Douora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN Associate Editors 
RICHARD ALDEN HOWARD 

CARROLL EMORY WOOD, JR. 


VOLUME 57 


1955 


Che New England Botanical Club, Ine. 


8 and 10 West King St.. Lancaster, Pa. 
Botanical Museum, Oxford St., Cambridge 38, Mass. 


Douora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN Associate Editors 
RICHARD ALDEN HOWARD 

CARROLL EMORY WOOD, JR. 


Vol. 57 January, 1955 No. 673 


CONTENTS: 


A Revision of the North American Genus Sabatia (Gentianaceae). 
Boben E e Soa Oe i a a 0s ee 1 


Ilex glabra and a New Station for Kalmia latifolia in New Hamp- 
shire. A. R. Hodgdon...... 01 eerie eee rrr 34 


A New Variety of Solidago ulmifolia. Arthur Cronquist......... 36 


The New England Botanical Club, Ine. 


8 and 10 West King St., Lancaster, Pa. 
Botanical Museum, Oxford St., Cambridge 38, Mass. 


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JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 57 January, 1955 No. 673 


A REVISION OF THE NORTH AMERICAN 
GENUS SABATIA (GENTIANACEAE)! 


Rosert L. WILBUR 


THE genus Sabatia is restricted in distribution to continental 
North America and the West Indies. It is frequently encoun- 
tered throughout much of its range, which extends from Nova 
Scotia south as far as the mountains of Hispaniola and the 
central plateau of Mexico and from the Atlantie seaboard 
westward to the prairies of Oklahoma and central Texas. All 
of the members of the genus recognized in this paper, seventeen 
species and two varieties, are to be found in the southeastern 
United States. The distribution of most of them is either 
entirely restricted to that area or is largely centered there. 
Most of the entities are either found in or are restricted entirely 
to the Coastal Plain, the oldest portions of which have been 
avallable to the occupancy of land plants only since the close 
of the Cretaceous. 

This genus is believed to be an ancient one whose intrageneric 
divisions appear to have been clear-cut and well-defined for 
a long period of time. Even the species seem, in most instances, 
to be genetically so well isolated from their closest relatives as 
to show no indication of hybridization even when flowering 
in close proximity to one another. These are almost certainly 
indications of great age. 

This group of plants has not been monographically treated 
since the time of Grisebach (1845) although it has been subjected 


1 Part of a dissertation submitted in partial fulfillment of the requirements for the 
degree of Doctor of Philosophy in the University of Michigan. I should like to take 
this opportunity to express my appreciation for the friendship, encouragement and 
guidance of Dr. Rogers McVaugh throughout this investigation. 


2 Rhodora [Vor. 57 


to numerous regional interpretations. The present treatment 
is based upon the examination of more than 6500 specimens 
borrowed from over thirty herbaria? and the prized opportunity, 
made possible largely through the assistance of Professor H. H. 
Bartlett, director of the Botanical Gardens of the University 
of Michigan, of studying all but one of the species in the field. 


HISTORY OF THE GENUS 


Plants now considered to belong to Sabatza were first described 
as species of Gentiana in Gronovius’ “Flora Virginica” (1739). 
Linnaeus, having assisted Gronovius in that work, was personally 
familiar with the specimens upon which the two polynomials 
were based and later provided binomials for both of them in 
“Species Plantarum” (1753). Two additional species, based 
upon specimens collected by Kalm, were described for the 
first time in that work. Three of these species were placed 
by Linnaeus in the genus Chironia together with four very 
different species from the Old World while the remaining species 
known to him was assigned to the genus Swertia. Later he 
transferred one of the species that he had formerly assigned to 
Chironia to Chlora and thus the four different species known 
to Linnaeus were treated as representatives of three different 
genera. 

Adanson (1763), resisting what seemed to him to be the ex- 
cessive generic “lumping” of Linnaeus, recognized many pre- 
Linnean genera and initiated some new ones as well. Sabatza, 


2 The abbreviations employed in referring to these herbaria are those of Lanjouw 
and Stafleu (1952) and are as follows: 


BRU Brown University NCU University of North Carolina 

CHARL Charleston Museum NO Tulane University 

CU Cornell University NY New York Botanical Gardens 

DENN Dennison University OKL University of Oklahoma 

DUKE Duke University OKLA Oklahoma A. & M. College 

F Chicago Natural History PENN University of Pennsylvania 
Museum RUT Rutgers University 

FLAS Florida Agr. Exp. Sta. SMU Southern Methodist Univ. 

GA University of Georgia TAES A. & M. College of Texas 

GH Gray Herbarium TENN University of Tennessee 

K Kew (part of collection) TEX University of Texas 

KSC Kansas State College UARK University of Arkansas 

MICH University of Michigan UC University of California 

MO Missouri Botanical Garden Us U. S. National Herbarium 

MT Université de Montréal WIS University of Wisconsin 

NA U. S. National Arboretum WVA West Virginia University 

NCS North Carolina State College YU Yale University 


1955] Wilbur,—The North American Genus Sabatia 3 


one of the latter, was entirely based upon “Gentiana Gron. 
virg. 29" which Linnaeus had called Chironia dodecandra. 
Later authors frequently ‘corrected’? Adanson’s spelling of 
the generic name to Sabbat?a since it was proposed in commemora- 
tion of an Italian botanist called Liberato Sabbati. In the 
"Familles des Plantes" the generic name Sabatia is invariably 
spelled with a single “b” while the botanist’s name always 
appears as Sabbati. One can therefore only conclude that 
Adanson deliberately chose to spell the generic name as Sabatia 
and, according to Article 82 of the International Code of Botan- 
ical Nomenclature, his choice in this matter is binding. 

Pursh (1814) accepted Adanson’s genus but defined it to 
include all of the American species which had been placed by 
most earlier authors in the genus Chironia. Grisebach (1839, 
1845) provided a very complete account of the eleven species 
and one variety that were known to him in his two monographic 
treatments of the Gentianaceae. These monographs were 
very similar in regard to the species of Sabatza, with the excep- 
tion of the disposition of S. gentianoides. In the first account 
this species was considered to form a monotypic section while 
six years later its distinctive characteristics were warranted 
to be of generie status and, furthermore, it was then placed in a 
different subtribe from that of its former congeners. 

Gray's understanding of those species found within the 
United States was summarized in the ‘‘SSynoptical Flora" (1878) 
in which he recognized thirteen species and two varieties. 

Blake (1915) provided a formalized system of classification 
for the genus by arranging the twenty-two species and one 
variety accepted by him into named subgenera, sections and 
subsections. This was in large part merely the application 
of subgenerie names and validating diagnoses to the groups 
of species that long had been associated together under the 
synoptical and indented keys of earlier authors. 


GENERIC RELATIONSHIPS 


Sabatia was associated in Gilg’s (1895) treatment of the 
family in the tribe Gentianeae, subtribe Erythraeinae together 
with seventeen other genera. Most of the genera of the Ery- 
thraeinae are composed of but few species and more than half 


4 Rhodora [Vor. 57 


of the genera are known only from the Old World. None of 
them seem particularly closely related to Sabatia. This genus 
has been placed most often near Centaurium Hill ( — Erythraea 
Borkh.) from which it may be distinguished by many characters. 
'The following key employs some of their more obvious dif- 
ferences. 


Corolla rotate, the lobes much longer than the tube; stigma deeply 2-lobed, 
the lobes at least one-third the length of the style, at anthesis spirally twisted 
about one another but uncoiling and spreading apart with time; anthers 
either half-twisted laterally or becoming circinnately coiled or recurved 
after pollen discharge... SABATIA. 

Corolla salverform, the lobes equal to or shorter than the tube; stigma capitate 
or but barely creased, the divisions scarcely discernible, position unchanged 
with age; anthers becoming tightly spirally coiled in the manner of a cork- 
screw after pollen discharge... esee CENTAURIUM. 


SYSTEMATIC TREATMENT 
SABATIA Adans., Fam. Pl. 2: 503. 1763. 


Pleienta Raf., Fl. Tell. 3: 30. 1837. 

Neurola Raf., New Fl. 4: 92. 1838. 

Erect, glabrous, annual or perennial herbs. If perennial, sometimes 
stoloniferous and either arising from a short, erect caudex or from a short 
to elongate, almost always branched rhizome. Root-system of the an- 
nuals either a slender tap-root or a cluster of fibrous roots, that of the 
perennials a cluster of wiry to fleshy roots. Leaves mostly cauline but 
basal leaves also present in some species, decussate (or in aberrant speci- 
mens whorled), entire, sessile or somewhat amplexicaul, membranous 
to thick and turgid or even fleshy. Branches opposite or alternate. 
Inflorescence cymose, the cymules either compactly subeapitately clus- 
tered or loosely arranged, being often reduced to one or two flowers, or 
grouped into large, showy, compound dichasia. Flowers perfect, actino- 
morphic. Calyx 5-12(-14)-parted, gamosepalous, the tube campanulate 
to somewhat turbinate, membranous, the lobes minute to foliaceous, 
varying in length from shorter than the calyx-tube to many times longer. 
Corolla rotate, 5-12(-14)-parted, the tube urceolate to cylindrical, 
short but equaling or more commonly exceeding the calyx-tube; corolla- 
lobes usually widely spreading, dextrorsely contorted in bud. Stamens 
5-12; filaments slender, inserted on the upper edge of the corolla-tube, 
alternating with the corolla-lobes, at first erect but strongly curving to 
one side after pollen discharge; anthers basally attached, linear to oblong, 
slender to stout, dehiscing laterally by means of longitudinal slits, after 
pollen discharge becoming either laterally twisted or circinnately re- 
curved or revolute. Ovary bicarpellate, unilocular; carpel margins 
slightly intruding into locule and forming 4 parietally located placental 
lobes each bearing numerous scattered ovules. Style slender; stigmatic 


1955] Wilbur,—The North American Genus Sabatia 5 


branches 2, linear to slightly spatulate, tightly spirally twisted at anthesis 
and bent to one side, later becoming erect, untwisting and exposing the 
densely papillate stigmatic surface. Capsule globose, ovoid, or cylindrical. 
Seeds numerous, globose or somewhat flattened, densely pitted with 
small depressions. TYPE SPECIES: Sabatia dodecandra (L.) BSP. (Chironia 
dodecandra L.) 


KEY TO THE SECTIONS 


Flowers usually long-pedicellate or, if short-pedicellate, the subtending bracts 
small and scale-like; flowers either solitary or loosely cymose; anthers 
becoming circinnately recurved after pollen discharge; plants, if annual, 
pentapetalous, if perennial, either pentapetalous or pluripetalous. 

I. EUSABATIA. 

Flowers sessile or very nearly so, the subtending bracts large and leaf-like; 
inflorescence of few to several closely associated flowers, appearing capitate; 
anthers slightly twisted laterally even prior to pollen discharge, never be- 
coming circinnately recurved; plants annual and pluripetalous. 

II. PSEUDOCHIRONIA. 


I. Section EvsABATIA Griseb., Gen. et Sp. Gent. 120. 1839. 


Subg. Eusabatia (Griseb.) Blake, Rnopora 17: 56. 1915. 

Annuals or perennials. Flowers either pentamerous or plurimerous 
(only the perennials 7-12(-14)-merous); typically long—pedicellate or at 
least not appearing to be sessile and capitately arranged with the clusters 
possessing large foliaceous bracts. Anthers becoming revolutely coiled 
or recurved after pollen discharge. TYPE species: Sabatia dodecandra 
(L.) BSP. 

The fifteen species belonging to this, the larger, section of the genus 
show considerable diversity and may be arranged readily into several 
groups of species (subsections). 


KEY TO THE SUBSECTIONS 
A. Calyx-tube very prominently 5-nerved and thinly membranous-winged ; 
the lateral nerves of the calyx-lobes much more strongly developed than 
the midnerve; flowers pentamerous; annuals......... (C) CAMPESTRES. 
A. Calyx-tube smooth or finely nerved and lacking membranous wings; the 
midnerve of the calyx-lobes almost equaling or even more strongly 
developed than the lateral nerves; annual or perennial and flowers either 
pentamerous or plurimerous. 
B. Perennials with elongate rhizomes; some species stoloniferous; either 
pentamerous or plurimerous. 

C. Branches typically opposite throughout; flowers mostly 5-parted, the 
centermost in the cymules occasionally 6-parted; corolla white and 
lacking yellow “eye” at base of lobes; pedicels usually less than 1 
em. long; stolon and basal rosettes never present. . . (A) DIFFORMES. 
C. Branches typically alternate (at least the uppermost); flowers 5-12 
(-14)-parted (3 species 7-14-parted, the other species 5-6(-7)- 
parted); corolla typically pink or rose and with a yellow “eye” in 
the throat and base of the lobes; pedicels mostly over 1 em. long; 

stolons and basal rosettes present in some species. 
(E) DODECANDRAE 


6 Rhodora [Vor. 57 


B. Annuals or, if perennials, the caudex short, erect, and much-branched; 
rhizomes and stolons never present; flowers pentamerous. 
D. Upper branches typically opposite; basal rosette present at some stage 


of life-cycle and often persistent; annuals........ (B) ANGULAREs. 
D. Upper branches typically alternate; basal rosette lacking; either 
annuals or perennials. ..................... (D) CAMPANULATAE. 


(A) Dirrormes subsect. nov., Subsect. Angulares Blake 
in part, RHopora 17: 56. 1915. 

Robust perennials arising from a thick, gnarled, often much-branched, 
elongate rhizome bearing numerous fleshy roots about 2 mm. in diameter. 
Branches typically opposite. Leaves drying thick-chartaceous, margins 
revolute. Corolla white. TYPE species: Sabatia difformis (L.) Druce. 


The two species associated in this subsection are strongly 
differentiated morphologically from the annual species of sub- 
section. Angulares with which they were formerly placed. In- 
cluding these perennials with the radically different species 
belonging to the Angulares would be placing undue emphasis 
upon the single character of an opposite-branching pattern 
that is typical of both groups while underestimating features 
such as the perennial habit and fleshy roots which seem to me 
to be of greater phyletie significance. The habitat oceupied 
by the perennials is typically that of savannas and ditches, 
whereas the annuals of the Angulares are generally inhabitants 
of drier situations. 

The three entities forming subsection Difformes are a con- 
spicuous floristic feature of savannas or pine-barrens, or the 
marshy borders of low hardwood forests throughout their com- 
bined extensive range. "These plants are, with the exception 
of a very few Piedmont stations at the outer margin of that 
province, entirely restricted to the Coastal Plain, ranging from 
central New Jersey southward throughout much of peninsular 
Florida and westward into Louisiana. The three populations 
do not possess mutually exclusive areas but occupy ranges in 
the nature of overlapping ares somewhat radially arranged 
about what may have been the ancestral home of the group, 
the ancient Appalachian land-mass. It may be speculated 
that the original population, which was perhaps polymorphic, 

3 Subsectio Difformes. Herbae perennes, rhizomatibus crassis, contortis, saepe 
multiramosis radicibus carnosis ca. 2 mm, diam. Rami plerumque oppositi. Folia in 


speciminibus siccis crasse chartacea, marginibus revolutis. Corolla alba. Species 
typica, Sabatia difformis (L.) Druce. 


1955] Wilbur,— The North American Genus Sabatia 7 


migrated from Appalachia onto the Coastal Plain. The frag- 
ments of the original population that reached the new home 
continued to evolve and to spread laterally along the Coastal 
Plain. When they had reéstablished partial contact with their 
close relatives, genetical and other barriers had developed 
which maintained the identity of the three entities. 


KEY TO THE SPECIES OF SUBSECTION DIFFORMES 
Leaves and upper portion of the stem not glaucous; calyx-lobes typically more 
than 4 mm. long and often 10 mm. long or longer; upper bractlets linear, 
those subtending complete (i.e. 3-flowered) cymules 4-5 mm. or more in 
length; corolla-lobes almost invariably longer than 7 mm. and often up to 
15 mm. and occasionally longer, their color often turning saffron or orange 
upon drying, especially along the veins; plants rarely taller than 9 dm. 
high; the main stem and branches becoming strongly angled or even quadrate 
above from 4 internodal ridges; ranging from central New Jersey southward 
and westward into western Florida..................... 1. S. difformis. 
Leaves and upper portion of the stem glaucous; calyx-lobes rarely as long as 
3 mm.; upper bractlets minutely scale-like, even those subtending complete 
cymules less than 3 mm. long; corolla-lobes 4-7 mm. long, color remaining 
white or creamy-white upon drying; plants more robust, mostly taller than 
9 dm.; the main stem and branches terete throughout, not becoming angled 
or quadrate; ranging from central Georgia and ne. Florida westward into 
LOG. ou iure bEL RAE re ru Ee NE E. 2. S. macrophylla. 


1. Sabatia difformis (L.) Druce Swertia difformis L., Sp. Pl. 1: 
226. 1753. Chirenia lanceolata Walt., Fl. Car. 95. 1788. C. cymosa 
Lam., Tabl. Encyc. et Méth. Bot. 1: 479. 1791, not Burm., Fl. Cap. 5: 
1768. C. paniculata Michx., Fl. Bor.-Am. 1: 146. 1803. (C. venosa 
Muhl., Cat. Pl. Am. Sept. 24. 1813. Sabbatia paniculata (Michx.) Pursh, 
Fl. Am. Sept. 1: 138. 1814. S. paniculata æ latifolia Pursh, l.c. S. 
paniculata 8 angustifolia Pursh, l.c. S. corymbosa Baldw. ex Ell, Sk. 
Bot. S. C. & Ga. 1: 283. 1817. S. corymbosa var. angustifolia Ell., l.c. 
S. lanceol. [lanceolata Walt.] Raf., Fl. Tell. 3: 30. 1837, without basi- 
onym. S. cymosa [Lam.] Raf., l.c. without basionym. S. cymosa (Lam.) 
G. Don, Gen. Hist. 4: 207. 1838, as to basionym only. S. lanceolata 
(Walt.) T. & G., Man. ed. 1. 356. 1848. S. difformis (L.) Druce, Bot. 
Exch. Club & Soc. Brit. Is. 3: 423. 1914, as to basionym only. 

Perennial herb (25-)45-80(-105) em. tall, 2-5 mm. in diameter, 1- 
several stems arising in a cluster from a gnarled, stout, branched rhizome 
4-6 mm. in diameter. Stem stiffly erect, hollow, more or less terete 
below, becoming angular and sometimes almost quadrate above from 
the internodal ridges. Branches typically opposite, occasionally alter- 
nate at some nodes, ascendent at an angle of (10—)20-40(—50) degrees, 
generally restricted to the upper tenth or third of the stem, or less com- 
monly in very robust plants branching almost throughout the entire 
length of the stem. Principal branches typically 3—4-times branched 
and together forming a compact, corymbose, flat-topped to convex 


8 Rhodora [Vor. 57 


inflorescence. Numerous fleshy roots descending from the rhizome, 
these 8-12(-20) em. long, 1.5-3 mm. in diameter, usually bearing slender, 
fibrous, lateral roots. Leaves thick, rather succulent, drying somewhat 
thick-chartaceous, venation obscure except for the rather prominently 
elevated midvein and often the lateral pair of veins, or in the broader- 
leaved types with 2 lateral pairs prominent. Basal rosette none; the 
lowermost cauline leaves borne submerged or subterranean, at time of 
flowering usually represented only by scars, but when still present, often 
modified into appressed, obtuse, oblong bracts 2-3 em. long. Upper 
cauline leaves strongly ascendent, linear to lanceolate or even somewhat 
oblong or ovate-lanceolate to strongly ovate (especially in peninsular 
Florida), usually acute, rarely obtuse, somewhat revolute and both 
scarious-margined and -tipped, usually rather strongly clasping, or merely 
sessile in the narrower leaved types, (0.9—)1.8-4(-6.2) em. long, (3-) 
4-14(-22) mm. wide, usually not more than twice exceeded by the in- 
ternodes and usually less; internodes occasionally 3-5 times or more 
longer than the nearby leaves. Pedicels slender, erect, finely ridged, 
2-8(-15) mm. long. Calyx-tube only slightly ridged, rather shallowly 
campanulate, crateriform or even turbinate, 1-2(-3) mm. long; calyx- 
lobes linear to somewhat subulate, very slender, (2-)4-10(-14) mm. 
long, 2-8(-10) times the length of the tube, more or less outwardly arching 
at anthesis, erect in bud. Corolla-tube narrowly cylindrical, (2.5-) 
3-5(-6) mm. long; corolla-lobes typically 5, (the centermost flower 
of the central cymules often 6-parted), oblong, weakly spatulate, elliptic, 
obtuse to acute, wide-spreading, (5-)7-15(-21) mm. long, 2.5-6(-8) mm. 
wide, entirely white even at the base, but often turning orange or saffron 
upon drying, especially along the veins. Filaments 2-3 mm. long, 
pale-yellowish to almost white, slender; anthers slender, lanceolate, 
bright yellow, 2-3 mm. long. Style 2-5 mm. long; stigmatic lobes 
slender, linear, 2-5 mm. long. Capsule oblong-cylindrical, 4-8 mm. 
long, 2.5-4 mm. wide. TYPE LOCALITY: "In Virginia." Type: Clayton 
171 (British Museum); photograph of the type (GH!). DISTRIBUTION: 
Savannas and pine-barrens along the Coastal Plain from (perhaps Long 
Island) central New Jersey southward throughout much of peninsular 
Florida and westward into western Florida. Map. 1. 


The only species with which S. difformis might be confused 
is S. macrophylla, which belongs to the same subsection and 
which shares with it the distinctive features of a stout, gnarled 
rhizome bearing numerous, thick, fleshy roots, characteristically 
opposite branches, thick leaves and white 5(6)-parted corollas. 
However, S. difformis may be readily distinguished from S. 
macrophylla by numerous striking differences among which are 
the absence of a glaucous bloom, the almost invariably longer 
calyx-lobes, bractlets, and corolla-lobes; the very common 
change of color to orange or saffron upon drying, the stem angled 


1955] Wilbur,— The North American Genus Sabatia 9 


or quadrate above, and the typically smaller stature. The 
ranges of the two species overlap in Georgia and in Florida; 
but throughout the main part of the area occupied, they are 
not compatriots. (See Maps 1 and 2) 

S. difformis and another typically white-flowered, opposite- 
branched species, here called S. quadrangula but referred to by 
recent authors as S. paniculata, have been closely associated 
together in many treatments of the genus and often the two are 
misidentified. Gleason’s (1952) account of the species that has 
been passing as S. paniculata fails to distinguish clearly the two 
species. S. paniculata, according to Gleason, is “similar to 
S. difformis in habit, stature, and inflorescence” and is “very 
close to S. difformis in describable characters . . ." The two 
species may be easily distinguished and the differences are such 
that I feel that the two belong to different species-groups. 
The features of S. difformis mentioned in the preceding para- 
graph are in strong contrast to those of the other species which 
is a rosulate annual with a strongly 4-sided stem and membranous 
leaves. 

The synonymy of this species is extensive and there has been 
much confusion and misunderstanding throughout the litera- 
ture regarding these names. It therefore seems advisable to 
discuss each synonym briefly. 

Swertia difformis L. is based upon the Gronovian “Gentiana 
foliis linearibus acuminatis, pedunculis, longissimis nudis unifloris 
oppositis" which in turn rested upon Clayton’s collection 171 
and also apparently two other specimens or perhaps descriptions 
sent by Clayton. These last two references apparently played 
no part in Linnaeus’ concept and it is very unlikely that they 
entered to any extent into that of Gronovius as they presented 
characteristics that were in direct contrast to those described 
in the principal account. They certainly both belonged to 
at least a different species. The specimen in Clayton’s her- 
barium is, as is shown by the photograph in the Gray Herbarium 
or by Blake’s sketch (Rhodora 17: pl. 112. 1915.), the uppermost 
portion of the rhizomatous perennial possessing characteristically 
opposite branches and a_ white, 5(6)-parted corolla. The 
specimen is peculiar in that the lateral branches, all six of them, 
are very much elongated and bear but one flower each. The 


10 Rhodora [Vor. 57 


inflorescence hence is very loose. The terminal flower is six- 
parted and this feature is shown very clearly in the photograph. 
The original description little suggests the plant that it is now 
known to depict and it is therefore not surprising that its proper 
disposition was but so recently accepted by most American 
botanists. The diagnosis, however, is excellent as a description 
of the somewhat atypical type specimen which is the opposite- 
branched perennial which was known throughout the latter 
half of the nineteenth century as S. lanceolata. Druce, in 
transferring the epithet, referred it to what has been called S. 
elliottii, an  alternate-branched, white-flowered annual. In 
this he was following either the “Synoptical Flora" or the 
"Index Kewensis." Clayton's specimen has nothing to do with 
that species. The alternate-branched annual, furthermore, has 
not been found north of South Carolina; the opposite-branched 
perennial is known from southeastern Virginia, where Clayton 
served as a clerk of the court of Gloucester County for many 
years. 

Card (Ann. Mo. Bot. Gard. 18: 262. 1931.) in his monograph 
of Frasera listed Swertia difformis as a synonym of Frasera 
caroliniensis Walt. This mistake has been previously and 
emphatically pointed out by St. John (Am. Midl. Nat. 26: 5. 
1941.). Card was no doubt following Rafinesque (Med. Fl. 
1: 196. 1828.) who listed this name among others as being syn- 
onymous with his substitute name, F. verticillata. Merrill 
(1949) also equates Frasera verticillata in part to Sabatia difformis. 

Of the seven fragmentary specimens of Sabatia still represented 
in Walter's collection two match the brief description of Chironia 
lanceolata and are certainly referable to the opposite-branched 
perennial. The association of Walter's name with the perennial 
has been almost invariably the rule from the time of Pursh (1814), 
who, as stated in his Preface, had examined Walter's collection. 
The combination of Walter's epithet with Sabatia can perhaps 
first be credited to Torrey and Gray in Gray's Manual (1848) 
who, after describing the plant, listed S. corymbosa Ell. as a 
synonym. ‘The only name listed in synonymy under S. corym- 
bosa, when it was originally described, was Walter’s Chironia 
lanceolata. ‘The synonymy of this species presented by Grisebach 
(1839, 1845) was complete and these accounts were doubtless 


1955] Wilbur,— The North American Genus Sabatia 11 


the basis for Gray’s treatment. It is to be remembered that 
Gray also examined Walter's collection in 1839. 

Rafinesque (Fl. Tell. 3: 30. 1837.) may have intended this 
transfer when he listed ''lanceol." in an enumeration of the 
genus Sabatia. Many of the names in this list were never 
described as the promised monograph never appeared. For 
S. lanceolata (as lanceol.), there were no basionym or description 
so even listing the name in synonymy may not be warranted. 

A photograph of the type of Chironia cymosa Lam., which is 
located in Paris, was lent by the Gray Herbarium and it also 
proves to be a specimen of S. difformis. The specimen as is 
shown by the label was one of Fraser’s from “eastern [South] 
Carolina." 

Rafinesque (Fl. Tell. 3: 30. 1837.) in the same list of names 
in which 'anceol." appears, also included cymosa and again 
without basionym or description. It is possible that a transfer 
was intended. 

This epithet, cymosa, was transferred formally by G. Don 
(1838) with the necessary basionym. The description that 
accompanies the name, however, applies only to S. quadrangula, 
a very different species. 

Chironia paniculata Michx. is discussed more fully under 
S. quadrangula, a species with which it has long been confused. 
The original description was brief and the few clues presented 
there have been either overlooked or misunderstood. The 
species was known to Michaux from Georgia and ''Carolina." 
The photograph at the Gray Herbarium of the collection in 
Michaux’s herbarium labeled ‘‘Chironia paniculata Georgia," 
illustrates an excellent specimen and one which is unmistakably 
S. difformis. Pursh transferred Michaux’s epithet to Sabatia 
and noted the synonymy with both Swertia difformis and Chironia 
lanceolata. He had seen the specimen upon which Swertia 
difformis is based in Bank’s herbarium and was acquainted 
with authentic material of Walter's species. In addition Pursh 
designated two varieties, which, as I interpret the descriptions, 
are merely the broad- and narrow-leaved variations that are 
common among representatives of this species. However, 
Gray (1878) and following him Blake (1915) considered Chironia 
paniculata B angustifolia to be, excluding C. lanceolata listed 
in synonymy by Pursh, what is here called S. quadrangula. 


12 Rhodora (Vou. 57 


Chironia venosa Muhl. was relegated to the synonymy of the 
opposite-branched, white-flowered perennial by Torrey as early 
as 1824. Those names published for the first time in Muhlen- 
berg’s Catalogue are so briefly characterized that Merrill and 
Hu (Bartonia No. 25: 22. 1949) stated that they should be 
considered as nomina nuda unless a basionym were included. 
The information presented with Chironia venosa was that the 
corolla was white and veiny and that the plant was a perennial 
found in “N. Caes. Ten." S. difformis is not known from 
Tennessee, nor is any other perennial or any other typically 
white-flowered species. It is found in New Jersey (N. Caes.). 
Otherwise there can be but little doubt that Chironia venosa is 
the same as S. difformis. The specific epithet aptly describes 
the orange discoloration especially noticeable along veins of the 
corolla-lobes in some dried specimens. 

Sabbatia corymbosa Baldw. ex Ell. is the last of the new names 
proposed for the white-flowered, rhizomatose perennial. Au- 
thentie specimens from Elliott’s herbarium prove the correctness 
of treating this name as a synonym of S. difformis. This sheet 
apparently bears two collections, which may be taken perhaps 
as types; one with ovate leaves apparently representing the 
typical element and the other with narrower leaves, probably 
a representative of his S. corymbosa var. angustifolia. This 
specific name was generally employed in designating the peren- 
nial species until the combination of S. lanceolata was made in 
1848. 


REPRESENTATIVE SPECIMENS:—NEW YORK: Nassau Co., Hempstead Flats, 
Mackaness, 1937 (No). Since no other specimen of this species has been seen 
or has been reported from the state and since the region is rather well-known 
botanically, error in labeling is suspected. NEW JERSEY: Burlington Co., 
about 1.5 mi. se. of Atsion, Fogg 5663 (F, GH, PENN); Ocean Co., Tom's River, 
Mackenzie 2774 (MO, US). DELAWARE: Sussex Co., 0.5 mi. w. of Concord, 
Tatnall 3129 (GH). MARYLAND: Worchester Co., Stockton, Rusby, Aug. 1889 
(NY). VIRGINIA: Sussex Co., sw. of Wakefield, Fernald, Long & Clement 
15840 (GH, MO, US). NORTH CAROLINA: Bladen Co., 4 mi. n. of Elizabeth- 
town, Wilbur 2900 (micn); Harnett Co., about 3 mi. se. of Dunn, Wilbur 2897 
(MicH); New Hanover Co., Carolina Beach, Godfrey, PL. EX. GRAYANAE 974 
(F, GA, GH, MICH, MO, MT, NCS, NCU, NY, OKL, PENN, SMU, TENN, TEX, WIS, 
WVA, US). SOUTH CAROLINA: Berkeley Co., about 6 mi. ne. of Summerville, 
Wilbur & Webster 2854 (mMicu); Georgetown Co., 12 mi. n. of Georgetown, 
Godfrey & Tryon 65 (DUKE, F, GH, MO, NY, PENN, TENN, US). GEORGIA: 
Charlton Co., below Trader’s Hill, Small, 12-15 June 1895 (F, Ny); McIntosh 
Co., 1.5 mi. n. of Darien, Cronquist 5375 (Ga, us) Thomas Co., about 1 mi. 


