherae oblongae, 0.5-0.7 mm. 
Carpella matura falcata, ca. 7 mm. longa, stipite ca. 3 mm., 
ventre lanceolato ca. 3.5 mm. longo et 1.0 mm. lato, nervo 
ventrali fere recto vel parum concavo. Stigma ca. 0.3 mm., 
modo Thalictri clavati; stylus idem. Floret Junio et Julio— 
CHINA, KWEICHOW: T’siang 5662 pro parte, Tze-min-an. Tuyun, 
in dense shade, flower white, alt. 880 m., July 7, 1930 (NY, 
TYPE). Fia. 13, a and b. 

14. T. acutifolium (Handel-Mazzetti), stat. nov.—T. clava- 
tum DC. var. acutifolium Hanp.-Mzt. Anz. Wiss. Wien, Math.- 


1944] Boivin,—American Thalictra and their Allies 365 


nat. KI. 63: 8, 1926.—Plania 25-65 cm., glabra vel raro foliolis 
infernis pubescentibus. Radices nigricantes plus minusve tuber- 
osae. Folia basilaria 2-3-ternata, caulinaria saepius desunt, 
inflorescentiae 1—2-ternata. Foliolae dentatae saepius et ovatae 
et basi rotundae et apice acutae, ca. 5 cm. longae. Sepala 
obovata vel spathulata, ca. 2.0 mm. longa. Stamina 7-9 mm 
Filamenta multo latiora quam antheris. Antherae oblongo- 
lanceolatae ad oblongo-lineares 0.8-1.3 mm. Carpella matura 
ca. 5.5 mm. longa, stipite 2.0-2.2 mm., ventre lanceolato 3.0- 
3.2 mm. longo et 0.6—0.8 mm. lato, nervis lateralibus 3 in utroque 
latere, nervo basin recto vel parum convexo. Floret Martio 
ad Junium.—CHin ee Dunn 2316, expedition to central 
Fokien, April to ees 1905 (G); Dunn 2456A, expedition to 
central Fokien, April ‘to June, 1905 (G); Chung 7761 (NY). 
HUNAN: Handel-Mazzeiti 11,178, prope urbem Wukang, in monte 
Yiin-schan, a eon schisto argilloso, fl. pall.-violac., alt. 600— 
1300 m., June 7, 1918 la syntype); Wang-Te-Hui, March, 1919 
(US). KWANGTUNG: Ko 652,821, Yu-yen, in field, June 8, 1933 
de Tso 20,585 and 20, 651, Lok Chong, May, 1929 (NY). 
Fig. 14, a-c. 

Thalictrum clavatum DC. evidently belongs to the same section 
as all these eastern Asiatic species but it is not especially closely 
related to any of them. As far as it is possible to judge, with 
only the original description at hand, the fruits of T. tenerum 
Huth might resemble those of 7’. clavatum more than those of 
any other species except T. mirabile Small. 


15. T. microgynum Lecoyer in Hook. Ic. Pl. 18: 1766, 1888.— 
Cuina, HUPEH: Wilson 12, western Hupeh, April, 1900 (NY, 
US); Henry 3992, tp near Ichang, 1885-1888 (G, 1soTYPE). 
SZECHWAN: Faber 135 (NY); Farges 496, district de T chen-Kéou, 
alt. 1400 m., juin (NY). xwaneTuna: 7'so 20,651, Lokehong 
district, May 24, 1929 (NY). Fre. 15. 


The plate acccompanying the original description is a very 
au representation of the fruiting stage of this species. 

T. unguiculatum, sp. nov. Planta semimetralis. Folia 
beaten biternata. Folia caulinaria desunt. Folia inflores- 
centiae ternata. Petioli et petioluli pubescentes. Foliolae cor- 
datae acutae dentatae, nec lobatae, inferne pubescentes. In- 
florescentia paniculata. Sepala viridia, 3.0-4.5 mm. longa, 
conspicue trinervata, basi appt se lamina 
orbicularia. it ape matura viri mm. lo emery 


longo et 0.8-1.0 mm. lato, nervis obscuris, stylo ca. 4 mm., 
stigmate ca. 0.2 mm. ad ventrum styli inserto. Floret vero- 


366 Rhodora {OcroBER 


similiter Junio.—Cuina, KWEIcHOW: T'stang 5662 pro_parte, 
Tze-min-an, Tuyun, in dense shade, alt. 880 m., July 7, 1930 
(G, Type). Fig. 16, a and b. 
th ies Gueguenii, sp. nov. Planta gracillima, viridula, 
omnino glabra, 1.0-2.5 dm. Caulis tenuis. Folia basilaria 
biternata, inflorescentiae ternata. Caulinaria desunt. Foltolae 
fere orbiculares crenatae 1-2 cm. Flores pauci, saepius 3-5. 
Sepala alba spathulata ca. 2.5 mm. longa. Stamina ca. 5 mm. 
Filamenta insensibiliter clavata, fere latitudinis pees sans ag 
Antherae ellipticae, 0.4-0.8 mm arpella matura ayane nee ca. 
mm. longa, se ot ca. 3.5 mm., ventre lanceolato ca. 3.2 mm. 
longo, 0.6-0.8 mm. lato, stigmate sessili et discoideo. Floret 
Aprili ad Junium _—CHINA, KIANGSI: Gressit 1455, Hing San, 
ie border of Fukien, grassy rocky yt alt. 1300 m., June 23, 
6 (G). HuNAN: Tsang 23,486, Yi Chang district, Ping T’ou 
Shan, Pai Mu village, silt, swamp, ‘fair ly common, Apr. 1-11, 
1934 (G, Type; US, ISOTYPE); Tsang 23,392, Yi Chang district, 
P’ing T’ou Shan, Pai Mu village, steep slope, sandy soil, thicket, 
fairly common, flowers white and fragrant, March 21-29, 1934 
(G, US). Fie. 17, a-c 


Dedicated to EMILE Glee GufGuEN, a former teacher of 
mine, of Belgian origin, a self-effacing, conscientious hard worker. 
To him I owe a good deal of what training and information were 
inculeated into me while going through college; to him I also owe 
a deep and permanent interest in science. 


18. T. mrraBILE Small, Bull. Torr. Bot. Club, 27: 277, 1900. 
Planta glabra gracilis 15-45 em. Radices nigricantes ad originem 
tuberosae. Caulis basus saepius valde incurvatus, quia planta 
illa sponte crescit in locis praecipitosis et lateraliter inseritur in 
fissuris murorum saxosorum. Sepala alba obovata vel spathu- 
lata, 1.5-2.5 mm. longa. Stamina 2-4 mm. longa. Filamenta 
crassiora quam antherae. Antherae globosae vel ovoideae, 

mm. aria saepius 3. Stigma discoideum vel ellipti- 
cum, 0. 1-0.4 mm. longum, insertum ad ventrum styli. Carpella 
matura 5.5-7.5 mm. re stipite 2.5-3.0 mm., ventre obovato 


vel semiobovato 2.5-4.0 mm. longo et 1.0-1.2 ‘am. lato to, nervis 
lateralibus 3 in sete  etwe. nervo ventrali fere recto vel 
paululum convexo, isaac multo convexiore. Floret Maio, 


(ANS); Smith, H Jae Town : Me n Co., Cowbell 
Hollow, east of Berea, July 7, 1937 (F, G , NY, US); Smith & 
H odgdon 3928, Wayne Co., Beaver Cr eek, ‘southwest of Monti- 


cello, shady dripping. ledge, July 12-14, "1937 (ANS, G, US). 
Auapama: Mohr, Winston Co., dripping rocks of mts., May 1, 


1944] Boivin,—American Thalictra and their Allies 367 


1882 (US); Mohr, Winston Co., ad saxas madidas, in sylvis 
rea ome radicibus tuberosis, May 4, 1881 (F); M. ohr, Winston 
, Davidsons Creek, dripping rocks in the mountains, May 1, 
1881 (US); R. M. Harper 3400, Franklin Co., Di smal Branch, 
under sandstone cliffs on damp sandy bank, June 17, 1935 (A NS, 
G, N-ND, NY, US); Earle 2212, Little Mountain, near Moulton, 
under sandstone bluff, June 25, 1899 (NY, Type and 1soryPeE). 
Ig. 18, a-e. 
lt is surprising that this plant has not been collected at inter- 
vening localities in Tennessee. Fruits and stamens of the Ken- 
tucky specimens tend to be larger, but all characters overlap to 
some extent. 


19. T. TuBERIFERUM Maximovicz, Bull. Acad. Imp. Se. St. 
Pét. 22: 227, 1877 [1876].—Nipron, HONDO: T'schonoski, prov. 
Senano, 1864 (G, syntype); Arimoto, Mt. Iwate, July 15, 1903 
(G); Shimotsuke, Nikko, Aug. 8, 1910 (US); Enander, prov. 
Shimatsuke, in radicibus montis Nantai-San in ripa lacus 
Chutzanji, 14 km. procul a Nikko, Aug. 14, 1913 (G); [Watanabe?] 
Shinshu, Togakushi, June 10, 1894 (G, US); [Wat tanabe?}, 
Shinano, Togakushi-san, Aug. 18, 1893 (G): idem, Aug. 20, 
1893 (US); Hogg (NY). Corea: Komarov, distr. Sam-su, 
fluvium Jalu, trajectus Shanjen, julio 7, 1897 v. (G). Manp- 
CHURIA: -Komaron, prov. Kirinensis, dist. Omoso, trajectus 
Sey irine julio 24, 1896 v. (NY); Mazximowiez, — estuarium 
ae oe 1860 (G, syntype); Jankowski, ad. fl. Sedemi (NY). 


ge T. WaTANABEI Yatabe, Bot. Mag. Tokyo, 6: 307, 1892. 
ae HONDO: [Watanabe]?, Tosa prov., Nanokawa, July 
1892 (US, ISOTYPE?) ; Mi garnet Tosa prov. Nanokawa, 

i 11, 1893 (G). Fie. 20. 


I have at hand a fairly large number of Japanese plants, the 
labels of which are all in the same fashion, style and handwriting, 
but there is no collector’s name on any of them. However, one 
of them, 7. acteaefolium Sieb. & Zucc. from Nanokawa, Tosa, 
has a second label which reads: “Plants of Japan, Coll. K. 
Watanabe, Purchased 1892”. I presume all these specimens 
were collected by K. Watanabe, hence I have cited them all as 
[Watanabe?]. If this assumption is right, there is an IsoTYPE of 
T. Watanabei Yatabe at the Smithsonian Institution and perhaps 
also at the Gray Herbarium, for the labels of those two specimens 
would read exactly alike were it not for the year, This simi- 
larity, enhanced by the fact that the two sheets resemble each 


368 Rhodora |OcroBER 


other as plants from the same collection usually do, and the fact 
that the original description and its accompanying plate could 
have been drawn from these specimens without being in any 
way different,—all this strongly suggests that both specimens 
are isotypes. 

T. Hayatanum Koidz. seems to be closely related to T. Wata- 
nabei Yatabe, but I have not seen any material of the former. 


21. T. Chiaonis, sp. nov. Planta glabra 20-40 em. Radices 
nigricantes variis modis tuberosae. Foliolae saepius ellipticae, 
crenatae vel dentatae. Inflorescentia corymbosa vel, si elongata, 
flores in ramis singulis corymbosae. Sepala obovata ca. 2.5 mm. 
longa. Stamina 3.0-3.5mm. Filamenta latitudinis antherarum 
vel latioria. Antherae oblongae vel oblongo-lanceolatae, 0.4— 

mm. Stigma 0.3-0.6 mm., discoideum vel ellipticum, 
insertum ad ventrum styli brevissimi, vel ovarii apicalis. Car- 
pella matura 6-8 mm. longa, stipite 1-3 mm., ventre lanceolato 
4 mm. long. et 0.8-1.5 mm. lat., nervis lateralibus 1-3 in 
utroque latere, nervo ventrali plus minusve convexo, dorsali 
convexiore. Floret Junio Julioque—Corera: Uno 23,694, Kan- 
rasan Saisyuto, Zenranando, alt. 1700 m., Aug. 13, 1938 (G). 
CHINA, ANHWEI: Ching 8593, Wang Shan, July 15, 1925 (US). 
KIANGSI: Chiao 18,719, Kuling, along rocky slope, alt. 3400 ft., 
July 27, 1928 (NY, rypx; US, isotype); Steward 2495, Lu Shan, 
rocky ridge, July 17, 1922 (NY, US). Fic. 21, a and b. 

22. T. iInrEGRILOBUM Maximovicz, Bull. Acad. Imp. Se. St. 
Petersb. 32: 477, 1888, reeditum Mél. Biol. 12: 714, 1888.— 

IPPON, YEZO: Miyabe, prov. Hidaka, Samani mountain path, 
June 19, 1884 (G). 

23. 


The status of this species is open to question and the material 
at hand is not satisfactory. 


Sectio OmMaLopnysa Turcz. ex Fisch., Mey. & Lall., Index 
Sem. Hort. Petrop. 1: 40, 1835—Plantae pubescentes pilis 
brevibus capitatis. Flores perfecti, filamentis staminum parum 
apice dilatatis albis. Carpella stipitata valde compressa, cum 
matura, nervis rugosis attamen nunquam elevatis in costis. 


1944] Boivin,—American Thalictra and their Allies 369 


Rostrum deflexum ad tergum. Species typica: T. sparsiflorum 
Turez., ut patet e descriptione originali. 

It seems likely that 7. Przewalskii Max. belongs here although 
we do not have any herbarium material which would permit 
checking this particular point. 

25. T. SPARSIFLORUM Turez.—Planta elegans plus minusve 
pubescens, 30-180 (saepius 75) cm. alta. Pubescentia translucida 
et minutissima e pilis capitatis. Radices fibrosae. Folia 3-4- 
inert basilaria aetate florendi saepius desunt, caulinaria saepe 

5 in plantis singulis, plus minusve petiolata, superioribus exceptis, 
ista vero cum foliis etiam inflorescentiae omnibus semper sessilia 
sunt. Stipulae auriculatae erosae. Stipellulae desunt. Foliolae 
inferne paululum pallidiores, superne glabrae, circuitu variabili, 
saepius tamen basi vel rotundae vel cordatae, apice vel crenatae 
vel trilobatae. Inflorescentia foliosissima. Flores albi. Sepala 
elliptica 3-4 mm. longa. Filamenta apice paullum incrassata 
saepius dimidium latitudinis antherarum. Antherae ovatae vel 
oblongae parum si vero mucronatae. Ovaria laminaria stipitata. 
Stigma 0.5-1.0 mm., una cum stylo 1.0-1.5 mm. Carpella 
matura valde compressa semi-obovata, nervis lateralibus non- 
nunquam ramosis nec tamen sinuosis. Rostrum deflexum ad 
tergum et paululum incurvatum. Floret junio julioque ali- 
quandiu augusto etiam. 

The type is at Leningrad and was collected in Dahuria by 
Turezaninow. Not seen. The four following varieties seem to 
be worth recognition, although their characters overlap to some 
extent and notwithstanding an occasional intermediate specimen. 
The latter are never hard to classify, for each variety has a 
clear-cut geographic distribution and no two varieties occur in 
the same region. 

25. T. SPARSIFLORUM malig var. typicum. T. Aaa. on 
Turezaninow ex Fisch., Mey. & Lall. Index Sem. Hort. Petrop 
1: 40, 1825, . 7. sec eee ex Lec. Bull. Soc. Bot. Bele. 
24: 317, 1885, nomen nudum pro synon., nec T. contortum L. 
Sp. Pl. 1: 547, 1753.—Pedunculi omnes ex axillis foliorum 1-3- 
ternatorum. Filamenta 4.5-6.0 mm. Antherae 0.8-1.0 mm 
Carpella matura glabra ca. 8 mm. longa stipite ca. 1.5 mm., 
ventre ca. 4.5 mm. longo et ca. 2.0 mm. lato, apice acuto, nervo 
dorsali concavo. = 6 eatin, YENISSEISK: Schipezinsky 75, distr. 
Krasnojarsk, ad fl. Kas., 18 junii 1927 (NY). AMUR: Kom arov 
724, vallis anes prope stationem Liubavisky, 14 junii 1895 v. 

). Fie. 25, a and b. 


26. T. spARSIFLORUM Turcz., var. Richardsonii (Gray), stat. 
nov. T. clavatum sensu Hooker, Fl. Bor.-Amer. 1: 2, 1829, nec 


370 Rhodora [OcroBER 


sensu DC. Syst. 1: 171. 1817. TJ. Richardsonii Gray, Amer. 
Journ. Sci. 42: 17, 1842.—Pedunculi omnes ex axillis foliorum 
1—3-ternatorum. Filamenta 3.0-4.5 mm. Antherae 0.5-1.0 mm. 
Carpella matura glabra 7.5-10.0 mm. longa, stipite 1.2-3.0 mm., 
ventre 4.5-6.0 mm. longo et (2.3)-2.5-(3.0) mm. lato, apice 
truncato, nervo dorsali recto vel parum convexo. —“CANADA”: 
Franklin Expedition (G, paratype of T. Richardsoniz). ALBERTA: 
Hitchcock 12,070, Athabasca Landing, July 28, 1914 (US); 
Brinkman 4169, Slave Lake Distr., ead River (Salteau), 
wettish spots in forest, June 6, 1929 (NY); Richardson, Slave 
Lake, Portage la Loche (G, fragment of rypx of T. Richardsonit); 
Raup, eastern edge of Caribou Mountain apes about 114° 9’ 
W., 58° 51’ N., muskeg along creek, July 12, 1930 (G). BritisH 
Cotumsia: Kay, Lake Ailoknajik, 1883 '(G); McCabe 7678, 
Omineca River, Germansen Landing, i in boggy ground i in alder, 
cottonwood and small spruce woods, flowers dull white, not red, 
July 4, 1940 (UC); Anderson 7566A, Hazelton, open woods, 
June 19, 1925 (N-ND). YUKON: Eastwood 224, Dawson, 
Bonanza. Creek, June 13, 1914 (G); Macoun 58,344, mouth of 
Bonanza Creek, Aug. 11, 1902 (F, NY); Williams, mouth of 
Klondike, July 9, 1899 (NY): Tarleton 64, Five Finger Rapids, 
July 5, 1899 (NY). Auasxa: Coville & Kearney 2345, aa ie 
aay 20, 1899 (G); yevie 128, Kodiak, 1910 (NY); Z. & 
H. B. Loo k, Olga Bay, wet places i in valleys, aly 5, 
1938 ‘(N- ND, NY); e H. Loof 42, Kodiak, Alitak, June—July, 
1937 (G); Eyerdam 19, Kodiak, Sitkalidak Isl., Port Hobron, 
July 27, 1931 (NY); Harrington ch Nagai I., San born .. 1901" 
1872 (ANS, G, NY); Unalaska (NY); Kellogg 211, Unalaska, 
subalpine, 1867 (ANS, G); McKay, Nushagak, island ‘above tide- 
water, June 25, 1881 (ANS): Horne, Karluk, sparse but common 
in rich lower banks, July, 1901 (NY); Scamman 210, Fairbanks, 
Aug. 10-20, 1936 (G); Chapman 42, Anvik a end of island 
in Anvik River opposite mission, Aug. 15, "190 6 (G); Anderson 
6654, Hope, wet woods, June 8, 1941 (G); Anderson 2 A 293, 
Boat Harbor, Lynn Canal, in grass near beach, Aug. 11, 1925 
(G); Anderson 91 6, Matanuska, July 8, 1931 (G); Anderson 7079, 
Manly Hot Springs, July 5, 194 1 (G); Mexia 2185, McKinley 
National Park, hills west of Wonder Lake, in damp ‘hollow, alt. 
800 m., Aug. 4, 1928 (ANS, G, NY); Scamman 1609, Curry, June 
7 1940 (G); Martel 76, Fox Bay, stream-bank, Aug. 2, 1935 (G). 

BERIA, PRIMORSK: Eyer dam, Kamtchatka australis, Savoiko, 
29 julii, 1928 (G); idem, 31 julii 1928 (F); Tiling, Ajan (G). 

T. Richardsonii Gray: fragment of type and a paratype at the 
Gray Herbarium. Hooker and Gray at first called this plant 7. 
clavatum DC. Having seen true 7. clavatum DC. during a trip 
in the Southern Appalachian upland, Gray proposed the name 


1944} Boivin,—American Thalictra and their Allies 371 


Deraits oF FLOWERS oR Fruits oF THALICTRUM, all X 4. 
(For explanation see end of paper.) 


372 Rhodora [OcroBER 


T. Richardsonii for T. clavatum sensu Hooker. But later on, 
probably after receiving a specimen of the same species from 
Ajan, Gray, in Pl. Wright. 2: 8 (270), 1853, dropped T. Richard- 
sonit in favor of T'. sparsiflorum Turcz. Fia. 26. 

27. T. SPARSIFLORUM Turcz., var. saximontanum, var. nov. 
T. utahense Greene ex Rydb., Fl. Rock. Mts., 289, 1918, nomen 
nudum ex synonymis.—Nonnunquam pauci pedunculorum ex 
axillis bractearum, ceteri ex axillis foliorum 1-3-ternatorum. 
Filamenta 3.5-4.5 mm. Antherae0.5-1.0mm. Carpella matura 
pubescentia 6.0-8.5 mm. longa, stipite (0.3)—-0.5-(1.0) mm., 
ventre 4.0-5.5 mm. longo et (2.2)—2.5-(3.2) mm. lato, summo 
truncato vel obtuso, nervo dorsali fere rele nervis lateralibus 
nonnunquam undulatis ic ramosis.—Montana: Rose 61, Carbon 
Co., near Red Lodge (CA). Ipano: Rogleston 138, 980, Custer 
Co., Challis Forest, Sey as Creek, Taylor’s ranch, ca. 1940 m. ‘ 
July 29, 30, 1917 (G); E & L. B. Payson 2109, Teton Co., 
hills southeast of Victor, en creek-banks, alt. 6200 ft., July 
24, 1920 (CA, G); Macbride 593, Elmore Co., Trinity, creek- 
banks, alt. 4500 ft., Aug. 12, 1910 (G); ;M acbride & Payson wa 
Blaine Co., Camas Prairie, in shade of creek-bottom, alt. 5700 
ft., June 29, 1916 (G); FE. B. & L. B. Payson 2018, Fremont Co., 
Henry Lake, creek-banks, alt. 6000 ft., July 14, 1920 (CA, Gy; 
Macbride & Payson 3283, Custer Co., ’ Bear Creek, wet creek- 
banks, alt. 6000 ft., July 18, 1916 (CA, G); Henderson 3372, 
Blaine Co., near Ketchum, July 23, 1895 (CA). Wromina: L. 
Williams 1671, Grand Teton National Park, Cascade Creek, 
moist bottoms, "alt. 7500 ft. , July 7, 1934 (CA): L. Williams 855, 
Grand Teton National Park, in timber, moist places, alt. 7000 
An Sees 14, 1932 (CA, G); ‘A. & E. Nelson 6364, eer 

, Lewis River, on the banks of the riv er, Aug. 8 9 (G, 
fs ‘of var. saximontanum = b:O. ER R. Williams 3186, Big Horn 
Co., Shell Creek, near the pe th of Ant telope Creek, sandy 
rocky creek-bottom, alt. 8500 ft., June 30, 1936 (G); A. Nelson 
7970, Albany Co., Little “pathhem Middle Fork, in wet willow 
thickets, very common, Aug. 6, 1900 (G); Goodding 408, ee 

oO 30 


(G); Merrill & Wilcox 913, Lincoln Co., Leighs Lake, Jacksons 
Hole, damp soil, July 24, i901 (G); A. Nelson 1548, Cummins, 
alt. 8000 ft., July 30, 1895 (G); A. Nelson 1494, Cummins, July 
29, 1895 (G). Cotorapo: Parry 76, from the headwaters of 
Clear Creek and the alpine ridges lying east of Middle Park, 
1861 (G); Parry 9, Rocky Mountain Flora, 39°-41°, 1862 (US); 
Parry 76, 39°-41°, alpine and subalpine, 1862 (G); Hall & 

Harbour 9, Becky Mts., ne 39°-41°, 1868 (G); Vasey 7, 9, 
Rocky Mts., Grand Lake , 40°-41°, 1868 (G, US); Zobel, Middle 


, 


St. Vrain fives July 12, 1939 (CA); Hapeman, Poudre Canyon, 


PE LO er See) Se SRT Go en le ee ee 
ca . — ci 


| 


PTA SE ORS BTS RE Ls RINT I ACE Cl OLR ans 
iui ead Nie tie 


1944] Boivin,—American Thalictra and their Allies 373 


alt. 7500 ft., Aug. 24, 1986 (CA); Walker 478, San Miguel Co., 
Norwood con Reece north slope by bite alt. 7000 ft., Aug. 17, 
1912 (G); Crandall 33, near Georgetown, along stream, alt. 
847.4 ft. baah: Taly 20, 1892 (G); Chueh Clear Creek Co., 
Brookvale, Yankee Creek, thicket, June 22, 1918 (G); Patterson, 
vicinity of Georgetown, Bard Creek near Empire, June 28—Aug. 
7, 1875 (F). Uran: M. E. Jones 5731a, 5731b, Sevier Co., tog 
Lake, a alt. aan ft. _ Aug. 3, 1894 (US); idem 5789, Aug. 6-7 (N 
US); & L. B. Pa ayson 4970, Summit Co., Uintah Oh = 
Selatan: Fork of Bear River, in deep shade near stream-bank, 
alt. 8800 ft., July 15, 1926 (G); Goodman & Hitchcock 1482, 
Summit Co., Uintah Mts., E. Fork of Bear River, meadow, edge 
of small stream, alt. 8900 ft. , July 9-13, 1930 (G); Graham 10, 087, 
Uintah Co., between Paradise Park and Chepeta Lakes, lodge- 
pole-spruce. woods, along stream, alt. ” nk ft., See 20, 1935 
(G). OREGON: Cusick 3357 a, Wallow p of 
North Catherine Creek, very rare, alt. sect 4500 ft, ‘ne 31, 
1910 (G). Fie, 27. 

SPARSIFLORUM Turez., var. nevadense, var. nov. 
Pedunculi nonnunquam pauci ex axillis bractearum, plurimi vel 
omnes ex axillis foliorum 1—3-ternatorum. Filamenta ca. 4mm 
Antherae 0.7-1.0 mm. Carpella matura pubescentia 7.5-8. 0 
mm., stipite millimetri, ventre 4.5-6.0 mm. longo, 2.2-2.5 mm. 
lato, summo rotundo vel obtuso, nervo dorsali convexo vel 
summo convexo et basi concavo.—NEVADA, ESMERALDA: 
Duran 3354, ee Mountains, Trail Canyon, alt. 9800 ft., July 
20, 1932 (CA, G, UC). Caxirornia: Lemmon, Webber Lake, 
Oct., 1890 (UC); Campbell, Kearsarge Pass, on east side, July 
29, 1916 (CA); Michaele, Yosemite, Pohono ‘trail, July 19, 1922 
(CA); Redfield, prope lacus Tahoe, in paludibus, Sept. 1, 1872 
M). CALIFORNIA, LASSEN: Nutting & Baker, Pine Creek, July 
12, 1894 (UC). sHasta: Eastwood 191 8, Lassen Butte region, 
Battle Creek meadows, Aug. 22-26, 1912 (G, M, UC); Austin, 
Big Spring, 4—6 ft. high, weak, supported by bushes, July 4, 1878 
(G); Brown 6101, near Lassen Buttes, alt. 6000 ft., Aug. "1-15, 
1897 (M); Austin, near Lassen Peak, Warner Valley, 1879 (G). 
BUTTE: Copeland, Jonesville, above Butte Creek House, alt. 
1800 m., Aug. 26, 1930 (UC). sierra: Sonne, Little Truckee 
River, fls. July 10, 1892, fr. Aug. 7, 1892 (UC). NEVADA: 
Heller 7056, on the Truckee River, one mile above Truckee, 
July 30, 1903 (G, type; M, UC, isorypEs). PLACER: Sonne, 
road to Hot Springs, fl. June, 1896, fr. July, 1896 (NY). ELDO- 
RADO: Hillman, Lake Tahoe, Bijou, Sept. 15, 1894 (UC); Crum 
3011, two miles southeast of Echo Lake, Echo Summit, alt. 
7500, ft., subalpine zone, wet soil, shade, July 3, 1939 (UC). 
TUOLUMNE: Mason 680, Sierra Nevada Mts., Lyell Fork, Aug. 1, 
1923 (UC). mariposa: Babcock 281 (1097), Yosemite Nat. 


374 Rhodora [OcroBER 


Park, Illilouette Canon, on first branch of Illilouette Creek, 
above the falls, growing in bogs, abundant here but apparently 
not wore in the middle Sierra Nevada, alt. 1955 m., June 28, 
1912 (CA, G, M, UC). Mono: Wright, Mono Lake, July 1 18, 
1917 (CA). ‘MADERA: Congdon, Upper San Joaquin, Aug. 1 
1895 (UC). Fresno: Hall & eet ye 142, Pine Ridge, alt. 5300 
ft., June 15-25, 1900 (UC). «nyo: Rose 35,462, Ibex Park, E. 
slope, meadows, alt. 10,300 ft., July 24, 1935 (CA); Peirson, 
Rock Creek Lake Basin, margin of lake on east side of canyon 
and at base of Transverse Ridge, alt. 10,700 ft., Aug. 5, 1940 
(UC); Peitrson, Rock Creek Lake Basin, near Ruby Falls, in wet 
locations, alt. 10,900 ft., Aug. 19, 1933 (UC); Alexander & 
Kellogg 2581, Big Pine Creek, trail between 2nd and 3rd lakes, 
in rank vegetation near small creek, July 16, 1941 (UC); Hal- 
perin 579A, Rock Creek, alt. 10, 000 ft. July 31, 1932 (CA); 
Ferris 8866, Loch Leven Fork of Bishop Creek above North 
sore dry slopes in lodgepole pine forest, July 21, 1934 (UC). 
RE: Hopping 352, Kaweah River Basin, Round Meadow, 
Sept. 16, 1905 apt Hopping ae, Kaweah River Basin, Giant 
Forest, July 20, 1904 (UC); J. T. Howell 1 ‘f — Chagoopa 
Plateau, Sky Parlor Meadow, Aug. 1, 1942 (C A): J T. Howell, 
15,700, East Lake, July 30, 1940 (CA); Cronquist 2132-A, 
Sequoia Nat. Park, 3 miles from Giant Forest, Crescent Meadow, 
alt. about 7000 ft., June 27, aoe (M). SAN ay L. 
Wheeler 1186, San Bernardino ., 5. Fork Santa Ana R. sunny 
stream-side, alt. 8400 ft. , Aug. oe 1932 (CA); — 2842, San 
Bernardino Mts., Bear Valley, alt. 6700 ft., Aug. 1 1, 1902 (G, M); 
Abrams & M cGregor 765, San Gorgonio Mt., Dollar Lake ‘Can- 
yon, alt. 9000-9500 ft. , July 12, 1908 (G): 8 . B. & W. F. Parish 
1 eg San B. Mts., Bear Valley, Aug., 1882 @); Munz 12,676, 
B. Mts., South Fork of Santa Ana River, in meadow, a ut 
ls etc., foetid plant, alt. 8300 ft., Aug. 21, 1932 (M, UC); 
Munz 1 0, 617, San B. Mts., Bluff Lake Meadow, shade of willows, 
with foetid odor, alt. 7400 ft., July 5, 1926 (UC); Munz 6267, 
San B. M k. of Santa Ana, common in wet meadows, alt. 
8000 ft. , Aug. 25, 1922 (UC); M. E. Jones, San B. Mts., Bear 
Valley, alt. 6600 ft. , July 19, 1900 (CA, UC); Johnston, San B. 
Mts., Bluff Lake, shaded margin of meadow, alt. 7500 ft., July 
5, 1924 (G); J. T. Howell 2760, San B. Mts., Riverside Municipal 
Camp, Mill Creek Road, moist partially shaded stream-bank, 
7300 ft., July 10, 1927 (Ca); Grant 1287a-6351, San Gorgonio 
Mt., July 24, 1904 (CA, U C); Anderson grade south of Bear 
Lake, a soil, alt. 7000 ft. , July 24, 1935 (C00). RIVERSIDE: 
sua toe 2259, San Jacinto Mt., in sylvis, alt. 6000 ft., julio, 1928 


Se Annie Creek, idee shade, alt. 5000 ft., Jul 26, 1935 
(CA, G, NY). Fic. 28. 2 


a te etal tat a a ts eres ee ee) eS a ee ne Ke 
i mi 3 ant oy mae cy wot 7 i Ging amit on a ail ie 
ea - 7 ~ 


1944} Boivin,—American Thalictra and their Allies 375 


Sectio TripETRIUM DC. Syst. 1: 169, 1817. Tripetriuwm (DC.) 
Bercht. Eg Presl, O Priroz. Rostl. 1: 14, 1823. Subg. Tripetrium 
(DC.) Reich. Cons. Regn. Veg. 192, 1828. Ruprechtia Opiz, 
Sez. Rost. Kve. 86, 1852.—Filamenta staminum dilatata. Car- 
pella matura valde tri-quadrialata, longe stipitata, rostro ad 
ventrum deflexo. saeiite brevius. Species typica Thalictrum 
aquilegifolium L. sit 

All names nipees for this group are clearly based on T. 
aquilegifolium L. De Candolle himself included two species in 
this section, and one of the two may be a synonym of T. aquilegi- 
folium L. This species was also proposed as a standard species 
for the genus Thalictrum, but there was an earlier and better 
proposal: 7’. foetidum L. (See p. 347 for discussion of this topic). 
This section is not represented in America. 

aU. aK phi ok pp L., var. Sa yes Beck, f. Cornuti 
(L.), stat. no T. Cornuti L. , sp. PL, 1: 545, 1753. T. cana- 
dense Miller, Gardener * Diet. ed. 8 Pk ie confertum Moench, 
Hort. Marburgensis, 297, 1794. Leucocoma canadensis (Miller) 
Nieuwland, Amer. Midl. Nat. 3: a Atos ei gatas BS dca 
differt filamentis stamin albis. Typ 
herbarium * ra at the 1 British er aieired a Nenpat History. 
Fic. 29, a a 

There is in hee no species corresponding to Cornut’s 
description, and the plant is not represented in the Linnaean 
Herbarium, Lecoyer, however, saw the specimens under that 
name in the herbaria of the Hortus Cliffortianus, of Vaillant, of 
Tournefort and of the Academy of St. Petersburg, and states 
that they all belong to 7. aquilegifolium L. Thus it seems that 
the plant cultivated by Cornut was later widely distributed in 
European botanical gardens and, although he left no herbarium 
specimens, we still have good material by which to interpret 
what he meant. But this has been strongly contested by many 
authors, 

In 1635, when Cornut published his Thalictrum canadense, 
Canada had been explored from the Gulf of Saint Lawrence to 
the Great Lakes, a region where six species grow: 7. alpinum L., 
T. polygamum Muhl., T. dasycarpum Fisch. & Lall., T. revolutum 
DC., T. confine Fern, and T. dioicum L. 

If one reads Cornut’s description he will see that the height of 
the plant (bicubitalis) eliminated 7. alpinum L. The color of 
the stamens (mille alba filamenta ostendunt) eliminated T. 


376 Rhodora [OcroBER 


alpinum, T. confine and T. dioicum. The number of sepals per 
flower (singuli in quinque folia divisi) eliminated all but T. 
alpinum. The flowering time (julio mense singula filamenta 
dum deficiunt) eliminated T. polygamum, T. revolutum and T. 
dasycarpum. And the form of the fruit (triangularisque formae, 
extante quadam per singulos angulos epiphysi membranea) not 
only eliminated all six species but sufficed alone to characterize 
T. aquilegifolium L. 

If we now turn to Cornut’s plate, the stipellules, which are 
very clearly represented indeed, eliminate all eastern North 
American species. But the habit and the flowers suggest 7’. 
polygamum. The habit and the flowers also suggest 7’. aquilegi- 
folium as well. It is true that the latter has perfect flowers and 
there is not a word about the ovaries in the text nor are they 
represented in the drawing. But anyone who has seen 7’. aqui- 
legifolium L. in bloom has not seen any ovary unless he suspected 
their presence and carefully dissected a flower. For the ovaries 
are few, filiform, very small and hidden in the middle of the 
flower amidst the bases of the stamens. If anyone sees the plant 
in bloom and comes back to it a week or two later to find no 
trace of the stamens but only already well developed fruits, he 
will certainly get the impression which Cornut seems to have 
intended the following words to convey: “julio mense singula 
filamenta dum deficiunt, in totidem semina degenerant’’. 

De Candolle first seems to have realized this error and he 
proposed 7’. corynellum for the American plant hitherto called 
T. Cornuti L. In 1910, Greene! protested against dropping 
Linnaeus’ name to replace it by the very ambiguous one coined 
by Muhlenberg. He suggested that Cornut might have received 
his material through Dr. M. Sarrazin and, by mistake, described 
the flowers of T. polygamum but the fruits of T. aquilegifolium. 
This point of view was endorsed by Farwell? and others. 

As pointed out above, not only the fruits, but also the flowers 
and the foliage of T. canadense Corn. differ from those of T. 
polygamum Muhl. And Dr. Sarrazin’s manuscript, a copy of 
which is preserved at Saint-Hyacinthe near Montréal, reads as 
follows: 


1 Leaflets, 1: 51, 1910. 
2 Papers Mich. Acad. Sci. 26: 10, 1941. 


1944] Boivin,—American Thalictra and their Allies 377 


178 Thalictrum Canadence, caule purpuras 
a diets florum stomiedlons albis I. RIL H. 271. 


Il oro sur ne bords des riviéres, dans les prairies. 


Cornut having written in 1635, this point needs no further dis- 
cussion. Indeed, Dr. Sarrazin lived from 1659 to 1734. 

Greene also suggested that, Cornut’s plant having been 
described from cultivated material, it might be possible to find it 
in European botanical gardens. Well, what Linnaeus, Tourne- 
fort and Vaillant had and called 7. canadense Corn. was T. 
aquilegifolium L., but Bernard de Jussieu had in his herbarium 
both 7. aquilegifolium L. and T. polygamum Mubhl. labelled T. 
canadense Corn. 

T. canadense Miller. The intention of Miller was certainly 
not to describe a new species, for he calls it 7. canadense Cornut. 
Only through the rules of nomenclature is this name attributed 
to Miller. 

Thalictrum confertum Moench. Described from plants culti- 
vated in a botanical garden, no type being known to exist. 
Moench gives 7. Cornuti L. as a synonym and his description 
agrees perfectly well with 7. aquilegzfolium L. in bloom. 

hus it seems probable that, through exchanges between 
botanical gardens, the stock of seeds which furnished T. Cornutt 
L. also furnished 7. canadense Miller and T. confertum Moench. 
At any rate, it seems to be the only sensible explanation. 

In his herbarium, now at the Smithsonian Institution, Mohr 
had a small fragment of 7. aquilegifolium L. purportedly collected 
by Prof. Riddell in 1839 in Ohio and it was identified as Thalic- 
trum dioicum L., var. stipitatum T. & Gr. Undoubtedly it was 
from cultivated plants. 

(To be continued) 


AMERICAN THALICTRA AND THEIR 
OLD WORLD ALLIES 
BERNARD BolvIN 
(Continued from page 377) 

Subgenus Lecoyerium, subg. nov. Inflorescentia paniculata, 
rarissime subcorymbosa. Flores dioici vel polygami vel monoici 
vel in paucis perfecti. Sepala 4—(6), superantur o 
tempore anthesis. Species typica Thalictrum dioicum 

Sectio Cincinneria sect. nov. Sect. Camptogastrum, b. Fost 

carpa Prantl, Nat. Pfl. 3, 2:66, 1888.—Plantae elatae; in speciebus 
nonnullis nitentes in proximis, ‘glabrae vel puberulentes circa nodos 
aetate florendi. Folia percomposita, inflorescentiae numerosa, 
folios subrotundis, apice plus minusve trilobatis vel grosse 
dentatis. Pedunculi longissimi et pendentes aetate fructuum 
maturandorum, brevissimi tamen aetate florendi. Flores oe 
carpellis 1-4, staminibus purpureis antheris mucronatis. 
Viridia (an semper?). Stigmata saepius truncata aetate relies pes 
maturandorum. Fructus plus minusve recurvatus vel fere rectus, 
plus minusve compressus vel, si sectus, rotundus, nervis rugosis, 
T’. impexo nostro excepto, stipitatus, nec costatus. Species typica 
Thalictrum cincinnatum nostru 

Height attributed herewith ae the different species is based 
mainly on measurements of a few nearly complete individuals 
which have been folded until they fitted the Procrustean bed of 
an herbarium sheet. Collector’s observations were available 
only in the case of T. Mannii Hutchinson and on two herbarium 
Sheets, one of 7. innitens nostrum and one of JT. Steinbachii 
nostrum. 

The underground system of all the species of this section is 
unknown. There seems to be no fixed flowering season for the 
African species, at least those from central Africa. 


392 Rhodora {NOVEMBER 


CLAVIS AD FLORENTIA 
a. Antherae il 8-0.9 mm., sepala 1.3-2.0 mm. longa, filamenta 
a b 


b. Stigma A 15-3. Omm..... é. 
Filamenta ca. 1.5 mm., sepala 1.3-1.5 mm. longa....30. T. aduncum. 
é. Filamenta ca. 2.5 mm., sepala 1.5-2.0 mm. longa... .32. 7’. innitens. 
Bt UI oi tos cea wv eae ee ee rhynchocarpum. 
a. Antherae 1.2-3.0 mm., sepala 1.2-6.0 mm. longa... .d. 
d. Carpella singula raro gemina in floribus singulis. . . .e. 
e. Sepala 2.0-3.0 mm. longa... .f. 
Sy Witte. Oh RU oi sas i ee ia 37. T. Steyermarkii. 
f. Filamenta 3.0-4.5 mm..... 
g. Stylus una stigmate ca. { 5 mm., stamina 13-19 in 
POTTIUE DE. 0 es es ee te se = 33. T. Chapini. 
g. Stylus omit stigmate 2.0-4.0 mm., stamina 5-8 in 
SWANS COMMON <5 28 fo se ek bse ES . TT. impexum. 


e. Sepala ca. 5.0 a MN ey ne eine fe cee ee 35. T. Stolzit. 
d. Carpella 2-4 in floribus singulis. .. .h 
h. Sep 3.5 mm. | ae 

z. Stylus una stigmate 4-7 mm.... . Ge 

4. Depere 20-30 mm. LONRA, 6c ok a 33. T. Chapinii. 

] O03. i RS ei es 36. T. Steinbachit. 
7. Stylus una hyp 8-13 mm..... 

na cum stigmate ca. 13 mm....... 37. T. Steyermarkit. 

k. Stylus ae stigmate 8-10 mm........ sb seas 38. TT. cincinnatum. 

h. Sepala ee NR a ee es 39. T'. macrocarpum. 


Criavis AD FRUCTUS FERENTIA 
et es PR is i er ay a 2d a 34. 7. impexum. 
a. Carpella stipitata....b. 
b. Carpella solitaria, rarissime gemina. .. . 
lla compressa, ad tergi medias valde recurvata..30. 7. aduncum. 
c. Venter carpelli —— si vero recurvatum.... 
a. Carpelia 3-4 mm. lata. . i... on. secs ee 37. T. Steyermarkii. 
d. Cavdis Leite lata... .. 
e. Venter carpelli palsesebetiat ad extremas parum 


PROUT VA oS ee ee 31. T. rhynchocarpum. 
e. Ventrum carpelli lanceolatum vel ee 
f. Stylus una stigmate 3.5-4.5 mm.............. T. innitens 
i. ne una stigmate 5.0-6.0 mm............ 33 Chapini 
b. Carpella nunc ier nunc gemina, nunc ternata, in 
eadem planta... . ‘ 
g. Africanum, ca‘ TANOOOINE RG I. eee oe 33. T. Chapinn. 
g. Americana vel pyrenaicum, carpellis semiovatis vel 
ovatis.... 


h. Venter carpelli parum recurvatum 3-5 mm. lon 
36. 7. Steinbachit. 
h. Venter carpelli 5.0-7.5 mm. longum, nervo dorsali con- 
_ exo ve el recto, tamen medias parum concavo. .. .t. 
4. ae debiles ee pendentes, uisi os 


lantae ela nflorescentia copiose ramosa. . . -J 
j. CRU CD Wi ies oe ai an Be xs 37. T. Steyermar 
de RODOD AUP NN ce ec ves s. - 38. T. pret 
i. Pedunculi rigidi imetrales 


ascendentes. Plantae semi 
orescentia simplex vel fere simplex... .39. T. “‘macrocarpum. 


T. Stolzii Ulbrich being known only from flowering material 
has been, consequently, omitted from the second key. 


1944] Boivin,—American Thalictra and their Allies 393 


As the fruit reaches maturity, the stigma is likely to break off 
somewhere along its length. The accompanying illustrations all 
represent such shortened stigmas, but unbroken ones are usually 
to be found on each herbarium sheet. Vegetative characters 
vary but little through the section, exception being made of 7. 
macrocarpum Gren. 


30. T. aduncum, sp. nov. Planta metralis vel paululum 
altior. Folia bg foliali parvis. Pedunculi ca. 10 cm., 
fructubus maturis. Sepala 1.3-1.5 mm. longa. Stamin a 5-6. 


Filamenta ca. 1.5 mm. Miike oblongae, 0.8-0.9 mm. nat ath 
apice obtusae, breviter acuminatae. Ovaria solitaria. Stylus 
una stigmate 1.5-2.0 mm., aetate maturandi 4.5-6.0 mm. 
Fructus 13.5-15.5 mm. longus, compressus, ad medias valde 
recurvatus, stipite 2.5-3.0 mm., ventre ca. 7.0 mm. longo, ca. 
2.5 mm. lato, nervis sivaphediue —Conco: Humbert 7448, 
montagnes & |’ ouest du lac Kivu, forét 4 l’W de ae alt. 
2000-2400 m., février-mars 1929 (G, TYPE). Fig. 3 

31. T. RHyNcHOCARPUM Dill. & Rich., Ann. Sci. Nat. ser. 2, 
14: 262, 1841. T. lonpebanmiatens Hochst. & Steud. ex Steud. 
Nom. Bot. 2: 676, 1841 ut synonymon, nec Age rues ee 
dunculatum Sennen, Bull. Soc. Bot. Fr. 73: 642, tT. 
chymocarpum Dill. ex Walp. Ann. Bot. Syst. 2: 5, 1861. gh 
Mannii Hutchinson ex Hubshinashs & Dalziel, Fl. W. Trop. Afr. 
1: 66, 1927.—Planta sesquimetralis usque bimetralis (an metralis 
usque ad quadrimetralis?). Folia ca. 5-ternata, foliolis 0.3-2.0 

diametro. Sepala 1.4-1.8 mm. longa. Stamina 5-10. 

Antherae ia ca. 0.9 mm. Filamenta ca. 3.0 mm 
Stylus una stigmate 4.0-5.0 mm., aetate maturandi 4.5-5.5 mm. 
Carpella matura 15.5-18.0 mm. longa, parum recurvata in apice 
stipitis 2.5-5.0 mm. et ad apicem ventris oblanceolati nee com- 
pressi 8-10 mm. longi et 1.5-1.8 mm. lati, nervis simplicibus.— 
Nierrtia: according to Hutchinson, in the Pgs Mts. sig 
at Buea and Fernando Po (type locality of T. M ABY 
SINIA: Schimper 472, Berg Semajata, 7300 f. rey 14, ‘1862 (G): 
Schimper 1137, inter Endchedcap et Schoata, 1838-1842 (G); 
ipo 1183, Gerra, 7800 ft. (F). BELGIAN otilty Linder 
: 6, Mt. Ninagongo, 8000-9000 ft., Feb. 16, 1927 (G). Fic. 
4 a— —d. 


The type of this species from Abyssinia, is presumably pre- 
served at Paris and has not been seen, but the description is 
clear enough and, the species being the only one found in the 
mountains of Abyssinia, there is no doubt about its interpreta- 
tion. The illustration in A. Richard Tent. Fl. Abyss., tabula 2, 
confirms this interpretation. On the herbarium sheet of Schim- 


394 Rhodora {NOVEMBER 


per 1137, there is the following manuscript note: “Thalictrum 
rhynchocarpum. Ach. Rich. in Ann. sc. nat., Bot., XIV (1840) 
p. 262 (publ. le 24 Mars 1841)”. 

Thalictrum longepedunculatum Hochst & Steud. was not given 
a new description when published, it was simply a new name for 
T. rhynchocarpum Dill. & Rich. 

T. chymocarpum Dill. is probably an error of spelling or 
printing. 

T. Mannii Hutchinson. Described from flowering specimens 
from Nigeria, a region from which I have not seen any Thalic- 
trum. The characters stressed by the describer seem to fall 
within the range of variation of 7. rhynchocarpum Dill. & Rich. 
The stamens and the fruit, however, are too briefly described to 
enable one to form a definite opinion about the status of this 
species. As described by Hutchinson, it is a plant 1 to 4 meters 
high. As far as records go, this is the tallest plant in the whole 
genus. 

32. T. innitens, sp. nov. Planta ca. 1.5 metrorum. Folia 
usque ad septies ternata. Pedicelli ca. 10 em. aetate fructuum 
maturandorum. Sepala 1.5-2.0 mm. longa. Stamina 5-6. 
Antherae ovoideae breviter mucronatae, ca. 0.8 mm. Filamenta 
ca. 2.5mm. Ovaria solitaria, stigmate 2.0-3.0 mm. usque ad 4.5 
mm. accedente cum carpella maturant. Fructus compressus 
rectus vel parum recurvatus 10-14 mm. longus, stipite 1.5-2.0 
mm., = ovoideo-lanceolato 6.5-7.5 mm. longo, 2.0-2.4 mm. 

lato arcuantibus simplicibus.—Nyassa: Buchanan 64, 

1801 (i US); "Stolz. 148, Station Kyimbila, Bundali Gebirge, 1300- 
eter Seehshe, Nov. 25, 1907 (G, US). RHopeEsiA: 
Swynnerton 352, Chipete forest-patch, a climber common on 
forest outskirts, ‘alt. 3800 ft., 1906 (US). Transvaau: Drakens- 
berg. Pilgrim’s Rest (F 69 381). ORANGE: Cooper 1038, in woods, 

862 (N ter Nata: Wylie (J. M. Wood’s no. 8128), ’Wahawag 
Mtn. alt. 6-7000 ped., March 22, 1901 (US). Caps: Murray 
598, in densis sylvis “Kabousie”, alt. 3500 ft., flor. dec. (G, 
TYPE; F, isoTyPE). Fic. 32, a-d. 

It is doubtful whether this plant really is a climber. The 
herbarium specimens show no evidence to that effect. The stem 
is hollow, thin and rather easily crushed and seems an inappro- 
priate support for a plant of this size. The large and abundantly 
branched inflorescence with its pendulous fruits is likely to 
become more or less entangled in the branches of adjoining 
shrubs, thus giving the plant the additional support needed. 


ra 


1944] Boivin,—American Thalictra and their Allies 395 


This undoubtedly holds true for most of the species of the 
§ CINCINNERIA, especially the tallest ones. 

33. T. Chapinii, sp. nov. Planta circa sesquimetralis. 
Folia yee ae quinquies ternata. Pedicelli 5-10 mm. Sep 
2.0-3 onga. Stamina 13-19. Antherae oblongo-lanceo- 
latae 1. 4-9. i mm. , mucrone 0.2-0.7 mm. Filamenta 3.2-4.5 mm. 
Ovaria nunc singula nune gemina, nunc ternata. Stigma, 
aetate fructus maturandi, 5.0-6.0 mm. Carpella matura parum 
recurvata compressaque, 13.0-14.5 mm. longa, stpite 1.5 mm., 
ventre ca. 6.5 mm. longo, ca. 2.0 mm. lato, nervis arcuantibus 
simplicibusque. —Coneo: Chapin 386, ‘slope of Mt. Karisimbi, 
Kivu district, alt. 11,000 ft., rather common about Kabara, 
about same level, June 19, 1927 (NY, type); J. P. Chapin 518, 
Kivu district, mts. s. w. of Lemera, lower Ruzizi Valley, alt. 
9600 ft., July i7, 1027 ANY}. Kenya: Mearns 1850 & 1414, 
western slopes of Mount Kenia, along the trail from West Kenia 
Forest Station to summit, in the “giant heath” zone, at about 
3630 meters, Sept. 21- 27, 1909 (1350:US; 1414 NY, US); 
Mearns 1670 & 2820, western slopes of Mount Kenia, along the 
trail from West Kenia Forest Station to summit, bamboo ssn 
at about ge meters, Sept. 28—Oct. 7, 1909 (1670: F, US; 2320 
US). Fia. 33, a-d. 

Although flowering material of 7. Chapinii and T.. innitens is 
distinguished easily indeed, these species become more or less 
confluent as they mature their fruits. 


34. T. impexum, sp. nov. Planta verosimiliter circa bi- 
metralis. "Palia gi usque ad sexies. Pedunculi 5-9 cm 
Sepala 2.0-2.4 mm. longa. Stamina 5-8 in floribus singulis. 


Antherae chiniae-Rnaeantel: Ovaria singula rarius gemina, 
— 2.0-4.0 mm., aetate maturandi, 4.0-5.0 mm. longo, 
nd 


"| 
bo 
fund 


909 (595: Pi, 630: US, 630, 652, Type: US 630, 653 
ISOTYPE). Fic. 3 a-d. 

This species is pei for this section because of its nearly 
sessile fruit. When in flower it closely recsembles T. Chapinii 
which occupies about the same range at higher altitudes. 


396 Rhodora [NOVEMBER 


35. T. Strotzim Ulbrich, Notizbl. Berl. Gart. 10: 916, 1930. 
Herba perennis erecta ad 1.25-1.50 m. alta parce ramosa caule 
—§ mm. crasso, omnino glaberrimo fusco-violaceo. Folia 
caulina in vaginam fere 35 mm. longam 15-20 mm. latam dila- 
tato, usque ad laminae ramificationem + 12 cm. longo glaber- 
rimo; lamina 3-4-pinnata foliolis + 3 mm. petiolulatis circuitu 
orbicularibus vel reniformibus margine grosse crenatis crenis 
subapiculatis, 10-12 mm. longis, 10-15 mm. latis, glaberrimis, 
supra atroviridibus, subtus glaucescentibus, nervis reticulatis 
supra inconspicuis, subtus prominentibus. Inflorescentia satis 
parva decomposita contracta paniculata. Flores satis magni 
lilacei pedicello brevissimo, 2-4 mm. longo, glaberrimo instructi 
Leah basi bractea lineari-lanceolata + 4 mm. longa membra- 
, 0.5-1 mm. lata glaberrima lilacea munita; sepala ovalia 
vel "Tncecovedia + 5 mm. longa, + 3 mm. lata, obtusa, glaber- 
rima + quinquenervia. Stamina +8 mm. ‘longa filamentis 
filiformibus tenerrimis glaberrimis superne vix dilatatis sparsis 
(5-10), anthera linear-oblonga 1-1.5 mm. longa. Ovaria 
(plerumque in flore unicum, rarius 2) brevissime stipitata vel 
subsessilia glaberrima anguste lineari-lanceolata cum stylo + 5 
mm. longa in stylum fere 3 mm. longum rectum sensim attenuata; 
stigma apice lateraliter decurrens fere 1 mm. longum. Fructus 
ignotus.—‘‘Ostafrika: Nyassa-Hochland Kyimbila im Bambus- 
se bei Bulongwa, ca. 2100 m ii. d. M. (blithend 17. September 
913—Ap. Srouz n. 2175!).”’ 


“Die Art steht in der Fruchtbildung augenscheinlich Th. 
rhynchocarpum Dill. und Rich. nahe, das gleichfalls nur wenige 
bis 1 Fruchtblatt . . . hat aber einen sehr lockeren, sparrig ver- 
zweigten Bliitenstand und sehr kleine griine Bliiten. Th. Stolzii 
ist leicht kenntlich an den grossen rétlichen oder lila Bliiten in 
einem (im vorliegenden Material) dicht zusammengezogenen 
Bliitenstande. Leider liegen keine bis zur Fruchtreife ent- 
wickelten Pflanzen vor; es ist jedoch anzunehmen, dass sich der 
Bliitenstand spiter in vielleicht ahnlicher Weise wie bei Th. 
rhynchocarpum streckt. 

“Es ist auffallig, dass unter dem sehr reichhaltigen Material, 
welches das Botanische Museum in Berlin-Dahlem aus dem trop- 
ischen Afrika erhielt, diese Art bisher nicht vertreten war; sic 
scheint demnach selten und bisher iibersehen zu sein. 

‘“‘Anmerkung: Im gleichen Gebiete: Kyimbila, Mkinga, Mwak- 
alila, Bergwiese in 2000 m ii. d. M. sammelte Ap. Strouz (8. 
Januar 1914 n. 2421!) eine Pflanze, die wohl gleichfalls hierher 
zu stellen ist. Diese ist nur etwa 40 cm hoch, unverzweigt mit 


1944] Boivin,—American Thalictra and their Allies 397 


kleinen Blaittern und meergriinen Blattchen mit meist dreilap- 
piger Spreite. Die Bliiten (es liegen nur Knospen vor) sind 
gleichfalls sehr gross und nach Angabe des Sammlers weiss, 
aussen rosa. Es handelt sich vermutlich nur um eine Lichtform 
von Th. Stolziz, doch ist das Material leider zu spiirlich, um eine 
sichere Bestimmung zu erméglichen.”’ 

The preceding text is a copy of the original description and 
comments. 

No material has been seen which could be ascribed to this 
species; however the characters of the flowers contrast strongly 
with all other species of this section. Ulbrich’s description of 
the inflorescence of 7. Stolzii applies equally well to the flowering 
stage of any species of this group. Similarly his description of 
the inflorescence of 7. rhynchocarpum Dill. & Rich. characterizes 
all fruiting material belonging to the § Crncrnneria. The 
leaves are described as “pinnata,” but this we much doubt. If 
true indeed, there should be no trouble at all to recognize the 
species at any stage of its life-cycle. The fruits are not described 
and it is difficult to guess what they look like. The only material 
at hand from Nyassa (Stolz 148, 25 Dez., 1907), bears immature 
fruits strongly suggesting those of T. snnitons but they are some- 
what narrower. This latter specimen is cited under 7’. innitens. 

36. T. Steinbachii, sp. nov. Planta ca. 3 m. alta. Pedicelli 
5-10 cm. Sepala ca. 1.8 mm. longa, Ovaria nervosa pauca, 
2-4 in flore. Stylus una stigmate 4-7 mm. Carpella matura 
recurvata semi-ovata, 6.0-8.5 mm. longa, stipite 1.5-2.0 mm., 
ventre 3.0-5.0 mm. longo, 1.5—-2.0 mm. lato, nervo dorsali concavo. 
Floret verosimiliter Decembri.—Botiv1a: Steinbach 869, Inca- 
chaca, Prov. Chapare, age Cochabamba, Feuchte Béschung, 
Meereshéhe 2200 m., Jan. 21, 1929 (G, Type and 1soTyPE; F, 

Y, ANS, wont. Fig. 36, a-c 

One specimen bears the following soak: ca 3 M. Ranksustande. 
This plant is probably no more of a climber than T’. innitens but, 
just as does the latter, probably gains additional support when 
its inflorescence gets entangled in the surrounding vegetation. 

37. T. SreyeRMARKII Standley, Field Mus. Nat. Hist. Bot. 
22: 230, 1940. Planta 1.0-2.5 metrorum, glabra nisi in earpellis. 
Folia nonnunquam metralia, ad septies ternata, saepius con- 
spicue stipellulata. Pedicelli ca. 5 em. Sepala. aguiay ver ae 
purascentia ca. 3 mm. longa. Filamenta ca. 8 mm. 
oblongo-lanceolatae ca. 3 mm., acumine 0.2-0.6 mm. "Stele 


398 Rhodora [NovVEMBER 


una cum stigmate ca. 13mm. Carpella matura firma 9-12 mm. 
longa, cinerea, pubescentia. Pubescentza pilorum translucidorum 
rectorum densorum arachnoideorum. Si pubescentia tactu 
deleta est, carpella purpurascentia revelantur. Nervi fructuum 
in costis sublati, plus minusve ramosi et anastomosi, minute ad 
summas sinuosi et nonnunquam gibbosi, nervus ventralis con- 
vexior et semper gibbosus, dorsalis rectus esset nisi ad medias 
parum concavitas. Stipes fructus. ca. 1 mm., venter 5,.5—-6.5 
mm. longus et 3.0—4.0 mm. latus, rostrum 2.5—-4.0 mm. directum, 
paries 0.3-0.5 mm. crassitudinalis. Verosimiliter floret No- 
vembri. GUATEMALA, SAN MARCOS: Steyermark 36,575, north- 
western slopes of Voledn Tajumulco, barrancos south and west 
of town of Tajumulco, moist slopes around seepage at base of 
barranco, herbaceous, 8 ft. tall, leaves blue-green above, grass- 
green beneath, follicles glaucous-green, alt. 2300-2500 m., Feb. 
25, 1940 (F, typz, mounted on two sheets); Standley 68,465, 
above San Rafael Pie de la Cuesta, Barranco Eminencia, wet 
meadow, common, alt. 2100-2400 m., March 14-15, 1939 (F, 
paratype); Steyermark 36,483, near southeast portion of Volcan 
Tacand, between Canjula and La Unién Judrez, herb 5 ft. tall, 
leaves membranaceous, grass-green above, silvery-green beneath, 
Feb. 22, 1940 (F, paratypes) ; Steyermark 36,989, lower to middle 
slopes of Volcdn Tajumulco, between Todos Santos and Finca 
El Provenir, slopes around quebrada, alt. 1300-3000 m., March | 
1, 1940 (F, paratype); Steyermark 36,191, between La Vega 
ridge along Rio Vega and northeast slopes of Voledn Tacand4, 
to 3 miles from Guatemala-Mexico boundary, in vicinity of San 

ael, shaded banks of slopes bordering escarpment of stream, 
herb 5 ft. tall, alt. 2500-3000 m., Feb. 20, 1940 (F, paratype). 
QUEZALTENANGO: Standley 83,523, south of San Martin Chile 
Verde, region of Las Nubes, damp densely forested barranco, 
herb 2 m. tall, alt. about 2250 m., Jan. 16, 1941 (F); Standley 
85,131, South of San Martin Chile Verde, region of Las Nubes, 
on white sand mountain side, damp thicket, herb 1.5 m. tall, 
alt. about 2250 m., Jan. 27, 1941 (F); Standley 85,680, between 
San Martin Chile Verde and Colombo, above Mujuli4, in damp 
dense mixed forest on white sand slopes, herb 1.5 m. tall, alt. 
about 1800 m., Feb. 1, 1941 ); Standley 83,652, south of San 

: 0 


Standley 85,140, south of San Martin Chile Verde, region of 
Nubes, ae thicket, herb 1 m. tall, Jan. 27, 1941 (F). Fie. 


Planta verosimiliter ca. 3 


1944] Boivin,—American Thalictra and their Allies 399 


ventre 5.0-7.5 mm. longo, 2.5-3.0 mm. lato, nervis parum ramosis 
et parum anastomosis nervo dorsali fere recto nisi esset ad medias 
parum concavus. Tempus florendi ignotum  verosimiliter 
vernale.—Botivia: Steinbach 9231, Incachaca, Prov. Chapare, 
Dept. Cochabamba, Wald Meereshéhe 2300 m., Febr. 18, 1929 
(G, TypE; ANS, F, NY, tsoryres). Fia. 38, a and b. 


Judging from the herbarium fragments, this species is probably 
as high if not higher than 7’. Steinbachit or even than 7. rhyncho- 
carpum Dill. & Rich. (7. Mannii Hutchinson). 


39. T. MAcRocaRPUM Grenier, Séances Publ. Acad. Sci. 
Besancon 117. 1838. T. majus Gren. Act. Soc. Linn. Bordeaux, 
: 2, 1836, nec T. majus Crantz, Stirp. Austr. 2: 80, 1763, nec 
T. majus Jacq. Fl. Aust. 5: t. 420, 1788, nec 7. majus Tenore, 
Syll. Pl. Vase. Fl. Neap. 265, 1831, nee 7. majus Dunn, Ind. Sem. 
Hort. Panorm. 32, 1880. —Planta omnino glabra semimetralis. 
Folia inferiora 1-3 in planta, 4-ternata. Folia caulinaria desunt. 
Folia inflorescentiae inferioris 3-ternata foliolis 3-5-lobatis. 
Folia inflorescentiae superioris ovata integra. Stipulae et 
stipellulae desunt. Inflorescentia simplex vel, si ad basem ramosa, 
ramis bi-tri-floriferis foliumque solitarium ferentibus. Pedun- 
cult 5-10 em., aetate fructuum maturandorum. Flores lutei. 
Sepala elliptica 5-6 mm. longa. Antherae ca. 3 mm., acumine 
1.0-1.5 mm. Ovaria 2-4 in flore. Stigma 8-9 mm., in primis 
rectum vel recurvatum, deinde incurvatum, alis ca. 0. '5 mm. lat. 
ied aoe matura valde compressa, stzpite 0.5-2.0 mm., ventre 
8.5-10.0 mm. longo et 3-4 mm. lato, nervo dorsali fere recto vel 
parum convexo, ventrali multo quam dorsali convexiore, nervis 
lntdrafibus conspicue ramosis et anastomosis. Floret Julio et 
ugusto.— France: Cosson, Basses Pyrénées, prope Eaux- 
Bonnes, in monte, montagne de Gourzi, 7 augusti, 1847 (G); 
Forestier, prés les Eaux-Bonnes, pic d’Anis, 5 aofit, 1841 (G); 
Philippe, Basses Pyrénées, col de Tortes, aotit, “i (G). Fria. 39. 


Sectio Camprocastrum Prantl, Nat. Pfl. 3, 2: 66, 1888. 
Radiz fibrosa. Folia saepe stipellulata.  Follslae haud peltatae. 
Pedicelli sub fructum recurvatum. Flores polygamo-monoici, 
plurimi perfecti, pauci masculi, raro perfecti omnes. Masculi 
nonnulli ad anthesim veniunt cum carpella perfectorum maturant. 
Antherae oblongae lanceolatae vel lineares, plus minusve acum- 
inatae. Filamenta purpurascentia nec clavata. Ovaria sessilia 
vel stipitata, saepius, si adsunt, 4-10 in floribus singulis. Steg- 
mata filiformia, nec alata. Carpella matura plus minusve com- 
pressa, nonnunquam stipitata, nervis plus minusve ramosis, 
saepe et anastomosantibus et pr eicta et gibbosis. Species 
typica Thalictrum gibbosum Lec. sit. 


Some of the species are RE in the text as having all 


400 Rhodora [NOVEMBER 


flowers perfect, but this is a point still imperfectly known, for 
when the ovaries start enlarging the plant still goes on blooming 
but these later flowers are always staminate. This stage seems 
not to be represented for all species of this section. However, 
T. Venturii and T. inuncans are definitely known as always 
having perfect flowers. 