1955] Wilbur,—The North American Genus Sabatia 13 


n. of Coolidge, Duncan 8468 (FLAS, GA, MO). FLORIDA: Dade Co., Biscayne 
Bay, Rusby (MicH) [Since otherwise not known south of Lake Okeechobee, 
this station seems questionable.] Highlands Co., near Sebring, Small, Small 
& Dewinkeler, 17 July 1924 (FLAS, GH, NY, US); Lake Co., Eustis, Nash 849 
(cU, F, GH, MICH, MO, MT, NY, US); Manatee Co., Bradentown, Tracy 7541 
(cU, F, GH, MO, NY, US, WIS); Okaloosa Co., 4 mi. e. of Crestview, Webster & 
Wilbur 3592 (micu); Polk Co., 16 mi. sw. of Kissimmee, Wilbur & Webster 
2637 (micH); Santa Rosa Co., e. bank of Blackwater River across from the 
town of Milton, Webster & Wilbur 3579 (Micu); Walton Co., near DeFuniak 
Springs, Curtiss 5906 (CU, F, FLAS, GH, KSC, NCU, NY, SMU, US); Washington 
Co., 1 mi. e. of Caryville, Webster & Wilbur 3601 (micH). 


2. Sabatia macrophylla Hook., Compan. Bot. Mag. 1: 171. 1836. 


Perennial herb (50-)90-120(-140) em. high, 2-7 mm. in diameter, 
usually with several stems arising in a cluster from a stout, gnarled, 
branched rhizome up to 1 em. in diameter and often 10 cm. or more in 
length. Stem stiffly erect, hollow, terete throughout, without ridges 
or fine lines extending between the nodes, strikingly glaucous above. 
Numerous white to reddish-brown fleshy roots often 15 cm. or more in 
length and about 2 mm. in diameter descending in clusters from the 
rhizome with occasional slender, fibrous, lateral roots. Branches typically 
opposite, very rarely alternate, generally restricted to the upper tenth 
or quarter or, more rarely, throughout the upper half of the stem, as- 
cendent to rather strongly divergent forming an angle of 15-60 degrees 
with the stem, producing a compact, corymbose, flat-topped to somewhat 
convex inflorescence. Leaves thick, somewhat succulent, drying thickly 
chartaceous, strongly ascendent, very noticeably glaucous when fresh, 
ovate-lanceolate, lanceolate (sometimes very narrowly so), oblong, to 
ovate-oblong or even ovate, acute or rarely obtuse, scarious-mucronate 
tipped, margins conspicuously scarious and often slightly revolute, 
strongly clasping, (2.5-)3-6(-8.5) em. long, (0.5—)1-2.5(-4.5) em. wide, 
midvein prominently elevated beneath, 1—2 pairs lateral veins also often 
rather conspicuous. Basal rosette none; the lowermost cauline leaves 
absent at time of flowering. Internodes typically 3-5 times the length 
of the leaves. Inflorescence of numerous eymules corymbosely arranged 
forming a compact flat-topped to somewhat convex cluster. Leaves 
within the inflorescence abruptly reduced to acute, scale-like, usually 
non-chlorophyllous bracts 1-4 mm. long, the ultimate bracts less than 
2 mm. long; the flowering clusters thus appearing naked. Pedicels 
slender, erect or ascendent, 1-5 mm. long. Calyx-tube and lobes more 
or less colorless, non-chlorophyllous; the tube campanulate, scarcely 
ridged or lined by elevated vascular traces, 1-2 mm. long; the lobes tri- 
angular-dentate, subulate, or linear, erect, slightly spreading, or somewhat 
outwardly curved at the tip, or strongly recurved, (0.1—)0.2-2.5(-3.0) mm. 
long, 0.1-3.5 times the length of the calyx-tube. Corolla-tube narrowly 
cylindrical, white, (2-)3-3.5(-4) mm. long. Corolla-lobes oblong to ob- 
long-spatulate, obtuse, widely spreading at anthesis, (4-)5-7 (78) mm. long, 
2-3(4) mm. wide, entirely white or creamy-white, color not changing, or 


14 Rhodora (Vor. 57 


changing but little, upon drying. Filaments (1.5-)3-4 mm. long, white; 
anthers slender, lanceolate, creamy white to pale yellow, 1-2 mm. long. 
Style 1-3 mm. long; stigmatic lobes slender, 1-3 mm. long. Capsule 
globose or nearly so to ovoid, (3-)3.5-4(-4.5) mm. high, (2-)3-4 mm. wide. 

Small (1933) described S. recurvans and, according to him, 
this species differed from S. macrophylla by the characteristies 
presented below. 


S. macrophylla Hook. S. recurvans Small 
Calyx 2-3 mm. long, lobes Calyx 4-5 mm. long, lobes 
shorter than the tube. longer than the tube. 
Corolla-lobes 6-8 mm. long. Corolla-lobes 5-6.5 mm. long. 
Capsule 3-4 mm. long. Capsule 3.5-4.5 mm. long. 
Style and stigma about equal Style much shorter than 
in length. the stigma. 


Available to Small at New York were five sheets which here 
are considered part of the recurvans population. Two of these, 
Harper 1415 and 708, were so annotated by Small. Excellent 
specimens of Harper's 1415 are represented in several herbaria. 
The specimen of this number at New York is chosen as the 
lectotype. It, however, lacks fruit, which was described from 
Harper's 708. 

The specimens of S. recurvans available to Small at New York 
were too few to be significant in evaluating the supposedly 
distinguishing characters. This is especially true in regard to 
the overlapping size-ranges of the corolla-lobes and capsules 
published as additional evidence. "These overlapping ranges 
might be of some significance if based upon a large series of 
specimens but in this case furnished only “padding” to an 
otherwise scantily characterized entity. Even now few fruiting 
specimens are available but measurements of them indicate 
a range of 3-4.5 mm. in length of capsule in both species. The 
length of the corolla-lobes is equally useless as a distinguishing 
feature as the range of S. macrophylla was between 4-8 mm. 
and that of S. recurvans between 4—7 mm. The slight discrep- 
ancy would very likely disappear if a larger series of measure- 
ments were made. The length of the corolla-lobes is apparently 
strongly affected by environmental conditions and the season 
in which the flowers are produced. Flowers developing later 
in the season are generally much smaller than those appearing 
earlier in the year. Therefore, all of the characters with over- 


1955] Wilbur,— The North American Genus Sabatia 15 


lapping ranges presented by Small as supporting evidence of the 
distinetiveness of the two entities prove completely useless 
upon examination of more collections. 

'The ratio of the style to the stigmas was indicated by Small 
to be a reliable distinguishing feature. It was even used by 
him as a secondary key-character. However, the comparative 
length of these structures varies enough so that one may find 
either the stigma or the style longer on the same plant; con- 
sequently this character is not of taxonomic value. The com- 
parative length of the stigma to the style varies considerably 
in different stages of maturity. 

The calyx does possess features which seem to be valid; 
specimens can be assigned to one group or the other by the 
calyx characters and the two groups thus delineated occupy, 
as far as is now known, adjacent but distinct areas. Small’s 
very convenient hiatus of 1 mm. between the size-range of the 
calyx in the two entities disappears upon examination of a 
number of flowers. The overall calyx-length of S. macrophylla 
is 1.1-3(-4) mm., while that of S. recurvans is (2-)2.5-4.0(—4.5) 
mm. long. The overlap of these ranges is too great to be of 
much aid in identification. 

A comparison of the length of the calyx-lobes has much more 
promise, but that, too, fails to provide a clear-cut distinction 
that might be mechanically applied. The macrophylla-popu- 
lation has lobes (0.1—)0.2-1.2(-2) mm. long. Of the more 
than six hundred lobes measured in this group nearly sixty 
per cent were less than 1 mm. long, and over ninety-six per cent 
of the lobes were less than 1.2 mm. long. The few flowers 
with calyx-lobes longer than 1.2 mm. were always the centermost 
flower of the inflorescence or the oldest flower of the cymules 
closest the center of the compound inflorescence. Their lobes 
are often longer than those of other flowers on the same plant. 
Calyx-lobes of the recurvans-group measured (1—)1.5-3 mm. 
long with only five flowers (less than three per cent of the total) 
with lobes less than 1.5 mm. long and with more than seventy 
per cent of the nearly two hundred lobes measured 2 mm. long 
or longer. 

The ratio of the lobe/tube-length provides a somewhat 
sharper means of separation but even this provides no clear- 


16 Rhodora (Vor. 57 


eut division between the two populations. This ratio-range 
of the macrophylla-group is (0.1—-)0.2-1(-2) with all but five 
flowers possessing lobes equaling or less than the tube in length. 
These few flowers were again the centermost which are often 
strikingly different from the others in the inflorescence. The 
same ratio-range in the recurvans-population is (1—)1.3-3(-3.5) 
With only four flowers (2.3 per cent) equaling the tube and 
nearly forty per cent of the flowers possessing lobes two or more 
times greater than the tube. 

Even the feature of the recurved lobes is not restricted to 
the recurvans-group; some of the longer-lobed representatives 
of S. macrophylla, which are found in the extreme eastern portion 
of its range, show at least the beginnings of this tendency by 
possessing lobes slightly recurved at the extreme tip. 

The most useful criterion known to me for the separation of 
the two populations is that based upon the length of the calyx- 
lobes. Future study may demonstrate that the group known 
as S. macrophylla possesses pronounced clinal variation from 
west to east and that this variation is in the direction of the 
even more easternly population previously known as S. recurvans. 
The calycine lobes of the plants from western Florida seem 
longer on the average than those from Louisiana. The even 
more eastern S. difformis, the species most closely related to 
the above-discussed complex, is usually readily distinguished 
by calyx-lobes which are almost invariably longer than even 
those of S. recurvans. Again in regard to characters of the 
calyx there is no sharp separation between the two populations 
but the upper limits of S. recurvans overlap but slightly the 
relatively few calyx-lobes measured in S. difformis which were 
shorter than 4 mm. long. 

The material at hand may be readily assigned to one popula- 
tion or the other with little more than a glance at several calyces 
and the two groups have, as far as is now known, separate 
ranges. The two entities, here considered to represent weakly 
differentiated geographical varieties, are not distinguished by 
strong, or even exclusively delimiting morphological characters. 
Nor are the two varieties widely separated geographically as 
they are now known to occur at least within fifty miles of one 
another. 


1955] Wilbur,— The North American Genus Sabatia 17 


KEY TO THE VARIETIES 
Calyx-lobes erect or only slighty outwardly curved at the tip, equaling or less 
than the calyx-tube in length....... 2a. S. macrophylla var. macrophylla. 
Calyx-lobes strongly recurved, exceeding the calyx-tube in length. 
2b. S. macrophylla var. recurvans. 


2a. Sabatia macrophylla Hook., var. macrophylla. 

Calyx lobes (0.1-)0.2-1.5(-2) mm. long, triangular-dentate to linear, 
erect or only slightly outwardly curved at the tip, equaling or less than 
the calyx-tube in length (very rarely the centermost flower of the central 
eymules may have a ratio of lobe/tube greater than 1). TYPE LOCALITY: 
near Covington, Louisiana. Type: Drummond s. n. (K!). DISTRIBUTION. 
Savannas and pine-barrens and margins of low hardwoods from western 
Florida and Georgia westward to Louisiana. Map 2. 


REPRESENTATIVE SPECIMENS:—GEORGIA: Muscogee Co., Columbus, Boykin 
(BRU, GH, NY); Sumter Co., Americus, Harper, 30 July 1897 (GH). FLORIDA: Bay 
Co., 1 mi. n. of Lynn Haven, Webster & Wilbur 3626 (mica); Calhoun Co., 
2 mi. s. of Chipola, Thorne & M uenscher 8744 (cv); Columbia Co., Lake City, 
collector not stated (F); Escambia Co., 9 mi. w. of Pensacola, Webster & Wilbur 
3575 (MicH); Franklin Co., Apalachicola, Biltmore Distrib. Chapman Herb. 
4508a (an, NY); Gadsen Co., inter Quincy et Aspalga, Rugel, July 1843 (wo, 
ny); Gulf Co., Port St. Joe, Knight, 18 June 1936 (FLAS) ; Liberty Co., Bristol, 
West & Arnold, 23 July 1940 (Frias); Okaloosa Co., about 3 mi. e. of Crestview, 
Webster & Wilbur 3584 (micu); Wakulla Co., 1.5 mi. s. of Sopchoppy, Webster 
& Wilbur 3628 (micu); Walton Co., near Argyle, Curtiss 5931 (CU, FLAS, GH, 
KSC, MO, NCU, NY, SMU, US); Washington Co., 8 mi. s. of Chipley, Webster & 
Wilbur 3614 (micH). ALABAMA: Baldwin Co., 13.5 mi. w. of Foley, Webster & 
Wilbur 3530 (micu); Mobile Co., about 8 mi. w. of Mobile, Webster & Wilbur 
3481 (mic); Washington Co., 40 mi. n. of Mobile, Sargent, 18 June 1950 (Ga, 
NCS). MISSISSIPPI: Forest Co., Lake Shelby State Park about 14 mi. s. of 
Hattiesburg, Webster & Wilbur 3402 (micu); Harrison Co., Biloxi, Tracy 7006 
(F, MO, NY, US); Jackson Co., Ocean Springs, Pollard 1064 (cU, F, MO, NY, US); 
Perry Co., 9 mi. n. of Beaumont, Webster & Wilbur 3422 (micH); Stone Co., 
3 mi. s. of Wiggins, Webster & Wilbur 3439 (Micu). LOUISIANA: St. Tammany 
Parish, Covington, Drummond TYPE (k). 


2b. Sabatia macrophylla Hook., var recurvans (Small) comb. nov. 
Sabbatia recurvans Small, Man. SE Fl. 1049. 1933. i 

Calyx-lobes (1—)1.5-3 mm. long, linear to subulate, strongly recurved, 
typically exceeding the calyx-tube in length (less than 3 per cent of the 
nearly 200 lobes measured had a lobe/tube ratio of 1 or 1.2). LECTOTYPE 
LOCALITY: Moist pine-barrens east of Ocilla, Irwin Co., Georgia. Lecto- 
type: Harper 1415 (NY!. DISTRIBUTION: Savannas and pine-barrens 
of southern Georgia and northeastern Florida. Map 2. 


REPRESENTATIVE SPECIMENS:—GEORGIA: Brantley Co., 3 mi. e. of Nahunta, 
Wilbur & Webster 2748 (micu); Charlton Co., 8 mi. s. of Folkston, Wright 881 
(cv); Coffee Co. without exact locality, Harper 708 (NY, vs); Emanuel Co., 
e. of Swainsboro, Duncan 2677 (micu); Irwin Co., e. of Ocilla, Harper 1415 
(r, GH, MO, NY, US); Laurens Co., 10 mi. e. of Dublin, Pyron & McVaugh 750 


18 Rhodora [Vor. 57 


(G^, US); Worth Co., 3.5 mi. w. of Sylvester, Thorne 6363 (cU). FLORIDA: 
Baker Co., without exact locality, Curtiss 2224 (cU, F, Mo, NY, us, YU); Clay 
Co., without exact locality, T'odsen, 30 July 1939 (rtas); Duval Co., 15 mi. w. 
of Jacksonville, Wiegand & Manning 2566 (cu, au); Nassau Co., without 
exact locality, Knight, 1 July 1941 (Fuas). 


(B) Subsection ANGuLARES Blake, Rhodora 17: 56. 1915. 


Annuals with fibrous roots and with typically opposite branches. 
Leaves thin and membranous, margins not scarious. Basal rosette 
usually well-developed at some time during maturation of plant. TYPE 
SPECIES: Sabatia angularis (L.) Pursh. 


The three species comprising subsection Angulares, as here 
defined, are readily distinguished from one another. Asa group 
it is strongly differentiated from all other subsections of section 
Eusabatia. The annual habit separates them from subsections 
Difformes and Dodecandrae, the opposite branches and the 
typically 3-flowered cymules set them apart from the Campanu- 
latae, and the thin, merely finely ridged calyx-tube and, again, 
the opposite branches easily distinguish the Angulares from the 
Campestres. This subsection occupies the largest area of any 
within the genus. 

These three species and the other species reported as annuals 
in the paper have most often in the past been treated as biennials. 
Field observations and limited experience with some of the 
species in the greenhouse has led me to believe that they are 
all probably annuals but this point has not been definitely 
established. 


KEY TO THE SPECIES OF SUBSECTION ANGULARES 


A, Lower portion of the stem strongly 4-angled, quadrate, the corners with 
conspicuous membranously-winged margins. 

B. Corolla pink to roseate (very rarely white); pedicels elongate, 1 em. or 
more in length; lower cauline leaves usually broadly ovate to ovate- 
lanceolate, usually less than 3 times as long as broad; calyx-lobes 
usually 1.5 mm. or more in width (rarely as narrow as 1 mm.). 

3. S. angularis. 

B. Corolla white; pedicels short, less than 5 mm. long ; lower cauline leaves 
usually oblong, generally 4 or more times as long as broad; calyx-lobes 
less than 0.5 mm. wide.............2.......... 4. S. quadrangula. 

A. Lower portion of the stem terete and not bearing thin membranous wings. 
5. S. brachiata. 


3. Sabatia angularis (L.) Pursh Chironia angularis L., Sp. Pl. 1: 190. 
1753. C. angularis var. « latifolia Michx., Fl. Bor.-Am. 1: 147. 1803. 
Sabbatia angularis (L.) Pursh, Fl. Am. Sept. 1: 137. 1814. S. angularis 


1955] Wilbur,— The North American Genus Sabatia 19 


var. albiflora Raf., Med. Fl. 2: 77. 1830. nom. nud. S. angularis, var. 
elatior Raf., l.c., nom nud. S. angularis, var. latifolia Raf., l.c., nom nud. 
S. angularis, var pauciflora Raf., l. c., nom nud. S. angularis, f. albiflora 
Raf. ex House, Bull. N. Y. State Mus. 254: 566. 1924. S. angularis 
f. cleistantha Fern., RHopora 42: 474. 1940. 

Erect annual (15-)30-50(-90) em. tall, often robust and usually with 
the numerous branches presenting in plants developed under favorable 
conditions a bushily compact aspect. Stem strongly tetragonal, hollow, 
the corners conspicuously wing-angled by thin membranous extensions 
0.1-0.5 mm. wide. Stem green in younger portion becoming yellowish 
straw-colored below, (1-)2-3(5) mm. in diameter. Branches typically 
opposite, although ultimate-branching often alternate, usually by marked 
abortion of opposing bud, or generally alternate in depauperate plants, 
often numerous, rarely extending to base, but more typically restricted 
to upper third or half of stem, rather closely ascendent forming an angle 
of about 20-45 degrees with main stem. Branches composed of 1-10 
nodes in well-developed plants and forming a convexly-corymbose or 
pyramidal compact crown. Roots several, fibrous, 4-10(-20) em. long, 
(0.2-)1-3(-4) mm. in diameter, wide-spreading and usually shallowly 
situated. Leaves thin membranous, spreading to ascendent, drying 
thin, 5-7-nerved or fewer in those leaves transitional to bracts, (1—-)2-3.5 
(—4.5) em long, (0.8—)1-2.5(—3.5) em. wide, venation prominently elevated 
beneath. Basal rosette rarely present at anthesis, composed of spreading, 
shortly-spatulate, broadly oblong to obovate, obtuse leaves up to about 
4 em. long and often 2-3 em. wide. Lowermost cauline leaves usually 
absent at time of flowering, the lowest usually broadly ovate, always 
strongly clasping, obtuse, typieally nearly as broad as long, the middle 
narrower, more typically ovate-lanceolate, obtuse or more commonly 
acute, and gradually reduced and modified above to the 1-nerved linear 
bracts. Internodes generally about 1.5-3 times as long as the leaves. 
Inflorescence of corymbosely- or pyramidally-arranged cymules, either 
or both of the lateral branches sometimes suppressed.  Pedicel 5-sided, 
slightly wing-angled, (1—)2-3.5(—4.5) em. long, erect or strongly ascendent. 
Calyx-tube thin, shallow, 2-5 times exceeded by the corolla-tube, usually 
about 1.5 mm. long (1-2 mm.), with venation somewhat elevated. 
Calyx-lobes thin, narrowly linear, linear-oblong, to somewhat foliaceous, 
usually about 1 em. long (0.4-1.5 em.) and (1-)1.5-2.5(-3.5) mm. wide, 
ascendent in bud, wide-spreading at anthesis, usually exceeded by the 
corclla by 5-12 mm. Corolla-tube 4-7 mm. long, 2-4 mm. in diameter, 
cylindrical, pale greenish-white within and without. Corolla-lobes 
oblong, spatulate, or elliptic, obtuse, wide-spreading, (0.6—)1—1.8(-2.2) 
em. long, (2-)4-7(-9) mm. wide, greenish in bud turning white prior 
to full development and typically pink or roseate at anthesis or rarely 
white, coloration paler beneath, with an irregular, somewhat pointed, 
yellow or greenish-yellow area at base of lobe and in throat and usually 
bordered by a dark-red line. Filaments 2.5-4.5 mm. long, greenish- 
yellow to pale-yellow; anthers slender, bright yellow, 3-5 mm. long. Stig- 


20 Rhodora [Vor. 57 


matic lobes very slender, 3-6 mm. long. Style 4-6 mm. long, usually some- 
what less than length of stigmatie branches. Capsule cylindrical at ma- 
turity 5-9 mm. high, 4-6 mm. wide. Type Locarrrv: “In Virginia." Type: 
Kalm (in the Linnean Herbarium), but not seen. DISTRIBUTION: Margins 
of woods, fields, and prairies from southern New York south to northern 
Florida and west to Illinois and Texas. Occasionally introduced in 
Massachusetts and Connectieut. Map 3. 


This species is by far the most familiar representative of 
the genus because of its wide range, the greatest of any in the 
genus, and also because of the frequency with which it is en- 
countered throughout most of that range. "The characteristic 
features are the conspicuous tetragonal stem, the corners of 
which are strikingly membranously wing-margined; the typically 
opposite pattern of branching, the ovate-clasping leaves, the 
roseate corolla (although albinos are encountered rather rarely), 
and the shallow calyx-tube, the venation of which, although 
often slightly elevated, is not developed into a heavy costa. 
None of these characters alone is enough to distinguish the 
species but the combination of them in one plant is unique. 
In the western part of the range of S. angularis, it has been 
confused surprisingly often with S. campestris. This latter 
species may be readily distinguished from the former by its 
typically alternate branching-pattern and especially by the 
turbinate and comparatively long calyx-tube with strongly 
developed costae. In the southeastern region, it has been 
confused at times with S. brachiata, but is at once separable 
from that species which possesses an almost terete stem and 
oblong, non-clasping leaves. 

The species is remarkably uniform throughout its range. 
None of the trivial variants designated previously is here deemed 
worthy of recognition nor have any others been discovered. 
Although I have not seen even a photograph of the type of 
Chironia angularis B angustifolia, if it is still extant, I am ac- 
cepting for the time being the past interpretation that this 
entity is what is now called S. brachiata. Coordinate with 
this variety, Michaux published C. angularis x latifolia which 
is perhaps the typical element of S. angularis. The species 
remained unencumbered by the designation of minor varieties 
until Rafinesque. This author, after discussing S. angularis, 
stated that "it has some varieties: 1. Albiflora, 2. Latifolia, 


1955] Wilbur,— The North American Genus Sabatia 21 


S. MACROPHYLLA 
€ VAR. MACROPHYLLA 


Q VAR. RECURVANS 


S.DIFFORMIS 


S. ANGULARIS S.QUADRANGULA 


Mars1-4. Map 1, upper left; map 2, upper right; map 3, lower left; map 4, lower right. 


3. Pauciflora, 4. Elatior." None of these propositions is dis- 
cussed or mentioned by him further in any manner and hence 
are merely nomina nuda. Therefore, they are not available 
for transferral to any other rank. House, in providing formal 
nomenclature to designate the albino form, erred in transferring 
a name which “has no status under the Rules." For those 
who find it useful or satisfying to name such striking minor 
variations the epithet for the albino might best be written as 
Sabatia angularis forma albiflora House. Fernald described 


22 Rhodora [Vor. 57 


a teratological specimen as S. angularis forma cleistantha. It 
has not been seen again in any of the many hundreds of sheets 
of this species that I have examined. The branches are stunted 
and malformed bearing sessile or nearly sessile flowers which 
are aborted and modified in size and form as to be scarcely 
recognizable as flowers. Little is to be gained by formal designa- 
tion of such rare chance monstrosities. 


REPRESENTATIVE SPECIMENS:—CONNECTICUT: Locality not stated, Wright 
(yu). New York: Richmond Co., near New Dorp, Britton, 9 Aug. 1879 (vv). 
New JERSEY: Cape May Co., Cold Spring, Pennell 2128 (us). PENNSYLVANIA! 
Berks Co., 0.5 mi. wnw. of Gibralter, Berkheimer 2926 (GH, PENN). DELA- 
WARE: New Castle Co., near Wilmington, Canby, Aug. 1878 (MO). MARY- 
LAND: Harford Co., se. of Flintville, Adams & Hopkins 923 (cu, PENN). 
DISTRICT OF COLUMBIA: Chain Bridge, Earlanson & Martin 1210 (NA). 
VIRGINIA: Southampton Co., se. of Ivor, Fernald & Long 10784 (GH, Mo). 
NORTH CAROLINA: Buncombe Co., near Biltmore, Biltmore Herb. 3303a (F, 
GH, MO, NY, US); Durham Co., about 4 mi. ne. of Durham, Wilbur 2898 (micu); 
Halifax Co., Roanoke Rapids, Godfrey 5178 (DUKE, GH, US). SOUTH CAROLINA: 
Williamsburg Co., 2 mi. ne. of Lane, Godfrey & Tryon 399 (DUKE, F, GH, MO, 
NY, PENN, TENN). GEORGIA: Bartow Co., about 4.5 mi. s. 16° east of Alla- 
toona Dam, Duncan 8522 (FLAS, GA, GH, MO, TENN, US); Catoosa Co., 10 mi. 
w. of Ringgold, Cronquist 5610 (GA, GĦ, Us). FLORIDA: Jackson Co., Sneads, 
Knight, 29 June 1943 (FLAs). MICHIGAN: Kalamazoo Co., Hermann 9042 
(wrcH). OHIO: Coshocton Co., near Coshocton, Moldenke 13289 (OKLA, 
PENN, SMU). INDIANA: Morgan Co., about 2 mi. w. of Centerton, Friesner 
17008 (GH, NY, OKLA); Steuben Co., near Lake Gage, Deam, 11 Aug. 1903 
(GH, Us, wis, YU). ILLINOIS: Marion Co., Salem, Bebb 1860 (r, an, vv). 
WEST VIRGINIA: Cabell Co., overlooking Roland Park, Gilbert 778 (DUKE, F, 
GA, GH, MICH, MO, MT, NA, NY, OKL, PENN, RUT, TENN, Us, WIS); Putnam Co., 
near Hurricane, Gilbert et al., so. APP. BOT. CLUB DISTRIB. 146 (FLAS, MO, NCS, 
NY, OKL, PENN, TAES, TENN, WVA). KENTUCKY: Calloway Co., se. of Aurora, 
Smith & Hodgdon 4050 (au, Ny, us); McCreary Co., Cumberland Falls, 
McFarland & James, 2ND CENT. FL. KENT. 35 (DUKE, GA, MO, NY, PENN, TENN, 
WIS, WVA, US). TENNESSEE: Cheatham Co., Pegram, Svenson 10338 (an, 
TENN). ALABAMA: Lowndes Co., 23 mi. se. of Selma, Webster & Wilbur 3511 
(MICH). MISSISSIPPI: Amite Co., 5 mi. sw. of Gloster, Webster & Wilbur 3277 
(mica); Perry Co., 14 mi. w. of Beaumont, Webster & Wilbur 3405 (micn). 
Missouri: Howell Co., 4.5 mi. nw. of Willow Springs, Steyermark 23461 (F, 
MO). ARKANSAS: Pope Co., Nogo, Merrill 545 (TEX, UARK). LOUISIANA! 
Grant Parish, 15 mi. s. of Winnfield, Webster & Wilbur 3259( MICH). KANSAS: 
Cherokee Co., Hitchcock, Aug. 1896 (ksc). OKLAHOMA: Pushmataha Co., 
Antlers, Palmer 8315 (wo, wr, NY, Us). TEXAS: Smith Co., Swan, Reverchon 
3120 (MO, SMU). 

4. Sabatia quadrangula sp. nov.‘ 


Sabbatia cymosa in the sense of G. Don, Gen. Hist. 4: 207. 1838, as to 


4Sabatia quadrangula sp. nov. Herba annua rosulata, caulibus quadrangularibus, 
ramis oppositis, pedicellis brevibus, 1-2(-4) mm. longis, corolla alba, 5-partita. 
Specimen ty picum legit R. Wilbur (n. 2899) prope Durham, in Carolina Septentrionali, 
et in Herb. Universitatis Michiganensis conservatum, 


1955] Wilbur,— The North American Genus Sabatia 23 


Don’s description but excluding reference to Chironia cymosa Lam., 
a synonym of Sabatia differmis (L.) Druce. 


Sabbatia paniculata in the sense of authors after 1860 but not that of 
Michaux, the author of the basionym, nor of Pursh who made the 
combination. The earlier authors were describing the perennial 
now known as S. difformis (L.) Druce. 