The following key is based mainly on pubescence, notwith- 
standing the fact that the fruit has furnished the main characters 
on which to base each of the successive species. Only about one 
third of the herbarium specimens having mature fruits, and the 
flowers not being very much varied, it is not possible to build up 
a natural key which would be a satisfactory tool with which to 
identify herbarium material. Although this key is artificial it 
is still not possible to identify all flowering specimens, but it has 
been found that by comparing such flowering plants with only 
those species known to occur in the region of the plant to be 
named, it was usually possible to arrive at a satisfactory identifi- 
cation. This is why distribution-data have been given in the key. 
a. Ovaria 18-36 in flore. In Argentina.................. 71. TT. Venturi. 
a. Ovaria 0-10 in flore. ...b. 

b. Pubescentia diversa, aliqui pilorum aciculariformes com- 

ressi triangulares et pa ae recurvati. Staminum 
enta 15-17 mm. ta MU a ee cs o's 40. T. inuncans. 
b. Pili aciculariformes “ recurvati de ane Staminum fila- 
menta breviora. .. . 
¢. — nana chet -< alta, foliis imis plurimis, folium 


vel solitarium vel deest, folia inflorescen- 
tine vel 1-3, vel desunt. Foliolae parvae, 3-10 mm. 


longae. In ‘Hidalgo Ree ey we he cas 50. 7. pachuense. 
¢. hyn caulinaria plura.... 
d. Foliolae ~~ dentatae, acuminatae. Pubescentia 


e pilis tis stipitatis. In Guatemala...... 51. 7. Standleyt. 
d. Foliolae apice trilobatae vel grosse dentatae, nec acu- 
bescentia vel e pilis abs vel 


e. Foliolae inferne re a osr 
Bf 7 vie una cum stigma (-5) mm..... 
Filamenta 3-5 mm. Carodie aitiea 27-3. 5 
mm. lon, : 
h, Filamenta 4-5 mm. Carpella matura ca. 
mm. longa, nervis simplicibus nec re 
ae a PO COR. os 41. T. Galeotti. 
h. ey rang ca. 3mm. Carpella matura abd 
mm. longa, nervis ramosis et anastom 
santibus, nervo ventrali gibboso. In Bonork 
T. parvifructum. 
9. Brey 6-10 "aus Carpella matura 6-9 


ae at ee 
sis iene hn, Seon 


1944] Boivin,—American Thalictra and their Allies 401 


7, Antherae apice truncatae, vel acumine 0.1-0.2 
mm. Carpella matura gi . In Gua- 
tamale. fe ee eee 52. T. Johnstonii. 
7. Antherae acumine 0,5-1.0 
matura non gibbosa. In Peruvi, Bolivia, 
e ueeareh 5 ee ct oan s 43. T. decipiens. 
A bg ba una stigmate (4—) 5-15 
. Ca Lg matura nervo veuteadi nullo ee gib- 
k. Nervis lateralibus parum ramosis, nec sinu- 
osis, nec Peeper sina ager nce eo 
i Stylus una stigmate 4-10 m 
m. Carpella m ee ots 5 orth loneai In 
_,, Mexico ot Costa Rita: 2 sees. . 42. T. Hintonii. 
n. 


nta nonnunquam glauca. 
mE Peruvia, Bolivia et Meg 
43. T. decipiens. 
n. Carpella 3.0-3.2 mm. lata, = ger. 
sali basi concavo. Stigm 
ao - mH Planta cacti modo 
glau Pat cee: T. viridulum, 
is ide una stigmate 10-15 mm. In : Beri 
7; "icscelnascsaut 


ra 3 : 
Colombia et Ecuador.......... 70. T. podocarpum. 
o. Stipes ovarii 0-0.5 mm. gai matura 
4.5-8.0 mm, longa 


p. ne matura 6.5-8.0 mm. longa. In 
‘amaulipas, Nuevo Lada et San Luis 
Potosi i Rian Sees (ewe Pee ee 47. T. Deamit. 
p. C ura 4.5-5.5 mm. longa... .¢. 
q. Folia pare eel conspicue_petiolata. 
by glauca. n ua, 
Durango, Nuevo Leén et San Luis 
Re Bee wg Poa 48. T. grandifolium. 
q. Folia caulinaria sessilia in apice dilata- 
—— Pegging de a nullo modo 
In Guerrero....... 49. 7. Hernandezii. 
j. Carpella mature ees ventral 'gibboes, nan 
— saepius gibbosis. Gibbositas patet non- 
eater sr gates saepius tamen in carpellis 
r. Sangale mia . Pies mm. lon 
semimetralis minor. ae 
54. are parvifructum. 
r. GO matura 4.0-10.5 mm. longa... .8. 
lla matura mm. longa ; 


ey 
: 


402 Rhodora [NovEMBER 


_u, Petioluli plus age hd ascendentes. 
Se 


v, let matura 4-6 mm, longa, 
8-2.0 m. 


a 
ee 64. T. peninsulare. 
v. Carpella matura 4-8 mm. longa, 2-3 


0.2-0.5 mm. ongus. In Ver 
Cruz, Hidalgo, ees Puebla, 
Morelos et Oaxaca......... 65. TT. gibbosum. 


t. hay oe matura 7-8 mm. longa. 
w. Carpella matura saepius apice acuta, 
—8 mm. longa, 2-3 mm. lata, stipite 
0.5-2.0 mm. 


w. Carpella matura saepius apice obtuso, 
—7.5 mm, longa, ca. 3.3 mm. lata, 
oS 1.0-2.5 mm, Sepala 5-6.5 


s. C lla hacer 9.0-10.5 mm. longa. ee 
lombia, Venezuela et Ecuador. 
. TT. podocarpum. 
e. sae inferne plus minusve pubescentes. . . . 


mtia caulinaria e pilis opacis i Ghiitdadis 
acuminatis 1-4 millimetralibus. Foliolae in- 
ferne sparse dahane entes. In Oaxaca et 
WMO rs fae a ay a As 53. 7’. lanatum. 
x. Pubescentia caulinaria e ae millimetralibus vel 
brevioribus vel deest. . . . 
y. Pubescentia foliorum paginae inferioris e - 
omnibus capitatis... . 
z. Planta omnino pu hak carpellis 
ca. 6.5 mm. longis, 2.0-2.2 mm. ‘ae ps 
45. '. panamense. 


z. — lee super glabrae ae 3 fe hares. eis 
et rameales uniseriati et 


Jonga. In eee Ce T. Pennellii. 


6. C a glabra is * Vase Crus, ae 
Federal et Oaxaca........ 


T. Conzattit. 
6. Epes tet woe, cs ens 


1944] Boivin,—American Thalictra and their Allies 403 


«. Folia caulinaria conspicue _petiolata. 
Carpella matura 3.3-5.0 mm longa. 
In Tallse 0, Hidalgo et Maticankes 
. TT. pubigerum. 
e. Folia caulinaria. sessilia vel fere sessilia 
in apice dilatationis ae es Car- 
oe matura In 
xico, Michoacin et ot Puebla. 6 62. T. sessilifolium. 
y. Pubescentia e pilis omnibus crassiusculis 
acis coloratis, vel aliis crassiusculis 
er amg opacis obtusis. interdum acu- 
iieriey ot 
c. Carpets penta 7-3.0 mm. longa. Stylus 
una. sti te 3-5 mm. Planta semi- 
metralis Gel brevior. me Pindrancgs 55. T. parvifructum. 
¢. Carpella matura 3.5-8.0 mm. longa... .7. 


ruZ, go i 
Federal et Michoacin...... 56. “T. strigillosum. 
§. Carpella matura 7-8 mm. longa. In 
Bie toe a. Sa 4 oa oe T. Nelsonii. 
n. Foliolae inferne sparse pubescentes. .. . 
t. : oe _ mm. = Peruvia. . 66. 7. longistylum. 
. Stigma eee 
is Carpella ma matura. 5-6.5 mm. longa. 
n Peruvia....d. 
me Carpe matura stipite 0.5-1.0 
kw pin gL Ote. Ue ep wears 58. 7’. lasiostylum. 
X. Canela matura stipite ca.2mm __ 
o. T. rutidocarpum. 
C. la Sato 7-8 mm. longa. 
. a Oagnea Se 69. 7. Nelsonit. 


This section is restricted to the mountains of Mexico, Central 
America and South America. In general, the species are based 
on the different types of fruits, for the flowers offer but few 
specific characteristics. However, two species, 7. gibbosum Lec. 
and T. decipiens nostrum, show a high degree of variation in their 
fruits and they are pretty hard to characterize. A key to the 
flowering specimens would leave about 8 or 10 species, all 
glabrous, which could not be otherwise separated were it not for 


the fact that their ranges do not overlap in general. The inter- 


pretation of the different names already published is based on 
types or isotypes or syntypes or photographs of types or frag- 
ments of types. Only in two cases, 7. Hernandezii Tausch and 
T. lasiostylum Presl, was it necessary to form an opinion on the 


original description only. Lecoyer did not have the opportunity 
of studying the types of those two species and considered them 
aS synonyms. 


404 Rhodora [NOVEMBER 


Subsectio Simpricia, subsect. nov. Plantae glabrae, Thalictro 
inuncante nostro excepto. Ovaria 10 vel pauciora in flore. 
Carpella matura compressa nervis simplicibus vel parum ramosis 
anastomosantibus et reticulatis, nec gibbosis, raro sinuosis. 

40. T. inuncans, sp. nov. ’Planta verosimiliter 1.5-2.0 m. 
alta. Pubescentia caulis ramorum petiolorum petiolulorumque 
densa e pilis aliis capitatis translucidis, aliis crassiusculis tri- 
angularibus compressis recurvatisque. Foliolae superne g abrae, 
inferne puberulentes pilis Seager translucidis vestitae. Flores 
perfecti (!) glabri. Sepala ovata ca. 5 mm. longa. Antherae 
nonnunquam faleatae ca. 4 mm., vel apice truncatae, vel ob- 
tusae cum acumine 0.1-0.2 mm. Filamenta mm., con- 
spicue purpurascentia. Stylus una cum stigmate 6-11 mm. 
Stigma paullo et regulariter a ap incrassatum. Fructus mihi 
ignotus. Floret Decembri—Botivia, Tarisa: Fiebrig 2440, 
Tecumilla bei Tarija, alt. 2000 m., Des. 20, 1903 (G, Type and 
ISOTYPE: US, isotypr). Fia. 40, a-c. 

Although known only from one collection and at flowering 
time, this species should be easily recognized by its dense covering 
of very small recurved prickles, the only such case known to me 
in this genus. The filaments of the stamens are also longer than 
in any other species of Thalictrum. This species shows some 
relationship with T. Venturii, but its position is dubious. Both 
of these species have perfact flowers, obtuse or truncate anthers 
and somewhat clavate stigmas. 


41. T. Gatzorri Lecoyer, Bull. Soc. Bot. Belg. 24: 121, 1885. 
Planta omnino glabra altitudinis ignotae, verosimiliter metralis 
vel brevior. Flores forsan omnes perfecti. Sepala 3 3.5-4 mm. 
longa. Antherae 1.2-2.0 mm., oblongo-lanceolatae, in apice 
truncatae, vel acumine 0.1 mm. Filamenta 4-5 mm. Stylus 
una cum stigmate 3-4 mm. Carpella matura subsessilia vel 
sessilia, ca. 3.5 mm. longa, ca. 1.5 mm. lata, nervis lateralibus 
parum ramosis nec undulatis nec gibbosis, nervo. dorsali fere 
recto, ventrali convexo. Tempus florendi ignotum.—MExIco, 
VERA CRUZ: Galeotti 4541, pic d’Orizaba, fl. blanches, alt. 10,000 
p-, Juin-oct., 1842 (syntype at the Conservatoire Botanique ~ 
>i F, photograph of this syntype; G, syntype). Fie 


Naturally, I have not seen the syntype in Switzerland, but I 
cannot find anything which matches either the photograph or the 
syntype at hand. The other syntype is no. 4570 from Xalapa 
by the same collector. Judging from Lecoyer’s drawing, which 
naturally was made from one of those syntypes, and the photo- 


pe oy Ne See We ne RON RO a 


) ea MEAL eer ee ore Cercnen nee ner nr en ena, eee 


i 
i a 
Pa 
1 = 
be a 
hie 
ee 


1944] Boivin,—American Thalictra and their Allies 405 


graph at hand, the fruit resembles pretty much that of what we 
are calling 7’. Hernandezit Tausch, but it is much smaller. The 
anthers and stigmas also rank amongst the smallest in that 
section of the genus. 

42. T. Hintonii, sp. nov. Planta omnino glabra 40-120 cm., 
radicibus fibrosis. Stipulae parum dilatatae. Foliolae fere 
orbiculares, basi cordatae, apice trilobatae, lobis rotundis 
mucronulatis, nonnunquam figura Hepaticae americanae, varia- 
biles tamen. Inflorescentia 1-3 dm. longa, plus minusve secunda, 
pedunculis vel rectis vel parum recurvatis. Flores non satis 
notae, sepalis tamen ca. 4.5 mm. longis, antheris ca. 5 mm., 
acuminibus ca. 1.0 mm. Carpella matura fere orbicularia vel 
semi-ovata, 4.5-5.5 mm. longa, stipite 0.1 mm., ventre ca. 4 mm. 
longo 2.0-3.2 mm. lato, nervis parum ramosis nec gibbosis, nervo 
dorsali recto vel vario modo convexo, ventrali convexiore quam 
dorsali. Floret tempore mihi ignoto.—MExico, MExIco: Hinton 
6743, Temascaltepec district, Acatitlin, in the water, Oct. 12, 
1934 (US, type; F, G, M, NY, isotypes). GUATEMALA, 
QUEZALTENANGO: Standley 67,887, southwest of San Martin, 
Chile Verde, Cumbre de Tuilacdn, sandy Alnus forest, alt. about 
2400 m., March 8, 1939 (F). Costa Rica: Standley 35,247, near 
the Finca del Voledin de Turrialba, southern slope of Volcan de 
Turrialba, wet thicket, herb 3-4 ft., alt. about 2000 to 2400 m., 
Feb. 22, 1924 (US). Fic. 42. 

Dedicated to the late G. B. Hinton whose abundant and well 
prepared Mexican specimens form a large proportion of and are 
among the best of the material at hand. 


43. T. decipiens, sp. nov. Planta omnino glabra, 4-15 dm., 
plus minusve glauca, radice forsan tuberosa et solitaria. Inflores- 
centia saepius foliosissima. Pedunculi sub fructum recurvati. 
Sepala 3-7 mm. longa, elliptica vel orbicularia. Flamenta 6-10 
mm. Antherae 2.5-5.0 mm., acumine 0.5-1.0 mm. Stylus una 
cum stigmate 4-8 mm. Carpella matura obovata vel semiobo- 
vata, 6-9 mm. longa, stipite 0.5-1.5, ventre 3.5-6.5 mm. longo, 
2-3 mm. lato, nervis vel simplicibus vel ramosis anastomosanti- 
busque nec gibbosis, nervo dorsali fere recto vel plus minusve 
convexo, ventrali convexiore quam dorsali, rostro nonnunquam 
recto saepius tamen ad tergum deflexo. In Peruvia floret a 
Novembri ad Januarium, in Bolivia Novembri et Decembri et 
in Argentina a Septembri ad Decembrem.—PERU: Soukup 140 
(F); Ball, ex saxosis Andium, juxta pagum Chicla, 12—-13,000 p. 
8. m., apr. 21-23, 1882 (NY). sunin: Killip & Smith 22,125, 
near Huancayo, open rocky hillside, herb to 5 ft., alt. 3300-3500 
m., Apr. 26, 27, May, 25, 1929 (F, NY, US); Kalenborn 94, 
Vicinity of Oroya, among rocks, alt. 10-17,000 ft., 1918-1919 


406 Rhodora [NovEMBER 


(M, NY, US). uma: Killip & Smith 21,758, Rio Blanco, open 
Ailide, herb 2-4 ft., alt. 3000-3500 m., Apr. 15-17, 1929 (NY, 
US); idem 21,752, herb 1-2 ft. (F, NY, US). PAUCARTAMBO: 
Herrera 1047a, Hacienda Churu, alt. 3500 m., Jan., 1926 (US, 
typE; F, G, NY, 1sorypss). cuzco: Pennell 13 ,040, Quiquijana, 
grassy wayside, herb, alt. 3220 m., Apr. 23, 1925 (ANS, F); 
Soukup 40, Urubamba, entrada al valle Lares, Jan. 3, 1936 (F); 
Vargas 702, Hda Urco, alt. 2940 m., 1938 (F); Herrera 829, 
Sazaihuamén, hills, alt. 3200-3600 m., Dec., 1928 (F); Soukup 
254, near Puno, alt. 4000 m., Jan., 1936 (F). Boxrtvra: Kuntze, 
alt. 3600 m., Apr. 1-4, 1892 (NY). COCHABAMBA: Steinbach 
8799, valle de Cochabamba, alt. 2600 m., 27 Dez., 1928 (ANS). 
LA Paz: Buchtien 6289, Cotafia am Ilimani, an Wassergriiben, 
alt. 2500 m., Nov., 1911 (F, US); Bang 1314, vic. Sorata, 1892 
(ANS, F, G, NY, US); Rusby 601, Sorata, alt. 10,000 ft., 1886 
(ANS, G, M, Us). ARGENTINA, SALTA: Eyerdam & Beetle 
22,631, 14 km. west of Manuela Pedraza, Tartagal river, moist 
sand, stream -bank, semi-shade, perennial herb 1-1.5 m., alt. 
900 m., Oct. 26 , 1938 (G). [N. B. This specimen is the only 
one complete with underground system, a tuber about 3 inches 
long.| TUCUMAN: xe 930, dept. ee cave Buena, en una 
quebrada con agua, altura “ ‘la plant , flor amarilla, alt. 
700 m., Sept. 24, 050 (F, M, US); Venturi 3996, dept. Chicli- 
gasta, ‘Estancia ‘Las Pavas, en los prados, flor ’ amarilla, alt. 
1700 m., Die. 11, 1926 for 19257] (G, 08); Job 1425, Clavillo de 
Aconquija, flor verde, alt. 2800 m., 1937 (NY). CATAMARCA: 
Jérgensen 1304, El Candado, alt. 2700 m., Oct. 2, 1916 (G, M). 
cérDoBA: Hieronymus, am Ufer des Rio Primero bei Cérdoba, 
Nov. 17, 1877 (F, US); Kurtz 2707, Cérdoba, Rio I Cerro del 
Pueblo, Nov. 29-30, 1885 (NY); Job 429, La Falda, Cerro El] 
Charrito, alt. 950 m., Jan., 1936 (US); Kurtz 342, San Virente, 
~ umbrosis humidis frequens, Oct. 11, 1884 (NY); ’ Burkart 7198, 

a Reducién, Cérdoba, orilla de arroyos, hasta 2 m. de altura, 
Des. 27, 1935 (G, US). Fia. 43, a and b. 


This species is railed heterogenous but the herbarium material, 
although abundant, is rather unsatisfactory; most collections at 
hand are either unicates or made up of duplicates collected at 
different dates and localities; of a hundred or so herbarium sheets, 
only one had the underground system and even so it was dam- 
aged. Further segregations might well be possible, although I 
attempted it many times unsuccessfully. 


44. T. viridulum, sp. nov. Planta viridula omnino glabra 
nullo modo glauca. Folia caulinaria perfecte 5-ternata, petiolulis 
rectis. Flores polygamo-monoici, plurimi perfecti, pauci masculi. 
Sepala viridula ca. 4 mm. longa. Antherae acumen 0.5-1.0 mm. 


1944] © Boivin,—American Thalictra and their Allies 407 


Stylus una cum stigmate 8-10 mm. Carpella matura nullo 
modo gibbosa, ca. 7 mm. longo, stipite 0.7-1.0 mm., ventre 
5.0-5.5 mm. longo, 3.0-3.2 mm. lato, nervis lateralibus plus 
minusve ramosis anastomosantibusque, nervo dorsali basi parum 
concavo, apice fere recto, ventrali convexo, rostro ad ventrum 
deflexo. An floret Junio et Julio?—Pa ANAMA, CHIRIQUI: Sezbert 
204, valley of the upper Rio Chiriqui Viejo, vicinity of Monte 

irio, growing in rocky gorge, 1—1.5 m. tall, fls. green, styles and 
sa aee Dee ae alt. 1300-1900 m., June 27—July 13, 

1935 (NY, typr; G, N-ND, tsorypss). Fig. 44. 

45. T. ee Pintle Standley, Field Mus. Nat. Hist. Pub. Bot. 
22: 19, 1940. Planta 6-12 dm., caule ramis petiolisque plus 
minusve glaucis, omnino pubescens. Pubescentia e pilis jon nc 
translucidis. Folia caulinaria 4-5-ternata. Stylu 
stigmate ca. 8 mm. Carpella matura sigmoidea nec gibbosa, fe. 
6.5 mm. ae stipite ca. 1 mm. ventre ca. 4.5 mm. longo, 2.0— 
2.2 mm. lato, nervis lateralibus simplicibus vel parum ramosis, 
nervo dorsali baso concavo, apicali convexo; ventrali convexiore 
quam ag rostro ad ventrum deflexo. An floret Junio?— 
PANAMA, CHIRIQU{: Davidson 791, Boquete, flowers oe 2 
to 4 ft. baph alt. 4500 ft., June 26, 1938 (F, ryPE). Fia. 

46. T. Macbrideanum, sp. nov. Planta elata omnino Cane 
ad 12 dm. attingens et verosimiliter arcuans. Caulis subteres. 
Folia inflorescentiae estipellulata 3—4-ternata. Injlorescentia 
ampla paniculata (an secunda?), ramis divaricantibus gerbe 
foliosis. Peduncult 2-3 cm. ad aetatem.florendi, 4-5 et 
parum curvati cum fructus permatuerunt, nee sub Biche hme oo 
reflexi. Flores perfecti cum staminibus 16-20 et pistillis 3-6. 
Sepala quatuor, late ovalia, 5.0-6.5 mm. longa. Stamina fila- 
mento 6-8 mm., antheris 2.5 mm. oblongo-lanceolatis, mucrone 
0.3-1.2 mm. longo. Stylus una cum stigmate 12-15 mm. non- 
nunquam persistens. Carpella matura valde compressa, fere 
laminaria, semi-orbicularia, praca nec gibbosa, stipite ca. 2 
mm., ventre ca:’4.5 mm. lon o, ca. 2.5 mm. lato, nervo ventrali 
semi-circular, dorsali subrecto, “lateralibus conspicuis rugosis 
curvatis nec ramosis nec sinuosis, rostro ad tergum deflexo. 
Floret Junio.—Prrvu: Macbride 4466, Tambo de Vaca, June 10— 
24, 1923, in ee to 4 ft. high, pistils purple, anthers a 
alt. a about 13,000 ft. (G, Type: F, NY, US, isotypes). Fic. 


The type specimen is made up of three fragments evidently 
homogeneous, one with mature fruits, the second with half-open 
flowers, the third one larger, with flowers in full bloom. For 
this unusually fine herbarium specimen, we are thankful to the 
collector, J. Francis Macsring, and it is a pleasure to dedicate 
to him this new species. The length of the usually persistent 


408 Rhodora [NovEMBER 


style sets this species apart from any other Thalictrum we know 
of. And before the stamens reach the length of the sepals, the 
style is already full grown. 


47. T. Deamii, sp. nov. Planta omnino glabra. Sepala 
viridula ca. 25 mm. longa. Filamenta ca. 5 mm. Antherae ca. 
4 mm., acumine ca.0.5 mm. Stylus una cum stigmate 8.5-10.5 
mm. Carpella matura ovato-lanceolata, 6.5-8.0 mm. longa, 
stipite usque ad 0.8 mm., ventre 6-7 mm. longo, ca. 2.5 mm. lato, 
nervis rugosissimis valde anastomosantibus parum undulatis nec 
tamen gibbosis, nervo dorsali fere recto vel parum convexo, 
ventrali convexiore quam dorsali. Tempus florendi ignotum.— 
Mexico, TAMAULIPAS: von Rozynski 558, Sierra near San Lucas, 
Jamauve, July, 1982 (F, US). Nurvo Lré6n: Kenoyer 120, 
Monterrey, field, Sept. 18, 1937 (F). San Luis PoTost: Pringle 
5071, Las Canoas, June 23, 1891 (G, TyPK). Fic. 47. 


Dedicated to a botanist whom we admire very much for his 
patient and painstaking work, namely his herbarium specimens 
and his publications on the flora of Indiana. His Flora of 
Indiana contains the best key that was ever published to separate 
three very litigious species: T. dasycarpum Fisch. & Lall., T. 
polygamum Muhl. and T. revolutum DC. 


48. T. GRANDIFOLIUM Watson, Proc. Am. Acad. 23: 267, 1888, 
nec T. grandifolium Rose, Contrib. U. §. Nat. Herb. 5: 143, 1897. 
T. grandiflorum Watson ex Rose, Contrib. U. 8. Nat. Herb. 5: 
188, 1899, nee T. grandiflorum Maxim., Act. Hort. Petrop. 11: 
11, 1889, nec 7. grandiflorum Rose, Contrib. U. S. Nat. Herb. 5: 
188, 1899 .-—Planta verosimiliter semimetralis ad senior aoe 


parum jncurvatus, in apicem valde recurvatus. Flores omnes 
perfecti (?) vel plurimi pesioehs pauci masculi. Sepala 3-5 mm. 
longa. Filamenta 4.0-5.5mm. Antherae 3.5-5.0 mm., acumine 
0.5-1.5 mm. "Sip lus una cum stigmate 5-10 mm. Carpella 
matura suhatinitets. apice acuto vel saepius rotundo vel etiam 
prope rostrum retuso, 4.5-5.0 mm. longa, ventre ca. 4 mm. longo, 
2.5-3.2 mm. lato, nervis valde ramosis anastomosantibusque 
nervo dorsali aliquid convexo, ventrali convexiore quam do i. 
Floret a Junio per Julium ad ‘Augustum —MExIco, CHIHUAHUA: 
Pringle 1513, Sierra Madre, under cliffs, Oct. 17, 1887 (G, TYPE: 
ANS, CA sf, M, NY, UC, US, 1 SOTYPES): Nelson 6007, in the 
Sierra Madre, June 21-July 29 ."1899 (G, US); Hartman 761, 


FLOWERS OR Fruits oF THALICTRUM, all X 4, except 51a and 


4q Gn tae 
3 (For explanation see e end of paper.) 


410 Rhodora [NOVEMBER 


Pilares, Strawberry Valley, Sept. 22, 1891 (G); M. E. Jones, 
Soldier Canyon, alt. 6500 ft., Sept. 16, 1903 (CA, F, M, NY, UC, 
sie LeSueur 1323, Rio Negro, Aug. 29, 1937 (F): Townsend & 
arber 47, near Colonia Garcia, alt. 7500 Ft, June 23, 1899 (F, 
6 M, NY, US). puranao: EH. Palmer 357, Durango and 
vicinity, Apr. to Nov., 1896 (F, G, M, NY, UC, US). NUEVO 
LEON: C. H. & M. T. Mueller 899, Cieneguillas to Puerto Santa 
Ana, about 15 miles SW of Galeana, commonly scattered Ay 
dense moist wood, flower straw with lavender stamens, June 2 
1934 (F). san Luis Potosi: Pennell 17,910, Las Canoas, rocky 
limestone hill, flowers purplish, alt. 100-1200 m., Aug. 6-7, 1934 
ae ee Fic. 48. 
T. Hernanpezu Tausch ex Presl, Rel. Haenk. 2: 69, 1835. 
T. rabicutane Aschen ex Lecoyer, Bull. Soc. Bot. Belg. 24: 306, 
1885, nomen subnudum cum synonymis editum. Planta 
omnino glabra viridula nec glauca, metrum approximans. 
Flores polygamo-monoici, pauci masculi, alii perfecti. Sepala 
3-4 mm. longa, nonnunquam viridula. Filamenta ca. 5 mm. 
Antherae 3-3.5 mm., acumine circa millimetrali. Stylus una 
cum stigmate 8.5-10.0 mm. Carpella matura glauca semiovata 
ca. 5 mm. longa, stipite 0. 2-0. 5 mm., ventre ca. 4.5 mm. longo, 
3.3-3.6 mm. lato, nervis plus minusve ramosis anastomosantibus- 
que, nervo dorsali paululo convexo, ventrali conspicue convexiore 
quan dorsali. Floret Julio. —Mexico, GUERRERO: R. Q. sen 
, Taxco, rank-growing, in damp or ’ shady places, 1 meter 0 
2 al tall, , July 18, 1936 (G); idem, 237 and 238, July 12, 1937 (G). 


aa icc of this specific name is not quite certain. It 
might well apply to 7. gibbosum Lec. I have not seen the type, 
which is at Prague in the herbarium of the Museum. Lecoyer 
had not seen it either, but Dr. M. Willkomm studied it for him. 
The original description refers to a glabrous plant two feet high, 
from western Mexico, with ternate leaves, dioecious flowers and 
the nerves of the frail oblique and uneyen. There is no such 
Thalictrum amongst the thousands of specimens at hand. If the 
dioecious character is discarded, then it may be either 7’. gibbosum 
Lec. or the species just described above. Presumably Lecoyer 
had studied T. Hernandezii Tausch carefully enough to ascertain 
that it was not the same thing as his own T. gibbosum. Sensu 
Lecoyer, T. Hernandezii is what I am calling T. Nelsonii or T. 
strigillosum. I cannot agree with his interpretation, for 
Tausch very clearly says: “Caulis . . . una cum foliis flori- 
busque glaber’’. 


1944] Boivin,—American Thalictra and their Allies 411 


Subsectio Gibbosa, subsect. nov. Sect. Camptogastrum, d. 
Podocarpa PRANTL, Nat. P Pfl., 3, 2: 66, 1888. Plantae glabrae 
vel pubescentes. Ovaria 10° vel pauciora in flore 
matura compressa nervis ramosis anastomosantibusque saepius 
sinuosis et reticulatis, nervo ventrali, licet dorsali lateralibusque, 
gibbosis. Species typica Thalictrum gibbosum Lec. sit. 

0. T. pPacHUENSE Rose, Contrib. U. 8. Nat. Herb. 5: 188, 
1899. Planta 20-50 cm., ‘radicibus fibrosis, nonnullis istorum 
parum in locis incrassatis. Caulis basis subterraneus directus 
aphyllus, ad transitionem saepius ramosus et foliis numerosis 
munitus. Caulis aerius nonnunquam aphyllus, frequentius 
tamen cum folio caulinari solitario et cum 1-8 foliis inflorescen- 
tiae. Pubescentia deest, nisi interdum inferne, si adest e pilis 
incrassatis brevibus coloratis et in fructubus e pilis capitatis 
minutissimis translucidis. Foliolae 1 em. longae vel minores. 
Inflorescentia reducta. Pedunculi, fructubus maturis, paululum 
incurvati et sub receptaculo maxime recurvati. Flores omnes 
perfecti purpurascentes. Sepala 3-4 mm. longa. Filamenta 
5-6 mm. Antherae 2.5-4.0 mm., acumine 0.2-0.5 mm. Stylus 
una cum stigmate 5-9 mm. Carpella aietne 4-5 mm. longa, 
oblique clavata, interdum parum gibbosa, stipite 0.5-1.5 mm., 
ventre 3.4-3.6 mm. longo, 1.6-2.0 mm. lato, nervis sey en 
ramosis anastomosantibusque, nervo dorsali concavo. Flor 
Julio Augustoque.-—Mexico, HipaLeo: Pringle 6880, Sierra i 
Pachuca, open woods, alt. 9000 ft., July 16, 1898 (US, TYPE; 
AN _ US, 1soTyPEs); Pringle 9678, Sierra de 
Pachuca, alt. 9500 ft.., Aug. 22, 1902 (F, G, M, NY, US); Rose & 
Hay 5585, Sierra de Pachuca, 4 “is 21 & 22. 1901 (US). Type 


in foliolis supernis. Pubescentia e ‘pilis saepius paces inter- 
dum translucidis, aliis uniseriatis aliis stellatis stipitatisque. 
Folia 2—4-ternata. Foliolae profunde cordatae, apice acumina- 
tae, saepius parum as symmetricae, raro trilobatae, per marginem 
totam dentatae, dentibus serratis apice rotundis mucronulatis, 
sinubus acutissimis. Pedunculi variis modis recurvati. Flores 
polygamo-monoici, alii perfecti, alii masc Sepala oblonga 
6.0-7.5 mm. longa. Filamenta 7-8 mm. Antherae 4.0-4.5 mm., 
acumine 0.6-0.8 mm. Stylus una cum stigmate 9-11 mm. 
Carpella matura viridia carnosa obovata 4.5-5.5 mm. longa, 
stipite 0.2-1.0 mm., ventre 4.0-4.5 mm. longo, 2. 5-3.0 mm. lato, 
nervis obscuris copiose anastomosantibus, nervo dorsali ad basem 
et superne paululum concavo, ad medias parum convexo, ven- 
trali valde convexo. Rostrum dorsale ad superas insertum et 
deflexum adtergum. Floret Februario et Martio.—GuaTEMALa, 
SAN MaRcos: Standley 86,335, Barranco Eminencia, road 


412 Rhodora [NovEMBER 


tween San Marcos and San Raphael Pie de la Cuesta, in upper 
pest of the barranco pee Finca La Lucha et Buena Vista, 

wooded quebrada, herb 1 m. tall, rare, alt. 2500-2700 m., 
Feb. 6, 1941 (F); Steyermark 26 § 858, Rio Vega, near San Rafael 
and Guatemala-Mexico boundary, Voleén Tacanad, herb 5-8 ft. 
tall, leaves dull green above, grass-green beneath and rugose, 
sepals green, at bas e purplish, anthers pale yellow turning 
purplish, filaments light green, style purple, sepals erect, anthers 
expanding first, the pistils later, flowers olygamo-monoecious, 
stem-nodes purplish, alt. 2500-3000 m., Feb. 20, 1940 (F, TyPx); 
Steyermark 37,067, trail between Finca el Porvenir and San 
Sebastidn, Voledn Tajumulco, alt. 1300-1400 m., March 1, 1940 
(F, paratype); Steyermark 36,821, between town of Tajumulco 
and Tecutla (9 miles south and west of Tajumulco), northwestern 
slopes of Voledn Tajumulco, shaded thickets along quebrada, 
near top of barranco, herb 5-8 ft. tall, leaves membranaceous, 
rich green above, gray beneath, alt. 1800-2500 m., Feb. 27, 1940 
(F, paratype). QUEZALTENANGO: Standley 66 ‘346, mountains 
southeast of Palestina, damp dense forest, herb 5 ft., alt. about 
2700 m., Feb. 22, 1939 (F, paratype); Standley 66, 338, mountains 
southeast of Palestina, damp dense forest, tall coarse herb, alt. 
about 2700 m., Feb. 22. 1939 (F, paratypes); Standley 8), aie 
mountains southeast of Palestina, on old road to San Jus 
Ostuncalco, damp sandy hillside forest, herb 1-2 m an common, 
alt. 2550-2850 m., Jan. 21, 1941 (F); ‘idem 84, 258 ‘: Standley 
84,271, mountains southeast of Palestina a, on old r oad to San 
Juan Ostuncalco, damp sandy peng forest, herb 1- ‘L. 5 m. tall, 
Jan. 21, 1941 (F). Fig. 51, aa 

52. T. JonNsToNt Standley & Steyermark, Field Mus. Nat. 
Hist. Pub. Bot. 22: 229, 1940. Planta omnino glabra, 6 dm. 
alta vel altior. Florum plurimi perfect pauci masculi. Sepala 

.> mm. longa. Filamenta 7-10 m Antherae 1.2-3.5 mm., 
acumine 0-0.2 mm. Stylus una cum Calon anguste bialato 

—4 mm. Carpella matura ignota, fere matura 6-7 mm. longa, 
stipite ca. 1 mm., nervis lateralibus valde reticulatis, nervo 
dorsali ——_ concavo, ventrali gibboso. Floret Majo et 
Junio. GUATE ALA, HUEHUETENANGO: C. & E. Seler 2935, 
Facaltenango, a1 juni 1896 (G, NY, US). ex quicus: Heyde 
Lux 2977, Nebaj, alt. 6000 p., maj. 1892 (G, NY, US). SAN 
MARCOS: Salas 368, “Culantro de monte’”’, San Cristobal Cucho, 
alt. 1920 m. , June, 1923 (US). CHIMALTENANGO: XS rs lan: ston 
1 an Desconsuelo, June 20, 1940 (F, Type). Fie. 

T. LANATUM Lecoyer, Bull. Soc. Bot. Belg. 9A: 499, "1885. 
Plante omnino pubescens nisi per foliolas supernas et in floribus. 
Pubescentia e pilis 1-4 mm. flexuosis acicularibus opacis, densis- 
simis in caule. Carpellorum tamen pubescentia e pilis capitatis 
translucidis sparsis. Radix fibrosa. Flores polygamo-monoicl, 


1944] Boivin,—American Thalictra and their Allies 413 


oe perfecti, alii — Sepala 3. 2-4.5 mm. Filamenta 3.5- 
m. Antherae 2.5-4.0 mm., acumine 0.5-0.7 mm. Sarvs 
= Tabi, vet 3. 5-8. Omm. Carpella sic sessilia gibbosa 

longa, ca. 1.8 mm. lata, nervis sinuosis selvalaieaae 
rare aes Fi fere recto vel Spit Floret Majo et Junio.— 
Mexico, PUEBLA: Purpus wee Cerro de Gentile, southwest 
Puebla, rocks, Aug., 1907 (UC). oaxaca: Purpus 27140, 
vicinity of San Luis Tultitlanapa, Las Naranjos, May, 1908 
Gh, Ie , US); Galeotti 4575, Oaxaca (syntype in the 

Paris Herbarium: F, pho wa i. this syntype) Conzatti 


Gonzalez 1208, Huauchilla, alt. 2500 m., June 1 (G); Conzatti 
ot fs Acmmioe a de San Felipe, oe alt. 1700 m., junio 9 
7 (F). Fie 


oe pubescence is longer than in any other species of Thalic- 
trum. The fruits of 7. lanatum, T. parvifructum and T. Pennellii 
are the smallest in this section of the genus. Other syntypes are 
Galeotti 4548 pro parte, from Sola, and Andrieux 1834, from 
Mitla. Nothing of those two has been seen by me. 


54. T. Pennellii, sp. nov. Planta verosimiliter submetralis, 
caule basi, foliolis superne et floribus exceptis, pubescens 
Pubescentia translucida minutissima e pilis capitatis per foliola- 
rum paginam inferiorem, alio translucida densa e pilis uniseriatis. 
Radices ignotae. Pedicelli directi centimetrales, attamen sub 
receptaculum recurvati. Carpella matura glabra, subsessilia 
refracta ovata, ca. 3 mm. longa, ca. 1.8 mm. lata, apice acuto- 
rostrata, nervis lateralibus ramosis et anastomosantibus, nervo 
dorsali convexo, ventrali convexiore quam dorsali et saepius 
gibboso quidem. Tempus florendi ignotum, item flores.— 
Mexico, purRANGO: Pennell 18,505, El Salto, Aserraderos, rocky, 
andesitic, pineland canyon, alt. 2500-2530 m., Aug. 31, 1934 
(ANS, Type). Fia. 54. 


Dedicated to Dr. Francis W. PENNELL of the Academy of 
Natural Sciences of Philadelphia. 


55. T. parvifructum, sp. nov. Planta 3-5 dm., pubescens 
saepius per foliolas inferne, nunquam superne, interdum aliis 
locis. Pubescentia pilorum crassiusculorum opacorum colorato- 
rum nonnunquam cum pilis capitatis translucidis intermixtorum 
Flores polygamo-monoici. Sepala elliptica ca. 3 mm. longa. 
Filamenta ca. 3 mm. Antherae 2.5-3.0 mm., acumine 0.2-0.4 
mm. Stylus una cum gthasiats: 3.0-5.5 mm. Carpella matura 
rostrata glabra obovata subsessilia 2.7-3.0 mm. longa, 1.6-1.8 

mm. lata, nervis lateralibus ramosis undulatis anastomosanti- 
busque, nervo dorsali convexo, ventrali convexiore quam dorsali 
et gibboso. Floret Julii mense—Mexico, sonora: Pennell 


414 Rhodora [NovEMBER 


19,556, ridge south of Arroyo Gochico, east of San Bernardo, 
base of cliff near summit, alt. 1050-1150 m., Aug. 5-9, 1935 
(ANS, type); Gentry 1463, Rio Mayo, Bakachaka, upper ’Sono- 
rian, riparian, inconspicuous along the stream bank, a tolerant 
herb, July 5, 1985 (ANS, F, G, M, NY, UC, US). Fie 

56. T. STRIGILLOSUM Hemsley, Diag. Pl. Nov. 1, nhs 
papillosum Rose, Contrib. U. 8. Herb. 5: 188, ig 
jalapense Rose, Contrib. U. 8. Nat. “Herb, 8: 28, 1003. "eae 
nonnunquam omnino pubescens 2-10 dm. Pubescentia varia 
variabilisque semper tamen densa inferne in foliolis. Pubes- 
centia caulinaris, si adest, vel e pilis capitatis, vel e pilis milli- 
metralibus flexuosis acieularibus opacis, vel iisdem intermixtis. 
Pubescentia foliolarum superne e pilis capitatis cum adest, 
inferne e pilis crassiusculis ib eibues coloratis densissimis, non- 
nunquam pilis capitatis intermixtis. Pubescentia carpellorum 
vel deest vel e pilis capitatis translucidis. Radices fibrosae. 
Pedicelli parum incurvati, sub fructum valde recurvati. F lores 
plurimi perfecti pauci masculi. Sepala 3.0-6.5 mm. longa. 
Filamenta 4.5-6.0 mm. Antherae 2.5-4.0 mm., acumine 0.4-0.5 
mm. Stylus una cum stigmate 5-13 mm. ‘Carpella matura 
3.5-4.5 mm. longa, stipite 0.3-1.0 mm., ventre 2.5-4.0 mm. 
longo, 1.6-2.2 mm. lato, nervis gibbosis reticulatis, nervo dorsali 
fere recto, vel parum convexo, saepius mwas co ssi hes 
Floret Majo, Junio, Julio et ‘Augu sto.—MEXICcO, COAHUIL 
Palmer, 9, Lerios, Feb. to Oct. 1880 (ANS, US); Gregg 404, near 
Pave San Antonio, Sept. 2, 1848 (M’ . SAN LUIS PorTost: 

5361, Minas de San Rafael, May, 1911 (F, M, NY, UC). 

steak ieee ‘Rose 2658, near Monte Escobede, ‘Aug. 27, 1897 
(US, type of T. papillosum; G, US, isotypes). VERA CRUZ: 
Rose & Hay 5674, Mount Orizaba, July 25 and 26, 1901 (NY, 
US); Rose & Hay 6188, near Jalapa, Aug. 17, 1901 (US, type of 
T. jalapense; G, US, isotypes). HIDALGO: Coulte ter 652, Zimapan 
(G); Purpus, Ixmiquilpan, Sierra de la Mesa, July, 1905 (UC); 
Clokey 1855, Real del Monte, bank of stream, Oct. 1, 1910 (M, 

C-UC); Pringle 6929, Sierra de Pachuca, alt. 9000 ft., July 28, 
1898 (ANS, F, G, M, NY, UC, US). mextco and. DISTRITO 
FEDERAL: Rose, Painter & Rose 8440, Hacienda de la Encarna- 
cion, July 7, 1905 (G, NY, US); Rose & Painter 7185, neat 
Cima, Sept.’ 19, 1903'(G, NY, US); Pringle 6422, valley_of 
Mexico, Pedrigal (lava beds), 1-3 ft., alt. 7800 ft. ‘Aug. 1896 
ANS, F, G, M, NY, UC, US); Schaffner 11, vallée de Oi ies 
1870-1880 (G, NY); "Mexia 2685, Monte de Rio Frio, kilometer 
49, road from Mexico City to Puebla, pine forest on slopes, 12 
shade of thick pines, suffrutescent, spreading, up to 1 m., flowers 
yellowish, alt. 4000 m. , July 31, 1929 (F, M, NY, UC); Bourgeau 
276, vallée de Mexico, juin, 1865-66 (G, US, syntypes of T- 
strigillosum). MicHoAcAN: Arséne 5677, vicinity of Morelia 
alt. 2800 m., 1910 (US). 


1944] Boivin,—American Thalictra and their Allies 415 


The type of this species has not been selected. Other syn- 
types, not seen, are Andrieux 546 and Schaffner’s collection from 
Tacubaya. The types of the two species given as synonyms 
have not been seen either, but isotypes of both were available 
for study. Fie. 56. 

Except as to the type of pubescence and the size of the fruit, 
there is not much difference between 7. strigilloswm Hemsl. and 
T. lanatum Lee. They have, however, different ranges. 


. T. laeteviride, sp. nov. Planta 3-12 dm., omnino pubes- 
Stork ‘nisi in foliolis superne et in floribus. Pubescentia e pilis 
capitatis translucidis interdum tamen intermixtis in_foliolis 
infernis cum pilis opacis gist a capitatis. Radices fibrosae. 
Pedunculi parum incurv fructum valde recurvatum. 
Sepala viridula ca. 3 mm. ae Antherae 2-3 mm., acumine 

.3-0.4 mm. Stylus una stigmate 6-10 mm. Carpella matura 
sessilia 5.0-5.5 mm. longa, 3.5-3.8 mm. lata, nervis ramosis et 
reticulatis, nervo dorsali convexo, ventrali gibboso convexiore 
quam dorsali, rostro ad frontem deflexo. Floret Julio et Augusto. 
—Mexico, coanvuiLa: Johnston & Muller 463, Sierra del Pino, 
Vicinity of La Noria, open valley with scrub oaks and scattered 
pines, shaded arroyo-banks, 1 1-4 ft. tall,, Aug. 20-26, 1940. (G, 
TYPE); Stanford, Retherford & Northeraft 387, lat. 25° 3’, long. 
101° 18’, in arroyo on south slope of mountain, 24 kilo. northwest 
of Fraile, great variation of vegetation with many vines an 

other types of more southerly on plant 18-24 inches tall, 

alt. 2900 m., July 15, 1941 (G, M); R. M. Stewart 2291, north of 
La Noria, high central parts ‘of the caleareous Sierra del Pino, 
north of high eastern ridges, frequent on — sunny slopes, be- 
coming 8 dm. tall, Aug. 26, 1942 (G). 

T. LASIOSTYLUM Presl, Rel. amis 3: "20, 1835. Planta 
40-65 cm. glabra nisi in foliolis inferne. Pubescentia sparsa e€ 
pilis crassiusculis coloratis brevibus. Radix ignota. Flores 
verosimiliter polygamo-monoici. Stylus una cum stigmate 5.0- 
6.5 ren . long. Carpella matura 5.0-6.5 mm. longa, atipile 0.5-1.0 
mm., ventre 4-5 mm. longo, 3.0-3.5 mm. lato, nervis lateralibus 
alincns reticulatisque, nervo dorsali fere recto vel convexo, 
ventrali gibboso et convexiore quam dorsali. Tempus florendi 
mihi ignotum est—Prru, trma: Macbride & Featherstone 269, 
Matucana, perennial among large rocks on steep northern 
canyon slope, about 8000 ft., Apr. 12-May 3, 1922 (F, ae 
Pennell 14,388, Canta, open rocky slope, herb, alt. 2700-3200 m 
June 11-19, 1925 (ANS, F, G). Fie. 58. 


Type, not seen, in the herbarium of the Botanical Garden at 
Prague. Lecoyer assumed this specimen to have been errone- 


416 Rhodora [NOVEMBER 


ously attributed to Peru, that it was a Mexican specimen of T. 
Hernandezii Tausch. Indeed he had no other collection of T. 
lasiostylum Pres] from Peru and both species are no doubt closely 
related. But as we have now two other collections it becomes 
clear that they are probably different enough to be kept specifi- 
cally segregated. 

59. T. SUBPUBESCENS Rose, Contrib. U. 8S. Nat. Herb. 8: 28, 
1903. Planta submetralis vel sesquimetralis per foliolas inferne 
pubescens, aliis locis nunc glabra tum pubescens. Pubescentia 
densa e pilis capitatis translucidis. Flores polygamo-monoici, 
plurimi perfecti, pauci masculi. Sepala 3.8-4.5 mm. longa. 
Filamenta 5-6 mm. Antherae 2.8-3.5 mm., acumine 0.2-0.4 
mm. Stylus una cum stigmate 7-10 mm. Carpella matura 4-5 
mm. long., 2.3-2.9 mm. lat., stpite 0.5 mm. vel breviore vel 

eente, nervis lateralibus ramosis reticulatisque, nervo dorsali 

fere recto vel parum convexo, ventrali gibboso convexiore quam 

orsali. Floret Julio et Augusto.—Mextico, mexico: Pringle 
1896, Sierra de las Cruces, ao Lecre 3-5 ft., alt. 10,000 ft., 
Aug. 14, 1896 (US, rypn; ANS, F, G, M, NY UC; US, ISOTYPES) ; 
Pringle ‘11 ,917, bluffs of barranca below Ozymba, alt. 8000 it., 
Sept. 24, 1904 ‘(F, G, US). Fie. 59. 


Type not seen, but there is an abundance of isotypes at hand. 


60. T. puBIGERUM Bentham, Pl. Hartw., 3: 285, 1857. T. 
tomentellum Robinson & Seaton, Proc. Am. Acad. 28: 103, 1893. 
Planta submetralis vel subbimetralis omnino pubescens. Pubes- 
centia e pilis uniseriatis translucidis densissimis. Radices fibrosae 
in locis paululum tuberosae. Folia caulinaria petiolis 2-8 cm.., 
dilatatione petiolari exclusa. Sepala 3.7-4.5 mm. longa. Fila- 

t ? Antherae 2.0-3.5 mm., acumine 0.5-0.7 mm. 
Stylus una cum stigmate 5-7 mm. Carpella matura pubescentia, 
3.5-5.0 mm. longa, 2.0-2.5 mm. lata, stipite 0-0.5 mm., nervis 
rugosis sinuosis reticulatisque, nervo dorsali parum convexo, 
ventrali convexiore quam dorsali. Floret Majo, Junio et Julio. 
—Mexico: T. C. & E. M. Frye 2640, -actareae Desierto de los 
Liones, alt. 3000 m., May 20, 1939 (M C-UC); Plunkett 26, 

esert of the Lions, along stream, Roh lL July 8, 1932 (F). 
JALISCO: eg 2 9679, slopes of ge hae of Guadalajara, 
alt. 5000 ft., July 16, 1902 (F, G, M, NY, US); Pringle 2479, 
hillsides near Guadalajara, June 29, 1889 (ANS, F, G, M, NY, 

C, US); Rose & Painter 7360, near Guadalajara, Sept. 28, 1903 
(US); Rose & H ough 4744, near eyo July 5 and 6 6, 1899 (G, 

NY, UC, Ui HIDALGO: Pringle 7489, valley near Tula, June 
24, 1897 (G). MIcHOACAN: Pringle 4143, low lands about Lake 
Patzcuaro, July 15, 1892 (G, type of T. tomentellum; ANS, F, M, 
NY, UC, US, isotypes). Fre. 60. 


q 
a 


1944] Boivin,j—American Thalictra and their Allies 417 


The tyPx of 7. pubigerum Benth. is Hartweg 1591, ad fluviorum 
margines prope Lagos. It has not been seen. 


61. T. Conzattii, sp. nov. Planta metralis omnino pubescens, 
nisi in carpellis. Pubescentia e pilis uniseriatis translucidis. 
Flores polygamo-monoici, plurimi perfecti, pauci masculi. Sepala 

8-3 } ta 3.5-4.0 mm. Antherae 2.0-3.3 
mm., acumine 0.3-0.7 mm. Stylus una cum stigmate 5.0-6.0 
mm. Carpella matura glabra 3.5-4.5 mm. longa, stzpite ca. 0.5 
mm., ventre 2.9-3.5 mm. longo, 1.8-2.2 mm. lato, nervo ventrali 
gibboso, lateralibus gibbosioribus et reticulatis, dorsali paululum 
concavo, vel subrecto. Floret Julio et Augusto.—MeExico: 
VERA cruz: Bourgeau 2726, région d’Orizaba, 17 juillet, 1865— 
1866 (G); G. L. Fisher 298, Orizaba, alt. 4500 m., Aug. 9, 1924 
(F, M, US); Miuiller, Orizaba, 1855 (NY); Mohr, Huatusco, July, 
1857 (US); Mohr, Orizaba, in pratis et graminosis, sylvaticis 
(ANS); Mohr, Orizaba, julio, 1857 (US); Mohr, Orizaba, in 
sylvis montanis ad saxas calcares, julio, 1856 (US). MExIco 
and DISTRITO FEDERAL: Hinton 4200, distr. Temascaltepec, 
Comunidad, alt. 2480 m., July 6, 1933 (M); Hinton 3851, Temas- 
caltepec, Comunidad, by the river, alt. 2480 m., Aug. 5, 1933 
(G, NY); Langman 2509, 4 mile s. of Contadero, thickly wooded 
slopes, July 27, 1940 (ANS). oaxaca: Conzatti & Gonzalez 314, 
Etla, Canada de San Gabriel, alt. 3000 m., Aug. 8, 1897 (G, 
TYPE; US, 1soryez). Fic. 61. 2 

62. T. sessilifolium, sp. nov. Planta metralis vel altior omnino 
pubescens. Pubescentia e pilis uniseriatis translucidis. Folia 
caulinaria sessilia vel fere sessilia in apice dilatationis petiolaris. 
Sepala 4.2-5.0 mm. Filamenta 6.0-7.5 mm. Antherae 

m., acumine 1.0-1.2 mm. Stylus una cum stigmate 5-12 mm. 
Carpella matura pubescentia, 5-6 mm. longa, ca. 2.5 mm. lata, 
stipite 0-1 mm., nervis lateralibus valde reticulatis, nervo ven- 
trali gibboso, dorsali suberecto vel concavo. Floret Majo, 
Junio, Julio et Augusto.—Mexico, mexico: Harshberger 76, 
Salaza, Sierra de las Cruces, alt. 11,000 ft., Aug. 13, 1896 (M, 
TyPE; ANS, G, NY, US, 1sorypEs). MICHOACAN: Leavenworth 
250, west of Tancitaro, on edge of arroyo, herb 5-7 ft., alt. 6660 
ft., July 15, 1940 (F, M, NY). puxsia: Nicolas, rancho Posa- 
das, Hacienda Alamos, 1 aoft, 1909 (F, NY, US); Arséne & 
Nicolas 5168, prés Puebla, Hacienda Noria, sur l’Atoyae, alt. 
2170 m., 20 mai, 1910 (G, M, NY). Fic. 62. | 

63. T. refractum, sp. nov. Planta glabra metralis vel sesqui- 
metralis. Radices fibrosae. Rami petiolorum paululum in- 
curvati valde divarieati, saepius refracti. Flores polygami- 
monoici, plurimi perfecti, pauci masculi. Sepala 2.5-4.5 mm. 
Filamenta ca. 5 mm. Antherae 1.8-3.0 mm., acumine 0.5-1.0 
mm. Stylus una cum stigmate 5-10 mm. Carpella matura 


418 Rhodora [NOVEMBER 


6.0-6.5 mm., sizpite 0.5-1.0 mm., ventre 4-5 mm. longo, 3.0-3.2 
mm. lato, nervis lateralibus sinuosis reticulatisque, nervo dorsali 
parum concavo, ventrali gibboso valde convexo. Floret a 
mense Junii usque ad Octobris—CoLOMBIA, CUNDINAMARCA: 
Cuatrecasas 9664, Cordilleria Oriental, vertiente oriental, cerca 
de Machetd, hierba de 1 metro, calis verde blanquecino con 
venas violdceas, antheras amarillas, alt. 2200 m., June 29, 1940 
(US, Fae : Pennell 2398, 2-3 m. sw. of Sibate, fl. ’purplish-green, 
alt. 2700-2800 m., Oct. 13-15, 1917 (G, NY, US). Fie. 63, a 
an 

64. T. PENINSULARE (Brandegee) Rose, Contrib. U. S. Nat. 
Herb. 8: 28, 1903. 7’. vesiculosum Lec., var. peninsulare Brande- 
gee, Zoe, 4: 399, 1894. Planta omnino glabra submetralis. 
Radices fibrosae. Foliolae membranaceae discolores, saepius 
trilobatae, basi cuneatae raro rotundae. Flores polygamo- 
monoici. Sepala 2.5-3.5 mm. long. Filamenta 3-6 mm. An- 
therae 3.0-3.5 mm., acumine 0.5-0.8 mm. Ovarii stipes 0.5-1.0 
mm, Stylus una cum stigmate 5-8 mm. Carpella matura gib- 
bosa 4-6 mm. longa, st7pite 1-3 mm., ventre 3-4 mm. longo, 1.8- 
2.0 mm. lato, nervis lateralibus gibbosis ramosis anastomosanti- 
busque, nervo dorsali basi concavo, superne convexo, nervo 
ventrali convexo et gibboso. Floret Septembris mense. An 
etiam Augusti?—Mexico, BAJA CALIFORNIA: Brandegee, Saltillo, 
Sept. 17, 1893 (UC, rypr and tsotyrr); Brandegee, El Taste, 
Sept. 13, 1893 (G, NY, US, paratypes); Brandegee, Sierra de 
San Francisquito, Oct. 18, 1890 (UC); ees egee, Sierra de 
Laguna, Jan. 23, 1890 (UC); s 27,027, Laguna 
Mountains, The a 7 Sept. 22, 1930 (G, MM, NY, UC, ae 


vel trimetralis, omnino glabra, plus minusve glauca. Radices 
fibrosae. Flores polygamo-monoici, plurimi perfecti, pauci 


masculi. et 3-5 mm. Jigs Filamenta 3.5-6.0 mm. pal . 


anastomosantibus, nervo dorsali “Sei concavo, superne convexo, 
nervo ventrali gibboso. Floret a Maji mense usque ad Novem- 


1944] Boivin,—American Thalictra and their Allies 419 


bris.—MExXIco, MEXICO and DISTRITO FEDERAL: G. L. Fisher 394, 
Amecameca, July 24, 1924 (F, M, US); Hinton et al. 6412, dist- 
Temascaltepec, Crucero, 1.5 m. high, Aug. 28, 1934 (F, G, NY, 
US); Hinton 5283, dist. Temascaltepec, La Labor, oak woods, 
alt. 2100 m., Nov. 30, ra; (M); O. Kuntze 23,655, ’ Amecameca, 
alt. 8600 ft.. , Aug., 1904 (NY). MORELOS: G. L. Fisher 35, 229, 
Tepoztlan, alt. 7500 ft., July 14, 1935 (F, NY, US) ; Lyonnet 584, 
Huizilac, Aug. 1930 (G, M, NY, US); Rose & Painter 6966, near 
Cuernavaca, Sept. 10, 1903 (NY, US); R ose & Rose 11,121, near 
El Parque, Aug. 15, 1906 (US); Pringle 11,338, near Cuernavaca, 
Gautepec, limestone hills, alt. 4000 ft., Oct. 27, 1902 (G, US): 
Pringle 11,918, Cuernavaca, barranca bluffs, 4-6 't., alt. 5000 ft., 
July 20, 1904 (F, G, US); Pringle 13,771 Cuernavaca, wet 
places, 4-6 ft., alt. 5000 ft., July, 1906 (G, US); Pringle 651 1, wet 
barranca above Cuernavaca, 6-10 ft., alt. 6500 ft., Sept. 21, 1896 
(US, type of T. stipitatum Rose; ANS, F, G, M, NY, UG, US, 
isotypes) ; Pringle 6392, near Cuernavaca, wet ’ shaded places, 
five to eight feet high, stem leaves four to six, alt. 5000 ft., July 
28, 1896 (US, type of 7. grandifolium Rose and T. grandiflorum 
Rose; ANS, F, G, M, NY, UC, US, isotypes). peurBia: Amable 
& Arséne 2022, dist. Tepeaca, Acatzinco, alt. 2110 m., julio 1907 
(US); Arséne 1446, prés Puebla, barranca de l’Alseseca, Hacienda 
de Batan, prés de Totimehuacan, alt. 2120 m., June 13, 1907 
(G, US); Arséne 2258, dist. Tepeacan, La Barranquilla, alt. 2110 
m., July, 1907 (M US): Pringle 9541, Amozoc, bluffs of wie 
alt. 7000 ft., Sept. 10, 1901 (US, type of T. obliquum; F, G, M 
isotypes). OAXACA: Galeotti 4549 pro parte, Juquila del Sur 
(Paris Museum, rypE of T. A Falta: F, photograph of the type) ; 
Pringle 5821, cuesta de San Juan del Estado, Aug. 20, 1894 (G); 
E. W. Nelson 1579, valley of Oaxaca, alt. 6500-7800 ft., Oct. 3, 
1894 (US). Fia . 65. 

The types of none of the species listed above have been seen. 
However, I have at hand a photograph of one of them, furnished 
by the Field Museum, and numerous isotypes of the others. 
The material listed under the name 7. gibbosum Lec., is fairly 
heterogeneous, but all attempts to segregate it along satisfactory 
lines have been unsuccessful. 


66. T. LoncistyLuM HBK. ex DC. Syst. 1: 171, 1817. Planta 
12 dm., glabra nisi per foliolas inferne. Pubescentia sparsa e 
pilis crassiusculis, brevibus, opacis. Caulis teres flexuosus nodis 
inflatis, sulcatus, validissimus tamen nec facile compressione 
pertritus. Sepala 3-5 mm. Filamenta 4-5 mm. Antherae 
2.5-4.0 mm., acumine ca. 0.4 mm. Carpella stipitata gibbosa 
dorso_ basi concavo, parte apicali convexo. Stylus una cum 
stigmate 8-10 mm. Carpella matura ignota. Floret Aprili, 


420 Rhodora [NovEMBER 


Majo, et verosimiliter Martio—Prru: Dombey (Paris Museum, 
TYPE; F', isoTyPe and a photograph of the type). Lima: Mac- 
bride & Featherstone 409, Matucana, in shrubby thicket on 
western slope, 4 ft. high, bracts rose-green, anthers yellowish, 
alt. about 8000 ft., Apr. 12-May 2, 1922 (F, NY) 


It is possible that the flowering material at hand might belong 
to some species already described from the fruiting stage, but the 
material is still too scanty to elucidate this point. 


67. T. ruTrpocarPpuM DC. Syst. 1: 172, 1817. Planta 8-13 
dm., glabra, foliolis inferne exceptis. Pubescentia sparsa e pilis 
crassiusculis brevibus opacis. Caulis sulcatus facillime pertritus. 
Flores polygamo-monoici, plurimi perfecti, pauci masculi. 

epala 4-5 mm. longa. Filamentaca.6mm. Antherae ca. 3 mm., 
acumine 0.3-0.4 mm. Stylus una cum stigmate 3.5-6.5 mm. 
Carpella matura 5.5-6.0 mm. longa, stipite ca. 2 mm., ventre sub- 
rhomboideo ca. 4 mm. longo, ca. 2.8 mm. lato, nervis lateralibus 
paucis ramosis et anastomosantibus, nervo dorsali ad basem et 
superne concavo ad medias paululum convexo, ventrali gibboso et 
convexo. Floret Januario et Februario—PrErvu: Dombey (Paris 

useum, TYPE; I’, photograph and fragment of type). APURIMAC: 
Stork & Horton 10,693, prov. Andahuaylas, Pincos, rainy-green 
shrubland, humus over sand, annual herb 1.3 m., monoecious, 
general color of fl. variable green to purple, fr.an immature follicle, 
alt. 2700 m., Feb. 19, 1939 (F). HuaNcavetica: Stork & Horton 
10,268, prov. Tayacaja, Saleabamba, part shade, among shrubs, 
annual herb reaching 1 m., monoecious, stamens greenish yellow, 
sinter follicles partly mature, alt. 3250 m., Jan. 7, 1939 (F). 

1G. 67. 


ovata. Antherae 3.0-3.5 mm., acumine ca. 0.5 mm. Frlamenta 
ca. 7mm. Stylus una cum stigmate 7-10 mm. Ovarii stipes 
0.5-1.5 mm. Carpella matura 7.0-7.5 mm. longa, stipite 1.0-2.5 
mm., ventre 4.5-5.3 mm. longo, ca. 3.3 mm. lato, nervis laterali- 
bus ramosis anastomosantibusque, nervo dorsali ad basem con- 
cavo ad medias et superne parum convexo, ventrali convexiore 
quam dorsali et gibboso. Floret a mense Martii usque ad Julii. 
—KEcuapor: Spruce 5470, in Andibus, 1857-1859 (NY, syntype); 
Jameson (US); André 2100, San Juan, Quindio, 8 martii, 1876 
(F, NY); Jameson 130, Quitensian Andes at 11,000 feet of eleva- 
tion, July, 1859 (NY). picnincna: Firmin 157, Cerra del 
Cinto, Ugshu-Cinto, Camino, alt. 3250 m., Jul. 28, 1927 (US); 
Jameson, Pichincha, alt. 11,000 feet of elevation (NY); Fzrmin 
680, borde del camino de La Magdalena al Cinto, pl. algo frecu- 


ee 


1944] Boivin,—American Thalictra and their Allies 421 


ente en los matorrales, me 3100 m. [?], Oct. 25, 1928 (US). 


BOLIVAR: Penland & Summers 589, Hacienda Talahua, 1 1 m. tall, 
alt. 330 m., May 1, 1939 (F. bea Panes and caXar: Hitchcock 
21,674, between Cuenca and H a, moist shady ravine, alt. 


2700-3000 m. , Sept. 11-13, 1903 (Ue), ee eee mers 1014, 
Tipococha, up to 1% meters, alt. 3200 m., July 11, 1939 (F, US). 
Perv, cuzco: Herrera 3070, Sceiake! ‘Valley, “Pinco”, 1931 
(US). Fia. 68. 


Other syntypes not seen are Mandon, Sorata; Jameson 212; 
Hieronymus, Sierra Chica, Rio primero; Goudot, Bogota; Jameson, 
Artisana. These syntypes might be heterogeneous for they in- 
clude a far greater range than the one we have accepted. There 
seems, however, to be some material of this species from Bolivia 
amongst Rusby 501-502, but we do not dare try to disentangle 
ge mixture. 


T. Nelsonii, sp. nov. Planta elata omnino pubescens. 
Puhciecata: e pilis capitatis translucidis nisi in foliolis inferne ubi 
pili capitati cum pilis brevibus opacis truncatis intermixti sunt. 
Flores polygamo-monoici, plurimi -perfecti, pauci masculi. 
ee ca. 5 mm. longa. Filamenta ca. 8 mm. Antherae ca. 3 

m., acumine ca. 0.5 mm. Stylus una cum stigmate 6-7 mm. 
Carpella matura conspicue gibbosa, 7-8 mm. longa, stipite 1.5— 

2.0 mm., ventre ca. 6 mm. longo, ca. 3.5 mm. snes nervis laterali- 
bus gibbosis reticulatis, nervo dorsali ad basem concavo, ad 
medias et superne convexo, ventrali convexiore et gibboso. 
Floret a Junio usque ad Octobrem.—MeExico, oaxaca: Nelson 
1788, near Rayes, alt. 6700—-10,000 ft., Oct. 20, 1894 (G, TYPE; 
US, ISOTYPE); Conzatti & Gomez 2424, dist. Cuycatlan, Cuyamec- 
alco, alt. 1800 m. , junio 23, 1909 (F, US). 

. 'T. popocarrum HBK. ex DC. Syst. 1: 71, 1817. Physo- 
carpum dei m (DC.) Bercht. & Presl, O Priroz. Rostl. 1: 
15, 1823. T'. cordifolium Willd. ex Lec. Bull. Soc. Bot. Belg. 24: 
262, 1885, nomen nudum ut synonymon. Planta circa sesqui- 
metralis omnino glabra. Flores polygamo-monoici, plurimi 
perfecti, pauci masculi. Sepala ovata 4-7 mm. longa. Fila- 
menta 3- ntherae 3.0-4.5 mm., acumine 0.3-1.0 mm. 
Ovarii stipes 0. 5-1.5 mm. Carpella matura caesia valde _— 
pressa apice acuta 9.0-10.5 mm. longa, stipite 2.5-3.0 m 
ventre 6.0-6.5 mm. longo, 2.8-3.3 mm. lato, nervis latarabibas 
reticulatis nullo modo gibbosis, nervo dorsali ad basem concavo 
ad medias superneque vel fere recto vel parum convexo, ventrali 
convexo saepius gibboso. Floret Junio, Septembri, Decembri 
et aliis—VENEZUELA: Cardona 299, Tachfra, Péramo de Tamé, 
alt. 3100-3300 m., July, 1939 (US); Gehriger 180, Mucuru 
quebrada vel pueblo, en las orillas y vegas de un riachuelo, cerca 


422 Rhodora [NOVEMBER 


de la casas, planta de 1.50 m., n. v. Colanillota, alt. 2500 m., 
ages 18, 19380 (ANS, F, M, NY, US); Linden 362, hautes Andes 
ruxillo et de M erida, ‘depuis 4000 jusqu’a a 14, 500 pieds de 

Pc 1842 (F). a ecaang Bonpland (Paris Museum, TYPE; 
F, photograph and a few leaflets from the type); Weddell 3808, 
prov. Tomina, dépt. Chuquisaca, déc., 1845-janv., 1846 
Triana, Andes de Bogota, Quindio, vulgo “lutantillo” [?], alt. 
2000-2700 m., 1851-1857 (NY); Pennell 10,332, dept. Caldas, 
Rio San Rafael below Cerro Tatama, clearing along stream, alt. 
2400-2600 m., Sept. 7-11, 1922 (ANS, G, US); hb. Parseval- 
Grandmaison, Mt. Serrate ‘pres Bogota, 1861 (G); Pennell 6898, 
dept. of El Cauca, above Papayan, along stream, ee flowers 
greenish-white, alt. 1750-1800 m., June 19, 1922 (AN 8, G, US); 
Killip & Smith 16,017, dept. Santander, vicinity of Ps Vegas, 
thicket, coarse herb, sepals pee anthers yellow, alt. 2600-3000 
m., Dec. 21-23, 1926 (G, NY, US). Ecuapor: Jameson 202, 
Quito (US). Fia. 70, a on b. 

Subsectio Venturiana, subsect. nov. Flores perfecti ('). 
Antherae apice truncatae. Ovaria 18-36 in flore. Stigma paululum 
clavatum. Carpella matura globosa nervis sinuosis gibbosisque. 


T. inuncans nostrum might, perhaps, just as well belong to this 
group, but until the fruits of that species are known, it is not 
possible to settle this question. Its ovary is perfectly smooth, 
a fact suggesting that the fruit may not be gibbose at all, but 
this relation does not always hold true. 


71. T. Venturii, sp. nov. Planta omnino glabra, 5-9 deci- 
metrorum. Radices fibrosae paucae, parum tuberosae. Folia 
estipellulata, 3-4-ternata petiolulis arcuantibus, stipellis reduc- 
tis. Foliolae tenues. Inflorescentia foliosa, pauciflora, panicula- 
ta nec apice racemosa. Flores lutei perfecti (!), 6-12 in planta. 
Sepala ovalia, nec satis nota. Stamina 12-20 in flore. Fula- 
menta capillacea subrigida, 5.5-7.0 mm. Antherae luteae, ob- 
longo-lineares, apice squarrosae vel retusae, 2.0-2.5 mm. Pistilla 
numerosa 18-36 in flore, stipitata, inflata, stigmate 2. 5-6.0 mm., 
parum clavato, apice obtuso, aliquandiu persistente. Carpe 
matura numerosa ventre globoso 2.0-2.5 mm., nervis gibbosis et: 
sinuosis, nervo dorsali fere tam convexo quam ventrali. Floret 
Decembri.—ARGENTINA, SALTA: Venturi 10,026, dept. ee 
Alemania, bajo bosque’ alto, flor amarilla, Dic. 15, 1929 (G, 
trPE; M, NY, US, ISOTYPE). SEvCMEN: Venturi 7718, gers 
Burroyaco, Cerro del Campo, en el alisal, flor amarilla, Dec. 15, 
1928 (F, G, US); Venturi 3995, fea j “dah Chicligasta, 
aaa. Las poy en los prados, flor amarilla, Dec. 11, ve 

CATAMARCA: Jérgensen 130 ars), dept. Andal 
Candada (US). Fia. 7A. a-c. uns zal, 


1944] Boivin,—American Thalictra and their Allies 423 


Booties Pelteria sect. nov. Planta glabra, vel pubescens, non- 
unquam glauca vel lucida vel scabra, radicibus fibrosis. Folia 
+6 tarot: foliolis peltatis. Flores omnes perfecti vel poly- 
gamo-monoici, cum alii perfecti alii meres sunt. Pedunculi 
sub fructu reflexi. F%lamenta filiformia purpurascentia. Sepala 
ile Beer ise vel plus minusve purpurascentia. Stigmata 
in flore, decidua. Carpella matura compressa 
seunciuilatoealin: seg vel stipitata. Species typica Tha- 
lictrum peltatum DC. s 

This section is eee to the mountaineous regions of 
Mexico and Central America. Other species with peltate leaflets 
also occur in Eastern Asia, but they belong to another subgenus. 

In most sections of the genus, the vegetative characters are 
highly variable and furnish but few characters to distinguish 
species. Contrariwise, in this section a most satisfactory key 
can be drawn up entirely on vegetative characters. As for the 
flowers, they offer but few instances of good characters permit- 
ting separation of the species from one another. The fruits 
offer much better characters but they are deceptive in that the 
immature fruit may be larger, longer, more compressed, more 
recurved and less stipitate than the mature one. 

Full many a species of this section has been described as polyga- 
mous or dioecious. With the help of a good binocular, thorough 
examination was undertaken of all the flowering material and it 
failed to reveal any pistillate or staminate plant. In many 
species all flowers are perfect, in others each plant bears both 
perfect and staminate flowers. Staminate flowers tend to bloom 
later than the perfect ones. 

Subsectio Subpeltata, subsect. nov. Planta glabra vel 
pubescens vel scabra. Folia 3-6-ternata, petiolata. Foliolae 
prope marginem insertae 3-7-lobatae, vel grosse 3-7-dentatae. 
Flores omnes hermaphroditi, vel plurimi gee ok et pauci 
masculi in bate 


a. Pauci pil 1 illimetrales vel longiores 
pilorum aie ares semimillim ngi rage 


T. Lanxestert Standley, Field Mus. Nat. Hist. Pub. Bot. 
18, as: 1937. Planta glabra nisi per foliolas, verosimiliter 
metralis. Pubescentia e pilis capitatis vel crassiusculis brevissi- 
mis. Radix fibrosa. Antherae ca. 2.5 mm., acumine 0.5-1.0 


424 Rhodora [NOVEMBER 


mm. Filamentum ca. 3.5mm. Stylus una cum stigmate 8-10 
mm. Carpella matura compressa, glabra vel pubescentia, ca. 6 
mm. longa, ventre 4.5-5.0 mm. longo, 2.2-2.5 mm. lato, stipite 
0.5-1.0 mm., nervis lateralibus valde anastomosantibus nervo 
dorsali fere recto. Floret Junio.— Costa Rica: Brenes 14,506, 
prés San Ramon, pentes rocheuses du cerro de San Isidro, alt. 
1300 m., 14 junio, 1901 (G, US); Solis 297, Icsasu, cerro de 
Molti, alt. 1800 m., Aug., 1935 (F, M); Tonduz 8951, La Verbena, 
Aug., 1894 (US); C. H. Lankester 337, salen rocky slopes, 
July, 1919 (F, Type and isotype). Fia. 72, 

73. T. Torresii Standley & Boivin, sp. no os Planta metralis 
omnino ‘pubescens, nisi in ecarpellis cum pubescentia est e pili is 
capitatis. Folia caulinaria sessilia in apice dilatationis petio- 
laris. Sepala elliptica 2.5-3.0 mm. longa. Antherae ca. 3.5 mm., 
acumine millimetrali. Filamenta ca. 5.5 mm. Stylus una cum 
stigmate 5.0-7.5 mm. Carpella matura compressa sessilia ca. 

.5 mm. longa, ca. 2.0 mm. lata, nervis fere simplicibus, nervo 
dorsali concavo.—Costa Rica: Torres 187, Santa Lucia, Paraiso, 
terreno pedregoso, altura planta 1 m., tiene en las hojas una sus- 
tancia pegajosa, 28 julio, 1927 (US, TYPE); iene 1632, asaiige, 
alt. 1700 m., July 1, 1937 (F). Fic. 73 

74. T. GUATEMALENSE GC. DC. & re Contrib. U. S. Nat. 
Herb. 5: 88, 1899. T. peltatum, var. 8 hirsutum Loes., Bull. 
Herb. Boiss. 2° série, 3: 89, 1903. T. hondurense Standley ex 
Yuncker, Field Mus. Nat. Hist., Bot. 17: 362, 1938. Planta 
omnino pubescens, plus minusve scabra, 8-12--(20) dm. Pubes- 
centia e pilis aliis capitatis crassiusculis truncatis aliis elongatis 
flexuosis. Radix fibrosa. Folia caulinaria saepius 4-ternata 
petiolo brevi, vel sessilia in apice dilatationis petiolaris. Pedicelli 
plus minusve recurvati. Sepala elliptica 3.0-3.5 mm. longa. 
Antherae 4-5 mm., acumine 0.5-1.0mm. Filamenta 3.5—4.0 mm. 
Stylus una cum stigmate 6-9 mm. Fructus compressus 3.5-5.5 
mm. longus, ventre 3.0-4.5 mm. longo, 1.5-2.0 mm. lato, stipite 
0-0.5 mm., nervis parum ramosis reticulatisque, nervo ’ dorsali 
fere recto vel concavo. Floret Junio vel Julio vel Augusto.— 
Mexico, Mexico: Hinton et al. 4263, Temascaltepec, Tejupilco, 
llano, stalks 1 m., July 31, 1935 (M); Hinton 4530, Temascalte- 
pec, Laver in sears water, 2m. high, — 15, 1933 a: at 


(G, M, NY, US); Leavenworth & Hoogstraal 1282, Tancitaro 
region, two miles south of Uruapan, rocky la ry hirsu 
almost prickly, very common, alt. 5577 ft. a 29, 941 (F, ee 
Y). cHrapas: Matuda 4889, near Siltepec, Honduras 
encinal, alt. 1500 m., July 9, 1941 (M); Purpus 6798, Cerro del 


1944] Boivin,—American Thalictra and their Allies 425 


Boqueron, Sept., 1913 (UC); HE. W. Nelson 3236e, near San 
Cristobal, alt. 7000-8000 ft. , Sept. 18, 1895 (US). GUATEMALA: 
ayes, Las Vacas barranca, June 1860 (G, US). HUEHUETE- 
NANGO: C. & EL, Seler 3158, unter Gebiisch auf dem Coes von 
Zac-ulen bei Huehuetenango, 17 juni, 1896 (G, NY, US, iso- 
types of 7. peltatum var. 8 hirsutum); C. & E. Seler 2857, Checulé, 
auf den mit Busch u. Wald bewachsenen Steinfalzungen alter 
Bauten, 28 aug., 1896 (G, NY, US); Standley 81,212, Se emg 
west of Aguacatdn on the road to "Huehuetenango, oa 
herb 1 m. tall, alt. about 1950 m., Dec. 27, 1940 (F); Standen 
81,566, near Puente de Xinax6, along road 13 km. west of geal 
tenango, pine-oak forest, herb 1 m. tall or less, commo out 
of flower, alt. about 1800 m., Dec. 30, 1940 (F): Standley 81 8. 
along Aguacatdn road east of Huehuetenango at km. 13-14, 
damp forested quebrada, common, alt. about 1950 m., "Jan. 2, 
1941 (F); Standley 83,010, east of San Rafael Pitzal, near crossing 
of Rio San Juan Ixtén, — oak forest, herb 1 m. tall, common, 
alt. about 1730 m., Jan. 9, 1941 (F). Basa VERAPAZ: Cook & 
Doyle 254, Santa Rosa, May 28, 1904 (US); von Tiirckherm 1313, 
Santa Rosa, im Walde, alt. 5000 p., Juli, 1887 (F, G, US); von 
Tiirckheim 2307, ae ae Rosa und Patal, Wald, alt. 1600 
m., Juli, 1908 (F, G, NY, US). zapaca: Steyermark 29,676, 
Sierra de las Minas, dialed Rio Hondo and Finca Alejandria, 
moist forested slopes, alt. 1700-2000 m., Oct. 11, 1939 (F); 
Steyermark 29,654, along trail between Rio Hondo and summit 
of mountain at Finca Alej tas Lyrae slopes, rocky 
places, alt. 1000-1500 m., Oct. 11, 9 (F). CHIMALTENANGO: 
Standley 57,897, along road from Silene to San Martin, 
ies thicket, common but past fruit, herb 3-4 ft., alt. 1500-1900 
Nov. 25, 1938 (F); Standley oi, ot eode m, damp forest, 
heck 3 ft., alt. 1500-1900 m., Nov. 2 1938 (F); Standley 61,561, 
southeast of Patztiim, barranco de te Sierra, pine and oak orest, 
herb 3-4 ft., common, alt. about 2100 m., Dec. 31, 1938 (F); 
Standley 80, 039, southeast of Chimaltenango, along Rio Guaca- 
late, oak forest, common but past flowers, alt. about 1700 m., 
Dec. 14-23, 1940 (F). SACATEPEQUEZ: Gomez 794, Santiago, 
alt. 6500 p., 1891 (G, US, paratypes of T’. guatemalense) ; Stand 
58,980, cuesta de la Canas, above Antigua, damp thicket, nd 
4 ft., alt. about 1950 m., Dec. 6, 1938 (F); Standley 80,994, ‘along 
Rio ‘Guacalate, on road between Antegua and Chimaltenango, 
alt. about 1660 m., Dec. 23, 1940 (F). GuaTEMALA: Deam 6096, 
on top of mountain, growing among rocks, June 1, 1909 (G, US); 
Tonduz 657, eo de Se age dans les broussailles des collines, 
juin 1921 (G : US); Aguilar 249, 1939 (F). SANTA er 
Heyde & Lux sae Cenaguilla, alt. 4000 p., Sept., 1892 (G, M 
NY, US). savapa: Steyermark $2,641, Cerro Alcoba, just e east 
of Jalapa, oak woods, alt. 1300-1700 m., Dec. 2, 1939 (F); 


426 Rhodora [NoVEMBER 


Standley 76,810, 76,828, on hills northeast of Jalapa, rocky scrub- 
oak forest, herb 1 m. tall, alt. = se bt m., Nov. 10, 1940 (F). 
Honpuras, comayacua: Yu ncker, Dawson & House 5796, near 
Siguatepeque, on rocky hillside, otanes odut 4 ft. tall, more or 
less viscid and with pungent odor, roots a rich yellow color, June 
. 5p Pee 1936 (F, type; G, M, NY, US, isotypes of 7. hondurense). 

IG a-d 

The type of T. guatemalense C. DC. & Rose (Heyde 164) was 
stored away and could not be seen. 

Subsectio Eupeltata subsect. nov. Planta glabra vel pubes- 
cens e pilis multiseriatis translucidis, vel crassiusculis opacis 
truncatis, nonnunquam glauca vel valde lucida. Foliolae 
ampliores rotundae vel obovatae, petiolis circa tertias insertis. 
Flores polygamo-monoici, plurimi perfecti, pauci masculi, nisi 
Thalictro Treleasit nostro cujus omnes perfecti sunt. Species 
typica Thalictrum peltatum DC. sit. 


Whether the leaves are dull, shining or glaucous can not be 
relied upon as a specific character in this section, if we judge by 
what occurs in 7. Pringlei Wats. None of the other species has 
been collected at all stages throughout the year, but they are 
likely to follow a course of development parallel, if not similar, 
to that of 7. Pringlei. 


a. boli om ra, petiolate, petiolo ba o modo scabra....b. 
ee ee a tato....c. 


a Foliise ‘ieimbranaceae, marginem fere totam 


, crenis ca. 20 i = folio Beet pie vets 75. TT. Treleasit. 
d. oxigen coriacose, ad apicem grosse dentatae vel 
6s 


oe rT. Prine, hou reticulatum. 


b. F olia omnia, nisi inflorescentiae redu: sessilia in 
iat tationis 9 i Fo ie cu 
yi — 9 Be oliis caulinaribus et in inflores- 


Bee orek se he eS hoe EE 79. T. Roseanum. 


eee orci: Shep Os ye tise bees os 80. 7’. Arsenii 
g: pe carpella matura cae _ 
ge Ae ee ae > ee eee 81 jaliscanum. 
a. Planta scabra pubescens, nullo modo glauca... .h. : 
. a faciem eae § Sees Cee. 


hia i A Xi act aie echoes ee . 
er nt eae PSPS ee as Cee i. SP ee Awe Parone BE 
7 i pas si Ba oie a alia ‘i 


ae 


Seat Sey 


goa tro rai nisi 


1944] Boivin,—American Thalictra and their Allies 427 


75. T. Treleasii, sp. nov. Planta glabra circiter 6 dm. alta. 
Folia petiolata. Foliolae membranaceae per marginem fere 
totam crenatae, crenis 20 admodum. Sepalaca.4mm. Stam- 
ina 20-30 in floribus singulis. Filamenta 5-6 mm. Antherae 
4.0-5.5 mm. Acumen antherae 0.5-0.7 mm. Ovaria 3-6 in 
flore: Stylus una cum stigmate 6-9 mm. Fructum auctor non 
cognoscit, consimilem Thalictro Pringlet er esse videtur. 
Floret Junio.—Mexico, Gurrrero: Langlassé 1061, Sierra 
Madre, alt. 1000 m., juin 16, 1899 (G, Kat US ISOTYPE). 
Fie. 75, a and b 


Dedicated to Prof. W1LLIAM TRELEASE who in 1886 published 
a monographic paper on the genus as represented in America 
north of Mexico. 


76. T. pettatum DC. Prod. 1: 11, 1824, nec sensu J. N. Rose, 
Contrib. U. 8. Nat. Herb. 5: 186, 1899; nec T. peltatum Sessé & 
Moc., Fl. Mex. 134, 1894. Planta fere metralis, parum si vero 
glauca. Foliolae plus minusve coriaceae basi rotundae, ad 
apicem grosse dentatae vel crenatae, crenis vel ponrerne ® 3-10 in 
foliola. Sepala 3.3-4.5 mm. longa. Filamenta 5-7 m An- 
therae 1.9-4.0 mm., acumina 2.0-6.5 mm, subaequantes. 5 Onde 
0-5 in flore, stylo una cum stigmate 8-13 mm. Carpella matura 
ignota, verosimiliter (ex immaturis) ca. 5.0 mm. longa, 2.0 mm 
lata, semi-obovata, parum si vero stipitata, ista Thalictri Pringlet 
Wats. approximantia. Floret Augusto.— Mexico, Mexico: Hin- 

n 4547, distr. of Temascaltepec, Nanchititla, oak woods, Aug. 
18, 1933 (M, US); Alaman, [probably near Mexico city], 1811 
[or 1821?!] (vyPEi in Delessert Herbarium; F, photograph of same). 
MICHOACAN: Hinton 15,071, distr. Barroloso, Coaleoman, woods, 
alt. 13800 m., Aug. 7, 1939 (G). Fic. 76, a-c. 

DeCandolle’s original description applies well to all species 
of subsect. Eupeltata. In 1899, J. N. Rose (Contrib. U. S. Nat. 
Herb. 5: 186) with the help of C. De Candolle solved the problem 
by comparing specimens from five different species with the 
actual type. Since 7’. peltatum DC. had been collected but once 
until as late as 1933, it is not surprising that the name was 
applied to its closest related species, for which we are now pro- 
posing the name 7’. Roseanum. But as we now have a very good 
photograph at hand it becomes possible even to take measure- 
ments of the actual type, e. g.: sepal 3.3 mm., acumen 2.3 mm., 
anther 1.9 mm., filament 5.1 mm., stigma 10.5-13.0 mm., fruit 
5.0 x 2.1 mm., ete. 


1 Notes of the author. 


428 Rhodora [NovEMBER 


7. T. Prineper Wats. Proc. Am. Acad. 25: 141, 1890. 
Planta glabra, saepius sub florendi tempus caeca, deinde : saepius 
glauca cum fructus maturant, sub maturandi saepius lucida, 
posteaque lamina lucida exfoliatur, et planta pallida caeca 
revelatur. Radix fibrosa. Folia petiolata 2-4-ternata. Folli- 
olae margine vel integra vel sinuata vel crenata vel grosse 
dentata, dentibus vel crenis 0-7 in foliola. Sepala 3.0-4.5 mm. 
longa. “Stamina 30-50 in flore. Filamenta 3.0-3.5 mm. An- 
therae 3-4 mm., acumine 0.5-0.8 mm. Stylus una cum stigmate 

9 mm. Carpella matura subsessilia semi-ovata, nervis sim- 
plicibus vel ramosis et anastomosantibus, nervo dorsali fere 
recto vel parum concavo. Floret Junio Julioque.—MExIco, 
nayariT: M. E. Jones 22,843, Tepic, Feb. 15, 1927 (F). MExico: 
M. E. Jones 85, La Palma. June 9, 1892 (UC, US); Sarton, 
Tolucea Mts., 1852 (US). micHoacan: Leavenworth & Hoog- 
straal 1823,Tancitaro region, Apatzingan, common in open pine 
forest above Acahuato, sometimes in pure stands, alt. 3500—4000 
ft., Aug. 23, 1941 (F); Leavenworth 395, Tancitaro on the road 
from Tancitaro to Apatzingan, here 9/4, common on rocky 

anks in Boo woods from 4000-6000 ft., alt. 5000 ft., July 30, 
1940 (F, M, NY); Leavenworth & H oogstraal 989, Tancitaro, west 
of Santa Clara, erry bank near stream, open pine forest, alt. 

ft., July 11, 1(F, M). JALIsco: Pringle 2478, slopes of 
the barranca near Gasielios June 29, 1889 (G, TYPE; ANS, 


Guadalajara, in one a July "9, 1899 (US); Reke 4581, 
Guadalajara, Barranca Ibarra, alt. 1400 m., Nov., 1922 (US); 
Mexia 1412a, Sierra Madre Mtns., San Sebastian, trail to 

ascota, damp ravine in open pine forest, prominent on forest 
Hoos, alt. 1425 m., Jan. 7, 1927 (F, G, M, NY, UC). Fig. 77, 
a 


78. T. Private: Wats., var. RETICULATUM Rose, Contrib. U. 
S. Nat. Herb. 5: 188, 1899. Planta 35-40 cm., pubescens nisi 
in pagina superna foliolarum. Foliolae vel crenatae, vel repan- 
dae, vel integrae, per ambas paginas rugosae nervis conspicue 
reticulatis—Mexico, NAYARIT: Rose 1985, Territorio de Tepic, 
between Pedro Paulo and San Blascito, Aug. 6, 1897 (US, TYPE; 
G, 1soTyPE); Rose 1985a, Territorio de Tepic, ‘foothills between 
Pedro Paulo & San Blascito, Aug. 4, 1897 (US); Rose ile 
Territorio de Tepic, between Dolores & Santa Gertrudis, Aug. 7 
1897 (US). 

The type of this variety was stored away and hence it has not 
been seen. With better flowering material and fully mature 
fruits, this variety might prove specifically different from the 
typical form. 


1944] Boivin,—American Thalictra and their Allies 429 


74b 
() 75 
yA\\) 
75 


: 76a 76b76c 77> 87 86 89a 89b89c 
Deratts oF FLowers or Fruits oF THatictrum, all X 4. 
(For explanation see end of paper.) 


430 Rhodora [NovVEMBER 


79. T. Roseanum, sp. nov. T. peltatum Sessé & Moc. FI. 
Mex., 2: 134, 1894 nec 7. peltatum DC. Prod. 1: 11, 1824. 
Planta glabra, saepius conspicue glauca, metralis vel altior. 
Folia biternata, sessilia in apice dilatationis stipularis. Foliolae 
majores orbiculares, ad apicem crenatae vel grosse dentatae. 
Sepala 5.0-5.5 mm. longa. Antherae 2.2-3.2 mm., acumine 
1.0- mm. Filamenta 4-5 mm. Stylus una cum stigmate 
6.5-9.0 mm. Carpella matura compressa 5.6-6.5 mm. longa, 
stipite 0.5-1.5 mm., ventro 4.2-4.5 mm. longo, 2.2—-2.6 mm. lato, 
nervis honnunquam parum ramosis anastomosantibusque, nervo 
dorsali convexo, ventrali convexiore quam dorsali. Floret 
Julio mense. An etiam Junioque?—Merxico, MORELOS: Pringle 
7448, bluffs of barranca above Cuernavaca, alt. 6500 ft., Aug. 23, 
1897 (G, typy; M, US, tsotypses); Pringle 8261, blufis of moun- 
tain canyon above Cuernavaca, alt. 6500 ft., Sept. 30, 1899 
(ANS, F, G, M, NY, UC, US); Pringle 7205, bluffs of barranca 
rae Cuernavaca, alt. 6000 ft., Jul. 28, 1896 (G, US). Fia. 79, 


a-d. 

Dedicated to the late JoserpH Netson Rose who laid the 
foundations for our understanding of the Mexican species of 
Thalictrum. 

T. peltatum Sessé & Moc. is a heterogeneous species as can be 
easily seen by what is preserved of it at the Field Museum. How- 
ever the description seems mainly based on 7. Roseanum. The 
fragments can not be identified with certainty. 


prise d’eau, alt. 2000 m., 19 sept., 1909 (US); Arséne 5470 
vicinity of Morelia, Rincon, alt. 2000 m., 14 aa 1910 (G, 
TrPE; M, NY, US, 1sorypxs); Kenoyer A-482, Morelia, Aug. 17, 
ah rant pe tig - (F ij Be ise 8, Morelia, talus, 8 a 

» Arsene 13, Morelia, Rincd . 2000 m., 14 aodt 
1909 (F). Fig. 80, a-c. oe 556i 


1944] Boivin,—American Thalictra and their Allies 431 


Dedicated to Brother G. ArsiNE, an enthusiastic field-botan- 
ist. His Mexican collections are both abundant and well pre- 
served and form a high percentage of the herbarium material at 
hand for study. 


81. T. satiscanum Rose, Contrib. U. 8. Nat. Herb. 5: 187, 
1899. Planta glabra saepius valde glaucescentia, sesquimetralis 
usque bimetralis. Folia 2-6-ternata. Fiololae orbiculares vel 
obovales, ad apicem crenatae vel grosse dentatae. Sepala 3 

.5 mm. longa. Stamina ca. 30 in flore. Antherae 2.2-2.5 mm., 
acumine 1.2-1.7 mm. Carpella matura compressa, 5.5—7.5 mm. 
longa, ventre 5.0—-5.5 mm. longo, 2.0-2.5 mm. lato, stipite 0—-1.5 
mm. longo, nervis ramosis anastomosantibusque, nervo dorsali 

convexo, ventrali convexiore quam dorsali.—MrExiIco, MEXICO: 
Hinton 42 59, Temascaltepec, Telpintla, alt. 1840 m., July 28, 
1933 (G, NY); Hinton et al. 6557, Temascaltepec, Rincon del 
Carmen, edge of oak woods, 1.5 m. high, Sept. 6, 1934 (G, NY, 

US); Hinton 1078, Temascaltepec, Volcan, rolling prairie, 2 m. 
high, alt. 1410 m., July 18, 1932 (F); Hinton 6740, Temasealte- 
pec, Platanal, hill, 1.5 m. high, Oct. 11, 19384 (N Y). JALISCO: 
Rose & Hou gh 4786, near Tequila, July 5 and 6, 1899 (US); 
Goldsmith 28, pes of =e Colima, wooded hills, "alt. 5000 ft., 
July 13, 2 905 (G); Rose & Painter 7519, near Etzatlan, Oct. 2, 
1903 (NY, US); Rose ‘& Painter 7648, near Chapala, Oct. 5, 1903 
Lislh Diquet (NY): Mr. & Mrs. J. G. Lemmon 157, lake Chapala, 

5 (F, G, UC). Fie. 81, a-c. 


The type (Rose 2840) has not been seen but the original 
description is clear enough. One label states that the plant is 
3.4 m. high. Such a height is not impossible, but the way the 
number presents itself suggests the possibility of an error. 


82. T. currNnavacaNnuM Rose, Contrib. U. 8. Nat. Herb. 5: 
187, 1899. Planta omnino scabra, 7-12 decimetrorum, nullo 
modo glauca. Folia 2-3-ternata, petiolate, dilatatione petiolari 
reducta. Foliolae orbiculares, per marginem fe fere totam crenatae. 
Stamina ca. 30 in flore. Antherae 2.5-5.0 mm., acumine 0. 07-1. 2 
mm. Filamenta 4.0-5.5mm,. Stylus una cum stigmate 4-9 mm 
Carpella paaucbe compressa 3-5 mm. longa, 1.5-2.8 mm. lata, 


stipite 0-0.5 mm., nervis simplicibus vel parum ramosis an 
mosantibusque, nervo sc vse recto vel paululum convexo. 
oret Junio et Julio—Merxico, MorELOS: Rose 6860, near 


Cuernavaca, Sept. 8, 1903 NY, US); Pringle 7238, near Cuer- 
navaca, ea, rocky bluffs of barranca, alt. 5000 00 ft., July 26, 1896 
( F, G, M, sorypss); Pringle 6878, above Cuernavaca, 
lava fields, alt. 7000 ft., yt 27, 1898 (ANS, F, G, M, NY, UC, 
US, paratypes). Fie. 82, a-d 


432 Rhodora [NOVEMBER 


Type not seen, but there is an abundance of isotypes and 
paratypes at hand. 


83. T. cumRNavacaNuM Rose, var. supraglabrescens, var. 
nov., Morton in litt. Foliolis supra glabris nec scabris.—MEx- 
ICO, GUERRERO: Rusby 153, Limon Mtn., alt. 4000 ft., July 28, 
1910 (US, type and 1sotyPrE; NY, ISOTYPES) ; ; Lyonnet 304, Taxco, 
julio, 1929 (US); Kenoyer A-481, Taxco, July 20, 1938 (F). 

Sectio Heterogamia, sect. nov. Sect. Thalictrum, 1. Hetero- 
gama DC., Syst., 1: 172, 1817. Plantae glabrae vel pubescentes. 
Pubescentia, cum adest, pilorum capitatorum translucidorum. 
Flores dioici. Sepala dimorpha, floris masculi majora saepius 
colorata, foeminei minora viridia vel raro purpurascentia. Fila- 
menta imprimis lutea deinde nonnunquam purpurascentia, nec 
clavata sed filiformia et sub antheris paullulum dilatata. Stigma 
plus minusve bialatum. Species typica Thalictrum dioicum Li, 
sit. 


Group confined to Mexico, United States, Canada and the 
French Islands of St.-Pierre et Miquelon. 


Subsectio Debilia, subsect. nov. Plantae glabrae, foliis 
ternatisectis, radicibus tuberosis fusiformibus haud stoloniferae. ° 
Caulis basis decumbens. Folia inferiora anges nisi reducta 
velis vaginis, intermedia plurima conferta, superiora pauca 
sparsa, inflorescentiae minora. Stipellulae’ desunt. Inflores- 
centia elongata pauciflora paniculata. Carpella matura subses- 
rs recta haud compressa, ventre symetrico pariete membrana- 

eo, nervis simplicibus convexis nec sinuosis. Species typica 
Thalictrum debile Buckley sit. 


a. Carpella matura ovoidea; stigma 0.5-1.0 mm. Planta 
ah radicibus ng pie Se ge es Oia Ok races 84. T. teranum. 

a. sep oe -0 mm. Plantae decumbentes radicibus brun- 
4 Patil matura elliptico-lanceolata, 0.7-1.2 mm. lata....85. T'. debile. 
b. Carpella matura ellipsoidea, 1.5-2.0 mm, lata..... 86. T. arkansanum. 


84. T. rexanum (Gray) Small, Fl. Se. U. S. 446, 1903. T. 
debile Buckley, var. texanum Gray ex E. Hall, Pl. Tex. 3, 1873, 
nomen nudum; Gray, Syn. Fl. N. Am. 1: 18, 1895. Planta 
rigida 10-45 cm. Radices nigricantes, cum siccatae, haud 

tae irregulares. Sepala floris maris 1.7-3.0 mm., 
oe 0.7-1.5 mm. longa. Filamenta ca. 1.5 mm. Antherae 

2.0 mm. Stigma 0.5-1.0 mm. Carpella matura ovoidea, 
acpi 0.1-0.3 mm., ventre 2.7-3.7 mm. longo, 1.4-1.6 mm. lato, 
apice acuto, basi rotundo, nervis 6-8 in fructu. Verosimiliter 
floret Februario et Martio.—Texas: Wright 170, Fayette Co. 
Cumming’s Creek, May 10, 1849 (ANS, G); Hall, Houston, 
April, 1872 (F, paratype): Hall 3, Houston, moist prairies, 


aba Sh =e CE ST tag ah eh a Ee ERs 


1944] Boivin,—American Thalictra and their Allies 433 


March 28, 1872 (G, type; ANS, F, M, NY, US, 1sorypss). 
Fig. 84, a-f. 
DEBILE Buckley, Amer. Journ. Sci. 45: 175, 1848. 
Plania innitens, 10-40 cm. Radices brunneae, cum siccatae, 
costatae. Sepala floris maris obovata, ca. 1.7-2.0 mm., foeminei 
saepius nc nee ca. 1.00 mm. Filamenta ca. 2.0 mm oe 
therae 1.7—2.0 mm., acumine 0.1-0.3 mm. Stigmata 1.3-2. Om 
Carpella matura  oblongo-lanceelata, stipite ca. 0.1 mm., i 
per ambas apices rotundo 3.0-3.7 mm. longo, 0.7—1.2 mm. lato, 
nervis 8-10 in fructu. Floret Aprili —GrorGIA: Chapman, 
conc 1882 (F, G, US); Chapman, Mts. of Georgia (M, US). 
wr Nieuw wland, Blount t Co., Warnock Peak, April 12, 
1930 (N-ND); Buckley, Wilcox Co., woods, rich alluvial soil, 
ees April, 1840 (NY, US, paratypes) ; ; Buckley (ANS, 
G, paratypes): Buckley, April (M, eee uckley, Wilcox 
Co., near Allenton, rich woods, April, ps (M, TYPE); Buckley, 
April, 1841 (M, NY, paratypes). Pre. 8 
86. T. arkansanum, sp. nov. Pianta erecta 20-40 cm. 
Radices brunneae nonnunquam, cum siccatae, costatae. Sepala 
floris maris ovata vel elliptica 2. a 0 mm. longa, foeminei ovata 
1.0-1.5 mm. longa. Filamenta 2.0-3.0 mm. Antherae 1.8-2.3 
mm., acumine 0.1-0.4 mm. Stigma (1.5)-2.3-3.0 mm. Car- 
pella matura sessilia ventro ellipsoideo 3.5-4.5 mm. longo, 1.5- 
2.0 mm. lato, nervis 10-12 in fructu. Floret Aprili et Majo.— 
Arkansas: Canby, Sargent, Trelease & Bush, Fulton, rich woods, 
Apr. 18, 1901 (ANS); Bush 2445, Fulton, low ground, Apr. 26, 
1905 (M, TYPE); E. J. Palmer ies Hempstead Co., Fulton’ 
upland open woods, Apr. 8, _ ie ig og E. J. Palmer 7161, 
Fulton, upland woods Apr. 5 : 5 (NY . US); > fe $4 Kellogg, 
Arkansas Post, March 14, 1909 aD: Bush 51 1, Fulton, common 
in low ground, Apr. 4, 1900 (G, M, US); Nuitall (ANS); Bush 
2271, Toxetane:. in "rich woods, Apr. 8, 1905 (M). Fie. 86. 
Subsectio Sin sa, subsect. nov. Plantae glabrae, raro 
paululum pabendehien pilis capitatis. Radices tuberosae. 7a 
sparsa foliolis parvis. Stigma sagittatum. Carpe ella matura 
recta basi rotunda, apice acuta, nervis simplicibus sinuosissimis 
et crassiusculis, nec costata. Species typica Thalictrum pin- 
natum Wats. sit. 


a. Folia ternata vel raro subpinnata.. .. - 
b. Caulis basis procumbens aphyllus, folia ternats... 65.05% 87. TT. pudicum. 
b. Caulis rigidus erectus, ce oa har eis vel subpinnata, 
licet folia ramorum ternata. . .. 
c. Caulis a basi ramosus, auntie iciiosiotlinin, foliis 2-ternatis 
. T. madrense. 
ce. Caulis foliis caulinariis eigen a vel subpinnatis, ramo- 
rum plurimis vel omnibus a 
‘ id T. pinnatum, var. subternatum 
a. Folia pinnata, pinnis ternatis............-...++++5+> 8.22%. pinnatum. 


434 Rhodora [NOVEMBER 


87. T. pudicum Standley & Boivin, sp. nov. Plania glabra, 
5-25 cm., ramosa, ramis foliosis. Caulis gracilis erectus nisi 
basis procumbens. Folia ternata et fere omnia sessilia. Infe- 
riora desunt vel stzpulae istorum adsunt, quae stipulae oppositae 
lanceolatae et 2-4 mm. longae sunt. Stipulae foliorum fere 
omnes auriculatae et erosae. Pedunculi 2.5-5.0 cm. Flores 
singulae in ramis singulis. Sepala floris maris viridia ovata ca. 


of ¢ 
soil, open situation, less than 1 ft. high, alt. 6000 ft. re 19, 1941 
(F, type; M, 1sotypr). Fia. 87. 

88. T. MADRENSE Rose, Contrib. U. S. Nat. Herb. 5: 188, 
1899. Planta glabra ramosa rigida 20-30 cm., ramis foliosis. 
Ovaria tamen interdum puberula, pilis capitatis translucidis 
minutissimis. Folia ramorum 1—2-ternata cetera 2-(3)-ternata. 
Stipulae integrae. Pedunculi 1.5-3.0 cm. Flores in paniculis 
angustis caulem et ramos definientes. eA foeminei lanceo- 
lata 1-2 mm. longa. Stigma 1.0-1.5 mm. Carpella matura 
ventre 3-4 mm. — et 1.5-1.8 mm. lato. Planta mascula et 
tempus florendi ignota.—Merxico, puRANGO: Rose 3505, Sierra 
Madre, alt. 5200 ft., Aug. 15, 1897 (US, paratype). NAYARIT: 
Rose 2232, Territorio de Tepic, i in the Sierra Madre, near Santa 
Tec. Aug. 13, 1897 (US, typr; G, isoryrr). Fia. 88. 


Type not seen. 


89. T. prnnatum Watson, Proc. Am. Acad. 23: 267, 1888. 
Planta erecta rigida glabra, 2-8 dm. alta. Radices tuberosae. 
Folia caulinaria pinnata, pinnis ternatis vel simplicibus raro 
pinnatis. Stipulae integrae. Foliolae parvae, basae cuneatae, 
apicales saepius trifidae. Sepala floris maris elliptica, 3.5-6.0 
mm. longa, foeminei pee ars vel linearia 1-2 mm. longa. 
Filamenta 3-4 mm. Antherae 2.5-4.5 mm., acumine 0.3-0.7 
mm. Stigma 0.6-1.5 mm. Carpella matura 3.5-5.0 mm. eta 
stipite 0.2-0.4 mm., ventre 2.5-3.0 mm. longo et 1.5-1.7 m 
lato, nervo dorsali asso al higacho ventrali 5 ices: 
Floret Junio Julioque—Mexico, curmvanva: LeSueur 1059, 
Culebra Mts., Aug. 18, 1936 F. "G): S. S. White 2255, 4 miles 
northwest Chihuahua, Majalea (Pilares), alt. 6800 ft. , Aug. il, 
1939 (G); M. E. Jones, Sierra Madr = Mts., Soldier Canyon, alt. 
6600 ft., Sept. 16, 1903 (CA, UC); T ownsend & Barber 134, in 
the Sierra Madres, n near Colonia Garcia, alt. 7500 ft., July 15, 
1899 (F, G, M, NY, UC, US): ic ew 110, July 6 (F, M, NY, US); 
Shreve 9050, summit of Santa Clara Mts., in pine forest, alt. 
7100 ft., Aug. 13, 1939 (G); Pringle 1568, Sierra Madre, alt. 
7000-8000 ft., Oct., 1888 (UC); Pringle 1 887, base of the Sierra 


EN RES TP ee ee ee TR ea ee ae ee RE TD eT ETT 


1944] Boivin,—American Thalictra and their Allies 435 


Madre, pine pane, Sept. 20, 1887 (G, rypz; ANS, F, NY, US, 
ISOTYPES) ; Nelson 6087, in the Sierra _ Madre, June-July, 1899 
(NY, US); ps 7997, 6 mi. w. of C.G in open 
pine forest, alt. 7100 ft. ee 25, 1937 (F); Ean sue, Sen 
Juanito, July 20, 1938 (F): Hartman r72, near canyon de St. 
lego, moist places on high mountains, “Culantrio”, Sept. 17, 
me (G, US). sonora: Pennell 19, 611, Cerro Saguarivo, east 
f San Bernardo, on rocks, alt. 1500-1600 m., Aug. 7-8, 1935 
(ANS): Schott 3, Sierra del Najarito, June 28, 1855 (F). DURAN- 
Go: E. Palmer 381, Otinapa, July 25—Aug. 5, 1906 (F, G, M, NY, 
UC, US); Garcia 341 (US); Nelson 45641, r El Salto, alt. 
8000-8700 ft., July 12, 1898 (US); Pennell 18, ‘324, El Salto, dry 
open pineland, herb, ‘alt. 2570-2600 m., Aug. 28, 19384 (ANS). 
Iq. 8 
90. T. ye ATuUM Wats., var. subternatum var. A 
varietate typica differt folits caulinariis subternatis vel as 
ternatis foliolisque membranaceis. Plania humilior est, 1-4 
dm. alta, et crescit iri locis humidosioribus. Floret Julii mense.— 
Mexico, CHIHUAHUA: Gentry 1526, Sierra Charuco, Rio Fuerte, 
transition, herb scattered along rocks in spring in meadow, 
July 25, 1935 (F, trypr; ANS, G, M, UC, US, 1soryPss); LeSueur 
1211, Chuichupa, Aug., 1936 (G). DURANGO: Pennell 18,378, 
7" Salto, edge of marsh, herb, alt. 2530-2540 m., Aug. 28, 1934 
NS) 


Subsectio Dioica (Prantl), stat. nov. Sect. Camptogastrum, 
KE. Dioica Prantl, Nat. Pfl. 3, 2: 66, 1888. Plantae glabrae. 
Radices fibrosae. Caulis erectus rigidus. Stipulae integrae. 
Stipellulae desunt. Petioli articulati in insertione. Carpella 
matura_ sessilia inflata, ventre’ symmetrico nec compresso, 
pariete firmo, nervis simplicibus et directis. Species typica: 
Thalictrum diotcum L. ut patet e Prantl. 


This group is made up of only one very distinct and little 
variable species. 


91. T. piorcum L., Sp. Pl. 1: 545, 1753. 7’. laevigatum Michx. 
Fl. Bor.-Am. 1: 322, 1803, 7. pauciflorum Rafinesque, Atl. 
Journ. 1: 151, 1832, nee T. pauciflorum Royle, Ill. Bot. Him. 52, 
1839, nec T. ’ pauciflorum Schur, Verh. Sieb. Ver. Nat. 3: 84, 
1852, nec 7’. pauciflorum Steph. ex Lec., Bull. Soc. Bot. Belg. 24: 
302, 1885. TT. — L., var. genuinum Lee., Bull. Soc. Bot. 


Pian Greene, ‘Le. * 1908. r dioicum, L. var. ig ian ee 
Greene, |. ¢. 40, 1909. Leucocoma dioica (L.) Nieuwland, Amer. 
Midl. Nat. 3: 304, 1914.—Planta omnino glabra erecta rigida, 
3-8 dm. Radices fibrosae. Folia supraternatisecta, basilaria 


436 Rhodora [NOVEMBER 


0-3 in planta, caulinaria 0-1, inflorescentiae plurima. Folium 
inferius inflorescentiae petiolo 3-8 cm. Sepala floris maris 
ovata vel elliptica 2.5-4.0 mm. longa, foeminei ovata vel obovata 
1.8-2.2 mm. longa. Firlamenta lutea 3.5-5.5 mm. Antherae 
luteae oblongo-lanceolatae vel lineares 1.4-4.0 mm. longae, 
acumine 0-0.2 mm. Stigma 1.0-2.5 mm. Carpella matura 4-5 
mm. longa, stipite 0.3-0.4 mm., ventre ovoideo vel ellipsoideo 
3.4-4.3 mm. longo, 1.5-1.8 mm. lato. Nerv carpellorum 
maturorum parum a costis distincti, pariter arcuantes vel 
interdum nervus ventralis quam dorsali convexior. Floret 
Aprili, Majo, Junio, Julioque.-—“Canapa”: Kalm (Linnean 
Society Lond., TYPE ‘of T. dioicum; G, photograph). Unrrep 
STATES (without localization): Michaux, in Pensylvania et 
montibus Carolinae (Paris, National Museum, type of 7’. laevi- 
gatum; G, photograph). Saint-Prerre ET MIQUELON: 
Arséne 238, Saint-Pierre, sgh es de la ferme Maillard, lieux 
herbeux et ‘humides, 8 juillet, 1902 (NY); L. Arséne 263, Saint- 
Pierre, anse & Ravenel, lieux herbeux et humides, bois, 8 et 20 
juillet, 1900 (G). Shae Peers, Montreal, Mt. Royal, May 
11, 1941 (CA); C. H. Knowlton, ’ Missisquoi, Philipsburg, dry 
calcareous woods, 10-11, 1923 (G); Macoun & Malte 88,011, 
Hull, north end of Fai airy Lake, May 04 (C). 

Fernald, Penobscot Co., Orono, rocky woods, May 19, 1890 
(NE); C. A. E. Long 858, Knox Co., Rockland, rocky bank, 


Rand ‘é Robinson 458, Cheshire Co., Troy, Gap Mt., becky 
_ woods, June 13, 1898 (G, NE). Vermont: C. H. Knowlion, 
—- Co. , Newbury, slaty woods, ned 17, 1932 (NE); E. E. 


shire oe Mount pert dry open eer “May 30, 1919 
(NE); @. Gilbert, Newton, Pine Grove, May 6, 1894(G). RHODE 
Istanp: Leland, Johnston Co., Snake Den, May 13, 1899 (NE); 


1944] Boivin,—American Thalictra and their Allies 437 


a B. Simmons, Newport Co., Middletown (NE). Connecti- 
uT: Woodward, New London Co., Rachel’s Hut, rocky woods, 
Nii 27, 1905 (NE at. Spalding, Hartford Co., East proper eh 
May, 1920 (NE); Safford, New Haven Co., New Haven, East 
Rock, June 2, 1884. (US). New York: 0. P. Phelps 160 ee 461, 
Canton, woods, May 19, 1914 (G); Burnham, Washington Co., 
% mile east of Tripoli schoolhouse, rocky copse, May 13, 1914 
(G); McVaugh 4107, Columbia Co., Ghent, moist rocky banks, 
May 1, 1986 (G); Raup 8105, Orange Co., Black Rock Forest, 
ravine of Canterbury Brook, May 18, 1937 (G); Eames & M et- 
calf 2301, Tompkins Co., Ithaca, gravelly soil, side of gully, 
May 7, 1904 (G); E. B. Ba rtram, Monroe Co., Rochester, moist 
woods, May 15, 1910 (G); Peck, ‘St. Lawrence Co., Rossie, June 
10, 1909 (US). New JERsrEy: ‘Carey, Bergen, ravine, May 24, 
1842 (G); Fogg 12,313, Somerset Co., Sourland Mts., about 4 m. 
n. w. of Blawenburg, July 7, 1937 (G); B. Long 39,9838, Hunter- 
don Co., Treasure Island, steep wooded meng along Delaware 
oe May 21, 1933 (G). PrNNsyYLvaAntia: Fogg 6502, enters 
Co. f Swa rthmore, wooded atten along creek, May 17, 
1934 (G); Heller, rig ya see on the Conestoga near Binkley’s 
Bridge, in limestone, May 1901 (G); Galen 24, Lancaster Co., 
Apel May. 1885 (G); Wal 643, Elk Co., 3 mi. W. of serene 
ag roadside, June 2, 1940 (G); Fender 1500, Union Co., 
of Lewisburg, along West Branch of Susquehanna Riven 
pie 15, 1936 (G); E. B. Bartram, Chester Co., Frazer, April 23 
& 30, 1910 (G); Fogg 14,478, Bradford Co., 1.5 mi. n.-nw. of 
Wyalusing, along Susquehanna River, wooded crest of cliffs, 
June 8, 1938 (G). District or Cotumsra: T. A. Williams, 
Rock Creek, May 1, 1898 (G). West Virarnia: Eggleston 4344, 
White Sulphur Springs, May 17-18, 1909, (G). Virernia: 
Small, Smyth Co. E. Marion, on Stalie’s Knob and Stalie’s 


Hot Spri rings, woods , May 1 1916 (G): Ae Hd. Curtiss, Bedford 
Co., Apel 15, 1871 (G); ott 14,589, Giles Co., 214 mi. ne. of 
Mountain Lake Por , Salt Pond Mt., ‘alt. 3800 ft., June 22, 1938 
ref Churchill, Rockbridge Co., Goshen, rocky woods, April 28, 

5 (M). NortH CAROLINA: "Ashe, Mitchell Co., Frank re, 
May 11, 1893 (NC); Peattie 1667, Polk Co., Tryon, in cool 
mountain woods, April 20, 1922 (NC); Barksdale, Bat Cave 
Mountain, April 19, 1937 (NC). SoutH CAROLINA: House 2064, 
Oconee Co., Tamassee Falls, May 5, 1906 (M); House 1856, 
Oconee Co., Clemson College, near Seneca River, low woods, 
April 16, 1906 (M ). Groraia: Leeds 2012, Union Co., 1 mile n. 
of Fro ogtow n Gap on branch of Helton Creek, June 1, 1934 
(ANS); Churchill, Lookout Mountain, near the line between 
Tennessee and Geo orgia, April 19, 1906 (G). Onrarro: J. 
Macoun 33,609, Niagara, May 9, 1901 (C, G); T. M. C. Taylor 


438 Rhodora [NovEMBER 


et al. 2277, Algoma District, Carp River, mixed woods, bottom- 
lands, July 15, 1935 (C); J. Macoun 72,515, near St. Thomas, 
woods, June 24, 1907 (C, type of 7. dioicum var. adiantinum) ; 
J. Macoun 83, 608, Lake Huron, Pt. Edward, June 12, 1901 (G, 
NY). Micuican: Greenman 3356, Sawyer Beach, Hemlock 
Dune, May 5, 1912 (G). Onto: Clavenger, Columbus, June, 
1903 (US); Gleason, north of Columbus, rich wooded hillsides, 
May 13, Sto (G); Webb 496 & 497, Portage Co., Nelson Ledge, 
May 3, 1902 (G). Inprana: Deam 16,861, Noble Co., about 4 
mi. northwest of Kendallville, in woods, June 6, 1915. (G). 
Kentucky: Demaree 11,252, ah ie Co., South Portsmouth, 
fertile river-hillsides, April 21 1935 (M); W. A. Anderson 407, 
Fayette Co. , Elklick, March 26, 1927 (G); "Blumer 238, Louisville, 
River Road, woods, ’ April 2, 1933 (G); W..A. Anderson 9, Hen- 
derson Co., Robard, April, 1923 (G). TENNESSEE: Ruth 888, 
Knoxville, "Tennessee River, bluffs, May 20, 1894 (G); Ruth, 
Knoxville, rocky woods, April, 1896 q M); Eyles 7718, Obion Co., 
near Walnut Log, bluff, April 25, 1941 (G). ALABAMA: Mohr, 
Huntsville, Mount Sano, i in sylvis umbrosis, May 8, 1881 (F); 
Mohr 66, near Huntsville, Mount Sano, upper districts, rich 
woods, May 1, 1881 (US); Harper 3840, Jackson Co., southeast 
of Scottsboro, ‘northwest slope of Sand "Mountain, among lime- 
stone rocks in woods, April 17, 1935 (G, NY, US). ILLINOIS: 
Pease 11,831, Champaign Co., Urbana, “moist bank near Crystal 
Lake, April ; 27, 1909 (G); Gleason §27, Urbana, hillside, April 28, 
1899 (G). MINNESOTA: Pammel 204, Cass Lake, abundant in 
pine woods with Hard Maple and Basswood, July 28, 1925 (G); 
M. L. Grant 2703, Clearwater Co., Itase a Park, near campus, 
woods, sand, ie! 9, 1929 (Q); Sandberg 1148, Itaska Lake, along 
stream, July 6 ; 1 (US). Iowa: Pammel, Ames, College Park, 
May 28, 1927 (ANS); Fink, Fayette, prairies, 1894 (G); F ink, 

o1 (US); Fink ee Fayette Co., May 15, 1894 (US). Muis- 
a Steyermark 11,569, Wayne Co., west of Grenville, along 
St. Francis River, July 9, 1936 (M); ‘Steyermark 11,220, Barry 
Co., south of Shell Knob, Smith Pond, wooded lime slopes, 
mg 26, 1936 (M); E. J. Palmer 35 ,966, Benton Co., Fredonia, 
rich woods along bluffs of Osage River, May a 1929 (G); E. J. 
Palmer 22,826, Stone Co., Sarr limestone bluffs of the James 
River, rich woods, May 27, 3 (M); Bush 753, Jackson Co., 
Courtney, May 2, 189 6 (M, US) i. Bush 98, Sheffield, common on 
rocky hill, May 12, 1898 (G,M US). 


Specimens collected in July ‘a Saint-Pierre are in flower, while 
in Georgia and Alabama this species matures its fruits in May 
or June. No specimens were seen either from eastern Québec or 
from the Maritime Provinces. We do not know if there is any 
type in existence of 7. pauciflorum Raf. 


Ge 


1944] Boivin,—American Thalictra and their Allies 439 


Subsectio Incurvata subsect. nov. Plantae stoloniferae gla- . 
brae vel pubescentes. Pubescentia si adest e pilis capitatis 
minutis translucidis. Folia supraternatisecta. Injlorescentia 
aphylla vel, parum si foliosa foliis sessijibus, paniculata, apice 
acuta. Flores dioici, rarissime polygami. Sepala dimorpha, 
floris masculi majora. Filamenta colorata. Carpella matura 
subsessilia costata apice incurvata, nervo dorsali convexiore 

m ventrali, pariete saepius incrassato. Nervi carpellorum 
costis parum distincti. Species typica Thalictrum venulosum 
Trel. sit. : 


a. Carpella matura haud compressa... .b. ; f 

b. Stigma (1.0-) 1.5-2.0 (-2.5) mm., venter carpelli maturi 
mm. longus, 1.8-2.4 mm. latus........... ---92, T, venulosum. 

b. Stigma (2.0-) 2.5-4.0 (-5.0) mm., venter carpelli maturi 

4-6 mm. longus... .c. 

c. Venter carpelli maturi 2.5-2.7 mm. latus............ 93. 7’. confine. 
c. Venter carpelli maturi 2.0-2.3 mm. latus. 
T. confine, var. Greeneanum. 


a, Carpella matura paullum compressa. . . 05. T. confine, var. columbianum. 


92. T. venvuLosuM Trelease, Proc. Bost. Soc. Nat. Hist. 23: 
302, 1886. TT. campestre Greene, Erythrea, 4: 123, 1896. T. 
Lunellii Greene, Midl. Nat. 1: 102, 1909. T. thyrsoideum 
Greene, Mid]. Nat. 1. c. 1909. TT. thyrsoideum sylvanum Lunell, 
Amer. Midl. Nat. 2: 157, 1912. Leucocoma Lunellii (Greene) 
Lunell, 1. c. 4: 361, 1916. JL. thyrsoidea (Greene) Lunell, |. c. 361, 
1916. L. thyrsoidea sylvana (Lunell) Lunell, 1. c. 361, 1916. 
Planta 20-75 em. Folza basilaria 1-2, caulinaria 1-3 in planta. 
Pedunculi fructuum 0.3-1.5 em., conspicue ascendentes vel ap- 
pressi, saepius gemini. Sepala floris masculi 3.5-4.0 mm. longa, 
foeminei ca.2mm. Filamenta 3-4 mm. Antherae (2.0)—2.5-3. 

m., acumin . 0.1 mm. Stigma (1.0)-1.5-2.0-(2.5) mm. 
Carpella matura stipite 0-0.3 mm., ventre 3-4 mm. longo et 
1.8-2.4 mm. lato. Floret Majo, Junio, Julioque-—NorTHWEST 
America: Franklin Expedition (G, tTrpn of T. venulosum); 
Richardson 207 (G, paratype). Quésec: D. Potter 380, James 
Bay, mouth of the Abitibi River, bank opposite Allen Island, 
June 30, 1929 (G); Roy, comté de Labelle, Bellerive, 9 juin 1935 
(G). Onrarto: C. S. Williamson 2028 & 2032, vicinity of Fort 

illiams, Kaministiqua, Aug. 2, 1912 (ANS); D. Potter 379, 


SIN: Griscom, Sawyer Co., R. R. right of way north of Hayward, 
June 11, 1928 (G). Manrropa: E. L. Greene, near Carberry, 
prairies, 1890 (G-ND, type of 7. campestre; G, photo of type; 
NY, isotypes); Scamman 2892, Riding Mountain National Park, 
Clear Lake, alt. 2016 ft., Aug. 29-Sept. 2, 1941 (G); Macoun & 
Herriot 69,804, 6 mi. east of Forest, June 19, 1906 (G); Herriot 


440 Rhodora [NovEMBER 


69,803, Portage La Prairie, May 31, 1906 (G). MINNESOTA: 
Chandonnet, Ottertail Co., Ottertail Lake, prairies, June 8, 1910 
(N-ND). Norta Dakota: Stevens 17 77, Cass Co. , Fargo, open 
places in woods, June 30, 1936 (G, F); ees ’ Benson Co., 
peninsula of Lake Ibsen, in woodlands, May 30, 1910 (NY, syn- 
type of T. thyrsoideum sylvanum); idem, May 28 (N-ND, syn- 
type of T. thyrsoideum sylvanum); idem, May 24 (F, NY, syn- 
types of T. thyrsoideum sylvanum) ; vere Ward Co., Mi not, 
along Mouse River, in timber, July 1, 1909 (G-ND 14394, type 
of 7. Lunellii; G, ‘photo of type; N-ND, isotype) ; Lunell 76, 
Benson Co., Lees, in dry soil on the prairies, June 10, 1909 
(G-ND, type of T. thyrsoideum; G, photo of type; N-ND, NY, 

isotypes). SoutH Dakota: i Pal mer 27,198, Lawrence Co., 
Boulder Canyon near Deadwo rocky ground along brook, 
June 12, 1929 (G); Hayward pea Teh 1840, 2060, 2200, 2304 
& 2410, Black Hills, various localities, 1927 (F). SASKATCHE- 
WAN: Bourgeau, 1857-58 (G, paratype of T. venulosum); 
Macoun 2952, Crane Lake, June * tg (G); Macoun & H Aad 
69,802, Tramping Lake, Aug. 4 6 (G); ‘J. Macoun 12,316, 
near Prince Albert, lat. 53°, ja 1896 (G). Auperta: J. 
Macoun 59,624, Burnt River, north of Peace River, open prairies, 
July 19, 1903 (G); Moodie 9386, Red Deer Valley, vicinity of 
Rosedale, low ground, alt. 2200-2500 ft., June 1, 1915 (F, G); 
Raup 2480, 2488, 2434, 2486a, 2439, 2440, fue & 2444, Wood 
Buffalo Park, various localities, 1928-29-30 (G). MOonrTANaA: 
B. J. Jones, Gallatin Co., Sedan, June ig 1901 (G); Flodman 
485, Gallatin Co., East Gallatin Swamps, alt. 5000 ft. , July 24, 
1896 (US). Wromine: Clifford Richardson, Alkali Co., Branch 
of Sandy R., 1878 (G, paratype of 7. venulosum); A. Nelson 
7307, Albany Co., Chug Creek, on the open bottoms, June 29, 
190 Ci Schubert, Albany Co., Bear Creek about 2 miles 
from Eagle Peak, and 4 miles from Laramie Peak, Aug. 22, 1899 
(US) ; Forwood, Wind River, July 24, 1882 (G). 

British CoLuMBIA: Raup & Abbe 3557, north bank of Peace 
— at ap oe Flats, dry bluffs, about 56° 8’ N., 120° 40’ W., alt. 

-, June 12, 1933 (G); McCabe 674, Chezacut, very abun- 

dent ne in dry si tes, Aug. 30, 1983 (UC); S. Brown 960 & 972, 
North Branch Saskatchewan, ‘Kootenay Plains, June 17, 1908 
(G). Fig. 92, a-e 

93. T. CONFINE Fernald, Ruopora, 2: 232, 1900. T. purpur- 
ascens L. sensu DC. Syst. 1: 174, 1817. T. purpurascens L., @ 
monoicum DC. Syst. 1: ae 1817. T. dioicum X pur purascens 
Trel., Proc. Bost. Soc. Nat. Hist. 23: 301. 1886. Planta 25- 
120 em. Folium basilarium vel solitarium vel deest, caulinaria 
1-5 in planta. Pedunculi fructuum AG a —1.5-2. 5-(3. 0) cm. 
ascendentes. Sepala floris maris 2.5— longa, foeminei 
1.5-2.4mm. Filamenta 3.0-4.5-(5.5) er invA ntherae (2.0)-3.0- 


1944] Boivin,—American Thalictra and their Allies 441 


4.0 mm., acumine ca. 0.4 mm. Stigma (2.0)—2.5-4.0-(5.0) mm. 
Carpella matura stipite 0.1-0.2 mm., ventre 4-6 mm. longo et 
2. . lato, nervis nonnunquam ramosis ate im) 
Foret Junio vel Julio et interdum Augusto.—LaBRapor: 
Fowler, Lower East Main Coast, Aug. 20, 1892 (US). Gratis: 
D. Potter 382, James Bay , East’ Main, marshy river bank, July 
11, 1929 (G); Williams - Fernald, Témiscouata Co., Riviére- 
du-Loup, rocky bank of the St. Lawrence, Aug. 2, 1902 (G); 
Rousseau 26,423, comté de Rimouski, pointe du Vieux-Bic, sur 
les schistes, 30 juin, 1927 (G); Fernald & Pease 25,074, Matane 
Co., Ste. Félicité, Sattaera July 17, ler (G): Victorin & 
Rolland 18,777, archipel de Mingan, fle au Mart rteau, sur les 
rivages calcaires, 15 juillet, 1924 (G); Victorin 4288, y Paring 
cap a |’Ours, Aug. 23, 1917 (G); Victorin 28,156, Boucherville, 
bordant le rivage de oo des files, avec Carex intermedia et 
Carex nutans, 1 sept., 8 (G); Victorin 3233, St.-Eustache, 
Aug., 1916 (G): Ou ca 7 Fassett 18,100, Gatineau Co., Hull, 
June 29, 1934 (C); C. H. Knowlton, Philipsburg, dry open woods, 
shale ledges, July 18, 1931 (G). New Brunswick: Chalmers 
hg Restigouche, Eel River, woods, July 29, 1876 (C, 2 
release T. dioicum X purpurascens; G, sam e note); J 
M. acoun 21,136, along the St. John, above pr deere July 3 
1899 (C, G); Williams, Collins & Fernald, Gorge of Aroostook 
River, Andover, talus from calcareous cliffs, July 17, 1902 (G); 
Pease & Edgerton 27, weet i ig Mangerville, alluvium by St. 
John R., June 29, 1939 (G 3G Hay, St. John, Aug., 1890 YG). 
MAINE: Fernald, ’ Aroostook Co., ees Fairfield, fruits from plant 
in river-thicket, ‘Sept. 19, 1900 (G, TYPE of T. ‘confine; e; C 69,609, 
ISOTYPE); Fernald 215, Fort Fairfield, abundant in alluvial 
thickets, fl. June 6, 1901, fr. July 16, 1902 (G, NE). VERMont: 
Eggleston 4869, Ferrisburg, Gardener’ s Island, July 17, 1909 (G); 
Flynn, Chittenden Co., Burlington Bay, rocky shore of Lake 
Champlain, Sept. 8, 1909 (G, NE); C. H. Knowlton, Franklin Co., 
Lake Champlain, Swanton, sandy shore, July 24, 1926 (NE); 
C. H. Knowlton, Grand Isle Co., Grand Isle, swampy woods near 
Lake, Faly 24, 1935 (NE). New York: House 1 10,323, Essex Co., 
Port Henry, rocky shore of Lake Champlain, July 12, 1924 (G, 
US). rman J. Macoun 2956, Ottawa, Rideau Hall, ao rneen 
Aug. 8, 1894 (C, paratype); D. ’Potter $81, James Bay, 
River, ‘north end of Saw Pit Island, marsh, July 1, 1929 (G): 
Pease & ji stty 23,557, Thunder Bay District, W. side of Jack 
Fish Bay, beside a brook, July 8, 1933 (C, G); Taylor, Losee & 
Bannan 905, Thunder Bay District, Sibley Tp., Talus Bay, 
gravelly beach, June 24, 1936 (C). Micuican: Fernald & 
Pease 3306, Schoolcraft Co., east of Manistique, glades and 
Openings in thicket bordering calcareous beach of Lake Michigan, 
July 9, 1934 (G). Manrropa: Bourgeau, Lake Winnipeg Valley, 


442 Rhodora [NovEMBER 


1857 (G); G. Gardner, nae Ba ae South Shore, 19 juin, 
1932 (ANS). Minnesota: Lakela 1408 be 1428, a June 
11 & 19, 1936 (ANS); idem, “ene 17, 1938 (G). Fria. a-f. 

Thalictrum dioicum X purpurascens Trel. also sue 1 oe 
coriaceum (Britt.) Small. Trelease referred to those plants being 
“hybrids of 7. diotcum L. with T. purpurascens L. or polygamum 
Muhl.”” No specimen has been found marked by Trelease T. 
dioicum X polygamum. Oddly enough, one of these supposedly 
hybrid plants comes from a region where the only other Thalic- 
trum yet known to occur is T. polygamum. 


94. T. CONFINE Fern. var. Greeneanum, var. nov. Planta 
30-70 em. Folia basilaria 1-3, caulinaria 1-2 in planta. Fuila- 
menta 2.5-3.2 mm. Antherae 2.5-3.0 mm., acumine 0.3-0.7 mm. 
Stigma 2.0-3.0 mm. Carpella matura ventre 4.5-5.0 mm. longo 
et 2.0-2.3 mm. lato. Floret saepius Julio, interdum Junio.— 
Cotorapo: Parry, Middle Park, July, 1864 (G, paratype of T. 
venulosum); Vasey 10B, Middle’ Park, 1868 (G, paratype of T. 
tioned Baker 561, Gunnison, alt. 7680 ft., July 23, 1901 (G, 

NY, US, ISOTYPES) ; idem 556 (G, US); "Hall & Harbour 8, 
lat. "39 O41? 1862 (F); Coulter, Weston’s Pass, alt. 11,000 ft., 
July 18, 1873 (US); Patterson, Middle Park, west end, damp 
plains, July 27, 1875 (F); B. H. Smith, Archuleta Co., Pagosa 
Springs, June 15, 1894 (ANS): Boi; Smith, Mineral Co., Wagon 
Wh Gap, July, 1882 (ANS). Urau: Pennell & ’ Shaeffer 
22,522, Wasatch Co., head of Daniels Canyon, northeast of 
Hebert, below Daniels’ , rocky slope, alt. 7900-8000 ft., July 1-2, 
1938 (ANS); Watson 9, ’ Wahsatch, Provo Canon, July 1869 (US). 


E. L. Greene left two unpublished names on sheets of this 
variety. 


95. T. CoNFINE Fern., var. columbianum (Rydb.), stat. nov. 
T’. fissum Greene, Pittonia, 4: 233, 1901. TT. columbianum Ryd- 
berg, Bull. Torr. Bot. Club 39: 320, 1912. T. occidentale Gray, 
var. columbianum (Rydb.) St. John, Fl. South. Wash. & Adj. Id. 
156, 1937. Planta 50-80 cm. Folia basilaria 0-1, catilinasin 
1-3 in planta. Pedicelli 0.5-2.5 cm. aetate fructuum maturan- 
dorum. Filamenta 3-6 mm. Antherae 1.5-4.0 mm. Stigma 
2.0-3.0 mm. Carpella matura ventre paullulum compresso, 3-6 
mm. longa et 1.8-2.5 mm. lata. Floret Majo, Junio Julioque.— 
Ipano: Mulford, De Lamar, alt. 7000 ft., July 7, 1892 (G, para- 
type of T. columbianum). Onrxcon: Cusick 1187, 1886 (NY, 
paratype of T. columbianum); Cusick 1837, Mts OB Oregon, & 
form not common in the mountains, Jone: "July, 1886 (G, NY, 
paratypes of 7. columbianum); Henderson 9056, Harney 
Steins Mts., near Sheep Camp, moist bottoms in aspen woods, 


OE AES EE SEC eS oF F 


1944] Boivin,—American Thalictra and their Allies 443 


alt. 5500 ft., June 15, 1927 (CA); Henderson 9058, Harney Co., 
15 miles above Bu urns, up Silvies River, moist sunny flats, June 
13, 1927 (CA); Cusick, Stein’ s Mountains, at the head of Wild 
Horse Creek, July 14, 1898 (G-ND, type of T. fissum; G, photo- 
graphs of the type). Wasnincron: Elmer 599, Okanogan Co., 
Loomiston, Mount Chapaca, wide g ge in the meadow on the 
south slope, Aug., 1897 (NY, Type of 7. columbianum; BG, US, 
ISOTYPES); G. R. Vasey 155, 1399 (G, NY, US, paratypes of T. 
columbianum oH Henderson 2367, Yakima Co., June, 1892 (G); 
Piper 1467, Withman Co., Pullman, June 26, 1893 (G); Lyall, 
Pend d’ Oreille River, 1861 (G). 

BRITISH CouuMBIA: Raup & Abbe 2642, vicinity of Hudson 

eed south slopes of Peace River Valley, about 56° 1’ N., 121° 

3’ W., June 21, 1932 (G); ; McCabe oer, —_ a north of 
oe moist hollow in open range, June 13, UC); 
McCabe 8232, one see: ag southeast of Here Ha nis yet ope 
forest edge, border of bog, in - 1940 (UC). 

Subsectio Clavocarpa subsect nov. Plantae glabrae elatae 
rigidae. Species duae, alia teloitaes alia haud. Caulis saepius 
glauca. Folia, nisi inferiora, sessilia in apice dilatationis 
petiolaris. Inflorescentia paniculata copiosa. Sepala lanceolata. 
Stigma bialatum. eek orien os recta stipitata, nervis 
exsertis sublatis in costis obtusis, nervo ventrali convexiore quam 
dorsali. Species typica Thalictrum pases ove (Britt.) Small sit. 


Planta eeepc) a; anthers luteae vel viridulae; carpella — 


ven PAIWRIE 5 ats eG ey ena cx eh eben T. Steeleanum. 
Plan a bad Molentierk: stamina imprimis lutea, deinde purpur- 
ascentia; carpella matura ventre ovoideo.............. 97. T. coriaceum. 