Sabatia brachiata f. candida Fern., RHopora 39: 443. 1937 

Erect annual (15-)25-45(-75) cm. high, usually with but one stem 

arising from each rosette, but occasionally with two to several. Roots 
few to more typically numerous, slender, fibrous, usually widely spreading 
and shallowly situated, about 4-8 em. long and less than 1 mm. in diam- 
eter. Branches usually restricted to upper one-quarter or one-third 
of stem, occasionally arising from nodes of the upper half of the stem 
or rarely from even the lcwermcst nodes, usually rather strongly as- 
cendent, forming an angle of 15-30 degrees, or occasionally up to 60 
degrees, typically opposite along the stem and principal branches, the 
ultimate ramifications more commcnly alternate, the branches forming 
a flat-topped or eonvexly corymbcse loose to compact crown. Stem 
strongly quadrate, conspicucusly so below, 1-2(-3) mm. in diameter, 
strikingly although finely membranous wing-angled, wings thin, very 
low, less than 0.5 mm. in height in the lower portion of the stem and 
reduced above to the merest trace. Basal rosette present or absent at 
anthesis, even when present often not well-developed or conspicuous, 
when well-developed of spreading obovate to spatulate, obtuse to oc- 
casionally acute leaves tapering to an almost petiolate base, about 2-4 
em. long and 1.8 em. wide. Cauline leaves membranous, ascendent, 3- 
nerved, or only l-nerved in smaller leaves, somewhat revolute, (0.8-) 
1.6-3.2 (-5.8) em. long, (3-)4-8(-18) mm. wide, obtuse to acute, typically 
apiculate, usually narrowly to broadly oblong or lanceolate, usually 
about 3-5 times as long as broad but ranging from about 2-8 times as 
long as broad, typically at least the lowermost with a strongly clasping 
base. Internodes often equaling cr slightly exceeding the leaf, com- 
monly about 1-2 times as long, but ranging from 0.5-5 times as long as 
the leaves. Inflorescence of corymbosely to pyramidally arranged 
cymules, either one, both or none of whose lateral branches may be 
suppressed, flowers unusually closely associated in the cymule as the 
pedicels are always short, the flowers often sessile or nearly so, pedicels 
usually not more than 1-2 mm. in length, always less than 4 mm. long. 
Calyx-tube thin, 5 ridge-angled, about one-half as long as the corolla- 
tube, or about 2-3 mm. high, broadly turbinate, 1.5-3(-3.5) mm. long. 
Calyx-lobes thin, narrowly linear, (2-)4-8(-11) mm. long, usually about 
0.5 mm. or less in width, but very rarely lobes as broad as 1.5 mm., erect 
in bud, and apparently ascendent at anthesis arching upward between 
the corolla-lobes, usually exceeded by the corolla-lobes by about 2-5 mm. 
Corolla-tube (2.5-)4-5(-7) mm. long, white but appearing greenish as 
the ovary is visible through the thin translucent wall. Corolla-lobes 


24 Rhodora [Vor. 57 


usually oblong, or spatulate, or somewhat elliptie, usually obtuse but 
not uncommonly acute, (4.5-)6-12(-15) mm. long, (1.5-)3-4(-60) mm. 
wide, spreading, pure white, occasionally with or more commonly without 
the basal yellow patch, often turning saffron-yellow upon drying, es- 
pecially along the reticulate veins. Filaments slender, white, 2-4 mm. 
long; anthers pale yellow, 1.5-3 mm. long. Stigmatie lobes greenish, 
lanceolate to oblong, 2-4 times the length of white style, 0.5-1.5(-2) mm. 
long. Capsule cylindrical, 4.5-7 mm. long, 3-4 mm. in diameter. TYPE 
LOCALITY: About four miles northeast of Durham, Durham County, 
North Carolina, rype: Wilbur 2899 (MICH). DISTRIBUTION: Fields, pine- 
lands and drier savannas from southeastern Virginia south to central 
Florida. Map 4. 


This species has been known for almost one hundred years 
as Sabatia paniculata. This name, based upon Chironia panicu- 
lata Michx., has long been a source of confusion and error and 
has been applied by various authors to at least three different 
species. For almost the past century its application has been 
rather consistently restricted to the species which is here called 
S. quadrangula. 


Michaux's original description in full is as follows: 

paniculata. C. firmiter erecta: foliis lanceolato-linearibus; panicula multi- 
flora, brachiata, subfastigata; calyce subulato, corolla 
semibreviore. 

Obs. Caulis lineis 4 prominulis quasiquadrangulus: folia inferiora 
interdum vali-lanceolata; suprema subulato-setacea. Co- 
rollae laciniae oblongae. 

Hab. in Georgia et Carolina. 

To be sure, there is very little descriptive information here 
that would be sufficient to allow anyone to be certain or even 
relatively confident as to which entity Michaux was attempting 
to characterize. Many of the phrases such as “C. firmiter 
erecta . . . panicula multiflora, brachiata, subfastigata: calyce 
subulato, corollae semibreviore . . . Corollae laciniae oblongae" 
might be applied with equal propriety to several other species. 
Of the five species included by Michaux, the flower-color of 
the four others is described as being rose; that of C. paniculata 
is not given. "Those characters of the leaf that were mentioned 
are rather indicative of S. difformis (“. . . foliis lanceolato- 
linearibus . . . folia inferiora. interdum ovali-lanceolata; su- 
prema subulato-setacea . . ."). The features of the stem 
described in the original diagnosis, especially “caulis lineis 4 
prominulis quasi quadrangulus .’ are also a much more 


1955] Wilbur,— The North American Genus Sabatia 29 


apt description of the stem of S. difformis with its four elevated 
lines or ridges in the upper portion of the stem than it is of the 
very pronounced and strikingly 4-sided stem of S. quadrangula. 


It was not surprising, therefore, to find that Michaux’s col- 
lection, as shown by a photograph in the Gray Herbarium of 
the type, was a specimen of S. difformis. This species is a 
perennial with a large rhizome; the stem below is terete or with 
four lines or ridges superimposed upon it, and above the stem 
becomes somewhat quadrangular; the leaves are ovate-lanceolate, 
lanceolate, linear to setaceous in a gradually modified sequence 
from the base to the inflorescence; the corolla-lobes are often 
5 mm. or more longer than the longest lobes observed in S. 
quadrangula. The specimen in Michaux’s herbarium, as is 
witnessed by the very adequate photograph, is excellent and 
most of these features may be observed or interpreted from it. 
The underground portion of the plant, to be sure, is lacking. 
It is obvious that the lowermost part of the specimen (which 
can be seen even from the photograph to be terete or nearly 
so and approximately twice the diameter of the largest stem 
of the species that has most recently passed as S. paniculata) 
has been broken from a very substantial underground structure. 
Therefore Chironia paniculata is, both on the basis of its original 
diagnosis and upon the photograph of the specimen in Michaux's 
own collection in Paris, but a later synonym for Swertia difformis 
L. (1753), Chironia lanceolata Walt. (1788) and Chironia cymosa 
Lam. (1791). 

Pursh (1814) transferred Michaux’s epithet to Sabatia (as 
Sabbatia) listing Swertia difformis as a synonym, stating by 
way of explanation that this species “certainly is the long 
lost Swertia difformis, as the specimens in the Herbarium of 
Sir Joseph Banks, sufficiently prove." Pursh also formally 
named as varieties the broad- and narrow-leaved forms of this 
species with C. lanceolata Walt. as a synonym of the latter. 
Elliott (1817) rather hopelessly confused the white-flowered 
Sabatias nomenclatorially and the influence of his splendid 
"Sketch" was rather long-lasting. The very different S. brevi- 
folia Raf. (S. elliottii Steud.) was very well-described, no doubt 
for the first time, but called S. paniculata with the following 
note by way of explanation: “Though the description of Michaux 


26 Rhodora (Vou. 57 


applies more peculiarly to the S. corymbosa [S. difformis], yet 
as this species was definitely included, and is the only one to 
which the term paniculata is correctly applicable, I have referred 
to him here." This freedom of interpretation on the part of 
Elliott caused confusion that persisted for many years. Grise- 
bach (1839) pointed out the confusion but apparently felt that 
it would be best to apply the name S. paniculata in the sense 
of Elliott, who had been followed by all American authors, and 
to adopt the name S. corymbosa for the perennial species. His 
list of synonymy showed a very complete understanding of 
the nomenclatoral propositions that had been made for this 
last-mentioned entity for the names of Linnaeus, Walter, La- 
marck, Michaux, Pursh, and finally Baldwin (or Elliott) are 
arrayed there. Grisebach apparently included the opposite- 
branched annual, here called 5. quadrangula, in the same concept 
for a duplicate of the specimen of Beyrich cited by him under 
S. corymbosa is the earliest collection of S. quadrangula known 
to me. Chapman (1860) was the first to define S. paniculata 
in the sense that it has been applied in recent times. His action 
in this, as in most other matters, was heavily influenced by 
ray as 1s shown from a letter (in the National Herbarium, 
dated December 1883) from Chapman to Dr. J. H. Mellichamp 
which is here quoted in part. 


. . . I well remember the quarrel Gray and I had over it, I contending that 
the one we now call S. elliottii, our really only paniculate species, should bear 
that name. However, for the sake of uniformity as to our northern and 
southern Floras, I yield. 


The appearance of the Synoptical Flora (1878) firmly es- 
tablished the usage of S. paniculata as the name of the white- 
flowered, opposite-branched annual and this name has been 
so applied since that date. 

Rafinesque (Med. Fl. 2: 77. 1830.) published a description 
that in some ways strongly indicates this entity. This name 
has never been identified and I am at a loss to know what species 
Rafinesque had. Sabbatia nivea was described by him as having 
snowy-white, trichotomose flowers, a four-angled stem and 
oblong leaves which all would indicate S. quadrangula as well 
or better than any other species. However, the species was 
definitely stated to have been discovered by him in eastern 


1655] Wilbur,—' The North American Genus Sabatia 27 


Kentucky near the Cumberland River. There is no species 
known to me from that area which matches his description. 


G. Don (1838) transferred Lamarck's C. cymosa to Sabatia 
applying it to what is here called S. quadrangula. This usage 
was not followed by any other author. The photograph of the 
specimen upon which Lamarck’s species was based is again 
what is now known as S. difformis. 

Gray (1878) placed S. paniculata var. latifolia Pursh under 
the synonymy of what is now considered S. difformis while 
placing S. paniculata var. augustifolia Pursh under the species 
which I am calling S. quadrangula and which he called S. panicu- 
lata. In other words he felt that Pursh was including two 
species under S. paniculata and Gray also implied that Michaux 
had done so as well. I am aware of no evidence that has been 
presented that would warrant such a conclusion. 

The only name that has been published previously for this 
species, whose identity has been for so long known to American 
botanists, is S. brachiata forma candida Fernald. This name 
must be assigned to the synonymy of S. quadrangula rather than 
to S. brachiata as the type specimen, Fernald & Long 6346, is not 
an albino of S. brachiata as claimed by Fernald, but rather is 
a specimen of the white-flowered species, S. quadrangula. Its 
identity is shown by the stems which below are strongly quadrate 
with conspicuously membranous-margined angles and whose 
pedicels are 2 mm. or less in length so that the flowers appear 
almost sessile. The only specimen cited by Fernald besides 
the type was Seymour 33 which came from Sussex Co., in south- 
eastern Virginia as did the type-collection of Fernald’s form. 
Seymour's plant, seen only at the Gray Herbarium, is S. brachiata 
as is indicated by the stem being terete below and some of the 
pedicels being over 5 mm. long and many of them over 2 mm. 
long. The flowers are bleached on the herbarium specimen 
but, as the sheet was originally determined as S. paniculata, 
they were very likely white. I also have seen white flowered 
specimens of S. brachiata in the field. However, since the 
designated type of Fernald's form is what is here called S. 
quadrangula, it wil be necessary for those who feel it useful 
formally to designate such minor variations to provide another 
name for albino plants of S. brachiata. Fernald's name could have 


28 Rhodora [Vor. 57 


been elevated to specifie rank for the annual species with a 
quadrate stem instead of providing an entirely new name. This 
procedure would seem undesirable as Fernald's intention in 
describing the form is clear; his choice of type was unfortunate. 
Besides this consideration, isotypes are not available for that 
number. Therefore, a new name has been provided and a 
type designated of which there are many duplicates. 


REPRESENTATIVE SPECIMENS:—VIRGINIA: Chesterfield County, near 
Chesterfield Courthouse, Fernald & Long 6342 (an); Greensville Co., nw. of 
Taylor's Millpond, Fernald & Long 10785 (GH, PENN); Hanover Co., 1 mi. s. of 
Ashland, Ward, 8 Aug. 1885 (vs); Henrico Co., west of Elko Station, Fernald 
& Long 8818 (au); James City Co., south of Hotwater, Fernald & Long 8817 
(DUKE, GH); Pittsylvania Co., Falls Creek, Heller 1105 (wv, us); Prince 
George Co., w. of Prince George Courthouse, Fernald & Long 8816 (am); 
Sussex Co., ne. of Homeville, Fernald & Long 6341 (an, MO, NY, PENN, US). 
NORTH CAROLINA: Anson Co., 6 mi. w. of Wadesboro, Smith, 30 July 1884 
(au, us); Columbus Co., Nakina, Schallert, 30 June 1928 (oxra); Dare Co., 
Old Battle Ground, Bartley & Pontius 489 (Ny); Davidson Co., High Rock, 
Schallert 8660 (puKE); Durham Co., about 4 mi. n.e. of Durham, Wilbur 2899 
(micu); Forsyth Co., without exact locality, Schallert, 18 Aug. 1940 (MO, UARK, 
wis); Granville Co., Oxford, Godfrey 5464 (DUKE, GH); Johnston Co., 0.5 mi. 
n. of Cox's Mill, Foz 4942 (micn); Orange Co., Hillsboro, Gray & Carey, July 
1841 (au); Randolph Co., Asheboro, Hood 2399 (rLAs); Rowan Co., vicinity 
of Salisbury, Heller 130 (F, Mo, NY, PENN); Wake Co., just w. of Morrisville, 
Godfrey 49518 (DUKE, FLAS, GA, GH, MICH, NCS); Wayne Co., Goldsboro, 
Burlingame, July 1887 (BRU). SOUTH CAROLINA: Aiken Co., Vancluse, 
Eggert, 6 Aug. 1898 (Mo); Anderson Co., Long Branch Church, Davis, 29 July 
1919 (mo, TEX, US); Beaufort Co., Bluffton, Mellichamp, 1884 (r, vs); Berkeley 
Co., 8 mi. s. of Monks Corner, Wiegand & Manning 2557 (mixed sheets) (cv, 
aH); Georgetown Co., 12 mi. n. of Georgetown, Godfrey & Tryon 26 (DUKE, 
F, GH, MO, NY, US); Jasper Co., Ridgeland, Mohr, 1893 (mo); Lexington Co., 
8 mi. s. of Columbia, Godfrey & Tryon 1229 (mixed sheet) (xv); Williamsburg 
Co., Lanes Station, Burlingame, 20 July 1894 (BRU). GkoRGIA: Emmanuel 
Co., near Graymont, Harper 990 (au, wv, vs); Gwinnet Co., between the 
Alcovy River and No Business Creek, Small, 14 July 1893 (F, GH, MO, NY, US); 
Johnson Co., 2.5 mi. w. of Wrightsville, Pyron & McVaugh 3083 (GA); Mitchell 
Co., few miles e. of Pelham, Thorne 5078 (cu); Newton Co., 4 mi. n. of Coving- 
ton, Pyron & McVaugh 3036 (G^, NA); Sumter Co., without exact locality, 
Harper, July 1897 (Ny); Telfair Co., McRae, Biltmore Herb. 4511 (us); Wash- 
ington Co., 3 mi. n. of Harrison, Pyron & McVaugh 3104 (NA, NY). FLORIDA: 
Bay Co., Panama City, Knight, 6 July 1943 (Frias); Calhoun Co., without 
exact locality, Chapman (Mo); Dixie Co., near east limits of Jena, Beaman 280 
(mic); Franklin Co., Apalachicola, Chapman (au); Gulf Co., north of Port 
Saint Joe, Small, DeWinkeler & Mosier, 11 July 1924 (DUKE, FLAS, GH, MICH, 
MO, NCU, NY, PENN, TENN, WVA); Levy Co., Rosewood, Garber, June 1876 
(BRU, F, NY); Hamilton Co., near Jennings, Bright 3902 (wis); Taylor Co., 
between Perry and the Gulf of Mexico, Small, Small & DeWinkeler 11452 
(Ny, vs); Wakulla Co., prope St. Marks, Kugel May 1843 (mo, us); Walton 
Co., Freeport Mohr, 18 June 1880 (vs). 


1955] Wilbur,—The North American Genus Sabatia 29 


5. Sabatia brachiata Ell., Sk. Bot. S. C. & Ga. 1: 284. 1817.  Chironia 
angularis var. B angustifolia Michx., Fl. Bor. Am. 1: 147. 1803. Sabbatia 
concinna Wood, Class-Book ed. 2. 451. 1847. S. angustifolia (Michx.) 
Britton, Mem. Torr. Club 5: 259. 1889. 


Erect annual (10-)25-50(-67) em. tall, usually but one stem arising 
from each rosette, more rarely with 2 or 3. Branching usually restricted 
to the upper one-third or half of stem but in more robust plants with 
branches sometimes developing from the lowermost nodes, usually rather 
strongly ascendent forming an angle of 15-40 degrees with the main 
stem, rarely spreading to as much as 70 degrees, typically opposite, al- 
though rarely alternate along main stem or principal branches, or ultimate 
branches somewhat more commonly alternate, forming convexly corym- 
bose or somewhat pyramidal loose to compact crown. Stem 1-2(-4) 
mm. in diameter, terete, conspicucusly so below and smooth, not wing- 
angled, although sometimes noticeably lined or finely ridged. The stem 
above and upon the branches becoming subquadrate or even quadrate 
and bearing very slight wings at the angles. Roots few to numerous, 
slender, fibrous, spreading or descendent, about 4-8(-10) em. long, 0.5-1.5 
mm. in diameter. Basal rosette typically present at time of flowering, 
usually conspicuous, typically composed of numerous overlapping, spread- 
ing, usually broadly spatulate, rarely elliptie, obtuse to seldom acute 
leaves tapering strongly to an almost petiolate base, (8-)15-30(—45) 
mm. long, (6-)10-14(-18) mm. wide. Cauline leaves membranous, 
ascendent, 3-nerved or rarely only I-nerved in smaller leaves, (1—)1.5-3 
(—4.5) em. long, (3-)4-10(-16) mm. wide, often obtuse, especially below, 
or acute, commonly slightly callose-apieulate, oblong, with more or less 
parallel margins, or more rarely elliptie or somewhat lanceolate, tapering 
somewhat to base, at least usually not strongly or broadly clasping. 
Internodes usually about 1.5-3 times longer than the leaves, but ranging 
from but about three-fourths to about 4-times as long as the leaves. 
Inflorescence of corymbosely to pyramidally arranged cymules either 
1, both, or none of whose lateral branches may be surpressed.  Pedicels 
5-sided, noticeably thin-ribbed, (1-)2-8(-13) mm. long. Calyx-tube 
thin, noticeably but finely 5-ribbed, usually about half as long as the 
corolla-tube, (1—-)1.5-3(-4) mm. long, turbinate to campanulate. Calyx- 
lobes thin, narrowly linear, (4-)7-10(-15) mm. long, 0.3-1.0 mm. wide, as- 
cendent in bud, wide-spreading at anthesis, usually exceeded by the corolla 
lobes by 3-8 mm.  Corolla-tube (3-)4-5(-6) mm. long, 1.5-3 mm. in di- 
ameter, cylindrical, greenish within and withcut. Corolla-lobes oblong, or 
narrowly to broadly spatulate, usually obtuse, or occasionally tapering 
to an acute apex, (5-)7-14(-20) mm. long, (2-)3-6(-8) mm. broad, wide- 
spreading at anthesis, pale-pink to darker roseate, rarely white, with 
low triangular greenish-yellow area at base of lobe, usually bordered 
by reddish line. Filaments slender, pale yellow to nearly colorless, 
(1.5-)2-3(-4) mm. long; anthers bright yellow, usually about (2-)3(-4) 
mm. long. Stigmatic lobes slender, (2-)3-5(-7) mm. long; style 2-4 mm. 
long, usually less than the stigmatic branches in length. Capsule cylin- 


30 Rhodora [Vor. 57 


drical, 5-8 mm. high, 3-4.5 mm. in diameter. TYPE LOCALITY: “Grows 
in the middle and upper country of Carolina. Near Columbia," Lexing- 
ton Co., South Carolina. type: Herbemont s. n. (CHARL!). DISTRIBU- 
TION: Fields, dry and open oak and pine woods ranging from south- 
eastern Virginia south into Georgia and westward to southern Missouri 
and Louisiana. Map 5. 

This well-marked species has been rarely confused with any 
other, except occasionally with specimens of S. angularis or 
S. quadrangula (which for almost a century has been called 
S. paniculata). Both of these last-mentioned species are, however, 
readily distinguished from S. brachiata by their strongly tetra- 
gonal stems whose angles are strikingly membranous winged. 
S. brachiata is strongly contrasted to those two species by pos- 
sessing, especially below, a smooth, terete stem. Above and on 
the branches the stem is somewhat finely ridged and angled 
but not conspicuously so. The corolla of S. brachiata is typically 
pink or roseate and very robust specimens, especially in areas 
where 5. angularis is more commonly to be expected, have 
been rather often mistaken for that wide-spread species. Albinos 
of S. brachiata are rarely encountered, but when they are found 
or when the corollas become entirely bleached as they often 
do in drying, these plants are often mistaken for what has 
previously been called S. paniculata (= S. quadrangula). The 
type of S. brachiata forma candida (Fernald & Long 6346), 
is not S. brachiata, which normally has a rose-colored corolla, 
but S. quadrangula (S. paniculata in the sense of recent authors) 
which always has a white corolla. Fernald’s misidentification 
of the type-sheet is easily demonstrated: the lower portion of 
the stem of the type is very distinctly quadrate with winged 
margins which is a distinctive feature of S. quadrangula. The 
only other specimen cited with the original description of this 
"form" was Seymour 33 which is definitely a specimen of S. 
brachiata. A new name will be required by those who feel 
it essential to designate formally such minor variations. 

Chironia angularis B augustifolia, of Michaux, has been as- 
signed to the synonymy of this species at least since the ap- 
pearance of the Synoptical Flora, where it was so listed. Not 
having seen an authentic specimen from Michaux's herbarium, 
it is only tentatively that I place the name in the synonymy of 
S. brachiata. The original characterization of the variety was 


1955] Wilbur,—The North American Genus Sabatia 3l 


very brief ("8 augustifolia: folis quasi lanceolatis; supremis 
etiam linearibus") and certainly states nothing that would 
exclude forms of S. angularis from consideration or indicates 
clearly that the entity described was S. brachiata. Both this 
variety of Michaux's and his latifolia, which has been considered 
the sole element of Michaux's concept that is S. angularis as 
now interpreted, were arranged under the generalized diagnosis 
of the species. The stem in that account of the species is 
stated to be “marginato-quadrangulo” and this would seem 
to exclude the entity now known as S. brachiata. Perhaps 
Gray saw an authentic specimen at Paris and hence may be 
correct in his treatment; in any event the question is yet to be 
settled. 

S. concinna Wood is assigned to the synonymy of this species 
upon the basis of the original description which led Gray as 
early as 1856 to the same conclusion. It is perhaps of some 
significance that Wood himself later listed the name as a syn- 
onym of S. brachiata. The oXginal description follows: 


3. S. concinna. Wood (Nov. sp.) Elegant Star Flower. St. slender, sub- 
quadrangular, internodes 2—4-times longer than the leaves; branches oppo- 
site, sub-erect; lvs. linear and lance-linear, lower ones ovate, all acutish, 
sessile, panicle oblong; cal. segments linear, twice longer than the tube, 
twice shorter than the corolla; cor. 5-parted, segments oblong-obovate, 
obtuse, light purple.—Dry grassy prairies, Ia.! abundant. Stem a foot 
high, few or many-flowered. Leaves 9-12" by 1-3", flowers 15" diam., of 
a delicate blush purple, the star in the center yellow, bordered with green. 
Jl. Aug. 


Merrill (RHopora 50: 127. 1948.) concluded that “it is the 
same as S. campestris Nutt., not S. brachiata Ell. as the latter 
species does not occur in Iowa; Wood's type was from Iowa." 
However, Wood described the branches as opposite and the 
calyx-segments as linear and only half as long as the corolla 
and these features would exclude from consideration anything 
but an unusual specimen of S. campestris. If'the plant were 
S. campestris, it would hardly seem possible that the very peculiar 
and prominent costae of the calyx-tube could be overlooked 
in even the most cursory examination. I have seen no specimen 
of S. campestris from Iowa. Merrill placed considerable weight 
of evidence upon the locality of the plant but his interpretation 
of the abbreviation “Ia.” is certainly an error. In 1846 between 


32 Rhodora [Vor. 57 


the first and second editions of his Class-Book, Wood, enlarging 
the area to be covered by his flora, made a short collecting trip 
to Indiana. S. concinna was presumably seen upon that trip. 
The explanation of geographical abbreviations appearing in 


the second edition informs one that “the names of . . . states 
. are often abbreviated, and always in the same manner as 
in other works; thus . . . Ia. or Ind., Indiana, &c." In this 


same edition (1847, p. 3) the area of the flora was said to be 
"essentially the States lying north of the Ohio River and Mary- 
land." One also learns in Wood's ‘American Botanist and 
Florist" that the “geographical limits of the present flora are 
the same as those adopted in the Class-Book: viz, all the States 
of the American Union lying east of the Mississippi River." 
Although the area covered by Wood's publications was expanded 
in later years, they were never intended to cover the region 
west of the Mississippi River except indirectly. Wood meant 
Indiana by the abbreviation “Ia.” Fernald (Rhodora 47: 404— 
405. 1945) concluded as much from indirect evidence. 

S. brachiata is not represented from Indiana in any of the 
herbaria that I have studied and neither it nor S. campestris 
were included by Deam in his Flora of that state. In fact, 
Deam (1940, p. 1080) after considering the evidence and after 
finding that that species was not represented in either the Gray 
Herbarium or in that of the New York Botanical Garden defi- 
nitely excluded it from the known flora of the state. Fernald 
(1950) lists the range of the species as including southern Indiana. 
S. brachiata is not represented from Indiana, Illinois, or Kentucky 
in any of the collections that I have studied. The nearest 
known stations are southeastern Missouri and in Tennessee. 


The description of the stem as being ''subquadrangular" 
is also a bit puzzling if the name really is synonymous with 
S. brachiata, for that species is very conspicuously marked by 
a decidedly terete stem in at least the lowermost portion. The 
upper part of the stem and the branches do become rather 
strongly angled and, if S. concinna is the same as S. brachiata, 
this explanation probably accounts for that at first puzzling 
statement in the description. The disposition of Wood’s name, 
in the absence of authentic material, will remain tentative, 
but the evidence available indicates that it was S. brachiata. 


1955] Wilbur,— The North American Genus Sabatia 33 


REPRESENTATIVE SPECIMENS:—VIRGINIA: Princess Anne Co., Little Neck, 
Fernald & Long 4134 (GH, PENN); Prince George Co., just s. of Disputanta, 
Fernald & Long 8815 (GH); Sussex Co., northeast of Homeville, Fernald & 
Long 6344 (GH, NY, PENN, US). NORTH CAROLINA: Anson Co., 4 mi. n. of 
Ansonville, Boyce 1127 (Ncs); Beaufort Co., 7 mi. s. of Washington, Godfrey 
4400 (GH, US); Brunswick Co., 3 mi. n. of Bolivia, Wilbur 2888 (micu); Bladen 
Co., without exact locality, Biltmore Herb. 4511a (GH, MO, NCU, NY, PENN, US); 
Carteret Co., 3 mi. se. of Newport, Godfrey 48192 (Ncs); Cumberland Co., 
7.5 mi. n. of Fayetteville, Godfrey & For 49446 (DUKE, Ncs); Harnett Co., 
6 mi. e. of Cameron, For 2546 (ncs); Montgomery Co., 1 mi. n. of Ether, 
Wiegand & Manning 2543 (cu); Moore Co., 2 mi. s. of West End, Fox & 
Whitford 3903 (mica, Ncs); New Hanover Co., Wilmington, McCarthy, Aug. 
1885 (vs); Onslow Co., at Richlands, Godfrey 4475 (an, vs); Sampson Co., 
Roseboro, Godfrey 4532 (DUKE, GH, US); Scotland Co., 12 mi. n. of Laurinburg, 
Godfrey 5046 (DUKE, GH, US); Wake Co., Raleigh, Godfrey 4918 (GH, NCU, NY); 
Wilkes Co., up Pores Knob, Radford & Stewart 1743 (NCU). SOUTH CAROLINA: 
Chesterfield Co., near Cheraw, Ward, 25 June 1895 (Ny, us); Darlington Co., 
Society Hill, Canby, July 1878 (r, ny); Horry Co., without exact locality, 
Adams 30 (PENN); Lexington Co., 8 mi. s. of Columbia, Godfrey & Tryon 1229 
(GH, NY [a mixed sheet], vs). GEoRGIA: Bartow Co., 1.25 mi. e. of Emerson, 
Duncan 8548 (GA, MO); DeKalb Co., northern slope of Stone Mountain, Small, 
17 July 1893 (r, xv); Douglas Co., 10 mi. s. of Douglasville, Duncan 3648 
(micH); Pickens Co., near Jasper, Biltmore Herb. 4511a (us); Richmond Co., 
Augusta, Cuthbert, Aug. 1876 (RUT); Taylor Co., without exact locality, 
Pyron, 3 Aug. 1930 (DUKE). TENNESSEE: Coffee Co., near Manchester, 
Clebsch & Clebsch 4770 (TENN); Franklin Co., between Tullahoma and Estill 
Springs, Svenson 9211 (au); Grundy Co., east of Altamont, Svenson 7137 (aH, 
TENN); McNairy Co., without exact locality, Bain 431 (an, Ny); Van Buren 
Co., Falls Creek State Park, Shanks, Clebsch & Sharp 3431 (MICH, TENN); 
White Co., sw. of Clifty, Shanks, Clebsch & Sharp 2916 (TENN). ALABAMA: 
Mobile Co., about 8 mi. sw. of Mobile, Webster & Wilbur 3479 (mica); Wash- 
inton Co., Fruitdale, collector not stated, July 1904 (Mo). wississiPPI: Coving- 
ton Co., 1.5 mi. se. of Mt. Olive, Webster & Wilbur 3308 (icu); Forest Co., 
about 14 mi. s. of Hattiesburg, Webster & Wilbur 3397 (micu); Harrison Co., 
3 mi. s. of Saucier, Webster & Wilbur 3442 (mic); Jackson Co., Ocean Springs, 
Pollard 1083 (mo, NY, us); Jones Co., Laurel, Tracy 3356 (Ny); Pearl River 
Co., Poplarville, Tracy 1687 (Gu, vs); Pike Co., Holmesville, Wheeler (mo); 
Stone Co., 8 mi. s. of Wiggins, Webster & Wilbur 3440 (mica); Wayne Co., 
6 mi. nw. of Ala.-Miss. state line, Sargent, 18 June 1950 (OKL, NCS). MISSOURI: 
Butler Co., 12 mi. n. of Poplar Bluff Steyermark 11593, (MO). ARKANSAS: 
Faulkner Co., Conway, Haas 1629 (us); Drew Co., Ladelle, Demaree 22321 
(MO, OKLA); Lonoke Co., Grand Prairie, Demaree 22333 (MO, NY, OKL, OKLA); 
Prairie Co., DeValls Bluff, Demaree 22176 (mo); Pulaski Co., near Little Rock, 
Carpenter, June 1938 (UARK). LOUISIANA: Calcasieu Parish, Lake Charles, 
Daves, Aug. 1888 (F); Grant Parish, 4 mi. s. of Pollock, Webster & Wilbur 
3257 (MICH); Orleans Parish, New Orleans, Drummond 224 (an, x); Rapides 
Parish, 12 mi. se. of Hineston, Webster & Wilbur 3275 (micu); St. Tammany 
Parish, vicinity of Covington, Arséne 11741 (vs); Tangipahoa Parish, 6 mi. e. 
and 1 mi. n. of Hammond, Nease, 1945 (okL); Vernon Parish, 2 mi. w. of 
Leander, Webster & Wilbur 3228 (wicH); Washington Parish, Bogalusa, 
Cocks, 5 June 1917 (xo). 


(To be continued) 


34 Rhodora (Vor. 57 


ILEX GLABRA AND A NEW STATION FOR KALMIA LATIFOLIA 
IN New HawrsnuiRE.— The Inkberry, Ilex glabra (L.) Gray, 
has never been validly reported to occur naturally in New Hamp- 
shire. However, it has long been known to be present in Maine 
on Isle au Haut as well as in Nova Scotia. 

The frequency of stations for it in Essex County south of the 
Merrimack River, suggests that one might expect to find it 
in similar habitats along the coast in Rockingham County, 
New Hampshire. 