96. T. Steeleanum, sp. nov. Planta ane stolonifera 1.0- 
15 m. Foliolae majusculae, 2.5-7.0 cm. latae, interdum sub- 
quadratae. Sepala mascula 4-5 mm. longa, alate ovata ca. 
2mm.longa. Filamenta saepius lutea, interdum purpurascentia 
4.0-4.5 mm. Antherae saepius luteae, interdum viridulae, 
lineares, 3.2-4.5 mm., acumine 0.4-1.0 mm. — 2.0-2.8 


er, 

June 3, 1899 (ANS). MARYLAND: Steele, C. & O. Canal, pete! 
5 OR 903 ve nee bank of Potomac above D.C., July 23, 1902 
G, Y); , Potomac, near the Md. line, ‘Feeder "Dam 
Island, Suly 10, "1902 (N Y); ee idem, May 1 0 (G, NY); 
Steele, ‘bank of Potomac, above D Wa Cad 26, 1902 (G, NY); E. 
orris 1527 , Montgomery Co., Plummers Island, alluvial 
woods, May 18, 1902 (BG); Long ‘& ree 1292, Cecil Co., 


444 Rhodora [NovEMBER 


north of Conowingo, woods, June 1, 1913 (G, TYPE); idem 1287 
& 1289 (ANS); Hitchcock 12,902a, Great Falls, woods along canal, 
June 21, 1917 (G); Shriver, Cumberland, 1894 (NY). VuirGinia: 
Allard 2873, Fauquier Co., near Markham, woods at top of 
Rattlesnake Mountain, abundant and in full bloom, under- 
ground rhizomes a rich golden yellow, May 30, 1937 (G); #. L. 
Morris 2361, Falls Church, dark woods, May 1896 (BG); Camp 
1174a & 1174b, Shenandoah National Park, south and southeast 
of Skyland, alt. about 3500 ft., May 25, 1986 (NY); Rawlinston 
16, Madison Co., near Hoover’s Camp, June 22, 1934 (US); 
Wherry & Pennell 13,3844, Page Co., southeast of Luray, Stony 
Man Mountain, Aug. 24, 1927 (M). Fie. 96, a-d. 

97. T. corraceum (Britt.) Small, Mem. Torr. Bot. Club, 4: 
98, 1893. T. dioicum L., 8? stipitatum Torr. & Gray, FI. N. 
Amer. 1: 38, 1838. TT. dioicum L., var. stipitatum Lecoyer, Bull. 
Soc. Bot. Belg. 24: 142, 1885, pars. TT. diotcum L., var. cortaceum 
Britton, Bull. Torr. Bot. Club, 25: 136, 1898. 7. caulophylloides 
Small, Bull. Torr. Bot. Club, 25: 136, 1898. Planta glabra, 
haud stolonifera, 65-100-(150) cm. Foliolae (1)-3-(7) cm. 
latae. Sepala floris masculi 3.5-4.0 mm. longa, foeminei fere 
lanceolata 1.5-2.0 mm. longa. Filamenta 4.5-5.0 mm. An- 
therae 2.0-3.5 mm., acumine ca. 0.4 mm. Stigma 1.5-2.5 mm. 
Carpella matura stipite 0.3-5.5 mm., ventre ovoideo 2.3-4.0 mm. 
longo et 1.3-2.0 mm. lato, nervis paululum sinuosis. Floret 

unio, rarissime Majo vel Julio—Werst Vireinia: Brition, 
White Sulphur Springs, May 16, 1897 (NY). Vzireria: Small, 
Smyth Co., slope of White Rock Mountain, alt. 3500-4000 ft., 
June 21, 1892 (F, M); A. H. Curtiss, Bedford Co., 1871 (F, G, M); 
E. 8. & Mrs. Steele 146, near Luray, Stony Man Mountain and 
vicinity, alt. 3600 ft., Aug. 18, 1901 (G, M, NY, US); Steele, 
Augusta Co., vicinity of Augusta Springs, over meter high, alt. 
+ 1200, Sept. 8, 1908 (M, US). Norra Carona: M. A. 
Curtis, Bald Mt. (G, type of T. dioicum @? stipitatum Torr. & 
Gray); Small & Heller 268, Watauga Co., Blowing Rock Moun- 
tain, alt. 4000 ft., July 29, 1891 (NY, Type of 7. dioicum, var. 
coriaceum; US, 1sorypr); Porter, Mtns., June 28, 1880 (ANS, 
NY, paratypes of var. coriaceum) ; Small & Heller, Blowing Rock 
Mountain, alt. 4000 ft., June 10-20, 1891 (F, NY, paratypes of 
var. coriaceum); Small & Heller, eastern slopes of Blowing Rock 
Mountain, July 7, 1891 (F, paratype of var. coriaceum); idem 
491 (ANS, F, M, paratypes of var. coriaceum); Small & Heller, 
on the road between Blowing Rock and Shull’s Mill, June 16-17, 
1891 (F, M, NY, paratypes of var. coriaceum); Small & Heller 
491. Caldwell Co., summit of Stone Mountain, July 9, 1891 
(ANS, US, paratypes of var. coriaceum); House 4346, Transyl- 
vania Co., Pisgah Ridge, alt. 4000-5000 ft., July 4, 1909 (US); 
Biltmore Herbarium 6043a, Buncombe Co., slopes of Cedar Cliff 


ee 


i al mI Meas Penge foe pits 


1944] Boivin,—American Thalictra and their Allies 445 


Mountain, rich soil, May 24, 1898 (NY); Mohr, Grandfather 
Mountain, rich woods, uly 27, 1894 (US). GrorGIA AND 
Carouinas: Buckley, in mountibus Carolinae et Georgiae (M, 
paratype of 7. dioicum X purpurascens); Wright, N. Georgia, 
1875 (G). Kentucky: Kearney 290, Harlan Co., Big Black 
Mountain, Aug., 1893 (G, NY). TENNESSEE: Svenson 8309, 
Sevier Co., Thom as Ridge, south of Indian Gap, Aug. 12, 1935 
(BG): Ruth 1800, Blount Co., Cade Cove Mountain, mountain- 
sides, July, 1892 (NY, type of T. caulophylloides) ; Eggert, David- 
son Co., woods, July 13, 1897 (M, NY). Fie. 97. 
(To be continued) 


AMERICAN THALICTRA AND THEIR 
OLD WORLD ALLIES 


BERNARD BoIvin 
(Continued from page 446) 
Subsection woe alee subsect. nov. Planiae erectae rigidae. 
adix 


orsalis apex convexior quam ventralis apex. Species typica 
Thalictrum occidentale Gray sit. 


a. Crassitudo carpelli maturi dimidias latitudinis aequat. 
“lat nter ejusdem ovatus costatus, sesquies aye “ee 
u 


tie bees 
©) a Oe eee Oh oe ee ye Re Oe ee ee ee Ee ee ere, a OR OT 


5. nervosa, ter vel pes pager quam la lata... 
Carpella matura divaricata stipitata, ventro lanceolate. 
orescentia ampla ad 50 cm. pyret et s 7% josa.... 
. - Stigma 3 S648 tii. 6 ox ined cs 
mm 


aé ca. 3mm., acumine ca. 0.4mm. Stigma 3-4 mm. Carpella 
matura compressa et costata, valde Oa stipite 0.2-0.3 mm., 


454 Rhodora [DECEMBER 


ventre ovato 4-5 mm. longo, 2.5-3.0 mm. lato et ca. 1.5 mm. 
crasso, nervis a costis parum distinctis. Nervus ventralis con- 
vexior quam dorsalis. Tempus florendi a lectoribus ignotum, 
verosimiliter tamen Junium.—SovuTH Aa ieest hades P. Carr 135, 
Deadwood, shady woods, July 31, 1913 (C ae eee SF 
ISOTYPE ES); Hayward 847 (in part), Black Hills, Weed Creek & 
Dark Canyon, ¥1927 (F); Hayward 1331, Black Hills, Spearfish 
Canyon, lower 7 miles above fish hatchery, 1927 (F); Hayward 
1337, Black Hills, Deadwood, near Pinecrest Camp, 1 1927 (F). 
99. 'T. OCCIDENTALE Gray. T.dioicum L., var. paper Lec., 


Bull. Soc. Bot. Belg. 24: 142, 1885, pars, nec T. m, 6? 
stipitatum Torr. & Gray, Fl. N. Amer. 1: 38, 1838. or “deen 
L., var. oxycarpum T: Bot. Wilkes Expe d. 17: 212, 1874. 


Planta atin vel vibetviens. Pedunculi divaricati (1)-2-(5) 
em. Sepala floris masculi 3.5-5.0 mm. longa et foeminei 1.5-2. 
mm. longa. Filamenta (5)-6-7-(10) mm., purpurascentia. 
Antherae 2-4 mm. Carpella matura nervosa, vel nervo dorsali 
Sanaa et ventrali convexiore, vel dorsali basi concavo et apice 
con 

The an of the fruit is highly variable in this species and 
transitional forms to other species occasionally occur. This 
also seems to hold true for the Incurvata and Laminaria. These 
intermediates seem to point toward the following series: T’. 
venulosum—T. confine—T. occidentale—T. Fendleri—T. poly- 
carpum in which each species is most closely related to the two 
adjoining ones. The two varieties given in synonymy cannot at 
present be related to any of the following three. 


99. T. occIDENTALE Gray, var. typicu “as be apes 
Gray, Proc. Amer. Acad. 8: 372, 1873. Planta 60-120 ¢ 
Inflorescentia 20-50 cm. longa, foliosa, foliis nonnullis saticlntie 
et triternatis. Ovaria ventro lanceolato, interdum ato. 
Stigma 3.04.5 mm. Carpella matura divaricata, stipite 0.4-1.2 
mm., ventre lanceolato 6-10 mm. longo, 1.6-2.5 mm. lato. 
Floret Majo et Junio.— WasHINGTON AND OreEGoN: Hall, 1871 
(G, tyre; F, tsorrpe); J. Howell, Cascades, June, 1879 (G); 

J.& T. owell, near Cascades, in rich woods, May-June, 1880 
(ANS, F, US); T. Howell, Cascades, May, 1882 (NY); Sukadorf 

339, ‘Skamania Co ., near lower Cascades, May 30, 1886 (G); 
Epling 5533, Benton Co., Corvallis, May, 1922 (F); F. E. Lloyd, 

orest Grove, low w grounds, June 1, 1894 (NY); Henderson 8, 
Sandy River, warm rich hillsides, May 21-June 12, 1883 (NY); 
Henderson 87 7, near Columbia River, ie creeks, moist rather 
open woods, May 27, 1924 (G). G. 

100. T. occriDENTALE Gray, v. pitas var. nov. 
80-125 cm. Hila ee Adlecwsivonceth Se pedie )- 


Se ee ne TT. 


1944] Boivin,—American Thalictra and their Allies 455 


3-(5)em. Ovariaventrolanceolato. Stigma5-6mm. Carpella 
matura stipite ca. 0.5, caeteris ut praecedentis. Floret Junio.— 
British CotumBia: J. Macoun 77,395, Vancouver Island, 
Koksilah River, near Cowichau, July 30, 1908 (NY, TyPE; C, 
ISOTYPE); Newcombe 8, Pr. of Wales Isl., Karta Lake, 1901-02 
(F); Newcombe 416, Victoria, June 4, 1896 (F); Lyall, Vancouver 
Island (NY); J. Macoun, Vancouver Island, Nanaimo, June 10, 
1887 (US); Carter 157, Vancouver Island, Alberni, Roger Creek, 
creek-bottom, May, 1915 (G); Rosendahl 1988, Vancouver 
Island, Cameron River Valley, alt. 600 ft., June 28, 1907 (US); 
J. Macoun 22, Vancouver Island, borders of streams, July 10, 
1887 (G); C. B. Wood, Vancouver’s Island, 1859 (G); J. Macoun 
849, Sproat, woods, June 24, 1890 (C). 

101. T. occipenTALE Gray, var. PALOUENSE St. John, Fi. 
South. Wash. & Adj. Id. 158, 1937. 7. heterophyllum Nutt. ex 
Gray, Proc. Amer. Acad. 8: 372, 1873, ut synonymon dubium, 
nec 7’. heterophyllum Lej., Rev. Fl. Spa, 109, 1824, nec T. hetero- 
phyllum Schur ex Verh., Naturf. Ver. Bruenn, 15, 2: 20, 1877, nec 
T’. heterophyllum Turez. ex Ledeb., Fl. Ross. 1: 727, 1843. T. 
propinguum Greene, Fedde, Rep. Nov. Spec. 7: 254, 1909. 
T. rainierense St. John, Madrofio, 4: 114, fig. 1, 1937. Planta 


aspen groves, alt. 6000 ft., July 14, 1920 (CA, G); Piper 1468, 
Latah Co., Cedar Mountains, July 7 (G); Macbride 420, Owyhee 
Co., Silver City, slopes along streams, alt. 7000 ft., July 18, 1910 
(G). Wrommne: L. O. & R. P. Williams 3028, Big Horn Co., 
10-15 miles east of Kane, west slopes of the mountains, alt. 
8000 ft., June 19, 1936 (G); Goodding 1971, Carbon Co., Bridger 


456 Rhodora [DECEMBER 


Peak, moist timbered flats, Aug. 24, 1903 (G); L. 0. :& Bak, 
Williams 3639, Park Co., Beartooth Lake, in pine woods, alt. 
9000 ft., July ‘21, 1937 (G): idem 3562, Crazy Woman Creek, 
July 14” (G); E. 'B. & L. B. Payson 2999, Sublette Co., Gros 
Ventre Mountains, mg sre northeast of Bondurant, yao slopes, 
Aug. 13, 1922 (F, G, US); Williams & Pierson 652, Teton Co., 
vicinity ‘of Hoback Canyon, spring-bank, alt. 7000 ft., eee 19, 
1932 (CA, G). Nevapa: Nelson & Macbride 1936, Elko Co., 
Jarbridge, aspen copses, seo 7000 ft., July 6, 1912 (G, NY); 
Nelson & Macbride 2210, Elko Co., ‘ Mountain City, aspen 
copses, alt. 7000 ft., Aug. 14, 1912 (G, NY, US); Maguire & 
Plage one x nie along Ole Creek, woodlands, June 28, 1934 
(G). . S. Rose 36,476, Wallowa Co., 1 mi. s. Wallowa 
a alt. 4600 ft., July 18, i936 (CA); Thompson 13,340, Baker 

near Lornucopi ia, rocky slopes of Wallowa Mts., July 18, 
1988 (ANS, NY); Henderson 5154 & 5658, E. Grant Co., Austin 
Ranch, 1925 (CA, G); Nuttall, Columbia Woods (ANS, G, NY) 
with Thalictrum heterophyllum i in Nuttall’s handwriting). WasH- 
INGTON: Piper 2022, Mt. Rainier, rich meadows, alt. 6500 ft., 
Aug. 1-15, 1895 (G, isotype of T. rainierense); Piper, Walla 
Walla Co., Blue Mts., along streams in woods, July 17, 1896 
(G, ISOTYPE of oD: occidentale var. palouense) ; Thompson 6945, 
Okanogan Co., moist shaded slopes by road to Salmon Meadows, 
alt. 3500 ft. tome "35, 1931 (G); Thompson 77938, Chelan Co., 
Wenatchee Mts., below Stuart Pass, moist alder groves, alt. 
prea ons Sta 27-31, 1931 (ANS, G). British Cotumsia: J. 
M. M n 33,606, Tami i Hy Mt., Chilliwack Valley, alt. 5000 
ft., July 3 30, 1901 (G- ND, type of T. She nquum); Heacock 53, 
Emerald Lake, nakng he Path, alt. 4400 ft., June 29, 1904 (G, 
Y, US, p a or 7. propingwum); Shaw 970, in the Big 
Bend district, wien 118° 20’ W., 51° N., alpine meadow, alt. 
meet July 24, 1905 (G, shee NioCabe $08, aa River Road, 17 
north of Natal, edge of o of green spruce timber, 
Sept. 8, 1937 (UC); Raup é Abbe 3 3867, sige & Wicked R., near 
ye Peace, about 56° 4 N., 123° 39’ W., open woods, July 18, 
32(G). Fie. 101, a-e 


he type of T. viii Greene is not in the Herbarium of 
the Canaan Geological Survey as stated in the original descrip- 
tion; similarly with the types of T. tortuosum Greene, 7’. Mortont 
Greene and T. glaucodeum Greene. In his last letter to J. M. 
Macoun, Greene wrote: 

April 28, 1915. 

Another parcel of your Thalictrum went last night; only a small 

parcel, Uf shanks on Whdck T hast cues sake esate TEmMains. 


e588 ee ak ee 


2 Ae Oe Rice 


ae ee Ee 


1944] Boivin,—American Thalictra and their Allies 457 


That “small parcel” apparently was never sent, for 66 of the 
numbers of the Herbarium of the Geological Survey which are — 
represented in the herbarium at Notre-Dame University, are all 
missing in the National Herbarium of Canada. 


Subsectio Laminaria, subsect. nov. Plantae persaepius plus 
minusve stoloniferae. Carpella matura valde compressa vel 
laminaria, nervis rugosis, haud costata, vel, si costata, costis 
acutis. Crassitudo carpelli a dimidiis latitudinis ejus recedit. 
Species typica Thalictrum Fendleri Gray sit. 


a, Carpella haud reflexa, ovata ad lanceolata, viridia vel brun- 
nea, plus minusve pubescentia vel interdum glabra, nervis 
lateralibus raro ramosis et sinuosis, nunquam minute sinu- 
osis, nec anastomosantibus, reticulatis. Planta nunquam 
pruinosa nec caesia nec glauca. Cauli tamen interdum 
urpurascens. .. .b. : 
b. Nervi laterales 3 conspicui. Planta pubescens. ...... 102. 7. Fendieri. 
b, Nervus lateralis solitarius vel, si ternatus, medius con- 
spicuor et multo crassior quam duo alii. Plantae 
saepius glabrae. .. .c. 
¢. Carpella ventre ca. 2.5 mm. longo... .d. : Ste 
d. Nervi haud sinuosi T. Fendleri, var. Wrightit. 
d. Nervus ventralis, licet alii, valde sinuosi. : : 
04. T. Fendleri, var. sinuosum. 


ee ee '° 


c. Carpella ventre ca. 5 mm. longo. 
105. T. Fendleri, var. quadrinervatum. 
. pupae plura vel omnia reflexa, obovata vel semiobovata, 
plus minusve pruinosa glauca, nervis lateralibus saepius 
ramosis anastomosantibus reticulatis minute sinuosis. 
Planta glabra pruinosa et plus minusve glauca vel caesia. 
Ovaria ventre compresso orbiculari et nunquam rugoso. 
106. ZZ. polycarpum. 


Practically, 7’. polycarpum Wats. is always glabrous while 7. 
Fendleri Engelm. is always more or less pubescent, at least within 
the range of the former. Ovaries, fruits and under surface of 
upper leaves always show this character clearly. The fruits of 

ese two species are endlessly variable. Some of these varia- 
tions are more frequent in one part of the range, but they are 
liable to be found anywhere throughout the range and none of 
them is clearly cut from the other variations. However, three 
variations of 7. Fendleri have a geographic range of their own 
and outnumber the intermediate forms. 

102. T. Fenpiert E . ex Gray, Pl. Fendl. 5, 1849. T. 
Fendleri, var. ehicaineces Tel Piso. Bost Soc. Nat. Hist. 23: 
304, 1886. 7. platycarpum (Trel.) Greene, Pittonia, 1: 166, 
1888, nec. T. platycarpum Hook. f. & Th., Fl. Ind. 1: 13, 1855. 
T. hesperium Greene, Pittonia, 2: 24, 1889. polycarpum 


458 Rhodora [DECEMBER 


ness var. Maceo (Greene) Jepson, Fl. W. Midd. Cal. 202, 
T.0 m Greene, Fedde, Rep. Sp. Nov. 7: 254, 1909. 
r ious Teabere Fl. "Rocky Mts., 290, 1918, nec T’. stipi- 
tatum Rose, Contrib. U. S. Nat. Herb. 8: 28, 1903. TT. Fendleri, 
var. hesperium (Greene) plates Fl. Calif. 1: 530, 1921. Planta 
pubescens, raro subglabra vel glabra etiam, nunquam pruinosa 
nec caesia nec glauca, caulis tamen interdum purpurascens, 60— 
150 em. Radix plus minusve stolonifera. Sepala erosa, floris 
maris ovata vel elliptica 3-5 mm. longa, foeminei ovata vel 
rhomboidea vel late lanceolata (1.0)—1.5-(2.0) mm. longa. 
Filamenta 4.0-7.5 mm. lutea. Antherae oblongae vel lineares, 
luteae, 2.2-3.4 mm., acumine 0.1-0.8 mm. Stigma 1.5-4.0 mm. 
Ovaria dense viridia, ventro ovato ad lanceolato, saepius densis- 
sime pubescente. Carpella matura haud reflexa, ovata ad 
lanceolata, viridia vel brunnea, en minusve pubescentia vel 
ae glabra, stipite (0.1)-0. 5-(2 (2.0) mm., ventre 2.7-9.0 m 
longo, 1.8-4.5 mm. lato, nervis lateralibus raro ramosis et sinuo- 
sis, nunquam minute sinuosis, nec anastomosantibus, nec 
reticulatis, nervo ventrali hice quam dorsali. Floret ab 
Aprili ad Septembrem.—Texas: Ferris & Duncan 2569, Jeff 
Davis Co., Davis Mts., hewccai Peak, July 9-12, 1921 (CA, 
M); Havard. 138, Jeff Davis Co., Limpia Mts., July, 1883 (G); 
Moore & Steyermark 3566, Culbertson Co., Guadalupe Moun- 
tains, McKittrick pvnhee’ ’ shaded rocky woodland, alt. 2000 m., 
July 22, 1931 (G, M). Wyromine: Tweedy 4244, Carbon Co., 
forks Battle Creek, alt. 7200 ft., Aug. 15, 1901 (US, type of T. 
omissum; NY, i sotype); A. Nelson 766 0, "Alban ny Co., Tie City, 
in a canon, Jay 20, 1900 (G); Payson & Armstrong 3608, Lincoln 
Co. , Alpine, on the Snake River near the Idaho boundary, Wolf 
Creek, — groves, July 25, 1923 (G). Cotorapo: Hall & 
Harbour 8, lat. 39 °—41°, 1862 (BC, G, M); E. L. Greene 598, 
Golden City &e., 1871 (G, poate of . Fendleri var. platy- 
carpum); F. E. & S. 8. Clements 248, El Paso Co., Minnehaha, 
alt. 2600 m., 1 julii, Sap (NY, type of 7. pr noes G, toot peel 
Baker, Earle & Trac 343, Montezuma Co., W. La Plata Mts., 
Chicken Creek, scaerien, alt. 9000 ft., July 6, 1898 (G); "Rollins 
1808, Las Animas Co., 26 —- northwest of Trinidad, 2 miles 
north of the Pe rgatore River, dry hillside, stems single, alt. 
7500 ft., July 3, 1937 (G); romero Clear Creek Co., Brookvale, 
road Creek Canyon, June 17, 1918 (G); Hermann 5399, Garfield 
, Trapper’s Lake, n. shore, open spruce-fir grove, a alt. at ae 
ine 29, 1933 (G). Uran: M. E. Jones 1194, Salt Lake C 
Wahsatch Mts., Alta, alt. 10,000 ft., Aug. 5, 1879 (CA); Good. 
man & Hitchcock 1891, San Juan Co. , Abajo Mts., n. slope, ’ beside 
small stream, alt. 8500-11,000 ft. , July 1-2, 1930 (C A): E. B. & 
be BB. Payson 4883, iia Co., foothills of Uinta Mts., near 
Mill Creek, dry clay slope, alt. 8200 ft. , July 4, 1926 (G, US); 


1944] Boivin,—American Thalictra and their Allies 459 


McKelvey 4204, Washington Co., Zion Natl. Park, Zion Cajon, 
alt. 4000-5000 ft., May 7, 1934 (G). Nevapa: Clokey 5462, 
Clark Co., Lee Cafion, gravelly flat, with Pinus scopulorum and 
Populus aurea, alt. 2670 m., Aug. 1, , G, UC); M. E. 
Jones, Humboldt Co., East Humboldt Mts., alt. 10,000 ft., Aug. 
13, 1897 (UC); P. B. Kennedy 1923, Washoe Co., Hunter Creek, 
elev. 6000 ft., Aug. 2, 1912 (CA, G); Baker 1823, Ormsby Co., 
head of Fall Creek, alt. 2460 m., July 15, 1902 (G, NY). New 
Mexico: Standley 4257, Pecos River National Forest, Windsor 
Creek, alt. ca. 8600 ft., July 8, 1908 (G, M); Wright 1851 [Wright’s 
mss.: 151, Grant Co., hillsides of Coppermine Creek, 2-4 ft. 
tall, Aug. 4] (G, NY, US); Metcalfe 248, Socorro Co., Mogollon 
Mts., on Mogollon Creek, alt. ca. 8000 ft., July 17, 1903 (G, M, 
UC, C-UC); Wooton 228, Lincoln Co., White Mts., alt. 6300 
ft., July 28, 1897 (M, UC); Fendler 13, 1847 [Fendler’s mss.: 13, 

anta Fe, Creek-valley, shady places, margin of irrigation ditches 
at the foot of perpendic. rocks, 13th June-Ist July in flower, 
19th July in fruit. Flowers dioecious] (G, TYPE and ISOTYPES 
of T. Fendleri; ANS, M, NY, 1soryrss); J. M. Bigelow 963, San 
Antonita, Camp B, mt. arroyos, Oct. 9, 1853 (G, NY, US, para- 
types of T. Fendleri var.? polycarpum). Arizona: Blumer 3309, 
Rincon a eames Spud Ranch, rocky places, alt. 2260 m., Aug. 

M 


US); Munz & Johnston 8696, San Bernardino Co., 1 mile south 
of Oak Glen, alt. 4500 ft., July 17, 1924 (G); Lemmon, San 
Bernardino Co., woods near Grayback, July, 1879 (G, paratype 
of T. Fendleri var. platycarpum); A. Gray, Santa Barbara, Feb.— 
ay, 1885 (G, paratype of 7. Fendleri var. platycarpum) ; 

oulter (G, US, paratypes of T. Fendleri var.? polycarpum). 
OREGON: Cusick 2086a, Eastern Oregon, 1898 (G); Eastwood & 

owell 1614, Lane Co., McKenzie Highway, near Blue River, 


460 Rhodora [DECEMBER 


Apr. 17, 1984 (CA); Constance, Lane Co., Eugene, Young’s 
Grove, Apr. 12, 1924 (G); Henderson 9057, Harney Co., Stein 
Mts., above Fish Lake, in aspen woods, alt. 6500 ft., July 20, 
1927 (CA); M. E. Peck, 14,202, Stein Mts., above Alberson, 
along stream, alt. 7000 ft., July 4, 1925 (ANS). Mexico, BAJA 
CALIFORNIA: Orcutt, n. Lower Cal., Topo Cafion Mts., July 30, 
1883 (F). Fre. 102, a-e. 


Trelease did not authenticate every Thalictrum he saw at the 
Gray Herbarium where he prepared his monograph of the genus; 
only four herbarium sheets bear his identification. These are 
the specimens he cited as T. venulosum. On about half a dozen 
other sheets are to be found pockets containing a smaller pocket 
on which he briefly copied out the label of the specimen, probably 
in order to be able to replace those pockets where they belong 
after the drawings were made, for these pockets contain dissected 
fruits, presumably the very ones from which the illustrations of 
his monograph were drawn. 

It seems that, at the Gray Herbarium, Dr. B. L. Robinson 
went over Trelease’s work, wrote the new species-covers needed 
and pasted all the revision labels as Trelease would have done, 
for all the names written by Robinson on sheets or covers in the 
genus Thalictrum are in perfect accordance with Trelease’s treat- 
ment of that genus. Nearly all of the Thalictra bearing a revision 
label in Robinson’s handwriting were collected prior to 1886. 
Very few indeed of the specimens collected after that date were 
revised by Robinson. The later specimens were obviously not 
accessible to Trelease for his monograph. Furthermore, a few 
specimens might have been collected prior to 1886, but mounted 
and revised by Robinson only later on. These can not be dis- 
tinguished from those Trelease had on hand in 1886, and they 
introduce some uncertainty as to the correspondence of Robin- 
son’s labels with Trelease’s opinions in the genus. However, 
in the absence of any better criterion, the revision of the former 
has been considered as representing the opinion of the latter, 
provided the specimens were collected prior to 1886 in a locality 
within the range given by Trelease in his monograph of that year. 

There is no authenticated specimen of 7. Fendleri, var. platy- 
carpum Trel. except for one specimen from the Smithsonian 
Institution which bears the following note in Trelease’s hand- 
writing: “7. Fendleri v. platycaroum or nearly that’, but six 


1944] Boivin,—American Thalictra and their Allies 461 


specimens at the Gray Herbarium bear a revision-label in 
Robinson’s handwriting, stating that these are J. Fendleri 
Englm., var. platycarpum Trel. All six are from California and 
were collected in or before 1885. In view of this variety having 
been published in 1886 with California given as its range, it 
seems likely that these labels represent Trelease’s opinion rather . 
than Robinson’s. The Tyrr was selected from among these six 
specimens. It bears in a pocket a fruit dissected by Trelease 
and this apparently is the very fruit from which he made the 
drawing he published to illustrate 7. Fendleri var. platycarpum 
in the paper where he published that variety. A drawing was 
made of this fruit (our fig. 102d) and one can not fail to note some 
similarity of pattern of this drawing to Trelease’s drawing. Such 
sinuose lateral nerves are rather exceptional in 7. Fendleri. 

103. T. FenpLERI Engelm. var. WRIGHTII Sao Trel., Proc. 
Bost. Soc. Nat. Hist. 23: 304, 1886. T. i Gray, Pl 
Wright. 2: 7 (269). 1853. Planta saepius ga a omnibus 
partibus minor, haud stolonifera, (25)—50-(90) cm. Stigma ca. 

mm. Carpella matura ovata ca. 2.5 mm. longa, nervis haud 
sinuosis, nervo dorsali convexo, ventrali convexiore quam dorsali. 
Nervus lateralis singulus, vel, si ternati, nervus medius conspicuor 
et multo crassior quam duobus aliis intocmneare Floret Au- 
gusto et interdum Julio vel Septembri.—Arizon . E. Jones, 
Santa Rita Mts., alt. 4500 ft., hte. 24, 1903 (CA A. UC); Darrow 
& Arnold, Santa Rita Mts., alt. 5500 ft., Aug. 23, 1936 (UC); 
Harrison & Kearney 8907, Santa Rita Mts., ae 20, 1932 (US). 
Mexico, CHIHUAHUA: Barlow, Sierra Madre ridge besos Rio 


Ww f Min ca, dry 
rocky 7 ay herb, alt. 2900-2400 m., " Gaps, 16-17, 1934 “(ANS): 
LeSueur 1211, Chuchichupa, Aug, 1936 (F); Pringle 1131, 
ie Mts., alt. 7300 ft. Bree 


fete 
oe 
ee 
ae 
Ss 
3 
oS 
Bo 


_ NY, 
Sonora: Hartman 1 21, Los Pinitos, alt. 6100 f , Oct. 12, 1890 
(G, NY, US); Wright 834, mountain ravine at sacita Cruz, Sept., 
1851 (G, TrPE; ANS, G, M ky UC, 1sorypEs); Mearns 1605, 
summit of San Jose Mes Aug. 3 , 1893 (US); S. ’S. White 3081, 
e las Estacas, July 30, 1940 (G). sInALoa: Penne li 


‘ along 

on pineland, alt. 1800-1900 m., Aug. 29-30, 1935 (ANS); 

Gentry 6266, Sierra Surotato, Ocurahui, Pine Forest area, steep 

ft. ist shady canyon slope with mixed dominants, alt. 6000-7000 
» Sept. 1-10, 1941 (ANS, M, NY). Fie. 103. 


462 Rhodora [DECEMBER 


The number 834 in the manuscript of Wright is an Artemisia. 
But it is known that Gray changed Wright’s collection-numbers. 
The manuscript of the latter enumerates three collections of 
Thalictrum: 

178. Thalictrum, Cummings’ Creek, May 10, 1849. 
151. eager rum, a of Coppermine Creek, 2-4 ft. tall, 
ug. 4, 1 
639. Thatictrum, "Mowxtaia ravines at Santa Cruz, Sept. 
23, 1851.5 

At the Gray Herbarium there are three herbarium sheets of 
Wright’s collections. The labels read as follows (Gray’s hand- 
writing is in italics, the printed caption in roman): 

First specimen: 


=) 


le, 
Texas, Mr. Charles Wright. 


Second specimen: 


No. 833 C. Wright, np N. Mex. 1851. 
Thalictrum Fendleri, E ngelm 


Third specimen: 
No. 834 C. Wright, vip N. Mex. 1851. 
tit n. 


Thalictrum Wri 
Santa Cruz, Sonora. 


Comparing these with what Gray published in Pl. Wright. 2: 
7 (269), 1853, it seems clear that these three specimens correspond 
respectively to Wright’s field numbers 178, 151, 639. 


104. T. Fenpieri Engelm., var. sinuosum, var. nov. Planta 
similis praecedenti, fructubus tamen nervis sinuosis. Floret 
Augusti mense.—Merxico, cHrHuAHUA: Goldman 126, - 
Parral, alt. about 6500 ft., Sept. 20, 1898 (G, US). puRAN 
Pringle 18,701, Sandia Station, Oct. 12, 1905 (G, US); Pennell 
18,448, north of Cueva, Metates, ravine at warcealh alt. 2600- 


TYPES). SAN LUIS POTOSf: Scha er 26, ex sini“ San 
Potosf, in sylvis montibus San Miguelito, 1876 (G). Fic. 104. 


to a manuscript at the Gray Herbarium Rema ts I. M., Field Notes 
of Foon Wright, 1940), this ri ae Se County, Texas 
*Eodem. Grant County, 
3’ Eodem. exico 


1944] Boivin,—American Thalictra and their Allies 463 


105. T. FenpLert Engelm., var. quadrinervatum, var. nov. 
Planta pubescens verosimiliter metrum superans. Stigma ca. 

mm. Carpella matura ventro ca. 5 mm. longo et ca. 2.5 mm. 
lato, nervis rectis vel sinuosis, ventrali convexiore quam dorsali. 
Nervi laterales carpelli vel tres vel singulus in utroque latere. 
Si tres, intermedius quam duo alii multo conspicuor et crassior 
est. s ignoti—MEeExiIco, BAJA CALIFORNIA: Wiggins & 
Demaree 4941, Sierra San Pedro Martir, La Encantada, margins 
of meadow and adjacent slopes, alt. 2200 m., Sept. 20, 1930 (G, 
Type; F, NY, UC, , IsoTYPES). Fie. 105. 

106. T. potycarpum (Torr.) Watson, Proc. Am. Acad. 14: 
288, 1879, nec T. polycarpum Loret, Bull. Soc. Bot. Fr. 6: 16, 
1859, ut nomen provisorium alternativumque editum. T. 
Fendleri Engelm., var.? polycarpum Torr., Pac. R. R. Rep. 4: 
5 (61), 1857. 7. caesium Greene, Fl. Franc. 3: 309, 1892, nec 
T. caesium Blocki, Oest. Bot. Zeitschr. 37: 286, 1888. T. 
bernardinum Greene, Fedde, Rep. Nov. Sp. 7: 252, 1909. T. 
campylopodum Greene, 1. c. 253, 1909. 7. coreospermum Greene, 
l.c. 1909. 1. lentiginosum Greene, 1. c. 1909. T. papyraceum 
Greene, 1. c. 1909. 7. ametrum Greene, Muhlenbergia, 5: 129, 
1909. 1. latiusculum Greene, 1. c. 130, 1909. 7. magarum 
Greene, 1. c. 1909. 7. mendocinum Greene, 1. c. 129, 1909. T. 
letocarpum Greene, 1. c. 130, 1909, sicut nomen provisorium et 
alternativum praecedenti editum, nec 7. leiocarpum Fries, 
Linnaea, 29: 731, 1857 sive 1858. TT. polycarpum Wats. var. 
caesium (Greene) Jepson, Fl. Calif. 1: 530, 1921. Planta 
stolonifera omnino glabra, saepissime cauli, ramis, ramulis, 
fructubusque pruinosis vel glaucis vel caesiis, 50-150 em. Se- 
pala floris maris elliptica 3-5 mm. longa, foeminei orbicularia vel 
ovata vel saepius late ovata 1.0-2.5 mm. longa. Filamenta 5-6 

Antherae 2-4 mm., acumine 0.1-0.5 mm. Stigma 1.5-4.0 

mm., filiforme. Carpella matura reflexa et valde compressa, 
hunquam costata, stipite (0.1)—0.2-(1.0) mm., ventre oborbicu- 
lari vel obovato-lanceolato vel saepius obovato 4-6-(8) mm. 
longo et (2.2)-3.5-(4.5) mm. lato, nervis lateralibus valde sinuo- 
Sis ramosis anastomosantibusque, nervo ventrali interdum gib- 
boso et semper quam dorsali convexiore. Floret Martio Aprili 
Majo et Junio.—CatrorNnia: Chesnut 33, Mendocino Co., 
Round Valley, alt. 440 m., May 20-June 20, 1898 (US, type of 
a of 


464 Rhodora [DECEMBER 


coreospermum); Alderson, San Diego Co., Witch Creek, April, 
1894 (G-ND, type of 7. magarum; G, photo of type); Greene, 
Chico, June, 1889 (G-ND, type of 7. caesium; G, photograph of 
the type; UC, NY, isotypes); L. E. Smith 306, Shasta Co., Pitt 
River, May 28, 1913 (CA); Bidwell, Chico, Apr., 1879 (G); idem, 
May, 1878 (G); Mrs. R. M. Austin, Plumas Co., 1877 (G); G. BR 
Vasey, Sancelito, 1876 (G) [These last four specimens were 
authenticated by Watson]; J. M. Bigelow, Napa Valley, 1853- 
1854 (G, type of T. Fendleri var.? polycarpum; NY, 1soTyPe). 
Orecon: J. Howell, Tualitin Plains, damp shady woods, along 
creek, July, 1877 (G); J. Howell, Sauvie’s Island, July 1877 (F, 
type of 7. papyraceum); T. Howell, along wooded streams, June 
1881 (ANS, F, isotypes of T. campylopodum); J. C. Nelson 1118, 
Polk Co., W. Salem, woods near river, May 8, 1817 (G); Hast- 
wood & Howell 1487, Douglas Co., 4 miles n. of Oakland, Apr. 13, 
1934 (CA); J. C. Nelson 2701, Marion Co., State School, low 
ground, June 21, 1919 (G); 7. Howell, Washington Co., by creeks, 
May, 1880 (F); idem, May, 1881 (ANS). Fie. 106, a-c. 

The validity of 7. polycarpum Wats. against the earlier T. 
polycarpum Loret has been argued pro and con many times. 
First, Greene noticed that there were two species called 7’. poly- 
carpum and proposed the name 7. ametrum to supersede T. 
polycarpum Wats. See Muhlenbergia, 5: 129, 1909. 

Second, L. C. Wheeler pointed out that T. polycarpum Loret, 
being a “nomen provisorium’”, was not validly published and 
thus could not invalidate a later homonym. Hence T. poly- 
carpum Wats. was the correct name for the plant discussed. See 
Ruovora, 40: 319, 1938. Third, Leon Croizat in Madroijio, 7: 
1, 1943, in an article which I find much harder to follow through 
than Loret’s “stiff french”, contested the following points: 1, 
that it has not been made clear yet which of Loret’s names, 7’. 
polycarpum and T. multiflorum, is a “nomen provisorium”. 
that T. polycarpwm Loret was published as a synonym of T. 
multiflorum Loret, hence it is invalid (Amsterdam Code, Art. 40). 
3, that T. polycarpum Loret is illegitimate, invalid, has been 
treated as a synonym on taxonomic grounds, and is an earlier 
homonym. ‘Thus, according to Article 61, it renders illegitimate 
T. polycarpum Wats. 

To this one may answer: 1, that both T. multiflorum Loret and 
T. polycarpum Loret are “nomina provisoria”. If not, what 1s 
the meaning of “nomen provisorium”? There does not exist 
yet any officially published definition of that term, but in the 


1944] Boivin,—American Thalictra and their Allies 465 


meantime we will take it in its common, everyday sense, hoping 
that it will not turn out that the 1935 International Congress 
had some esoteric meaning in mind. 2, that T. polycarpum 
Loret was not published as a synonym of T. multiflorum Loret, 
but as a “nomen alternativum”. As ruled by the 1935 Congress, 
when names are published as ‘‘nomina alternativa” that does not 
render them invalid, but they are invalid if published as ‘nomina 
provisoria”. Hence both T. polycarpum Loret and T. multi- 
florum Loret are not validly published. See Journ. Bot. 74: 75, 
1936. 3, that Article 61 mentions 4 conditions as necessary, 
that a homonym might be invalidated by another homonym, 
namely, that a homonym must be “previously and validly pub- 
lished for a group of the same rank based on a different type 
[italics mine]’. 7. polycarpum Loret does not fulfill the second 
of those conditions. Hence it cannot render T. polycarpum 
Wats. illegitimate. Article 61 speaks of “illegitimate names” 
and of names “treated as synonyms on taxonomic ground” only 
to specify that those two qualities are of no effect in the solution 
of the problem. 

4 and 5, Cronquist and Weatherby, in two different articles 
published on the same page, both pointed out that “nomina 
provisoria” have been ruled out as not being validly published, 
and that the other half of Article 61, cited by Croizat, explicitly 
states that an earlier homonym must have been validly published 
in order to invalidate a later homonym. Now, Croizat himself 
admits that 7. polycarpum Loret was not validly published, 
hence there is no reason why we should not keep T. polycarpum 
Wats. (Madrofio, 7: 83, 1943). 

The type of T. Fendleri Engelm., var. ? polycarpum Torr., is 
evidently not at New York, but at the Gray Herbarium. The 

of the original description suggests that Coulter’s plant 
should not be selected as a type. It is represented by two sheets, 
one at the Gray, the other at the National Herbarium, and both 
were originally labelled 7. dioicum L. in John Ball’s handwriting. 
The specimen at the Gray bears the following successive anno- 

tations in Gray’s handwriting: 
> 


Fendleri 
T. polycarpum Torr. = Fendleri var. 
?. polycarpum 


466 Rhodora [DECEMBER 


Both specimens are densely pubescent individuals of T. 
Fendleri and they agree only in part with the original description 
of T. Fendleri, var. ? polycarpum Torr. 

J. M. Bigelow’s collection from San Antonita, New Mexico, is 
represented at the Gray Herbarium, at the New York Botanical 
Garden and at the Smithsonian Institution. All three are good 
ordinary 7. Fendleri with a large, open and compound panicle 
and with pubescent fruits and leaflets. Although cited by 
Torrey, this collection does not agree at all with his description. 
The specimen at New York is labelled Thalictrum Fendleri 
Engelm. in Gray’s handwriting and so is the specimen at the 
Gray Herbarium, but to this one Gray himself added later 
fere var. Wrightii. The third specimen is labelled Thalictrum 
Fendleri Engelm. in Torrey’s handwriting. The New York sheet 
has the original label: Camp B, San Antonita, Octo. 9, 1853, 963 
Thalictrum sp., mt arroyos, JMB. 

At the Smithsonian Institution there is also a specimen of T. 
polycarpum which was first labelled Thalictrum dioicum L. ?, but 
Torrey changed it later to polycarpum. It was collected by E 
Samuels in Sonoma Co., California. Although authenticated and 
agreeing with the original description, this specimen was not cited. 

J. M. Bigelow’s collection in Napa Valley is represented both 
at the Gray Herbarium and at the New York Botanical Garden. 
The New York specimen is identified Thalictrum Fendlert 
Engelm. in Torrey’s handwriting. It is an immature pistillate 
plant. The label of the specimen at the Gray Herbarium reads 
as follows: 


carpum 
Napa Valley Calif. n. sp. Torr. 
Bigelow eal 


Gray’s handwriting is indicated by italics, Torrey’s by bold- 
face characters. This latter specimen is made up of a staminate 
inflorescence and a complete pistillate specimen with immature 
fruits. This Bigelow collection is very clearly T. polycarpum 
(Torr.) Wats. and agrees perfectly well with the original descrip- 
tion of T. Fendleri Engelm. var.? polycarpum Torr. The speci- 
men to be selected as a TYPE is evidently J. M. Bigelow’s speci- 
men at the Gray Herbarium, from the Napa Valley collection. 


1944] Boivin,—American Thalictra and their Allies 467 


92a92b 


Ze 


iW 
106c > 107b | 
= 107a 1076 Na 
\ 


i" Q | 


aay." 108c 109a 


0b 


10la 10lc 102d Sa 


Derams or Frowers or Fruits or THALICTRUM, all X 4; except 10le, 
X \, and 110e and 113a, these X 3. 


(For explanation see end of paper.) 


468 Rhodora [DECEMBER 


rf 
It is the only specimen which is at the same time cited, authenti- 
cated and agreeing well with the original description. It is also 
the most complete specimen and a well preserved one. This 
problem had already been outlined by L. C. Wheeler in Ruopora, 
40: 319, 1938, but no definite conclusion-was given. 

When Watson published his 7’. polycarpum he gave T’. Fendlerz, 
var.? polycarpum as a synonym “in part”, without specifying 
which of Torrey’s specimens were included. But the description 
of Watson eliminates both Coulter’s and J. M. Bigelow’s New 
Mexican collections; only the Napa Valley specimen agrees with 
the description. Thus 7. polycarpum must be considered as a 
new status for 7’. Fendleri Engelm. var. ? polycarpum Torr. “ 
part including the type specimen” and there is no need to select a 
new type for T. polycarpum, but we must write 7. polycarpum 
(Torr.) Wats. 

L. C. Wheeler has expressed a different opinion in RHopora, 
40: 317, 1938. To him it is not clear what of Torrey’s variety 
Watson included in his species. He thus naturally comes to the 
conclusion that one must write 7’. polycarpum Wats. and must 
select the type of that species independently from the type of 
Torrey’s variety. To this we may say that Torrey’s variety 
included two different species but his and Watson’s descriptions 
agree well only with the Napa Valley collection. Watson had 
all three syntypes at hand and could easily settle that point. 

Anyhow, there are at the Gray Herbarium 5 collections of T. 
polycarpum (Torr.) Watson authenticated by Watson himself. 
All agree well with the description, are within the range given, 
and were collected from 1876 to 1879. As a matter of fact, all 
5 labels are entirely in Watson’s handwriting. If one should 
follow Wheeler’s opinion, one of those 5 specimens should be 
taken as the type. There is ample choice: one specimen is made 
up of 2 staminate inflorescences, another of a pistillate inflores- 
cence, a third one of two pistillate and one staminate inflores- 
cence, a fourth is the upper half of a plant with fully mature 
fruits, and the fifth one, the best specimen, a nearly complete 
plant with also fully mature fruits. The latter is the only spect- 
men cited by Watson. Its label reads as follows: 

Thalictrum polycarpu 
(T. Fendlert, var. saree ey Torr.) 


1944] Boivin,—American Thalictra and their Allies 469 


Damp shady woods along creeks, 
near Tualitin Plains, Oregon 
J. Howell, July 1877. 


Oddly enough, Wheeler selected J. M. Bigelow’s Napa Valley 
collection as the type of T. polycarpum, the very type of T. 
Fendleri Engelm. var. ? polycarpum Torr. I see no reason to 
write T. polycarpum Wats. rather than T. polycarpum (Torr.) 
Wats., if the type of both units is the same specimen. 


Sectio Leucocoma (Greene), stat. nov. Subg. or Sect. Leu- 
cocoma Greene, Leaflets, 2: 54 and 56, 1910, subaivisio on 3 


. me 
coma Rydb. Fl. Rocky Mts. 108, 1917. Plantae haud stoloni- 
ferae, sed radicibus fibrosis et numerosis. Flores polygami, sexu 
variabiles modo mirabili. Sepala erosa maris majora. Fila- 

menta alba in nonnullis clavata. Antherae fere ovoideae ad 
tins linearee: pallido-fuscae, apice truncatae vel acumine 
brevi. Stigma breve plus minusve bialatum. Carpella p ariete 


This section is vophuestad from Labrador to British Columbia 
and also in Newfoundland, Saint-Pierre et Miquelon and in the 
United States except in Mississippi, California, Utah and 
Nevada. 

A key to the species of the section is not a workable one if it 
takes into account the full variation of each species. With 
more than a thousand herbarium sheets at hand one finds that 
occasional specimens will agree with the species to which they 
belong but for one character. To take such specimens into ac- 
count would render the description so full of perhaps, sometimes, | 
rarely, or, even, that they would not be descriptions any more. 
The following keys and descriptions are built upon specimens 
which I consider typical. 


CLAVIS AD FLORES MASCULOS FERENTIA 


a. Pubescentia pilorum og hag vel rarissime deest; a 
therae 1.6-2.8 mm.; foliolae coriaceae et margine 


minusve revolate.. 6s cis so a ea ts 107. TP, revolutum, 
* -s atia vel dest vl piioram uniseriato Soe 
& Antherne 0.5-1.4 mm.; filamenta apice dilatata sub an- 
eris co 


eee ee 


470 Rhodora [DECEMBER 


c. F Cretia gt merit et pe 4 integrae 0.5-1.5 cm. longae. 
1 


brae. oe 3.0-5.0 mm..... d. 
d. tae oa prelate ees. 4s. 108. 7. macrostylum. 
Filamenta debilia. Planta reclinans........109. T. Fabeatundies, 
c. Foliolae fere omnes apice trilobatae 1.5-4.0 c m. longae. 
Plantae — pubescentes. Wilaneate 3.5-8.0 
MM, . 2. e. 
e. Antherae ca. 1 mm BP pis gracillimi. Inflor 
centia cuibalate ilamenta 3.5-5.0 mm. .110. ks polygamum. 
e. puiberee ca. 1.2 mm.; unculi crassiusculi; 


.0-8.0 mm.; gp omnia subco: 
1 a. Berean ery fal hebecarpum. 
b. Antherae 1.5-3.2 mm..... 


f. 
J ae 1.5-2.5 mm. Plantae pubescentes. .. .g. 
. Antherae 1.5-2.0 mm. Titonmentia | pice rotunda. 
ygamum, var. intermedium. 
g. Antherae 1.8-2.2 mm. Planta fate ata 

ee et margine revoluto. amebicooas Peer 
Pes Se OLIN CORN US Eb de ees T. dasycarpum. 

> Antheras 2.2-3.2 mm. Plantae ee ius ae 
114. TT. dasycarpum, var. hypoglaucum. 


CLAVIS AD FLORES FOEMINEAS FERENTIA 
a, Foliolae glabrae saepe integrae 0.5-1.5 em. longae. . . .b. 
—) 1.0 (-1.5) 


b. Stigma (0.7- mm. Foliolae hee mits integrae 
saepius apice chip” cy has coriaceae...... 7. macrostylum 
b. Stigma 1 liolae fere omnes integrae, saepius 
ee bo a a ee Soe: ve T. subrotundum 


. Inflorescentia panic ata apice acuta.... 
e. Foliolae coriaceae pubescentes. ee ata’ 2-3 mm 

113. “|. dasycarpum. 

e. Foliolae saepius membranaceae. —— .5-5.0 mm 
114. T. dasycarpum, var. penance 

d. Inflorescentia apice rotunda vel " subcorymbosa ape 

f. Inflorescentia subcorymbosa, pedunculis cr. ante 

Lit... ZT. aplninaen: vax hebecarpum. 
f. Inflorescentia paniculata................... 430, 2 emcee? 
112. 7. polygamum, var. intermedium. 


CLAVIS AD FRUCTUS FERENTIA 


a. — —. — bi tiahen ‘Gaim nervis valde gw 
fa) rae et inte ti 
pee ve Res jena egies htnnen, IO T subrotundum. 


‘ ng omnia SMe Lae Receptaculum ad basem 
capitis fructuum. .. . 

c a apice aeuininata + incurvata nervis valde 

sinuosis; stigma ca. 1.0 mm.; foliolae glabrae, inferne 
albicantes, ca. 1 cm. longae................ 108. 1’. macrostylum. 

c. Stigma 2-3: mm. Plamtas de dense pubescentes et ase: 
Ch. 2db O08, JONG a oe 113. 7. dasycarpum. 

b. Receptaculum ad centrum capitis fructuum. Carpella 


se ee 


1944] Boivin,—American Thalictra and their Allies 471 


d. Bybeeceate pilorum rigidorum capitatorum, rarissime 
dee Foliolae coriaceae margine valde revoluto 
107. a revolutum. 
d, Pubescentia saepius adest pilorum ee et - 
seriatorum. Foliolae saepius membranaceae.... 
e. Inflorescentia subco. osa pedunculis paced eae 
Saas ventre saepius oblanceolato 
111. T. polygamum, var. hebecarpum. 


anuenios quam dors: 
. T. dasycarpum, var. hypoglaucum. 
4; — carpelli ovatum vel obo vatum vel ellipsoi- 
deum vel lanceolatum, stipite 0.2-0.5 (-1.0) mm 
110. . ?. Sac 
112. T. polygamum, var. intermedi 

107. T. REvoLuTum DC. Syst. 1: 173, 1817, nec T. revolutum 
ong Oestr. Bot. eae 23: 254, 1873. T. revolutum DC., 


Risiohen Willd. ex ee a Bull. Soe. Bot. le ve 253, 1885, 
. oh depts on ex Trel., Proc. Bost. Soc. Nat. Hist. 
23: 3 , ex a dette ae Cornuti L., var. macrostylum 
Shuttly. j in Binal & Heller, Mem. Torr. Bot. Club, 3: 8, 1892, 
ut synonymon editum. T. Cornuti L., var. brevifolium Rugel in 
Small & Heller, 1. c. 9, 1892, nomen ut synonymon editum nec 
T. Cornuti L., var. brevifolium Shuttlw. in sae Syn. Fl. North 
Am. 1, part 1: 17, 1895. 7. amabile Greene, Am. Midl. Nat. 
2: 294, 1912. 1. revolutum DC., f. glabra isicl Pennell, Bartonia, 
12: 12, 1931. Planta circa metralis, fere semper pubescens per 
foliolas inferne, sepala, pedunculos et ovaria. Aliter semper 
glabra. Foliolae coriaceae margine valde revoluto, saepius 
obovatae, apice trilobatae, lobis saepius rotundis. Sepala ovata 
vel oblanceolata 3-4 mm. longa, floris foeminei 2.0-3.5 mm. 
onga. Filamenta debilia paululum apice incrassata, sub ee 


oblongo-lanceolatae ad oblongo-lineares 1.7-2.8 mm., acumine 
0.2-0.5 mm. Stigma 2.0-3.5 mm. alis saepius conspicuis et pilis 

minutissimis, in nae saepius incurvatum. Carpella matura 
stipite 0.2-0.5 mm., ventre ellipsoideo_ad lanceolato. Recepta- 


Julio.—Quesec: J. Macoun 72,578, our Co., Percé, Aug. 30, 
1907 (C, G).. ‘ah aa caetoen: Fernald & Weat herby 16,807, 
Barnstable Co., Falmouth, roadsides in Aud north of Wood’s 


Middlesex Co., Stoneham 
rocky woods, June 18, 1894 (NE); W. Boott, south end of Mystic 


Pond, wood, daly, 1869 (G); Pease 3782, Essex Co., N. Andover, 
(NE Ru ODE ISLAND: 


). 
J. F. Collins, Pevidanes. n. of Cat Swamp, June 26, 1892 (G); 


472 Rhodora {[DEcEMBEK 


Lownes & Collins, Washington Co., South Kingston, July 11, 
1930 (NE); Fernald 9518, Providence Co. .. Lincoln, Wilbur 
Crossing, dry thicket, July 17, 1913 (NE). CoNnNecticuT: 
vee by 6779, Tolla nd Co. sy Somers, roadside thicket, June 22, 
5 (NE); Bi issell §4, Hartford Co., Southington, dry rocky 
ok frequent, June 15, July 10, July 30, 1893 (NE); Wood- 
ward, Franklin, dry bank, June 24, 1911 (G). New Yorx: Muh- 
lenberg 795 (ANS): ; Eames, Randolph & beg dein 12, — Ontario 
o., Phelps-Waterloo townline, Sept. 6, 1919 (G); B urnham, 
Warren Co., Lake George village, Sept. 3, 1897 (G); Bi 
25,018, Albany Co., so. of Kamer, sand plains, Aug. 24, 1937 
(NY). New JERSEY: Austin, Closter, June 5-15 (G, ae of 
T. purpurascens var. ceriferum) ; Austin, ( Closter, 1866 (G, para- 
ype of 7. purpurascens var. ceriferum); A ustin, Ne. New J ersey, 
uplands (G, paratype of 7. purpurascens var. ceriferum) ; Austin, 
Palisades, 1858 (G, F); Austin, Closter, June, 1865 (F, paratype 
of T. purpurascens var. ceriferum). PENNSYLVANIA: Pennell 
8956, Chester Co., Nottingham, serpentine barren, Sept. 14, 
1916 (ANS, paratype of 7. revolutum, f. deviate Small, Perry 
Co., vicinity of Marysville, June 25, 1888 (F); Schweinitz, 
Bethlehem (ANS); Meredith, Bucks ond e 14 miles west of 
Union School House, open hedge-row, May 30, 1921 (G); Muhlen- 
berg 598 (Willdenow Herb., paratype of 7’. polygamum 2) DEL- 
AWARE: Commons, near Centreville, banks of streams, July 5, 1872 
(ANS). Manryzann: J. D. bbe Garrett Co., Oakland, copses, 
borders of glades, July 13, 83 (G). DistTRIctT OF CoLuMBIA: 
Steele, Chain Bridge flats, ‘Tuly 4, 1904 (US); L. F. Ward, June 
23, 1878 (F, NY). West Vircinia: Miilspaugh 336, 1890-1899 
(F); Gilbert 482, Cabell Co., Pleasant Valley, open oak woods, 
July 8, 1936 (G, NY); Core, ‘Monongalia Co., near Halleck, July 
1}; 1931 (G). Virani: Fernald & Long 11 835, Greensville Co., 
Emporia, bs deciduous wooded slope, May 11, 1940 (G); 
Small, Smyth Co., Walker Mountains, Shannon nae alt. 3 
ft. , June 20, 1802, G (F, G, ese ee ap Rockbridge Co., Glasgow, 


June 1, 1891 (M). TH Carouina: Small & Heller 264, on 
road between Blowing. Rock and Shull’s Mill, June 16-17, 
1801 (ANS, F, M, NY, US); mine ees Polk Co., Tryon, open 


sides, May 22, 1899 (M); Churchi Ul, Madison Co., Hot 
Springs, June 1, 1899 (M). Sours Caroxina: Saint Andrews, 
May, 1855 (G); B. E. Smith, Darlington Co., Lauther’s Lake, 
low a _ 3, 1940 ged House 2226, Oconee Co., Fort 
May 24 906 (BG, M , US). Gronera; — 2018, Lumpkin 
Co., Blo od M ock Gap, moist edge of exposed ’rock slide, 
June 2, 1934 (ANS): r Cutlber 516 & 551, Augusta, May 17, June 
18, 1901 (NY); Pollard & Mazon 521, Dou ghe rty Co., vicinity 
of Albany, Aug. 13, 1900 (G, US); Churchill, Chickamauga Park, 
near Lookout Mountain, May 8, 1906 (G, M); Green, Macon 


1944] Boivin,—American Thalictra and their Allies 473 


(ANS, labelled 7. Cornuti, var. brevifolium Rugel and paratype 
of T. macrostylum). Fiorina: Chapman (G, NY); Rugel, prope 
St. Marks, inter frutices (Magnolia-Chamaerops), jun. 1843 
(NY, labelled 7. Cornutt var. macrostylum and paratyp 7: 
macrostylum). Ontario: J. Macoun 23,615, Amherstburg, July 
31, 1901 (G). Onto: Wilkinson 228, Mansfield, waste places, 
June, Aug., 1895 (US, type of 7. amabile); Moseley, Erie Co., 
Perkins, July 8, 1895 (G); Lea, Hamilton Co., June 14, 1838 
(ANS). Inprana: Peattie, Porter Co., Tremont, sandy field and 
along roads in the dunes, Sept. 4, 1920 (G). Kenrucxy: Smith 
& Hodgdon 3865, Monticello, Beaver Creek, rich wooded slopes, 
July 12-14, 1937 (G); Short, Lexington (ANS); McFarland 
James 5, Whitley Co., 3 miles west of Corbin, low swampy field 
near edge of ditch, July 25, 1941 (M). TEnNEssEE: Ruth 1808, 
Knoxville, groves and open woods, June, 1897 (NC, NY); 
Svenson 10,564, Cheatham Co., Pegram, dry shale, Aug. 22, 1940 
(BG); Eggert, Dickson Co., near White Bluff, Aug. 19, 1897 (M). 
AvaBaMa: Eggert, De Kalb, Collinsville, June 29, 1897 (M). 
Inurnors: Gleason 9131, Champaign Co., Champaign, wet 
prairie along railway, July 3, 1940 (NY); Gleason, Urbana, wet 
woods, June 27, 1906 (G); Greenman 3688, vicinity of Chicago, 
June 15, 1911 (G). Mussourr: Standley 9802, Webster Co., 
vicinity of Rogersville, thin woods, Sept. 3, 1912 (US) ; Letterman, 
near Allenton, 1893 (ANS, M, US); Emig 268, Elmont, May 23, 
1914 (M). Fie. 107, a-f. 

The specimen from Percé is only tentatively identified as T. 
revolutum. It is from far out of range and in a very different 
floristic area. 

The correct name for this species is still unsettled. Two 
nomina dubia antedate 7’. revolutum and when better understood 
might replace it: T. purpurascens L. and T. pubescens Pursh. 


474 Rhodora [DECEMBER 


(0.7)—1.0-(1.5) mm., alis obscuris. Carpella matura haud com- 
ae nec costata, ‘abrupte stipitata et apice incurvato, stipite 
a. 0.5 mm., ventro ca. 3 mm. longo et 1.5 mm. lato, nervis gins 
iin et no nnunquam sinuosis. Receptaculum ad_ bas 
capitis fructuum. Floret Junio, Julio et Augusto. bias pho 
Fernald & Long 9050, Henrico Co., west of Elko Station, sphag- 
nous springy swales bordering Whiteoak Swamp, Aug. 17, 1938 
(G); idem 8711, July 23, 1938 (G, M); idem 8710 (G); Fernald, 
ong & Smart 5778, on headwaters of Blackwater River, abou 
3 miles southeast of Petersburg, swampy woods, June 25, 1936 
(G); L. F. & F. R. Randolph o boreae Anne Co., Pungo, open 
swampy land along West Neck Creek, June 29, 1922 (G). 
NortuH CAROLINA: whee & Holler, ia Co., n. of a 
swamp, June 25-26, 1891 (NY, Type and isotypes of T. m 
stylum; F, ISOTYPES) ; idem 428 (ANS, NY); Rugel, Swanane ie 
pratis vallis, Aug., 1841 (G, T. Cornutt var. brevifolium) ; Heller 
1015, ape wha Co., near Hickory, June 23, 1893 (ANS, F, G, M, 
NY US); Biltmore Herbarium 1 024, Flat Rock, low grounds, 
June 6, 1905 (NY, US); Wiegand & Manning 1216, Martin Co., 
June 21, 1927 (); Godfrey 4384, Beaufort Co., ’ Washington, 
marsh, June 9, 1938 (G); Godfrey 4875, Hyde Co., ’ Swanquarter, 
June 9, 19 < (GQ); Godfrey 4449, Cartaret Co. , Newport, marsh, 
oe 10, 193 8 (G); Feri 4318, Pisgah Forest, "alt. 2500 ft., June 
29, 1909 (US); Peattie 1317, Polk Co., The Sho als, low wet woods, 
Aug. 20, 1921 (F); Peattie 1313 & 1313 13A, Polk Co., east of Colum- 
bus, edge of a M. agnolia virginica swamp, Aug. 20, 1921 (NC). 
The two herbarium sheets on which Small and Heller picked 
up the two synonyms given in the original description belong to 
T. revolutum DC. 
109. T. subrotundum, sp. nov. Planta omnino gishes, 


Godfrey & Tryon 121, Georgetown Co., 4 miles west of i 
town, creek-bottom dustak rich lowland woods, June 27, 1939 
(G, NY); Eggert, Aiken Co. , sandy swamps, Aug. 8, 1 1898 (M); 
Godfrey & Tryon 584, Berkeley Co., 3 miles southeast of Pine- 


atatnndbeaneaenenemaneaantiie ini. 


1944] Boivin,—American Thalictra and their Allies 475 


e 
of Muckalee Creek, Aug. 2, 1902 (NY, US). Fiorrpa: Berg, near 
Tallahassee (NY). AwtaBama: Earle, Lee Co., Camp Hill, 
June 23, 1897 (M, NC, NY); Harper 3525, Autauga Co., about 
2 miles southeast of Booth, swamp of Bridge Creek, June 15, 
1936 (ANS, BG, F, G, M, N-ND, NY, US). Fic. 108, a-d 
110. T. pouygamum Muhl. ex Sprengel, Syst. Veg. 2: 671, 
1825. T. polygamum Muhl. Trans. Amer. Phil. Soc. 3: 172, 


T. divergens Link, Enum. Hort. Berol. 2: 92, 1822. T. harsutum 


b 


Syst. 1: 174, 1817. T. altissimum Greene, Leaflets, 2: 58, 1910, 
nec 7’. altissimum Wender, Flora, 9: 358, 1826, nec T. altissimum 
Thomas ex De Massas, Ann. Sc. Nat. ser. 2, 9: 369, 1838. 


Farwell, Pap. Mich. Acad. Sci. Arts & Lett. 26 (1940): 11, 1941. 
Planta statura variabili, saepius pubescens. Pubescentza pilorum 
uniseriatorum flexuosorum. Foliolae membranaceae variabiles 
(1.0)-2.5-(7.5) em. longae, obovatae vel oblongae, apice trilo- 
atae lobis integris. Inflorescentia paniculata, apice rotunda, 
pedunculis tenuibus. Filamenta rigida 3.5-5.0 mm., apice 
clavata et sub antheris constricta. Antherae saepius oblongae 
ca. 1.0 mm. long. Stigma 0.5-2.0 mm., alis obscuris et pilis 
crassiusculis. Carpella stipite 0.2-1.0 mm., ventre saeplus ovato 
vel obovato vel ellipsoideo, nervis raro parum sinuosis. Floret 
Junio, Julio et Augusto.—NewrounpLanp: Fernald, Long | 

Fogg 269, Bay of Islands, southern slope of Lark Mountain, 


476 Rhodora [DECEMBER 


peaty and turfy subalpine sap gag ee 1, 1926 (G); Howe & 
Lang 1006, Bay St. George, Aug. 5-7, 1901 (G); H. Bishop 319, 
near Bonne hago Neddy Harbor, worded banks of stream, Aug. 
28-30, 1928 (G). ‘“Canapa”: Kalm [?] (Linnaean Society 
Lond., veraty pe of T. dioicum; G, photograph). Saint-PIERRE 
et MrquEto : L.-Arséne 264, Cha apeau de Miquelon, 31 juil., 
1901 (G); L. A fsing 239, Langlade, vallée de la Belle-Riviére, 
aotit, 1901 (G). QuEBEC: Victorin, Rolland, Brunel & Rousseau 
17; 47, Percé, sur les corniches de conglomérat, 24 juil., 1923 
(G); J. Macoun 66,630, 66,631 & 66,682, Cap a l’Aigle, Aug., 
1905 (G); Senn 396, near Wakefield Lake, — July 24, 


1938 (G). Prince Epwarp Istanp: J. Macoun 869, Tignish, 
July 25, 1888 — type of T. glaucodeum; G, rpliotoerenhe of 
ame); "J. Mac Mt. Stewart, July, 1888 (US); Fernald, 


Bartram, Long & St. John 7498, Mt. St tewart, springy ditch by 
railroad, July 30, 1912 (G). New Brunswick: Rousseau & 
Bonin 32,047, junction of Restigouche and Matapedia Rivers, 
gravelly banks, hed oa 1929 (G); Chadbourne, King’s Co., 
Rothesay, July—-Aug 1883 (G); Fowler, Kent Co., Bass River 
(F). Nova Satta: tdi Macoun 19, 006, Cape Breton Island, 
Baddeck, thickets, July 28, 1898 (G-ND, type of 7. tortuosum; 
G, photographs of the same); ; Fernald & Long 23,8658, Yarmouth 
Co., Parr Lake, thicket at upper border of cobbly beach, Aug. 
12, 1921 (G); Fernald & Long 23,852, Digby Co., Little Meteghan 
Lake, thicket, Aug. 9, 1921 (G); Perry, Wetmore, Hicks & Prince 
10,140, Antigonish Co., Salt Springs, along brook, Sept. 11, 1925 
(G). aero B. L. Ro binson, Rangeley Lakes, Middle Dam, 
Aug. 2, 1903 (G); Solar Penobscot Co., Orono, rocky bank, 
July 12, "1802 (G, NE); Fernald 8, Aroostook Co., St. Francis, 
low thickets, Aug. 15, 1893 (G). New Hampsuire: A. H. Moore 
4082, Coos Co. , Colebrook, foot of Lombard Hill, east of Monad- 
nock phate shaded r oadside, July 20, 1907 (G); ; Batchelder, 
Cheshire Co., Biniatond, shore of Sandy Pond, Sept. 3, 1916 

E). Ver RMONT: Day 11 , Bennington Co., Manchester, June 
rb July ocean (G, NE): Williams, Ripton, Bread Loaf Inn, 


J . Massacuusetts: Churchill, Berkshire Co., 
heficld. low ground, July 19, 1920 (NE); Fern 
18,470, me wich, Spring rg low thicket 


a Ruope Isuanp: Fernald, Kent roe ., Warwick, ao! 
ae June 25, 1910 (G, NE): Ware & Fernald. d, Washin 

esterly , boggy swale north of Babcock Pond, Aug. 31, 

1919 (NE); below East Providence, sta . 1929 (NE). Con- 

NEcTicUT: Bissell, Southington, July, 1897 (G-ND, fragment 

from type of T. Bissellii; G, tay Saas of this fragment) ; 

Meredith, Litchfield Co., Kent Falls, brookside, Aug. 2, 1927 


1944] Boivin,—American Thalictra and their Allies 477 


(ANS); Wright, Meriden, July 21, 1879 (NE). New York: 
Lucy 227, Chemung Co., Elmira, July 2, 1896 (G-ND, fragments 


Oswego, Mud Pond, swampy woods and _ thickets overlying 
Silurian sandstones, Aug. 23, 1922 (G). New Jersny: EL. 