In October 1950 some fragments of Inkberry were mailed to 
me from Seabrook accompanied by a brief diseussion of the 
plants' whereabouts, the inference being that the station was 
within the boundaries of New Hampshire. 

Not until 1954, was it convenient to arrange a visit to the 
Ilex-colony. But in March of that year, though temporarily 
viewed with some quite undeserved suspicion as a vandal, I 
was permitted to see the somewhat straggling and recently 
molested colony, and was even encouraged to bring away a 
small living plant. The herbarium specimen which unfortu- 
nately is meagre, though quite adequate for verification, is 
deposited in the University of New Hampshire Herbarium. 
The colony definitely is in New Hampshire, but rather less than 
a half mile from Massachusetts. 

It will be interesting to watch the progress of this colony. 
Recent growth of shading Pinus Strobus in the immediate 
vicinity suggests vegetational changes perhaps detrimental 
to the plant’s growth. Possibly in earlier times the colony 
was more vigorous and extensive and what we have now is a 
mere remnant of a much larger stand. Out of respect for the 
wishes of the owners in the matter, more precise information 
concerning the plant's whereabouts will not now be published. 

Kalmia latifolia L. is one of numerous species which have 
mostly disjunet ranges in New Hampshire. Reaching the 
northeastern limit of its range in south-central and southeastern 
Maine, Mountain-laurel becomes increasingly more frequent 
as one approaches its more or less continuous area beginning in 
New Hampshire, west of the Merrimack River and including 
those upland townships which are adjacent to Massachusetts 
or which lie not many miles to the north. Unlike many other 


1955] Hodgdon,— Ilex glabra 35 


species of southern distribution, Kalmia latifolia does not extend 
northward appreciably in the Connecticut Valley. Fernald? 
summarizing botanical studies made in western Cheshire County 
during three summers, reported it from only one station. He 
described it as “Forming an extensive thicket by the Ashuelot 
River, Gilsum," while H. G. Jesup? mentioned only the well 
known Squam Lake Station. Kalmia latifolia is a normal 
constituent of the acid upland forests of southwestern New 
Hampshire becoming only occasional in the different type of 
forest of the Connecticut Valley. To the east of the Merrimack 
River, Mountain-laurel again is very localized in the dominant 
White Pine Region of southeastern New Hampshire. Specimens 
are represented in herbaria from favorable sites in southeastern 
New Hampshire where locally the plant is luxuriant, in the 
townships of Pelham, Windham, Sandown and Barrington. 
Doubtless it occurs elsewhere in this region but to most persons 
living there, Kalmia latifolia is either totally unknown as a 
native or is exceedingly rare. 

Farther northward in New Hampshire as in Maine the plant 
becomes less frequent. Reports from creditable observers have 
reached me of colonies in New Durham and Effingham. These 
I have not visited, but an herbarium specimen from Chase 
Hill in Albany, confirms the existence of a vigorous colony 
there with which I am familiar. 

In central New Hampshire, there is the isolated colony near 
Squam Lake in Holderness, evident from the highway and 
familiar to all. This I had always believed to be the most 
outlying stand in this part of New Hampshire. But in mid- 
July 1954, in company with Mr. Radcliffe Pike, I visited Peaked 
Hill Pond, in Thornton, Grafton County, to investigate a report 
of “Rhododendron” in the vicinity. A half-acre of ''Rhodo- 
dendron" near the southern shore turned out to be two or more 
acres of Mountain-laurel. The colony occurs in deciduous 
woods, with a scattered admixture of large hemlocks and is 
somewhat away from the water on the southern and southeastern 
sides of the pond. "Phe growth of laurel is noticeably dwarf, 
most of the plants not exceeding 3 or 4 feet in height, in marked 


1 RHODORA, 3: 235, September 1901. . . 
? A catalogue of the Flowering Plants and Higher Cryptogams found within about 
thirty miles of Hanover, New Hampshire, 26 (1891). 


36 Rhodora [Vor. 57 


contrast to some of the colonies in southeastern New Hampshire 
which contain plants averaging much higher than a man’s 
height. Here at nearly 1200 feet altitude in an area exposed 
to winds from the north the necessity for protection by deep 
snow in severe winter weather may account in some way for 
the low stature of the plants. Peaked Hill Pond is rather less 
than 30 miles northeast of Hanover and not more than 15 
miles, as the crow flies, from the area of Kalmia latifolia at 
Squam Lake. Specimens from Thornton are deposited in the 
Herbaria of the University of New Hampshire and the New 
England Botanical Club.—A. R. HopGDON, UNIVERSITY OF NEW 
HAMPSHIRE. 


A NEw VARIETY OF SOLIDAGO ULMIFOLIA.—Solidago ulmifolia 
Muhl., var. palmeri Cronq. var. nov. Caulibus pilis patentibus 
obtectis. 

TYPE: Palmer 24111, shaded sandstone slopes, north side, near top of 
Magazine Mt., Logan Co., Arkansas, October 14, 1923; deposited at the New 
York Botanical Garden. ADDITIONAL SPECIMENS: Arkansas: Garland Co., 
Palmer 29092, 29194; Demaree 11001, 20498, 20512, 21836. Logan Co.: 
Palmer 24180; Demaree 8072. Perry Co.,: Demaree 20145. Pope Co.: 
Demaree 19876. Yell Co.: Demaree 15942. Alabama: County undetermined: 
Buckley s.n., October, 1838, without further data; Biltmore Herb. 15008, 
Avondale. 

The var. ulmifolia, with the stems essentially glabrous below 
the inflorescence, is largely replaced in the Ouachita region of 
Arkansas by the var. palmeri. The existence of two specimens 
of var. palmeri from Alabama in the herbarium of the New York 
Botanical Garden indicates that the hairy-stemmed phase of 
the species is not purely local, however, and the limits of its 
range remain to be determined. 

Several of the specimens (including the type) here cited as 
S. ulmifolia var. palmeri were annotated by K. K. Mackenzie 
with an apparently unpublished binomial using the epithet 
palmeri in conjunction with the genus Solidago. "The treatment 
of the new variety was inadvertently omitted by me from an 
earlier paper dealing with eastern American goldenrods, with 
the result that the name as used on page 428 of volume 3 of 
Gleason’s Illustrated Flora is technically a nomen nudum.— 
ARTHUR CRONQUIST, THE NEW YORK BOTANICAL GARDEN. 


Volume 56, no. 672, including pages 261-286, was issued 10 January 1956, 


Dodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club. by 
REED CLARK ROLLINS, Editor-in-Chiet 


ALBERT FREDERICK HILL ~. 
STUART KIMBALL HARRIS l 

RALPH CARLETON BEAN Associate Editors 
RICHARD ALDEN HOWARD ( 

CARROLL EMORY WOOD, JR. 


Vol. 57 February, 1955 No. 674 
CONTENTS: 

Studies in the Hippocastanaceae, I. Variation within the Mature 
Fruit of Aesculus. James W. Hardin..................... 37 

A Revision of the North American Genus Sabatia (Gentianaceae). 
Robert L. Wilbur (continued from page 33)................. 43 

Epifagus virginiana. Samuel C. Palmer........... cee cece cece 71 


Calycera balsamitaefolia in the United States. Julian A. Steyer- 
mark. and Floyd A. Swink... reaa ua aiena raah 72 


The New England Botanical Club, Jne. 


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Botanical Museum, Oxford St., Cambridge 38, Mass. 


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Rbodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 57 February, 1955 No. 674 


STUDIES IN THE HIPPOCASTANACEAE, 
I. VARIATION WITHIN THE MATURE 
FRUIT OF AESCULUS 


JAMES W. HARDIN! 


A wIDE variation in fruit size and shape is readily noticeable 
from observations of an Aesculus tree or shrub in late summer 
(Fig. 1). Upon closer examination it becomes apparent that 
this variation is due in part to the number of maturing seeds in 
the ovary. 

The ovary is typically 3-locular and 3-carpellate with two 
ovules in each locule. These ovules are oriented differently, 
the lower descending with a dorsal raphe, and the upper hori- 
zontal or more often ascending with a ventral raphe. Each 
fruit, therefore, is potentially six-seeded. 

The literature is not clear in regard to the number of seeds 
usually found in the mature fruit. Gray (1849) stated that 
the locules are one seeded by the abortion of one ovule in each, 
and by suppression the fruit is often 1- to 2-celled and 1- to 2- 
seeded. Pax (1895) said that the mature fruit is mostly 1- 
seldom 2-locular with one seed. Rendle (1925) also stated 
that only one seed per fruit develops due to the crushing of 
two out of three cells by the considerable growth of the one 
ovule. Chapman (1897) on the other hand described the fruit 
as being l- to 3-seeded. The more recent literature presents 
the same information: Fernald (1950)—3-seeded or usually 
by abortion 2- or I-loeular and 2- or 1-seeded; Lawrence (1951)— 
usually 1-loeular and I-seeded; Gleason (1952)—seeds by 


! My sincere thanks go to Professor Rogers McVaugh, University of Michigan, for 
his helpful suggestions during the preparation of the manuscript. 


38 Rhodora [Vor. 57 


abortion only one and subglobose, or 2 and semiglobose, or 
rarely 3 with flattened sides. Summarizing this information 
then, one might expect to find usually one, or possibly up to 
three, seeds per fruit. 

Since there seems to be as much variation in the literature 
as in the fruit, and more than three seeds per capsule are often 
found, it was thought necessary to make some actual counts 
in conjunetion with a much broader study of the entire genus. 

Collections of fruit from six species were made during the 
summer of 1954 and studied from the point of view of the num- 
ber of mature seeds, the position of these seeds within the 
capsule and the number of locules present. "The result of this 
analysis is presented here. 

Each fruit was examined and the number of mature seeds 
and their position recorded. In Table 1, "top" and “bottom” 
refer to the ascending and descending ovules respectively. The 
fifteen position-combinations were determined from the abortive 
ovules more easily than from the mature ones since the former 
more nearly retain their original position after the great en- 
largement of the seeds. More combinations could be recognized 
if, e.g. with 3 and 4 seeds, it were determined whether one or 
both ovules mature within the same locule. This information 
was kept only for 2-seeded combinations—both seeds in the 
same or different locules. Also given in Table 1 is the number 
of fruits found with a particular combination and the percentage 
based on the total fruits examined for each species. 

From three trees of A. hippocastanum L. growing on the 
University of Michigan Campus and from two trees along a 
highway in Franklin County, Ohio, a total of 107 fruits was 
collected and examined. Table 1 indicates that the 1-seeded 
fruit was the most common type and that the ratio of “top” 
to *bottom" was essentially 1:1. In the 2-seeded type the most 
frequent combination was the 1 — 1 diff., meaning one ascending 
and one descending seed but in different locules. 

An analysis was made of 153 fruits of A. glabra Willd. which 
were collected from a small population in Washtenaw County, 
Michigan. The l1-seeded capsule was again found to be the 
most common (Table 1) and maturation of ovules occurred in 
essentially the same numbers for ascending and descending. 


Plate 1205 


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39 


in the Hippocastanaceae 


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40 Rhodora [Vor. 57 


Aside from percentages which differ between A. hippocastanum 
and A. glabra, the most obvious difference is the occurrence 
in A. glabra of fruit with four, five and six mature seeds. Only 
one fruit each of the 5- and 6-seeded type was found, but it is 
significant that this condition does occur, even though seemingly 
rare for the species as well as the entire genus. 

Fifty five capsules of A. octandra Marsh., from a population 
in Haywood County, North Carolina, were studied. Over 
half were of the 1l-seeded type, although 2-, 3- and 4-seeded 
forms were found in decreasing frequencies. 

A total of 145 capsules of A. sylvatica Bartr. were examined 
from two populations, one from Clarke County, Georgia and 
the other introduced from Georgia and now growing at the 
University of Michigan Botanical Garden. Again there was 
a high percentage of l-seeded fruits, but also relatively high 
percentages of 2- and 3-seeded forms were found. 

A very small and probably clonal population of A. parviflora 
Walt. was visited in Early County, Georgia during June 1954. 
Mature fruits were not available, but a population sample 
of partially mature fruits was collected. Analysis of these 
showed only the one-seeded type. This lack of variation is 
of interest since the population is probably one clone which 
occuples an area of nearly 400 square feet and includes approxi- 
mately 75 trees. 

Sixteen capsules were kindly sent from a small population 
of what is tentatively considered A. discolor Pursh growing 
near Old Sutherland Springs, Wilson County, Texas. It was 
unusual to find no 1-seeded fruits among this collection. Over 
half were of the 2-seeded type and the rest 3- and 4-seeded. 

Considering all six of these species together, percentages are 
given (Table 1) for the genus based upon the random samples 
totaling 504 capsules. The emphasis which previous authors 
have placed upon the 1-seeded type is not justified on the basis 
of these figures which show that this form is present in barely 
more than one half of the total. Over one quarter of all fruit 
was of the 2-seeded form and just under one eighth had three 
seeds. "The most significant deviation from the literature is the 
nearly 4% of the total which contained four to six seeds. 

Correction should be made of the error found in the literature 
regarding the interpretation of the number of locules present 


1955] Hardin,—Studies in the Hippocastanaceae 41 


in the mature fruit. Regardless of the number of seeds maturing, 
and the crushing of the locules by the excessive growth of the 
seeds, each of the locules remains at least partially distinct 
with its abortive ovules in their original position. The typical 
fruit then, is always 3-locular regardless of whether 1-seeded 
or 2- to 6-seeded. 

Exceptions to this 3-locular condition were found in two of 
the species. Four-carpellate, 4-locular and in maturity 4-valved 
fruits of A. hippocastanum were found among the collections 
(Fig. 2). Also 2-locular and 2-valved fruits of A. sylvatica 
were found (Fig. 3). Of the 107 capsules of A. hippocastanum 
examined, thirteen of them, or 12%, were 4-locular. The 
majority of these were of the l-seeded type, only a few having 
2 seeds. Each of the four locules was distinct at maturity and 
each contained the expected two ovules. Only two capsules 
of A. sylvatica out of the 145 examined were 2-locular. One 
of these was l-seeded and the other 2-seeded. The expected 
total of four ovules was present in both cases. The anomalous 
condition of 2 or 4 locules is rare, or at least infrequent in A. 
hippocastanum. Just what evolutionary significance these 
conditions have, if any, is not yet known. 

At the outset of this study, the question arose as to whether 
only the ascending or descending ovules matured, or if both, 
in what proportions. The answer is shown in Table 1. A 1:1 
ratio was found in the 1-seeded forms. In the 2-seeded and 
3-seeded capsules only slightly more of the ascending ovules 
matured. These ratios show that the position of the maturing 
ovule in respect to ascending or descending is entirely random, 
and abortion is not due to a morphological degeneration of a 
portion of the ovary or placenta. 

The cause of abortion in the majority of ovules is not known. 
According to Gray (1849) and Rendle (1925) the pressure from 
the maturing ovules is the important factor. If this were true, 
one would expect to find some degree of enlargement in all six 
ovules. Fig. 4 illustrated one of the few cases seen where pressure 
was effective in retarding the growth of ovules. In this case 
the lower two ovules did enlarge greatly until presumably 
stopped by pressure from the top. The smaller two were not 
true abortives since they expanded to full size after the capsule 
was opened. In practically every other case, the abortive 


42 Rhodora [Vor. 57 


ovules were 1-4 mm. in diameter—the size being nearly constant 
within a single capsule. Even when one mature seed was found 
in a locule, the second and abortive ovule in that locule showed 
no signs of being crowded. The “pressure” theory then, does 
not seem to be the answer in the majority of cases. 

Incomplete fertilization could possibly be the determining 
factor. If this were true, then the unfertilized ovules would not 
ordinarily enlarge, which is found to be the case in these capsules. 
This incomplete fertilization could arise because of an incomplete 
pollination, or more probably caused by some morphological, 
physiological or genetic sterility factor. The answer to this 
question involves a major research problem in itself. 

The phylogenetic significance of seed number in the genus 
Aesculus is not completely understood at this point. If one 
extreme in number (l-seeded or 6-seeded) is more primitive 
than the other in terms of evolution of the family, genus or 
species, the interpretation may be made only after a more 
complete study of the related taxa within the Sapindales. 


DEPARTMENT OF BOTANY, UNIVERSITY OF MICHIGAN, 


LITERATURE CITED 

CHAPMAN, A. W. 1897. Flora of the Southern United States. Ed. 3. 
Amer. Book Co., N. Y. 

FrERNALD, M. L. 1950. Gray’s Manual of Botany. Ed. 8. Amer. Book 
Co., N. Y. 

Gurason, H. A. 1952. The New Britton and Brown Illustrated Flora of 
the northeastern United States and adjacent Canada. Vol. II. Lan- 
caster Press, Pa. 

Gray, Asa. 1849. The genera of the plants of the United States. Vol. II. 
George Putnam, N. Y. 

Lawrence, G. H. M. 1951. Taxonomy of Vascular Plants. Macmillan 
Co., N. Y. 

Pax, F. 1895. Hippocastanaceae. In Engler and Prantl, Natürl. Pflanz- 
enfam. III (5): 273-276. 

RENDLE, A. B. 1925. The Classification of Flowering Plants. Vol. II. 
Cambridge. 


1955] Wilbur,—The North American Genus Sabatia 43 


A REVISION OF THE NORTH AMERICAN 
GENUS SABATIA (GENTIANACEAE) 


Rosert L. WILBUR 
(Continued from page 33) 


C. Subsection campresTREs Blake, Ruopora 17: 56. 1915. 


Neurola Raf., New Fl. 4: 92. 1838. 

Annuals, branches typically alternate, at least above. Basal rosette 
never present. Leaves either thin and membranous or thick and suc- 
culent, margins not scarious. Calyx-tube conspicuously ribbed or 
costate by the prominent development of the fused lateral traces and 
bearing a thin membranous wing. TYPE SPECIES: Sabatia campestris Nutt. 


This subsection contains but two species which are obviously 
closely related to one another and which are strongly and unique- 
ly distinguished from the other groups by the prominently 
developed calycine costae. In addition, the typically alternate 
branching pattern sets this small species-group apart from 
subsection Angulares, to which it is probably most closely related. 
Also, the Angulares possess a very well-developed basal rosette 
at some stage in their life-cycle although it is not always per- 
sistent at time of flowering; the Campestres are never rosulate. 
An additional indication, besides the aspect of the plants, the 
texture of the leaves, and perhaps the habit, of the relationship of 
this group to the opposite-branched Angulares is the pronounced 
quadrate stems in the species of both subsections (at least above) 
which are margined along the angles by a thin, hyaline membrane. 

The distribution of the Campestres is almost entirely west 
of the Mississippi River where the subsection occupies an ex- 
tensive area. S. campestris accounts for the bulk of the sub- 
sectional range. Sabatia arenicola is very obviously related 
to S. campestris and is believed to have been derived from that 
species or from its precursor. S. arenicola is at present known 
only from the Quaternary littoral deposits along the beaches 
from Louisiana to southern Tamaulipas. 


KEY TO THE SPECIES OF SUBSECTION CAMPESTRES 


A. Leaves and calycine lobes thin and membranous, neither succulent nor 
heavily cutinized; leaves broadest near the base, usually strongly clasping 
the stem, the midvein at least prominently elevated beneath; calyx-lobes 
usually 2-6 times longer than the calyx-tube, generally 4-8 times longer 
than broad; corolla-lobes typically equaling or exceeding the calyx-lobes, 


44 Rhodora [Vor. 57 


usually longer than 1 em. and wider than 7 mm.; stigmatic lobes over 5 
mm. long; anthers longer than 2 mm.; widespread in prairies and fields 
from Texas to Mississippi and north to Illinois and Kansas. 
6. S. campestris. 
A. Leaves and calycine lobes thick and succulent, heavily cutinized; leaves 
broadest above the base, tapering somewhat to the sessile, nonclasping 
base, venation obscure and flush with the surface; calyx-lobes usually less 
than twice the length of the calyx-tube, generally less than 4 times as long 
as broad; corolla-lobes usually equaled or exceeded by the calyx-lobes, the 
corolla-lobes usually less than 1 em. long and narrower than 7 mm.; 
stigmatic lobes less than 5 mm. long; anthers shorter than 2 mm.; restricted 
to the strand and dunes from Louisiana southward into Mexico. 
7. S. arenicola. 


6. Sabatia campestris Nutt., Trans. Am. Phil. Soc. n. s. 5: 197. 1835. 
S. nervosa Raf., New Fl. 4: 92. 1838. Neurola arkanzica Raf., l. c., 
an alternate name. Sabbatia formosa Buckl., Proc. Acad. Phila. 1862: 7. 
1863. S. campestris f. albiflora Moore, Proc. Ark. Acad. Sci. 1: 26. 1941. 

Erect annual (6—15-30(-40) em. high; stems 1-3 mm. wide, strongly 
quadrate throughout, the corners winged by a thin hyaline membrane, 
0.1-0.2 mm. high. Branches usually few or absent from the lower half 
of the stem, but when present sometimes opposite, typically alternate 
throughout, usually widely divergent, often forming angles of 60-90 
degrees with stem, giving plant a loose, strikingly divaricate or even 
geniculate aspect or, more rarely, forming a fuller, more compact, corym- 
biform crown, or in unfavorable situations sometimes unbranched. 
Root system of few to several fibrous roots or of a slender, wiry tap-root 
usually 5-10 em. long bearing several slender laterals. Leaves thin, 
membranous, neither succulent nor heavily cutinized, primary venation 
apparent even in living material and at least the midrib noticeably 
elevated on the lower surface, conspicuously 3-5-nerved in dried speci- 
mens. Lowermost leaves rarely conspicuously different in either texture 
or shape from those at the middle or upper nodes, except usually some- 
what smaller in size, rarely persistent at anthesis; or very rarely quite 
different in being broadly elliptie to oblong, tapering to the base and 
up to 8 em. long, above narrowly lanceolate to broadly so, or ovate- 
lanceolate, rarely somewhat oblong or even ovate, typically broadest 
at or very near the base, generally broadly clasping, usually those at 
the lower nodes obtuse while the upper are acute, the tip callose-mucronu- 
late, (0.8-)1.5-3.0(-4.0) em. long, (0.3-)0.8-1.5(-2.0) em. wide, usually 
about (1-)1.5-2(-3) times as long as broad. Internodes (1-)2-5(-10) 
em. long, usually about 1.5-4 times the length of the leaves. Inflorescence 
usually of loosely-arranged, divergent, reduced, 1-2-flowered cymules 
forming a more or less open corymbose cluster. Peduncles (or pedicels) 
wiry, rigid, pentagonal, (1-)2-5(-10) em. long, bearing fine hyaline 
membranous wings at the angles. Calyx-tube broadly campanulate, 
usually about 1-1.5 times as long as broad, (3-)4-6(-8) mm. long; strongly 
pentagonal with 5 thin, membranous costae or ribs elevated about 1 mm. 
with fine hyaline margins. The intercostal tube very thin, membranous, 


1955] Wilbur,—' The North American Genus Sabatia 45 


translucent, appearing green due to ovary within. Calyx-lobes thin, 
membranous, not heavily cutinized, strongly spreading at anthesis, 
(1.5-)2-4(-6) times longer than the tube, acute, apiculate, linear, (0.4—) 
1.0-2.2(-3.2) em. long, usually 4-8 times longer than broad, strongly 
resembling the leaves in texture. Corolla-tube usually 1-2 mm. longer 
than the calyx-tube, colorless except for the extension of the yellow 
"star" patches in the uppermost portion. Corolla-lobes typically equaling 
or exceeding the calyx-lobes, occasionally shorter, ascendent to rather 
strongly spreading, oblong, elliptical, or most commonly broadly obovate 
or spatulate, acute to obtuse, (1.0-)1.3-2.0(-2.3) em. long, (0.4-)0.6-1.3 
(—1.5) em. wide, rose to pale pink or rarely white with a 3-4 mm. by 1-1.5 
mm. greenish-yellow patch at the base of the lobe, bordered by a white 
area and sometimes by a more densely roseate region. Anthers bright 
yellow, narrowly linear, 2.5-5(-6) mm. long; filaments white to pale 
yellow, slender, 3-6 mm. long. Style slender, white to pale yellowish, 
3-4 mm. long; stigmatie branches greenish at first, turning yellow at 
maturity, 5-8 mm. long. Capsules 5-9 mm. long, either equaling and 
included within the calyx-tube or exserted about 2-4 mm. TYPE LOCALITY: 
“in the open prairies of Arkansas and Red River." Type: Nuttall's 
collection, presumably in the Herbarium of the Philadelphia Academy 
of Science, but not seen. DISTRIBUTION: Fields and prairies from Illinois 
to Kansas south throughout most of the eastern half of Texas and east- 
ward to Mississippi. Very rarely introduced farther east. Map 6. 


This species is easily recognized by its prominently ribbed 
calyx-tube together with its thin, membranous leaves. 

Nuttall's type, or authentic material known to have been 
in his possession when the original description was made, has 
not been seen by me but his diagnosis is so ample as to leave no 
doubt as to the plant he had at hand. As yet but four species 
of the genus are known from Arkansas and of these Nuttall's 
excellent description fits but one. The original collection is 
stated to have been made “in the open prairies of Arkansas 
and Red River." 

If one were to accept the date of publication that appeared 
with the works of Nuttall and Rafinesque describing this species, 
a different conclusion than that adopted here would be reached 
as to which name has priority. Nuttall’s “Collections towards 
a Flora of the Territory of Arkansas" appeared in Vol. 5 of 
the “Transactions of the American Philosophical Society." 
It has therefore been dated as 1837, the year that that volume 
was apparently completed. Foster (RHopomRA 46: 1506-157. 
1944.) has pointed out that that date is certainly later than 
the publication and distribution of Nuttall's contribution. 


46 Rhodora [VoL. 57 


W. J. Hooker was familiar with Nuttall's account and cited 
it and descriptions provided in it on several occasions. The 
original diagnosis of S. campestris was quoted by Hooker (Comp. 
Bot. Mag. 1: 171. 1836.) and its place of publication cited to 
the page. Since Sprague (Kew Bull. 1933: 362-364.) has 
provided very precise dates for the publication of the various 
parts of Hooker's "Companion to the Botanical Magazine," 
it is certain that the whole of Nuttall's Flora, and not just the 
first twenty-one pages, as implied by Foster, was published and 
had reached England some time prior to the publication of 
Vol. 1, No. 6, of the Companion, which appeared January 1, 
1836. The four parts or volumes of Rafinesque's New Flora 
were issued, not in 1836 as stated on the title page, but at various 
dates between very late in 1836 and late in 1838. This has been 
demonstrated rather conclusively by Barnhart (Torreya 7: 
177-181. 1907.). The names of Sabatia appeared in the part 
which appeared in 1838. 

The Rafinesquian names here considered as synonymous 
with S. campestris were all published in one paragraph and were 
intended for the same entity. His account is quoted in full as 
it seems the surest way to prove the identity of the plant de- 
scribed by him and also to explain his peculiar nomenclature. 

975. Sabbatia ? nervosa Raf. ramose pauciflore, branches anceps, leaves 
ovate lanceolate trinerve acute, peduncles axil. and terminal, calix base 
campanulate 5 gone, segments linear elongate nervose, corolla as long as calix, 
segments broad obovate, trinerve at base—anonymous sp. of Nuttal, which 
deserves perhaps to be a genus Neurola Raf. by calix 5 gone corolla nerved, 
stamens rolled up but not twisted, style declinate and with linear stigmas as 
in Sabbatia. From Arkansas, 8 inches high, leaves small remote semi-uncial, 
flowers large over uncial incarnate ? Neurola arkanzica will be a better name. 

This description, which alone is rather convincing, coupled 
with the information that the specimen was one of Nuttall’s 
collections from Arkansas leaves no doubt as to the identity 
of the plant. Nuttall found but two species on that journey, 
S. campestris and the well-known S. angularis. Rafinesque 
(Aut. Bot. p. 55. 1840.) was still uncertain as to the proper 
rank to bestow upon this species for he referred to it as '*Sab- 
balia nervosa or Neurola arkanzica . . . fine subg. perhaps a 
Genus . . ." 

The final proposition of specific rank made for this species 
was that of Buckley. The specimen upon which this was 


1955] Wilbur,— The North American Genus Sabatia 47 


based has not been seen by me but again the description is such 
that determination is certain. "The original publication reports 
it to have been collected in Llano Co., Texas, from whence 
numerous specimens of S. campestris have been examined. 
'That is the only species of the genus known from Central Texas. 
Buckley's proposal was promptly relegated to synonymy by 
Gray (Proc. Acad. Nat. Sci. Phil. 1862: 166. 1863.) whose regret 
as to the “hundred worse than useless synonyms" published 
with S. formosa, especially after the caustic criticism strongly 
reminiscent of an even later Fisher Professor of Natural History, 
was doubtless shared by Buckley. 

Merrill (Rnopona 50: 127. 1948.) equates S. concinna Wood 
with S. campestris rather than with S. brachiata to which Gray 
long before had reduced it and to which Wood himself later 
considered it synonymous. The evidence that it is anything 
but S. brachiata is not convincing and the name is discussed 
more fully under that species. 


REPRESENTATIVE SPECIMENS:—ILLINOIS: DuPage Co., Hinsdale, Smith 497 
(F). MISSISSIPPI: Covington Co., about 13 mi. nw. of Hattiesburg, Webster & 
Wilbur 3359 (micH). Mrissounr: Christian Co., 2 mi. s. of Saunderds, Steyer- 
mark 23232 (F, MO, NY, TENN, WIS); Jasper Co., near Asbury, Palmer 34661 
(MO, NY). ARKANSAS: Benton Co., 9 mi. ne. of Springdale, Moore 350339 
TYPE of S. campestris f. albiflora Moore (UARK); Bradley Co., Warren, Demaree 
19268 (MO, NY, OKLA). LOUISIANA: Calcasieu Parish, vicinity of Lake Charles, 
Allison 279 (Ny, vs); Winn Parish, 10 mi. w. of Winnfield, Webster & Wilbur 
3261 (MIcH). KANSAS: Bourbon Co., 4 mi. w. of Fort Scott, Horr E165 (DUKE, 
FLAS, NCS, OKL, OKLA, PENN, SMU, US); Cherokee Co., Baxter Springs, Oyster 
5445 (F, MO, NY); Montgomery Co., 5 mi. ne. of Caney, Rydberg & Imler 408 
(KSC, MO, NY, US). OKLAHOMA: Cleveland Co., Norman, Demaree 13112 (Gu, 
MO, NY, OKL, US); Comanche Co., near Cache, Stevens 1328 (GH, MO, NY, OKL, 
OKLA, US); Creek Co., 7 mi. n. of Sapulpa, Ownbey 1606 (GH, MICH, MO, NY, 
OKL, PENN, TEX, US); Johnston Co., near Mannsville, Griffith 3474 (GH, Mo, 
NY). TEXAS: Brazoria Co., Columbia, Bush 76 (an, Mo, NY, US); Calhoun Co., 
Port Lavaca, Tharp, 22 May 1930 (DUKE, KSC, MO, MT, OKLA, TAES, TEX, WVA); 
Frio Co., near Moore, Palmer 33868 (Gu, Mo, NY, vs); Harris Co., Houston, 
Hall 508 (BRU, F, GH, NY); Llano Co., Enchanted Rock, Tharp, 11 June 1930 
(GH, MO, NY, SMU, TAES); Parker Co., Weatherford, Tracy 8045 (cU, F, GH, MO, 
NY, TEX, US); Rockwall Co., ca. 30 mi. e. of Dallas, Webster & Wilbur 2960 
(MrcH); Travis Co., Austin, Tharp, 16 May 1939 (MICH, MO, NA, NY, SMU, TEX). 