1930 (G); Greenman 368, Randolph Co., Big Run, Sept. 14, 1904 
(F, G). Virernta: Fernald & Long 8267, Surry Co., James 


Co., river banks, May 3, 1932 (G); Biltmore Herbarium, Madison 
Co., near Marshall, banks of Big Ivy Creek, Aug. 8, 1898 (G, M, 
Y, US). Gzorara: Ruth 10, Blue Ridge, swamps, July 10, 


ensum; G, photographs of the type and of one isotype) ; idem, 
june 12, 1909 (G-ND, paratype of 7. perpensum). Outo: F. EB. 


Leonard 87-11214, Elynia, July 14, 1887 (US). INDIANA: 


=o 


|! 
Rene, Floyd Co., New Albany, Aug. 23, 1923 (N-ND). 


Kentucky: Smith & Hodgdon 3929, Wayne Co., southwest of 


Monticello, Beaver Creek, shady ledge, July 12-14, 1937 (G); 
Smith & Hodgdon 3695, Rockcastle Co., between Berea and Mt. 


478 Rhodora [DECEMBER 


Vernon, slough, July 8, 1937 (G, NY); Short 2, alluvium of the 
Kentucky River (ANS); Lea 3, Kenton Co., Banklick Creek, 
1834-44 (ANS). TreNnNeEsSEE: Kearney 602, Cocke Co., near 
Lemon’s Gap, Sept. 3, 1897 (G-ND, type of T. perelegans; M, 
NY, US, isotypes) ; idem 60214 & 603, Sept. 8 (M, NC, NY, US); 
Svenson 4050, Morgan Co., Rugby, mossy banks of stream, 
Aug. 19, 1930 (BG). Fig. 110, a-c. 

Thalictrum polygamum Muhl. is antedated by two other 
validly published names and perhaps by four. Although we 
know that this name is not the right one for the species, it seems 
preferable to keep it until we make sure of the right name to take. 

Gray gives 1813 (Cat. Plant. Amer. Sept., p. 54) in his Synop- 
tical Flora as the right date for the valid publication of T. poly- 
gamum Muhl. He considered the two words “smooth, polyga- 
mous” as a valid description. Actually these were not intended 
to be a description, but common names, that is ‘smooth meadow- 
rue, polygamous meadow-rue”, as stated at the beginning of the 
Muhlenberg’s Catalogue. The first valid publication is, then, 
Sprengel’s in 1825, later than 7. corynellum DC. 1817, T. diver- 
gens Link 1822, T. pubescens Pursh 1814, and T. purpurascens 
L. 1753. 


111. T. poryeamum Muhl., var. HEBECARPUM Fern., RHODORA, 
10: 49, 1908. T. leucocrinum Greene, Ott. Nat., 24: 29, 1910. 
T. zibellinum Greene, |. c. 30, 1910. 7. labradoricum Greene, 
1. c. 53, 1910. TT’. terrae-novae Greene, 1. c. 52, 1910. T. cana- 
dense Miller, var. hebecarpum (Fern.) House, Bull. N. Y. State 
Mus. 254: 346, 1924. Planta robustior saepius pubescens et 
sesquimetralis. Inflorescentia ampla subcorymbosa, pedunculis 
crassiusculis et nonnunquam incurvatis. Filamenta saepius 5.0- 
6.5 mm. conspicue clavata. Antherae ca. 1.2mm. Stigma 1.5- 
3.5 mm. Carpella matura saepius oblanceolata. Floret Julio 
et Augusto.—Lasrapor: H. Bishop 318, Petty Harbor, 52° 25’ 
N., 55° 40’ W., sphagnous spruce woods, July 12, 1928 (G); 
Fernald & Wiegand 3438, Blanc Sablon, by brooks, Aug. 6, 1910 
(G). NewrounpLanp: Robinson & Schrenk 187, St. John’s, 
Rennie’s River, rocky banks, Aug. 4, 1894 (US, type of Tr. 
terrae-novae; ANS, F, G, NY, isotypes); Fernald & Wiegand 3437, 
Port Saunders, gravelly shore, Aug. 6, 1910 (G); Fernald & Long 
28,274, Bard Harbor Hill, meadow below limestone escarpment, 
Aug. 21, 1925 (G). Qursec: Williams & Fernald, Témiscouata 
Co., Riviére du Loup, gravelly thicket by the Saint Lawrence, 
Aug. 2, 1902 (G, typr of T. polygamum var. hebecarpum); 
Fernald, Bartram, Long & St. John 7492, Magdalen Islands, 
Grindstone Island, Grindstone, wet clearing, July 22, 1912 (G); _ 


1944] Boivin,—American Thalictra and their Allies 479 


Richardson 867, Lake Mistassini, wet meadows around springs 
and along rivulets, July 15, 1870 (C); Churchill, Terrebonne Co., 
Lac Tremblant, July 18, 1922 (G). Prince Epwarp Isuanp: J. 
Macoun, Mt. Stewart, Aug. 17, 1888 (G). New Brunswick: 
J. D. Smith, Campobello Island, July 17-Aug. 20, 1888 (US, 
type and isotype of 7. leucocrinum); F. T. Hubbard, Shediac 
Cape, rich meadow, with alder, July 21, 1914 (G); Fowler 2, St. 
Andrew’s, July 3, 1900 (G); Fernald & Long 13,641, Carleton Co., 
Woodstock, gravelly river-thickets and bushy terraces, July 14, 
1916 (G). ova Scotia: J. Macoun 21,134, Sable Island, not 
uncommon on the old land, July 24 & Aug. 2, 1899 (G, isotype of 
T. zibellinum); Perry & Roscoe 217, St. Paul Island, between 
Petries Pond and White Spring, Aug. 3, 1929 (G); Long & Linder 
21,256, Yarmouth Co., Tusket, peaty spruce and alder thickets 
bordering swales, July 15, 1920 (G). Marne: C. A. E. Long, 


111, a an 

This variety is not a well cut unit but a rather well marked 
extreme commoner than the typical in northern parts of the 
range of the species. The following variety exhibits a similar 
behavior but the relation between the staminate and the pistillate 
plants has not been made yet. A third variety is perhaps worth 
recognition in the southern Appalachian upland (T. altissimum 
Greene and T. perelegans Greene). 


112. T. ponycamum Muhl., var. intermedium, var. nov. 
? T. viride Greene, Leaflets, 2: 56, 1910. Planta metralis pu- 


480 Rhodora (DECEMBER 


bescens. Foliolae subcoriaceae et revolutae. Inflorescentia pa- 
niculata. Antherae 1.5-2.0 mm. Ruope Istanp: F. S. Collins 
2854, Bristol, July 12, 1918 (NE). _ CONNECTICUT: Woodward, 


The bos: of T. viride Greene has ait been seen by me. 


113. T. pasycarpum Fischer & Lall. ex Fisch., Mey. & Lall. 
Ind. Sem. Hort. Petrop. 8: 72, 1842. T. Co rnuti L., var. @ 
hg Mey. & Lall. 1. c. 72, 1842. T. virginicum Drege. ex Lec., 

Bull. Soc. fe Belg. 24: 323, 1885, nomen ex synonymis. T. 
purpurascens L., var. dasycarpum (Fisch. & Lall.) Trel., Proc. 
Bost. Soc. Nat. Hist. 23: 301, 1886. T. vegetum Greene, Midl. 
Nat. 1: 103, 1909. T. albens ’Greene, Amer. Midl. N : 292, 
1912. T. Moseleyi Greene, |. c. 294, 1912. T. Nortoni i Greene C. 

, 1912. 


Nieu wland, Amer. Midi. Nat. 3: 324. 1914. L. albens (Greene) 
Lunell, Amer. Midl. Nat. 4: 361, 1916. L. vegeta (Greene) Lunell, 
l. c. 1916. Planta metralis et ’ pubescens. Pubescentia ag 8 
densa pilorum uniseriatorum et flexuosorum. Rami et folia 
plus minusve conferta. Foliolae coriaceae, margine revoluto, 
oblongae, apice trilobatae, lobis acutis. I nflorescentia paniculata 
ae acuta. Filamenta capillaria et sub antheris paululum 
dilatata nec constricta, ca. 4 mm. Antherae 1.5-2. = mm., 
oblongo-lines aria, acumine 0.1-0.4 mm. Stigma 2-3 m Re- 

eptaculum ad basem cnapiie fructuum. Carpella. ‘tint 
stipite 0.1-0.3 mm., ventre saepius ovato, apice nonnunquam 
incurvato, nervis — sinuosis. Floret Junio Julioque.— 
Russia (cultivated): x horto botanico petropolitensi (US, 
presumably from type ae in the garden). NortH AMERICA: 
el Expedition te Ontario: C. F. ee 1 at 


& Leslie 2280, Al goma District - tke adenehon birch 
association, July 35, 1935 (C)j Teor Losee & Bannan 904, 
Thunder Bay District, Marie Louise Creek, along —— Aug. 
13, 1936 (C); W. S. 62, 298, Aine vanary, July 15, 904 (C). 
Micuican: W. F. Wight 118b, Alle egan Co., along Fes Rea R. 
at Allegan, Sept. 11, em (US, type, on two sheets, of T. Wights- 
anum); Barlow, M. ette Co. ., Turin, by flowing water, July 8, 
1901 (G); Fernald & ibe 307 Houghton Co., southwest of 
Houghton, border of swamp, July 3, 1934 (G); 'L. H. Bailey, 


1944] Boivin,—American Thalictra and their Allies 481 


near Lansing, June 13, 1886 (G). Onto: Webb 491, Portage Co., 
Garrettsville, June 24, 1901 (G); Moseley, Erie Co., Oxford Tp., 


W. Edmonds, Vilas Co., Eagle River, 1926 (NY); Fassett & 

Wilson 4301, Buffalo Co., Fountain City, sandy R. R. embank- 

ment, Aug. 25, 1927 (G); Schuette, Brown Co., Dach’s Creek, 

July 11, 1895 (G, NC). Ixxrots: Bebb, Fountaindale, 1867 
; G 


5, 19 
Pammel & Zimmerman 275, Feinback, June 23, 1925 (G); Ball 2, 
Ames, July 18, 1896 (G). Missouri: E. J. Palmer 18,991, 


: r 
north-facing wooded bluff of Little River, July 5, 1933 (M 
ortH Daxora: Lunell, Ramsay Co., Devil’s Lake, in woods, 
July 16, 1902 (G-ND, type of T. vegetum; G, photographs) ; 
Lunell, Devil’s Lake, July 1, 1905 (G-ND, paratype get 


Hayward 1694, Black Hills, Hot Springs, mixed prairie, 1927 (F). 
Nepraska: Tolstead 5138a, Valentine, along the banks of the 
Niobrara river, July 27, 1936 (G); F. Clements, St. James, June 
24, 1893 (G, US); Rydberg 1413, on South Fork of Dismal River, 
in meadow, Aug. 14, 1893 (G). Kansas: J. B. Norton 4a, Riley 
Co., 1896 (US, type of T. Nortoni; G, M, eer 
White, Cowley Co., June, 1898 (M); Rydberg & Imler 44, a 
Co., between Olathe and Pleasanton, June 18, 1929 (NY). 
SaskatcHewan: Bourgeau, 1857-1858 (G); Herriot 69,8014, 
Yorkton, damp thickets, July 6, 1906 (G). Anperta: Herriot 


Ipano: Leiberg 1576, Clarks Fork Valley, below Weeksville, alt. 


482 Rhodora [DECEMBER 


650 m., Aug. 23, 1895 (G). Wyomine: F. L. Bennett 827, Black 
Hills, Sand Creek Canyon, shady flood-plain, alt. 3800 ft., June 
24, 1938 (CA); A. Nelson 8338, Laramie Co., Badger, thickets 
on ’ river-banks, July 1, 1901 (U 8). Cotorapo: Cowen 27 & 82, 
Fort Collins, river-bank, alt. 5000 ft., July 1, 1895 (G, US). 
New Mexico: Standley 13,541, Colfax Co. wf vicinity of Ute 
Park, meadow along creek, alt. 2200 to 2900 m., Aug. 22, 1916 
(US); Arséne “e x 17 ,431, vicinity of Watrous, alt. 1950 
m., Aug. oT, 6 (CA, US a: Vasey, Las Vegas, 1881 (US). 
ARIZONA: Haugh, havin, July 14 (US). Fie. 113, a and b. 
A study of the polygamo-dioeciousness of this species was pub- 
lished by J. H. Schaffner in the Ohio Journ. Sci. 20: 25, 1919. 


T. pasycarpum Fisch. & Lall., var. hypoglaucum 
(Rydbers), stat. nov. 7. hypoglaucum Rydb. Brittonia, 1: 88, 
1931. 1. macrostigma Torr. ex Trel., Proc. Bost. Soc. Nat. Hist. 
23: 301, 1886, ut synonymon. T. amphibolum Greene, Fedde, 
Rep. Nov. Spec. 7: 255, 1909. 7. dasycarpum Fisch. & Lall., a 
hypoglaucum Steyermark, RHopora, 40: 178, 1938. Planta 
saepius glabra. Foliolae saepius membranaceae. Filamenta 
4-7 mm. Antherae 2.2-3.2 mm. Stigma 2.5-5.0 mm. Recep- 
taculum ad centrum capitis fructuum. Capella matura recta 
ventre lanceolato et nervo ventrali paullulum convexiore quam 

orsali. Floret Majo et Junio. Mussourr: Standley 9298, 
Greene Co., creek-bottom, vicinity of Ash Grove, Aug. 24, 1912 
(US); £. J. Palmer 36 ,684, Johnson Co., Columbus, thickets, 
limestone hills and low woods, June 21, 1930 (G); Bush 4, 
Independence, June 26, 1895 (NY); Steyermark 5 5938, Nodaway 
Co., Parnell, att open woods of Bunker Hill, June 20, 1938 (F). 
peop EP soln ech Fort Cobb and Fort Arbuckle, 
68 (NY, U; stigma); Bush 2479, Miller Co., woods, 
kon 27, Joos a) : Fabiett 19,760, Hempstead Co., Hope; April 
se 1938 (G). Lovtstana: Hale, Alexandria, 1840 (NY, US, 7. 
acrostigma). Sours Daxota: T. A. Williams, Big Stone, Aug. 

7. 1895 (US); Hayward 135, Spearfish Canyon, streamside, Aug. 
, 1926 (NY); Brenckle 41 -27, east of Malette, river-bottom, in 
oods, June 16, 1941 (G). NEBRASKA: Bates, Red Cloud, June 
iL 1906 (G); E.S. Bacon, Neligh, June 15, 1896 (G). KANsas: 
Hal, ees 1870 W); Kellerman, Manhattan, 1887 (M). OKta- 
“Pega Washin ngton Co., Copan, in moist 

shady mt rear a2 13 (G); Merrill & ican 546, Platt 
National Park, June 3 1935 (F, US); G. W. Stevens 920, ‘Cadd ty) 
Co., Hinton, bottoms of Devil’s. Canyon, June 15, 1913 (G, US); 
J. Clements '11 ,083, Comanche Co., Fort Sill, June 2, 1916 (G). 
aoe Hall 2, Dallas, creek-banks, 1872 (NY, vyPE of T. hypo- 
ucum; F, G, M, US, ISOTYPES) ; Eggert, Mowie, woods, June 

13, 1898 nn NY, paratypes of T. hypoglaucum) ; Lindheimer, 


1944] Boivin,—American Thalictra and their Allies 483 


Houston, April, 1842 (G, US); Lewton 57, Victoria Co., Victoria, 
March 23, 1905 (US). Montana: Spragg 369, Square Butte, 
canyons, July 15, 1901 (G); Watson 7, near Frenchtown, Aug. 4, 
1880 (G, US); M. E. Jones, Ravalli, Middle Temperate Life Zone, 

: (CA, UC). Coxrorapo: Cowen 32, 
Fort Collins, river-bank, alt. 5000 ft., July 1, 1895 (G); Bethel, 
Platte, July 31, 1916 (US). Arizona: J. W. Lead 1511, White 
Mountains, meadows, July 30, 1935 (US); Rusby, Oak Creek, 

1883 (ANS, F, G, NY, US). WasHINeTON: Kreager 377, Box 
- Canyon, Aug. 2, 1902 (US). Brivis Cotumsia: J. Macoun 
10,059, Waterton Lake, thickets, July 27, 1895 (C); J. Macoun 
860, Warm Springs, open places, July 5, 1890 (C). Fria. 114, 
a and b. 

The carpel is not always perfectly closed in the genus Thalic- 
trum. Near the base of the style the ventral nerves are free for 
a short distance and not infrequently there is a transition in the 
tissue of the carpel between the two ventral nerves. This is more 
easily seen in nearly mature fruits of T. dasycarpum Fisch. & 
Lall. var. hypoglaucum. 


SPECIES HAUD SATIS COGNITA 


foliolis ambitu suborbicularibus, subquadratis v. rhomboideis, 
raro usque infra medium 3-sectis, plerumque antice trilobis v. 
tricrenatis, glabris; inflorescentiis 1-2-floris; sepalis 3.5 mm. 
ongis, albis, subtus violaceo-maculatis; antheris ovatis obtusis; 
acheniis sessilibus anguste lanceolatis, longitrorsum striatis, 
rostro uncato excepto cr. 3 mm. longis, 0.7 mm. latis. 
Caules debiles, verisimiliter plantis aliis incumbentes, inferne 
usque 1.5 mm., medio vix supra 0.5 mm. crassi, teretes v. obtusan- 
guli. Stipulae semiovatae, inferae usque 4 mm. longae, superiores 
sensim decrescentes, margine saepius dentatae, marcescentes. 


feste nervosa. Flores primarii (terminales) 10-12 mm., secun- 
darii (ex axilla folii simplicis v. bracteiformis abeuntes) usque 

em. longe pedicellati. Sepala non unguiculata, ovata v. 
ovalia, apice obtusa, 2 mm. lata. Stamina er. 10; filamenta 2 


484 Rhodora [DECEMBER 


Hab. in Haiti in Morne la Selle prope fontem Riviére Blanche 
de Jacmel 1800-1900 m. alt., m. Aug. flor. et fruct.: Christ n. 
1848, 1848b. 

Obs. Species hujus generis prima ex Antillis cognita, pecu- 
liaris, habitu foliorum Th. Fendleri Engelm. et Th. Wrighti 
Gray (ex. America sept.) in memoriam revocans, sed ab iis 
statura, inflorescentia, antheris, acheniis omnino diversa. 

The preceding text is a copy of the original description. I 
have not seen any material of this species but it seems to differ 
from the continental species and probably belongs to the Debilia. 


NOMINA DUBIA 

T. purpurascens L., Sp. Pl., 1: 546, 1753. 7. dioicum L., 
var. « purpurascens ens (L.) Povanoling Fl. Can. 1: 5, 1862. 

Of the type of this species in the herbarium of Linnaeus I have 
seen only the negative of a 35 mm. photograph. It is the upper 
half of a staminate plant collected in early bloom and looks 
pretty much like 7’. revolutum DC., but could be just as well 7’. 
polygamum Muhl. 

T. puspescens Pursh, Fl. Amer. Sept. 2: 388, 1814. T. 
polygamum Muhl., var. ‘pubescens (Pursh) K. C. Davis, Minn. 
Bot. Stud. 2: 514, 1900. 

The descriptions of Thalictra given by Pursh in his Flora are 
all more or less obscure to me and there are a few too many. I 
have seen five of Pursh’s collections from the B. S. Barton her- 
barium. But that did not elucidate the problem. I suspect the 
type to be in the Delessert Herbarium and to belong either to 
T. polygamum Muhl. or to 7. revolutum DC. . 

T. CAROLINIANUM Bose and var. (ht tee She tha Syst. 1: 
174, 1817, nee T. carolinianum Walter, Fl. Car., 157, 1788. 

I have not seen any typical material of this aes and its 
variety, and Lecoyer, who has seen the type specimens in the 
Delessert herbaritim, considers them as synonyms of 7’. polyga- 
mum and T’. dasycarpum, but his account is not very pagan | 
De Candolle’s description i is not very clear, but as the name 
carolinianum Bose is already preoccupied by one of Walter’s s 
species, its correct interpretation is of little one ret 

T. mecacarpum Torr. Cat. Pl. Fremont Exped. 87, 1845. 
nomen nudum. 7. megacarpum Torr. ex Rydberg, in Rocky 


1944] Boivin,—American Thalictra and their Allies 485 


Mts. 290, 1918. TT. occidentale, var. megacarpum (Torr.) St. 
John, Fl. South. Wash. & adj. Idaho, 158, 1937. 


The type of this species I have seen, but it greatly puzzles me. 
The foliage is typical of 7. occidentale Gray var. palouense St. 
John; its inflorescence is pretty much like that of T. confine 
Fern. var. columbianum (Rydb.) nostrum, and the fruit, although’ 
closer to certain forms of 7. Fendleri Engelm., also recalls that 
of T. occidentale Gray var. typicum. Such a specimen I consider 
as being either abnormal or of hybrid origin. 

Other names have been discarded for various reason but none 
of them is likely to be entitled to supersede any of the names: 
adopted in this paper. The list of these names follows: 


Sect. campronotum Prantl, Nat. Pflanzenf. 3, 2: 66, 1888. 

ect. CAMPTONOTUM, ¢. PETALOIDEA Prantl, lc. 

Sect. CAMPTOGASTRUM, C. PLATYCARPA Prantl, 1. ¢. 

aa pasa pg DC. Syst. 1: 187, 1817, nomen provisorium. 

T. Cornutt L., var. MONOSTYLA [author?], Bot. Zeit. 3: 218- 
219, 1845, peitiegts nudum 

T. PURPUREUM K. C. Davis, Minn. Bot. Studies, bt 1900, 
nomen ex synonymis Thalictri purpurascentis L., 7. pur- 
pureum Schang ex Pall., N. Nord. Beitr., 6: 42. 


EXPLANATION OF FIGURES 
Fias 2—25 (p. 351), all X 4 


Fig. 2. Taaticrrum Hvuirenu Boivin: — Everman : 
ALPINUM sees var. Typicum: a) sepal, Blaisdell. 800; b) stamen, » a 
a 1935; s) ovary, Pease & Smith 26, 761; @)! ) brac peduncle and fruit, 
Fidiz, Aug. 11, 1910. 5. TT. aALPINUM L., var. sT rraTum ‘Yabe: a) ovary, 
Rock 17 si; ) fruit, Takemoto 591. igs ie ee AICALENSE Turez.: fruit, Hsia 
2310. 9. AICALENSE Turcz., var. ama Boivin: fruit, Fang. 
3619, 10. ee crtyeracrnaee Cc. ot Robinson: a: fruit, OV illiams 1187. 11. 

Aw coor tht yaa gg Akg Mazi cz, Amur. 12. T. cLAVA og ee a) 
uncle and sepal, M agee, Tgadortone’ one; .) peduncle and anther, id.; ¢) seed, 

er, Cullowhee; d) section of fruit with — M oe ckstone; e) peduncle 

fruit, id. id. Pca T. pecurnatom Boivin: Lam 0 Teint onde Mozset b) fruit, 

Hand.-Mazz.) Boi andel-M 
11,173; b) stamen, ks 3c) ae & Ko &2, S21. (The Mane “ ee tee aatted 
byerror.) 1 oe ener s fruit, Henry $982. 16, 'T. UNGUICULATUM 
Boivin: ayes fruit, id. — - yr ae Boivin: ‘a) 
Sepal, Teang. 28 $186; b) 0) ether Ps c) frui T. MIRABILE Small: 
sanen, Bath Hodgdon 39 6) fruit, gry £2 frit Earle 2212. Fig. 18cis 
esctiorally Mg x Tw i, ry. ntaber f oa. shoatil: Nan seen 
aie a , 
1892. 21. T. Caraonts Boivin: a) stamen, res a0 18,71 719; ides = id, 23; T. 
Secoeat rte Lec.: fruit, Wilson 492. 25. T. + Var. 
CUM: a) stamen, Komarov 724; b) fruit, id. 


486 Rhodora [DECEMBER 


Fics. 26-43 (p. 371), all x 4 


Fic. 26. T. sPARSIFLORUM Turcz., var. RICHARDSONII (Gray) Boivin: fruit, 
Kellog 211. 27. T. SPARSIFLORUM Turez., var. SAXOMONTANUM 
36. 


g 
fruit, Nelson 6864, 28. T. sPARSIFLORUM Turcz., var. NEVADENSE Boivin: 
fruit, Heller 7056. 29. T. "AQu UILEGIFOLIUM L,: a) section of fruit and seed, 
ae Chablais; b) peduncle and fruit, id. 30 D ivin: a) sepal, 
en, id.; c) ovary id.; d) peduncle and fruit, id. 31 
guciceocasie D ich.: a) stamen, Linder 2116; b) sepal, id.; c) ovary, 
id.; d) peduncle and fruit, Schimper 472 . INNITENS Boivin: a) sepal, 
= on en, id.; c) ovary, id.; d) peduncle and fruit, Owan 598 
T. Cuarini Boivin: a) sepal bar 6; b) stamen, id.; c) ovary, id.; d) 
peduncle and fruit, Mearns Le - IMPEXUM Boivin: a) sepal, Curtis 
39; b) stamen, id.; c) Rak gh 8 : d) fru sig dyiey 630. 36. T. STEINBACHII 
Bovis a) sepal, Ste inbach 8. ‘apodiond .; ¢) fruit, id. 37. T. Sreyer- 


MARKII Standley: a) ovary, hte 85 140; i fruit,id. 38. be paso ie 
Boivin: a) ovary, Steinbach 9231; b ruit id. 39. T. MACROCARPUM 
fruit, Cosson, Gourzy. 40. a INUNCANS Boivin: a) sepal, Fiebrig 2440; 'b) 
ovary, id.; c) stame n, id. . T. Gateorri Lec.: a) stamen, Galeottt 4541; 
ovary, id.: 3c) fruit, id. Fig. ‘41a and 41b 4 were drawn Pasa actual eee 
fig. 4le from a photograph of the type. 42. T. Hivronm Boivin: 

inton 6743. 43. T. pecirrens Boivin: a) ovary, Herrera 1047a; b) fruit, ad 


a 44-69 (p. 409) all x 4, except 5la and 68a, these 1/5 


Fig . T. virmuLum Boivin: fruit, Seibert 204. 45. T. PANAMENSE 
Standiey: ‘fruit, Destin Vol 46, "Macs CBRIDEANUM Boivin: io sepal, 
Macbride 4466; fag pemgrre ng a A! __. id.; d) fruit, id. 47. DeaMIii 

le 507. 


Boivin: ae Pring GRANDIFOLIUM Wa f: nik Jones, 
Soldier Canyon Muamesi Tausch: fruit, Abbott 288. 50. T. 
PACHUENSE —= ge Beg 96 51, STaNDLEYI Steyermark: a) 


leaflet, Sta tandley 

Stand. & Stever.- ay ace ars | = Pang sb) stamen, Johnston 1643; 

¢) nearly orga a T 
had seo fabled 


E gee : 

ovary, id.; d) peduncle and fruit, id. 65. T. crsnosum Lec.: — and 
fruit, pabonoe 6611. 67. T. aesroasesren DC.: fruit, Dombey, } Drawn 
from a photograph of the type. 68. T. vesicuLosum Lec.: fruit, Penland 
Saanies 1014. 69. T. Netsonn Boivin. a) ovary, Nelson ‘1788; '» fruit, id. 


Fics. eae (p. 429), all x 4 
Fic. 70. T. popocarrum HBK.: a) ovary, a & Smith 16,017; hey 
pecenee and fruit, id. 71, T. pa hs Boivin: a) peduncle and stam 
enturt 10,026; b) ovary, id.; ¢) fruit, id. 72. T. LANKESTERI Standley: ) 


ret ey & Boivin: a) sepal, Torres 187; b) stamen, id.; ¢) 
ovary, id.; d) fruit,id. 74. T.c MALEN DC. & ) sepal, 
— 7b) stamen, id.; c) o , id.; d) fruit, Hayes, Las V: 75. T, Tar 
Me > , Langlassé 1061; b) = ie T rarer DC 


sepal e ; : -: 
eee anal nton 4547; b) stamen, Alaman, Mexico; c) ov: ton 4547. 
Fig. 76b wae drawn from the photograph of the type... 71. I. Sree gee nr Wate.: 


1944] Boivin,—American Thalictra and their Allies 487 


a) ape, Pringle on a — id.; c) ovary, i id.; d) fruit, Leavenworth & 
Hoogstraal 989. EANUM m Boivi n: a) a "Pringle 7205; b) stamen, 
id.; c) ovary, iaste This; a) fruit, Pringle 8261. 80. T. Arsenu Boivin: a) 

sepal, Arséne, Jaripeo; b) stamen, ‘Arsen e 5470; c) fruit, id. 81. T. JaLisca- 

num Rose: a) sepal, Hinton ton 4259; b) stamen, id.; c) fruit, Hinton 6557. 82. 


. DEBILE ick: : fruit, ile Alabanik 86. . ARKANSANUM Boivin: 
fruit, Bush 2445. 87. T. pupicum Standle ey & Boivin: portion of stem with 
stipule and base of petiole, piety & H oogsiraal 1 1 an 88. T. MADRENSE 
Rose: fruit, Rose 2232. 89. T. prnnatum Wats.: 4 al of staminate flower, 
Townsend & Barber 134; b) sepal of pistillate flower, id a fruit, LeSueur 1059. 


Fias. 92-114 (p. 467), all X 4; —— bos < 1/5, and 110c and 113a, 
t x 


Fic. 92. T. vencLosum Trel.: a) sepal of ee pt rit — 
b) ovary, id.; c) sepal of staminate flower, Potter 0; d) stamen, Herriot 
69,803; e) fruit, Scamman 2892. 93. T. connie Fern : a) se fg of pistillat 

: wi 


; age Joivin: a) sepal of pistillate 
gl Morris 1527; b) sepal of staminate flower, Steele, Feeder pale Island, 
July 10; c) stamen, Steele, Feeder Dam Island, May 10; d) fruit, 

yo ibe 1292. 97. T. contaceum (Britt.) Small: ar aes Bedford Co. 


Engelm.: a) peduncle and stamen, Metcalfe 248; b) section of fruit and seed 

Munz 8696; ¢) fruit, id.; d) fruit, Parish 1483; e) fruit, Heller 11, 669. 103. 

T. Fenpierr Engelm., var. WricuTu (Gray) Trel.: fruit, Wrig Wright 834. 104. 
elson 


Fenpiert Engelm., var. QUADRINERVATUM spaah es fruit, Wiggins & Demaree 
4941. : Sige adm (Torr.) Wats.: sepal of pistillate flower, 
aces 2 & Harford 3; b) stamen, Hansen 563; ne fruit, Smith 306. 107. T. 

REVOLUTUM DC.: a) stamen, Pease 3782; b) sepal of s staminate flower, id.; ¢) 
sepal ey pistillate flower, Rich, Stoneham; d) ovary, id.; e) seed, Fern ald & 
Wea 16,807; {) frait, id. 108, T, MacrosryLum Small & Heller: a) 
sepal of pistillate flower, Small & Heller, Hickory; b) sepal of staminate flower, 
Small & He “ . tam & 


Heller 1015. 109. T. susroruNDUM Boivin: a) ovary, 
b) stamen, id.; c) etal and fruit, id. 110. T. pouycamum Muhl.: a 
stamen, Burnham, pes ipoli; ye os Robinson, Rangeley Lakes; ¢) head of 
mature carpels, M uenscher & Clausen 4649. 111. T. POLYGAM 


: Fern.: a) s og Tans 
fruit, Fernald & Long, 28,274. 112. T. roLYaaMoM Muhl., var. INTERMEDIUM 
ivin: stamen, Woodward, Franklin. 113. T. DASYCARPUM Fise 
a) head of mature carpels, Lunell. Lake Ibsen; b) stamen, Mos tak Oxfo 
114, T, vere ae isch. & tak. . HYPOGLAUCUM om (Rydb,) Bi Boivin: a) 
ventral side of an immature fruit ae the fully expanded sti and the 
small hole in the = of the ovary near the base of the style, uous 920; b) 
stamen, Palmer 3. 


es 
pie 
hi 


i TEN iia NNNINRNSS —nikiicaiiiiiisiiaiin,. - 
ne 


OC EE 7 hast, 


INDEX 
New scientific names are printed in full-face type 


Anemone, 347 
onells, 347 
Aquilegi: 
Raatanis, 162 


Clavata, 360 
>. ir alma 360 
Coptis 


Hepatica americana, 405 
Homothalictrum, 350 


Leucocoma, 346, 347, 469; albens, 
480; canadensis, 375; dasycarpa, 
480; dioica, 435; Lunellii, 439; 
cadre 439; thyrsoidea sylva- 

, 439; vegeta, 


Physocarpa, 346 

Physocarpon, 346 

Physoc Bocarpum, 346, 360; clavatum, 
um, 421 

ei ay 346 


Ruprechtia, 346, 375 
Sumnera, 346, 347, 360; clavata, 363 


Thalictraceae, 347 
Thalictrum mabe. Lecoyerium, 341, 
g 


. Physo- 
348, 360, 361; sect. 
Thaletrun 349, l. Heterogama, 


432; t. Tripetrium, 348, 375; 
pag Clavooarps, 349, 443; 
subsect. Compressa 349, 453; 
subsect. Debilia, 349 432, 484; 
subsect. Dioica, 349, 435; subsect. 
rays 348, 426, 42 7; subsect. 

uina, 360; su t dibboos, 
348, “411: subsect. poner 349, 


subsect. 
349, 454, 457; subsect. Simplicia, 
34 ; osa, 349, 


80; alpinum, 337, 341, ) 302, 
353, 375, 376, var. elatum, ‘ 
9, var. gaspense v 


pallidum, 353, 355, var. pudicum 
53, 355, var. stipitatum, 358, nary 
var. typicum : altiss 
mum, 475, 479; amabile, 471, 473: 
am hibolum 


444, 
» 392, 395, 436 486; cheilanthoides, 
7: Chiaonis, , 368, ; 


chymoca rpum. cincin- 


393, 394; 
~ natum, 391, 392, 398, 486; clava- 


tum, 342, 361-365, 369, 370, oie; 
ar. acutifoli - colum- 


490 


zattii, 402, 417, oat cordifolium, 
421; coreanum, 361 3, coreo- 


oxycarp a. p 

cens, 484, ray idoitatend, 444, "454, 
var. stipitatum, 377, 444, 454; 
oss 4 ame um, 442; dioi- 


purpur: 4 2, 

445; divergens, 475, “ATR: doming- 
, 483; uriusculum m, 356, 357; 
elegantulm = 357; Esquirolii, 
ri, 360; Fe an 


468, 469, beac, Henle gr 468, 
var. quadrinervatum ; 457, 463, 
487, 457, 

487, Pate * Wright at 457, 461, 466, 
487; filamentosum, 362, 363, 85; 


eottii 

50; gibbosum, 399, 
402, 403, ages 411, fs, 419, 486; 
glauc 47 ’ ; glau- 
cum, 338, { 359, 360; grandiflorum, 
408, 418, 419; grandifolium, 341, 
a 408, — 419, 486; graveolens, 


INDEX 


486; hirsutum, 475; hondurense, 
424, 426; Hultenii, 340, 352, 485; 
ypoglaue cum, 342, 482; ichang- 
361, 85; impexum, 
391, ” 392, 395, "486; integrilobum, 
362, 368; innitens, 391, 392, 394, 
395, 397, 486; inuncans, 339, 400, 
2°, 486; isopyroides, 350; 
jalapense, 414; jaliscanum, 426, 
431, 4 Aptarere ; John- 
stonii, 348, 401, 412, 486; kemense, 
-* eg aig Berea 478; lae 
4 6; lae evigatum, 
1 


464; majus, 352, 

393, 394, 399; marginatum, 358 

megacarpum, : en 
i 6 


“ee 


485; monoense, 356, : 
456, 475, 477; Moseleyi, Se ot 


434, 487, var. su 


INDEX 491 


433, 435; platycarpum, 457; podo- 
carpum, 401, 402, 421, 486; podo- 


polygamum, : 338, 
442, 469-472; 475, 477, 478, 484, 
0, 


485; rainierense, 455, 

AB6: yetractuns, 401, 417, 486; 
revolutum, 375, 376, 4 408, 469-471, 
473, i Saag spl q. glabrum, 471, 
72: ubglabrum, 471; 
Fhynehocarpu, 339, 392-394, 396, 
pl 486; Richardsonii, 370, 
Tae cin omen , 360; Rose- 

, 430, 487; rotundi- 


lorum, 356; sessilifolium, 4 

417, 486; sis vireo 475; simplex, 
350; sparsiflorum, 340, 341, = 
372, var. nevadense, 


rier 369, 485; specios' 

359, 360; squarrosum, 350; Sta mr 

leyi, 339, 400, 411, 486; Steelea- 
: Stei 


sum, 

subpubescens a6, 416, 48 

ee rte 474, 487; suspen- 
tene: 


a: 
o 


utahense, 372; vegetum, 480, 
Venturii, 400, 404, 422, 486; ‘venu- 


Wightianum, 480; Wri 
, 484; zibellinum, 478, 4 


Trautvetteria, 34 


Fy ; 
5 
: 


Reprinted from Ruopora, Vol. 47, January—March, 1945 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
3 OF HARVARD UNIVERSITY 


CLIII / 


RUELLIA IN THE EASTERN UNITED STATES 


M. L. FERNALD 


Dates or IssuE 


poets 1-38 and Plates 839-850... ....----- sere i 
. Pages 47-63 and Plates 851-860. .......5-+-++ RC ee Y 3 Haesunald 1945 
Pages 69-90 and Plates 861-875: .......--¢-+-200r terete 19 Mareh, 1945 
pr! BOTA . 
Soy CEIY wc 
eer” Fa” 


AUG 17 1945 


wer ee ~ & 


Reprinted from Ruopora, Vol. 47, January—March, 1945 


CONTRIBUTIONS FROM THE GRAY HERBARIUM 
OF HARVARD UNIVERSITY 


CLIII 


RUELLIA IN THE EASTERN UNITED STATES 


M. L. FERNALD 


pases 1-88 and Plates 839-850... 0-00 --0s rr 11 January, 1945 
mim, eae 
Pages 69-90 Late a ° 19 March, 1945 


‘silliest. cunpiemiiaiiaiiiibas ii ae 
. EEE _——_ @—-_ — 
poets SU es se mes . eeenneiateineenaeenntee 


OONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD 
UNIVERSITY—NO. CLIII 


RUELLIA IN THE EASTERN UNITED STATES 
M. L. FERNALD 
(Plates 839-875)! 


Aways a difficult genus to understand, Ruellia, as it occurs 
in the eastern United States, is quite as baffling as elsewhere. 
Field-experience for several seasons in eastern Virginia has 
demonstrated to Mr. Bayard Long and me that too often plants 
growing side-by-side will differ very strikingly in the degree of 
pubescence or its length on stem or calyx and that, to some ex- 
tent, the specimens from a single small colony have to be arbi- 
trarily sorted in order that those of a single number may be 
tolerably uniform. The accumulated collections have been al- 
lowed to wait, with the hope that some ambitious student would 
care to hunt for stable characters. Finally, however, the 
familiar “let Fernald do it” has prevailed and I have found it 
necessary to seek fundamental characters in the plants of the 
“Manual range”. In doing so it soon became evident that, 
since names originally given to plants of Georgia, Carolina, 
Virginia, Kentucky or Arkansas were being currently used for 
species growing only far from the type-localities, it would be 
necessary to consider all species occurring fro 
coast to Nebraska, Kansas and eastern Texas. Happily for me 
the polymorphic Mexican and Texan series centering about /. 

luci defrayed through a gift from Mr. Bararp Lone. 


m the Atlantic 


Th i £ + 1 
: tho pie 


2 Rhodora [JANUARY 


nudiflora does not extend into the area specially covered, and the 
relatively simple R. Drummondiana and R. Parryi are likewise 
excluded from the present study. I have been concerned only 
with the species which are definitely eastern or which extend 
eastward into the “‘ Manual range”’. 

In studying these relatively few species it has become evident 
that much recent publication upon them has rather glibly passed 
by the need even to look up the original descriptions of the earlier 
recognized species. Thus, although R. humilis Nutt. was de- 
scribed from “rocks in the upland forests and prairies” of 
Arkansas, a plant with oblong-ovate and sessile leaves, the name 
has been transferred by Small and others to a species with rosu- 
late and chiefly spatulate leaves occurring, to quote Small, in 
“Sandy soil, Coastal Plain, Fla. to Miss. and Ga.”’; and the latter 
author says “Stem 1-3 cm. tall’ but correctly identifies as R. 
humilis, in his sense, specimens up to ten times that height! 
Again, R. hybrida Pursh came from Savannah, Georgia, but 
Small restricts it to ‘‘ Pinelands, Fla.”’. As striking a case as any 
is R. parviflora (Nees) Britton, given by Small the range ‘‘Sandy 
soil, ~oantes Plain and adj. provinces, Fla. to Tex., Ind. and 

; the case striking because Dzpteracanthus caliontts. 7. 
nena Nees, basis of Britton’s binomial, came from far 
away from the Coastal Plain, “Ad Ky. fluvium, Julio (Short! 
in herb. Hook.)”’. Incidentally, Nees described it with petioles 
3-6 inches long—‘coroll& vix pollicari, foliis paulo longiori — 
petiolo (3-6-pollicari).”” Such a description can hardly apply 
to a plant with petioles rarely 1.5 cm. long! Nevertheless the 
misbegotten name, R. parviflora (Nees) Britton, promptly came 
into vogue for almost anything, with either petioled or sessile 
leaves and no matter where it grew, until Blake revived KR. 
caroliniensis (Walt.) Steud., which was based in a roundabout 
way upon Walter’s clear description of a characteristic Carolina 
plant. When R. caroliniensis came to the front it replaced R. 
parviflora as a blanket-name for anything, just as, still earlier, 
the name R. strepens L. had covered almost every species in the 
United States. 

The greatest complexity is in the two series of variations which 
together make up the northernmost or, at least, the most north- 
ern-ranging species, R. humilis and R. caroliniensis, for a number 


1945] Fernald,—Ruellia in the Eastern United States 3 


of strictly southern species are relatively stable. The latter 
series of plants, relatively comforting to work with, includes R. 
pedunculata Torr., a clear-cut species radiating from the Ozarkian 
region southward into Louisiana and eastern Texas; two habitally 
similar but morphologically distinct new species, one of pine 
barrens of the Coastal Plain from’ South Carolina (possibly 
Virginia) to Louisiana, the other centering on the Blue Ridge 
and Alleghenies; another newly described species concentrated 
in southern Florida; R. ciliosa Pursh (as I interpret him), the 
characteristic rosulate-leaved plant of central Florida, north to 
South Carolina and west to Louisiana, the species treated errone- 
ously by Small as R. humilis Nutt.; the fleshy plant of sub- 
tropical Florida, R. succulenta Small; and the local and amazingly 
clear long-flowered and large-fruited R. noctiflora (Nees) Gray, 
occurring from eastern Georgia to northwestern Florida, thence 
to southern Louisiana. There is no trouble in separating these 
relatively stable species nor in promptly recognizing the conti- 
nental R. strepens. 

The great difficulty is to find stable characters within the two 
polymorphous species, R. caroliniensis and R. humilis. The 
latter, characterized by somewhat uniform sessile or essentially 
sessile leaves, many flowering nodes and a strong tendency to 
bushy branching, is found through much of the prairie region, 
but in some of its variations it extends eastward to the Blue 
Ridge and in one variety even to northwestern Florida. In some 
areas the corolla may be only 2 cm. long, in others up to 8 cm. 
Whether the corolla is short or long the leaf-outlines present two 
parallel trends, on the one hand elliptic-oblong to oblong- 
lanceolate and (the larger ones) only 1—2.5 cm. broad, on the 
other hand ovate, oval-oblong or elliptic and 2-4 em. broad. 
Furthermore, the pubescence varies from a copious wide-spread- 
ing hirsuteness to glabrescence (the latter especially on the 
Cumberland Plateau). Search at long and baffling periods 
through eight months has failed to bring to light any stable 
characters. The series is certainly an unstable one. I have, as 
the best I can yet do, suggested its recognition as a number of 
more or less segregated geographic varieties with some minor 
forms. 

Similarly with R. caroliniensis, trials of first one character, 


4 Rhodora [JANUARY 


then another, from June to February, have led to optimistic 
moments, these always followed by despair. At one time the 
reactions of the seeds in the diverse trends within the mass of 
material looked like a hopeful character. In Ruellia (at least in 
all of ours) the thin and discoid seeds are superficially very simi- 
lar. When soaked for a few seconds in water they give an im- 
mediate reaction. They promptly send out from the margin a 
continuous film of mucilage and the flat surfaces become covered 
with minute processes. When watched for a minute or two they 
broaden the marginal band of mucilage and the minute processes 
prolong as flexuous mucilage-spiracles, these visibly shooting out 
(as seen by a microscope) and often suggesting slender flames. 
Eventually (after many minutes) the whole surface is plush-like 
and the marginal band breaks up (on drying) into innumerable 
slender spiracles; and finally the spiracles somewhat collapse, 
fall together as domes and eventually subside. During a full 
week of soaking and watching the behavior of seeds, with thou- 
sands of objects to study, I had great hopes that the different 
species and varieties would yield in the behavior of moistened 
seeds something diagnostic. When, however, the whole series 
was comparéd I was forced to give up; the possibility that, in 
spite of its history and development, taxonomy, through Ruellia, 
was entering the field of experimental physiology, was dis- 
couragingly abandoned. I have, then, as the best I can do, 
treated the polymorphous Ruellia caroliniensis as a series of 
geographic varieties, some with pretty definite areas of develop- 
ment, others overlapping, and within these varieties, minor local 
forms in which the abundance, scarcity or length of pubescence 
strikingly differ. 

In this unusually protracted study, in view of the small number 
of species involved, I have had not only the accumulated ma- 
terial of the Gray Herbarium but that of several of the larger 
herbaria and of a number of smaller ones, to the officers or owners 
of which I extend my cordial thanks for the use of their material. 
Throughout the work, both in bibliographic details and in making 
the plates I have had the unlimited and most valuable aid of 
Dr. Bernice G. Scnusert. Even if the keys and descriptions 
fail to convey my ideas of a still perplexing group her photo- 
graphs will make quite clear the chief diagnostic characters. 


1945] Fernald,—Ruellia in the Eastern United States 5 


In the citations, specimens which are in the Gray Herbarium 


are rarely designated as being there, then by G. The other 


herbaria cited are as follows: 


E. Lucy Braun (Braun) 
DuKe University (Duke) 
F. W. HunneweE .t (F. W. H.) 
Missourt BOTANICAL 
GARDEN (Mo) 
New York Botanica 
GARDEN (NY) 
University oF NorTH 
Carouina (NC) 
UNIVERSITY OF PENNSYLVANIA (Pa) 


PHILADELPHIA ACADEMY 
oF ScreNcEs (Phil) 

Dr. P. O. SCHALLERT 
(Schallert) 

Unitep States NATIONAL 
Arporretum (USN 

Unitep. States NATIONAL 
Hersarium (US) 

VIRGINIA POLYTECHNIC 
InstiTuTE (VPI) 


As I understand these species they are as follows. 


Key to RuUELLIA IN THE EASTERN UnitEepD STATES 


a. Main iy terminating in a prolonged i leafy tip, “~ 
1-few at leafy-bracted summits of bran ches or 0 
getatene from the median or jeer ee or ibe pian 
inflorescence much 
b. rte chiefly iematodiienr: bracts of loose cyme linear 
r linear-lanceolate; garden-escape in the Gulf States. Oe 
R. Brittoniana. 
b. Leaves. Lereyeay oblong, elliptic wh ov peduncles 
y dilated bracts; indige 
c. Calyx-sogments — = iance-linear, flat tb the tip, 
mm psule glabrous; larger 
ees with Shades 718. em. ne and 3-9 cm . broad. .2. 


ce ee 


R, strepens. 


road. over branching, t the branches 
simple or forking, 0 bo simple to but slightly 
branched; peduncles from axils 0 


Ww ase slender 
Calyx, abel — ae pilose with slender tpped 
ubesce of Ozarkia 
‘naa nol peruse popes rues 3. R. pedunculata. 
and partially 
ersed slender cystoliths Toker? and ee 28 
oasta 
wakioul: plant of southeastern cere 
d, Stem sew or with say ascending branches; peduncles 
1-flowered, ae or 2(-4) n 
+ aime 


Ba 
& 
28 


linear, 0. 71. 
thence to apex, minutely Rampant ont ae 

eastern Piedm 
pilose-hirtellous ; Appalachian r SR Purehiond. 


species ee eh a EY eet ee 


6 Rhodora : [J ANUARY 


a, Main axis with sessile or very short-peduncled momeruies: or 
owers in the axils of the upper (sometimes median and 
lower) leaves, or the terminal pairs 0 of leaves snreted and 


é. segs ee gas flat, lanceolate, 2-4 mm .. wide, shorter than 
slightly exceeding ac att ‘flowers chie fly or 
wholly cleistogamous, with small closed corollas; wides 
anging erect simple or bat slightly’ ereet-branchin 
ecntinental WES feces as 2a.. R.. strepens, rae cleistantha. 
e. Calyx-segments narrowly gd with croloaged pup slender 
sailors st bristleform tips mostly overtopping the cap- 
; flowers. rarely cleistogamous (except in nos. 5a. 
and 67. mostly showy and expanding.... 
5 Plant arn dimorphic or hetersincrehic: the vernal 
(sometimes later) Pye ao rect and simple or slightly 
branched, poe 1-3 sho leat from axils of 1-3 
ot leaves; later trailing or decumbent 
branches prolonged and bearing dense glomerules ; Of 


anit eisbrons capsules; leaves elliptic, oblong, sie meitet 
olate or narrowly obovate, 1.5—5.5 cm. long, definitely 
short-petioled, firm, with one or bak joie often 
cee covered with slender Sher : ee 


z Plant 1 sak strongly or not at all ina ag flowers all o 
nearly all with expanded corollas or, if most or all 


broad; veins of leaves and usually the 
ae odes white-villous; species of spathenr Costal 
be ae agen ee she kek Cale ws a Aedes od diene we 7. &. ciliosa. 


not ap 

ret epee in no. a fhe wth only minutely 
tem fleshy, often hollow, streak wile simple 
or with strong] ascending branches; leaves fleshy, 


ob 


verglades region of southern Florida....8. R. succulenta. 
h. Stem fe fleshy, solid, firm, eubeteant to rarely 
; leaves membranaceous to firm, - sat 
eshy, rarely cong el linear-lanceolate to broadly 
—— ob long, vee or ovate, pono ig 


Dag 
H 
oF 
= a 
Bs. 
an 
Te 
c 
or 
omy 
eg 
af 
2" 
bene) 
a 
5 
“Hl: 
Ss 


); prim 
with 3-10 remote pairs of saw Sane selade 
to ce- or elliptic-oblong submembrana- 


1945) 


longis 0.5-2 em 
foliosis 
bus; bracteis bracteolisque lineari- vel anguste lanceolato-su ; calycis 


subrigidis an 
hirtellis s supra 


Fernald,—Ruellia in the Eastern United States 7 


ceous puberulent or ey hirtellous leaves 

ng to apex; flowers 1—few from 1—4 upper 

nodes; calyx 2.5-4. yak long, cinereous- 

puberulent or minutely hispidulous; corolla 
c 


m. 
R. noctiflora. 


habit; gine narrowly oblong-lanceolate to 
broa dly Ov: , coriaceous, with blunt 
a ane isch tine, often anal to coarsely 
0 ~ rs on well dev 


glabrous, 1.2-1.5 cm - long; plant of Great 

Aas ins, eastward to Blue Ridge and, an'/ 

western PIGndA, <5. 55 0. 

1. tae. or ras least thé: nrinclial ones, tapering te 

definite petioles (up to 2 gee long), spreading to 

ascending; primary axis 4 flowering 

nodes or, if more, with pais often undulate- 
entate. 

Upper internodes para fi abbreviated, villou 5; 
Thisaaite, puberulen gla brescenti upper 
pairs of leaves approximate or crowded; 

sho 


R. humilis. 


near : 

ae the plant with senonos d villous upper eae 
ternodes, le say! paths ca. R. caroliniensis. 
Upper internodes ongate, ai nutely _pilose- 
p of leaves distant; 


Se 8 6h 97s 


becca u 
-above; capsules pi dhSecsery 
: oa. , Purshiana, forma claustroflora. 


‘Rurtura BrirrontaAna Leonard, emended and validated. 
Steins 
early or 
1 Validated 
RveLiia re hatc Leonard, caulibus subligneis adscendentibus 3-10 dm. altis, 
einde ram 


sually several, subligneous, 3-10 dm. high, subterete, 
quite — glabrous, soon ‘branching, saboowriboscly 


by the following Latin diagnosis: 


eari-lanceolatis utrinque 
m. 


. latis integris vel obscure undulatis; infl 


pedunculis rigide adscendentibus quam foliis subtendentibus 1/3-1/2 breviori- 


segmen 


anguste Janceolatis attenuatis ad apicem © rsaaeigeinie elabris ve. Sees 


libulif. xtus glabro ~s Slee supra 0.9-1.5 cm. lato, 


limbo (expanso) 2.5-5 em. lato; capsulis lan 


longis, retinaculis 14-24; seminibus sa bortiousaetiie 2-2.5 mm. diametro. 


8 Rhodora [JANUARY 


mm. long: corolla Panels ong; cylindric Sele 8-13 
mm. ; the broadly erhinaibalifotin throat pli to but 
aan pilose without, 1.2-2 cm. long, 0.9-1.5 em. broad at 


summit; limb (laid o pen) 2 5-5 em. bead, with sed rounded 
lobes; capsule lance-fusiform, pate 2-2.7 om. long; retinacula 
14-24; seeds ore en 2-2.5 n diameter.—Journ. Wash. 
Sci. xxxi. 96, fig. 1 (1 041) 9 wana ctaknodil but accepted 
oe here validated as ‘ho type, Cryphiacanthus angustifolius Nees 
in DC. Prodr. xi. 199 (1847) in part (the Galeotti specimen from 
Jalapa); not R. angustifolia Swartz, Prodr. Veg. Ind. Occ. 93 
(1788). R. spectabilis Britton in Ann. N. Y. Acad. Sci. vii. 192 
(1893), without diagnosis, only as the Mexican plant was poles 
in his rhage He the type, Cryph. angustifolius Nees, 1. c. 
not as to Par cer ey cited; not R. spectabilis ese ne 
Gard. Dict. iii. 334 (1886). R. malaco <efiiea sensu Small, Man. 
Se. Fl. 1229 (1933), not Greenman in Proc. Am. Acad. xxxiv. 
572 (1909).—Native of eastern Mexico; cultivated and spread 
we eben — meres cultivated ground, borders of ditches, 
fro orida. FLoripa: open places, Arcadia, 
918, Small, ae O00 (NY) as R. patios: Glen St. Mary, 
June, 1923, C. R. Stevens (Mo); roadsides, Ft. Myers, Aug., 
W. M. Buswell (NY) as R. malacosperma. ad ps 
ee gees soil, Houma, Aug. 31, 1913, E. C. Wurzlow (Mo 
US) as R. spectabilis, hole 10, 1913, Wurzlow (NY), ‘dentified ‘ae 
Small as R. malacosp a, by Leonard as R. Tweediana Griseb. 
TEXAS: Sadaed fron: gente San Antonio, Sept. 20, 1901, 
Bush, no. 864 (Mo); orm poe near Polytechnic, Oct. 10, 1916, 
A. R. mb ere (US) as tabilis; Houston, July 10, 1934, 
Sok 11,330, as R. Pond na. SAN Luis Potosi: wet ledges 
hake tog “Micos, July 31, 1891, Pringle, no. 5043, as R. Tweediana; 
eeveig ie rocky sand near river, alt. 200 "feet, near Axtla, 
June 27, 1942, J. N. Weaver, no. 658, as R. Tweediana, corrected 
to R. Brittoniana. Vera Cruz: in the bed of the Colobozo near 
Tantoyuca, April, May, 1858, Ervendberg, no. 104, identified mY 
Asa Gray as Orvphsecoieas angustifolius, by Leonard as 


1945) Fernald,—Ruellia in the Eastern United States 9 


Tweediana; along streams, Tenera, Zacuapan, Dec., 1912, Purpus, 
no. 6162, identified by Leonard as R. ediana; Barranca de 
Panoya, Sept., 1919, Purpus, no. 8409, identified by Leonard as 
R. Tweediana. GuaTEMALA: cultivated, vicinity of Quirigué4, 
Dept. Izabel, Standley, no. 24,307, as R. malacosperma, another 
number (72,225) cited by Leonard as R. Brittoniana in publishing 
that unclarified name. N. B. e Mexican material in the 
Gray Herbarium only here cited. PLATE 839. 

So far as I can find the specific characters of Ruellia Brittoniana 
have never been clearly stated, except partially by Nees who, 
under the name Cryphiacanthus angustifolius, had two quite 
distinct but by him undifferentiated species, and partially by 
Small, who described and had before him R. Brittoniana but 
misidentified it as R. malacosperma. In fact, the entire history 
of the names and the recognition of specific lines in the two 
plants, inadequately treated by Nees in 1847 as a geographically 
bi-centric species, Cryph. angustifolius, is one of discreditably 
opportunist shiftings, without any evident attempt at clarifica- 
tion. The original treatment of the two confused species by 
Nees in DC. Prodr. xi. 199 (1847) was 

C. ancustiroiius, caulescens, foliis lanceolato-linearibus acutis 
integerrimis sessilibus glabris, pedunculo subbifloro folio breviore, 

calycis laciniis subulatis scabris. 2| Ad Xalapa (Galeotti! in h. 

ook.), Entre Rios (Tweedie!). Corolla pollicaris. Capsula 9 lin. 
longa, lanceolata, 16-20-sperma. (v. in h. ) 

The first separation of the Mexican (Xalapa or Jalapa, Galeotti) 
plant and the Argentinian (Tweedie) elements was in 1879 when 
Grisebach, in his Symbolae ad Floram argentinam, in die Abhand- 
lungen Kéniglichen Gesellschaft der Wissenschaften zu Gdtt- 
ingen, xxiv. 259 (1879), took out the Argentinian element, as 
Ruellia Tweediana Griseb. : 


late v. ovato-lanceolata —E [i. e. Prov. Entrerios, as explained by 

Grisebach on his p. 4, Nees having originally cited “Entre Rios”’]. 

In other words, Grisebach, concerned only with the flora of 
Argentina and not the Galeotti specimen from Xalapa (or Jalapa) 
in the state of Vera Cruz, Mexico, based his Ruellia Tweedia ea 
(without adequate description) on the Tweedie specimen cited 
by Nees. Since Grisebach did not use trinomials but regularly 
designated varieties as “‘var.’’ (see the synonym of R. geminiflora” 


10 - Rhodora [JANUARY 


Kunth given by him on the same page as ““R. geminiflora var. 
humilis Gr.”) it may reasonably be inferred that his typonym of 
R. Tweediana “Syn. Cryphiacanthus angustifolius Tweedianus 
Ns.” was intended to mean the Tweedie element of C. angusti- 
folius, Grisebach thus leaving out the Galeotti plant, which, by 
the very sensible but commonly ridiculed “doctrine of residues”’ 
remained as true C. angustifolius. If it be argued otherwise the 
result is somewhat the same, for there was already a Ruellia 
angustifolia Swartz (1788), so that R. T weediana is the first valid 
name under Ruellia. Hemsley, in his monumental Biologia 
Centrali-Americana (Botany), ii. 508 (1882), seems to have 
suspected that the Jalapa (Mexican) plant was not identical 
with the Argentinian element, for he entered 


Ruellia tweediana, Griseb. Symb. ad Fl. Arg. p. 259? 
Cryphiacanthus angustifolius, Nees in DC. Prodr. xi. p. 199, saltem 

pro parte 

SourH Mexico, Jalapa (Galeotti). Hb. Kew. 

The Argentine plant may be a different species. 

But Hemsley did not note any specific differences. Neither 
have those who have rather easily and very carelessly rushed 
into print with substitute-names. Thus, when, in 1893, in an 
enumeration of plants of Paraguay, Britton substituted for 
Cryphiacanthus angustifolius Nees the name Ruellia spectabilis, ° 
he obviously intended the plant of temperate eastern South 
America, not the Mexican element, but Britton’s item was 0 
hastily prepared that it is evident that he did not stop to gain a 
clear understanding of specific lines, nor had he gone carefully 
into the literature. His treatment was as follows: 

Ruellia spectabilis, Britton. 

Cryphiacanthus angustifolius, Nees in D.C. Prod., xi, 199, not 

Ruellia angus{tlifolia, Sw. 

Caballero (461). January. 

This species has branching stems 10-12 em. high, linear, sessile leaves, 
and flowers larger than in no. 323 [Ruellia Morongii Britt., new name 
for Cryphiacanthus acaulis Nees, not R. acaulis R. Br.], otherwise much 
the same. Occurs on the railway track. 

Since Cryphiacanthus angustifolius (in its inclusive sense) con- 
sists of erect, caulescent plants up to 1 m. high, with remote 
pairs of linear-lanceolate to broadly lanceolate leaves and 
glabrescent to only minutely glandular-hirtellous calyx about 
half the length of the capsule or shorter, while Cryphiacanthus 


1945] Fernald,—Ruellia in the Eastern United States 11 


acaulis Nees, basis of Ruellia Morongii, is, as described by Nees 
and partly by Britton and as shown by all specimens, acaulescent, 
“Habitus Primulae” (Nees), with the basal rosulate leaves 
oblong-ovate, -obovate or subspatulate, and the calyces copi- 
ously villous-hirsute and equaling to exceeding the capsule, 
Britton’s characterization of the two as “much the same”’ indi- 
cates a rather offhand understanding of a few conspicuously 
different species. Dr. Britton also overlooked the fact that, in 
his well known Dictionary of Gardening, iii. 334 (1886), George 
Nicholson had published a R. spectabilis (Hook.) Nicholson, 
based on Dipteracanthus spectabilis Hook. Bot. Mag. t. 4494 
(1850), an Andean plant of a different section, with flowers ses- 
sile in the axils of ovate leaves, ete. Furthermore, Britton 
evidently overlooked R. Tweediana Griseb. (1879), the name he 
should have taken up for the Paraguayan as well as Argentinian 
Cryphiacanthus angustifolius. Had he looked up Grisebach’s 
Symbolae he would not, on the same page with his R. spectabilis, 
have published as new R. Tweedyt (Nees) T. Anderson in Herb. 
Kew., based on Blechum Tweedyi Nees, for, by the rules promul- 
gated and followed by Britton, the substantive-genitive personal 
names (such as Tweedyi) and the adjectival forms (such as 
Tweediana) could not both be used. By the International 
Rules, vigorously fought by Britton, his combination R. Tweedyi 
(Nees) T. Anderson ex Britton is rescued. Whether the type of 
R. Tweedyi is a Ruellia is much more doubtful. The genus 
Blechum is so very different from Ruellia that it would be sur- 
prising if Nees, who monographed both genera, did not know it. 
Incidentally, but of real importance, the original Blechum 
Tweedyi came from Panama. It would be very surprising, to 
say the least, if the same species (even if not a Blechum, a genus 
apparently unknown in Paraguay) were found also in Paraguay. 

The next step in the tortuous history of the much abused 
typonym, Cryphiacanthus angustifolius Nees, was when Leonard, 
noting that Britton had slipped in publishing a second Ruellia 
spectabilis, gave, in Journ. Wash. Acad. Sci. xxxi. 96, fig. 1 
(1941), another name: 

ia brittoni . NOV. Fig. 1 
joint, peukeee Soar DC. Prodr. 11: 199. 1847. 
I 788. 
Ruellia spectabilis Britton, Ann. New York Acad. 7: 192. 1893; not 
Nichols, 1886. 


12 Rhoders [JANUARY 


A single cultivated plant, from Guatemala, was cited and a 
very characteristic figure of the Mexican plant, with long- 
attenuate upper leaves inclined to overtop the subcorymbiform 
inflorescence, was given as fig. 1. Leonard gave no statement of 
characters nor any indication as to whether he was accepting 
Cryphiacanthus angustifolius in the original inclusive sense of 
Nees or whether he restricted it to the Mexican element left to 
stand for it when Grisebach withdrew the Argentinian element as 
Ruellia Tweediana. It is unfortunate, if he intended R. Brit- 
toniana (Britton having published only on the Paraguayan plant) 
to stand exclusively for the quite different North American 
species, that he did not give any word of clarification, for the 
North American species, at least in the Gray Herbarium and the 
Britton Herbarium, had been annotated by Leonard as R. 
Tweediana. Incidentally, had he looked up the first publication 
of R. spectabilis he would have found that its author was Nichol- 
son, not “Nichols”. Only by accepting the possible and perhaps 
probable interpretation that, by removing the Tweedie element 
from the mixed originals of Nees as R. Tweediana, Grisebach had, 
by the “doctrine of residues”’, left the Mexican element as true 
Cryph. angustifolius—only by this interpretation can we possibly 
save for the Mexican plant the inappropriate name R. Brittoniana, 
which may or may not have been intended for it. I am following 
this interpretation merely in order to avoid publishing still 
another name and thus further increasing the confusion. If the 
alternative reasoning were adopted the Mexican species (culti- 
vated and naturalized eastward to Florida) would require a new 
and clearly applied name, since none of the authors, from Grise- 
bach on, who have hastily proposed new names in this relatively 
simple pair of species, has recognized the elementary requirement 
of sound taxonomy, of accurately defining their species and ex- 
plaining what they meant. 

I have stated above what I consider the specific characters of 
Ruellia Brittoniana as here validated, and in PLATE 839 its diag- 
nostic characters are shown. In pLaTE 840 I have shown some 
of the differential characters of R. Tweediana; and in the following 
paragraph I indicate some of its other claims to recognition as an 
endemic species of temperate eastern South America, the name 
R. Tweediana thus being validated: 


1945] Fernald,—Ruellia in the Eastern United States 13 


R. Tweepiana Grisebach, caulibus glabris; foliis inferioribus 
lanceolatis vel lanceolato-ovatis , Supernis lanceolatis subacutis 
vel obtusis saeco ciliolatis, majoribus 5-12 cm. ionate in- 
tegris vel undulatis; inflorescentiis elongatis subthyrsoideis; 
calycibus glautiiloso hictallis segmentis lineari-subulatis a, 
atis ad apicem acutum hirtum; vbr 3-4 em. longis e 
valde pilosis, fauce supra 5-10 mm. lato; capsulis AE BeolAe 
fusiformibus 2—2.5 cm. longis abu 40. 

Since Ruellia leiidoktien has been mnistalean by Small and his 
followers for R. malacosperma Greenm. it should be noted that 
the latter species differs in the following characters: young inter- 
nodes of stem and young leaves villous-hirsute, becoming glab- 
rate; leaves oblong or elliptic-lanceolate, often undulate-dentate, 
the primary ones slender-petioled, strigillose-lineolate; calyx 
lineolate, with lance-attenuate sharp-pointed segments becoming 
1.5-2 em. long; corolla 3.5-5 em. long, essentially glabrous with- 
out; capsule 2.5-3 cm. long; seeds orbicular, broadly obovate or 
elliptic, 2.8-3.3 mm. long. The plant called R. malacosperma by 
Small (as represented in the Britton Herbarium) is not that 
species; all that I have seen of it belongs to the frequently 
cultivated R. Brittoniana. 

2. R. strepENS L., as emended by L. in nk 1. Stems 1-few 
from a knotty rhizome, 0.2-1.1 m. high, simple or with few 
ascending branches, Signage shale. Enc pilose (often in 


14 Rhodora [JANUARY 


elxxii. (1791); Willd. Sp. iii. 363 (1800); Pursh, Fi. Am. Sept. ii. 
420 (1814); LeConte in Ann. Lye. N. Y.i. 1 40 (1824) ; and later 
os Ayia Dip teracants strepens (L.) Nees in Linnaea, 
xvi. 292 (1842) and in DC. P rodr. xi. 121 (1847), including var. 
ae ahi Nees, 1. c. (1847), see Loghhgerry Haley l. ¢, (122 
(1847) and var. strictus (Nees) Nees, 1. c. (1847), in ee D. 
strictus Nees in Linnaea, xvi. 293 (1842). R. biflora Balbis ex 
Nees in DC. Prodr. xi. 122 (1847) in synonymy, nomen only. 
R. foliosa Schweinitz ex Nees |. c. in synonymy, nomen only. 
R. oblongifolia Kinn ex Neen. ]. c. in synonymy, nomen only. R. 
vincaeflora ex Nees, 1. c. in synonymy, nomen only.—Low 
woods, bottomlands, wooded swamps, etc., chiefly in basic or 
calcareous soils, South Carolina to eastern Texas, northeast and 
north to north-central New Jersey, southern Pennsylvania, 
central Ohio, Indiana, Illinois, southern Iowa and eastern Kansas. 
Fl. mid-M ay—July (rarely — —Octob er). The following, from a very 
much larger series, are characteristic. New JERsEyY: New 
Brunswick, ‘‘common”’ June, 1894, F. H. Blodgett (NY). PENN- 
sylvania: LANCASTER Co.: rich wooded hillside along Conestoga 
ie 1 mile south of Bausman, Louise F. A. Tanger, no. 3270 
(Pa, Phil); banks of Conestoga, near Lancaster, 1838, W. W. 
raat (Phil), June 17, 1859, Porter: near Columbia, S. W. Knipe 
Phil); ‘fon an excursion to ‘Safe Harbor”, June 18, 1859, Joseph 
Crawford (Phil). CUMBERLAND Co.: creek-bankside, Camp ill 
Borough, H. L. Plasterer, as D. E. and Dorothy Wade, no. 1727 
(Pa). FRANKLIN CO.: Mercersburg, June, 1844, Porter (Phil). 
DELAWARE: without statement of locality, 'N’ uttall (Phil). 
MARYLAND: CECIL co.: Bald Friar, July 4, 1907, EF. B. ates el 
hee MONTGOMERY CO.: banks of Potomac, June 6, 1881, 
J. D. Smith; nehere Falls, C. S. Williamson; High I Island, June 6, 
1881, C. S. Sheldon (US). District oF CoLUMBIA: Potomac 
Flats, Chain Brides, June 13, 1897, Kearney (NY); Canal 
District, June 9, 1897, Steele (US). West VIRGINIA: JEFFERSON 
Co.: along Shenandoah River, near Charlestown, R. F. Martin, 
= 200 (USNA). caBELt co.: dry hillside, Huntington, Gilbert, 
0. 123 (Mo, Pa). wayYNr - Buffalo Creek, Plymale, no. 445. 
poate ISLE OF WIGHT Co.: base of rich calcareous wooded 
slo opes by Burwell’s Bay, sae River, below Rushmere (Fergus- 
son’s Wharf), Fernald & Long, no. 13 463. PRINCE GEORGE CO.: 
swampy woods, bottomland of Powell’s Creek, Garysville, 
Fernald & Long, n nos. 8472 and 8854. cLARKE co.: Castleman 
Ferry, June 13, 1937, O. M. Freeman (USNA). FREDERICK CO.: 
low woods, Cedar Creek, Meadow Mills, Hunnewell, no. 13,712. 
SHENANDOAH CoO.: along stream, north of Short Mt., Allard, no. 
5087. PAGE Co.: rats soe eastern foothills of Massanutten 
Mountain, W. H. Camp, no. 1386 (NY). ROANOKE co.: Roanoke 
River, south of Roaioks Small & Heller, no. 431 (Phil). MoNntT- 


1945] Fernald,—Ruellia in the Eastern United States 15 


GOMERY Co.: limestone soil, edge of woodlands just north of 
Price’s Station, A. B. Massey, no. 5065. SourH CaRoLiNa: 
BERKELEY CO.: oes oe May, H. W. Ravenel rie 
co.: Aiken, Sept. 17, 1885, H. W. Ravenel. Grorata: withou 
stated locality: ‘of te Savannahs”, ie Conte Phi; “Mts. 2 
Georgia’, Chapman (US). WALKER co.: dry round, Chicka- 
mauga Park, May 25, 1911, J. R. Churchill. euetinom from 
Polk County, Florida, Mrs. J. M. Milli gan (US) were presumably 
of ape ees Chapman canhadoe ‘Florida”’ in his stated 
range, but his “‘ R. strepens’’ was made up of several other species. 
There is no raietal from Florida in the Britton Herbarium, rich 
in specimens from that state.) OHIO: ATHENS CO.: Atl ens, 
J. P. Drushel, no. 6623 (Mo). LAWRENCE Co.: open woodlands, 
Coal Grove, "Biltmore Herb., no. 4500 (US). FRANKLIN CO.: 
Columbus, Sullivant MONTGOMERY co.: Dayton, July 7, 1879, 
L. V. Morgan (US). BUTLER CO.: moist woods, Oxford, June 30, 
1910, Cuerhalts (Mo). HAMILTON co.: near Cincinnati, June 15, 
18 879, C. G. Lloyd (NY, aay open woods on hillsides, Anderson’ s 
Ferry, June 14, 1905, Braun (Braun). (Specimens with 
the label “Plants of ee Gk Ohio’’, and marked as from Lucas 
County, bear the memorandum “Obtained from Mr. Burger of 
Toledo; never found it in northern Ohio”’.) INDIANA: WELLS. 
Co. : moist banks of Wabash River, June 21, 1905, Deam (US). 
MONTGOMERY CO.: Crawfordsville, June 23, 1892, Rose (US). 
MARION Co.: woods along White River, Scott McCoy, no. ee 


viGo co.: Terre Haute, June 1, 1889, B. W. Evermann (US). 
BARTHOLOMEW Co.: creek-bottom north of Elizabethtown, Deam, 
no. 34,266. JEFFERSON co.: Hanover, 1874, Coulter (Phil). 
KENTUCKY: ScorT Co. : along Elkhorn Creek, Stamping Ground, 
J. W. Singer, no. 219 (US). FRANKLIN Co.: open woods, Farm- 
dale, June 16, 1879, H. R. Bassler Mars SPENCER CO.: near 
High Grove, E. L. Braun, no. 3259 (Braun). WARREN CO.: 


Mo). 
oa a: 345, LYON me ; Kuttawa, Eggleston, no. 4524 


TENNESSEE: KNOX CO.: woods, Knoxvi e, Ruth, no. 591 
(Mo) and 721 (US), Pence no. “1, 301 (Phil). genta co.: 
Chickamauga Park, May 27, ae R. Churchill. DAVIDSON 


co.: bluffs below Nashville, ienianes Herb., no. 4500¢ (US); 
Nashville, Eggleston, no. 4444: limestone cliff of Cumberland 
River, Nashville, thera: no. 11,411 (Phil). eee co.: 


limestone bank, Erin, E. B. Harger, no. 7866. LAKE Co.: dense 
shade of sandy woods, Reelfoot poet < R. Sieolion, no. 2159 
(US). ALABAMA: JACKSON co.: dry Stevenson, Biltmore 


Herb., no. 4500 (US); Sand ioeniai ies Herb., no. 45004 


16 Rhodora [JANUARY 


(US). CLARKE co.: dry copses and hillsides, Thomasville, April 
27, 1888, C. Mohr, as R. ciliosa, var. ambigua (US). Misstss1prt: 
LEE CO.: Tupelo, 1914, Henshaw (U SNA). OKTIBBEHA Co.: low 

woods north of | Starkville, C. A. & Una F. Weatherby, no. 6309. 
ILLINOIS: KANKAKEE CO.: Kankakee, C. C. Crampton, no. 212 
(US). PEORIA co.: rich rg Peoria July, 1903, F. EF. Mc- 
Donald. HANCOCK co.: Augusta, S. B. Mead. MACON CO.: 3 
miles east of Decatur, lobey. ¢ ox 2486. PIKE Co.: Mississippi 
levees, East Hannibal, June 3, 1913 (Mo). RICHLAND Co.: 
Parkersburg, June 9, 1902, Robt. Ridgway (US). MARION co.: 
Salem, June, 1860, M.S. Bebb (P hil). sv. CLAIR co.: woods, St. 
Clair Co. , Sept. 20, 1878, Eggert; Cahokia, June 11, 1890, AS 
Hitchcock (Mo); East Carondelet, June 4, 1875, Eggert (Mo, US). 
JACKSON 0.: black rich soil, bottoms of Big ‘Muddy and Crab 
Orchard Creeks, John McCree, J¥.; D0. 775 (Mo.). Iowa: 

904, J. 


0). LINCOLN co.: rich soil, Winfield, June 7, 1916, John 
Davis, no. 1406 (Mo). Marton co.: Scipio Bluffs, north of Hanni- 
, John Davis, no. 1491 0). ST. LOUIS co.: Creve Coeur 
Lake, June 12, 1914, M. W. Lyon (Mo); Meramec Highlands, 
June 13, 1909, W.W. Ohlweiler (Mo); Allenton, G. W. Letterman, 
many collections (Mo); rich woods, Allenton, May 29, 1918, 
J. R. Churchill. serrerson co.: D eSoto, June, 1887, ok 
Hasse; hee Jo. Hi; ieee p no. 2005 (Mo). FRANKLIN CO.: 
Pacific, Greenman, no. 3895 (Mo). SHELBY co.: rich woods near 
pale Palmer & Steyermark, no. 40,908 (Mo). BUTLER CO.: 
w woods along Mud Creek, northwest ‘of Rombauer, Steyermark, 

si 11,420 (Mo). Boone co.: Rock B Bridge, er 1926, H. 
Ricket fone PHELPS Co.: Jerome, June 1, rie 8 H. Kellogg, 
4 (Mo). SALINE co.: Sweet Springs, Pde 20, 1886, Wm. 
7, lai (Mo). GREENE co.: Springfield, June 11, 1887, J. 8%. 
Blankinship (Mo). stone co.: rich hillside woods, Galena, 
E. J. Palmer, no. 5774 ory US). pavigss co.: dry banks, 
Pattonsburg, Bush, no. 13,587. JOHNSON co.: rich woods 
and thickets, limestone bill Columbus, EZ. J. Palmer, oi 

36,697 ( Mo). BARRY CO.: barrens, Shell Knob, Bush, 
15,596 (Mo). Jackson co.: Independence, Bush, no. 39 (Pa), 
CASS CO.: bottoms, June 23, 1864, G. C. Broadhead (Mo). JASPER 
co.: woods, Webb City, Bush, no. 528 (Mo), H. J. Palmer, no. 
528 (Mo). ARKANSAS: CRITTENDON CO.: bottomland, Hulbert, 
Demaree, no. 11,372.’ PHILLIPS co.: Crowleys Ridge, Helena, 

le 


Demaree, no. 19 ,240 (NY, Mo). putaski co.: Little Rock, 
H. E. Hasse (NY); swampy Arkansas bottoms, Little Rock 
Demaree, no. 17,321 (Mo). KANSAS: LEAVENWORTH Co.: woody 


ravines, Fort Leavenworth, June, 1854, F. V. Hayden (Mo). 
WYANDOTTE Co.: low woods, May 30, 1897, K. K. Mackenzie 


1945] Fernald,—Ruellia in the Eastern United States 17 


(NY). CHEROKEE Co.: woods along Shoal River, near Schimmer- 
oo Pk., no. 20,178, from Kansas State College (NY). RILEY 
: low woo ods, JB. Norton, no. 387; Manhattan, June, 1886, 


Sagar dasa Se mouth of Illinois River, Goodman & Barkley, 
RE Co.: low woods, Poteau, E. J. Palmer, no. 
3075 (Mo). nak co.: Sapulpa, June 2, 1924, Cie: Willkame 
KAY co.: woods, Tonkawa, G. W. Stevens, no. 1869. OKLAHOMA 
co.: wooded creek-bottom north of Edmond, Waterfall, no. 1975. 
POTTOWATTOMIE CO. : in small valley, St. Louis, Mortimer Faulkner, 
no. 106 (Mo). MurRRay co.: Davis, W. H. Ennig, no. 683 (Mo). 
Texas: without stated locality, Drummond, no. 259 (cited by 
Nees under his Dipteracanthus strepens, vars. strictus and peduncu- 
latus and under several other species and varieties!); Coombs 
Branch, Reverchon (Mo). DALLAS co.: woods, Dallas, May 6, 
ey Beverchon (Mo); moist woodlands, Dallas, Biltmore Herb, 
no. eee vicinity of Dallas, Mc ary R. Stephenson, nos. 
91 Gk a (US). TARRANT CO.: rich woods near Trinity River, 
Lake Worth, Ruth, no. 318 (Pa, Phil, US). Frorr BEND Co.: 
Richmond, W. L. Bray, no. 118 (US). Puate 841; map 1. 
Forma CLEISTANTHA (Gray) 8S. McCoy. Flowers and 
abundant fruits borne in sessile or subsessile glomerules from the 
upper and often from most pat of the axils, es stem only 
occasionally branching: calyx-segments often more pubescent 
and much shorter than te but Tittle longer than eles ae 
relatively small, usually reduced to a slender closed tube 
em. long, pale ‘to creamy, but sometimes partially ede 
or even eee and terminal: capsules abundant, usually plumper 
and shorter than in typical form of ane) retinacula mostly 6 
or 8: seeds suborbicular to elliptical, 3-4 mm. a Oe —Am. Bot. 
xlili. 24 (1937). Dipteracanthus paca tig Engelm. & Gray in 
Bost. Journ. Nat. Hist. v. (Pl. a amigo , 49 (1845). 
Hygrophila illinoiensis Wood in Bull. Torr. Bot. Cl. v. 41 (1874). 
Var. cleistantha rok Syn. Fl. N. Am. ii. 327 «iszs). R. oc 


EST VIRGINIA:, “OHIO co.: thickets near Wheeling Creek, east of 
Wheeling, July 22, 1909, Mac Elwee (Phil). VirGINIA: CHARLES 


18 : Rhodora [JANUARY 


city co.: alluvial woods along is ate Harrison Point, 
' Fernald & Long, no. 9150. PRINCE GEORGE COo.: wooded swamp 
by James River, south of Indian Point, Fernald & Long, no. 


. Riv 
Hunnewell, no. 17,872 (stems, deeply covered by freshet-silke 
subligneous, with strong branches, and shortened and firm 
leaves). ROCKBRIDGE co.: Natural Bridge, Sept. 14, 1907, 
E. B. Bartram (Phil). Onto: FRANKLIN co.: Gahanna, Oct. 19, 
1903, O. E. Jennings. WARREN CO.: moist rich soil along Little 
Miami River, South Lebanon, H. B. Harger, no. 8010. INDIANA: 
GIBSON Co.: low woods bordering Eggwood Pond, Deam, no. 


: SON CO.: woode 
hapin, Wherry & Pennell, no. 13, 673 (Phil). EDMONSON CO.: 
wooded alluvial flat of Green River, Mammoth Cave, #. L. 


SEE: SHELBY Co.: Memphis, Oct. 20, 1850, Fendler. ALABAMA: 
WITHOUT STATED LOCALITY: Buckley (paratype of Dipteracanthus 
micranthus Engelm. & Gray). Ler co.: “N. W. of Lee Co.”, 
June 24, 1897, F. 8. cae (NY). ILLINOIS: CHAMPAIGN CO.: 
Urbana, Oct. 4, 1880, A. B. Seymour (Duke). wasasH co.: Mt. 
wo: 1874, J. Schneck: (isotypes of Hygrophila illinoiensis 
. HARDIN CO.: low woods, Elizabethtown, FE. J. Palmer, 
mete 17 5023 (Mo). Iowa: HENRY co.: Mt. Pleasant, Ec dhs Mills, 
no. 1854 (Mo). Missouri: ST. CHARLES CO.: Watson, Wm. 
Trelease, no. 453 (Mo). st. Louis co.: St. Louis, Sept. 1845, 
Engelmann (paratypes of Dipteracanthus micranthus). IRON CO 
moist shady ground, Iron Mountain, Sept. 1897, Colton Russell 
(Mo). MISssIssIPPI co.: rich swampy woods, Three States Timber 
Tract, southwest of Wolf Island, Steyermark, no. 8761 (Mo). 
OZARK Co.: thickets along creek, near Bakersfield, E. J. Palmer, 
no. 32,872 (Mo). TANEY Co.: woods, Swan, Bush, no. 697 (Mo). 
SULLIVAN CO.: Pawpaw Junction, Sept. 15, 1893, Bush (Mo). 
HICKORY Co.: low woods along Pomme de Terre River, northeast 
of Elkland, Steyermark, no. 24,514 (Mo). stv. cLAR co.: low 
woods around White Sulphur ‘Spring, Steyermark, no. 24,401 
(Mo). pawuas co.: base of slope apy Mangua River, south- 
west of Long Lane, Steyermark, no. 24,231 (M a) VERNON CO.: 
low open woods along creek, near Deerfield, Palmer & Steyer- 
mark, no. 42,140 (Mo, NY). mcponaxp co.: low ground, Noel, 
almer, no. 4069 (Mo, US). ARKANSAS: MARION CO.: 
bottoms of White Ri iver, Flippin, Demaree, no. 20,640 (Mo, pre 


(US). carRoxt co.: Eureka Springs, E. J. Palmer, no. 4439 
(Mo). HEMPSTEAD co.: woods, Fulton, Bush, no. 984 (Mo). 
LAFAYETTE Co.: Spirit Lake, AA. ee Heller, no. 4118. 


ens 


me i mnmiieiiiniiiimmniietn, _.ommmamguimeimmmiiiemmamm., _ceupmmmmmmmantn = 


| 
| 
| 
| 
| 


1945] Fernald,—Ruellia in the Eastern United States 19 


LOUISIANA: WEST FELICIANA PARISH: deciduous woodland, 
Catalpa, Pennell, no. 4308 (NY). OKLAHOMA: ROGERS CO.: 
Aa Nata Bush, no. 429 (Mo). JOHNSTON CO.: open woods near 
Tishomingo, . Houghton as Stevens, no. 3342. PAYNE oo 
Stillwater, Eugene Blea. no. 90 (Mo). TEXAS: WITHOU 
STATED LOCALITY: Drummond, no. 202 Gere en hat following). 
HARRIS Co.: rich shaded bottoms around H n (data wit 
sheet in Herb. Mo), Lindheimer, Fasc. I], no. 300. (type ay 
isotypes of GREEN micranthus). BRAZORIA CO.: WoO 
Columbia, Bush, no. 1842 (Mo); San Bernardo, June 28, 1923, 
Tharp. JACKSON co.: Lavaca River, Aug. 29, 1941, Tharp. 
PLATE 842, 

* As originally published in Species Plantarum (1753) Ruellia 
strepens (from strepo, to rustle, presumably from the dehiscing 
of the capsules) was a mixture. The plant of the Linnean Her- 
barium, which Linneaus had before him, has not been available 
and cannot be until after “‘the duration”. Neither can I discuss 
the specimens cited in other Linnean works. The name was 
taken over from Ruellia strepens, capitulis comosis of Dillenius, 
Elth. ii. 330 (misprinted by L. as 300), t. 249, fig. 321 (our 
PLATE 863), a wholly different plant from that here treated, one of 
the species (our no. 11) with relatively low hirsute stem, pubes- 
cent oblong leaves, dense glomerules of relatively small flowers 
crowded in the upper axils, and the calyx-segments narrowly 
linear. The confusion prevailed for some decades (before and 
after 1753) but in 1771, in his Observationes in Species Plantarum 
cum Emendationibus et Animadversionibus, Mantissa Altera, pp. 
315 et seq., Linnaeus redefined Ruellia strepens (p. 422) to stand 
only for the present species «|. Peduncult oppositi, 
laterales breves, triflori. prion 2 oppositae, etiam 2 sub 
singulo flore lateral: Calyx 5partitus, lanceolatus” etc. He 
thus threw out the wholly different plant of Dillenius (with 
abundant fruit which, when pressed, promptly rustles) and re- 
stricted the name to the showy-flowered and usually infertile or 
only weakly fertile typical R. strepens which, except in the cle- 
istogamous state (unknown to Linnaeus), rarely gets a chance 
to rustle! Schkuhr, Willdenow, Pursh, LeConte, Torrey, Gray, 
Engelmann, Nees, Britton and all others have consistently 
adopted the redefinition made in Mantissa Altera, and only con- 
fusion would result if the pre-Linnean and confused application 
of the name were forced. Our species, preeminently of calcare- 


20 Rhodora [JANUARY 


ous bottomland and bases of limestone bluffs, with great con- 
centration in the Mississippi Basin (map 1), pushes down to the 
Atlantic area along the Susquehanna, Potomac, James, Santee 
and Savannah River systems. It is on the upper Roanoke, and, 
presumably, search may bring it to light farther down that 
valley, even in northeastern North Carolina. In the great ac- 
cumulation of material before me, from some of the more repre- 
sentative larger herbaria, there is no evidence that it is common 


in either North or South Carolina; and from Georgia I have seen | 


it only from tributaries of the Tennessee (thence the Mississippi) 
River, although (since it has been found at Aiken) it is probably 
along the Savannah in Georgia. The Ruellia strepens, capitulis 
comosis of Dillenius was raised from seed sent from Carolina 
and flowered in 1726: ‘Nata fuit haec species e seminibus 
Carolinensibus, & Septembri mense primum floruit anno 1726, 
sequentibus autem annis tota fere aestate’. This plant (our 
no. 11), abundant in eastern North and South Carolina as well 
as reaching eastern Virginia, is inclined to grow in slightly dry and 
rather acid soils. 

It is a very striking fact that the accumulated material before 
me shows forma cleistantha regularly and abundantly fruiting, 
while the typical form, with few peduncles from a few median 
axils and few showy, expanded flowers, is largely sterile. Of the 
270 sheets of typical R. strepens before me only 12 (42% per cent.) 
show 1 or 2 developed capsules (PLATE 841, Fig. 4); all of the 168 
sheets of forma cleistantha are loaded with fruit or show the 
possibility of it. 

When he reduced Gray’s var. cleistantha to the rank of a form 
Mr. Scott McCoy reported on plants brought into the garden: 
“Each June it bloomed as the species and each fall it bore 
cleistogamous flowers as does the so-called variety cleistantha 
Gray.” Further checks should be made in other regions for, if 
all the material in the United States National Herbarium, the 
Torrey and Britton Herbaria of the New York Botanical Garden, 
the herbaria of the Missouri Botanical Garden, the Philadelphia 
Academy of Sciences, the University of Pennsylvania and several 
smaller collections, added to the representation in the Gray 
Herbarium, can be taken (and I believe it can) as a fair average, 
there are some very important characters distinguishing the two 


; 
) 
| 
| 
| 


1945] Fernald,—Ruellia in the Eastern United States a1 


plants which one would not expect to find if forma cleistantha 
always develops from individuals which early in the summer 
were typical R. strepens. Typical showy-flowered R. strepens 
bears 1-3 flowers on few leafy-bracted peduncles from the median 
axils; forma cleistantha has the flowers more densely crowded in 
nearly sessile glomerules, usually from many, including the upper, 
axils. Of the 270 sheets of typical R. strepens before me 145 
(nearly 54 per cent.) have elongate median peduncles 2-10 cm. 
long; when rarely such plants fruit (July 25, Va., Fernald & Long, 
no. 13,463; July 22, Ky., Braun, no. 3259; June 25, Mo., Hasse, 
no. 1094; July 7, Mo., Steyermark, no. 11,420; July 13, Mo., 
Palmer, no. 8275; August 27, Kans., Norton; etc.) they show no 
incipient sessile glomerules in the upper axils, such as one would 
expect if they always change to forma cleistantha. The peduncles 
are still there, up to autumn. Of the 168 sheets of forma cle- 
istantha only 20 (11.5 per cent.) have such peduncles (not counting 
branches with subsessile glomerules). Furthermore, very many 
specimens with only glomerulate fruits in the middle and upper 
axils were collected pretty early in the season, June 17-August 18 
(W. Va., McElwee; Ky., Price; Ill., Eggert; Mo., Palmer, no. 1310; 
Ark., Heller & Heller, no. 4118), their fruiting period overlapping 
the flowering period of typical R. strepens. It must be evident, 
then, that not always do typical early and showily flowering 
plants of R. strepens change late in the season into forma cle- 
istantha; if they did so a much larger percentage of the latter 
would retain the elongate peduncles of the former, and the 
former, late in the season, would regularly bear crowded fruits 
in the upper axils. The problem is a promising one for the ex- 
perimenter. Do the abundant seeds of the cleistogamous plant 
reproduce only the cleistogamous form or do they equally yield 
the typical showy-flowered and largely infertile plant? Carefully 
checked and numerous cultures are necessary before we can say 
with finality. 

. R. pepuncunata Torr. Stem 1-7.5 dm. high, slender, 
firm, obtusely quadrangular or subterete, puberulent or minute 


: . 
habit: leaves ovate to ovate-oblong or lanceolate, short-petioled, 
tapering from slightly above base, pale green, puberulent, 


22 Rhodora [JANUARY 


entire or very shallowly undulate; those of primary axis (above 
the rounded or obovate lowest ones) 3-11 em. long and 2—4.5 em. 
broad, the rameal smaller: ees solitary at tips of simple 2- 
bracted peduncles or loosely cymose on the ee the cymes, 
when developed, 2-several-flowered: calyx ments linear- 
filiform, 0.5-1 mm. wide at base, thence baaititg to very slender 
often flexuous tips, in maturity 1-3 om. long, closely ¢ age 
hirtellous. with Bender upped spreading pubescence: coro 
blue-violet, 2.5-5.5 em. long; the ce tube about ound 
the ampliate ‘brat OvAEY a and capsule ee 
the capsule 1-2 cm. long; a tinea iaialky 6 or 8: seeds orbicu- 
lar or suborbicular, cinereous, 2:5-3.5 mm. dati —R. 
pedunculata Torr. ex Gray, Syn. Fl. N. Am. iit, 325 (1878).— 
‘Woods, bluffs, rocky slopes, Baars open fields, etc., in calcare- 
ous to circumneutral soil, western n Louisiana and eastern Texas, 
north to southern Illinois, eastern and south-central Missouri 
and eastern Oklahoma. The dolléwing are representative. 
ILLINOIS: JACKSON co.: Murphysboro, Benke, no. 4648 (US); 
mesophytic woods, ne June 20, 1903, ’ Gleason; dry up- 
land or rocky woods, Grand T Tower, Gleason, nos. 1793, 2654, 
2655 (all as R. strepens) ; dry rocky. pneeione hillsides, ‘Grand 


Tower, Gleason, no. 9007 (NY). JOHNSON Co.: rocky, tei 
Tunnel Hill, June 27, 1902, eee Sehneck (NY), tol fae 
sandstone bluff, Cobden, May 1902, F. S. E tiny). 


(In Herb. Duke Univ. the ere is a pacman of R. ooduacuieie: 
bearing a label, “Ruellia strepens, Nees. oe Ill. Oct. 4, 1884, 
W.” with the hea ading “Herbarium of Bhi n B. Waite” and, 
print dod above it, “Herbarium of A. Be Bomoni?. Since R. 
strepens is well known from the region ‘of Ur bana, where fre- 
quently collected, while there is no other evidence of R. peduncu- 

ata from northeast of the southwestern corner of Illinois, it is 
probable that in the wanderings of this material some transfer o 
abels has occurred]. Missouri: JEFFERSON Co.: rich woods in 
ravine, southwest of Crystal City, =e teks no. 1357. Leenel 
dry copses, DeSoto, May 30 1887, Has e (Mo, US). SsTE 
GENEVIEVE CO.: Bloomdale, J: #: 'Kellogg, no. 2004. CAPE 
GIRARDEAU Co.: wooded limestone slopes, Hickory Ridge, west 
of Delta, Sesernaek no. 20,811 (Mo). puNKLIN co.: Campbell, 
unc 0. 34 


ncommon, Bush, 3 (Mo, NY). Manson co.: rocky open 
woods, near Fredericktown, E. J. er, no. 31,608 (Mo) 
Ww co.: low woods in Happy Hollow, north of Kime; Steyer- 


no. 11,422 (Mo). tron co.: Arcadia, Greenman, no. 3750 (Mo, 
Phil); rocky open woods near Iro nton, LH. J. Palmer, no. 18,111. 
REYNALDS Co.: cherty slopes, south of Oates, Steyermark, no. 


neem 


TT " 


-_ —".' - ins - ahneites 
oa — mien a oe a _— fo “ai - ~ -~ 


1945] Fernald,—Ruellia in the Eastern United States 23 


19,724 (Mo). carrer co.: rocky woods, Van Buren, J. H. 
Kellogg, no. 15,300 (Mo). PHELPS co.: Jerome, June 11, 1914, 
Kellogg (Mo). SHANNON Co.: rocky woods, Monteer, Bush, nos. 
6401 and 6401 A. tTExas co.: wooded limestone slopes at base 
of bluffs along oe ae River, southeast of Prewitt Spring, 
Steyermark, no. 7 (Mo). oREGON co.: stony grove, Thayer, 
F. W. Pennell, no. ot ,521 (Phil). ozarx co.: cherty limestone 
slopes on top of bluff ‘along White River, northeast of Dormio, 
Steyermark, no. 10,417 (Mo). povuauas co.: limestone glade and 
cherty open woods, between Roosevelt and Richville, i i 
no. 19,165 (Mo). Miho co.: open hillside, west of 

O. E. Lansing, no. 3020. LACLEDE co.: cherty pokes of Pine 
Creek Hollow, ued of Nebo, Steyermark, no. 25,159 (Mo). 
WEBSTER a limestone outcrops, south of Fo rdland, Steyer- 
mark, no. 239 (Mo). TANEY CO.: common in woods, Swan, 
Bush, n " 336 (Mo, US); open rocky ground near Gretna, EF. SA 
Palmer, no. 19,224. sToNnE co.: dry rocky hillside oy. "James 
Riv r, E. fi Palmer, no. 5831 (Mo, US). BARRY Co.: dry woods 
Senna Eagle Rock, Sept. 24, 1896, K. K. Mackenzie (Mo, NY); 

Eagle Rock, Bush, nos. 78 and 1551 (Mo, NY, US). MCDONALD 
co.: dry ground, ’ Bush, no. 283. ARKANSAS: CRAIGHEAD CO.: 

open sandy soil, Jonesboro, F. W. Pennell, no. 11,510, as R. 
one Sale open woods, Bono, Demaree, no. 3519 sae 


(NY, Phil). LONOKE Co.: fallow fields, Carlisle, Demaree, no. 
17,516 (Mo, NY). prREw co.: woods, Monticello, Demaree, no. 
14,969 (Mo, NY). cee os Hino sit Conway, as R. 
ciliosa, Flora A. Hass, no. 1746 (US). AND co.: near Hot 
Springs, Runyon, nos. 1142 (NY) and 1430 ‘(US ). PULASKI CO.: 

low ridges, Fort Roots, Demaree, no. 17,301 (Mo, NY); Little 
Rock, Demaree, nos. 17, 325 (Mo, N a and 17,516 (Mo, NY). 
POPE CO.: Nogo, Geo. M. Merrill, no. 342 (Mo y. eae Co.: 

southeast of piven ane 3, 1912, Mabel P. Hollister (US). 


CARROLL Co.: dry ground, Eureka Springs, HZ. J. Palmer, 
nos. 4378 (Mo, US) and i 0,483 (NY). HEMPSTEAD CO.: near 
cNab, Greenman, no. 7 (Mo). FRANKLIN co.: rocky hill- 


side, Ozark, F. W. aire no. 10, Es (NY, Phil). HOWARD Co.: 
Baker Springs, Oct. 5, 1909, Ey aes ellogg. BENTON co.: 1889, 
N. ata k (NY). WASHINGTON co.: Savoy, May 18, 1922, 
ET. Wherry (US). smBasTIAN co.: Fort Smith, 1853-4, J. M. 
Bigelow, wal fete (US). LovistANA: WITHOUT CITED STATION: 
Hale, IsoTypk. NATCHITOCHES PARISH: dry wee pound. 
Natchitoches, E. J. Palmer, no. 7511 (Mo, NY, U 
LANDRY PARISH: dry woods, Opelousas, Carpenter & Hale (US). 
JEFFERSON DAVIS fies knolls in low prairies, Welsh, E. J. 


24 Rhodora [JANUARY 


Palmer, no. 7649 (US). OKLAHOMA: LE FLORE CO.: woods, near 
Page, G. W. Stevens, no. 1423. McCURTAIN CO.: woods near 
Idabel, H. W. Houghton as G. W. Stevens, nos. 3625 and 3638. 
TEXAS: BOWIE co.: near Texarkana, A. ie & E. G. Heller, no. 
4171 (Mo, NY, US). HARRISON co.: woods, Mar shall, Bush, 
no. 781 (Mo). cass co.: rocky svoddiaaid: Hughes Sheinae Bilt- 
more Herb., no. 10,6799 (US). CHEROKEE co.: dry sandy ground, 
Jacksonville, x. J. Palmer, no. 8600 (Mo, ; LS). ANDERSON 
co.: Palestine, April 19, 1895, E. N. Plank k (NY). UPSHUR CO.: 
sandy woods, Big Sandy, May 28, 1901, Reverchon (Mo); com- 
mon in sand, Big Sandy, Reverchon, no. 2535 o, NY). SAN 
AUGUSTINE CO.: eas woods, Geo. L. Crocket (US). HARRIS CO.: 
Houston, 1917, Ada Hayden. PLATE 843; MAP 2. 

In view of its very definite characters it is remarkable that the 
earlier collectors seem not to have secured Ruellia pedunculata 
and that it was not described until 1878. It is not improbable 
that Nees included it in his complex and chiefly tropical Cryphi- 
acanthus barbadensis. In his treatment in DC. Prodr. xi. 197 
(1847) Nees gave the broad range of the latter as tropical Amer- 
ica, thence to Virginia, Carolina and Texas (‘‘In Americae 
calidioris . . . inde a prov. Virginia, Carolina et Texas”’) 
but under the citation of specimens he gave nothing from the 
United States. Since his C. barbadensis had long peduncles with 
cymes, subovate leaves, and subulate-acuminate calyx-segments, 
the Texan element was presumably R. pedunculata. The rep- 
resentatives of the latter in Virginia and Carolina are the two 
following, only the first of which has ‘‘pedunculis subcymosis”’. 

R. pinetorum, sp. nov. (tab. 844), planta habitu R. pe- 
dunculatae; caule 1-3 dm. alto puberulo obtuse quadrangulato 
vel subtereto subsimplice vel divergenter ramoso vel ramosissimo 
internodiis elongatis; foliis oblongis vel elliptico-lanceolatis 
breviter petiolatis obtusis vel subacutis subcoriaceis minute 
lineolato-puberulis vel care pes tesa vel undulatis, majoribus 
2-3.8 cm. longis 0.8-1.8 cm. latis; pedunculis axillaribus 

3 cm. longis 1-3-floris, biravtedtis oblongis seers ogi 
brevioribus; calycis laciniis lineari-acicularibus 3-2 ¢ 
longis a basi 0.5-1 mm. latis attenuatis dses eystolithos 


em. longis; seminibus orbicularibus 3 mm. diametro.—Low pine 
barrens of the Coastal Plain, South Carolina (possibly Ae sat 
to northern Florida and Louisia ana, apparently local. 

CAROLINA: HORRY Co.: low pine barrens, July 28, 1936, Pr. 6. 


emt ee A 
2 enna 
“ need 

- eeeahtaateeanaeennt en ST 

| remnant, 

oo cceceneteeneneeeemeonnndllltememememeeneennentill 


Pa ae = 


2, 
of Ruetuia. Map 1, R. sTREPENS; 2, R. PEDUNCULATA; 


? 


R. , , 
, ? HIANA , 5, Re HETER ? , 


, . 
PANSA*‘ 13 R LINIENSIS, Var : 
H . es : 

, ILIS, ar. CALVESCENS; 14, fe CAR ’ 
? - UM V 0. TYPICA 


CHELO?! V ; . CAROLINIENSIS, V 
. 7 R. CAROLINIENSIS, ar. N LLA , 7 : : 
, V 


26 Bhodora [JANUARY 


parahed no. 800, TYPE in U.S. Nat. Herb. FLorIDA: CALHOUN 
low grounds, Iola, May, 1896, Chapman, three specimens, one 
shaman one marked oh. a. ”” with entry of an unpublished 
name which appears in American herbaria on sheets of at least 
two other and uke different species (therefore unwise to take 
up), the third marked “sp. nov. affin. R. pedunculata” (Mo). 
ALA ASHINGTON Co.: Fruitdale, July, 1904, as R. peduncu- 
lata, Southern Floral Seuest oe (Mo ). MISSISSIPPI: WAYNE CO.: 
Wayn renal he 8-9, 1896, C. L. Pollard, as R. eae no. 
1220 (Mo, US). ane CO.: Cuevas, Sept. 8, 1900, 
Lloyd & ite no. 346 (NY), LovurIsIANA: ST. TAMMANY PARISH: 
‘Covington, Sept., 1919, as R. parviflora, G. eee no. 11,687 
(US). ORLEANS ’parisH: New Orleans, 1832, T. ’Driimmond, 
nos. 257 in part (as R. strepens); 258 in part and 259 in part (as 
R. longiflora), the numbers inextricably confused, two of them 
appearing on one label. ‘cALCASIEU PARISH: vicinity of Lake 
Charles, May 28, 1904, and other dates (not given) in 1904, 
Andrew Allison, nos. 57, 261 and 297, all as R. pedunculata 
(all US). Map 3. 
Ruellia pinetorum is the southeastern Coastal Plain representa- 
tive of R. pedunculata and most of the few specimens seen were 
identified with that species which centers on the Ozark Upland. 
Chapman correctly understood it as a new species of this relation- 
ship but, as explained, the name he proposed but did not publish 
has been entered as a wholly new name on many sheets of at 
least two other species and should not be taken up. The finest 
material is that in the National Herbarium collected by Mr. 
Tarbox in low pine barrens of Horry County, South Carolina. 
I am, therefore, treating this as the type. In the chiefly Ozark- 
ian R. pedunculata the leaves are more ovate, the primary ones 
3-11 cm. long and 2-4.5 cm. wide; in R. pinetorum the leaves are 
oblong to elliptic-lanceolate and only 2-3.8 cm. long by 0.8-1.8 
em. wide. In R. pedunculata the broad bracts, especially in the 
simpler-stemmed plant with peduncles bearing solitary terminal 
flowers, nearly equal to greatly exceed the calyx; in R. pinetorum 
the narrow bracts are much shorter than the calyx. In R. 
pedunculata the calyx is copiously hirtellous with divergent 
sharp-tipped trichomes; in R. pinetorum glabrous or nearly so 
and closely invested with elongate and partially imbedded cysto- 
liths. In R. pedunculata the corolla-tube and the. ampliate 
throat are subequal in length; in R. pinetorum the tube is much 
longer than the less ampliate throat. In R. pedunculata the 
capsule is cinereous-puberulent; in R. pinetorum glabrous. 


1945] Fernald,—Ruellia in the Eastern United States 27 


The very few specimens assembled indicate that Ruellia 
pinetorum is a very local plant. Now that attention is called 
to it, it is hoped that fuller material will become available. As 
noted under R. pedunculata, it is probable that this is the plant 
intended by Nees when, in DC. Prodr. xi. 197 (1847), he noted 
his quite different tropical Cryphiacanthus barbadensis as ex- 
tending northward to Virginia and Carolina. No other plant 
known in the East satisfies his ‘‘pedunculis subcymosis petiolo 
longioribus vel et folium aequantibus superantibusve”. We do 
not now know R. pinetorum from Virginia but so many species 
are now known to “jump” from eastern South Carolina or south- 
eastern North Carolina to southeastern Virginia that R. pine- 
torum may well (before the destruction of most of the pine 
barrens) have been one of them. Really quite as closely related 
to the Ozarkian R. pedunculata, as is the Coastal Plain R. pine- 
torum, is the following beautiful species which centers on the 
Appalachian Upland. 


5. R. Purshiana, sp. nov. (ras. 845 et TaB. 846, FIG. 3), 
planta habitu plantae simplicissimae R. pedunculatae; caule 
simplice recto vel ramis erectis paucis gracile 1.5-6 alto 


cinereo-puberulo internodiis elongatis; foliis membranaceis 


.7-1.2 mm. latis apice attenuatis minute cinereo-pilosis vel 
-hirtellis deinde 1.6-2.8 cm. longis; corollis 3-5 cm. longis 
caeruleo-purpureis vel pallide purpureis vel albescentibus, tubo 
cylindrico 1.5-3 em. longo, fauce ampliato supra 0.8-1.4 cm. 
diametro, limbo (expanso) 3—4 em. lato; capsulis minute strigoso- 
hirtellis vel pilosis 1.5-1.8 em. longis; retinaculis 8.—R. ciliosa, 
var. hybrida Gray, Syn. Fl. N. Am. ii!. 326 (1878) in part only. 

. parviflora sensu Britton in Britton & Brown, Ill. Fl. ed. 2, m. 
241, fig. 3891 (1913) at least as to fig., not R. parviflora (Nees) 
Britt. (1901) at least as to basonym, Dipteracanthus ciliosus, 
var. parviflorus Nees (1842)—Dry to moist woods, bluffs, 
granitic or calcareous slopes, etc., western Maryland, south 
along the mountains and locally on the Piedmont to eastern 
Virginia, central South Carolina, Georgia and Alabama. Mary- 
LAND: FREDERICK co.: W. E. A. Aiken, as R. strepens, altered 


28 Rhodora [JANUARY 


to R. caroliniensis ee VIRGINIA: FREDERICK CO.: woods, 
Cedar Creek, June 2, 1929, Hunnewell, no. 11,135 (F WH): 
limestone cliffs, Cedar Creek, June 5, 1936, Hunnewell (VPI); 
both as R. caroliniensis, var. parviflora; Meadow Mills, June 9, 
1935, O. M. Freeman (USN A), as R. pie ROCKINGHAM CO.: 
Paul’s Fort, Frederick Pursh (Phil). RocKBRIDGE co.: Natural 
Bridge, May 28 and 29, 1909, HZ. B. Bartram, as R. pare viflora, 
one sheet (Gray) changed by later student to R. humilis ‘ ‘Pursh’’, 
another (Phil) to R. caroliniensis; ‘ he tl June 1, 1891, J. R. 
Churchill, as R. ciliosa, var. ambigua (Mo). BOTETOURT CO.: 
Indian Rock, June, 1887, H. E. Wetherill, as R. strepens (Pa). 
daagetpoag co.: shaly banks, vicinity of Mendot a, L. G. Carr, 
. caroliniensis. ROANOKE CO.: Decne May 29, 
1000; Brown, Hogg, Vail, Timmerman, Britton & Britton, as 
R. ciliosa, var. ambigua ( Y); wooded limestone Bs along 
Roanoke River at Dixie Caverns, July 6, 1942, C. E. Wood, Jr., 
no. 3673, as R. caroliniensis. BEDFORD CO.: July 8, i871 Bod. 
Curtiss, one of the several quite dissimilar dhaets marked by 
Gray as his R. ciliosa var. ambigua, one of the Curtiss specimens 
tagged by a later student as R. caroliniensis, the other as R. 
hybrida. AMELIA CO.: mnie 5, 1937, J. B. Lewis, no. 626, as R. 
jlpans ibe parviflora (VP I). HENRICO CO.: "Richmond, De 
as R. speceusouieia (US). NortH CAROLINA: ORANGE 
00. : pire poem Upper New Hope Creek, Duke Forest, May 27, 
1932, Blomquist, no. 4911, as fig caroliniensis (IT uke); New Hope 
Creek, Duke Forest , May 20 1933, Blomquist & Oosting, no. 
3364, as R. parviflora (Duke); dry bank near University Lake, 
on Neville’s Cree k, northwest of Chapel Hill, May 29, 1940, 
Radford & Stewart, no. 654a, as R. ciliosa (NC). GUILFORD CO.: 
acer High Point, May 22, 1902, Biltmore Herb., no. 147184, as 
R. parviflora (N Y, US). ForsyTH co.: Salem , Schweinitz, as R. 
strepens, altered by others, first to R. ciliosa, later to R. carolini- 
at woods, Winston-Salem, Aug . 20, 1921, P. D. Shal- 
lert, as R. adie. May 30, 1934, Schallert; no. 6! 509, as R. strepens 
(Schallert). RUTHERFORD CO.: Cuba, June 27, 1887, L. W. Lynch, 
no. 36, as R. strepens (NC). MADISON Co.: ’ Marshall, May 28, 


N. H. E. SONY). Lee. co.: Keowee, May 20, 1906, H. D. 


House, no. 2171, as R. parviflora (NY). GEORGIA: WITHOUT 
STATED LOCALITY: banal wa: of Georgia, R. 
ciliosa, var. ambigua (Mo). BURKE CO.: woods, Shell Bluff on 


Savannah River, ‘April 23, i936, Leeds & Harper, no. 2756, as R. 
parviflora (Phil). OGLETHORP co.: granite outcrop west of Lex- 


1945] Fernald,—Ruellia in the Eastern United States 29 


ington, May 28, 19384, Francis Harper, as R. parviflora (Phil). 
DEKALB CO.: Stone Mountain , May 23, 1897, Henry Eggert (Mo); 
mixed woods, Emory University campus, ‘April 30, 1936, Don 


Silver Creek, May 11, 1899, Biltmore Herb., no. 8494, as R. ciliosa 
(TYPE in Herb. U. 8. National Herb.). TENNESSEE: KNOX CO.: 
Knoxville, May ‘14, 1889, Lamson-Scribner, as R. strepens, 
changed by later students to R. ciliosa and to R. parviflora (US); 
woods and groves, Knoxville, July, 1897, Ruth, no. 9572, as R. 
strepens (NY); woodlands, Knox Co unty, May, 1898, Ruth, no. 
737, as R. strepens (NY ‘i ALABAMA: BLOUNT CO.: rocky wood- 
lands, Bangor, May 20, 1902, Biltmore Herb. no. 14,718, as 
ctliosa, var. parviflora (US). JEFFERSON CO.: Birmingham, a 
24, 1901, ee. Rosle, as R. parviflora (NY). Map 4. 

orma claustrofiora, f. nov. (TAB. 846, Fic. 1 et 2), floribus 
in glomerulis axillaribus aggregatis, glomerulis ad nodos omnes 
gestis; corollis tubulosis clausis 4-8 mm. longis apice dense pilosis; 
capsulis numerosis.— VIRGINIA: sin tai hee lag LOCALITY: 1843, 
Gray a een Ys as ee strepens. ROCK E CO.: “ex umbrosis 


Betiged to o R. parviflora; Natural Bridge, Sept. 4, 1885, 


COCKE CO.: within three miles of Wolf Creek Station, Aug. 31, 
1897, Kearney, no. 863, as R. ciliosa, var. hybrida (Mo and NC), 
TYPE in Herb. Missouri Botauicel Garden. 


Ruellia Purshiana, named for Frepertck Pursu, who first 
collected the species in the mountains of Virginia, is, when as- 
sembled from the very miscellaneous covers in which it has been 
confused, under 11 misidentifications, with no less than 7 species, 
stands out as a remarkably definite species of the Appalachian 
Upland. Although this is doubtless the plant chiefly intended 
by Gray when he conceived his R. ciliosa, var. ambigua, “‘as if & 
hybrid between R. ciliosa and R. strepens, with the aspect of the 
latter, but the calyx of the former”’, it can not be overlooked that 
Gray promptly last his bearings in applying the name R. ciliosa, 
var. ambigua, for sheets carrying the printed annotation-slip 
“Syn. Fl. N. Amer.” and marked by Gray as R. ciliosa, var. 
ambigua belong to no less than five species: the present one (in 
Herb. Gray); a Floridan sheet in Torrey’s Herbarium containing 
at least three species, none of them like anything else included by 
Gray in his var. ambigua; and a mixed sheet in Herb. Gray, con- 


30 Rhodora [JANUARY 


taining a sprig of R. pedunculata Torr. from Arkansas and the 
top of a plant of the very different Texan R. Drummondiana 
(Nees) Gray (this mixed sheet later misidentified as R. ‘‘caro- 
liniensis”’). I have not located the Kentucky plant included by 
Gray under R. ciliosa, var. ambigua. Even though we can infer 
that by his description and note Gray meant chiefly the plant I 
am here calling R. Purshiana, it is evident that, as he originally 
labelled specimens, the name R. ciliosa, var. ambigua was em- 
phatically a nomen ambiguum. The name has subsequently been 
further misapplied. I am, therefore, assigning to the species 
with somewhat the aspect of R. strepens but with much more 
slender calyx-segments a new name, typified by a characteristic 
sheet of specimens. ; 

That Ruellia Purshiana simulates very extreme plants of R. 
strepens with short leaves there can be no question. Some speci- 
mens of the latter, especially those from upland and dry habitats, 
consequently with greatly reduced stature and abbreviated 
leaves, are superficially similar (such specimens as the following: 
dry ground, Chickamauga Park, Georgia, May 25, 1911, Churchill 
(G); open woodlands, Coalgrove, Ohio, Biltmore H erb., no. 4500° 
(US); dry ground near Chattanooga, Tennessee, May 27, 1911, 
Churchill; and dry soil, Stevenson, Jackson County, Alabama, 
Biltmore Herb., no. 4500* (US)). There the resemblance stops, 
except for the possibly significant fact that both species have the 
showy flowers mostly solitary on few axillary peduncles and 
rarely producing fruit, and also have cleistogamous forms with 
glomerules of several highly fertile flowers in many of the upper 
axils. In R. strepens the stem is glabrous, pilose in lines or rarely 
over the whole surface; the lanceolate calyx-segments are: flat to 
the tip, 2-4 mm. broad, and conspicuously villous-ciliate; and 
the capsules glabrous. R. strepens is usually a plant of rich 
calcareous woods, oftenest on wooded bottoms, with its greatest 
concentration in the Mississippi Basin (Map 1). R. Purshiana 
(MAP 4) is a plant of dry rocky or upland habitats, sometimes on 
limestones but often (as on Stone Mountain) in granitic or some- 
what acid soils. Its stem is closely cinereous-puberulent; its 
calyx-segments linear and only 0.7-1.2 mm. wide below the 
middle, thence tapering to almost thread-like tips, and its sur- 
faces are densely cinereous-hirtellous. The capsules. too. are 


1945] Fernald,—Ruellia in the Eastern United States 31 


closely and minutely hirtellous. R. Purshiana was illustrated, 
erroneously as R. parviflora, in Britton & Brown, Ill. Fl. ed. 2, 
iii. fig. 3891. It certainly has nothing to do with the type of 
k. parviflora, which rests upon Dipteracanthus ciliosus, var. 
parviflorus (see p. 2). 

In its minutely puberulent and slender stem, its slender calyx- 
segments and pubescent capsule Ruellia Purshiana is similar to 
the simpler-stemmed and least floriferous states of the chiefly 
Ozarkian R. pedunculata (map 2); but in these least branching 
plants of R. pedunculata (PLATE 843) the peduncles are soon 
widely divergent; the linear-acicular calyx-segments taper from 
base to apex; the showy flowers are quite fertile and regularly 
followed by capsules. So far as we know R. pedunculata does 
not have a cleistogamous form. 

Although Ruellia Purshiana has often been identified as R. 
ciliosa Pursh, R. caroliniensis (Walt.) Steud., R. parviflora (Nees) 
Britton, R. hybrida Pursh, and even as R. humilis N uttall, such 
identifications merely reflect the general lack of clarity regarding 
specific characters and the current and still inevitable vagueness 
about the early-proposed species. The earliest of these names or 
their basonyms is Anonymos caroliniensis Walt. Fl. Carol. (our 
no. 11). Although no specimen now exists as type of Walter’s 
species, his remarkably detailed description, ‘‘caule tetragono 
hirsuto; foliis . . . hirsutis, . . . ; floribus sessilibus 
purpureis”’, surely does not apply to R. Purshiana. R. ciliosa 
Pursh (our no. 7) from near Savannah, was a branching plant, 
with subsessile ovate-oblong (really, apparently, obovate) leaves 
with margins and veins ciliate with long white hairs, the calyx- 
Segments four times shorter than the corolla-tube (calycis laciniis 
subulatis tubo corollae quadruplo brevioribus). Such a plant 
could not be R. Purshiana; in fact, when Pursh twice collected 
the latter in the mountains of Virginia, he did not venture to 
name it. R. hybrida Pursh (see discussion under no. 11) also 
from Savannah, is quite as remote: much branched (ramosissima) 
and hirsute with white hairs, the oblong leaves densely hirsute, 
etc. R. humilis Nutt. (our no. 10), an essentially sessile-leaved 
and usually freely divergent-branched plant, originally from 


Arkansas, has nothing to do with R. Purshiana; and as to R 


parviflora (see p. 2), that ill-defined name goes back to Dip- 


32 Rhodora [JANUARY 


teracanthus ciliosus, var. parviflorus Nees, from the Kentucky 
River, and with an impossible description for any North American 
member of Ruellia: with petioles 3-6 inches long—foliis paulo 
longiori petiolo (3—-6-pollicari)! Only by substituting for “peti- 
olo”’ the word “pedunculo”’ could one make much sense out of 
Nees’ diagnosis (if his plant was a Dipteracanthus and related to 
D. ciliosus) but even then his var. parviflorus remains wholly 
vague. It is not a good basis for the name of a common species 
with petioles rarely 1 cm. long, and surely it was not R. Pur- 
shiana. 

6. R: heteromorpha, sp. nov. (Tas. 847 et 848), planta di- 
morpha vel plus minusve heteromorpha. CAULIBUS VERNALIBUS 
1-3 basi plerumque decumbentibus jam adscendentibus simplici- 
bus vel divergenter ramosis 0.4-4 dm. altis puberulis plus minusve 
patenter villoso-hirsutis, nodis 3-5 (—10), internodiis 0.5-5 cm. 
ongis; foliis membranaceis obovatis vel ellipticis vel oblanceo- 
latis breviter petiolatis integris vel obscure undulatis plus 
minusve lineolato-strigillosis supra villoso-strigosis basin versus 
villoso-ciliatis subtus strigoso-hispidis glabratisve, laminis ma- 
turis 1.5-4.5 cm. longis 0.8-2.5 cm. latis; pedunculis perbrevibus 
axillaribus ad 1-3 nodos superiores bracteatis; bracteis oblongis; 
calycis segmentis lineari-attenuatis vix 1 mm. latis villoso- 


cleistogamicis 1-3 cm. longis, vel corollis expansis reductisque; 
seminibus subrotundis 2.5-3.5 mm. diametro. R. hybrida sensu 
Small, Fl. Se. U. 8. 1084 (1903), presumably not Pursh, Fl. Am. 


and Everglades, north to east-central FLORIDA: VOLUSIA CO.: 
moist pine barrens, near Seville, Aug. 1, 1900, A. H. Curtiss, no. 
6701A (US), July 30, 1909, Curtiss, no. 6701, as R. ciliosa, var., 
vernal fl. BREVARD Co.: scrub, south of Eau Gallie, Dec. 1, 
1919, Small, Britton & De Winkeler, no. 9200 (NY, US), later 
state, as R. parvi CIE 


sis, with an unpublished varietal name); Indian River, Edw. 
Palmer, no. 348 (Mo), as R. strepens. DE SOTO Co.: dry gravelly 


1945] Fernald,—Ruellia in the Eastern United States 33 


E CO.: Oa March 18, 1907, ellogg, vernal fl., as 
R. ciliosa, later annotated as sindaserihed var. of R. caroliniensis; 


Jeanette P. Standley, no. 70, vernal fi., as R. humilis later 
annotated like the last (G) and as R. parviflora (US) and as an 
ied species (Phil); in pineland, vicinity of Fort Myers, 
May 4, 1916, J. P. Standley, no. 425, vernal 8), as R. 
humilis, later annotated as k. parviflora; in pineland, Mullock 
Creek District, about 8 miles southeast of Fort Myer, iar 
June, 1917, J. P. Standley, no. 444, vernal f., 

annotated (G) as an unpublished var. fe R. snl ehienais: “Phil 
as an unpublished species, and (US) as R. parviflora; sandy pine 
woods along road to Coconut, April 4, 1930, Moldenke , no. 968, 
vernal fl. (Duke, Mo, NY). DaApDE co.: dry sandy soil among 
palmettos, Buena Vista, Jan. 17, 1930, Moldenke, no. 426 (Duke, 
Mo, N Y), as R. hybrida, vernal fl. and i: pinelands near Little 
River, Feb., 1917, W. E. Safford (US), as R. hybrida, later 
annotated as R. parviflora; Miami, April 4-7, 1898, Pollard & 
Collins, no. 233 (US), as R. humilis, later annotated as R. parvi- 
flora, April 1, 1903, vernal fl., V. L. Britton (US), as R. hybrida, 
later annotated as an unpublished var. of R. caroliniensis, 
March 14, 1917, H. B. Meredith (Phil), vernal fl. passing to 
later stage, as R. ciliosa, later annotated as an unpublished 
species; pinelands between Miami and Kendall Station, Nov. 5, 
1906, Sm all & nosivbi no. 2603 (NY), later stage, as R. hybrida, 
later. Soactated as s unpublished var. of R. caroliniensis; pinelands 
etween Cocoanut Grove and Cutler, Oct. 31-Nov 903, 
Small & Carter, a 1273 (NY as R. hybrida, Phil as R. parviflora, 
the former later annotated as an undescribed species), later 
Stage; pinelands about Sykes Hammock, Everglade Keys, 


a 


sis—rypE of R. heteromorpha in Herb. N. Y. Bot. Gard., vernal 
fl., passing to later stage; pinelands about Goodburn Hammock, 
Everglade Keys, June 31, 1915. Small & Mosier, no. 6381 (NY); 
pinelands between Cutler and Longview Camp, Nov. 9-12, 1903, 
Small & Carter, no. 1097, later state, labeled and annotated 
like the last: : pinelands between Long Prairie and Camp Longview, 
Oct. 31, 1906, Small & Carter, no. 2695 (NY), later state, labeled 
and annotated like the last; ‘between Peter’ s and Long Prairie, 
Nov., 1906, J. J. Carter, no. 173 (Phil), as R. parviflora, _— 
annotated as an undescribed species; pinelands about Humbugu 
Prairie, Feb. 28, 1915, Small & Mosier, no. 5555 (NY), fate! 
State, as R. hybrida, later annotated as a var. of R. caroliniensis; 
pinelands between Peter’s Prairie and Homestead, Nov. 10, 


34 Rhodora [JANUARY 


1906, Small & Carter, no. 2701 (NY), later state, named as the 

last; pinelands about Ross-Costello Hammock, Everglades Keys, 

June 24, 1915, Small, Mosier & Small, no. 6552, later state, as 

R. hybrida. MONROE Co.: woods, Pine Key, Blodgett, later state; 

pinelands, Big Pine Key, May 1, 1917, Small, no. 8142, vernal 
. a8 R. humilis. Map 5. 

Although most often, following Small, identified as Ruellia 
hybrida Pursh, it can hardly be overlooked that R. heteromorpha 
is a species of southern Florida, most abundant in the subtropical 
Everglades and Keys regions, that its branches, as soon as de- 
veloped, trail and greatly elongate, that the lower surfaces of its 
petioled leaves are glabrescent or nearly glabrous from the first 
(except for lineolate cystoliths and closely appressed strigae) 
while the upper surfaces are strigose, and that the tube of its 
fully expanded corolla is usually twice as long as the calyx- 
segments. fk. hybrida, from Savannah, Georgia, was described 
as erect and very much branched, hirsute with white hairs; the 
subsessile oblong leaves densely hirsute; the calyx-segments scarcely 
shorter than the corolla-tube. That is not a good diagnosis of the 
present species. Furthermore, Savannah is about 180 miles (a 
significant distance) north of the northernmost known station for 
R. heteromorpha; and it is surely significant that Nees in DC. 
Prodr. xi. 123 (1847), indicating by a mark of affirmation that 
he had seen the Pursh material, cited as belonging with it Drum- 
mond’s nos. 258 and 259, both of which belong in the essentially 
sessile- and oblong-leaved copiously white-villous-hirsute plant 
which Nuttall described from Arkansas as R. humilis, a species, 
moreover, which is unknown in eastern Georgia. Nevertheless, 
one can hardly pass by this identification by Nees, especially 
since the description of R. hybrida does not closely suggest R. 
heteromorpha. In view of his lack of material and his “lumping” 
of many different American plants as so-called varieties of his 
all-inclusive Dipteracanthus ciliosus, one can hardly accept. the 
identification of Nees as final. Similarly, R. ciliosa Pursh, 
originally described as with subsessile, ovate-oblong leaves with 
margins and veins long-ciliate with white hairs, and also from 
Savannah, was by Nees, who evidently saw the original material, 
matched with the broader-leaved and longer-flowered extreme of 
the species which Nuttall described as R. humilis. Several of 
the numbers cited by Nees are before me. They are uniform and 


| 
| 
| 
| 


1945] Fernald,—Ruellia in the Eastern United States 35 


have nothing to do except generically with R. heteromorpha nor 
with any plant known from the Savannah region; R. parviflora, 
as already sufficiently emphasized (see p. 2), was a misbegotten 
name for some plant (surely not the present) with “petioles 3-6 
inches long’’; while R. caroliniensis goes back to Walter’s re- 
markably clear description of a plant extending hundreds of 
miles north of R. heteromorpha, with distinctive characters which 
do not belong to the latter: ovate-lanceolate and hirsute leaves; 
throat of corolla campanulate, etc. When many of the speci- 
mens were annotated by an earlier student as an unpublished 
new species, they were given a name previously used by Nuttall 
for a quite different species (with rosettes of spatulate leaves, 
the plant I take to be Pursh’s R. ciliosa, the R. humilis sensu 
Small, not Nutt.), but identical material of the same numbers 
was likewise annotated with 2 or more additional names. Under 
these circumstances it seems wiser to avoid publishing names so 
vaguely understood by their author and to start anew. 

In its vernal showily flowering state, with simple or subsimple, 
erect stems (PLATE 847, FIGs. 1-3) Ruellia heteromorpha is so un- 
like the later state of the plant (PLATE 848, Figs. 1-3), with trail- 
ing and elongate heavily fruiting and freely branching stems, 
and with smaller or even closed and cleistogamous flowers, that 
the two might be mistaken for two species. Transitions between 
the two states are, however, very numerous. | 

7. R. crt1osa Pursh. Rosulate copiously villous plant, either 
with main axis abbreviated (0.5-5 cm. long) or elongate _ to 
dm. high, the internodes copiously white-villous, the pairs of 
leaves in the abbreviated plants crowded, in the more open and 
elongate individuals the 2-5 pairs remote, the stem often di- 
vergently branched, especially at base: lower subrosulate leaves 
oblong, oblong-obovate or oblong-oblanceolate to lance-spatu- 
late, prolonged to subsessile bases, obtuse to rounded at summit, 
copiously lineolate and more or less villous-hirsute, the larger 
ones 2-10 cm. long and 1-3 em. broad; the median leaves often 
much longer; the upper reduced and often crowded, both median 
and upper more definitely petioled, their margins often undulate: 
flowers mostly solitary in the axils, short-peduncled, the heavily 
villous peduncle terminated by 2 small oblong to lanceolate 
bracts: calyx-segments linear-acicular, copiously villous, 1.5-2.5 
em. long: corolla bluish or lavender to nearly white, 2.5-5.5 cm. 
long; the slender tube 1.3-3 em. long, the slenderly obconic 
throat 5-10 mm. in diameter at summit, the expanded limb 


36 Rhodora [JANUARY 


2.5-5 cm. broad: capsule 1.3—2 es long, glabrous; retinacula 6 
or 8: seeds suborbicular, about 3 mm. in diameter.— 
ae ii. 420 (1814), not most aps auth. Dipteracanthus ciliosus 
h) Nees in Linnaea, xvi. 204 (1842), not in DC. Prodr. 
122 (1847), which was a hopeless mixture. R. humilis sensu 
Small, Fl. Se. U. S. 1084 (1903), not Nutt.—Dry pine barrens 
and sands, South Carolina to central Florida and southeastern 
arg SoutH CAROLINA: lage STATED LOCALITY: 
ab. in Carolina”, H. M. Altorp (Mo). DARLINGTON CoO.: 
sandy soil in Sheep Pasture Savaninh x Hartsville, B. E. Smith, 
no. 1651 (NC), dwarf state, as R. humilis; sandy soil between 
Darlington and Hartsville, B. nt Smith, no. 916 (NC), tall state, 
as R. hybrida Pursh. SUMTER Co.: pine barrens near Cane 
Savannah, Witmer Stone, no. 496 (Phil), as R. humilis. Groreta: 
WITHOUT STATED LocaLity: Nuétall, as an unpublished new 
xe (Phil); Boykin (Phil). camprENn co.: lowland, Grace 
llert, no. 13,127 (Schallert). CHARLTON co.: “Extreme 
eciteht conditions” Gopher ware Mere ag near Sterling 
Branch, Jean Sherwood Harper, no. 419 (Phil), as R. humilis. 
WORTH CO.: vicinity of Poulan, Polterd i; Maxon, no. 574, as R. 
humilis, later annotated (G) correctly as R. ciliosa. MITCHELL 
co.: Camilla, S. M. Tracy, no. 3532 (Mo), as R. strepens. FLORI- 
DA: DUVAL Co.: dry pine barrens, A. H. Curtiss, no. 1944*, as an 
unpublished var. of R. strepens, in some herbaria altered to R. 
ciliosa, in others to R. humilis; Jacksonville, 1875, A. H. Curtiss 
(US), correctly as R. ciliosa, annotated as R. humilis; dry pine 
barrens, Fredholm, no. 5085, as R. ciliosa, var. longiflora Gray. 
ST. JOHNS CO.: near St. Augus tine, June, 1883, T. F. Seal (Phil). 
ORANGE Co.: sand, Winter Park, Apri il, 1900, A. M. Huger (NY), 
as R. humilis; sandy pine barren, Lake Brantley, C. 8S. William- 


A. J. Pieters, no. 120 (US). LAKE co.: Okahumpha, wong 
1888, Isaac Burk (Phil); vicinity of Eustis, Nash, 183. 
eageee co.: dry sandy woods, Brookeville, M plidsite: no. 
8 (NY). pasco co.: high pineland, Blanton, Barnhart, no. 
phe (US), as R. humilis. HILLSBORO CO.: Tampa, June—Aug., 
1898 M. Ferguson ne MARION Co.: turkey-oak woods, 
east of Citra, April 14, 1940, W. A. Murrill (Mo), as R. humilis. 
ALACHUA Co.: Gainesville, G. S. Miller, no. 437 (US) as R. 


W. A. Murrill (Mo), as R. humilis. suwANEE Co.: Live Oak, 
S. M. Tracy, no. 6684 Ree as R. humilis. GADSDEN CO.: in 
pinetis, prope Quincy, Mai—Jul. 1843, Rugel (NY), as R. arene 
grassy, white sandy dry field under. pine, River Junction, Wze- 
gand & Manning no. 2927, as R. humilis. Liperry co.: Bristol, 
Chapman (Mo, US). caLHoun co.: Wewahitchka, Chapman 
(Mo). HOLMEs co.: Ponce de Leon, June 20, 1905, J. M. Mac- 


BS Et, A —.neemnmnSmtE 


1945} Fernald,—Ruellia in the Eastern United States 37 


farlane (Pa). hea co.: dry open woods, Crestview, A. H. 
Curtiss, no. 6489, in part, as R. humilis. OKALOOSA Co.: near 
Camp Pinchot, Fon 21, 1928, O. M. Freeman (USNA), as R. 
humilis. ALABAMA: WITHOUT STATED LOCALITY: R. Haines, from 


5 (US), as R. humilis. WasHINGTON co.: Fruitdale, July, 
i001. 86 Floral fies Co. (Mo), as R. humilis. BALDWIN CO.: 
dry woods, Magnolia Springs, June 25, 1909, Schallert (Duke); 
dry open woods, Point Clear, 1896, Mohr (US): dry pine barrens 
olles 4 mile west of Elberta, R. M. Harper, no. 3795. MOBILE 

dry pine woods, Mobile, July, 1877, (US); pine 
tag Spring Hill, E. W. Graves, no. 498 (? or 948), in part 
only (Mo, US), as Re. humilis. MUIssissiprt: cme co.: Heidel- 
ae Tracy, no. 3324 (US), as R. humilis. WAYNE co.: Waynes- 
boro, Pollard, no. 1221 (Mo, NY, US), as R. humilis. HARRISON 


flora. LOUISIANA: ORLEANS PARISH: New O rleans, Drummond. 
POINTE COUPEE PARISH: dry sandy soil, Red River, Hale. PLATES 
849 and 850; map 6. 