7. Sabatia arenicola Greenm., Proc. Am. Acad. 34: 569. 1899. S. 
carnosa Small, Fl. SE. U. S. 927. 1903. 

Erect annual (3-)10-20(-32 or up to 45 in greenhouse-grown plants) 
em. high; stems rigid, 1-3 mm. in diameter, succulent when young but 
becoming less so with maturity, more or less terete in the lowermost 
internodes but soon becoming strongly quadrate above and the corners 


48 Rhodora [Vor. 57 


winged by a fine, hyaline membrane about 0.1-0.3 mm. high that is 
barely discernable in dried material. Branches most commonly arising 
from near the base and then often opposite but becoming alternate 
above, or in more loosely developed plants absent from the lower half 
of the plant and alternate throughout, strongly ascendent or at first 
widely divergent (up to 60 degrees) and, in plants grown under the 
normal highly exposed environmental conditions, typically presenting 
an almost globose form or a compact corymbiform to convex crown. 
Root-system most commonly a strongly developed tap-root or almost 
as commonly with several almost equally developed fibrous roots; these 
4-20 cm. long and bearing few to several slender laterals. Leaves thick, 
widely spreading, strikingly succulent, shiny, heavily cutinized, venation 
almost completely obscured in living material, not at all elevated on 
either surface, appearing 3-5 nerved in dried specimens, the lowermost 
narrowly spatulate to somewhat elliptie, strongly tapering to the almost 
petiolate base, above becoming elliptical to ovate-lanceolate or rarely 
somewhat oblong, usually broadest one-fourth to one-third the length 
from the base or even near the middle, and somewhat tapering to the 
sessile, non-clasping base, (0.7-)1.2-1.8(-2.7) em. long, (2.0-)5.0-8.0 
(-13.0) mm. wide, usually about 1.5-4 times as long as broad, obtuse 
to acute with a minute, non-chlorophyllous callosity forming a mucronu- 
late tip. Internodes (0.3-)1.0-2.0(-8.0) em. long, usually 0.5-2 times 
the length of the leaves but in more loosely developed plants not uncom- 
monly up to 4 or more times as long as the leaves. Inflorescence of 
corymbosely to pyramidally arranged cymules compactly clustered or 
borne loosely in plants grown under less rigorously exposed conditions. 
Cymules 1-2-flowered. Pedicels often somewhat succulent, rigid, 
pentagonal, bearing 5 fine, hyaline, membranous wings, 0.5-3(-7) em. 
long. Calyx-tube broadly campanulate to urceolate, usually about 
twice as long as broad, (0.3—)1.0-2.0(-8.0) mm. long, becoming broadly 
turbinate in fruit, strongly pentagonal due to thick, somewhat succulent, 
costae or ribs about 0.5-0.7 mm. high, the internal portion of which is 
chlorophyllous, the outer finely hyaline margined. Intercostal tube very 
thinly membranous, colorless but translucent, thus appearing green due 
to the ovary within. Calyx-lobes thick, succulent, heavily cutinized, 
erect in bud but becoming strongly spreading at anthesis, (0.5—)1—1.5(-3) 
times longer than the tube, (0.3-)0.5-1.2(-2.0) em. long, oblong to most 
commonly linear-lanceolate, usually about 3 times as long as broad, 
acute, minutely hyalinely callose-tipped, becoming prominently nerved 
when dried, in general very similar to leaves in texture. Corolla-tube 
exceeding calyx-tube as well as the ovary by about 1-2 mm., lower portion 
colorless, the upper colored by extension of the yellow patches at lobal 
bases. Corolla-lobes more or less equaling or somewhat shorter than 
the calyx-lohes, or rarely longer, ascendent to strongly spreading, broadly 
spatulate, obovate, or oblong, obtuse, (0.5-)0.7-1.0(-1.3) cm. long, 
3-7.5 mm. wide, deeply roseate or pure white, both forms with a 2-3 mm. 
long, 1.5 mm. wide basal, oblong, yellow patch, which in roseate forms 
is often bordered by deeper shade of red. Anthers bright-yellow, oblong, 


1955] Wilbur,—The North American Genus Sabatia 40 


1.5-2 mm. long; filaments white to pale-yellow, slender, 2-4 mm. long. 
Style slender, colorless to pale greenish-yellow, 2-3.5 mm. long, stigmatic 
branches green to greenish-yellow, 2.5-4 mm. long. Capsule 5-9 mm. 
long, equaling and included within the calyx-tube or exserted by 1-3 mm. 
TYPE LOCALITY: “on damp sands of seacoast near Tampico, State of 
Tamaulipas, Mexico." Type: Pringle 6808 (Gu!). DISTRIBUTION: Beaches 
and dunes from Louisiana southward to at least the southern part of 
the State of Tamaulipas, Mexico. Map 7. 


The pronounced succulence of this species is perhaps its most 
striking characteristic and that by which it is most readily 
distinguished from the closely related but perfectly distinct 
S. campestris. 


Many of the differences noted between the two species of 
subsection Campestres are such that one might expect them to 
be merely response of a highly plastic organism to an environ- 
mental extreme. Succulence and diminution of the various 
organs and structures and the assumption of an almost globose, 
compact form are features that logically would be expected 
among the littoral members of a species of wide environmental 
variation. Sabatia arenicola, found along the strand and dunes 
from Louisiana to at least Tampico in Mexico, possesses such 
characteristics and accordingly suspicion is raised as to its status. 


The few specimens of S. campestris observed in the field in 
almost similar situations, although stunted or dwarfed, failed 
to show the peculiarities characteristic of the littoral species. 
Furthermore, herbarium specimens of plants that were un- 
doubtedly S. campestris, as indicated by their morphological 
characters, were collected, according to their accompanying 
labels, from what was approximately the environment of S. areni- 
cola. ln addition, numerous plants of S. arenicola grown in the 
greenhouse from seeds collected at various stations along the 
Texas coast retained the pronounced succulent nature so charac- 
teristic of the littoral species. The greenhouse-grown plants were 
larger, more elongate, and much more diffuse than the majority of 
specimens of S. arenicola seen by me either in the field or in the 
herbarium but are approached to some extent by those growing 
in nature in the more sheltered areas. The principal distinguish- 
ing characters mentioned in the key remain constant and the 
plants of the two species grown side by side are readily separ- 
able. 


50 Rhodora [Vor. 57 


S. BRACHIATA 


S. CAMPANUL ATA 


Maps 5-8. Map 5, upper left; map 6, upper right; map 7, lower left; map 8, lower 
right. 

Two specimens were annotated by Small as S. carnosa. These 
are the only representatives of the littoral species that were 
available to him at New York prior to 1903 when his Flora 
was published. None of the specimens on either of the sheets 
is as much as 3 dm. tall, the maximum height mentioned in 
the original description. However, it is likely that these were 
the only examples seen by him before publication; they were 
certainly the only two annotated by him as that species and one 
of them should be chosen as the type of the name. The only 
collection of the two that matches the statement “‘often branched 
at the base" is that of A. Schott taken on the beach at Brazos 


1955] Wilbur,— The North American Genus Sabatia 51 


Santiago near the mouth of the Rio Grande on the expedition 
led by Major Emory to survey the Mexican boundary. The 
other specimen annotated as S. carnosa by Small is one of 
Drummond’s but no other information, other than that it was 
from Texas, is presented on the accompanying label. However, 
the great similarity in aspect and in state of preservation strongly 
suggests that it is a duplicate of these specimens seen at both 
the Gray Herbarium and Kew which bear Drummond’s collection 
number ‘59’ and which came from the “Rio Brassos" (Rio 
Brazos) Texas. The New York specimen may well be taken 
as the lectotype of S. carnosa. These specimens, the basis of 
Small’s S. carnosa, can in no way be distinguished from Green- 
man's S. arenicola which has four years priority. 

This species and S. campestris are usually readily separable 
if the specimens are adequate. A few collections have been 
seen, however, that indicate that hybridization may be taking 
place between them in the region where they come into contact. 
Careful observation in the field should do much toward settling 
this suspicion. 

REPRESENTATIVE SPECIMENS:—LOUISIANA: Jefferson Parish, Grand Isle, 
Cangemi & Andrus, 13 June 1931 (Ny); Terrebonne Parish, Timbalier Island, 
Tharp, 29 July 1929 (mo, TEX). tTExas: Aransas Co., n. of Rockport, White- 
house 18232 (smu); Brazoria Co., Rio Brazos, Drummond 59 (GH, K, NY); 
Calhoun Co., beach near Port O'Connor, Webster & Wilbur 3134 (wich); 
Cameron Co., beach at Brazos Santiago, Schott 27 (F, Ny), about 3 mi. w. of 
Boca Chica, Lundell 8635 (Gn, MICH, NY, SMU); Galveston Co., beach on Gal- 
veston Island, Webster & Wilbur 3153 (micu); Harris Co., Morgan’s Point, 
Palmer 11966 (Mo, Us); Kennedy Co., El Toro Island, Tharp 49218 (TEX); 
Kleberg Co., Kingsville, Sinclair, 1940 (Tex); Matagorda Co., beach beyond 
Sargent, Webster & Wilbur 3152 (micH); Nueces Co., north end of Padre 


Island, Webster & Wilbur 3092 (micH); San Patricio Co., 10 mi. s. of Aransas 
Pass, Whitehouse 18213 (swv). 


Mexico: TAMAULIPAS: Washington Beach, Runyon 461 (aH, Ny, us); San 
Jose, LaSueur 349 (TEX); vicinity of LaBarra, 8 km. e. of Tampico, Palmer 297 
(F, GH, US); Tampico, Pringle 6808 (BRU, F, GH, NY, UC, US). 


D. Subsection CAMPANULATAE Blake, RHopona 17: 56. 1915. 


Annuals or perennials with either slender tap-root, several fibrous 
roots, or an erect, branched, perennating caudex. Branches typically 
alternate. Corolla 5-parted, rose, pink, or white. Calyx smooth or 
finely ribbed, not wing-costate. TYPE SPECIES: Sabatia campanulata (L.) 
Torr. 

The species forming this subsection are not of uniform habit of growth 
as three of the species are annuals, while one, the type, is a perennial 


52 Rhodora [Vor. 57 


with a subterranean system unique in the genus. This is the only species- 
group showing diversity in life-form. The branching pattern in this 
group is typically alternate, but this character is highly variable, as it 
is throughout most of the genus. This subsection is not set off by one 
strong characteristic or even by a series of lesser ones. Segregation of 
the annuals, leaving S. campanulata to form a monotypic subsection 
characterized by its shortly branched, erect, caespitose caudices, would 
result in two subsections, each certainly more homogeneous than the 
present one. For the moment at least, I am hesitant about erecting 
additional subgeneric units as their utility is defeated when they become 
so numerous as to be composed of only one or two species. The genus 
is already, for one so small, burdened with a large number of subgeneric 
units, 


KEY TO THE SPECIES OF SUBSECTION CAMPANULATAE 


A. Caespitose perennial from branched, erect caudex; calyx-lobes usually 
more or less equaling the corolla in length......... 8. S. campanulata. 

A. Annuals with either a tap-root or several fibrous roots; calyx-lobes usually 
less than three-fourths the length of the corolla. 

B. Calyx lobes linear to almost filiform, usually more than 8 mm. long; 
corolla usually roseate, rarely white; style 2 mm. or more in length; 
branches usually not numerous, mostly restricted to the upper half of 
the stem; nodes along the main axis not numerous, usually less than 
15; root-system typically of several, almost equal, fibrous roots. 

C. Leaves drying thin and smooth, the uppermost equaling or exceeding 
the stem in diameter; corolla-lobes usually less than 1.5 cm. long; 
calyx-tube usually turbinate or narrowly campanulate; known from 
Mexico, Cuba, the Bahamas, Florida west to Louisiana and north 
to southern Massachusetts. ........ 0.00000 eevee 9. S. stellaris. 

C. Leaves drying thick and very noticeably rugose; above typically less 
than the stem in diameter; corolla-lobes usually longer than 1.5 
cm.; calyx-tube usually broadly campanulate; known only from 
Cuba and peninsular Florida.................. 10. S. grandiflora. 

B. Calyx-lobes setaceous or subulate, less than 8 mm. long; corolla white; 
style 1 mm. long or less; branches usually numerous and typically 
arising from throughout the length of the stem; nodes very numerous 
and typically more than 20 along the main axis; root-system of one 
prominently developed, wiry tap-root with several slender laterals. 

11. S. brevifolia. 


8. Sabatia campanulata (L.) Torr., Fl. N. & Mid. U.S. 217. 1824. Chir- 
onia campanulata L., Sp. Pl. 1: 190. 1753. C. gracilis Michx., Fl. Bor. 
Am. 1: 146. 1803. C. campanulata 8 gracilis (Michx.) Pers., Syn. Pl. 
1: 282. 1805. Sabbatia gracilis (Michx.) Salisb., Parad. Lond. t. 32. 
1806. S. campanulata f. albina Fern., Ruopora 18: 151. 1916. 5$. 
Tracyi Gandoger, Bull. Soc. Bot. Fr. 65: 61. 1918. S. campanulata 
var. gracilis (Michx.) Fern., RHopora 39: 444. 1937. 

Caespitose perennial from an erect, short, much-branched, underground 
caudex 1-4 em. long. Stems few to numerous, erect, (15-)30-60(-90) 
cm. high, 1-3 mm. in diameter, terete but often strongly ridge-angled 


1955] Wilbur,— The North American Genus Sabatia 53 


from fine elevated internodal lines, becoming hollow below. Branches 
typically alternate throughout, or occasionally those at lower nodes 
opposite, rarely opposite even above; usually restricted to the upper half 
of the stem but occasionally arising from throughout the length, strongly 
divergent forming an angle with the stem of 50-80 degrees or, especially 
in the northern portion of its range, strongly ascendent forming angle of 
about 20-40 degrees; few to numerous, slender, bearing but few nodes 
(more common in the northern portion of the range) or several. Root- 
system of few to many, slender, but apparently succulent, roots descending 
from the caudex, about 5-10 em. long and 1 mm. thick bearing few to 
several very slender, fibrous laterals. Leaves ascendent to very com- 
monly (especially the upper) strongly spreading, midvein noticeably 
elevated beneath, venation otherwise obscure, margins somewhat thick- 
ened and slightly revolute, drying thick, somewhat chartaceous, smooth 
or slightly rugose, obtuse to acute, broadly sessile or somewhat tapering 
in broader leaves to the base, the lowermost rarely ovate; the lower 
cauline (1-)1.5-3(-4) em. long, (1-)2-7(-12) mm. wide, usually 5-12 
times as long as broad, narrowly lanceolate, oblong, or linear gradually 
reduced above to very narrowly linear or even filiform; those of the 
branches narrowly linear to filiform. Internodes usually 1-2 times the 
length of the leaves. Flowers apparently solitary, borne on alternate, 
or rarely opposite, ascendent to widely divergent, slender branches 
bearing 1 to several nodes, or in more obvious cymules which are often 
reduced. Pedicels (2-)4-7(-9) em. long, slender, wiry, slightly angled. 
Calyx-tube shallowly turbinate or more rarely somewhat campanulate, 
smooth or nearly so, not conspicuously nerved, scarious, usually about 
as broad as long, (1—)1.5-2.5(-3) mm. long. Calyx-lobes linear to very 
narrowly so or very commonly acicular or setaceous, erect to more com- 
monly widely spreading, (6-)7-17(-23) mm. long, typically less than 0.5 
mm. wide but occasionally as wide as 1 or even 2 mm., usually 5-15 
times as long as the calyx-tube, generally nearly equaling the corolla 
in length, or but several millimeters shorter, rarely exceeding it. Corolla- 
tube cylindrical, about 2-3 times as long as the calyx-tube or exceeding 
it about 2-3 mm., colorless to white or pale greenish-yellow, (2-)3-5(-6) 
mm. long. Corolla-lobes wide-spreading, oblanceolate, oblong, or 
elliptic, acute to obtuse, (0.6-)0.9-1.8(-2.4) em. long, (3-)4-7(-9) mm. 
wide, usually about 2-4 times as long as wide, rose, pink, or rarely white 
with an usually unlobed yellow area at base of lobe 2-3 mm. long, often 
bordered by a dull red irregular line. Anthers slender, bright yellow, 
(2-)3-4(-5) mm. long; filaments pale yellow, slender, 2-4 mm. long, 
usually shorter than the anthers. Style 2-5 mm. long, usually shorter 
than its branches, green to greenish-yellow; stigmatic lobes 3-7 mm. 
long, pale-green turning yellowish (with pollen?) at maturity. Ovary 
pale-green, slightly exserted from the corolla-tube. Capsule cylindrical, 
5-7 mm. long, 2.5-4 mm. in diameter, usually 1.5-2 times as long as 
wide. TYPE LOCALITY: “in Canada." This plant is not known from 
farther north than Massachusetts. type: Kalm (Photograph of the 


54 Rhodora (Vou. 57 


Type in the Linnaean Herbarium seen in the collection of the Gray Her- 
barium). DISTRIBUTION: Peaty bogs and savannas along the Coastal 
Plain from Massachusetts south to northern Florida and westward to 
Louisiana and Arkansas; also found in the southern Appalachians and 
locally in Kentucky and Illinois. Map 8. 


Fernald recognized two populations within this species. The 
supposed differences as defined by him are presented below. 


campanulata gracilis 
Primary cauline leaves oblong- Lower cauline leaves linear, the 
linear to lanceolate. upper ones very narrowly so. 
Pedicels naked or only slightly Pedicels mostly leafy-bracted. 
bracted. 
The linear calyx-segments (except in The linear-acicular calyx-segments 
in small secondary flowers) 1-2 6-14 mm. long. 
em. long. 
The corolla-segments 1-1.7 cm. long. The corolla-segments 6-14 mm. long. 
Branches erect or strongly ascendent. Branches more divergent. 
Upland and Piedmont region, Ga., On the Coastal Plain from Florida 
and Ala., n. to Va., and Ind., to Louisiana and se. Va. 
and on or near Coastal Plain to 
se. Mass. 


Many of the specimens studied could be assigned to one or 
the other group with a fair degree of assurance by applying 
these criteria. Yet there are a number of puzzling collections 
which aroused a strong suspicion as to one's ability to determine 
"correctly" a large series of specimens if one were unaware 
of their geographic origin. For example, Deam (1940) considered 
the material from southwestern Indiana to be a representative 
of the typically Coastal Plain population while Fernald (1950) 
mentioned Indiana only in the range of the campanulata-group. 

Fernald (Ruopona 34: 27. 1932.) originally had considered 
the plants found along the southern Coastal Plain to be “a 
rather well defined species" although admitting that “. . . it 
may be better to treat it as a southern Coastal Plain variety . . ." 
After encountering specimens in southeastern Virginia that 
possessed some of the supposed distinguishing characteristics 
of both varieties, Fernald (Rhodora 39: 444. 1937.) concluded 
that “. . . S. gracilis should be treated as a geographical vari- 
ety." Those intermediate specimens combined the presumably 
characteristic narrow leaves and calyx-segments of the southern 
Coastal Plain population with the longer lobes of both calyx 
and corolla of the more northern collections. Since numerous 


1955] Wilbur,— The North American Genus Sabatia 55 
large-flowered specimens had been observed throughout the 
supposed range of the gracilis-population, the several hundred 
available measurements were tallied for comparison and are 
presented below. The specimens from southeastern Virginia 
which Fernald considered atypical as they showed features 
of both populations were not included. 


LENGTH OF CALYx-LonBEs LENGTH OF CoROLLA-LOBES 
No. of lobes No. of lobes No. of lobes No. of lobes 
mm. gracilis-range | campanulata-range gracilis-range | campanulata-range 

6 6 0 1 0 

7 23 5 4 0 

8 30 6 4 1 

9 42 18 16 7 
10 40 22 34 16 
11 50 31 44 38 
12 60 42 63 38 
13 45 36 52 43 
14 45 20 36 38 
15 27 19 32 19 
16 19 Lr 17 7 
17 10 19 15 0 

18 TA 9 10 0 

19 1 5 4 2 
20 4 2 2 0 
21 2 1 0 1 
22 0 1 0 0 
23 0 1 e 0 
24 0 0 0 1 


These hundreds of measurements indicate that, although 
the smallest flowers were found on the specimens from the 
gracilis-range and the largest from the range of the more northerly 
population, no reliance can be placed upon the length of either 
the calyx- or corolla-segments as a distinguishing characteristic 
between the two supposed varieties. These measurements 
show that the range of flower-size of the plants from the Coastal 
Plain is considerably greater than that known to Fernald and 
for diagnostie purposes the range of the length of the lobes of 
either floral series is so broadly overlapping as to be unworkable. 

Nor am I able to detect any consistent differences in the shape 
of the calyx-lobes between those specimens of the Coastal Plain 
south of Virginia and those from the Appalachian region and 
the Costal Plain north of Virginia. Numerous specimens from 
Florida and the other states of the supposed range of gracilis 
have linear calyx-lobes presumably characteristic of the northern 
variety. 


56 Rhodora [Vor. 57 


By ''pedicels" Fernald apparently meant lateral branches 
bearing flowers. "These branches in the Massachusetts material, 
with which Fernald had field experience, are typically short 
and rather strongly ascendent and their one or two nodes bear 
bracts that are foliose in comparison to the narrowly linear to 
even filiform bracts of the typical southern coastal material. 
A tendency for opposite branching in the upper part of the stem 
is also more pronounced, especially in more robust specimens, 
and then the unreduced cymose pattern of inflorescence is some- 
what more apparent. These rather striking characteristics 
found with considerable constancy in the few New England 
stations are far less apparent in the specimens seen from the more 
southern part of the range of the supposed typical variety. 
Numerous specimens from the mountainous regions of Georgia, 
Alabama, the Carolinas, Tennessee and Kentucky, as well as 
Deam’s Indiana station, possess elongate, widely divergent 
branches with several to numerous nodes bearing narrowly 
linear to even filiform bracts. The extremes of leaf-type from 
the two ranges are often most striking and clear-cut but many 
specimens within the range attributed to one of the varieties 
would be assigned to the other upon the application of this 
criterion. These considerations have convinced me that it 
would be difficult to recognize varieties in this species in spite 
of some rather strong tendencies which are somewhat geograph- 
ically segregated and whose extremes are of rather striking 
difference in aspect. 


Gandoger based the description of S. tracyi upon the specimen 
in his herbarium of Tracy 6468 collected at Biloxi, Mississippi. 
I have not seen his type but four specimens of the same collection 
number are located at the Gray Herbarium (2 sheets), New York 
Botanical Garden, and the National Herbarium. They are 
presumably isotypes. Gandoger described his specimen as an 
annual and this would relate or equate S. tracy? with S. stellaris, 
the only annual of subsection Campanulatae known to the state. 
The presumed isotypes, however, are perennials with narrowly 
oblong-linear to linear leaves gradually reduced above and on 
the branches to filiform bracts; their branches, which are alter- 
nate, are elongate, bear numerous nodes, and are very strongly 
divergent. There is still a trace of rose-pink color on the corolla- 


1955] Wilbur,— The North American Genus Sabatia 57 


lobes of at least three of the sheets seen although even on those 
sheets most of the flowers appear yellowish-white. Gandoger’s 
species was described as having white flowers which together 
with the more slender leaves and mucronate sepals supposedly 
distinguished it from S. gracilis. S. tracyi is synonymous with 
S. campanulata, or, if one prefers to recognize what now seems 
to me to be a variation too weak to maintain, to S. campanulata 
var. gracilis (Michx.) Fern. It may be matched by a majority 
of the specimens collected along the Gulf and Southern Atlantic 
Coastal Plain. 


REPRESENTATIVE SPECIMENS:—MASSACHUSETTS: Barnstable Co., Barn- 
stable, Fernald & Svenson, PL. EX. GRAYANAE 479 (in many herbaria); Nan- 
tucket Co., Almanac Pond, Williams, 30 July 1911 (aH); Plymouth Co., Pem- 
broke, Foster, 10 Sept. 1884 (aH). NEW YORK: Nassau Co., East Rockaway, 
Bicknell 7089 (Ny); Suffolk Co., Oakdale, Ferguson 7775 (NY). NEW JERSEY: 
Ocean Co., e. of Silverton, Fogg 4908 (GH, NY, PENN, TENN). PENNSYLVANIA: 
Bucks Co., Tullytown, MacElwee, 28 July 1894 (aH, NY). DELAWARE: Kent 
Co., Felton, Canby, July 1878 (us). MARYLAND: County unknown, Delaware 
Beach, Hood 2370 (FLAS). DISTRICT OF COLUMBIA: Virginia Highlands, 
Griggs, 17 July 1923 (us). VrRGINIA: Greensville Co., ne. of Gaskins, Fernald 
& Long 13421 (GH, MO, TENN, US); Sussex Co., about 4 mi. nw. of Homesville, 
Fernald & Long 6351 (GH, NY, PENN). NORTH CAROLINA: Columbus Co., 2 mi. 
w. of Freeman, Wilbur 2882 (micu); Henderson Co., 1 mi. s. of East Flat 
Rock, Oosting 1790 (puKE); Northampton Co., near Margarettsville, Heller 
1160 (CU, F, GH, KSC, MO, MT, NY, PENN, US). SOUTH CAROLINA: Anderson Co., 
Belton, Smith, 23 July 1881 (cH, vs); Georgetown Co., 12 mi. n. of George- 
town, Godfrey & T'ryon 66 (DUKE, F, GH, MICH, NY, PENN, TENN, US). GEORGIA: 
Bartow Co., 4.5 mi. s. 16° east of Allatoona Dam, Duncan 8521 (FLAS, GA, 
MO, SMU, TENN, US); Thomas Co., about 1 mi. n. of Coolidge, Duncan 8466 
(FLAS, GA, MO, TENN). FLORIDA: Bradford Co., near Lawtey, Murrill 528 
(DUKE, MO, US); Holmes Co., Bonifay, Curtiss 6481 (GH, MO, NY, SMU, US); 
Washington Co., 8 mi. s. of Chipley, Webster & Wilbur 3615 (MICH). INDIANA: 
Daviess Co., about 4 mi. n. of Washington, Deam 52742 (F, GH, MO, NY); 
Jasper Co., about 3 mi. se. of Wheatfield, EK, 30 July 1940 (GĦ). KENTUCKY: 
Fayette Co., Lexington, Short, 1831 (Ny); Whitley Co., n. Jellico, Braun, 21 
July 1933 (GH, NY). TENNESSEE: Coffee Co., n. of Manchester, Svenson 9219 
(DUKE, GH, MO, NA, NY, WIS, US); Franklin Co., near Tullahoma, Sharp, 
Clebsch & Fairchild 9938 (DUKE, GA, NCU, OKLA, SMU, TENN, TEX, US). ALA- 
BAMA: Conecuh Co., about 9 mi. w. of Evergreen, Webster & Wilbur 3514 
(mich); DeKalb Co., Lookout Mountain, Biltmore Herb. 4510g (us); Mobile 
Co., about 8 mi. sw. of Mobile, Webster & Wilbur 3480 (MICH). MISSISSIPPI: 
Covington Co., 1.5 mi. se. of Mt. Olive, Webster & Wilbur 3336 (wicm); 
Forest Co., 18 mi. nw. of Beaumont, Webster & Wilbur 3424 (micu); Jackson 
Co., Ocean Springs, Pollard 1010 (cv, F, GH, MO, NY, US). ARKANSAS: Lonoke 
Co., Grand Prairie, Demaree 22329 (wo); Pulaski Co., Little Rock, Harvey, 
July 1882 (UARK). LoUuiSIANA: Calcasieu Parish, Lake Charles, Cocks, Sept. 
1906 (No); Tangipahoa Parish, 3 mi. e. of Robert, Correll 9307 (DUKE, Mo, 
TEX). 


58 Rhodora [Vor. 57 


9. Sabatia stellaris Pursh, Fl. Am. Sept. 1: 137. 1814. Chironia 
amoena Raf., Med. Repos. II. 5: 359. 1808. not Salisb., Prodr. 137. 
1796. C. stellata Muhl., Cat. Pl. Am. Sept. ed. 2, 23. 1818. C. stel- 
laris [Pursh] Eaton, Man. ed. 2, 204. 1818. Sabbatia maritima Raf., 
Med. Fl. 2: 77. 1830. S. amoena (Raf.) G. Don, Gen. Hist. 4: 207. 
1838. S. stellaris 8 pumila A. Gray ex Griseb., Prodr. 9: 49. 1845. 
Eustoma maculata Benth., Pl. Hartw. 292. 1848. Sabbatia gracilis 8 
stellaris [Pursh] Wood, Am. Bot. & Flor. 267. 1870, without basionym. 
S. nana Featherman, Rep. Bot. Surv. S. & Cent. La. 71. 1871. 5$. 
maculata (Benth.) Benth. ex. A. Gray, Proc. Am. Acad. 22: 438. 1887. 
S. Palmeri A. Gray, Proc. Am. Acad. 22: 438. 1887. S. stellaris f. 
albiflora Britton, Bull. Torrey Club 17: 125. 1890. S. simulata Britton, 
Bull. N. Y. Bot. Gard. 3: 448. 1905. S. Purpusii Brandegee, Univ. 
Calif. Pub. Bot. 4: 275. 1912. Sabatia amoena f. albiflora (Britton) 
Fern., HnHopon4 34: 26. 1932. Sabbatia campanulata var. amoena 
(Raf.) Monachino, Torreya 41: 99. 1941. 


Erect annual (2-)15-50(-80) em. high; stem 1-4 mm. in diameter, 
terete to strongly angled due to several somewhat irregularly disposed 
fine lines or ridges extending between nodes, pith parenchymatous in 
young living specimens but becoming hollow in older or dried specimens. 
Branches almost always alternate, very rarely one or so nodes with op- 
posite branches, usually restricted to upper half or two-thirds of the stem 
but sometimes arising throughout the length and even from the base, 
typically strongly divergent, forming an angle of about 30-70(-85) de- 
grees with the stem, few to numerous, slender, wiry, bearing few to 
numerous nodes. Hoot-system of several to numerous, usually slender, 
but occasionally thickened, fibrous roots 2-10 em. long, 1-3 mm. in 
diameter or occasionally with one principal root and several laterals. 
Leaves ascendent, above often closely appressed to stem, succulent, thick, 
rubbery in texture, midvein conspicuous and elevated beneath, venation 
otherwise obscure; after drying leaves commonly darken, usually thinly 
membranous or occasionally the lowermost slightly rugose, the midvein 
alone usually prominent. Lower leaves broadly to narrowly elliptic, or 
even linear, rarely spatulate or obovate, acute to rarely obtuse, often 
apiculate, usually tapering to both ends, typically about 5-10 times as 
long as broad, (0.5-)1.5-3(-6) em. long, (2-)3-8(-15) mm. wide. Upper 
leaves more narrowly elliptie to linear, the uppermost sometimes very 
narrowly so to even almost filiform, usually 7-15(-20) times as long as 
wide, 1-4 em. long, (0.5-)1.5-3(-5) mm. wide, typically exceeding the 
diameter of the stem or at least equaling it. Internodes usually 1-2 
times the length of the leaves. Flowers appearing solitary but usually 
arranged in very loose and reduced eymules and these sometimes aggre- 
gated in loose clusters. — Pedicels slender, straight, ascendent to divergent, 
(1-)4-10(-15) em. long, usually about 0.5 mm. in diameter. Calvx-tube 
turbinate, or occasionally even narrowly campanulate, usually rather 
gradually narrowing to the base, smooth or but very faintly nerved, thin, 
somewhat scarious, usually 1-2 times as long as broad, (1.5-)2-4(-6) mm. 