Among the very diverse plants which have been identified with 
Ruellia ciliosa Pursh the present species seems to me most likely 
to be what he had from Savannah. His description, emphasizing 
the subsessile ovate-oblong leaves (remembering that Pursh used 
“ovate’’ for truly ovate and for obovate outlines) with long white 
ciliation, the bracts lanceolate, the subulate calyx-segments one 
fourth the length of the corolla-tube, was better for this plant 
than for most of the species (with ovate or lanceolate and petioled 
leaves) to which the name has been applied. Pursh’s account 
was as follows 


fates. 1 md ee ry 


ciliosa. 4. R. erecta, ramosa ; 


sa: 


albis longe ailtaae rabbis lanceolatis 
brevibus, calycis laciniis sehen tubo 
corollae quadruplo breviori 
Near Savannah, Georgia. 2. fy. v.s. in Herb. 
Enslen 


Since this is the éeiy plant which approximately satisfies the 
diagnosis and since it extends northward across South Carolina, 
I am retaining the name Ruellia ciliosa for it. When the actual 
type can be examined some revision may be necessitated. In 
DeCandolle, Prodr. xi. 122 (1847) Nees introduced a confusion 


38 Rhodora [JANUARY 


which has lasted for nearly a century, by citing under Dipteracan- 
thus ciliosus a number of Drummond and other specimens, from 
Texas to Missouri, which belong to the wholly different R. 
humilis Nutt. The latter, of which a type or isotype is before me, 
is a definite and wide-ranging inland species, quite unknown on 
the Atlantic slope of South Carolina and Georgia, with essentially 
sessile, oblong, ovate or lanceolate leaves with none of the su 
spatulate or obovate tendency of R. ciliosa and never, so far as I 
have seen, with the slightest rosulate tendency. The type of R. 
humilis was from Arkansas. Nevertheless, Small, clearly de- 
scribing as R. humilis only the dwarfer plants of R. ciliosa, gave 
it, as R. humilis, the range: ‘‘Georgia and Florida to Mississippi”, 
thus excluding the type-region. By Small’s account, his “R 
humilis” had the “Stems very short, 1-3 em. long’. The ma- 
jority of collections show some individuals with stems 0.3-3 dm. 
high, many of them from Florida; I cannot make out how Small 
disposed of them, for their leaf-outline and extreme villosity keep 
them out of his other categories. 


(To be continued) 


nea NR a RM 
ener eeeencenmcmanaael x — 


RUELLIA IN THE EASTERN UNITED STATES 
M. L. FERNALD 
(Continued from page 38) 

In northwestern Florida there occurs a localized variety which 
differs from wide-ranging Ruellia ciliosa in suppression of the 
pubescence, the internodes being merely cinereous-puberulent, 
and the calyx-segments, though long-ciliate, covered on the back 
with cinereous pulverulence. This should be called 


aoa c Rm tI orm RCA — mins RTT ET (mea i ane 


48 Rhodora [FEBRUARY 


Var. cinerascens, var. nov., caulis internodiis cinereo-puberulis; : 


calycis laciniis dorso cinereo-pulverulentibus margine villoso- 

ciliatis NORTHWESTERN FLORIDA: WALTON CO.: dry open 

woods, Crestview, July 22, sy A. H. Curtiss, no. 6489 in part, 

distrib, as R. humilis (Mo, N US (TYPE in U.S. Nat. Herb. )). 
SCAMBIA Co.: Fisherville, oa Pensacola, June 17, 1905, J. M 

M varies (Pa). 


R. succuLenta Small. Erect or nearly so, with simple 
sale and fleshy stems or erect branches 1.5-7.5 dm. high, the 
internodes sparingly puberulent or generally glabrous, purplish: 
leaves succulent, purplish, oblanceolate, narrowly obovate or 
oblong, narrowed to definite ticles glabrous or minutely 
hirtellous and glabrate, undulate, the surfaces more or less 
lineolate; larger blades 3-6 cm: long and 0.7—2 cm. wide: flowers 

1-3 on very short aban axillary peduncles: bracts small, 
ae ge or oblong, glabrous: calyx-segments linear-filiform, 

ong, glabrous or sparsely ciliate: corolla blue-purple, 
y 54.5 cm. long, or the latest ones smaller; the slender tube 
1.5-2.5 em. long, the slenderly campanulate throat 0.7-1 cm. in 
diameter at summit, the expanded limb 2. em. broad: capsule 
5 eves rarely strigose, 1-1.5 cm. long, longer or orb 
shorter than calyx.—Bu i. N. Y; Bot. Gant iii. 437 (1905).— 
ine ah region of southern FLoripA: pEsoTo co.: sandhills, 
~ Park to Sebring, Small & DeWinkeler, no. 9047 (NY). 

co.: hammocks, Myers, Hitchcock, no. 261. DADE Co.: 

bordee of wet prairie, Bay Biscayne, A. t Curtiss, no. 5500E; 
near Cutler, A. A. Eaton, no. 257; in everglades near unfinished 
railroad grades, between Coconut Grove and Cutler, Small & 
Carter, no. 1721 Lee el eer pal of edits Small & 


(NY); Rages about Sykes Hammock, Small, Mosier & Small, 
no. 5659 (Du NY); hammocks on prairie east of Florida 
City, Se: nee "8080 0 (NY); near the Homestead Trail, Small, 
no. 225 6 (NY ); west of Camp Jackson, Small & Wilson, no. 1844 
(NY); Cues Jackson to Camp Longview w, Small, Carter & Small, 
no. 3490 ibis basta gas and vicinity, Saffo rd & M osier, no. 
207 (US). OE everglades, intersecting Long Key, 
Small & Nea cn 3017 (NY). PLATE 851; MAP 7. 

In its stiffly upright habit, glabrescent and more or less fistulous 
stems, fleshy purplish leaves and relatively short calyx Ruellia 
succulenta is not likely to be confused with more northern species. 


rd 


1945] Fernald,—Ruellia in the Eastern United States 49 


Small contrasted it with R. “parviflora”, but that name, as cur- 
rently used, applies to as diverse an aggregation of plants as can 
be imagined, the name, as originally applied, belonging to nothing 
readily identifiable. In some characters, especially in habit and 
foliage R. succulenta might, superficially, be mistaken for the next. 


9. NOCTIFLORA (Nees) Gray. Stems erect or arched-ascend- 
ing (rarely divergently pole from base—presumably after 
injury), 1.5-6 dm. high, cinereous-puberulent or minutely 
cinereous-pilose, glabrescent, ‘with elongate internodes and 3-10 
pairs of leaves on main axis: leaves narrowly lanceolate to lance- 
or elliptic-oblong, submembranaceous, Ratna: or minutely 
hirtellous, ee to rae or subacute apex and to sessile to 


of leaves, very short, gnome erulent; the 2 cinereous 
bracts linear-lanceolate and undulate-dentate: calyx-segments 
slenderly linear, cinereous-puberulent or minutely cone 
.O-4.5 em. long: ¢ orolla bluish to nearly white, expandin 

the night, 6-11 cm. long; slender tube 4.5-8 em. long; ihe 
slender throat only 1-2 cm. long and 5-10 mm. thick; the ex- 
panded limb 3.5-6 cm. ae capsule cinereous-puberulent, 
2.25-3.5 cm. long.—Syn. Fl. N. oe ii!, 326 (1878). Cee ey 
LeConte in Ann. Lye. iv Y. 1. ae not HBK. (1817). 


wet pine barrens, evidently local, eastern Georgia to north- 
western Florida, ‘west to southwestern Louisi ~ a. GEORGIA: 
WITHOUT STATED LocaLiTy: LeConte (Mo, NY); savannas, 
LeConte? (Phil); these perhaps portions of original collec 
tions, at first called R. longiflora (not R. longiflora Richard, 
1792) but on en sheets altered i R. A LeConte in his 
seo a poptrs ‘“‘TInhabits in the savannahs of the Altamaha”’. 
RICHMON : “Altamaha, Bath”, LeConte (Phil). Fistor co.: 
near Sonbis Lalioite (NY); wet peaty pine barrens, 14 miles 
southeast of Ludowici, one Co. & Manning, no. 29 928 (errone- 
ously noted as from “Lon Co.”). MCINTOSH CO.: at sea-level, 
about Darien Junction, “Jane 35-27, 1895, J. K. Small (NY). 
FLORIDA: WITHOUT STATED LOCALITY: Florida, LeConte (Phil); 
Florida, Chapman, eal collections (Mo, NY, Phil, US); 
est Florida, Chapman, rota begreer age WAKULLA CO. 


“Dizygandra noctiflora Shuttl. , In campis graminosis, 
1 Sunbury, not eg on modern maps of cur, was thus described in Lippincott’s 
Gazeteer (1856): ecayed town of Liberty co., Georgia, on the Medway River, 


about 30 miles S. Mycteare of Savannah 


50 Rhodora [FEBRUARY 


inter St. Marks et Port Leon, Florida, legit Rugel, Jun. 1843 
(Mo, NY), 1soTyPes of Dipteracanthus noctiflorus Nees. FRANK- 
LIN CO.: Apalachicola, Chapman, with note, ‘‘The flowers open 
in the night and fall off by morning”; low prairie, Apalachicola, 
1867, B. F. Saurman; ae gra y Pie barren, Apalachicola, 
Chapman, distr. by Biltmore Hans . 4501*; low pinelands, 
Port St. Joe to Aneinsticoln. Small, Small & DeWinkeler, no. 
11,417 (NY, US). Atasama: low meadows on the coast, Porters- 
ville, Sept., 1892, Mohr (US)—locality not definitely located, 
since Portersville in DeKalb County is not on the coast. CULL- 


ss m ek 

1892, Mohr (NY, US). Mussissrprt: Deer Island, A. B. Seymour, 
no. 197 (loosely branched, presumably injured, the oe 
leaves with unusually long petioles). HARRISON CO.: Bilo 
June 22, 1899, S. M. Tracy, some specimens as no. wpe others as 
no. Mo, NC, NY, US). HaANcocK co ne_ barren, 
Prete & Schery, no. 51 (Mo); Bay St. Louis, foal Is (NY). 
LOUISIANA: CALCASIEU PARISH: vicinity of Lake Charles, 1904, 
y oa Allison (US). Puiarns 852 and 853; map 8. 

Ruellia noctiflora is one of the most distinctive and, at the same 
time, highly localized American species. In view of the great 
amount of collecting in the Southeastern States, it should, if at 
all common, be better represented in herbaria. LeConte’s state- 
ment, when he originally published it as R. tubiflora, that it 
“Tnhabits in the savannahs of the Altamaha’’, suggest localiza- 
tion. When he renamed the species Dipteracanthus noctiflorus, 
Nees, as usual, had very mixed ideas, for he included with the 
LeConte plant and that from St. Marks, Florida, material from 
Texas of the wholly different R. humilis Nutt.; in fact he made 
R. humilis a variety of D. noctiflorus, and he went so far in his 
general misinterpretations as to note that the wonderfully differ- 
ent Dipteracanthus noctiflorus “vix a D. cilioso Purshii differret”’. 
Aside from the minutely canescent pubescence which covers most 
parts of R. noctiflora, this species has the longest corolla and calyx 
of any of our species, and its very long (up to 3.5 cm. long) 
ee is cinereous-puberulent. 


R. numitis Nutt. Stem often at first simple, soon with 
ed ing to horizontally divergent or reclining branches; 
the main axis 1-7 (-8.5) dm. high; the rather short internodes 
berate! or can pant ioe to pkneinemen quires 

with 4-12 pairs of leaves longer than the internodes, and 4-10 
mse floriferous: leaves coriaceous, often hirsute to ’ villous on 


1945] Fernald,—Ruellia in the Eastern United States 51 


nerves and margin, oblong or oblong-lanceolate to ovate, _— 
or subsessile, nearly uniform or but slightly reduced upward: 
flowers few in the axils: bracts lanceolate, oblong or elliptic: 
calyx-segments 1.5-2.5 cm. long, linear-attenuate, mostly 
hirsute- to villous-ciliate: corolla 2-8 ¢ . long, lavender to 
bluish (rarely white), the slender tube 0. 75 cm. long, the throat 
campanulate; corolla reduced, closed and tubular in rare cleisto- 
gamous individuals: an a aarreabias 1.2-1.5 cm. long: ~ 
suborbicular, 3-4 m n diameter.—A wide-ranging polymor- 
phous species, of Teh: the following are the more sanidenat 
varieties. 


ong. 
oe leaves of gore! axis elliptic-oblong to ae 
lanceolate, 1-2. road, obtuse to subacute. 


aiverwent heiress oe pes Var. typica. 


rounded at summit, 2.5-4 cm. bro a eee oes 100. Var. expansa. 
a. Younger internodes of stem nae glabrescent or with 
only few scattered hairs on the angles; leaves glabrous, 
piabiessaiit or only soni ely short-hirsute on 
neath and margin, narrowly e ce Boo ashy to oblong- 
lanceolate, the otf ones 1-3 cm. broad and 2-6 em. long; 
corolla 2-3.5 em. long, its tube 0.7-2.3 cm. long; plant 
chiefly of Cinnbertand TIMOR 2 oe ee 10f. Var. calvescens. 


10a. Var. typica. R. hirsuta Ell. Sk. ii. 109 (1822), not Vell. 
(1790). R. humilis Nutt. in Trans. Am. Phil. Soc. v. 182 (1837). 
Dipteracanthus noctiflorus, @. Mania Ne ees in DC. Prodr. xi. 123 
< mae - least as to basonym. R. ciliosa, var. ice Gray, 
. Am. ii!. 326 (1878) in part only. liosa, var. 
“thle (N ut.) Britton in Trans. N. Y. Acad. Ae ix. 185 (1890), 
at least as to basonym.—Dry prairies, rocky slopes, open woods 
etc., southern and eastern Iowa to Texas, east to the Mississippi 


land, western Virginia, ssee a 0 stern Alabama. 
The following are characteristic (all, unless eeu called 
R. ciliosa Purs NIA: FRANKLIN ‘abundant 


h). 
and common”, iaatentes July 24, 1896, Thos. C. Porter 


52 Rhodora [FEBRUARY 


(NY, Phil), variously as R. ciliosa, R. ciliosa, var. parviflora and 

R. strepens, var.; dry ground, among grasses, near Baker Cavern, 

south of Williamson, Hans Wilkens, no. ey Be Mercersberg, 
co 


1846, Ruel (Phil), as R. strepens. MARYLAND: WASHINGTON 
Potomac River, Williamsport, Aug. 1849, Porter (Phil), as 
hg eaagy ‘stre repens. Wrst VIRGINIA: HARDY co.: Lost 


River, Aug. 18, 1931, Core (NY). Virernta: GILEs co.: Ripple- 


August 4E 1937, ‘Lena Artz (Mt ie OHIO: CUYAHOGA CO.: near 
Blue Rock S$ prings, Cleveland, oe 000 no. 1542. INDIANA: 


ry, 8 

n, R. M. Tryon, Je no. 2698 Deke. as R. caroliniensis. 
ots co.: rocky woods between Duncan and New Albany, 
Deam, no. 13,995 (NY). CRAWFoRD co.: rocky, exposed wooded 
hillside southwest of Milltown, Deam, no. 16,422. KrntTucKY: 
LOGAN Co.: limestone ledges and barrens, near Russelville, EZ. J. 
Patines no. 17,758 (Mo), . ie parviflora (transition to var. 


calvescens). TENNESSEE: DAV : copses, vicinity of Nash- 
ville, Gattinger; West Nashville, Mosleson, no. 5160. FRANKLIN 
co.: Cumberland Mts ; Cowan, Ruth, no. 564 (US). ALABAMA 


FRANKLIN CO.: small flat cedar glade just north of Isbell, R. M. 
Harper, no. 3887 (US, VPI), as R. caroliniensis (transition to 
var. calvescens). MISSISSIPPI: LOWNDES Co.: sandy, open places, 
banks of Tombigbee River, Columbus, June 15, 1892, C. Mohr 
(US), as R. ciliosa, var. hy brida. WARREN CO.: near Vicksburg, 
Paul J. Schallert, Jr. no. 13197 (Schallert). ILLINOIS: COOK CO.: 

limestone cliffs, Le emont, July 20, 1912, H. L. Braun (Braun). 


). 

co.: river-bank, vicinity of Kankakee, C. C. Crampton, no. 568 
(US). RICHLAND co.: southwest of Calsmin. Robt. Ridgway, no. 
3183, in part (Phil). KENDALL co.: Yorkville, oy doar 1884, 
T. E. Boyc LEE co.: Dixon, July, 1861, Wm. B oot (Phil), as 
Dipteracanthus ciliosus, correctly annotated as R. haonilis. WASH- 
tincTon co.: Ashley, F. Beckwith, no. 47 (Mo). JACKSON CO.: 

gumbo soil, woods, Bie Muddy River, John McCree, Jr., no. 833 
(Mo). UNION CO.: ona V tp of Lick Creek, Oct. 7, 1939, E. 
Anderson & Wm. Bauer (Mo). Iowa: cosa 3 CO,! Lyons am 
mel, no. Je te saab co.: Mt. Pleasant, C. R. Ball, no. 1564 


ohn Davis, no. 978 (Mo). RALLS CO.: ie hillsides, west of Hanni- 
bal, John Davis, no. 4646 (Mo). prKE co.: Eolia, Aug. 26, 1916, 
John Davis (Mo). ST. LOUIS co.: Meramec Highlands, July 29, 


1945] Fernald,—Ruellia in the Eastern United States 53 


1905, A. G. Johnson (Mo); West Webster, Uphof, no. 3803 (Mo). 
JEFFERSON co.: Cedar Hill, John H. Ke llogg, no. 1185 (Mo). 
FRANKLIN Co.: Pacific, Sept. 18, 1910, Moses Cratg (Mo). BUTLER 
co.: woods, Bush, no. 3739 (Mo, NY, US), as R. parviflora. SHAN- 
NON CO.: Bush, no. 70, identification correctly altered to R. humilis. 
COOPER CO.: rocky woods, Bush, no. 15,137 (Mo). MORGAN co.: 
rocky woods, Bush, no. 15, 147 Mo). WEBSTER Co.: upland lime- 
nae glade, north of Forkner’s Hill, Steyermark, no. 23,854 (Mo), 
s R. caroliniensis. OZARK Co.: rocky 4 ground (dolomite), 
Bacies of “Bald Jesse”, near Gainesville, Z. J. Palmer, no. 33,073 
(Mo). pores co.: Cole Cam mp, Trelease, no. 439 (Mo). BARRY 
CO.: s, Bush, no. 15,007 se as R. parviflora. CLAY CO.: 
Phxadier z fee 29, 189 qd: A. C'. Magruder (Mo), as R. ciliosa, 
var. humilis. JACKSON CO.: rocky barrens, Greenwood, Bush, no. 
9745 (Mo, a as R. parviflora; Rocky Bluff, south of pute: 
pendence, K. K. Mackenzie, no. 1065 (NY), as R. ciliosa, var. 
humilis. iyo co.: chert barrens, Joplin, E. J. Palmer, no. 
1309 (Mo) and no. 18,451 (NY), the latter as R. sin a 
NEWTON Co.: chert barrens, Reding’s Mill, #. J. Palmer, no 
1532 (Mo). McDONALD co.: July 24, 1893, Bush (Mo). ARKAN- 
SAS: WITHOUT STATED LOCALITY: “on rocks in the upland forests 
and prairies’, Nuttall, Type or isotype (NY), the specimen 
marked by Britton R. ciliosa Pursh. cuay co.: woods, Moark, 
Bush, no. 3746 (Mo), as R. pt ees a. LONOKE Co.: fallow fields, 

Carlisle, Demaree, no. 17,613 (Mo), as R. caroliniensis. PULASK 
CO.: rocky ridges, ae Mt., Demaree, no. 19,797 (Mo), ie 
R. caroliniensis. CARROLL co.: rocky slopes, Eureka Springs, 
E. J. Palmer, no. 4426 (Mo). BENTON co.: Decatur, 1899, E. N. 
Plank (NY). JOHNSON co.: ridges, Knoxville, Demaree, no. 
19,933 (Mo), as R. caroliniensis. FRANKLIN Co.: low ridges, 
ranch, Demaree, no. 17,786 (NY), as R. caroliniensis. YELL 
co.: dry rocky ridges, Tones Mt., Dardanelle, Demaree, no. 
20,006 (Mo, NY). scorr co.: dry low hills, Mansfield, Demaree, 
no. 18,171 (NY). LOUISIANA: CALCASIEU PARISH? Lake Charles, 

. Tracy, no. 3468 (NY). Kansas: Fy ican co.: 6 

west of Pittsburg, Rydberg & Imler, no. 164 (NY). LABETTE 60. 
bluffs north of Oswego, Rydberg & I wiler, no. 359 (NY). MONT- 
GOMERY Co.: northeast of Caney, Rydberg & Imler, no. 401 (NY). 
OKLAHOMA: OTTAWA Co.: woods, Miami, G. W. Stevens, no. 2304. 
MCCURTAIN CO.: woods, near Idabel, H. W. Houghton as G. W. 
Stevens, no. 3633. semen co.: dry knoll, near haa a G. W. 
Stevens, no. 1931. YNE CO.: ny) Rane Waugh, no. 144 (Mo); 
sandy clay soil, oth of Sele Robert Stratton, no. 141 
(Mo). Exuis co.:sand, Arnett, L. F. Locke, no. 25 (US). Texas: 

WOOD co.: sands, Mineola, yest (Mo). JEFFERSON CO 
west of Beaumont, April 11, 1921, E. T. Wherry (US), correctly 
identified. HARRIS CO.: prairies, ’ Laporte, Reverchon, no. 3938 


54 Rhodora [FEBRUARY 


(Mo), as R. parviflora. Brazos co.: College Station, R. G. 
Reeves, nos. 167 and 168 (US). pauuas co.: dry places, Dallas, 
July, 1877, Reverchon (NY). BELL co.: dry woods, near Temple, 
S. E. Wolff, no. 2288 (US). TARRANT Co.: sandy post-oak woods 
between Grapevine and Ft. Worth, heey & Lundell, no. 9516 
(US). FAYETTE co.: Crawford, 1892 (Mo). Lavaca co.: Hal- 
a G. L. Fisher, no. 100 (US). Bexar co.: bank of 

elot ae northwest of San Antonio, Sister Mary Clare 
Mite. no 9 (NY). KERR Co.: hillside woods, Lacey’s Ranch, 
E. J. Pale, no. 9994 (Mo). Puarss 854 and 855; MAP 9. 

The minor form ay white corollas is 


Forma alba (Steyermark), comb. nov. £&. caroliniensis, forma 
alba Steyermark in RuHopora, xli. 585 (1939). —Typr from 
Missouri: prairie-slopes above limestone bluffs along Long 
Creek, 1144 miles south of Shae Caldwell County, Steyer- 
mark, no. 3058 (IsoTyPpE Mo 

Although the description of forma alba, as a form of Ruellia 
caroliniensis (a strikingly different species not found in Missouri), 
gave no indication that it is an albino of the sessile-leaved 
species of the prairie-region and not at all of R. caroliniensis, the 
isotype is quite like typical R. humilis except for its white corolla. 


Forma grisea, f. nov. (raB. 856) caulis internodiis griseo- 
puberulis vel cinereo-pilosis, Me hirsutis.—Scattered through the 
range of var. typica. OHIO: PICKAWAY co.: Aug. 9, 1912, R. 
Lebo (Phil). finmioia® aSEOGN co.: gravelly hillside, 

ower, Aug. 28, 1900, Gleason, no. 1803. MussourRt: 
eeiea co.: Jerome, Oct. 5, 1913, John H. Kellogg, no. 157 (Mo). 
JACKSON Co.: dry open bank, Kansas City, Sept. 15, 1916, Ralph 
Hoffmann (Mo). McDONAL .: dry ground, uncommon, July 
24, 1893, Bush, no. 281 Geren | bs | Herb. Gray.; ISOTYPE in Herb. 
Britt tt.). LOUISIANA: JEFFERSON DAVIS PARISH: knolls in low 
prairie, May 17, 1915, E. J. Palmer, no. 7649 (Mo). CALCASIEU 
PARISH: Lake Charles, Aug. 7, 1897, S. M. Tracy, no. 3469 (US). 
OKLAHOMA: OTTAWA CO.: pasture near eee Aug. 31, 1937, 
Rk. Fosberg, no. 14,289 (Penn). EXAS: VAN ZANDT CO.: 
Grand Saline, Oct. 18, ——, sais Mo). 

Var. frondosa, var. nov. (ra AB. 857), vencbopiac valde villoso- 

irsutis; axis primarii foliis majoribus ovatis vel ovali-oblongis 
vel late ellipticis, 3.5-8 cm. longis 2-4 em.. hee a 
obtusis; corollis 3-4.5 (-5) em. longis tubo 1.2-3 em. longo.— 
Southeastern Nebraska to eastern Texas aad Louisiana, “cant 
ward to south-central Pennsylvania and western Virginia; speci- 
mens, unless otherwise noted, originally called R. ciliosa, in a 
few cases corrected to R. humilis. PENNSYLVANIA: FRANKLIN 
co.: Chambersburg, Aug. 27, 1897, Thos. C. Porter. V1rGINIA: 


i ome et ree e 
sated - at: CT: eeemenaant 


1945] Fernald,—Ruellia in the Eastern United States 55 


SHENANDOAH Co.: rocky (limey) woods, near Strasburg, Hunne- 
well, no. 12,037 (FWH), as R. caroliniensis; limestone barrens 
near Strasburg, June 6, 1936, Hunnewell (VPI), as R. carolinien- 
sis; damp thicket near Meadow Mills, Hunnewell, no. 17,837 
FW 0, F 


WYTHE Co.: banks, Reed Creek, June 28, em as 
es: es site tie co.: Euclid Heights, Cleveland, Green- 
man, 353; r Blue Rock PpRngs, mea oe Greenman, 
nos. 1541 and 1542 (U S). GREEN CO.: a, July 10, 1883, H. 


A. Young; open bottoms, Yellow aes July 10, 1935, Demaree, 
no. 11,486 (Mo, Phil); ’ Cedarville, July, 1906 "J. F. Cleven nger 
(US). MonTGomERY co.: Dayton, John W. VanCleve (Phil), as 
R. angele CLERMONT Co.: sandy soil, gravel terrace, Mul- 
ford, June 17, 1916, H. L. Braun (Braun). HAMILTON Co.: gravel 
banks, ligeeat Fernald, July 19, 1914, #. L. Braun (Braun). Inp1- 
N CO.: prairie north of "Tab, August 31, 1916, Deam, 
a “art 613 (NY): high bank S Pine Creek, west of Kramer, July 
8, 1918, Deam, no. 25,843. MARION co.: abundant among tall 
weeds between Indianapolis se Carmel, Aug. 8, 1942, 
Friesner, no. 17,202 (type in Herb. Gray. ; ISOTYPE in Herb. 
Butler Univ.), distrib. as R. caroliniensis; same station, August 5, 
1944, Friesner, no. 18,579 (topotypes).. HARRISON co.: bluff, 
south side of Buck Creek, southeast of Corydon, June 15, 1940, 
Friesner, no. 14,369 (D NY), as R. caroliniensis. KENTUCKY: 
PENDLETON C0.: dry fea bank, south of Peach Grove, July 8, 
1941, EZ. L. Braun, no. 4037. ‘TENNESSEE: DAVIDSON Co.: West 
Nashville, May 26-27, 1909, Eggleston, no. 4436 (Phil). Ittrots. 
COOK co.: Chicago, Se H. Babcock (US); Flossmoor, Aug. 1, 1909, 
Greenman, no. 2816. DUPAGE co.: introduced along roadside, 
Naperville, Aug. 1, 1897, L. M. Umbach (US). WILL co.: ro ad- 
side, Romeo, June 25, 1896 (Phil), July 26, 1897 “(US), Ss) 
WINNEBAGO CO.: Rockford, July 11, 1926, Pammel & Fisk, 
293 (Mo). RICHLAND co.: Larchmount, Aug. 1, 1914, Robt. 
idgway. CHAMPAIGN Co.: vicinity of Urbana, Aug. 10, 1910, 
Steele (US). sTarxk co.: gravel slope, near Wady Petra, July 9, 
1900, V. H. Chase, no. 673 (Phil). HANCOCK CO.: Augusta, 
1845, S. B. Mead, as R. strepens. sv. CLAIR co.: dry ground, 
June 28, 1875, Henry Eggert (Mo). Iowa: cLinTon co.: Clinton 


(Lyons), Sept. 4, , Pammel, no. 77. WARREN CO.: Indianola, 
July 6, eer Paina Welbus & J acques. MISSOURI: CLARK C 
pene u ush (Mo). st. LOUIS co. collines arides, 


6205 (Mo). NODAWAY co.: geet ngs, n 
Parnell, June 20, 1938, eae no. no. 5936 (Mo). TACKSON ee 


56 Rhodora [FEBRUARY 


dry grounds, Waldo Park, June 10, 1896, K. K. Mackenzie (US). 
JOHNSON Co.: sandstone hillside, Warrenburg, Aug. 27, 1916, G. 
W. Stevens, no. 4106 (NY). GREENE co.,: northeast of Spring- 
field, Aug. 29, 1911, P. C. Standley, no. 8390 (US). JASPER co.: 
chert barrens, northwest of Joplin, Aug. 16, 1908, E. J. Palmer, 
no. 1309 (Mo). mMcpona.p co.: dry open ground, Noel, Sept. 7, 
1913, E. J. Palmer, no. 4164 (Mo); chert slopes west of Noel, 
May 31, 1938, Steyermark, no. 562 (Mo), as R. caroliniensis. 
ARKANSAS: FAULKNER Co.: rocky glade, Guy, Sept. 4, 1934, 
Demaree, no. 10,963 (US). puLAsKi co.: Little Rock, July, 1886, 
H. E. Hasse (NY). carrout co.: Eureka Springs, July 17, 1898, 
N. M. Glatfelter (Mo). LovtstaNa: RAPIDES PARISH: pine woods, 
vicinity of Alexandria, June 6, 1899, C. R. Ball, no. 655, in part 

0, US). NeBRASKA: LANCASTER CO.: roadsides, Lincoln, Aug. 
=e H. J. Webber (US). Kansas: pouaLas co.: Lawrence, 

en 


through wooded area, southeast corner of county, July 23, 1937, 
. B. Jacobs, no. 134 (NY). cowLEy co.: 1895, C. M. Gould 
NY). OKLAHOMA: ELLIS Co.: east of Harmon, June 17, 1932, 


em. broad and 3-6 em. long, obtuse to subacute: corolla 5-8 em. 
ong; = tube 3-5 cm. long.—R. ci tosa, var. longiflora Gray, 
yn. 


as R. ciliosa, more rarely as R. humilis, ILLINOIS: RICHLAND CO.: 
Parkersburg, June 14, 1902, Robt. Ridgway (US). MACOUPIN co.: 
Piasa, Aug. 5, 1905, G. E. McClure (Mo). WILLIAMson co.: dry 
soil, Bush, John McCree, Jr. no. 881 (Mo). Missourt: MARION 
co.: dry soil, near Mark Twain Cave, John Davis, no. 4101 (Mo). 
LINCOLN Co0.: Silex, John Davis, no. 3887 (Mo). st. Louis co.: 
Windsor Springs, July 1, 1890, A. S. Hitchcock (Mo). JEFFERSON 
co.: stony hills north of Hematite, July 7, 1891, Henry Eggert, 
(Mo); dry limestone glade southwest of Crystal City, Steyer- 
mark, no. 1114 (Mo), as R. caroliniensis; Festus, July 11, 1925, 
Woodson (Mo); south of Festus, Mildred M athias, no. 700 (Mo). 
WASHINGTON CO.: Potosi, July 24, 1885, Frederick Wislizenus 

0). COOPER co.: rocky barrens, Bush, no. 13,673 (Mo). 


1945] Fernald,—Ruellia in the Eastern United States 57 


GREENE Co.: dry hills near Fulbright Spring, P. C. Standley, no. 
9567 (US). TaNrEyY co.: Forsythe, Trelease, no. 440 (Mo), as 
R. ciliosa, var. longiflora. JACKSON Co.: rocky hillside, June 15, 
1864, G. C. Broadhead (Mo). BENTON co.: Mora, Trelease, no. 
438 (Mo). ST. CLAIR CO.: rocky woods, Osceola, Bush, no. 12, 819 
(Mo). JASPER co.: Aug. 16, 1893, Bush (Mo), as R. ciliosa, var. 
longiflora. Amesarais: CRAIGHEAD CO.: open woods, Lake City, 
Demaree, no. 3391 (Mo). SHARP Co.: Hardy, Greenman, no. 
1524 Mo). pete co.: prairie-thickets, Mist, Demaree, no. 
15,096 (Mo). FAULKNER co.: rocky hillsides, Guy, Demaree, no. 
10,963 (NY). conway co.: Petit Jean Mt., Morrilton, June 25, 
1937, John K. Edwards (Pa). LOGAN CO.: rocky, dry situations 
at 2800 ft., Magazine Mt., Demaree, no. 17,720 (Mo, “yet as 
R. caroliniensis SEBASTIAN co.: Fort Smith, 1853-4 


eo Herb. (NY). NATCHITOCHES PARISH: dry open ati) 
Soecte mag mee E. J. Palmer, no. Pee Y); dry sandy ground, 
Chopin, EF. J. Palmer, no. 7565 (Mo); long-leaf pine pendhills, 
fake 20, 1930, Caroline Dorman (NY). RAPIDES PARISH: Alex 
andria, Josiah Hale (NY). JEFFERSON DAVIS PARISH: knolls on 
low prairies, Welsh, E. J. Palmer, no. 7649 (NY). marisa 4 
PARISH: low grassy soil, 1 mile east of Lake Charles, D. S. & H 
Correll, no. 9652. KANSAS: WITHOUT DEFINITE LOCALITY i 
tween Neusha and Red Fork, Sept., 1849, Marcy Exped. RILEY CO: 
Manhattan, Pek 29, 1884, M. A. Carleton (Mo). LYON co.: ae 
poria, July 13, 891, E. C. Sm ith (Mo). MONTGOMERY CO.: 
prairie, one September 16, 1900, Frank W. Johnson (NY). 
SEDGWICK co.: Wichita, 1892, H. R. Rose. OKLAHOMA: CRAIG CO.: 
north of Vinita, G. J. Goodman, no. 3047. osaGeE co.: dry knoll, 
Pawhuska, G. W. Seta no. 1931. CREEK Co.: fag ‘bi it oe 
428 (Mo). TExas: WITHOUT STATED LocALITy: Drum , No 
219 and 220 (amas ean) and 258 (nos. 220 and 258 praing Hae of 
 Rboeceacaae! reas Torr. & Gray); Lindheimer, no. 158; 
everchon, no. 725 (Mo). TYLER CO.: Jct south of Woodville, 
Lundell & leandats no. 11,544 (US). HARDIN co.: east of Camp 
Jackson, Cory, no. 19, 793. HOUSTON sos cm open ground, La- 
texo, EZ. J.P. Palmer, no. 12, sig eT NY). WALKER co. seme agy of 


co.: Dallas, June, 1874, Reverchon, no. 410, PARATYPE; vicinity 


58 Rhodora [FEBRUARY 


rocky ground, Aug. 10, 1924 (NY)). TRavis co.: Austin, Tharp, 
no. 1384 (US). Bastrop co.: McDade, Va. Collins, no. 266. pxE 
witT co.: hillside, July 30, 1941, Marguerite Riedel. BEXAR Co.: 
hills north of San Antonio, Sept. 1, 1900, Henry Eggert (Mo). 
TOM GREEN Co.: Knickerbocker Ranch, Dove Creek, May, 1880, 
Frank Tweedy (US), as R. ciliosa, var. longiflora. PLATE 858; 
MAP 


Lakes, Chapman (Mo). Mrcutcan: old specimen marked b 
Asa Gray “ Michigan coll.”. Wisconsin: RocK co.: Beloit, 1860, 


(US). Intinots: coox co.: Chicago, July 7, 1873, H. H. Babcock 
(US); prairie, Streator, Aug. 11, 1929, Howard K. Henry, no. 130 
(Pa); between the Sag and Palos Park, Sept. 1, 1908, Caldwell & 
Greenman, no. 3584 (Mo); Riverside, June 29, 1871, H. H. Bab- 
cock (NY), July, 1888, E. L. Sturtevant (Mo). wit co.: Joliet, 
July 4, 1900, H. P. Skeels (USNA). wINNEBAGo co.: Fountain- 
dale, M. S. Bebb, as Dipteracanthus strepens. WoopFORD CO.: 
Minonk, Aug. 13, 1895, Chas. Thom (USNA). PEORIA Co.: Peoria, 

68, J. T. Stewart (Phil); dry barrens, Peoria, June, 1887, F. E. 
McDonald; dry prairies, Peoria, July, 1903, and July, 1904, F. £. 
McDonald. Mason co.: Havana, Aug. 17, 1904, Gleason. HENDER- 


MACON CO.: openings in timber, Stevens Creek at Wabash, July 1, 
1915, I. W. Clokey, no. 2429 (TyPE in Herb. Gray; 1sorypEs in 


. : : us ee ~ Ree, _ tenn, ey annem eee osememmeme si Pon 
4 


1945] Fernald,—Ruellia in the Eastern United States 59 


1877, A. B. Seymour (Duke). st. cLAIR co.: woods, Belleville, 
July, 1846, Th. Hilgard, Jr. (Mo)—plant 8.25 m. high. RANDOLPH 
co.: Red Bud, June 30, 1888, L. H. Pammel (Mo). Muississiprt: 
OKTIBBEHA CO.: Agency, May 31, 1897, S. M. Tracy, no.. 3203 
(NY). Iowa: JOHNSON Co.: Aug. 13; 1909, M. P. Somes, no. 
vg (US). VAN BUREN CO.: open woods, Bentonport, July, 1920, 

LE. W. Graves, nos. 1687 and 1957 (Mo). WwayNE co.: July, 1885, 
R. KB. (Phil). GREENE Co.: Jefferson, July 24, 1867, J. A. 
Allen. MaApIsON co.: Peru, June 20, 1897, D. E. Hollingsworth, 
no. 577. RINGGOLD Co: Mount Ayr, July 24, 1926, Ada Hayden 


1925, L. H. Pammel, no. 733. MissouRI: MARION. Co.: dry woods, 
Hannibal, Aug. 7, 1912, John Davis (Mo). PIKE co.: dry mea- 
dow, near Clarkesville, June 15, 1914, John Davis, no. 2677 

0). ST. LOUIS CO.: limestone hill, Pacific, June 8, 1918, MGS 5 
Churchill (Mo). WASHINGTON CO.: Irondale, June 30, 1893, 


.2 Toc 
July 6, 1914, E. J. Palmer, no. 6205. PHELPS CoO.: ss 
June 16, 1914, John H. Kellogg east iy OONE Co.: banks 
Missouri River, Rocheport, July 4, H. W. Rickett (Duke. 
WRIGHT CO.: open woods, east of Manchold, June, 1911, O. E. 
Lansing, no. 3180. MORGAN co.: rocky woods, June ee 1934, 
Bush, no. 13,708 (Mo, Pa). HICKORY Co.: cherty limestone 
slopes, east of Jordan, July 10, 1934, Steyermark, no. 13,216 (Mo). 
GREENE Co.: Springfield, July, 1904, P. C. Standley (U8). JACK- 
SON co.: Independence, June 10, 1894, Bush, no. 349; barrens, 
Independence, July 6, 1900, Bush, no. 775 (Mo, US) and several 
other nos. (one of them with the penciled memorandum, “type 
of R. ciliosa”, a difficult proposition to defend since Pursh’ s 
species was from Savannah, Georgia!). BARRY Co.: fields, 
Eagle Rock, June 29, 1897, Bush, no. 506 (Mo). JASPER Co.: 
dry prairies, Carthage, July 13, bai Bush, no. 356 (Mo). 
MCDONALD co.: dry ground, Bush, no. 282 (NY), as R. ciliosa, 
var. three ARKANSAS: WITHOUT STATED LOCALITY: Arun- 
holdt, no, 24. pRArRIE co.: Hazen, Grand Prairie, June 29, 1941, 
Tenipres. no. 23,304 (Mo), as R. caroliniensis. CARROLL Co.: 
Eureka Springs, ‘July 7, 1898, N. M. Glatfelter (Mo). BENTON 
CO.: prairie-like thickets, Willow Springs, Aug. 9, 1941, Demaree, 
no. 22,380 (Mo), as R. caroliniensis. LOUISIANA! RED RIVER 
PARISH? J. Hale as R. etre Ell. NATCHITOCHES PARISH: eed 


60 Rhodora [FEBRUARY 


ELLIS co.: Cyril Zeller (Mo). OKLAHOMA: OTTAWA CO.: pasture 
near Narcissa, Aug. 31, 1937, F. R. Fosberg, no. 14,289 (Pa). 
at aay co.: rocky woods west of McAlester, May 27, 1920, 
W. Pennell, no. 10,600 (Phil). PAYNE Co.: Stillwater, June 
3 1926, Robt. Stratton, no. 622 ( slg oe soil north of Still- 
water, June 6, 1937, Darrell McLean n, no. 3 (Phil). BRYAN CO.: 
Colbert, June. 15, 1891, C. S. Sheldon, no. 14 (US), as R. ciliosa, 
r. longiflora. CLEVELAND CO.: Norman, Sept. 25, 1914, 
fa no. 522 (Mo), as R. parviflora. CADDO CO.: ‘between Fort 
obb and Fort Rebuekle 1868, Edw. Palmer (US), as ae 
canthus strepens. on i co.: Fort Sill, May 29, 1916, 
Joseph Clemens. TEXAS: NEWTON CO.: Autreville, "1857, 'C. GC. 
Fosberg (Phil). camp pa Pittsburg, Sept. 13, 1923, Tharp 
(Phil). Greee co.: July, 1939, C. L. York. woop co.: post-oak 
woods, July 6, 1924, Tharp (US). HARRIS co.: barren bluffs, 
Houston, Sept., 1 843, Engelmann (Mo). DALLAS CO.: vicinity of 
Dallas, July 3, 1929, Mary R. Stephenson, no. 96 (US). PARKER 
ce : Weatherford, July 4 1902, S. M. Tracy, no. 8078. TRAVIS 
: Austin, July 20, 1940, Tharp (transition to var. Sy gldtae 
och co.: near Corpus Christi, March, 1894, A. 
(NY). COMANCHE co.: DeLeon, June 16, 1941, Tharp. eb a 
Var. calvescens, var. nov. (TAB. 860), caulibus 1-4 dm. altis, 
amosis, ramis plerumque divergentibus, internodiis glabris vel 
glabrescentibus vel sparse breviterque hispidis; foliis glabris vel 
glabrescentibus vel ad venas remote hirtellis, anguste elliptico- 
oblon vel oblongo-lanceolatis, majoribus 2- 
0. 7-2.3 em. latis; corollis 2-3.5 cm. longis, tubo 0.7—2.3 cm. 
longo.—Cumber land Plateau of Kentucky and Tennessee, over- 
lapping slightly into southern Indiana and southern Ohio, 
locally in Alleghenies of northwestern Virginia, Great Smoky 
untains of eastern Tennessee and mountains of northern 
Georgia and Alabama; specimens, unless otherwise noted, dis- 
tributed as R. ciliosa or R. aie re hog i a IA: FREDERICK CO 
aa seine Middletown, Sept. 1 1, Hunnewell, no. 17, 361, 
as Dyschoriste oblongifolia (ieadteassiotaien ea in e). SHENAN- 
DOAH co.: dry limerock barrens, 1 pis forthanat of Strasburg, 
Aug. 30, 1927, Wiegand & Manning, no. 2934; rocky field near 
anda, July 27, 1942, Rane; aa 17 721 (FWH), as 
Dyschoriste oblongifolia. GEORGIA: WALKER CO.: Chickamauga 
Park, Biltmore Herb., no. 849 (US): Onto: HIGHLAND CO.: 
Sept., eng figex M. Roads, September, 1928 (US). InpIANA: 
CRAWFOR : in shallow soil on washed limestone slope, jus 
north of Leavenworth Oct. 5, 1920, Deam, no. 33,429 (Pa). 
KENTUCKY: WITHOUT STATED nega 1916, Anna King, no. 
310 (Duke and, in Part US); hills of Kentucky River, Aug. 15, 
1895, H. Garman & J. N. Rose (TYPE in Herb. Gray; ISOTYPE in 
U. 8. Nat. Herb. ), distrib. as R. strepens. FLEMING co.: dry soil 


cg ity = 


1945] Fernald,—Ruellia in the Eastern United States 61 


east of Hillsboro, Aug. 21, 1940, #. L. Braun, no. 3364 (Braun). 
ROBERTSON Co.: dry slopes, limestone soil, west of Blue Licks, 
July 8, 1941, Z. L. Braun, no. 4044. NIcHOLAS Co.: open woods, 
Blue Lick Springs, Sept. 17, 1936, F. J. Scully, no. 814 (US), as 
R. parviflora; dry soil, open red cedar, south of Fairview, Sept. 
22, 1939, HE. L. Braun, no. 2684 (Braun). scotr co.: dry woods, 
Stamping Ground, July 27, 1931, J. W. Singer, no. 145 (US), as 
. parviflora. PULASKI co.: bluffs near Burnside, Aug. 22, 1903, 
Biltmore Herb. no. 849* (US). NELSON co.: prairie patches, 
Balltown, July 22, 1940, #. L. Braun, no. 3248 (Braun); dry 
limestone slope, prairie patches, west of Bardstown, Aug. 18, 
E. L. Braun, no. 4085. HARDIN co.: thin soil over limestone, 
Howe Valley, Sept. 6, 1927, Wherry & Pennell, no. 73,642 (Phil). 
HART CO:. sandy roadside, 2 miles southeast of Horse Cave, 
Sept. 4, 1930, Svenson, no. 4418. Grayson co.: Leitchfield, 


TENNESSEE: SEVIER co.: between Gatlinburg and Maryville, 
July 27, 1936, W. C. Coker (NC). witson co.: dry cedar glades, 
Lebanon, Aug. 11, 1900, Biltmore Herb. no. 8498 (US). DAvID- 
SON co.: cedar barrens, common, Gattinger (Mo); Nashville, 
Aug., 1879, Gattinger (Mo); West Nashville, May 26-27, 1909, 
Eggleston, no. 4436 (US), Sept. 24-25, 1909, Eggleston, no. 5160 
(NY, Phil). corree co.: near Manchester, Aug. 14, 1899, 
Biltmore Herb., no. 849 (US). RUTHERFORD CO.: near Lavergne, 
Aug., 1897, Henry Eggert (Mo). FRANKLIN co.: Cumberland 
Mts., Cowan, July, 1898, Ruth, no. 556 (NY), no. 590 (Mo). 
ALABAMA: MADISON Co.: ad montes, prope Huntsville, Oct., 1843, 
Rugel (NY), with unpublished but quite appropriate varietal 
name. Map 13. a 

When sorted out from the very diverse species with which it 
has been confused, Ruellia ciliosa, R. caroliniensis, and even the 
wholly different R. strepens and the apocryphal R. parviflora, 
Ruellia humilis becomes a very consistent species of wide inland 
range. By the singular and not very edifying fatality which has 
obscured the clarification of even the simpler of our species of 
Ruellia the great bulk of material of the continental R. humilis 
(including its varieties) has very generally passed as R. ciliosa 
Pursh, its type from Savannah, Georgia, although there is no 
evidence in any of the larger American herbaria that the present 
species approaches Savannah nearer than the Shenandoah Valley 


62 Rhodora [FEBRUARY 


of Virginia, the Great Smoky Mountains of Tennessee, the 
mountains of northwestern Georgia, and the northwestern corner 
of Florida. Conversely, although R. humilis was described from 
‘rocks in the upland forests and prairies’ of Arkansas, the name 
R. humilis has been consistently used by Small and his followers 
for true and strikingly different R. ciliosa Pursh and given a range, 
“Sandy soil, Coastal Plain, Fla. to Miss. and Ga.’’, through a 
country without the dry limestone rocks where true R. humilis 
prevails, and entirely excluding the type-region of the latter! 

There is great probability that Ruellia hirsuta Ell. Sk. 11. 109 
(1822) was typical R. humilis Nutt. Elliott, familiar with the 
more eastern plants, went beyond the stated limits of his Sketch 
to describe a remarkable new species from “near the Alabama 
River in dry soils”: “‘ Hirsute, branching, leaves oval-lanceolate, 
nearly acute, sessile, segments of the calyx subulate, hispid, a 
little longer than the tube of the corolla Tea re ewe 
very hirsute. Leaves . . . almost hispid’. Elliott’s type is 
lost; furthermore his name was antedated by R. hirsuta Velloso 
(1790). 

As the TYPE or IsoTYPE of Ruellia humilis (until a possibly 
more authentic type is found) I have taken an Arkansas speci- 
men from Nuttall, in the Torrey Herbarium. This (PLATE 854, 
Fic. 1) is of the relatively short-flowered series with narrow 
leaves, a plant (Map 9) prevailing west of the Mississippi, south- 
ward into Texas, but scattered eastward even to the mountains 
of south-central Pennsylvania, western Maryland and western 
Virginia. This plant passes through obvious transitions into a 
coarser and broader-leaved extreme of similar range (map 10), 
which might perhaps be considered an ecological variety were it 
not that in their morphological characters they exactly parallel 
the two plants with greatly prolonged corolla-tube (the longest 
corolla of any of our species except the southeastern Coastal 
Plain R. noctiflora). Although the very long-flowered plants 
have similar areas of development, the narrow-leaved var. longi- 
flora (MAP 11) is more restricted, its eastern range stopping 
essentially at the Mississippi, with northern limits in southern 
Illinois and Missouri; the broader-leaved var. erpansa (MAP 12) 
with outposts eastward to northwestern Florida, northeastern 
Alabama and in Indiana and Wisconsin and, farther west, 


1945] Whelden,—Marine Algae of the Monterey Peninsula 63 


extending well into Iowa. These two plants, although merging, 
seem fairly well defined. In Texas and Louisiana var. longiflora 
has often been mistaken for the more eastern Coastal Plain R. 
noctiflora. It is promptly distinguished, however, by its greatly 
developed villous-hirsute pubescence, R. noctiflora being puberu- 
lent; by its short and strongly hirsute-ciliate calyx-segments, 
those of R. noctiflora prolonged (2.5-4.5 em. long) and puberu- 
lent; and by its short and glabrous capsule, the very long 
(2.25-3.5 em. long) capsule of R. noctiflora pubescent. 

As extreme as any variation of Ruellia humilis is var. calvescens 
(mar 13), concentrated on the Cumberland Plateau, with 
colonies along the Alleghenies and the Great Smokies. Smaller 
in all parts than most members of the species and nearly glab- 
rous, it has been mistaken (by the writer among others) for 
Dyschoriste oblongifolia (Michx.) Ktze. In their more upright’ 
extremes the two plants are often superficially similar, but the 
technical differences of calyx-segments and anthers hold. Fur- 
thermore, when the ripe seeds of R. humilis, var. calvescens, like 
those of all our species of Ruellia, are moistened they promptly 
exude mucilage which soon takes the form of prolonging and 
streaming spiracles which finally give the seed a more or less 
plush-like surface. Many tests of seeds of Dyschoriste oblongi- 
folia have shown no such mucilage-spiracles there. The latter 
species, furthermore, is a plant of southern Coastal-Plain pine 
barrens and fall-line sands, northward to South Carolina (the 
old and doubted basis for it as Virginian, in the Gray Herbarium, 
being a specimen with copied—not original—label, reading 
“Southern States. Virginia’’), not of calcareous upland. 


(To be continued) 


1945] Fernald,—Ruellia in the Eastern United States 71 


elongate floriferous ascending basal branches; median 
leaves oblong, oblong-lanceolate, ovate or elliptic, sub- 
equal, subcoriaceous, often undulate-dentate...... 11]. Var. dentata. 


lla. Var. typica. Stem when well developed 1.5-7 dm. high, 
simple, or with divergently to horizontally spreading elongate 
branches mostly flowering at tips, obtusely angled, canescent- 
pilose to copiously white-villous; middle and lower internodes 
greatly elongated (5-18 cm. long), the uppermost greatly. abbre- 
viated: principal leaves elliptic, subrhombic, oval, ovate or 
ovate-lanceolate, membranaceous, slender-petioled, obtuse to 
acute, white-villous to villous-hirsute or canescent-pilose be- 
neath when young, closely strigose above; the blades 3-10 cm. 
long and 1.5—4 cm. broad, with somewhat undulate surface, thus 
crumpled or puckered in drying: glomerules very short-peduncled, 
trom 1-3 approximate or subapproximate upper nodes and at 
tips of longer branches, 2-many-flowered: bracts oblong-lanceo- 
late to narrowly elliptic, nearly equaling to shorter than calyx: 
calyx-segments linear-setaceous, white-villous or canescent- 
pilose on back, villous-ciliate, 1.3-2.3 cm. long, usually con- 
siderably shorter than corolla-tube: corolla bluish-lavender, 
2.5-5 cm. long; the slender tube 1.3-2.7 em. long, the campanu- 
late-obconic throat 1-1.5 cm. thick at summit; expanded limb 
2.5-3.5 cm. broad: capsule often densely pilose-hirtellous, 
sometimes glabrous, 1.4-1.7 cm. long.—R. strepens, capitulis 
comosis Dillenius, Hort. Elth. ii. 330, t. 331 (1732). KR. strepens 
L. Sp. Pl. 634 (1753), in part, not L. Mantiss. Alt. 422 (1771). 
Anonymos caroliniensis Walt. Fl. Carol. 168 (1788). Pattersonza 


Lyc. N. Y. 
(1824), in part only; Small, FI. Se. U. S. 1084 (1903) and Man. 
1229 (193 
eme 


change of spelling of both names) directly from J. F. Gmelin; 
Blake in Ruopora, xxii. 134 (1915), in part, excluding R. ciliosa 
and R. carol. var. parviflora (Nees) Blake and all synonyms 


anus Nees 1. ec. (1847). R. ciliosa, var. hybrida (Pursh) Gray, ae 
Fl. iit, 326 (1878), in part—Sandy woods, Florida, north to 
eastern South Carolina; Tennessee, Kentucky ‘bu ou R. 
Arkansas (unless otherwise noted, specimens erat hillside, 40- 


70 Rhodora [Marcu 


pono Ss ne 
upper more or less approximate nodes chiefly 
longer nein those boner f entire or not very dentate; 1 or 2 
(rarely 3 or 4) nodes of the main axis floriferous, the low- 
pecshonasbosings ones from closely crowded to as (-15) cm. 
mple, only rarely branched... . 
d. oan rsaseings of the uppermost noden broadly ‘ieselete 
o oval, ovate or elliptic, if lance-subacuminate 1.5- 
4.5 cm. ‘broad; calyx-segments glabrous or nearly so on 
ck; summit of herigaod of eal ve 7-1.5 cm. broad, 
expanded lim limb 2.5-4 cage 
. Leaves m Cates oock “phabie, a ovate or elliptic 
to broadly lanceolate... .f. 
Fae stoutish to slender, 1-5 mm. thick at base, 1-9 
dm. high; larger leaves 5-12 cm. long, if obtuse 
snatie Tis ore than 7 cm. long... .g 
g. Leaves ee ispid on veins beneath; calyx- 
segmen 
inten aa stem (or some of them) copiously 
spreading-hirsute............... d. Var. membranacea. 
teterncdee aon eoe teh to glabrescent, at most 
with remote elongate trichomes 
1 Var. membr., forma breviberbis. 


° 
y 
BS 
= 


g. Leaves glabrous beneath. 
Calyx-segments oliates: internodes of stem se 
aly acini upper surfaces of leaves 
ally strigose........ 1lf. Var. einen ‘forms hypopsila. 
Calyx. Hossa hl eciliate; internodes glab 
pe glabrous; By tog aces of leaves sabeode 
essentially so........ llg. Var Se ae forma laevior. 
f. Stem fiform, 1-2 Pia thick at abe, 1-2 (-4) dm 
; leaves membranaceous, elliptic to oblong, 
= larger ones 32-6 em. long, obtuse to rounded at 


rar malar — internodes (or some of 
them) of a piously villous-hirsute with 
aversen’ Bae ss es Es heard rod nanella. 
alyx-segments palin: internodes glabresce 
or merely puberulen oe Var. ena , forma eciliata. 


e0 
lance-ovate, subacuminate; stem rigid, 4.5-9 


Wes a Se 11j. Var. cheloniformis. 
d, iene leaves of the uppermost acme lance- 
olate to Loved eg 0.5-1.5 C Cg em. broad; stem 
slender, 1.5-7 dm. ; calyx-segments canescent- 
pilose to glabrescent; throat of corolla 5-10 mm. broad 
at summit, expanded li .5 em. broad..... 11k. Var. salicina. 


c. ~~ of flowering s summit gradually much reduced i in — 
es 0 


all but upper internodes elongate; stem frequently with 


1945] Fernald,—Ruellia in the Eastern United States 71 


elongate floriferous ascending basal branches; median 
leaves oblong, oblong-lanceolate, ovate or elliptic, sub- 
equal, subcoriaceous, often undulate-dentate...... 11]. Var. dentata. 


lla. Var. typica. Stem when well developed 1.5-7 dm. high, 
simple, or with divergently to horizontally spreading elongate 
branches mostly flowering at tips, obtusely angled, canescent- 
pilose to copiously white-villous; middle and lower internodes 
greatly elongated (5-18 cm. long), the uppermost greatly. abbre- 
viated: principal leaves elliptic, subrhombic, oval, ovate or 
ovate-lanceolate, membranaceous, slender-petioled, obtuse to 
e, white-villous to villous-hirsute or canescent-pilose be- 
neath when young, closely strigose above; the blades 3-10 cm. 
long and 1.5—4 em. broad, with somewhat undulate surface, thus 
crumpled or puckered in drying: glomerules very short-peduncled, 
trom 1-3 approximate or subapproximate upper nodes and a 
tips of longer branches, 2—-many-flowered: bracts oblong-lanceo- 
late to narrowly elliptic, nearly equaling to shorter than calyx: 
calyx-segments linear-setaceous, white-villous or canescent- 
pilose on back, villous-ciliate, 1.3-2.3 cm. long, usually con- 
siderably shorter than corolla-tube: corolla bluish-lavender, 
2.5-5 cm. long; the slender tube 1.3-2.7 cm. long, the campanu- 
late-obconic throat 1-1.5 em. thick at summit; expanded limb 
2.5-3.5 em. broad: capsule often densely pilose-hirtellous, 
sometimes glabrous, 1.4-1.7 cm. long.—R. sérepens, capitulis 
comosis Dillenius, Hort. Elth. ii. 330, t. 331 (1732). R. strepens 
L. Sp. Pl. 634 (1753), in part, not L. Mantiss. Alt. 422 (1771). 


-) : i . e 
wrongly ascribed to Walter, p. 167 (instead of 168) with a brief 
summary of Walter’s generic diagnosis. 2. hybrida Pursh, Fl. 
Am. Sept. ii, 420 (1814); LeConte in Ann. Lyc. N. Y. i. 140 


ed. 2 0 
Walt.”, without further reference, this obviously copied (with 
change of spelling of both names) directly from J. F. Gmelin; 

lake in RHopora, xxii. 134 (1915), in part, excluding R. ciliosa 


6. hybridus (Pursh) Nees 1. c. 123 (1847), in part. D. Mitchilli- 
anus Nees ihe 5847), R. ciliosa, var. hybrida (Pursh) Gray, 
Fl. iit. 326 (1878), in part.—Sandy woods, Florida, north to 
eastern South Carolina; Tennessee, Kentucky and _ 
Arkansas (unless otherwise noted, specimens distributed as R. 
hybrida). Souru CAROLINA: LANCASTER CO.- rocky hillside, 40- 


72 Rhodora [Marcu 


Acre Rock, west of Taxahaw, Dorothy Huntley, os 256 (Duke). 
HORRY Co.: low woods near Myrtle Beach, July 13, 1932, Coker 
& Totten (NC). GEORGETOWN CO.: sandy Done Brookgreen 
Gardens, F. G. Tarbox, no. 567 (US). WILLIAMSBURG CO.: ric 
woods, 10 miles southeast of Gourdin, bd & Tryon, no. 433. 
BERKELEY ig ¥: Santee Canal, H. W.R enel, as R. strepens, 
changed by Asa Gray to R. ciliosa, var. PAS. CHARLESTON 
go; Charleston, June, 1902, Wm. Palmer (US), as R. parviflora. 
BEAUFORT CO Bluffton, "1872, Mellichamp (Mo, US); St. 
Helena, 1878, ‘Laura Towne (Pa), as R. ciliosa. GEORGIA: 
WITHOUT STATED LOCALITY: Chapman, CHATHAM Co.: Savannah, 
Mrs. Say. FLORIDA: WITHOUT DEFINITE LOCALITY: E. Florida, 
Chapman (NY), as Dipteracanthus Mitchillianus. DUVAL CO.: 
dry rich ground near Jacksonville, A. H. Curtiss, no. 4764 (Mo, 
Y, U8), . strepens, one sheet changed to A ee 
another to R. parviflora; St. Nicholas, May 12, 189 H. 
Lighthipe (NY), Re R. ciliosa, April 20, 1897, J. R. Churchill: 
vicinity of May and Jacksonville, 'H. ea Keeler (NY), as 
R. ciliosa; pasate Fredholm, no. 5150, as R. cilvosa. st. 
JOHNS co.: May 20-22, 1885, G. C. Whitlock. or co.: dry oak 
woods, Green Cove Springs, April 14, 1939, W. A. Murrill 
Mo), as R. ciliosa. PUTNAM CO.: Crescent City, March, 1880, 
G. Marten (Phil). ice co.: open pine woods I mile north 
of Newberry, E. Perot Walker, no. 1862 (Phil), as R. humilis; 
high hummock, Burnett’s Lake, April 30, 1939, W. A. Murrill 
(Mo); Gaineaville, 1887, M. F. Price, as R. ciliosa, var. piletgae! 
May 12, 1897, Joseph Crawford (Phil. ), as R. ciliosa; hamm 
near Gainesville, May 31, 1937, W. A. Murrill (Mo), as R. ilaaed: 
VOLUSIA CO.: Port Orange, F. C. Straub, no. 120, as R. czliosa, 
var. ambigua; sandy soil, Ormond, April 23, 1903, ‘B.A. Purdie, 
as R. ciliosa, altered to R. humilis; Log growth by road, north 
of Ormond, May 16, 1943, E. H. Butts & Oakes Ames ( Ames); 
in shade or half-shade, Ormond Beck May 12, 1943, Butts & 
le Signe dry soil, Crescent City, June 28, 943, pe oS 
Butt rviflora. LAKE Co.: vicinity of Eustis, N ash, nos. 
42 (Us) 7 1801, Tabs. 1894, A. S. Hitchcock (Mo); pinelands 
east of Eustis, J. K. Small, no. 8667 (NY), as R. parviflora; dry 
wood-border, "Hiawatha Lake, Wiegand & Manning, no. 2932, 
as R. parviflora. SUMTER co.: Lake oe March 29, 187 9, 
J. D. Smith (US), as R. strepens. HERNANDO CO.: rich open 
woods, Brooksville. April 9, 1927, Hugh ON eill (Mo). POLK 
co.: Peace Creek, March, 1 1880, J. D. Smith (US), as x strepens, 
dry pineland, —) 12, 1894, L. B. Ohlinger (Mo), as R. strepens. 
OSCEOLA CO.: sandy soil, Kissimmee, April 5, 1936, “M ary L. 
ri cee | (Duke). ep TEE CO.: Manatee, 8. M. Tr , no. 
675 , US). DADE co.: pine woods near Rockdale, J. L. 
Fennell, no. ae (USNA). Kentucky: owrNn co.: about 1 


: -_ actinic —_-<eniiinaniiane  —reatiti———ni —, R —  —  — — TLS —y 


1945] Fernald,—Ruellia in the Eastern United States 73 


mile north of Scott Co. line, EZ. L. Braun, no. 3204 (Braun). 
JEFFERSON CO.: 3 miles south of Louisville, July 5, 1892, L. S. 
Bergman, in part (Mo), as R. ciliosa. caLLOWAY co.: upland 
oak woods just west of the Tennessee River, Gleason, no. 8950 
. TENNESSEE: CocKE co.: Newport, June 6, 1925, O. M. 
Freeman (USNA). KNox co.: Knoxville, Ruth, no. 101, as R. 
strepens. FRANKLIN Co.: Cowan, Biltmore Herb., no. 849° (US), 
as R. ciliosa; woods north of Sheridan, June 6, 1897, H. Eggert 
(Mo). pDAvipson co.: Ridge Top, July 13, 1897, Eggert (Mo). 
SHELBY co.: Memphis, Fendler, as R. ciliosa. ARKANSAS: ST. 
FRANCIS CO.: Crowley’s Ridge, Forrest City, Demaree, no. 15,137 
(Mo), possibly, when more material available, to be placed 
elsewhere. Puiatses 861-863; MaP 14. oo. 
11b. Var. semicalva, var. nov. (TaB. 864), var. typicae simil- 
lima; caule vix villoso-hirsutis; foliis subtus glabrescentibus 
vel sparse strigosis; capsulis plerumque glabris.—Of much wider 
range, from central Florida to South Carolina, locally to south- 
eastern Virginia, westward to eastern Texas. VIRGINIA: SOUTH- 
AMPTON Co.: rich woods, Violet Hill, near Devil’s Elbow, June 
23, 1936, Fernald, Long & Smart, no. 5922 (type in Herb. Gray; 
IsoryPEs in Herb. Phil. Acad., etc.), as R. ciliosa; about Franklin, 
Heller, no. 953 (US), as R. ciliosa. NortH CAROLINA: CHATHAM 
co.: dry soil, Silver City, Biltmore Herb., no. 849! (US), as R. 
ciliosa. COLUMBUS Co.: grassy woodland at Lake Waccamaw, 
Godfrey & Shunk, no. 4170, as R. parviflora. NEW HANOVER CO.- 
Wilmington, June, 1894, J. M. Macfarlane (Pa). SouTH Caro- 
LINA: GEORGETOWN Co.: shady, weedy waste place, Georgetown, 
Godfrey & Tryon, no. 1683. WILLIAMSBURG CO.: rich woods, 10 
miles southeast of Gourdin, Godfrey & Tryon, no. 433. BERKELEY 
co.: railroad right-of-way, south of Moncks Corner, Be 
Martin, no. 1130 (USNA), as R. parviflora. CHARLESTON CO.: 
near Charleston, Aug. 1886, L. R. Gibbes (NY), as R. parviflora. 
ANDERSON co.: dry rich woods, Anderson, John Davis, no. 9193 


O. M. Freeman (USNA). Fiortpa: LAKE co.: clay soil, vicinity 
of Eustis, N oe no. ai. sumTER co.: March 29, 1879, J. = 
Smith (US), as R. strepens. cITRUS co.: moist shaded rock 
north of Pineola, H. J. Oosting, no. 185 (Duke). Brevarp co: 
Coco, J. K. Small, no. 8727. 
of Sanford, Moldenke, 


N 
no. 184 (Duke, Mo, NY, Pa), as R. parviflora. POLK CO.- sandy 


74 Rhodora [Marcn 


roadside, Winter Park, March 8, 1923, Hunnewell, no. 8732 
(FWH). HILLSBORO CO.: Tampa, May, 1876, A. P. Garber 
ese US), as Dipt. ciliosus. LEON Co.: near Tallahassee, Nik; 
g (NY), as R. parviflora. WAKULLA Co.: prope St. Marks, 
Mal 1843, Rugel, with an unpublished name (NY). CALHOUN 
co.: Iola, Chapman (Mo). ‘TENNESSEE: HAMILTON CO.: si 
fields, Sequach Valley, Biltmore Herb., no. 849" (US), as 
ciliosa. ALABAMA: DEKALB AND ETOWAH cos.: Lookout Mt, 
July 6 and 8, 1898, H. Eggert (Mo). ETOWAH CO.: Gadsden, 
1878, G. R. Vasey (US), a s R. ciliosa. CULLMAN CO.: woods, 
June and Sept., 1897, H. Bipers (Mo). HALE co.: Greensboro, 
1857, Sereno Watson, as R. ciliosa, var. ambigua. LEE CO.: dry 
pine ‘woods, J. D. Smith, no. 1940 (US), as R. strepens. MONT- 
GOMERY Co.: low abandoned field near Montgomery, June 18, 
1932, a K. Edwards (Pa). Dawuuas co.: 1879, Wm. Trelease 
(Mo ipt. ciliosus. CONECAH co.: Evergreen, Baker 
Earle, on 39 (US), as ar ciliosa. MONROE Co.: dry hills, July 23, 
1885, Mohr (US), as R. ciliosa. MOBILE CO.: Mobile, 1905, 
. Dukes, as R. parviflora; pine barrens, Spring Hill, £. 