1955] Wilbur,— The North American Genus Sabatia 59 


long. Calyx-lobes narrowly linear to almost filiform, ascendent or more 
typically wide-spreading, (4-)6-15(-22) mm. long, typically less than 0.5 
mm. but rarely as wide as 1 mm., usually 3-6 times as long as calyx-tube, 
in general considerably shorter than the corolla, rarely equaling or ex- 
ceeding it, apparently slightly hyaline-scarious margined. Corolla-tube 
about 2-3 times as long as wide, usually exceeding calyx-tube by 1-3 mm., 
translucent but appearing greenish due to ovary, (3-)4-6(-8) mm. long. 
Corolla-lobes strongly spreading, elliptic, oblong, spatulate, or obovate, 
obtuse to somewhat acute, (0.5-)1—-1.5(-2) em. long, (2-)4-8(-10) mm. 
wide, rose, pink, or more rarely white, with a usually irregularly 3-lobed 
yellow area at the base often bordered by a distinct, bright or dark red 
line which in turn is sometimes bordered by a white area of variable 
width. Anthers linear, yellow, (2-)3-4(-5) mm. long; filaments pale 
yellow to greenish, (1.5-)2-3(-4) mm. long. Style 2-4 mm. long, usually 
about half the length of its branches; stigmatie lobes 3-8 mm. long. 
Capsule from almost globose to cylindrical, (4-)6-8(-14) mm. long, 
(3-)4-5(-6) mm. wide, usually 1.6-2 times as long as broad. TYPE 
LOCALITY: “in salt marshes: New York, New Jersey &c." DISTRIBUTION: 
Salt marshes and sandy (usually littoral) places from southern Massa- 
chusetts south along the coast and throughout much of Florida and west 
into Louisiana. Also known from the Bahamas, western Cuba and the 
central plateau of Mexico. Map 9. 


Sabatia stellaris is the most variable species of the Campanulatae 
but within this wide range of diversity there appears to be no 
sharply differentiated population that might be more naturally 
treated as a subspecies or a variety. Rather, there is an almost 
imperceptible change and as a consequence specimens from the 
northern limits of the range are often strikingly different from 
those seen at the extreme southern portion of its distribution. 
Typically, plants collected in southern Florida will be taller, 
more profusely branched, often from near the base, and these 
branches will be much more elongate, wiry, and slender. The 
leaves, for the most part, will be narrowly linear and only the 
uppermost with indications of a more elliptical or spatulate 
nature. The northern material will be on the average smaller, 
less branched, and then the branching usually restricted to 
the upper portion of the stem; the branches usually short and 
bearing few nodes. Also the leaves in most cases will be linear 
only in the very upper portion of the stem and otherwise will 
often be broadly elliptical. The sum of these differences results 
in plants of very dissimilar appearance. However, examination 
of an ample series taken from throughout the range will convince 


60 Rhodora [Vor. 57 


one, I believe, that these differences are not sufficient for the 
recognition of subspecific categories. Rather these differences 
should in part be attributed to the great diversity in environment 
encountered through the range of more than twenty degrees 
of latitude and especially the difference in the length of the 
growing season between the Massachusetts coast and the shores 
of the Caribbean. 

After initially attempting (1903) to distinguish between 5$. 
campanulata and 5. stellaris, Small abandoned the effort and 
thereafter applied the former name to specimens of both 
species in the Southeast. The confusion between the two 
species is understandable due to the often misleading distinguish- 
ing criteria published in most works previous to the detection 
of the strikingly obvious differences by Bicknell (Bull. Torrey 
Club 42: 30-32. 1915.) and especially by Fernald (RHODORA 
18: 145-147. 1916.). "Thereafter there should have been little 
diffculty in distinguishing the two. However, most of the 
southern material of S. stellaris has passed as S. campanulata 
or the larger-flowered specimens as S. grandiflora due to the 
influence of Small’s work. Sabatia campanulata is not known 
in peninsular Florida or from the West Indies. 

Gleason's treatment (1952) illustrates the confusion that 
still exists in this species-group. He ascribes a salt or brackish 
marsh habitat to both S. campanulata and S. stellaris and includes 
the West Indies in the range of the former species rather than 
that of the latter. Furthermore he states that S. stellaris 
“and S. campanulata constitute the local extremes of a variable 
population which may represent only one species or as many 
as four. Even in our range numerous intermediate forms 
occur and their assignment to a species depends on the general 
prominence of characters of which the proportion of the petals 
is probably the most reliable. In the southeastern part of our 
range and in the southeastern states both species pass into forms 
with lvs. all linear (S. gracilis (Michx.) Salisb.)." "The true 
picture is by no means as confusing or as complex as Gleason 
suggests. Sabatia stellaris and S. campanulata are separable by 
strong morphological features and the two species occupy 
mutually exclusive habitats. 

It should be pointed out that while S. stellaris is rather closely 
confined to the littoral region in the northern part of its range, 


1955] Wilbur,— The North American Genus Sabatia 61 


specimens from inland peninsular Florida are fairly numerous. 
In spite of the information available in the literature, S. stellaris, 
at least in the southern part of its range, 1s not restricted to 
a salt-marsh or littoral environment and in fact is at least more 
commonly collected in inland sandy areas. 

Britton (Bull. N. Y. Bot. Gard. 3: 448. 1905.) and later 
Britton & Millspaugh (1920) recognized two species as occurring 
in the Bahamas, S. campanulata and the supposedly endemic 
S. simulata. This latter species was to be distinguished from 
what has been called S. campanulata by 1ts more slender habit 
and smaller white flowers. All the material seen from the 
Bahamas has proven to be annual and consequently the larger- 
flowered plant with the rose-pink corollas is definitely not S. 
campanulata; it is S. stellaris. There is no break in flower-size 
between the pink and white flowered specimens although the 
latter are on the average smaller. Several of the larger-flowered 
specimens (2.5 cm. or more in diameter) had been designated 
by Britton as S. simulata apparently on the basis of the flower 
color. The stigma of S. simulata was described as ''spatulate" 
in contrast to the "oblong-linear" stigma of “S. campanulata.” 
I was unable to discover any consistent difference between 
the stigmas of the specimens. There was the usual variation 
dependent upon age. S. simulata is, I believe, merely a white- 
flowered form of S. stellaris and is conspecific with the pink- 
flowered specimens of the Bahamas. These color-phases are 
indistinguishable from much of the material from southern 
Florida and, for the present at least, all had best be treated as 
but another trend in the variation apparent in this species 
throughout its extensive range. 

One name deserving brief discussion 1s S. nana of Featherman, 
which is not listed in “Index Kewensis." The description is 
here quoted in full: 


Sabbatia Nana.—Stem simple, low, somewhat angled. Leaves small, 
sessile, the lower spatulate lanceolate, the upper linear lanceolate. Lobes of 
the corolla five to six, one third longer than the narrow linear calyx lobes. 
Root perpendicular, slender. Flowers rose-colored. Stem three to four 
inches high. Blooms in August. 

Habitat—Marshy soil of Grande Isle. 


The diagnosis is certainly brief but the shape of the leaves, the 
‘narrow linear” calyx-lobes two-thirds as long as the corolla, 
the habit and the habitat all together make it quite certain 


62 Rhodora [VoL. 57 


that S. stellaris was the plant being described. This Louisianan 
record is at or near the westernmost extremity of the species’ 
present known range in the southeastern states. 

Type-specimens of S. maculata, S. palmeri, and S. purpusii 
have all been examined and compared both with the few addi- 
tional Mexican collections of the genus that have been made 
and specimens of the widespread species of eastern North Amer- 
ica. Two of the species seemed very different from each other 
and at first I thought that I could easily distinguish them both 
from any others in the genus. Sabatia palmeri, on the other 
hand, very quiekly seemed to be indistinguishable from 5. 
maculata. After examining more material, especially from 
Florida, it became obvious that any of the Mexican specimens 
could be matched or very nearly matched by a considerable 
number of plants of 5. stellaris. I am certain that I could not 
distinguish a large unlabeled series of specimens made in Florida 
and another large series made in Mexico. Therefore, all three 
names are here treated as synonyms of S. stellaris, a wide- 
ranging, variable species. 

The blackened patches at the base of the corolla-lobes of the 
type of S. maculata are very striking and the epithet chosen 
by Bentham points out this feature. However, the dark black 
color ean almost certainly be attributed to discoloration upon 
drying and has been noted in many specimens found as far 
north as New Jersey. The type itself seems indistinguishable 
from many specimens from Florida of which Cuthbert 1525 
(FLAS) from Manatee Co. is a good example. The type of 
S. palmeri is very similar in appearance to that of S. maculata. 

S. purpusit is indistinguishable from many of the wiry Floridian 
specimens. For example, Pennell’s collection (18048) from the 
state of San Luis Potosí seems remarkably similar to the collection 
of Hood 3565 (FLAS) among others from western peninsular 
Florida. In the Mexican specimens a tendency was noted 
for the calyx-tube to be more cylindrical with sides more parallel 
rather than strongly tapering and whose length is typically 
in the upper range of that of the specimens from the eastern 
United States. Also, the few capsules seen in the Mexican 
specimens of the purpusii-aspect appeared to be more slender 
and longer than those of most of the specimens seen from the 
principal area of distribution of the species. 


1955] Wilbur,—The North American Genus Sabatia 63 


The disjunct distribution of this species is somewhat perplexing. 
Fernald (RHopora 18: 146. 1916.) stated that the range of 
S. stellaris was from “Massachusetts to eastern Mexico" and 
at that time he considered the plants called by some S. purpusiz 
as part of the wide ranging species as is shown by his annotation 
labels. In the most recent edition of Gray’s Manual the western 
limit of the range was stated to be Louisiana. The apparent 
absence of this species from the Texas coast and its eventual 
reappearance in eastern Mexico far south of the border and, 
most surprisingly of all, in the central plateau of that country 
is not easily explained. It was with considerable reluctance 
that attempts to distinguish these species were abandoned but 
there seemed to be no alternative. The absence of the plant 
from Texas is in particular inexplicable as there seem to be 
enormous stretches of territory offering an environment apparent- 
ly similar to that in which it is found so abundantly in the 
Atlantic and Gulf Coastal Plain. 


REPRESENTATIVE SPECIMENS:—MASSACHUSETTS: Bristol Co., Dartmouth, 
Hervey (au); Dukes Co., Tisbury, Cushman, July 1911 (wis), RHODE ISLAND: 
Washington Co., Westerly, Weatherby 3311 (NCS). CONNECTICUT: Fairfield 
Co., Stratford, Eames, 16 Aug. 1895 (NCU, NY, US). NEW YORK: Kings Co., 
Barren Island, Svenson 8242 (GA, GH, MO, PENN, TENN); Richmond Co., 
vicinity of New Dorp, Small, 2 Aug. 1890 (F, GH, MO, NA, NY, US, YU); Suffolk 
Co., mouth of Wading River, Muenscher & Curtis 6420 (cU, GH, MT, NY, Us). 
NEW JERSEY: Cape May Co., Cold Spring, Pennell 2179 (an, Mo, vs); Salem 
Co., about 4 mi. s. of Canton, Fogg 6054 (GH, PENN, TENN). DELAWARE: 
Sussex Co., 2 mi. s. of Rehoboth, Larsen 989 (GA, PENN, US). MARYLAND: 
Caroline Co., between Choptank River and Bethlehem, Killip 7278 (an, vs). 
VIRGINIA: Princess Anne Co., along Back Bay, Fernald & Long 11113 (DUKE, 
GH, MO, NY, US); Surry Co., Hog Island, Fernald & Long 12774 (GA, GH, MO, 
US). NORTH CAROLINA: Brunswick Co., near Southport, Biltmore Herb. 8304a 
(GH, Mo, NCU, NY, Us); New Hanover Co., Fort Fisher, Godfrey 4726 (DUKE, 
GH, NCU, NY, US). SOUTH CAROLINA: Georgetown Co., Pawleys Island, 
Godfrey & Tryon 306 (DUKE, F, GH, MO, NY, PENN, TENN, US); Horry Co., 
about 2 mi. sw. of Ocean Drive, Wilbur 2901 (micH). GEORGIA: Chatham Co., 
Tybee Island, Harper 924 (an, Mo, NY, US); Glynn Co., Sea Island, Cronquist 
5356 (FLAS, GA, GH, NO, SMU, US). FLORIDA: Brevard Co., Titusville, Nash 
2308 (F, FLAS, GH, K, MICH, MO, US); Lee Co., vicinity of Fort Myer, Standley 
12 (F, GH, MO, NY, US); Levy Co., 1 mi. se. of Chiefland, Webster & Wilbur 
3639 (micu); Monroe Co., Big Pine Key, Small 8148 (DUKE, GH, NCU, NY, 
PENN, WVA); Orange Co., about 2 mi. e. of Fort Christmas, Moldenke 210 
(DUKE, MO, NY, PENN, US); Sarasota Co., Longboat Key, Tracy 6808 (F, MO, 
NY, US). ALABAMA: Mobile Co., saline marshes, Mohr, Aug. 1892 (NY). 
MISSISSIPPI: Harrison Co., Cat Island, Tracy, Earle & Seymour 101 (cu, an, 
MO, NCU, WIS). LOUISIANA: Plaquemines Parish, Breton Island, Tracy & 
Lloyd 343 (cv, F, GH, MO, NY, US). BAHAMA ISLANDS: Abaco Is., Brace 1793 
(xv); Andros Is., Brace 7116 (rF, NY, vs); Cat Is., Britton & M illspaugh 5780 


64 Rhodora (Vor. 57 


(r, NY); East Bimini, Howard 10274 (smu, us); Eleuthera Is., Britton 6427 
(F, GH, NY, US); Great Bahama Is., Brace 3518 (F, Ny, us); New Providence 
Is., Britton 58 [ryeE of S. simulata Britt.] (Ny); Curtiss 200 (F, GH, MO, NY, 
US). CUBA: Without more exact locality but very likely from Pinar del Rio, 
Wright 412 [sheets sometimes mixed with S. grandiflora] (BRU, GH, NY, US). 
This is the same number but from a different collecting trip as the type of 
S. gracilis var. cubensis Griseb. (= S. calycina). MExico: Coahuila: Cuatro 
Cienagas, Marsh 2068 (F). | Michoacán: 60-70 mi. e. of Lake Pátzcuaro in bog 
4 mi. w. of Hidalgo, Hitchcock & Stanford 7196 (GH, vc). Jalisco: Lagos de 
Moreno, Hartweg 1615 [rype of Eustoma maculata Benth.] (x); Rio Blanco, 
Palmer 668 [ryrE of S. palmeri Gray] (GH). Querétaro: Querétaro, Arséen 
10281 (vs). San Luis Potosí: Media Luna near Rio Verde, Palmer 80 (F, 
GH, MO, NY, UC, US) and Palmer 80.5 (GH, Mo, Ny, US); Minas de San Rafael, 
Purpus 5845 [TYPE of S. purpusit Brandegee at uc] (F, GH, MO, NY, UC); 
Hacienda de Angostura, Pringle 3810 (BRU, F, GH, MICH, MO, NY, UC, US); Las 
Tablas, Pennell 18048 (us). Tamaulipas: Jaumave, Rozynski 377 (F). 


10. Sabatia grandiflora (A. Gray) Small, Fl. SE. U. S. 928. 1903. 
Sabbatia gracilis var. grandiflora A. Gray, Syn. Fl. N. Am. 2: 115. 1878. 
Sabatia campanulata var. grandiflora (A. Gray) Blake, Rnopona 17: 52. 
1915. Sabbatia Alainii Marie-Victorin, Contr. Inst. Bot. Univ. Montreal 
No. 63: 73. 1948. 

Erect annual (15-)40-90(-110) em. high; stem (1-)2-5(-7) mm. in 
diameter, terete, although in dried specimens with several fine lines run- 
ning between the nodes. Branches, rarely absent, almost entirely re- 
stricted to the upper third or half of the stem and almost invariably 
alternate, typically strongly divergent, forming angles of about 50-70 
degrees, stiffly geniculate, bearing but few nodes. Plant appearing vir- 
gate from elongate, rigid internodes. Root-system of several to numerous 
fibrous roots 3-8 em. long, 1-3 mm. in diameter. Leaves erect, at least 
above closely appressed against the stem, succulent, drying thick, rigid, 
rugose-roughened, venation obscure in both living and dried material; 
the lower spatulate, elliptie, oblong-linear or even linear, obtuse to acute, 
(1-)2-4(-5) em. long, (2-)4—7(-10) mm. wide, typically about 5-10 times 
longer than wide, usually several times exceeding the typically short, 
0.5-3 em. long internodes; gradually to abruptly reduced above to very 
narrowly linear or even filiform, the width noticeably less than the diam- 
eter of the stem, (1-)3-5(-12) em. long, (0.5—)1-2(-3) mm. wide, generally 
20-60 times as long as wide, hyaline or callose-apiculate. Internodes 
above typically greatly elongate, often 2-5 times as long as the leaves. 
Inflorescence of (1-)2-3(-4) or rarely more flowers in reduced cymules; 
the flowers borne on slender, but rigid, elongate, terete pedicels (2-)4-8 
(-12) em. long and about 0.5 mm. in diameter; the flowers thus appearing 
solitary. Calyx-tube campanulate, usually broadly so, the sides more or 
less parallel, abruptly contracting to base or but gradually tapering in 
the lower half, typically 1—1.5(-2) times as long as broad, 2-4(-6) mm. 
long, generally very smooth or but very finely lined, the wall thin and 
somewhat hyaline-scarious especially with age. Calyx-lobes erect to 
spreading, very narrowly linear or filiform, (0.6—)1—2(—3) em. long, usually 


1955] Wilbur,—' The North American Genus Sabatia 65 


S. GRANDIFLORA 


S. BREVIFOLIA 


Maps 9-12. Map 9, upper left; map 10, upper right; map 11, lower left; map 12, 
lower right. 
about 3-6 times as long as the tube, typically less than three-fourths as 
long as the corolla, rarely equaling or exceeding it, the tip callose-apiculate. 
Corolla-tube cylindrical, (3-)5-7(-8) mm. long, exceeding calyx-tube by 
about 2-4 mm., usually 1.5-2 times as long as wide, apparently colorless. 
Corolla-lobes strongly spreading, oblanceolate, obovate, broadly spatulate, 
oblong, or elliptic, usually obtuse or more rarely acute, (1.3-)1.8-2.5(—3) 
cm. long, (5-)7-12(-15) mm. wide, deep rose to pale-pink or white, with 
oblong basal vellow area 2-4 mm. long, rather irregularly lobed or toothed, 
and in the roseate forms usually bordered by an intense red line or area. 
Anthers linear, dark yellow, (3-)5-7(-8) mm. long; filaments yellow, 
slender, 2-4 mm. long. Style 2-5 mm. long; stigmatie lobes (4-)6-8(-9) 
mm. long. Capsule usually narrowly cylindrical, (6-)8-10(-15) mm. long, 
4-5(-7) mm. wide, generally 2-3 times as long as broad. TYPE LOCALITY: 
“Coast of E. Florida." Lectotype: Palmer 430 (GH!) from ‘Indian 
River." DISTRIBUTION: Everglades, pinelands and sandy places in penin- 
sular Florida and western Cuba. Map 10. 


In spite of the numerous striking characters which distinguish 
this species from any other, too much reliance has been placed 


66 Rhodora [Vor. 57 


upon floral size as the criterion for the recognition of S. grandi- 
flora. As a consequence one finds under this name in most 
herbaria a miscellaneous assemblage of large-flowered specimens 
belonging to subsection Campanulatae. Small never pointed 
out the distinctive characteristics of S. grandiflora and a con- 
siderable number of the specimens labeled or annotated by him 
are merely large-flowered S. stellaris. The identity of these 
incorrectly determined specimens is shown by the smooth, 
thin texture of the dried leaves which above, although narrowly 
linear, are typically as broad or broader than the diameter 
of the stem. Also these large-flowered specimens of 5. stellaris, 
which have been so often mistaken for S. grandiflora, are generally 
more profusely branched, especially below the middle of the 
stem. S. grandiflora may be readily distinguished from S. 
stellaris by its thick rugose-roughened or wrinkled leaves which 
above are almost filiform and less in width than the diameter 
of the stem. Furthermore, it is typically a larger and coarser 
plant with branching more or less restricted to the upper third 
or half of the stem. Another fairly reliable feature that may 
be utilized in recognizing S. grandiflora is that its calyx-tube 
is most often broadly campanulate, while that of S. stellaris 
is more typically narrowly turbinate. That much-relied-upon 
character, the length of the corolla-lobes, is at best a subsidiary 
one because of the considerable overlap in the size of flowers 
between the two species. 

In spite of the inferences that might be derived from the 
synonymy, the relationship of S. grandiflora is closer to S. 
stellaris than to S. campanulata (= S. gracilis). Numerous 
differences might be listed to distinguish S. grandiflora from 
S. campanulata with which it was originally and even later 
considered to be a variety. However, the most obvious and 
perhaps most fundamental is that the latter is a caespitose 
perennial while S. grandiflora, like S. stellaris, is an annual. 
The perennial species is not known in peninsular Florida. S. 
stellaris has been collected throughout the state, while in Florida, 
S. grandiflora is almost entirely restricted to the peninsular 
portion. Judging from the number of mixed sheets of S. stellaris 
and S. grandiflora, there is considerable overlap in range of 
habitat between these two species if their environmental toler- 
ances in Florida are not exactly the same. 


1955] Wilbur,—The North American Genus Sabatia 67 


In the United States other than in Florida, S. grandiflora 
is represented only from Louisiana by a specimen attributed 
to Josiah Hale deposited in the Gray Herbarium. This col- 
lection is S. grandiflora but more recent collections from Louisi- 
ana seen in the herbaria under this name have always proven 
to be large-flowered specimens of S. stellaris. Confusion in 
labeling should be considered a strong possibility for this speci- 
men reputedly from Louisiana. 


After the short original diagnosis, Gray presented the col- 
lections known to him in the following manner: ‘coast of E. 
Florida, Leavenworth, Buckley, Palmer & c.". Theirs are the 
only specimens available to him now preserved in either the 
Gray Herbarium or that of the New York Botanical Garden; 
who the other collectors were is unknown to me. Palmer’s 
collection, made in 1874, is definite and all four sheets at the 
three different herbaria bear the information “Indian River, 
East Florida" as well as the number ‘430.” These specimens 
are also equal to or better than the others and, in spite of being 
collected after a varietal status had been decided upon (with 
another epithet) based on the other collections, Palmer's col- 
lection (no. 430) at the Gray Herbarium is here designated the 
lectotype. 

The recently proposed S. alainii from Cuba here is considered 
to be but a synonym of S. grandiflora; its distinctness has cer- 
tainly not as yet been demonstrated. The original description 
of this supposedly distinct Cuban entity was accompanied by 
the following discussion in reference to its relationship with 
S. grandiflora. ‘Cette espèce se rapproche du Sabbatia grandi- 
flora par sa grande corolle, ses curtes feuilles, l'absence d'oeil 
jaune, sa capsule plus petite." The meaning of this is not 
altogether clear. It would seem as though a word or so had 
been omitted in printing and that the first two characters men- 
tioned were meant as features shared in common by S. grandi- 
flora and the newly proposed species and that the second two 
phrases were supposed distinctions. The stated absence of 
the yellow eye in the throat of the corolla-tube and at the base 
of the lobes of S. alainii is not borne out by examination of 
either the type material, distributed isotypes or information 
(“white with light yellow eye") provided on the label of Britton 
et al. 7166 which was collected in the same region. The few 


68 Rhodora [Vor. 57 


capsules available were smaller but did not appear to have 
fully developed. The Cuban representatives of this group 
are, I believe, indistinguishable from those of Florida. 


REPRESENTATIVE SPECIMENS:—FLORIDA: Alachua Co., Earlton Beach, 
Loucks, 5 Sept. 1927 (Frias); Brevard Co., between Cocoa and Lake Poinsett, 
Rhoads, 23 May 1937 (Frias); Broward Co., Ft. Lauderdale, Small & Wilson 
1785 (Ny); Charlotte Co., without exact locality, Frye, 17 May 1946 (Frias); 
Clay Co., Keystone Heights, West, 29 June 1930 (FLAs); Collier Co., 2 mi. s. of 
Immokalee, Sheehan 28 Feb. 1919 (Ny); Dade Co., w. of Camp Jackson, 
Small & Wilson 1991 (Ny); DeSoto Co., Lacy, West, 23 Sept. 1938 (FLas); 
Duval Co., without exact locality, Fredholm 5 (au); Glades Co., Lakeport, 
Lovett 264 (DUKE); Hardee Co., Limestone, Kirk, 1 July 1942 (FLAs); Hendry 
Co., se. corner of county, Davis, 13 Sept. 1947 (FLAs); Hernando Co., without 
exact locality, Hitchcock 1283 (v, Mo); Highlands Co., near Brighton, McFarlin 
10913 (GH, PENN); Hillsborough Co., Tampa, Garber, May 1876 (BRU, F, NY, 
us, YU); Indian River Co., near Felsmere, Small 8917 (Ny); Lake Co., Eustis, 
Nash 768 (cv, F, GH, MICH, MO, NY, US); Lee Co., about 8 mi. se. of Fort Myers, 
Standley 446 (F, GH, MO, MT, vs); Levy Co., Bronson, Watson & Murrill, 18 
June 1939 (rLas); Manatee Co., near Palma Sola Bay, Cuthbert, 23 June 1916 
(FLAS); Marion Co., Lake Kerr, West & Arnold, 21 July 1935 (rLas) Martin 
Co., Stuart, Atwood 1917 (cv); Okeechobee Co., n. of Fort Drum, West, 22 
Apr. 1946 (FLAS); Orange Co., near Oakland, Curtiss 6624 (CU, GH, MO, NY, 
us); Osceola Co., Kissimmee, Singletary 136 (pUKE, Ncv); Palm Beach Co., 
along the Palm Beach Canal, Small 8250 (FLAS, NY, TENN); Pasco Co., near 
Zephyrhills, Hood 3483 (rr4s); Pinellas Co., near St. Petersburg, Deam 1901 
(MicH); Polk Co., n. of Davenport, Hood 3546 (FLAS); Putnam Co., Welaka, 
DeVall, 23 June 1939 (rLas); Sarasota Co., Myakka River State Park about 
18 mi. se. of Sarasota, Wilbur & Webster 2502 (wicum); Volusia Co., near 
Ariel, Moldenke 180a (Ny, PENN); County unknown, Indian River, Palmer 
430 [LeEcTOTYPE of S. gracilis var. grandiflora] (F, GH, NY). CUBA:—Pinar 
Del Rio: au bord des lagunes de Santa Maria, San Luis, Marie-Victorin & 
Alain 369 [rve& of S. Alainii] (an, wr); Borders of Laguna Santa Maria, 
Britton, Britton & Gager 7166 (Ny, us). Province not stated, Wright 412 
[sheets sometimes mixed with S. stellaris] (BRU, NY, US). 


11. Sabatia brevifolia Raf., Atl. Journ. 1: 147. 1832. S. Elliotii 
[Elliottii] Steud., Nom. Bot. Ed. 2. 2: 489. 1841. S. paniculata B Elliottii 
[Steud.] Wood, Am. Bot. & Flor. 266. 1870, without basionym. 

Erect annual (15-)30-60(-70) em. high; stem (1-)2-3(-4) mm. in 
diameter, terete, but usually finely ridged or lined by slight elevations 
extending between the nodes, pith hollow, at least below. Branches 
typically alternate, always so above, but in robust, profusely branched 
plants occasionally with few to several branches of the main stem and even 
near base of principal branches opposite; in large well-developed plants 
usually branching throughout the length of the stem, in smaller plants 
often restricted to the upper half of the stem, strongly ascendent to more 
commonly widely divergent, usually forming an angle with the stem of 
(20-)40-60(-80) degrees, slender, very wiry, usually bearing numerous 
nodes, the internodes not elongate; the aspect of the plant strikingly 
wiry-virgate. Root-system a pronounced slender or occasionally thick- 


1955] Wilbur,—The North American Genus Sabatia 69 


ened tap-root 2-8 em. long, (1-)2-3(-4.5) mm. in diameter, bearing few 
to several very slender laterals. Leaves ascendent and apparently ap- 
pressed on the main stem and the more or less vertical branches but on 
strongly divergent branches usually strikingly upwardly secund and often 
appearing faleate, midvein alone easily detected and that elevated be- 
neath, apex very minutely apiculate, the lowermost somewhat obovate, 
oblanceolate, oblong, elliptic, or linear, (0.5-)1-2(-3) em. long, (1-)2-5(-7) 
mm. wide, typically obtuse or sometimes acute, usually tapering to the 
sessile base, generally 3-6 times as long as wide; those from about the mid- 
dle of the stem narrowly oblong or more commonly linear, 1.5-2.5 em. long, 
1-3 mm. wide, acute; the upper cauline leaves and those of the branches 
narrowly linear to filiform or even subulate, gradually reduced to about 
3-5 mm. long and less than 0.5 mm. wide, and almost invariably less than 
2 mm. wide or twice the diameter of the stem but often the same width, 
usually 10-25 times as long as broad. Internodes usually 1-2 times the 
length of the leaves, occasionally greater or sometimes, especially near the 
base, less. Flowers appearing solitary but arranged in reduced often 
1-2-flowered usually secund eymules on the upper side of the divaricate 
branches. Pedicels very slender, wiry, erect (1—)2-4(-5) em. long. 
Calyx-tube turbinate, sides sloping to the base, smooth or more commonly 
noticeably finely nerved, internerval wall thin, often scarious, about 1—2 
times as long as broad, (1-)1.5-2.5(-3) mm. long. Calyx-lobes setaceous 
or subulate, ascendent to wide spreading, (3-)4-7(-8) mm. long, tapering 
from about 0.5-1 mm. base, usually 2-4 times as long as the calyx-tube, 
usually not exceeding half the length of the corolla. Upper half of corolla- 
tube usually cylindrical, tapering below middle to the base, about 1.5-2 
times as long as wide, usually 1-2 mm. longer than the calyx-tube or 
about 2 times as long, (2.5-)3.5-4.5(-5) mm. long. Corolla-lobes spread- 
ing, elliptic, oblong, oblanceolate, or broadly spatulate, (0.6-)0.9-1.3(-1.8) 
em. long, (2-)3-5(-7) mm. wide, usually obtuse but not uncommonly 
somewhat acute, white with a greenish-yellow to yellow patch at base of 
lobe. Anthers linear, yellow, (2-)3-5(-6) mm. long; filaments greenish- 
yellow to pale yellow, 1-2 mm. long. Style 1 mm. long or less; stigmatic 
lobes 3-5 mm. long. Capsule cylindrical, 3-6 mm. long, 2-4 mm. wide. 
TYPE LOCALITY: Florida. Type: unknown. Described by Rafinesque 
along with thirty other species from Florida *'seen in gardens and herbals.” 
DISTRIBUTION: Pinelands and savannas from southeastern South Carolina 
along the Coastal Plain throughout most of Florida and west at least to 
Alabama. Map 1l. 


This species was described originally by Elliott with the 
acknowledged aid of Baldwin but the name applied to it was 
Sabbatia paniculata. This erroneous nomenclature was gen- 
erally employed even after it was understood to be a mistake. 
Chapman (1860) was persuaded by Gray to take up the sub- 
stitute name first proposed by Steudel nearly twenty years 


70 Rhodora [Vor. 57 


before. Elliott’s application of Michaux’s epithet which initi- 
ated the confusion is curious in view of his understanding of the 
subject as shown by the following quotation: 

Though the description of Michaux applies more peculiarly to the S. 
corymbosa, yet as this species was certainly included, and is the only one to 
which the term paniculata is certainly applicable, I have referred to him here. 

This change in concept of the species passing as S. paniculata 
was pointed out by Grisebach (1839) but he chose to continue 
the use of the name in the sense of Elliott. Steudel simply 
substituted the epithet elliottii (as Elliotii) citing in synonymy 
"S. paniculata Ell. (non Pursh)." The original spelling of the 
name need not be retained in a case such as this as it was clearly 
an unintentional orthographic error. Steudel, in listing Stephen 
Elhiott’s name, spelled the name, “Eliot.” It seems hardly possi- 
ble that anyone would carry the prerogative of modifying a name 
when latinizing it to a bibliographic listing; the change must 
have been unintentional. Wood's solution to the confusion 
that had enveloped the nomenclature of the white-flowered 
sabatias was to reduce S. elliottii to varietal status of S. panicu- 
lata. This indicated no more than a vague bibliographie rather 
than biological familiarity with the plants. 

Nine years before Steudel's publication of S. elliottii, Rafin- 
esque in a paper describing over thirty species from Florida be- 
longing to various genera ''seen in gardens and herbals” published 
S. brevifolia, the original description of which is here quoted 
in full: 


8. Sabbatia brevifolia Raf. Stem dichotomus filiform leaves short subulate 
acute, flowers terminal white, calyx shorter than corolla setaceous, segments 
of corolla obovate. Near to S. brachiata and stellaris. 

This description, although very brief, seems very apt in 
pointing out the stronger features that characterize the species 
that has long been called S. elliottii. Even in the absence of an 
authentic specimen I feel no hesitation in taking up Rafinesque's 
name other than that caused by the regret of seeing a well- 
established name replaced by one long-forgotten. It would be 
difficult to present a better description in so few words of the 
plant first characterized by Elliott than Rafinesque’s attempt. 
Every descriptive phrase fits Elliott's plant very well. 


REPRESENTATIVE SPECIMENS:—NORTH CAROLINA: without locality, Delile 
(Nv). Since this species is not otherwise known north of Charlestown, S. C., 


1955] Palmer,—Epifagus virginiana 71 


its presence in North Carolina seems most doubtful SOUTH CAROLINA: 
Beaufort Co., Bluffton, Mellichamp, 1886 (r, Mo, Ny, vs); Charleston Co., 
Adams Run, Godfrey & Tryon 1544 (DUKE, F, GH, MICH, MO, NY, PENN, TENN, 
us); Hampton Co., about 4 mi. se. of Hampton, Wilbur & Webster 2826 
(MrcH); Jasper Co., Ridgeland, Mohr, Nov. 1893 (Mo, NY, US). GEORGIA: 
Camden Co., w. of St. Marys, Small, DeW inkeler & Small 10545 (DUKE, WVA); 
Charlton Co., Okefenokee Swamp between Chesser Island and Camp Cornelia, 
Thorne & Ford 2064 (cv, Ga); Coffee Co., without exact locality, Harper 681 
(NY, vs); Pierce Co., 3 mi. n. of Blackshear, Thorne & Norris 6202 (cu, GA); 
Ware Co., 5 mi. se. of Waycross, Wilbur & Webster 2731 (MICH). FLORIDA: 
Brevard Co., Merritts Island, Curtiss 2227 (F, GH, MO, Ny, us); Clay Co., 
about 2 mi. n. of Orange Park, Moldenke 162 (DUKE, MO, NY, PENN, US); 
DeSoto Co., 4 mi. w. of Arcadia, Webster 4224 (micu); Duval Co., near Jack- 
sonville, Curtiss 5114 (cU, F, GH, NCS, NY, US); Franklin Co., Apalachicola, 
Biltmore Distrib. Chapman Herb. 903b (GH, MO, NCS, NCU, NY, PENN, US); 
Hernando Co., between Brookside and Bayport, Jones 43 (cu, vs); Hills- 
borough Co., Tampa, Nash 2422 (F, GH, MICH, Mo, MT, NY, US); Indian River 
Co., near Felsmere, Small 8902 (Fuas, GH, NY, US); Jackson Co., 4 mi. e. of 
Marianna, Wiegand & Manning 2559 (cu, au); Lake Co., near Cassia, Hood, 
1 Sept. 1911 (FLAs, GH); Lee Co., s. of Fort Myers, Moldenke 909 (DUKE, Mo, 
NY, PENN, Us); Liberty Co., near Roy, Wiegand & Manning 2558 (cv, an); 
Manatee Co., Bradenton, Tracy 7080 (CU, F, FLAS, GH, NY, PENN, TAES, US); 
Orange Co., about 2.5 mi. ne. of Apopka, Wilbur & Webster 2650 (MicH); 
Palm Beach Co., Earman, Randolph 26 (cv, GH); Pinellas Co., about 4 mi. w. 
of St. Petersburg, Deam 2840 (micu, vs); Putnam Co., 10 mi. se. of Inter- 
lochen, Fox 5700 (micu, Ncs); Seminole Co., Lake Monroe, Garber, Mar. 1896 
(BRU, F, US, YU); Volusia Co., Beresford, Hood, 21 Sept. 1910 (FLAs, Mo). 
ALABAMA: Mobile Co., between Theodore and Hollander’s Island, Pennell 
4512 (NY, PENN). LOUISIANA: without locality data, Featherman (mo). 
This species should not be included in the flora of the state without a better 
substantiated record. It seems likely that there is confusion in labeling in 
regard to this collection. 
(to be concluded) 


EPIFAGUS VIRGINIANA.—On the northern exposure of a 
deciduous woods, consisting chiefly of oaks and beeches, there 
is at Swarthmore, Delaware County, Pa., a large colony of 
Epifagus which is butter-yellow in color. Nearby a number 
of hemlocks are growing, but this plant is not found under 
these, but confines itself to the deciduous trees. A year ago 
I counted 300 of this yellow Epzfagus without materially changing 
my position, and this year (’54) I counted 175 plants along 
a wood’s lane within the space of 100 feet. A fair estimate 
of the plants on this northern hillside would be more than 1000. 

The plant resembles the species closely except for color. 
If any difference is to be noted I think the yellow form tends 
to be more bushy and is perhaps more fleshy. However, there 


72 Rhodora [Vor. 57 


are numbers of single-stemmed plants scattered about. Some 
of these are pure yellow, but others show touches of reddish 
brown on the stem scales. 

The usual color-formof E. virginiana does not appear within 
the area occupied by the main growth of the yellows, but as the 
yellows diminish along the edges the usual form appears. "The 
whole area is about an acre in extent. 

'The concentration of the yellow forms on this hillside suggests 
a common origin for them, but does not explain the absence 
of the usual form. So many plants of this one color within the 
limited area indicated with the usual form of the species on the 
periphery offers à nice problem of distribution and might provide 
an interesting population study for the appropriate person.— 
SAMUEL C. PALMER, SWARTHMORE, PA, 


CALYCERA BALSAMITAEFOLIA IN THE UNITED STATES.— 
Recently there were found in the Herbarium of the Chicago 
Natural History Museum two sheets of Calycera balsamitaefolia 
(Juss.) Rich., of the family Calyceraceae. "They were included 
in the J. T. Rothrock collection, when that herbarium was 
purchased by the Chicago Natural History Museum in 1909. 
Both sheets were collected on ballast at Kaighns Point (spelled 
Kaigns on the label) in New Jersey. This station is near Cam- 
den opposite Philadelphia. One sheet has the label of ‘Her- 
barium of Isaac Burk, Philadelphia; Pa.", the other sheet ac- 
companied by a label of the “Herbarium of University of Penn- 
sylvania." No date is indicated on either label. 

It is of interest to note this collection, because the species is a 
native of Chile, and is not mentioned in either the eighth edition 
of Gray’s Manual, Gleason’s New Britton & Brown, or any 
of the floras of New Jersey and Pennsylvania. Probably col- 
lected at the station given above in the latter decades of the 
nineteenth century or at the beginning of the twentieth century, 
the species apparently has never been re-collected either at the 
Kaighns Point station or elsewhere. Botanists from the vicinity 
of Philadelphia and New York should give special attention to 
ballast sites similar to the one where Calycera was found.— 
JULIAN A. STEYERMARK AND FLOYD A, SWINK, CHICAGO NATURAL 
HISTORY MUSEUM AND COLLEGE OF PHARMACY, UNIVERSITY OF ILLINOIS, 


Volume 57, no. 673, including pages 1-86, was issued 28 February, 1955. 


Hovova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


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REED CLARK ROLLINS. Editor-in-Chief 


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CARROLL EMORY WOOD, JR. - 


Vol. 57 March, 1955 No. 675 
CONTENTS: 
A Note on the Name Calamintha. Gordon P. DeWolf, Jr....... 73 


A Revision of the North American Genus Sabatia (Gentianaceae). 
Robert L. Wilbur (continued from page 71) ................ 78 


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Vol. 57 March, 1955 No. 675 


A NOTE ON THE NAME CALAMINTHA 
Gorpon P. DEWorr, Jn. 


THE genus Satureja, L. as interpreted in the 8th edition of 
“Gray’s Manual of Botany" includes plants that have been 
included in four genera: saTUREJA (S. hortensis L.); CALAMINTHA 
(S. Calamintha, (L.) Scheele — C. officinalis Moench; S. glabella 
(Michx.) Briq. = C. glabella (Michx.) Benth.; S. arkansana 
(Nutt.) Briq. = C. glabella (Michx.) Benth., var. angustifolia 
(Torr. DeWolf [Satureja glabella (Michx.) Briq., var. angusti- 
folia (Torr.) Svenson]; AciNos (S. Acinos (L.) Scheele = A. 
arvensis (Lam.) Dandy; and cLinopopium (S. vulgare (L.) 
Fritsch — C. vulgare L.). 

Study of material of species of this group known to be in 
cultivation has convinced me that these four taxa are not con- 
generic, and that current European practice of recognizing them 
as distinct genera should be followed. A fuller discussion of 
the taxonomic aspects of the problem has been published else- 
where (Baileya 2(4): 142-150. 1954 [Jan., 1955]). Here I should 
like to discuss a purely nomenclatural problem. 

In the 52nd volume of Fedde's “Repertorium specierum novar- 
um regni vegetabilis," Heft 2, pp. 144—161, 1943, there appears 
the second of a series of articles by Erwin Janchen entitled 
“Zur Nomenclature der Gattungsnamen." My attention was 
drawn to this paper by the inclusion in the 1952 edition of the 
"International Code of Botanical Nomenclature," p. 131, of 
the name Calamintha, Lamarck as a nomen conservandum pro- 
positum, the respective nomen propositum rejiciendum being 
Clinopodium, Linnaeus. It seems to me that certain data 
have been overlooked in the framing of this proposal, and hence, 
further, and perhaps fuller, discussion is in order. 


74 Rhodora [Vor. 57 


In the 1943 paper, p. 156, no. 7305, Janchen refers to a paper 
by Janchen and Neumayer, entitled “Beiträge zur Benennung, 
Bewertung und Verbreitung der Farn- und Blütenpflanzen 
Deutschlands" in the “Oesterreichische Botanische Zeitschrift," 
volume 91, 1942, pp. 209-298, in particular to p. 274. At this 
place the identity of Calamintha, Moench (1794) [non Lamarck! 
1778] and Clinopodium, Linnaeus is asserted, and their dis- 
tinetness from Satureja, Linnaeus is affirmed. Further, Janchen 
and Neumayer stated: *. . . Mit Rücksicht auf die Artenanzahl 
ist Calamintha gegenüber dem älteren Gattungsnamen Clinopodi- 
um auf der Ausnahmsliste zu schützen . . ." They also refer 
to a paper in the “Acta Horti Gotoburgensis," volume 13, 1939, 
pp. 335-380, in particular p. 349, in which essentially the same 
thought is expressed by Handel-Mazzetti. 


Objections to this proposal arise on two points: 


1. Calamintha, Lamarck (1778) is a later homonym of Calamintha, 
Miller (1754) Calamintha, Trew (1754) and Calamintha, Adanson 
(1763) (and/or Scopoli, 1772), and is synonymous with Calamintha, 
Miller and Calamintha, Trew. 

2. Calamintha, whether of Miller, Trew, Adanson, Scopoli, Lamarck, 
or Moench, did not contain any species then ascribed to Clinopodium. 
The two taxa were not confused until 1891 when O. Kuntze reduced 
Calamintha, Moench emend. Bentham to Clinopodium, Linnaeus 
emend. O. Kuntze. 


Reference to Pfeiffer's “Nomenclator Botanicus,” volume 1, 


part 1, of 1873 indicates that Calamintha was first used as a 
post-Linnaean name by Adanson, in the “Families des Plantes" 
of 1763, volume 2, p. 192, for the Linnaean genus Glechoma. 
This same usage was maintained by Scopoli in the first volume 
of the second edition of the “Flora Carnioliea" (1772), at p. 423, 
where synonymy and a good description of the single species 
[Glechoma hederacea] are given. The next entry should be to the 
usage of Lamarek in the second volume of the first edition of 
the “Flore Francoise," 1778, p. 393, no. 432. Pfeiffer missed 
this, however, so the next actual entry is to Moench's “Methodus 
Plantas . . ." of 1794. 

A note about Calamintha, Moench is perhaps in order. In 
1794 the “Methodus Plantas Horti Botanici et Agri Marburgen- 
sis, a Staminum Situ Describendi" of Konrad Moench was 
published. In this work, at least so far as the labiates are 


1955] DeWolf,—A Note on the Name Calamintha 75 


concerned, very good generic and specific diagnoses are given. 
It was used by Bentham during the preparation of his ‘‘Labia- 
tarum, Genera et Species," of 1832-36, and, in particular, 
Bentham accepted Moench’s circumscription of Calamintha 
over that of Lamarck. As published by Moench, three species 
were accepted: C. grandiflora, (L.) Moench; C. officinalis, Moench; 
and C. trichotoma, Moench = C. Nepeta, (L.) Savi. 

Meanwhile, Clinopodium had received a very uniform treat- 
ment. In 1753 Linnaeus included three species in his genus, 
viz., Clinopodium vulgare; C. incanum = Pycnanthemum incanum, 
(L. Michx., fide Bentham, "Labiatarum . . ." p. 327; and 
C. rugosum = Hyptis capitata, Jacquin (ex Jamaica) and H. 
radiata, Willdenow (ex Carolina), fide Bentham, “Labiatarum 

. ." pp. 104 and 108. Of these three species, Clinopodium 
vulgare has been consistently retained in the genus since its 
publication, and its immediate relatives have been placed with 
it. 

In 1754 Philip Miller published a three volume, octavo, 
abridgement of the last folio edition (? 6th, 1752) of “The Gar- 
deners Dictionary." In this work binomial nomenclature was 
not used, but the generic descriptions are well drawn!. We 
are, therefore, faced with the necessity of considering Miller's 
genera. We find that Miller defined a genus he called “Cala- 
mintha.” The generic circumscription is tolerably specific, and, 
of the six taxa listed, the first three are the same as those accepted 
by Moench. They are: Calamintha vulgaris, vel officinarum 
Germanicae, of Caspar Bauhin = C. officinalis, Moench; Cala- 
mintha pulegii odore, foliis latioribus, of Paul Hermann = C. 
Nepeta, (L.) Savi; and the Calamintha magno flore, of Caspar 
Bauhin = C. grandiflora, (L.) Moench. 

When I discussed the taxonomic aspects of this problem in 
Baileya (l.c.) the name Calamintha was ascribed to Philip Miller. 
This occurred because of ignorance of the fact that also in 1754 
Christopher Jacob Trew, in his Nuremburg edition of Elizabeth 
Blackwell’s “A Curious Herbal," which is usually referred to 
as the "Herbarium Blackwellianum," had used the name Cala- 
mintha. Trew validated his name by reference to the generic 
descriptions of pre-Linnaean authors, including Ray and Tourne- 


! After Tournefort, fide G. C. Druce in Rpt. Botanical Exchange Club of the British 
Isles 3: 426-428, 1913. 


76 Rhodora [Vor. 57 


fort. Further he gave two excellent plates: no. 166, Calamintha 
moniana — C. officinalis, Moench and no. 167, Calamintha 
officinalis — C. Nepeta, (L.) Savi. 

In the past there has been a good deal of discussion as to the 
advisability of rejecting all names published in works dated 
after 1753 which do not use binomial nomenclature. As late 
as 1935 a formal proposal? was made to the 6th International 
Botanical Congress that such be done, and a list of proscribed 
works be made. This seems, however, to have met with no 
success. We are left, then, with the tacit understanding that 
generic names, so long as validly and legitimately published 
after 1753, may be taken up even from works not using binomial 
nomenclature. 

In the present case we are concerned with homonymous and/or 
synonymous names, published in the same year, but in unknown 
sequence. It behooves us, then, to take up for purposes of 
nomenclature, the most adequately circumscribed of the two 
names. Miller provides a generie description in English, and 
cited, with description, also in English, six species. Trew gave 
no generic description as such, but validated his name implicitly 
by reference to previously effectively published generic deserip- 
tions, especially those of Ray and Tournefort. Trew also gave 
two excellent plates of the two species which he accepted and 
full specific descriptions in Latin and German. Therefore, 
for purposes of citation I am accepting Calamintha, Trew. 

Lamarck’s circumscription of Calamintha on the other hand, 
is not good, nor does he refer to descriptions by previous authors. 
The genus may be identified with certainty only by recourse 
to the included species. Of these, the first two, C. alpina and 
C. arvensis, are now placed in Acinos; C. cretica is referred to 
Micromeria; and only the last two, C. parviflora and C. montana 
are now retained, under earlier names, in Calamintha (C. Nepeta, 
(L.) Savi and C. officinalis, respectively, fide Bentham “Labia- 
tarum . . ." p. 387-388). 

Until the publication of O. Kuntze (Revisio Generum Plan- 
tarum, volume 2, pp. 513-516) in 1891, there was little con- 
fusion of the two taxa which may be distinguished taxonomieally 
as follows: 


? Wilmott, A. J.—Kew Bull. 1935 66. 
3 Little, E. L.—Madrono 7: 240—242, 1944. 


1955] DeWolf,—A Note on the Name Calamintha 71 


Calamintha Clinopodium 
Calyx tubular, straight. Ver- Calyx tubular, curved. Ver- 
ticellasters on relatively long ticellasters sessile or nearly so. 


pedicels, sub-secund. 


After the genera were firmly established, with good circum- 
scriptions, by Moench in 1794, in the “Methodus Plantas . . .," 
they were generally treated as taxa of correlative rank.  Ben- 
tham treated them as sections of Melissa, L. in the *Labiatarum 

. ." of 1832-36, and as sections of Calamintha, Moench emend. 
Bentham in the 12th volume of de Candolle's "Prodromus 
Systematis Naturalis . . ." of 1848. Briquet, in IV Teil, 3 
Abteilung, a, of" Die natürlichen Pflanzenfamilien" of 1897, 
treated them as sections of Satureja, L. Neither Bentham nor 
Briquet considered them synonymous, as did O. Kuntze and 
Janchen and Neumayer. 

Two facts emerge from this discussion. In the first place, 
Calamintha and Clinopodium, as originally proposed, and as 
generally used to 1891, apparently had no species in common. 
'There was, and is today, controversy over the rank of the taxa, 
but not over their identity or composition. In the second place, 
the proposal by Janchen is to conserve a poorly circumscribed 
later name over an adequately circumscribed Linnaean name. 
Further, the name proposed for conservation is both a later 
homonym and synonym of earlier names, facts not mentioned 
by the propositor. 

The “International Code of Botanical Nomenclature" (1952), 
at Article 24, stated: “. . . These names [for conservation] 
are preferably such as have come into general use in the fifty 
years following their publication, or which have been used in 
monographs and important floristic works up to the year 1890 

. ." Up to the year 1890 both Calamintha, Moench (non 
Lamarck) and Clinopodium, Linnaeus were in general use for 
the respective taxa concerned. "There was no confusion between 
them. 

Finally, if Clinopodium, Linnaeus is declared a nomen rejicien- 
dum, those who desire to recognize the taxon which has borne 
that name as a genus will be faced with the necessity of pub- 
lishing a new name for it. 


For these reasons it is recommended that the proposal for the 


78 Rhodora (Vor. 57 


conservation of the name Calamintha, Lamarck, and the rejec- 
tion of the name Clinopodium, Linnaeus not receive favorable 


actlon.— BAILEY HORTORIUM, CORNELL UNIVERSITY, ITHACA. 


A REVISION OF THE NORTH AMERICAN 
GENUS SABATIA (GENTIANACEAE) 


ROBERT L. WILBUR 
(Continued from page 71) 


E. Subsection DopECANDRAE subsect. nov.? 


Subg. Plurimaria Raf., Med. Fl. 2: 76. 1830, in part, not Plurimaria 
Raf. (as genus), Fl. Tell. 3: 31. 1837. 

Pleienta Raf., Fl. Tell. 3: 30. 1837, in part, an illegitimate name since 
its type, designated by Rafinesque, is the same as that of Sabatia. 
Sect. Pleienta (Raf.) Blake, Ruopora 17: 56. 1915, an illegitimate 

name. 
Dodecandrae Small, Man. SE. Fl. 1049. 1933, a category of undesig- 
nated status. 


Rhizomatose perennials with at least the secondary and very often the 
primary branches alternate. Strongly pronounced tendency towards 
plurimerous flowers ranging from 5-12(-14)-parted. Flowers typically 
large and pedicels usually longer than 1 em. TYPE SPECIES: Sabatia 
dodecandra (L.) BSP. 


This subsection contains but four species which, except for 
one species, are restricted to the Coastal Plain of the United 
States. S. calycina occurs in addition on at least two islands 
of the West Indies. 

S. calycina, I feel certain, has very little in common with the 
species of subsection Campanulatae with which it has been for- 
merly associated. Its proper relationship is more aptly shown 
by grouping it with the rhizomatose, pluripetalous species of the 
dodecandra-alliance. The strongly pronounced tendency for 
this species to possess flower-parts more numerous than five, 
the large, often foliose calyx-lobes, the often conspicuous, 
elongate rhizome, the broad leaves and the broadly campanulate 
calyx-tube are features of all or most of the dodecandra-group 

5 Subsectio Dodecandrae, Perennes rhizomatibus praeditae. Rami secondarii et 
interdum primarii alterni. Flores saepe plurimeres, inter 5-partiti et 14-partiti, 
plerumque in specie una 5-6-partiti, in speciebus aliis ca. 9- partiti, typice grandes, 


pedicellis quam 1 cm. longioribus. Species typica, Sabatia dodecandra (L. BSP. 
(Chironia dodecandra L.) 


1955] ^ Wilbur,—The North American Genus Sabatia 79 


and are present in none of the species with which it has formerly 
been allied. I do not feel that the Dodecandrae are so much 
more strongly differentiated than the other subsections as to 
warrant sectional rank which Blake accorded to them. 


KEY TO THE SPECIES OF SUBSECTION DODECANDRAE 


A Calyx- and corolla-lobes 5-6-parted or rarely up to 7; corolla-lobes (0.6—) 
0.7-1.3(-1.5) em. long; cauline leaves spatulate, tapering into a much- 
narrowed, sometimes almost petiolate base............ 12. S. calycina. 

A. Calyx- and corolla-lobes 7-13-parted, usually 9-12 in number, corolla- 
lobes (1.2—)1.6-2.4(-3.5) em. long; cauline leaves elliptic, lanceolate, 
or linear, the median and upper at least not strongly tapering into an 
almost petiolate base. 

B. Upper cauline leaves less than the diameter of the stem or but little 
wider, very narrowly linear, contrasting strongly with the broadly 
spatulate, rosulate, basal leaves; leaves thick and succulent, often 
drying somewhat rugose; the larger roots succulent; calyx-lobes 
strongly subulate, often succulent and even semicircular in cross- 
Secuont Mc vu PME ree ee mere re 15. S. bartramit. 

B. Upper cauline leaves considerably wider than the diameter of the stem, 
lanceolate, elliptic, or linear; basal leaves often absent, when present 
linear to lanceolate, and not strongly contrasting with the lower cau- 
line leaves; leaves drying smooth and usually thin in texture; roots 
fibrous, non-succulent; calyx-lobes linear, thin and flattened in 
cross-section. 

C. Calyx-lobes hyaline-margined; plants strongly stoloniferous; primary 
branches very often opposite; terminal flower short-pedicellate, 
usually considerably exceeded by the first internode of the lateral 
branches arising at the same node............ 14. S. kennedyana. 

C. Calyx-lobes not hyaline-margined; stolons usually lacking, if present 
neither numerous nor strongly developed; primary branches usually 
alternate; terminal flower typically long-pedicellate, usually about 
equaling or exceeding the first internode of lateral branch arising 
at the same node.............. cece eee eee eee 13. S. dodecandra. 


12. Sabatia calycina (Lam.) A. Heller, Bull. Torr. Bot. Club 21: 24. 
1894. Gentiana calycina Lam., Encyc. 2: 638. 1788. Chironia dicho- 
toma Walt., Fl. Car. 93. 1788. C. calycosa Michx., Fl. Bor.-Am. 1: 147. 
1803. Sabbatia calycosa (Michx.) Pursh ex Sims, Curtis’s Bot. Mag. pl. 
1600. 1813. S. gracilis var. cubensis Griseb., Mem. Am. Acad. 11: 521. 
1862. S. dichotoma (Walt.) Trelease ex Branner & Coville, Ann. Rept. 
Geol. Surv. Ark. 1888. 4: 204. 1891. Sabbatia cubensis (Griseb.) Urb. 
Symb. Ant. 8: 536. 1921. 

Perennial herb (although flowering the first year) (8-)15-40(-50) cm. 
high, with a slender to thick rhizome (1-)2-4(-10) em. long, 1-3 mm. in 
diameter. Stems usually solitary, or sometimes several and clustered, 
erect, rigid, smooth above or but very slightly ridged, hollow at least 
below, (1-)2-3(-4.5) mm. in diameter, branching usually restricted to 
the upper half or third of the stem but may arise from nearly throughout 
the entire length. Branches generally alternate but commonly opposite 


80 Rhodora [Vor. 57 


even above along the main stem, ascendent or more typically divaricate, 
further branching also strongly divergent, thus the plant typically pre- 
sents a strikingly geniculate aspect, branching usually restrieted to the 
second or third order, usually less than 20 em. long, bearing but few nodes. 
Root-system of several to numerous slender, fibrous roots arising from the 
rhizome or from the base of the stem. Leaves thin, drying very thinly 
membranous and hence venation conspicuous although only the midvein 
even slightly elevated beneath. Basal rosette lacking and the lower 
cauline leaves not conspicuously differentiated in shape from the median 
or upper leaves and usually the reduction in size is gradual and slight. 
Leaves ascending or more often strongly spreading, elliptical to broadly 
spatulate, mostly obtuse but still commonly acute, tapering into a con- 
spieuous, much-narrowed or even petiolate base, (1.5-)2.5-6(-10) em. 
long, (4-)10-18(-20) mm. wide. Inflorescence of usually reduced, 1-2- 
flowered cymules; the flowers appearing loosely arranged and even 
solitary. Pedicels slender, rigid, inconspicuously 5-7-angled, (1-)3-5(-6) 
em. long. Calyx-tube thin, smooth or with veins but very slightly 
elevated, somewhat scarious or translucent, shallowly crateriform to 
broadly campanulate, (1.5-)2-4(-5) mm. long. Calyx-lobes oblanceolate 
to spatulate, or rarely linear, often of unequal size, usually foliaceous, 
apparently enlarging after pollination, acute to obtuse, thin, membranous, 
(8-)10-25(-32) mm. long, (1-)1.5-6(-8) mm. wide. Corolla-tube 
cylindrical, (3-)4-5(-6) mm. long, 2-3 times as long as the calyx-tube, 
colorless or white to pale pink. Corolla-lobes equaling the calyx-lobes 
in number, typically 5-6-parted, but occasionally as many as 7, often 
exceeded in length by the calyx-lobes which are sometimes as much as 
twice as long, but not uncommonly equaling or even exceeding the calyx- 
lobes by 1-2 mm., oblong to oblong-spatulate, oblanceolate or elliptic, 
obtuse or acute, (6—)7-13(-15) mm. long, (2-)3-5(-6) mm. wide, white or 
more commonly pale rose to pink gradually giving way to white in the 
area above the triangular, pointed yellow patch at the base of the lobe. 
Filaments pale yellow, 2-3 mm. long; anthers bright yellow, slender 
2.5-3.5 mm. long. Style 1-2 mm. long; stigmatie branches 4-6 mm. long. 
Capsule almost globose or very broadly cylindrical, 6-10 mm. high, 5-8 
mm. in diameter. 

Type locality: Cette plante croit á la Louisiane.” Type: the original 
description states that its author saw it in the herbarium of Jussieu. A 
photograph in the Gray Herbarium, reputedly of the type, was of a speci- 
men in Lamarck’s herbarium. 

DISTRIBUTION. Ditches, riverbanks and swampy hardwoods from 
southeastern Virginia south along the Coastal Plain throughout most of 
Florida and westward into eastern Texas. Also known from eastern 
Cuba and the central Cordillera of Hispaniola, Map 12. 


In the West Indies this species has been called most often 
S. gracilis var. cubensis or S. cubensis. However in spite of the 
geographical isolation and the difference in physiographic 


1955] Wilbur,—The North American Genus Sabatia 81 


provinces, there has not been pointed out previously, nor have 
I been able to detect from herbarium specimens, any character 
that distinguishes the West Indian specimens from those from 
the south Atlantic and Gulf Coastal Plain. The original pub- 
lication of the insular representatives in a varietal status with 
the very different S. gracilis, which is but a later synonym of 
S. campanulata, by Grisebach, is especially surprising in view 
of that author's monographic treatment of the family. Grise- 
bach's varietal name was first treated as à synonym by Gray 
in the Synoptical Flora and this still seems to be the only de- 
fensible stand to take in view of the lack of any known mor- 
phological differences. Urban did not discuss the evidence, 
if any, that caused him to decide upon specific status. 

Lamarck's publication doubtless has priority over that of 
Walter as the portion of the former's work in which Gentiana 
calycina appears is usually dated as April, 1788. Fraser, who 
carried Walter's Flora back with him and arranged for its 
publication at his own expense, stated (1789, p. 5) that “after 
having resided in South Carolina and Georgia nineteen months, 
I returned to England in the month of March 1788. . . ." 
This would seem to make certain that it was at least several 
months after the appearance of Lamarck's name before Walter's 
Flora could have been published. 