Graves, no. 498, in sori (Mo). MISssIssIPrr: GRENADA CO.: 
moist wooded hillside, Pay West’s Lake, May 27, 1932, Vena 
Millsaps (NC), as R. ‘ciliosa. OKTIBBEHA CO.: Starkvill e, July 
26, 1890, S. M. Tracy, as R. strepens pale tO ccs habitally 
resembling var. dentata. LAUDERDALE Meridian, S. 
Tracy, no. 3271 (NY), resembling var. Sioa JACKSON 
co.: Ocean Bi 3 agg ses ih puri as Seymour & Earle, 
154, as R. ciliosa, var. ambig June 7, 1895, Skehan, as "R. 
ciliosa (eanuleting var. Ghelandeoniay. HARRISON co.: Biloxi, 

. M. Tracy, nos. 4944 (Mo —very mixed no., some 
specimens nearly var. salicina), 6434 (NY, Us), 6435 (G, Mo, 
NC, NY, US—some plants nearly var. salicina, others ap- 
proaching var. nanella), as R. strepens. copiaH co.: Brushy 
Creek, Crystal Springs, May 18, 1925 (US), as R. parviflora. 
LOUISIANA: ST. TAMMANY PARISH: segremae of Covington, Bro. 
Anect, no. 65 (US), as R. parviflora; Bro. G. Arséne, no. 12,240 
(US), as R. parviflora. IBERIA PARISH: moist open grassy woods, 
Avery Islan .S. & H. B. Correll, no. 9525. RAPIDES PARISH: 
Alexpndria, Hale ake as R. strepens, corrected by Britton to 
R. ciliosa. NATCHITOCHES PARISH: woods near Marthaville, 
May 5, 1893, fabs (US), as R. ciliosa. Trxas: Page Co.: 
sandy woods, Big Sandy, Reverchon, no. 2536 (Mo), as R. parvi- 
flora. LIBERTY CO.: eee woods, Dayton, E. J. Palmer, no. 7768 
(Mo), as R. parviflora (plant Heese large-leaved var. 
membranacea). HARRIS cO.: Houston, G. L. Fisher, no. 81 (US), 
as R. parviflora. PuaTE 864; risa or 

lle. Var. SEMICALVA, forma detonsa, f. nov., caule, foliis, 
calycis segmentibusque minute canescento-puberulis eis ecilia- 


1945] Fernald,—Ruellia in the Eastern United States 75 


tis.—FLoripa: roadside, Sebring, Highland Co., March 4, 1924, 
Hunnewell, no. 9044 (rypr in Herb. Hunnewell). 

lid. Var. membranacea, var. nov. (TAB. 865 et 866), caule 
1-9 dm. alto ad basin 1-4 mm. diametro, internodiis valde 
divergenterque hirsutis; foliis membranaceis 5-12 cm. longis 
supra strigosis subtus ad venas strigosis vel hirsutis, ovalibus, © 
ovatis, ellipticis vel late lanceolatis obtusis vel subacutis; calycis 
segmentis dorso glabris vel glabrescentibus longe ciliatis—R. 
parviflora sensu Small, Fl. Se. U. S. 1085 (1903) and sensu 
Britton in Britton & Brown, Ill. Fl. ed. 2, iii. 242 (1913), excl. 
fig., which is of R. Purshiana; not R. parviflora (Nees) Britt. 
Man. 854 (1901), at least as to basonym, Dipteracanthus ciliosus, 
var. parviflorus Nees.—Woods, openings and clearings, either 
caleareous or subacid, north-central South Carolina to south- 
central Tennessee, north to southern New Jersey, southeastern 
Pennsylvania, northern Maryland, West Virginia, southeastern 
Ohio and north-central Kentucky (unless otherwise noted, dis- 
tributed as R. caroliniensis, ciliosa or parviflora); usually com- 
moner than other vars. northward. New JERSEY: CAPE MAY 
co.: woods south of Mill Lane, north of Cape May City, Aug. 
10, 1918, W. Stone (Phil); north of Cape May, Sept. 13, 1901, 
C. S. Williamson (Phil); Cape May Court House, Aug. 6, 1909, 
Van Pelt (Phil); Cold Spring, Sept. 13, 1908, C. S. Williamson 
(Phil), July 18, 1909, Van Pelt, no. 12,199 (Pa). PENNSYLVANIA: 
YORK : below McCall’s Ferry, July 11, 1895, Joseph Crawford 
(Phil). DELAWARE: NEW CASTLE co.: Aug. 9, 1866, A. Commons 
(NY), as R. strepens. KENT CO.: moist woods west of Woodland 
Beach, R. R. Tatnall, no. 1462. sussex co.: Rehoboth, July 8, 
1908, C. S. Williamson (Pa, Phil); sandy woods east of Milford, 
July 16, 1896, A. Commons (Pa, Phil). MARYLAND: CECIL CO.: 

k Neck, June 24, 1923, W. L. Abbott (Phil); woods, high bluff 

Aug. 6, 1938, Mrs. L. R. Holmes (Pa); 

Leslie, July 15, 1891, Crawford (Phil); flats, Conowingo, July 29, 

1924, Crawford; Conowingo, July 25, 1907, J. J. Carter Leet 
NT 


(US). queen anne co.: Aug. 20, 1900, { 

TALBOT Co.: railroad ballast, northwest of St. Michaels, R. R. 
Tatnall, no. 386 (Pa); edge of woods, west of Easton, E. C. Earle, 
nos. 2251 (Pa) and 3698 (Pa). WICOMICO CO.: Salisbury, July 14, 
1904 J. Carter (Phi WORCESTER co.: roadside through 


(Duke). HarrorpD co.: low, moist woods, Creswell, July 28, 
1883, L. W. Brosius (Pa). BALTIMORE CO.: Baltimore, Leroy 
(NY). prince GrorGE co.: Hyattsville, Aug. 13, 1904, Steele 
(Mo). caLverRT co.: low swampy woods, North Bay, Chesa- 
peake Bay, Muenscher, no. 3867 (Mo). MONTGOMERY CO.: | 

f woods, July 7, 1901, M. W. Lyon, Jr. (US). WASHINGTON 


76 Rhodora [Marcu 


co.: island above Harper’s Ferry, Ff. W. Pennell, no. 2430 tre 
District oF CoLUMBIA: in vicinis Washington, June 29, 1884 
L. F. Ward; thickets near Ivy City, June 14, 1891, F. Blanchard 
(Mo), as R. strepens; open woodland, Washington and vicinity, 
July 16, 1896, Steele (NY, Mo); Congress Heights, Philip Dowell, 
no. 7012. West VIRGINIA: NICHOLAS CO.: dry bank, Peter 
Creek, Millspaugh, no. 596 (NY). MASON Co.: along Ohio 
River, near Pum Roy, E. Berkley, no. 9138 (Mo). CABELL 
0.: abandoned wooded pasture, Pleasant Valley, Gilbert, no. 
483. LINCOLN co.: near Mills School, EH. E. Berkley, no. 969 
(Mo). WAYNE co.: dry hillside, Buffalo Creek, Lewis Plymale, 
no. 95. a: gander and WYOMING Cos.: woods along Indian 
Ridge, E. Morvis, a 1172 (US). VrirGINIA: FAIRFAX CO.: 
Falls hah, June 23, 1872, J. J. Carter (Phil). ALEXANDRIA 
co.: Naucks, ewad : no. 2456 (Phil); Fort Myer, July 14, 1895, 
E. A. Mearns , as R. strepens. NORTHAMPTON CO.: ’ sandy 
woods back of the hanen: Savage Neck, Fernald & Long, no. 
5453. MIDDLESEX co.: bank of Rapahannock River at Gray’s S 
Point, A. B. Massey, no. 2115 (VPI). MATHEWS co.: sandy op 

soil along North End Branch, “Fort Nonsense”, south of Sela: 
Wherry & Pennell, no. 12, 616, in part (Phil). GLOUCESTER CO.: 
Beaver Swamp Creek, north of Gloucester, July 5, 1924, H. B. 
Meredith (Phil). yorK co.: steep wooded bank, York River, 
above Mt. Folly, Fernald & Long, no 13, 459. JAMES CITY CO.: rich 
a and slopes by James River, Grove La nding, southeast of 
Grove, Fernald & Long, no. 13 ABT: moist rich soil in woods, west 
aida of Jones Millpond, Williamsbure, Grimes, no. 3887. CHARLES 
cITy co.: wooded slope by James River, at “Four Oaks”, below 
Harrison Point, ag & Long,'no. 13, 149. PRINCESS ANNE CO.: 
rich deciduous woods east of Little Creek, Fernald & Long, no. 
4193; sandy woods, northern end of Knott’s Island, Fernald & 
Long, no. 4194, in part. NORFOLK Co.: Portsmouth, June, 1840, in 
part, Rugel (NY). ISLE OF WIGHT Co.: thicket back of sandbeach 


nos. 13,762 and 13,837; Heh calcareous wooded ravine near James 
River, ‘northwest of Chippokes, July 25, 1941, Fernald & Long, 
no. 13 ,458. SOUTHAMPTON co.: border of sandy woods south of 
Applewhite’ s Church, Fernald & Long, no. 13,153. SUSSEX CO.: 
border of woods near Nottoway River, Green Church Bridge, 
southwest of Owen’s Store, Fernald & ong, no. 12,472 (TYPE in 
erb. Gray.; ISOTYPE in Herb. Phil. Acad.). GREENVILLE CO.: 
bottomland woods along Meherrin River, southeast of Gaskins, 


erna ; 
Olivet Church, Fernald & Long, no. 14,021. ametia co.: J. B. 
Lewis, nos. 114 and 733 (VPI). ip occ co.: bottomland 


é meres — eile. eat, Nl, CCT. A CO, TT TT TTT a TT: LS 


¢45) Fernald,—Ruellia in the Eastern United States 77 


woods near Western Bridge, Meheus River, south of Edgerton, 
Fernald & Lewis, no. 14,498. CAMPBELL Co.: vicinity of Lynch- 
burg, July 1, 1802, Britton, Britton & ail (NY). BEDFORD CO.: 
July 8, 1871, _H. Curtiss (Mo, NY). RocKBRIDGE co.: ‘‘ Near 
the natural ree in rich soil along feo Pursh (Phil), correctly 
called ‘‘Anonymos caroliniensis Walt.”’; - Natural Bridge, Margaret 
P. Russell. Norta Carouina: PASQUOTANK co.: oak-pine-beech 
woods south of Elizabeth City, Wiegand & Manning, no. 2929 
TYRRELL CO.: open woodland, Columbia, Godfrey, no. 4322. PITT — 
Se pine woodland near Farmville, Godfrey, no. 4278. WARREN 
: Warrenton, July 19, 1924, H. B. Meredith (Phil). GRan- 
ee co.: wet meadow, Oxford, M. F. Buell, no. 1404 (US). 
DURHAM CO.: sided low woods, Duke Forest, Blomquist, no. 
367 (US). WAKE co.: sandy bank west of Raleigh, Wiegand & 
Manning, no. 2930. BLADEN co.: moist sandy soil, Biltmore 
Herb., no. 849%. ORANGE co.: woods near Chapel Hill, Aug. 17, 
1909, Coker (NC), June 23, 1933, Ms 7. Cameron (NC); dry 
open woods, Hollow Rock, C. c. Wilson, no. zo (Mo). CHATHAM 
co.: dry soil, Silver City, Biltmore H erb., . 849). FORSYTH 
co.: moist woods near Mt. Carmel, Coral 3 no. 2591 (Duke). 
SURRY CO.: moist shaded cove, Pilot Mt., R. M. Williams, no. 
464 (Duke). ROWAN CO.: vicinity of Salisbury i no. 140 
; Pha). — pee near Columbus, July 2 . 1897; Be: 
Townsend (US). ON CO.: open woods, bluk along river, 
Hot Springs, Coen = 34,230 (Duke). BUNCOMBE co.: Bi It- 
more, Biltmore Herb., no. 849», in part (US); Weaversville, 
July 8, 1933, H. B. Teague (Duke). SWAIN co.: partly shaded 
roadside near Cherokee eee rare, ft. A. McLean, no. 73 


of Batesburg, E. A. Me asa no. 309 (US), as R. en. 
GREENVILLE cCo.: summit of Paris Mt., July, 1896, J. K. Small 
Y). ANDERSON co.: damp soil, Anderson, John Davis, no. 


( . 
“dry ground” by B. F. Bush whose Missouri plant grows in dry 
ground). oekires co.: thickets, A. P. Anderson, no. 1222 (US); 
Clemson College, H. D. House, no. 2384 (NY). Gxoreta: 
CLARKE CO.: roadside, Athens, L. M. Perry, no. 1083, as &. 
humilis. Onto: mics co.: Letart, Sept. 10, 1985, C. H. _ Jones 
NY). Kentucky: LOCALITY NOT DEFINITELY PLACED ae 
river hills, mouth of Hickman Cr.”, July, 1832, C. W. Shor 
R. strepens (Phil), sent to Hooker who wrote in 1833: The bs 

uellia [var. nenetal with small flow® looks diff\—But Iam much 


puzzled with the Genus”. ROWAN Co.: prairie patch, Clark Mt., 


Braun, no. 1930 (Braun). MENIFEE CO.: oak woods, ry 
limestone slope, Red River valley, E. L. Braun, no. 2128 . 
BELL Co.: rocky woodland, Pineville, Pennell, no. 11 ,804 (Phil) ; 


78 Rhodora [Marcu 


sandy soil, slope of Pine Mt. at Frakes, HE. L. Braun, no. 1530 
Braun). ROCKCASTLE Co.: wooded gully, « south of Livingston, 
Smith & Hodgdon, no. 3785 (US). OWEN co.: about 1 mile 
north of Scott Co. line, E. L. Braun, no. 3004 (Braun) ; Wedel 
slope, Severn stata E. L. cy no. 4578 (Braun). WAYNE 
co.: Beaver Creek, E. L. Braun, no. 3099 (Braun). JEFFERSON 
co.: 3 miles south of Louisville, July 5, 1892, L. S. Bergmann 
(Mo). LARUE Co.: open cedar 'slope, limestone, Upton, a i 
Braun, no. 1970 (Braun). TENNESSEE: SEVIER CO.: 
Smoky Park, near Gatlinburg, 1932, Mrs. C. D. Walcott oy 
KNOX C0.: Knoxv ille, Ruth, no. 101, as R. strepens. HAMILTON 
co.: old fields, Sequachie Valley, Biltmor Sane no. 8495 (US); 
Chickamauga Park, May 25, Lik. Chur ll (Mo). FRANK- 
LIN co.: dry soil, Cowan, Biltmore Herb., no. at (US); woods 
north of Shernwood, June 1897, H. Eggert (Mo). CHEATHAM 
ner —— bluffs, tae Springs, Svenson, no. 42. Map 16. 
. Var. apenas forma breviberbis, f. nov. (TAB. 
8367, sal 4-6), var. membranaceae simillima, caulis internodiis 
puberulis vel Biabyeiooieisbus rare ee hirsutis. Scattered 
through the general range. MARYLAND: ANNE ARUNDEL CO.: 
mech Ridge, July 13, 1807, °F. H: ence lion (US). West Vir- 
GINIA: CABE o.: ‘dry field in clay soil, near Roland Park, 
Gilbert & Gilbert, no. 259 (VPI). Norra CAROLINA! POLK CO.: 
wet shaded ground, Tryon, D. C. Peattie, no. 1351 (NC). Gror- 
GIA: DEKALB Co.: Stone Mountain, July, 1-8, 1886, Small (TYPE 
in Herb. NY. Bot. Gard.). TENNESSEE: COCKE CO.: near 
Wolf Creek, Kearney, no. 863 (US). 
1lf. Var. MEMBRANACEA, forma hypopsila, , f. nov. (TAB. 867, 
FIG. 1-3), var. membranaceae simillima; cualia: internodiis plerum- 
que pubescentibus; foliis. subtus glabri ris vel subglabris, vix 
strigosis, supra strigosis; — segmentis. ciliatis. —Oceasional 
in Virginia and North Carolina. VrirGINIA: ELIZABETH CITY CO 
Old Point Comfort, Bent: 15, 1895, Britton (NY). NORFOLK 
co.: near Northw est, Kearney, no. 1565 (US). surRy co.: 
border of bottomland woods along Blackwater River, about 1 
mile southwest of Dendron, June 14, 1941, Fernald & Long, no. 
13,148 (TyPE in Herb. Gray.; ISOTYPE in Herb. Phil. Acad.), no. 
13 491 (fruit of preceding). soUTHAMPTON co.: dry sandy pine 
woods by Nottoway River, near Carey Dadi: Fernald & Long, 
no. 13,147; Borge wooded bottomland of Nottoway River: 
ypress Fernald & Long, no. 8470; wooded alluvial 
bottomland of hckoetn River, near Haley’s Bridge, Fernald & 
0. 8471. SUSSEX CO.: bottoml mland — yt: Nottoway 
ices cant of Huske, Fernald & Long, n 761.’ NortH 
AROLINA: ORANGE CO.: meadow below DurkarnGhage Hill 
Fade June 26, 1915, Scher & Totten, transition to next form 


1945, Fernald,—Ruellia in the Eastern United States 79 


lig. Var. MEMBRANACEA, forma: laevior, f:.nov. (TAB. 868), 
var. membranaceae simillima; caulis internodiis glabrescentibus; 
foliis utrinque glabris glabrescentibusve; calycis laciniis eciliatis 
vel vix ciliatis.—Loeal, Virginia and North Carolina. VrirGINia 
SOUTHAMPTON CO.: wooded alluvial bottomland of Meherrin 
River, near Haley’s Bridge, Fernald & Long, no. 9151. GREENS- 
VILLE Co.: bottomland woods along Meherrin River southeast of 
Gaskins, Aug. 3, 1941, Fernald & Long, no. 13,462 (TYPE in 
Herb. Gray.; 1soTypE in Herb. Phil. Acad.). Amexia co.: July 
1, 1936, J. B. Lewis, no. 114 (VPI). Nortu CaroLina: ORANGE 
co.: swamp of New Hope Creek, 5 miles east of Chapel Hill, 
June 30, 1931, H. R. Totten (NC); meadow 3 miles out from 
Chapel Hill, on Raleigh Road, 1931, T. N. Webb (NC). 

11h. Var. nanella, var. nov. (TaB. 869 et 870, FiG. 1), caule 
filiformi ad basin 1-2 mm. diametro 1-3(—4) dm. alto, internodiis 
valde divergenter villoso-hirsutis; foliis membranaceis, ellipticis 
vel oblongis majoribus 2-6 cm. longis; calycis segmentis 1.3-2 
em. longis, ciliatis dorso plerumque glabrescentibus.—Dry to 
moist woods, local, eastern Maryland to Kentucky, south to 
South Carolina; southern Mississippi. MARYLAND: CECIL Co.: 
loamy, wooded slope, Duffy Creek, 2 miles southeast of Cecilton, 
B. Long, no. 48,422 (Phil.); North East, Sept. 2, 1894, Joseph 
Crawford (Phil), transition to var. membranacea. KENT CO.: 
Chestertown, July 29, 1901, HZ. G. Vanatta (Phil), transition to 
var. membranacea. CHARLES co.: dense woods, Tompkinsville, 
Leonard & Killip, no. 838 (US). VIRGINIA: NANSEMOND CO.: 
near Suffolk, Kearney, no. 1718 (US); dry sandy woods above 
Nansemond River, east of Cahoon Pond, northwest of Suffolk 
July 23, 1941, Fernald & Long, no. 13,461 (ryPx in Herb. Gray.; 
ISOTYPE in Herb. Phil. Acad.). PRINCESS ANNE co.: Oceana, 
July 2, 1923, H. B. Meredith, in part (Phil), aoe to var. 


_ membranacea; dry, mixed woods, Little Neck, Fernald & Long, 


no. 4195, transition to var. membranace NortH CAROLINA: 
HALIFAX Co.: Weldon, July, 1875, Mac Elwee (Phil). MCDOWELL 
co.: Old Fort, June, 1872, Walter Faxon. ORANGE CO.: Chapel 
Hill, July, 1931, Vena Millsaps (NC). Sours CAROLINA: 
DARLINGTON co.: damp shady woods near Auburn, June 27, 
1909, Coker (NC). Kenrucxy: poor dry hill on the Kentucky 
River, 1832, C. W. Short (Phil), this material referred to Hooker 
(see note under Kentucky specimens of var. membranacea), with 
the following note: “Is this plant specifically distinct from R. 
strepens in another part of the Collection? Found on a poor dry 
hill on the Ky river. It flowers later than the former & is much 
more starved in its growth.’”’ MISSISSIPPI: JACKSON CO.. Ocean 
Springs, A. B. Seymour, no. 91 8 22, as R. ciliosa. HARRISON 
co.: Biloxi, Lloyd & Tracy, no. 347, in part (NY). Map. 17. 

i. Var. NANELLA, forma eciliata, f. nov. (TAB. 870, FIG. 2-4), 


80 Rhodora [Marcu 


var. nanellae simillima, internodiis glabrescentibus vel puberu- 
lentibus; foliis vix vel minutissime strigosis; calycis laciniis ecilia- 
tis.—Local, southeastern VIRGINIA: SOUTHAMPTON Co.: dry sand 
of open alluvial flat by Blackwater, oe southeast of Unity, 


July 4, 1942, Fernald & Long, n 412 (r TYPE in Herb. Gray.; 
1soTYPE in Herb. Phil. Acad. x aio per of pine and so ae 
at Round Gut, southwest of Franklin, Fernald & L no 


13,154. sussex co.: bushy clearing southeast of Siang ck: 
Fernald & Long, no. 13,145, taller plants, transitional to var. 
membr ee forma laevior. ater SIMPSON co.: Magel, 
. G. Holt, no. 5 (US), as R. hum 
11j. fase icienliekcda. var. nov. (TAB. 871 et 872), caule 
on vix — sparse ramoso 4.5-9 dm. alto basin versus 2-5 
dia o plus minusve hirsuto; foliis subcoriaceis firmis 
duris lancoolati vel lanceolato-ovatis subacuminatis, majoribus 
(supernis) 6-12 cm. longis 2-4.5 cm. latis, su tus ad venas 
a vel ‘ispidis supra strigosis; nodis floriferis 1—4 approxi- 
matis vel imis 3-15 cm. distantibus; glomerulis congestis; 
calycis laciniis dorso glabris vel ee hirsutis crear. corollis 
3-4.5 cm. longis.—Woods and clearings, north-central Sout 
Carolina and northern Georgia, north to southern cae jener, 
northern Maryland and northern Virginia, more rarely to 
Kentucky and southern Indiana. New JERSEY: CAPE MAY CO 
margins of fresh meadows east of Cape May Court House, Long, 
no. 6685 (Phil), appr. var. membranacea; Cape May Court 
House, 1886, Isaac Burk (Pa, Phil); woods, Cold Spring, July 
18, 1908, Van Pelt (P hil). DELAWARE: SUSSEX a eee 
July 8, 1908, Van Pelt (Phil). Maryianp: cEec : North 
East, July 20, 1890, J. B. Brinton (Pa), a as R. ipepeat “hk Neck, 


Mount, Pennell, no. 1591 (P a). MONTGOMERY co.: Little Falls 
Brook, "Pennell, no. 2439 (Phil). a MARYS CO.: Point Lookout, 
July 13, 1930, O. M. Freeman (USNA). District or COLUMBIA: 
Brookland, July 17, 1897, Carrie Harrison (USNA). VrirGInta: 
CLARKE CO.: woods, Boyce, Hunnewell, no. 10,742 (FWH). 
NORTHAMPTON CO.: dry woods, Savage Neck, July 19, 1936, R. 
- Tatnall, no. 3013 levee in Herb. Gray.). GLOUCESTER CO.: 
Beaver Swamp sas north of Gloucester, July 5, 1924, H. B. 
M. a Seah as R. parviflora. ELIZABETH CITY CO.: Hampton, 
July 22 vd: R. Churchill, (Mo), as R. strepens. JAMES CITY 
igs open ce in sre hardwood ravine, southwest of Williams- 
rg, Grimes, no. 4609 (NY); Jamestown, June 24, 1924, 
M. eelith (Duke). PRINCESS ANNE CO : Virginia Beach, July 3, 
1892, Britton, Britton & Vail (NY); pine woods, Virginia Beach, 
Ferna Long, no. oe ; sandy woods, Knott’s ee Fernald 
& Long, no. 4194, in part. NORFOLK co.: Port h, June, 
1840, Rugel (NY). Shenibn co.: low open caiend, thickets 


SD, eS TR 


1945] Fernald,—Ruellia in the Eastern United States 81 


and clearings just east of McKenney, Fernald & Long, no. 14,413. 
MECKLENBURG Co.: roadside-thicket, 6 miles north of Clarks- 
ville, F. R. Fosberg, no. 15,463. HALIFAX co.: old clearing, east 
of Dan River, Fosberg, no. 15,384. Norra CAROLINA: CAMDEN 
co.: low, moist bushy soil, near Shiloh, Correll, no. 2068 (Duke). 
WASHINGTON Co.: moist soil near Scuppermong, Correll, no. 191 
(Duke). GREENE CO.: pine woodland, Farmville, Godfrey, rey 
4278 (Duke, EL lees to var. membranacea. WAKE CO. 
pine woodland along Crabtree Creek, 8 miles Silnek. of 
Raleigh, Giahees no. 4991, in part. HARNETT co.: Buie’s Creek, 
June 29, 1938, Frances & davith Fount (NC). ROWAN CO.: 
Salisbury, June, 1872, LeRoy & Ruger, in part (NY). FORSYTH 
co.: woods, June 20, 1940, Schallert. ALEXANDER CO.: on bank, 


(US). BuNcomBE co.: Biltmore, Biltmore Herb. no. 849>, in part, 
mixed with var. membranacea (US). PouLK co.: Tryon, ‘July 22. 
1897, E. C. Townsend, in part (US). SouTH CAROLINA: FLORENCE 
CO.: ‘sandy, shady banks of Pee Dee River, near Mars Bluff 
Bridge, Wiegand & Manning, no. 2931, as R. parviflora. CHARLES- 
TON CO.: near a 1856, L. R. Gibbes (NY), as R. parvi- 
flora. OCONEE CO.: Clemson College, H. a House, no. 2384 


(US), as R. oat yhh a. GEORGIA: WITHOUT STATED LOCALITY: 
Boykin (NY), as Dzpteracanthus stesidaok: ‘Porter, 1846 (Phil) 
mid. Georgia, 1846,’ Porter, with the comment: “There is & 


good deal of confusion among the Ru elias”. CLARKE CO.: 
Athens, June 28, 1930, J. H. Pyron (Duke). GWINNETT CO.: 
as w River, near McGuire’s Mill, July 2, 1895, Small (NY), 
s R. Police changed by Small to R. parviflora; Thompson’s 
Mills, Allard, no. 225 (NY, US). FLoyp co.: Rome, July, 1888, 
Gerald McCarthy (US). INDIANA: JEFFERSON co.: Hanover, 
July, 1875, A. H. Young (NY). CRAWFORD ae cpg ues 
TUCKY: ROCKCASTLE co.: wooded gully so south ‘ot Livingston, 
Smith & Hodedan, no. 3758 (G), as RB. strepens. 
11k. Var. salicina, var. nov. (TA n, 873); ie abavles ve 
Sparse breviterque ramosi tenui 1.5-7 dm. alto canescenti- 
puberulo vel subvilloso; nodorum su 
anguste lanceolatis vel lanceolato-lineari re 
subtus glabrescentibus vel sparse strigosis; calycis segmen ' 
dorso canescenti-pilosis vel glabrescentibus ciliatis; corollis # 
cm. longis, fauce supra mm. diametro.—Northern Florida 
to eastern Texas, north, locally, to northern Georgia, Tennessee 
and southern Indiana. GEORGIA: pits ETT CO.: oe — 
Mills, Allard, no. 224 (US), as R. parviflora. FLOYD co.: decidu- 
ous woodland, Horseleg Mt., Pesnd no. 4099 (Pa). ee 
GADSDEN Co.: open dry woods of pine and oak, River Junction, 


82 Rhodora [Marcu 


Wiegand & Manning, no. 2933. LAKE co.: dense, moist wood- 
sat Hawkinsville, May 7, 1910, S. C. Hood (Mo). INDIANA: 
CLARK Co.: wet grassy al rocky shore, span es of Charleston, 
F. J. Hermann, no. 6730. TENNESSEE: DAVIDSON CO.: copses, 
vicinity of Nashville, Sept., 1885, Gatiitoe. SHELBY CO.: 
Normal, C. E. be no. 69 (US). ALABAMA: BLOUNT CO.: 
Blount Springs, May 5, 1898, C. F. Baker (NY), as R. peduncu- 
lata. MON co.: Perdue Hill, July 22, 1885, C. Mohr (US), as 
R. iisaais, 3 ie hybrida, this altered to var. ambigua, then the 
sheet, in spite of petioles 1.5 cm. long, annotated as the sessile- 
leaved R. humilis! MOBILE CO.: pine see and dry pastures, 
Spring Hill, H. W. Ganes, no. 948, in part (US)—the sheet also 
containing var. semicalva and R. ciliosa. MussisstPPt: CLARKE 
co.: Enterprise, S. M. Tracy, no. 3288 (NY), as R. ciliosa, 
altered é i. yes rviflora. Lae ANA: WITHOUT STATED LOCALITY: 
Steinhau RLEANS PARISH: New Orleans, Drummond, no. 257, 
as Divlaracantivus aia another, with no., as Calophanes 
humistrata. NATCHITOCHES PARISH: open groun nd, Natchitoches, 
June 10, 1915, FE. J. Palmer, no. 7945 (7 tyPE in Herb. N. Y. Bot. 
Gard.; isoryPEs in Herb. Mo. Bot. Gard. and U. 8. Nat. een 
RAPIDES PARISH: pine-barren hills, vicinity of Alexandria, C. R 
Ball, nos. 559 (US), as R. strepens, and 655, in part (Mo)—the 
atter mixed on a sheet with R. humilis var. frondosa and quite 
like the other sheets of no. 559 (presumably the former mixed in 
handling). IBERIA PARISH: moist open grassy woods, Avery 
Island, H. B. Correll, no. 9525. TExas: UPSHUR CO.: 
sandy ‘woods, Big Sandy, Reverchon, no. 2536 (Mo); damp sandy 
no = andy, Reverchon, no. 1402 (simulating small var. 
dentata). GREGG Co.: ric sandy woods, Gladwater, June 18, 
everchon (Mo). woop co.: damp woods, Mineola, Rever- 


sheets pap no., G), as R. strepens. map 19. 

11]. Var. dentata (Nees), comb. nov. Stem stiff and rela- 
tively strong, simple or more often with ascending branches, 
- main axis 1.5-9 dm. long, hirsute to puberulent; the lower 

median internodes shan eatin 4-9 nodes bearing glo merules; 
paves oblong, oblong-lanceolate, ovate or elliptic, subcoriaceous, 
—_ undulate-dentate, blunt or acutish, the lower and mi 
nes subequal, the upper reduced and definitely smaller; she 
occasionally -leistogamous; calyx-segments ciliate, 1.8-2.5 

ong.—Dipt acanthus ciliosus, }. dentatus Nees in De. Prove ia xi. 
123 (34) —Chi efly in dry’ woods and clearings, upland of 
South Carolina, western North Carolina and eastern Tennessee, 
north to Delaware, southeastern samp aeggonteg Maryland, 


1945] Fernald,—Ruellia in the Eastern United States 83 


July 11, 1895, Crawford (Phil). DELAWARE: NEW CASTLE 
north end of cedar scrub, near Taylor’s Bridge, July 28, 1801, 
Commons. SUSSEX CO.: oak copse, Rehoboth, Sept. 5, 1908, 
J. R. Churchill. MaryLAND: CECIL Co.: Conowingo, July 
1907, J. J. Carter (Phil). BALTIMORE Co.: near Baltimore, 7966 
P.V.LeR oy (NY). CALVERT co.: dry woods, Chesapeake Beach, 
Hunnewel ep 5666 (FWH). wicomico co.: Salisbury, July 14, 
1904, J. J. Carter (Phil). District or Co.tumsia: June 22, 
1902, Steele (Duke). West VIRGINIA: NICHOLAS co.: dry bank, 
Peter Creek, Millspaugh, no. 596 (NY), as R. ciliosa. VIRGINIA: 
FAIRFAX CO.: woods, Great Falls, Hunnewell, no. 7027 (FWH). 
JAMES CITY CO.: dry slope about 3 miles north of Williamsburg, 
. Menzel, no. 30, as R. strepens. HENRICO co.: Fairwa 
Ridge, near Richmond, July 10, 1928, F. H.W. PRINCESS ANNE 
: pine Senge Virgini ia Beach, Sept. 25 and 28, 1900, Wm. 
Pigces (US); Oceana, July 2, 1923, if. B. Meredith, in part 
(Phil); open clay at ’ border of woods, east. of Little Creek, 
Fernald & Long, no. 4192. NORFOLK co.: dry sandy roadside, 
Ocean View, July 3, 1923, H. B. M eredith (Phil). NANSEMOND 


of Zuni, Fernald & ipl no. 13, 460. SUSSEX CO.: ne sandy 
woods, thickets and clearings, north of Moore’s Mill, Fernald & 
Long, no. 6397; dry woods near Nottoway River, Green Church 
Bridge, northwest of Owen’s Store, Fernald & Long, no. 14,020. 
HALIFAX Co.: old clearing, east of Dan River, 12 miles east of 
Danville, F. R. Fosberg, no. 15,384 (Pa). ORANGE CO.: field, 
Orange, Killip, no. 13,248 (US). NortH CAROLINA: ORANGE 
co.: Arboretum and Campus of Univ. N. C., July 1, 1914, Coker 
(NC). PoLK co.: banks and meadows, “Valhalla”, Tryon, D. 
Peattie, no. 626 (NC). Mapison co.: Warm Spring, Aug. 23, 
1875, J. H. Redfield (Mo). rafeb ont ae rose B. L. 
Robinson, no. 59, as R. strepens; Biltm Bi 

849 US). ROWAN Co.: vicinity of Heilig’ s 5 Mill, Small & Heller, 
no. 139 (Phil, US). sacKSON CO.: near Dillsboro, Sept. 9, 1933, 


and along railroad, west of New Albany, Deam, no. 14,010. 
“S Braun, no. 


E. 
3132 (Braun). pPpuLAsKI co.: thin soil over limestone, south of 


L. 
Bowling Green, July 3, Sadie 
BALLARD CO.: os ous. Wickliffe, McFarland & Anderson, no. 


84 Rhodora [Marcu 


(Mo). TENNESSEE: BLOUNT co.: damp woods 2 miles east of 
Townsend, W. M. Benner, no. 5803 (Phil). Pxuares 874 and 
875; MAP 20. 


Of all the species of Ruellia in the eastern United States R. 
caroliniensis is the most baffling, for in different areas it is 
reasonably definable as geographic varieties, but, at least in 
eastern Virginia, plants with strongly spreading-hirsute stems 
and others with the internodes merely puberulent or short- 
pilose will occur side-by-side, while in other colonies may be 
found individuals with the leaves strongly hispid on the veins 
beneath almost mingled with individuals with the lower leaf- 
surfaces glabrous. In general the more northern series (vars. 
membranacea, nanella, cheloniformis and dentata) have the soft 
villosity or pilosity of stem or leaves less marked than in the more 
southern vars. typica, semicalva and salicina. In the three latter, 
furthermore, the leaf-blade is rather ‘‘full’”’, so that the pressed 
foliage commonly puckers or has a “‘tuck’”’ and the calyx-segments 
are commonly canescent on the back. In the more northern 
varieties, on the contrary, the leaf-blade is readily flattened in 
pressing and the calyx-segments are glabrous or only excep- 
tionally pubescent on the back. When well developed the stems 
of the southern vars. typica and semicalva tend to divergent 
branching, with the elongate branches floriferous at tip, but 
simple-stemmed plants somewhat obscure this character. In the 
northern series the branching, except sometimes (as in var. 
dentata) from the base, is weak, most plants having simple stems 
or few short and rarely floriferous branches. 

The commoner northern plants fall into the thin-leaved var. 
membranacea (with several minor forms based upon development 
or lack of development of trichomes on the leaves or of cilia on 
the calyx-segments), which passes insensibly, on the one hand, 
into the stouter, stiffer and firmer-leaved var. cheloniformis, on 
the other into the dwarf, very slender and small-leaved var. 
nanella, the latter, or something very like it, reappearing in 
southern Mississippi, where it passes into weak unbranched 
states of the sduthern var. semicalva. In the extreme South, 
furthermore, the narrow-leaved var. salicina, very definite in its 
extreme development, merges into var. semicalva and strongly 
suggests the narrowest-leaved extreme of the usually more 


1945] Fernald,—Ruellia in the Eastern United States 85 


northern var. cheloniformis. Var. dentata, with great reduction 
of leaves above, with the lower and median leaves nearly uni- 
form, with strong tendency to develop elongate and floriferous 
basal branches, and its great abundance of glomerules, is as 
definite as any of the varieties, but some specimens show evident 
transition to vars. salicina, semicalva, cheloniformis and mem- 
branacea. In short, the species, like the usually more western 
R. humilis, is evidently in a state of flux and I am unable to find 
in it really stable characters. 

The identifications of the past clearly show how perplexing 
have been the many variations of this most complex species, 
especially to those without any field-experience with it. It has 
been variously known or identified as R. strepens L. (our species 
2, PLATES 841 and 842), or R. pedunculata Torr. (our species 3, 
PLATE 843), mixed in collections with R. Purshzana (our species 
5, PLATES 845 and 846), identified as R. ciliosa Pursh (our species 
7, PLATES 849 and 850) or as R. humilis Nutt. (our species 10, 
PLATES 854-860). 

It was described and illustrated by Dillenius (our PLATE 863) 
and his description and plate were included in the R. strepens 
of Linnaeus (1753), but later excluded by him. In fact, Elliott, 
apparently not cognizant of the redefinition of R. strepens by 
Linnaeus (1771), was puzzled by the restriction of the latter 
name by authors who had Mantissa Altera. Describing in 
detail Ruellia caroliniensis, var. typica, as R. strepens, “the 
whole plant hairy . . . Calyx [-segments] . - - linear- 
lanceolate, the upper half almost setaceous, very hispid”, and 
correctly citing for his South Carolina plant the Dillenian plate 
and Anonymos caroliniensis Walter, 168, Elliott thus expressed 
his perplexity: “I know not how Pursh could have called [cor- 
rectly following the emended description of Linnaeus in 1771] 
the segments of the calyx lanceolate, they are very accurately 
represented by Dill. Hort. Elth. T. 249, F. 321, excepting that 
in number 5 and 6 the setaceous points are not sufficiently 
extended, but in number 1 from which he derived the epithet 
Comosa, the representation is very accurate.” one 

Although the Walter type of his Anonymos caroliniensis, p- 
168, basis of Ruellia caroliniensis, is lost, his generic description 
was good and his specific description well applies to the southern 


86 Rhodora [Marcu 


plant, which occurs abundantly in eastern South Carolina and 
about Savannah (see Map 14). Since the other strongly pubes- 
cent (“hirsute”) varieties (vars. membranacea, cheloniformis and 
dentata) barely reach the northern half of South Carolina from 
the north (see maps 16, 18 and 20) we are quite safe in identifying 
as Walter’s plant the variety which abounds in the region he 
best knew. R. caroliniensis (var. typica) was next described by 
Pursh as R. hybrida (1814), evidently from the strongly branch- 
ing state: “R. erecta, ramosissima, pilis albidis hirsuta; foliis 
; dense hirsutis, . . . calycis laciniis linearibus tubo 
corollae vix brevioribus. In sandy fields near Savannah, Geor- 
gia’”’, where our plant abounds. If it were not for the earlier 
name of Walter the correct name of this species would be R. 
hybrida Pursh, the name used for the canescent-villous extreme 
by Small and others. Nees, who sadly mixed the identities of 
our American species and varieties, again beautifully described 
typical R. caroliniensis as Dipteracanthus Mitchillianus (1847), 
emphasizing the stem “pube alb& densé incano pilisque patulis 
hirsutis”, the oval-oblong to oblong and obtusish leaves decur- 
rent into petioles, the blades ‘‘utrinque laxe hirsutis’’, the linear- 
setaceous calyx-segments ‘‘albo-hirsutissimis’”’ and about half as 
long as the corolla-tube. That was a first-rate description. 
Unfortunately, in the same treatment Nees published the ill+ 
described D. ciliosus, ‘‘y. parviflorus, corollA vix pollicari, foliis 
paullo longiori petiolo (3—6-pollicari)”, from Kentucky. As 
already sufficiently emphasized, a plant with petioles 3-6 inches 
long is quite impossible in Ruellia (at least in ours). Neverthe- 
less, the impossible description by Nees has been made the basis 
for a specific combination, R. parviflora (Nees) Britton, and for 
the varietal combination, R. caroliniensis, var. parviflora (Nees) 
Blake; and in recent years these ill-founded names have largely 
appeared on the labels of most of the varieties and forms of R. 
caroliniensis. 

The confusion in the use of names is here perhaps somewhat 
cleared. The presentation of the varieties and forms may later 
need revision. After some years of puzzling over the misbehavior 
of these inconstant trends, I have done the best I can with them. 
Future and wiser students may work out a better treatment. 


OL ee eee lee ee ee ee 


1945] . Fernald,—Ruellia in the Eastern United States 87 


EXPLANATION OF PLATES 


PLATE 839. RUELLIA BRITTONIANA Leonard: Fic. 1, summit of ce x1, 
from Hours, Louisiana, Wurz FIG. 2, summit of peduncle and calyx, X 4, 
from Wurzlow; FIG. 3, corolla, 1, from W urzlow; Fic. 4, calyx and capsule, 

rom Wurzlow 
LATE 840. ‘ EEDIANA Griseb.: Fics. 1 and 2, portions of branching 
plant, < 1, from Colonia Benitez, “ice oe Argentina, A. G. Schulz, no. 778; 


one basal bract removed, x 1, from near Sta e, — ay om ee 
Shacklette, ae 345; Fic. 3, flower, showing broad calyx-se 
no. 345; FIG. 4, long-peduneled fruit, < 1, from Powell’s , ay Gastivite, 
Virginia, Fernald & nih 2 

842. Ns, forma cLEISTANTHA (Gray) S. McCoy: tae be 
summit of characteristic plank; x 36, from se uth Lebanon, Ohio, EZ. B. 
no. 8010; Fic. 2, a terminal glomerule, X 1, from Mammoth Cave, Kentucky, 
E. L. Braun, no. 3611; | pot 3, node with a cleistogamous flower, < om 
Deerfield, Vernon Co., Missouri, Palmer & Steyermark, no. 42, 140; ri. 4, 
terminal pier ® with 2 (upper) small cleistogamous flowers, xX 2, from 
Joplin, Missouri, E. J. Palmer, no. ; Fic. 5, subterminal fruiting node, 


from Natchitoches, Louisiana, E. J. Palmer, no. 7511; Fic. 2, 
branching fruiting plant, x } , ; ; 
FIG. 3, portion of “branching inflorescence, X 1, from near Arcadia, Missouri, 
Greenman, no. 3764; Fic. 4, calyx and capsule, X 2, from Baker Springs, 
Howard Co., Arkansas, October 5, 1909, J. H. Kellogg; tee 5, calyx-segments 
and base of capsule, x 10, fr “an last specimen; FIG. 6, strigose surface of cap- 


R. PINETORUM Focuald: ric. 1, flowering and fruiting branches, 

, from TYPE; FIG. 2, portion of fruiting branch, X 1, fro ; Fic. 3, 
portion of leafy base, x 1, from Lake Charis. Jom Andrew Allison, no. 
; FIG. 5, surface of peduncle, 


S  PURSEIANA Fernald: rie. 1, md md of TYPE, X 


third node f base, with solitary flower, x 1, from Cedar Cree k, ederick 
0., Vi ae nnewell, no 11-135; Fic. 3, second or’ from an 


‘ u . 
? , Virginia, May 
solitary flower, X 1, from Natural Bridge on Dixie Caverns, Roanoke Co., 


. . 2, fro: 
Virginia, C. E. Wood, Jr., no. 3673; FIG. > slate of stem, X 4, from Wood, 
no. 3673. 


PLATE 846, Fics. 1 and.2. R. PURSHIANA, forma CLAUSTROFLORA Fernald: 
Fic. 1, summit of TY tbl xX 1; Fie. 2, uppermost ae with tiny cleistogamous 


A fro: ridge, V 
er ore on 7, Pyle, N Boman oma 
ca, rr rface, X 10, from Wood, no. ? 

ae shoving pase Ree ROMORPHA Fernald. PxuatTe 847, VERNAL 


: FIG. x m near Camp ens ascot do t of 
Small & Wilson, no. 1700; FIGs. 2 ‘and 3, flowering tips, pucean of in eeeande 


Srl Gare no; RT Tg! glas.” Paar 88, TATER cunTOGEY 
onroe Co., Florida, Smali, no ‘ 

STAGE: FIG. 1 aelion of a long fruitin soar —< 4: ee betwee n Peter's 
Prairie and Homestead, Florida, Sma & Carter, no. 2701; FI 


88 Rhodora [Marcu 


branchlets from more open fruiting branch, X 4, itis Ngee ors Coconut 
Grove and Cutler, Florida, Small & Carter, no. 1273; ria. 4, tip of — 
with two cleistogamous flowers, < 1, from Ross-Costello amanee , Dade 

Florida, Small, Mosier & Small, no. 6552; ric. 5, calyces and capsules, x 3 


LATES 849 and 850. R. crzi0sa Pursh: PLATE 849, Fic. 1, plant, 1, from 
Cape Savannah, Sumter Co., South Carolina, Witmer Stone, no. 426; Fics. 2 
and 8, plants, rom m Poulan n, Worth County, Georgia, Pollard & Maxon 


Florida, A. J. Pie eters, no. eee Fic. 3, basal branch, X 1, from Duval Co., 
Florida, A. H. Curtiss, no. 194 


from same collection as fig. 2; ric. 4, lower rane . leaf, oe cystoliths, 
xX 10, from Bay Biscayne, Honda, oz . de rtis 0. 5500E; as. 5 and 6, 
median aiid terminal flowering nodes, ries wc s a Hobe as a. te 
Fic. 7, calyx and fruit, < 2, from east or "plovida City, Dade Co., Ronin: 


‘8080. 

Puates 852 and 853. R. Noctirtora (Nees) Gray: PLate 852, ric. 1, base 
of plant, X 1, — southeast of Ludowici, Georgia, tegand & Mann ning, no. 
2928; FIG. 2, ring tip, < 1, from no. 2928 - ; Fia. 3, sonar mit, X 1, of IsoTYPE, 
from inter St. huis t Port Leon, Florida, Rugel; ¥ a. 4, flowering tip, X 1, 
from Apalachicola, Florida, 1867, Chapman; Fia. 5, gpm of internode and 
bases of leaves, < 4, fro m Apalachicola, Chasse ; FiG. 6, calyx and capsule, 
x 2, from iloxi, Mississippi, . M. Tracy, no. 6500. LATE 853, Fic. l, 
summit of flowering stem to show lon ng calyx-segments, x 1, 
bury, Liberty Co., Georgia, LeConte Te foe nae Arh or isotype of R. flora 

0 te); FIG. summit, X 1.8 t. Joe to Apalachicola, 
Florida, Small, Small and De Winkeler, no. ii “417: FIG. os bract, calyx-segment 


gmen 
and base of capsule, x 10, from Tracy, no . 6500; FIG. ’4, surface of capsule, 
x 10, from no. 

PLATES ei bag 855. R. HUMMUS Nutt., var. Typrca: Pate 854, ric. 1, 
TYPE Or ISOTYPE, X 1; FIG. 2, summit of primary axis, < 1, from Mercersburg, 
Pennsylvania, 1846, Ruel; Fria. 3, calyces ~ capsules, < 2, from Butler Co., 

ri, Bush, n 0. 3739. PLate 855, Fic. 1, median nodes of primary axis, 


x 'L 7 ren west of 3 Hannibal, Missouri Tal ohn Davis, no. 4646; FIG. 2, a ee : 
internode and bases of leaves, X 4, tea m Cooper Count; Missouri 
15,137; ric. 3, portion of fruiting branch, showing die a % 1. aes 


vicinity ee Nas ashville, Tennessee, Thsitenaber, 1885, G 

Puate 856. R. nurs, forma GRISEA Fern ss portion of TYPE, 
ining Gavan habit of the species, X 1; FIG. ‘3 eiatoeas and bases of 
leaves, X 4, from TYPE. 
- Pyare 857. Roa HUMILIS, var. FRONDOSA Fernald: ria. 1, Trypn, X 14; FIG. 2, 
summit of internode and bases of leaves, X 4, from Cleveland, Ohio, 1805, 
Starr; , flowering node, X 1, from Milfor i, Ohio, June 17, 1916, E. L. 
Braun; ria. 4, flowering a X_1, from Cham Pennsylvania 


,, 1897, T. C. Porter; ric. 5, flowering node, x 1, a m Corydon, Indiana, 
riesner, no. 14,369; FIG. “ fruiting node, x i from Clinton. m 
no. 77; FIG. 7, ¢ calyx and capsule, X 2, from Champaign, Illinois, A. "8. Pease 

no. 12, 420. 


ATE 858. R. HUMILIS, 
flowering branches, X 1, from Piasa, Illinois, 1905, G. EF. McClu ure; FIG. 2, 
summit of flowering ay <_1, from Wichita, Kansas, 1892, H. R. Peas 
FIG. 3, flowering summit, , x 1, fro m Huntsville, Texas, R. A. Biron, no. 377; 


EI TT aL LL OTL LON LE 


_ lower surfaces of leaves, x 10, from TYPE. | Fics. ay Creek, 


1945] Fernald,—Ruellia in the Eastern United States 89 


Fic. 4, summit of internode and bases of leaves, X 4, from Willis, 
L. . Warner; FIG. 5, bracts, calyces and fruits, X oF from north of rb 
Texas, Lundell, no. 11 ,679. 

PLATE 85 me var. EXPANSA Fernald: Fic. 1, flowering node, 
from peste < 1; rie. 2, median fruiting node of primary axis, X 1, from 
Oquawka, Illinois, pirat Fic. 3, internode and leaf-base, X 4, from 
McDonald Co., Missouri, Bush, n 0. 282: ge 4, calyx and capsule, x 2, from 
Hendrix, Illinois, August, 1904, 5 si 

PLATE ae R. nuM ILIS, var. gst Fernald: Fic. 1, portion of Iso- 
TYPE, X 1; Fia. 2, flowering tip, x 1, fro gaa paces Kentue ky, Biltmore 
Herb., no. ‘480K: FIG. 3, internode and leaf-bases, X 4, from near Manchester, 
Tennessee, Biltmore Herb., no. 849°; ric. 4, lower surface ‘of leaf, X 10, from 
Middletown, steal Co., Virginia, Hunnewell, no. 17,561; Fic. 5, fruiting 

ode no. 17, 561; FIG. 6, calyx and capsule, x 2, from ‘north of 
Leavenworth, Crrwiood Co., Indiana, Deam, no. 

PLATES 861, 862 and 8 63. R. sac ainaid orn (Walt.) $ Steud., var. TYPICA. 
Puate 861, Fic. 1, portion of cape: plant, X 1, from Ormond, Florida, 
H.A. Purdie; FIG. 2, dewering tip, X 1, from Brookgreen Carder Georgetown 
Co., South Carolina, F. H. Tarbox, n — 567; Fic. 3, summit of internode, X 4, 
from no. 567; Fic. 4, lower surface of leaf, x 10, from Green Cove Springs, 


ped, 
Loshon Co., Florida, E. P. Walker, no. 1862; ti a calyx se capsule, X 2, 
from east of Eustis, Florida, Small, no. 8667; 3, portion of calyx and 
capsule, X 10, from no. 8667. PLATE 863: copy of “the plate, slightly reduced, 
of R. strepens, capitulis comosis of Dillenius. 
Puate 864. R. CAROLINIENSIS, var. SEMICALVA Fernald: Fic. 1, TYPE 
3/7; eae 2, — n of internode, X 10, from TyPE; FIG. 3, lower surface of 
leaf, 0, from TYPE; FIG. 4, upper surface of leaf, X 10, from TYPE; FIG. 5, 
calyx Rak capsule, X 2, from Iola, caer | aise, 1896, Chapman; Fic. 6, 
calyx-segment and portion of capsule, X 10, ast, 

LATES 865 and 866. R. porate ese pring MEMBRANACEA Fernald: 
PuaTE 865, Fic. 1, Type, X 14; FIG. 2, upper surface of leaf, < 10, from Savage 
Neck, Northam mpton Co., Virginia, Fernald & Long, no. 5453; FIG. 3, — 
surface, x 10, from no. 5453; Fig. 4, flowering tip, X 1.7, from below Rush- 
mere, Isle of Wight Co., Virginia, foemai & ae Forbesagc a ery 5, ghee 

Fia. 6, frui 
of oe and papas x 10, ier Lopes nee cg 866, Summits HH 
show variations of aves, xX 1: Fie. 1, from Little Creek, Prine eee eae 
Virginia, Fernald & Long, no. 4193; Fi. 2, from Congress Heights, srs ric be 
Sag » ee eee no. 7012; ak 3, from Cold Sonne, Cape 


on. 
form 

pial “887, ak: ey R. CAROLINIENSIS, Var. MEMBRANACEA, 

HYPOPSILA Fernald: F Fic. 1, summit 1.7, of rypxE; Fics. 2 and 3, pile and 


Fernald: ria. 4, internode and leaves, X 1.7, from Wol Tenaane 
Kearney, no. 863; Fias. 5 and 6, upper and lower leaf-surfaces, X ‘10, from no. 


: Fernald: 
a E 868. R. CAROLINIENSIS, var. MEMBRANACEA forma LAEVIOR 
G.1 fruiting summit, X 1.7, of TYPE; Ape 2 and 3, upper oo niche “age 10 
ot leaf, X< 10, from TyPE; FIG. 4, portion of calyx-segment and ca : 
rom TYPE 
PLate ‘869. R. CAROLINIENSIS, var. ai feat, 10, shat ne yes ry 
Figs. 2 and 3, upper and lower surfaces of le *< 10, from TYPE; FIG. ‘ 
portion of calyx-segment and sa Ri x i irom TYPE. d base of 
TE 870, Fic. 1. R. CAROL : internode x Fernald: 


90 Rhodora [Marcu 


ric. 2, TYPE, X 1; FIG. 3, portion of leaf, bracts and internode, ee ha from 
TYPE; FIG. 4, caiiion ‘of calyx-segment and capsule, X 10, from 
Puares 871 and 872. R. CAROLINIENSIS, var. CHELONIF ORMIS. oaeeunld: 


1, fr 

upper ses ret leaf-surfaces, X 10, from Fernald & Long, no. 

gpa R. CAROLINIENSIS, var. SALICINA Fernald: Fie. 1, TYPE, xX 4/7; 
FIG. 2, whan ae of plant, X 1, from Louisiana, Steinhaur; FIG. 3, plant, x 1, 
from River Junction, Gadsden Co., Florida, Wiegand & Manning, no. 2933. 

TES and 87 ; CAROLINIENSIS, var. DENTATA (Nees) Fernald: 

PLATE 874, Fic. 1, small plant, Res from Broadwater si aed of ee 
Co. , Virginia, Fernald & Long, no. 13,460; Fic. 2, summit ne 
from no. 13,460; . 3, portion of fruiting ‘ahs x &, froin “Teloboth 
Delaware, Churchill. PLATE 875, Fic. 1, upper fifth of main axis, X 1, 
Marsh’s Millpond, Nansemond Co., Virginia, Fernald & Long, no. 14, 414: 
Fic. 2, summit of branch, X 1, from Sweeden, Edmonson Co., Kentucky, 

L. Braun, no. 3593. 


INDEX 


New scientific names are printed in full-face type 


Anonymos caroliniensis 31, 71, 77, 85 
Blechum, 11; Tweedyi, 11 


Calophanes humistrata, 82 
Cryphiacanthus acaulis, 10, 11; an- 


gustifolius, 8-12; angustifolius 
edianus, 9, 10; barbadensis, 


Dipteracanthus, 32; ciliosus, 32, 34 


36, 38, 50, 52, 74, 8. dentatus, 82, 
8. hybridus, 71, y. parviflorus, 2, 


Drummondii. 56, 57; hybridus, 81; 
micranthus, 17-19; Mitchillianus, 

, 72, 86; noctiflorus, 49, 50, 8. 
humilis, 51; spectabilis, 11; strep- 
ens, 14, 52, 58, 60, 82, var. calyci- 

us 4, y. Dillenii, 71, var. 
pedunculatus, 14, 17, var. strictus, 
14, 17; strictus, 14 


Dizygandra noctiflora, 49; tubiflora, 
Dyschoriste oblongifolia, 60, 63 
Hygrophila illinoiensis, 17, 18 
Pattersonia caroliniensis, 71 


Ruellia acaulis, 10; angustifolia, 8, 10, 


; biflora, ; Brittoniana, 5, 
8 ‘ 
’ 


(91) 


89, pl. 864, f. detonsa, 69, 74, var. 
typica, 25, 69, 71, 73, 84-86, 89, 
pls. 861-863; ciliosa, 3, 6, 238, 25, 
28, 29, 31-38, 47, 51-54, 56, 58-62, 
s. 849, 


ie : 
28-30, 72, 74, 82, var. cineras- 
cens, 48, var. humilis, 51, 53, var. 
hybrida, 27, 29, 52, 71, 72, 82, var. 
longiflora, 36, 51, 56-60, var. 


858, var. typica, 25, 51, 54, 88, 
pls. 854, 855, f. a, 54, f. grisea, 
51, 54, 88, pl. 856; hybrida, 2, 28, 
31-34, 36, 71, 72, 77, 86; longiflora, 
26, 49; malacosperma, 13; 
Morongii, 10, 1 


(eS) 
— 
Ww 
Sg 
ow 
- 
Ww 
= 
Wf 
a] 
a 
jor) 
(—) 
or] 
ine 
~J 
i 
~] 
A 


3 : 
flora, 49, 50, 88; Tweediana, 8-13, 
87, pl. 840; Tweedyi, 11; vincae- 


Rhodora 


Plate 839 


Photo. B. G. Schubert. 


acne BrrrroniaNna: FIG. 1, summit of plant, X 1; FIG. 
Fig. 3, corolla, X 1; FIG. 4, calyx and capsule, x 2. 


2, summit of peduncle and calyx, 


Rhodora Plate 840 


Photo, B. G. Schubert, 


tUELLIA TWEEDIANA: Figs. 1 and 2 


i , portions of branching plant, x 1; Frc. 3, summit of 
peduncle and base of calyx, x 4; FIG. ¢ 1 


4, tip of calyx-segment, < 10; ria. 5, corolla, X 1. 


Rhodora Plate 841 


Photo. B. G. Schubert. 


RUELLIA STREPE meg ric. 1, flowering median node, X 1; FIG. 2, ocep guar avon aka 
removed, < 1; Fic. 3, flower, showing broad caly ‘<-segniont, x "1G ong-pe 
fruit, x 1, 


Rhodora Plate 842 


Photo, B. G. Schubert. 


UELLIA a carci forma CLEISTANTHA: FIG. 1, summit of plant, < 24; Fria. 2, terminal 
glomerule, X 1; a3. ode with large cleistogamous flower, X 2; Fia. 4, terminal glomerule 
with two small setcpatatas flowers, X 2; Fia. 5, subterminal fruiting node, X 2; FIG. 6, 
calyx and open capsule, 


Rhodora Plate 843 


Photo. B. G. Schubert. 


RvUELLIA PEDUNCULATA: FIG, 1, small flowering plant, x 4/9; FIG. 2, aga 2 more branch- 
ing whine plant, x 14: FIG. 3, portion of branchi ng inflorescence, SK 4, ealyx and 
Capsule, x 2; FIG. 5, ih Etepee Leh and base of capsule, X 10; FIG. 6, Bai nirtaee of 
Capsule, x 10. 


Rhodora Plate 844 


fas 


a sm. 
me gee 


ee 


Thro 


oo 


a. hs 
ifr, 


Photo. B. G. Schubert, 
PINETORUM ge 1, flowering and fruiting — _ TYPE, X 1; FIG. 2, por- 
tion of fruiting bran na x1 - FIG. 3, portion of leafy base, x 1; calyx and 1 eapsule, Xx 2; 
Fria. 5, surface of peduncle, x 10; ria. 6, bases of malysaspiient na capsule, X 


Rhodora Plate 845 


Photo, B. G. Schubert. 


Ruewxia Purswiana: FiG. 1, portions of TYPE, X 7%; FIG. 2, third node from base, X 1; 
FIG, 3, § eta node from base, ' 1; FIG. 4, calyx and oa % 2: ria. 5, surface of a 
x 4. 


Rhodora Plate 846 


Photo, B. G. Schubert. 


RUELLIA ee - 3, summit of capsule, showing pilose surface, x 10. d 
R. Purs A, fo AUSTROF : Fic. 1, summit of type, <1; F1G. 2, uppermost node, 
with siualt sleiatciaceduns flowers (above) and capsule, x 2 


. 


Rhodora Plate 847 


Photo, B. G. Schubert. 


RUELLIA HETEROMORPHA, vernal stage: FIGS. 1-3, portions of flowering stems, X 1; FIG. 
4, summit of internode and base of leaf, X 4; F1a. 5, base of calyx, X 10. 


Rhodora Plate 848 


be 


Photo, B. G. Schubert. 


is cori sit EB S branche later SF pret stage: Fic. 1, portion of long fruiting branch, 
x 1; FIGs. d 3, branchlets, x } , tip of branchlet, with two cleistogamous flowers, 
x 1; FIG. 5, calyces and capsules, 9 


Rhodora Plate 849 


Photo. B. G. Schubert. 


Rvexta cruosa: FIGs. 1-3, plants, X 1; Fra. 4, summit of internode and base of cauline 
leaf, x 4; ria. 5, calyx and capsule, X 2. 


Rhodora Plate 850 


Photo. B. G. Schubert. 


R ee 
beat cILIosA: Fig. 1, plant, X 1; ria. 2, basal branch, X 1; Fic. 3, large-flowered 


Rhodora Plate 851 


24- pig. 2, fruiting branch, 
of leaves, : FIG. 4, lower surface of leaf, 
is. 5 and 6, median and terminal "hewetne nodes, X 1; 


RUELLIA SUCCULENTA: FIG. 1, portion of flow ering plant, x 3 
leave: 


X 1; Fie. 3, s 
ene cystoliths, ; FIG 
G. 7, calyx and fruit, X 2 


Rhodora 


Plate 852 


oan 
rrr 


Photo. B. G. Schubert. 


ae LLIA NOCTIFLORA: FIG. 1, base of plant, ; Figs. 2, 3 and 4, A? tips, 
G. 3, ISOTYPE); FIG. 5, summit of internode pa bases of leaves, xX 4; 
se sule, X 2 


x1 
1G. 6, calyx and 


Rhodora Plate 853 


* 
PN bed 


c 


* 


Ss 
+ 


fe 


Photo. B. G. Schubert. 


ELLIA NOC apowues 1, summit of flowering stem of probable TYPE - 
R, te 1; FIG. 2, fruiting summit, X 1.8; FIG. 3, bract, calyx-segment an 
base of capsule, NS 10; ric. 4, surface of capsule, pd 10 


Rhodora Plate 854 


Photo. B. G. Schubert. 


RUELLIA beep var. TYPICA: FIG. 1, TYPE Or ISOTYPE, X 1; FIG. 2, summit of primary 
axis, X 1; FIG. 3, calyces and capsules, x 2 


Rhodora Plate 855 


Photo. B. G. Schubert. 


i 
HLLIA HUMILIS, var. TYPICA: FIG. 1, median nodes of primary axis. < 15 . 2, summit 


of C... node and bases of leaves, X 4; FIG. 3, portion of fruiting branch, eine ‘diffuse habit, 
X1 


Plate 856 


~ X ‘SVAvET JO 


8 


as 


Bq PUB BpOUIE4Ul ‘Z "NIA tT Xx ‘ 


ad 


AL 


jo uory 


od ‘] ‘pla 


Vv 


" 
1G 


ISIMD VULIO} ‘SITUNOH VITIAOYY 


19QnyIg “YH “_ ‘olwyd 


Rhodora 


Rhodora Plate 857 


Photo, B. G. Schubert. 


Rue aye var. igo eae FIG. 1, portion of TYPE, X 7 G. 2, ee os st 
node and bases of leav X 4; Fias. 3-5, ’ flowering nodes, u FIG. 6 ruiting n 
oy Ag 


FIG. 7, calbx ate Ci seat 


Rhodora Plate 858 


Photo. B. G. Schubert. 


RUELLIA HUMILIS, var. LONGIFLORA: FIGs. 1-3, ee of flowering branches, 1 
FIG. 4, summit of iitersodes and bases of leaves, * "4: . 5, bracts, calyces and trisite, x < 


Plate 859 


Rhodora 


Photo, B, G. Schubert. 
RUELLIA HUMILIS, var. EXPANSA: FIG. I, flowering node, X 1, from tye eal 2, ee 
fruiting node, X 1; FIG. 3, internode and leaf-base, X 4; FIG. 4, calyx and capsule, x 


Rhodora Plate 860 


Photo. B. G. Schubert. 


RUELLIA HUMILIS, var. CALVESCENS: FIG. 1, portion of isorypr, X 1; FIG. 2, flowering tip, 
X 1; m1. cy internode and leaf-bases, X 4; FIG. 4, lower surface of leaf, x 10; ria, 5, fruiting 
fede x 1; Fic. 6, calyx and e capsule e, x2 


Plate 861 


Rhodora 


Photo. B. G. Schubert. 
RUELLIA CAROLINIENSIS, var. TYPICA: ria. 1, portion of flowering plant, X 1; FIG. 2, flower- 
ace of leaf, X 10; FIG. 5, lower 


ing tip, x 1G. 3, summit of internode, X 4; pig. 4, upper surf 
surface of ial x 10 


Rhodora Plate 862 


Photo. B. G. Schubert. 


LIA CAROLINIENSIS, var. TYPICA: FIG. 1, summit of plant, to show characteristic 
divergen ad ching, X 1; Fic. 2, calyx and Bahl X 2; Fic. 3, portion of calyx and 
capsule 


Rhodora Plate 863 


T COXbEA, 


na Ai ; : ae 
yg ?, ne Word fin O1S , capwlater COPA: 
cs 


Es ee 
ULIS COMOSIS 


Copy, slightly reduced, of the Dillenian plate of RUELLIA STREPENS, CAPIT 


Rhodora Plate 864 


Photo. B. G. Schubert. 


RUELLIA CAROLINIENSIS, var. SEMICALVA: FIG. 1, TYPE, X 2 7: BIG. 2, ‘etree of pus 
node, X 10; Fic. 3, lower, and FIG . 4, upper bartate: of leaf, x 10; Fic. 5, calyx and ¢ 
sule, X 2; FIG. 6, calyx- -segment and portion of capsule, x 10 


Rhodora Plate 865 


Photo. B. G. Schubert. 


1, TYPE, X } 2, upper, and F 
RUELLIA CAROLINIENSIS aah Doe MBRANACEA: FIG, 1, ~ 
, lower surface of leaf, ; ;. 4, flowering tip, 1 7: aa. 5, ‘eatitin of calyx and ¢ fe 
sule, < 10; ria. 6, fruiting scent oe: 


Rhodora Plate 866 


Photo. B. G. Schubert. 


RUELLIA CAROLINIENSIS, Var. MEMBRANACEA: FIGS. 1-3, summits of plants, to show leaf- 
rariation, X< ; 


Rhodors Plate 867 


Photo. B. G. Schubert. 


1, summit, X 1, 


¢ 


D IIENSIS, var SMBRANACEA, forma HYPOPSILA: FIG. 

RUELLIA CAROLINIENSIS, Var. MEMB A : er ep 

of ryPE; FIG. 2, upper, and FIG. 3, lower suriace OI lea’, 
Var. MEMBRANACEA, forma BREVIBERBIS: FIG. 4, internode and leaves, 

and Fic. 6, lower surface of leaf, X 


1.7: FIG. 5, upper, 


Rhodorz Plate 868 


Photo. B. G. Schubert. 


RUELLIA CAROLINIENSIS, sip Lag yptoaslonty EA, forma LAEVioR: FIG. 1, fru 
ing summit, x , Of TYPE . 2, upper, nd F aye 3, lower surface of leaf, 
x 10; Fie. 4, portion of caly X- ae mat and eine age 10 


Rhodora Plate 869 


Photo. B, G, Schubert. 


RUELLIA CAROLINIENSIS, var. prone LLA: FIG. 1, TYPE (2 plan - riG. 2, upper, and 
FIG. 3, iibiig surface of le af, ower-bud, ete Ase Tiant silty 2 calyx-segments. 
x aes 


xX 10; 4, 
10; 5, portion of caly uamicent and capsule 


Rhodorz Plate 870 


eee | 


Photo, B, G. Schubert. 


Jr + 
bellies CAROLINIE tae var. N. spe LLA! Fic. 1, reget ed ci base of _ pal, 
INIENSI : ELLA, lorma ECILIATA: FIG. 2, : (3 plants), X< i. 3, 
portion of leaf, br: kee ‘at poet I X 10; Fic. 4, ‘portion of sis and capsule, x 10 


Rhodora Plate 871 


Photo. B. G. Schubert. 


RUELLIA CAROLINIENSIS, Var. CHELONIFORMIS: FIG. 1, T 2: Fic. 2, corolla, X 1 
FIG. 3, calyx and capsule, X 2; FIG. 4, bract, caly nin i Saban of capsule, 16 


Rhodore Plate 872 


Photo. B. G. Schubert. 


RUELLIA CAROLINIENSIS, Var. CHELONIFORMIS: FIGS. 1 and 2, flowering summits, X 1; 
FIG. 3, upper and ria. 4, lower surface of leaf 


Rhodora Plate 873 


Photo. B. G. Schubert. 


RUELLIA CAROLINIENSIS, var. SALICINA: FIG. 1, TYPE, X 4/7; FIGS. 2 and 3, summits of 


plants, X 1 


Rhodora Plate 874 


Photo. B. G. Schubert. 


RUELLIA ne geen var. DENTATA: FIG. ¥ small plant, X 24; Fic. 2, portion of 
branch, X 1; FIG. 3, porti ion of fruiting branch, X %, to show dhasanlerets branching 


Rhodora Plate 875 


Photo. B. G. Schubert. 


RUELLIA CAROLINIENSIS, var. DENTATA: FIGs. 1 and 2, flowering summits, X 1