REPRESENTATIVE SPECIMENS:—VIRGINIA: Isle of Wight Co., Zuni, Fernald & 
Long 6349 (GH, PENN); Nansemond Co., near Suffolk, Kearney 1726 (us); 
Southampton Co., about Franklin, Heller 1114 (GH, MO, NY, PENN, US); Sussex 
Co., west of Lumberton, Fernald & Long 14386 (GH, MO, TENN). NORTH 
CAROLINA: Brunswick Co., just w. of Wilmington, Wilbur 2894 (micH); 
Chowan Co., 1.5 mi. e. of Edenton, Randolph 659 (cu, au); New Hanover Co., 
near Wilmington, Biltmore Herb. 3806c (Ny, vs); Onslow Co., 4.5 mi. s. of 
Jacksonville, For & Boyce 3718 (mica, ncs); Pender Co., n. of Castle Hayne, 
Foz & Boyce 3773 (mica, Ncs); Pitt Co., w. of Grimesland, Wiegand & Manning 
2560 (CU, aH). SOUTH CAROLINA: Berkeley Co., 10 mi. ne. of Moncks Corner, 
Godfrey & Tryon 872 (an, Ny, vs); Dorchester Co., along the Ashley River, 
Correll 5878 (DUKE, NA); Georgetown Co., 1.5 mi. w. of Andrews, Godfrey & 
Tryon 555 (DUKE, F, MO, NY, TENN, US); Horry Co., Myrtle Beach, Coker, 29 
July 1946 (xcv, Ny); Williamsburg Co., s. of Kingstree, W?egand & Manning 
2562 (cu, aH). GEORGIA: Baker Co., in western portion of county, Eyles 
7185 (DUKE, GH); Dougherty Co., vicinity of Albany, Pollard & Mazon 532 
(ny, Us); Early Co., about 4 mi. se. of Blakely, Harper 1910 (F, GH, MO, NY, 
US). FLORIDA: Alachua Co., River Sink, Murrill, 21 May 1939 (wo, vs); 
Columbia Co., without exact locality, Hitchcock, 1898 (r, Mo); Duval Co., 
near Jacksonville, Curtiss 4373 (F, Mo, Ny, vs); Lake Co., Eustis, Nash 2063 
(GH, MICH, MO, MT, NY, US); Levy Co., Rosewood, Garber, June 1886 (BRU, F, 


82 Rhodora [Vor. 57 


NY, Us, YU); Orange Co., 7 mi. se. of Fort Christmas, O'Neill 25 June 1925 
(FLAS, US); St. Johns Co., near Tocoi, Curtiss 2230 (CU, F, FLAS, GH, MO, NY, 
SMU, US, YU). ALABAMA: Mobile Co., Magazine Point, Dougan, 19 June 1914 
(MO). LOUISIANA: Calcasieu Parish, vicinity of Lake Charles, Allison 324 
(ny, Us); East Baton Rouge Parish, near Baton Rouge, Joor, 22 May 1874 
(F); Orleans Parish, New Orleans, Drummond 222 (Gu, K). TEXAS: Harris 
Co., Houston, Hall 510 (BRU, F, GH, MO, NY, US). CUBA:—Oriente Province: 
Monte Verde, Shafer 8719 (Ny, US); in Cuba Orientali, Wright 412, 1856-57 
(TYPE-NUMBER of jS. gracilis var. cubensis) (F, GH, MO, NY). DOMINICAN 
REPUBLIC: Cordillera Central, Prov. de la Vega, Constanza, in Cienaga de los 
Hoyos, c. 1200 m., Ekman 13919 (GH, MO, NY, US); prope Farabocoa, c. 
550 m., Fuertes, June 1912 (an); prope Constanza, T'ürckheim 3368 (au, Mo, 
NY, Us); Cordillera Central, Prov. de Azua, Valle del Yanque, Ekman 13700 
(NY). 


13. Sabatia dodecandra (L.) BSP., Prel. Cat. N. Y. 36. 1888. 

Perennial herb (10—)30-70(-100) em. high, with a slender to robust, 
often somewhat branched rhizome 4-10(-15) em. long, 2-4 mm. in diam- 
eter. Stems solitary or several arising in a cluster, erect, terete and 
smooth or somewhat angular and slightly ridged, hollow, (1—)2-3(-4) 
mm. in diameter. Branches usually restricted to the upper third or 
half of the stem, typically alternate, or more rarely, on unusually robust 
plants, the primary sometimes opposite; ascendent to strongly spreading, 
forming an angle of from 30-70 degrees with the stem, 5-15(-20) em. 
long, rigid. Root-system of slender, definitely non-fleshy, wiry roots 
5-10 em. long and generally considerably less than 1 mm. in diameter. 
Commonly, especially in the south, with one to few, slender, superficial 
rhizomes, or more rarely stolons, 2-10 em. long, 1-2 mm. in diameter 
bearing small rosettes of few to numerous, thin, slender, oblanceolate, 
obtuse to acute leaves 1.5—4 cm. long. Basal rosettes absent from the 
base of the aerial stem, the lower cauline leaves neither densely clustered 
nor strongly contrasted in either size or shape with those borne several 
nodes higher up on the stem. Cauline leaves spreading to ascendent, 
non-succulent, in texture drying from thin and membranous to even 
somewhat thick and chartaceous, smooth, not rugose, with 1-2 pairs 
of usually inconspicuous veins paralleling the midvein which is typically 
elevated beneath; leaves only gradually reduced above, in length ranging 
from twice exceeded by the internodes to 3-4 (or more) times surpassed 
by them, lanceolate, linear, elliptic, or oblong, acute to obtuse, clasping 
to merely sessile, (1.5-)2.5-4(-7) em. long, (4-)5-12(-20) mm. wide; 
the lowermost sometimes narrowly spatulate with long tapering almost 
petiolate base. Inflorescence almost invariably of l-several, reduced, 
1-2-flowered cymules but rarely complete with three flowers; the flowers 
loosely arranged and thus appearing solitary on erect, rigid, slightly 
angled pedicels (1-)3-6(-11) em. long and about 1 mm. in diameter. 
Calyx-tube crateriform, somewhat turbinate or campanulate, with 
nerves typically somewhat elevated, (1.5-)2-3(-4) mm. long, usually 
green but sometimes straw-colored. Calyx-lobes thin, flat, drying 
smooth, linear, narrowly oblanceolate or even spatulate, ranging from 


1955] Wilbur,— The North American Genus Sabatia 83 


rather inconspicuous to large and somewhat foliaceous, acute, (0.4—) 
0.8-1.8(-2), em. long, one-fifth as long to slightly exceeding the corolla 
in length, 1-2.5 mm. wide. Corolla-tube (4-)5-7(-8) mm. long, usually 
2-4 times the length of the calyx-tube. Corolla-lobes (7-)9-12(-13) in 
number, elliptic, oblanceolate, or oblong, acute to obtuse, (1.2-)1.6-2.4(—3) 
em. long, (3-)5-8(-10) mm. wide, usually not overlapping each other 
when fully expanded, deep rose-purple, rose-pink, pink, or more rarely 
white, with oblong to somewhat triangular yellow patch at base of lobe, 
the patch sharply 3-lobed or even irregularly toothed, usually bordered 
by thin red line except in albinos. Anthers bright yellow, slender, 3-5 
mm. long; filaments yellow, slender, 3-5 mm. long. Stigmatic lobes 
slender, 5-9 mm. long; style 3-5 mm. long. Capsule cylindrical, 6-10 
mm. high, 4-6 mm. in diameter. 


Some indication of the considerable variation within this 
species may be deduced from the fact that in the past fifty years 
three species have been segregated from it. The variation 
is such, that with but a few sheets upon which to base their 
conclusions, it is not surprising that the various authors have 
been convinced of the distinctness of the entities that they 
were publishing. The several hundred specimens of this species 
examined by me from nearly one hundred stations extending 
from Connecticut to Louisiana are quite diverse and at the same 
time it is all but impossible to define any of the various tendencies 
that have attracted the interest of botanists in the past. 

After the belated recognition of the specific status of the entity 
formerly called S. decandra in 1900, the next species to be segre- 
gated from the all-inclusive S. dodecandra (= S. chloroides) 
was S. foliosa Fern. in 1902. What eventually became a syntype 
of this species was sent to the Gray Herbarium by the collector, 
A. H. Curtiss, who afterwards distributed the duplicates as 
S. chloroides according to the determination of Fernald. The 
latter became more impressed with the features of the southern 
plant after observing similar variation in specimens collected 
by J. Donnell Smith in South Carolina. He described it as 
a species especially emphasizing the (1) more stoloniferous 
habit, (2) leaves equaling the internodes, (3) foliaceous calyx- 
lobes. The calyx-lobes in both the numerous duplicates col- 
lected by Curtiss and those of Smith are certainly large, ranging 
in size from one-half or more commonly three-quarters as long 
as the corolla-lobes or in a few instances even exceeding them. 
This character has proven most variable and the variation is 


84 Rhodora [Vor. 57 


so great within the same colony and even on the same plant 
that it is of but little diagnostic importance. For example, 
the variation of large series of specimens collected (Webster & 
Wilbur 3583) along the eastern bank of the Blackwater River 
just aeross from Milton, Florida, the locality from which one 
of the syntypes was collected, ranges rather uniformly between 
having calyx-lobes one-fifth to about three-quarters as long as 
the corolla-lobes. 

Small (1903) proposed another species based largely upon 
two collections from central Georgia which he called S. harperi, 
in honor of the original collector. The supposed distinguishing 
characters were calyx-lobes less than half the length of the corolla 
and supposed features of the leaf which are in reality highly 
variable. Small restricted the range of S. foliosa to Florida 
and Alabama which excluded one of the syntypes. S. harperi 
was said to range from South Carolina to Florida and west 
to Louisiana. In the previous month Harper, in his account 
(Bull. Torrey Club 3?: 338-339. 1903) of these same collections, 
referred to them as S. foliosa upon the authority of Fernald who 
had examined them. Harper pointed out at that time that 
“the two species (i.e., S. foliosa and S. dodecandra) are very 
closely related differing perhaps in habitat as much as in any 
other way" in that “S. dodecandra seems to be mostly a maritime 
plant, while the usual home of 5. foliosa is in creek and river 
swamps.” Fernald had informed Harper also that S. dodecandra 
was not represented in the Gray Herbarium from south of North 
Carolina and this restricted range was presented in the Seventh 
Edition of Gray's Manual. 


Blake (1915) concluded that S. foliosa and S. harperi were 
conspecific. At the same time he proposed a new species, 
S. obtusata, from central Georgia. The distinctive features 
were said to be oblong leaves with rounded apices and non- 
clasping bases which nearly equaled the internodes. He 
emphasized that S. foliosa (including S. harper?) had ‘‘consider- 
ably longer calyx-lobes” and the sketch accompanying the 
description of S. obtusata showing part of the type indicates 
that the calyx-lobes extend barely one-third the length of the 
corolla. None of the supposed differences pointed out by 
Blake is distinetive, as they are but minor variations which are 


1955] Wilbur,—The North American Genus Sabatia 85 


well within the range of variability of the southern population. 
Several of these features appear within the individual sheets 
taken from one stand. The diagnostic importance of clasping 
versus sessile leaves has been overemphasized in the past. 
Robustly developed plants throughout the range of the species 
may possess clasping or semi-clasping leaves. 


In the last edition of Gray’s Manual (1950), the range and 
habitat of S. dodecandra was stated as ‘‘saline, brackish or 
rarely fresh marshes and meadows, Fla. to La., n. on or near 
Coastal Plain to s. Ct." However, of all the many sheets that 
Fernald annotated, mostly in 1916, there is only one from south 
of North Carolina that he considered at that earlier date to be 
S. dodecandra and that, a very fragmentary specimen of Buckley 
from Alabama, seems quite inadequate for satisfactory determin- 
ation. In spite of a return to the extensive range for S. dode- 
candra by Fernald, it is not certainly to be inferred that he had 
necessarily abandoned any attempt to distinguish the entities 
since fresh-water habitats are implied by Fernald to be the 
exception for the species which is anything but the case in the 
South. 


I am convinced that it is impossible to recognize S. dode- 
candra, S. foliosa, S. harperi, and S. obtusata as four distinct 
entities regardlesss of rank. In general, the material examined 
from stations ranging from Connecticut to just north of Charles- 
ton, South Carolina, is characterized by having the internodes 
from half-again to twice or more as long as the leaves and this 
variety seems for the most part to be rather closely restricted 
to brackish habitats or at least seldom gets very far from the 
coast. The variety that occurs from South Carolina to Florida 
and westward to Louisiana is characterized by having shorter 
internodes which rarely exceed one and a half times the length 
of the leaves and more often than not are equaled or exceeded 
by them. Its habitat is inland along river banks and borders 
of ditches and streams. Most of the specimens examined have 
been easily separable by this morphological criterion and by 
several additional tendencies that are little more tangible than 
the “certain indescribable grace" of E. L. Greene. Rather 
unsatisfactory specimens were seen from St. Vincent Island and 
Tampa Bay, Florida, which have been tentatively determined 


86 Rhodora (Vor. 57 


as S. dodecandra, the only specimens so identified by me from 
south of South Carolina. From the southeastern corner of 
this last-mentioned state three collections are available and they, 
too, leave considerable to be desired in the matter of providing 
an ample basis for positive determination. There are few 
complete or even modern collections known from Alabama, 
none at all from Mississippi, and but very few from Louisiana. 

In spite of the obvious need for intensive fieldwork and an 
accumulation of numerous well-prepared collections from 
throughout the entire range of the species before this group 
will be satisfactorily understood, it is believed that the treatment 
suggested here will serve as à more natural and at the same 
time more useful arrangement until this necessary research 
is accomplished and may prove a better point of departure 
than previously has been available. 


KEY TO THE VARIETIES OF S. PODECANDRA 

Internodes usually exceeding the leaf in length, commonly 1.5 to 3 times 
longer than the leaf; stolons rarely present but if present then only weakly 
developed; plants usually of brackish habitats, coastal from Connecticut to 

South Carolina (and perhaps locally to Florida) 
13a. S. dodecandra var. dodecandra. 
Internodes usually equaled in length by the leaf or nearly so, commonly less 
than the leaf in length and very rarely exceeded by it as much as 1.5 times; 
stolons commonly present and often numerous and well developed; plants 
of river-banks, ditch and stream margins, inland from South Carolina to 
Louisiana... 6... eee 13b. S. dodecandra var. foliosa. 


13a. Sabatia dodecandra (L.) BSP., var. dodecandra 


Chironia dodecandra L., Sp. Pl. 190. 1753. Chlora dodecandra (L.) 
L., Syst. Nat. ed. 12. 2: 207. 1767. Chironia chloroides Michx., Fl. 
Bor. Am. 1: 147. 1803. Illegitimate as it was merely a substitute 
name. Sabbatia chloroides (Michx.) Pursh, Fl. Am. Sept. 1: 138. 1814. 
S. dodecandra (L.) BSP., Prel. Cat. N. Y. 36. 1888.  Pleienta leucantha 
Raf., New Fl. 4: 92. 1838. P. dodecandra (L.) Raf. ex B. D. Jackson, 
Ind. Kew. 2: 561. 1894. 

Rarely stoloniferous and stolons when present scarcely developed; 
internodes usually exceeding the leaf in length, commonly 1.5-3 times 
longer. TYPE LOCALITY: “in Virginia." Type: Clayton 120 (British 
Museum); Phototype seen in collection of Gray Herbarium. DISTRIBU- 
TION: Salt or brackish marshes from Connecticut south along the coast 
into South Carolina and perhaps locally to Florida. Map 13. 


REPRESENTATIVE SPECIMENS:—CONNECTICUT: Middlesex Co., Saybrook, 
Thompson, 20 Aug. 1891 (wv). NEw yorK: Rockland Co., Iona Island, 
M uenscher & Curtis 5939 (cv); county unknown, Long Island, White Mills, 


1955] Wilbur,— The North American Genus Sabatia 87 


Poggenburg, Aug. 1886 (GH). NEW JERSEY: Atlantic Co., Port Republic, 
Long 10486 (au); Bergen Co., Hackensack marshes, Eaton, Sept. 1860 (vv); 
Burlington Co., New Gretna, Chrysler, 30 Aug. 1926 (cu, RUT); Cape May Co., 
first of Fishing Creek Bogs, Walker 1653 (NA, us); Hudson Co., Granton, 
Sickle, 1 Aug. 1894 (us); Monmouth Co., Avon-by-the-Sea, Kaufman, 25 
Aug. 1904 (yu); Ocean Co., east of Silverton, Long 38148 (PENN). DELAWARE: 
Sussex Co., s. of Rehoboth Beach, Hood 2378 (FLAS). MARYLAND: Anne 
Arundel Co., Furnace Branch, Pitt, 5 Aug. 1902 (au); Caroline Co., between 
Choptank River and Bethlehem, Killip 7279 (vs); Cecil Co., ca. 1.5 mi. s. of 
Elkton, Long 57046 (an); Charles Co., Stump Neck, Turpin 474 (us); Harford 
Co., 0.25 mi. n. of Bush River Station, Shull 344 (an, Mo, Ny, Us); St. Marys 
Co., St. Marys River 3.7 mi. from St. Mary, Walker 3888 (us); Somerset Co., 
Kings Creek, Holmes, 24 July 1890 (us); Talbot Co., 5.5 mi. se. by s. of 
Easton, Earle 3075 (PENN, WVA); Wicomico Co., Salisbury, Smith 302 (wr, 
NY); Worchester Co., Stockton, Rusby, Aug. 1889 (Ny). vrRGINIA: Fairfax 
Co., near mouth of Occoquan Creek, Ulke, 1 Aug. 1910 (us); James City Co., 
about 5 mi. from Toano, Menzel 193 (an); Nansemond Co., s. of Reid’s Ferry, 
Fernald & Long 13423 (au, Us); Norfolk Co., near Northwest, Fernald & Long 
13997 (GA, GH, MO); Princess Anne Co., Cape Henry, Egler & Ryan 40-183 
(NY). NORTH CAROLINA: Carteret Co., between Core Creek and Adam's 
Creek, Hill 171 (puxe); Chowan Co., near Edenton, Godfrey 5346 (Gu, vs); 
Craven Co., 2 mi. sw. of James City, Randolph 862 (cv, an); Currituck Co., 
marsh at Sligo, Godfrey 5279 (DUKE, GH, us); Dare Co., Kill Devil Hills, 
Fox 144 (mica, Ncs); New Hanover Co., Carolina Beach, Biltmore Herb. 3305c 
(GH, MICH, MO, NCU, NY, PENN, US); Pasquotank Co., Elizabeth City, Kearney 
1996 (us); Tyrrell Co., Fort Landing, Radford 4622 (NCU). SOUTH CAROLINA: 
Charleston Co., north of McClellanville, Godfrey & Tryon 722 (DUKE, F, GH, 
MICH, MO, NY, PENN, TENN, US). FLORIDA: Franklin Co., St. Vincent Island, 
McAtee 1835A (vs); Hillsborough Co., Tampa Bay, Leavenworth (an, NY). 
'The material upon which these Florida stations are reported is rather unsatis- 
factory for certain identification but they seem to belong to the otherwise 
more northern population. 


13b. Sabatia dodecandra (L.) BSP., var. foliosa (Fern.) comb. nov. 


Sabbatia foliosa Fern., Bot. Gaz. 33: 155. 1902. S. Harperi Small, 
Fl. SE. U. S. 928. 1903.  Sabatia obtusata Blake, Ruopora 17: 54 pl. 
112. 1915. 

Commonly stoloniferous and the stolons often numerous and well- 
developed; internodes usually equaled or exceeded by the leaves, com- 
monly the leaves as much as 1.5 times as long as the internodes. TYPE 
LOCALITY: Muddy banks of the Blackwater River, near Milton, Florida." 
Syntype and lectotype: Curtiss 5928 (GH). DISTRIBUTION: River banks, 
ditches and stream and pond margins inland from South Carolina south- 
ward into northern Florida and westward into Louisiana. Map 13. 


REPRESENTATIVE SPECIMENS: SOUTH CAROLINA: Beaufort Co., Bluffton, 
Mellichamp, 1887 (us); Berkeley Co., 10 mi. ne. of Moncks Corner, Godfrey & 
Tryon 876 (DUKE, F, GH, MO, NY, PENN, TENN, US); Charleston Co., 17 mi. w. 
of Charleston, Duncan 5802 (a4); Colleton Co., Cottageville, Hunt 1648 
(CHARL, MICH,); Jasper Co., Ridgeland, Mohr, 14 Nov. 1893 (vs); Marion Co., 
east of Nichols, Wiegand & Manning 2561 (cv, an); Orangeburg Co., North 


88 Rhodora [Vou. 57 


Edisto River, Smith, 9 Aug. 1884 (r, GH, Mo, vs); Williamsburg Co., 6 mi. s. 
of Kingstree, Godfrey & Tryon 369 (DUKE, F, GH, MICH, MO, NY, PENN, TENN, 
Us). GEORGIA: Bullock Co., swamp of Big Lott’s Creek, Harper 964 [SYNTYPE 
of S. harperi] (GH, Mo, Ny, US); Candler Co., 6 mi. ne. of Metter, Pyron & 
McVaugh 718 (aa, us); Charlton Co., Traders Hill, Wright 878 (cv); Decatur 
Co., between Forest Falls and Bainbridge, Harper 1196 [SYNTYPE of S. harpert] 
(GH, MO, NY, vs); Dodge Co., between Copeland and Rhine, Harper 1876 
(F, GH, MO, NY, US); Early Co., about 2 mi. e. of Restler, Harper 1912 (F, Gu, 
MO, NY, US); Grady Co., about 4 mi. w. of Whigham, Wise, 30 June 1940 
(FLAS); Jenkins Co., about 3 mi. w. of Millen, Pyron & McVaugh 968 (Ga, Us); 
Laurens Co., 12 mi. s. of Wrightsville and 6 w. of Adrian, Pyron & McVaugh 
3056 (GA, NA); Long Co., 3 mi. sw. of Ludowici, Wilbur & Webster 2756 
(mica); McIntosh Co., Darien, Wiegand & Manning 2564 (cv); Mitchell Co., 
12 mi. ne. of Camilla, Thorne 5784 (cu); Montgomery Co., north of Mt. 
Vernon, Harper 1866 (r, GH, MO, Ny, vs); Telfair Co., near Lumber City, 
Biltmore Herb. 3305d (vype-number of S. obtusata] (Us). FLORIDA: Gadsen 
Co., sw. of Havana, Small, Small & DeWinkeler 11387 (Ny, us); Jefferson Co., 
2 mi. nw. of Lamont, Webster & Wilbur 3631 (M1cn); Madison Co., 1 mi. w. of 
Greenville, Wiegand & Manning 2565 (cu, aH); Nassau Co., Boulogne, 
Hume & West, 14 July 1933 (FLAs); Santa Rosa Co., near Milton, Curtiss 
5928 (cU, FLAS, GH-SYNTYPE of S. foliosa, KSC, MO, NCU, NY, SMU, US); Taylor 
Co., 14 mi. nw. of Perry, Webster & Wilbur 3635 (micH). ALABAMA: Mobile Co., 
Mobile, Harvey (us); Dallas Co., Selma, Biltmore Herb. 3305a (us); Washington 
Co., 2 mi. s. of Wagarville, Webster & Wilbur 3496 (MICH). LOUISIANA: 
Calcasieu Parish, Lake Charles, Tracy 3453 (F); St. Landry Parish, Opelousas, 
Carpenter (an); St. Tammany Parish, vicinity of Covington, Arséne 11706 (Us). 


14. Sabatia kennedyana Fern. Ruopora 18: 150. fig. 98e. 1916. 
S. Kennedyana, f. candida Fern., Ruopora 18: 151. 1916. S. Ken- 
nedyana, f. encycla Fern., Rnopona 24: 180. 1922. 


Perennial herb (15-)30-65(-80) em. high, arising from slender, easily 
broken rhizome about 2-3 mm. in diameter and up to 12 em. long. Stems 
usually solitary or occasionally 2 or more arising together, erect, stiffly 
rigid, terete, smooth, ridgeless throughout, hollow below, 2-4(-6) mm. 
in diameter. Branches typically restricted to the upper third or half 
of the stem, the primary, at least in well-developed plants, typically 
opposite, additional ramifications, when present, mostly alternate; 
usually rather strongly ascendent forming an angle of about 20—40(—50) 
degrees with the stem, generally 5-15(-30) em. long, slender, rigid, 
further branching also typically stiffly geniculate. Root-system a dense 
mass of non-fleshy, very slender, white to straw-colored roots 5-10 cm. 
long, 1 mm. or less in diameter. Typically with few to several or oc- 
casionally with numerous slender, decumbent stolons initiated as super- 
ficial rhizomes 2-8(-12) em. long, 1-2 mm. in diameter, bearing few to 
several slender roots and at the tip a rosette of numerous, ascendent to 
erect, usually very narrowly linear or oblanceolate leaves 10-15 times 
as long as broad, 2-6(-15) em. long, 3-8(-15) mm. wide, acute to acumi- 
nate, strongly long-tapering to the nearly petiolate base. Leaves thin, 
non-succulent, drying brittle, smooth, thinly chartaceous in texture, 
venation obscure except for the midvein which is somewhat elevated 


1955] Wilbur,— The North American Genus Sabatia 89 


beneath; the basal leaves, when present, and the lower cauline very 
similar in size and shape to those of the stolon-borne rosettes; the cauline 
strongly ascendent and only gradually reduced above, stems almost 
equally leafy throughout, typically lanceolate or narrowly linear, weakly 
clasping or sessile, acute to acuminate, slightly callose-tipped, (1.5—) 
3-5(-10) em. long, (2-)4-10(-16) mm. wide, usually about (5-)7-12(-15) 
times as long as wide, ranging from about (0.5-)2-4(—5) times as long 
as the internodes. Inflorescence composed of complete or reduced 
cymules; the terminal flower typically shortly pedicellate and usually 
greatly exceeded by the first node of the lateral branches. Flowers 
borne on erect, slender, smooth pedicels (0.5—-)1—5(-7) em. long and about 
1 mm. in diameter. Calyx-tube crateriform or shallowly campanulate, 
thin, smooth or rarely with a few nerves slightly elevated, (2-)3—4(-4.5) 
mm. long, green to almost straw-colored. Calyx-lobes linear, (5—-)6—10(—18) 
mm. long, 0.5-1.5 mm. wide, thin, very flat in cross-section, acute, slightly 
hyaline-margined, strongly ascendent, green. Corolla-tube cylindrical, 
5-8 mm. long, 2-3 times as long or about 2-4 mm. longer than the calyx- 
tube, pale yellow without, darker within. Corolla-lobes (7-)9-12 in 
number, (12-)18-24(-26) mm. long, (4-)7-10(-13) mm. wide, obovate- 
spatulate, cuneate-obovate, or more nearly oblong, tapering gradually 
to broad 3-5 mm. base, broadly obtuse or rarely somewhat emarginate, 
usually overlapping, rose-pink, pink, or rarely white, with an often 3- 
pointed oblong yellow patch usually about 4—7 mm. long, except in white 
flowers this area bordered by a dull red to reddish-brown line. Filaments 
slender, pale yellow, (2-)4-6(-7) mm. long; anthers linear, slender, 
bright yellow, 3-6 mm. long; style 2-6 mm. long; stigmatic lobes slender, 
(4-)5-7(-9) mm. long. Ovary rather conspicuously half-exserted from 
the corolla-tube. Capsule broadly cylindrical, 7-11 mm. long, 4-7 mm. 
in diameter. TYPE LOCALITY: “shore of Wequawket Pond, Centreville, 
Barnstable Co., Massachusetts." type: E. F. Williams s.n. (am!). 
DISTRIBUTION: Sandy and peaty margins of ponds and streams in southern 
Nova Scotia, eastern Massachusetts, Rhode Island, southeastern North 
Carolina and northeastern South Carolina. Map 14. 


S. kennedyana, recognized only in 1916 as being distinct from 
the widespread S. dodecandra, is a rather easily identifiable 
entity with as distinctive morphological features as any of the 
four species within the subsection. It is distinguishable by 
a combination of several strong tendencies rather than by one 
or even a few strong and constant features. These characters 
are: stems almost perfectly smooth and in well-developed 
specimens the primary branches typically opposite; the terminal 
flowers, usually short-pedicellate, typically being much exceeded 
by the first internode of the lateral branches; leaves thin, drying 
thinly chartaceous, brittle, smooth; corolla-lobes broadest near 
the tip, more or less obovate-spatulate, calyx-lobes linear and 


90 Rhodora [Vor. 57 


hyaline-margined, thin and flat in cross section; calyx-tube 
broad, thin, unribbed. Some of these characteristics are shared 
by other species but the combination is unique. In addition, 
the frequency of the stolon-borne rosettes and their development 
in size is much greater in this species than in any of the others 
within the subsection. "Therefore, in spite of the late date 
of recognition of the distinctness of this species, it appears to 
be a strongly characterized biological entity and not merely the 
result of too diligent a study of too few specimens. 


When originally deseribed, the plant was known only from 
Rhode Island and Massachusetts. Four years later Fernald 
discovered it in Nova Scotia where it is apparently fairly com- 
mon in Yarmouth County on the southern side of that peninsula. 
An even greater range extension for this species into south- 
eastern. North Carolina and northeastern South Carolina can 
now be reported. H. L. Blomquist, R. K. Godfrey and I made 
a large collection of what proves to be this species along the sandy 
bank of the Waccamaw River in Columbus County, North 
Carolina in the summer of 1951. After studying these speci- 
mens, it was then possible to recognize that a few other collec- 
tions from the same general area which previously had been 
rather uncritically passed over as merely abnormal S. dode- 
candra were in reality this species which I had not expected 
to see from south of the fresh-water ponds of southern New 
England. Disjuncts between the eastern Carolinas or south- 
eastern Virginia and Delaware, New Jersey or New England 
are numerous but examples matching that of S. kennedyana 
are certainly much less common. One species with a somewhat 
similar distribution as shown by existing records is Scirpus 
longi? Fern. This sedge is at present known from one station 
in eastern North Carolina, again from New Jersey to Mas- 
sachusetts, and finally reappears in western Nova Scotia. Fer- 
nald (Rhodora 45: 55. 1948.) wrote that this species of Scirpus 
“has shown itself to be an old Coastal Plain type which, like 
so many other species, became isolated in Nova Scotia before 
the late Tertiary or early Pleistocene submergence of the con- 
tinental shelf." 


The possibility that the name S. kennedyana may be synony- 
mous with Chzronia decandra Walter is included in the discussion 


1955] Wilbur,— The North American Genus Sabatia 91 


of S. bartramii and need not be repeated here. Walter’s name 
had best be treated, it is believed, as a nomen dubium. 


REPRESENTIVE SPECIMENS:—NOVA SCOTIA: Yarmouth Co., Gravelton, 
Fernald & Long 22266 (F, GH, MO, NY, PENN, YU, US). MASSACHUSETTS: 
Barnstable Co., Harwich, Fernald pL. EX. GRAYANAE 387 (in many herbaria) ; 
Bristol Co., Dartmouth, Sturtevant, 26 July 1889 (cu, F, Mo); Essex Co., 
Newburyport, Chamberlain, June 1899 (Ny); Norfolk Co., Weymouth, Seymour 
4373 (DUKE, WIS); Plymouth Co., in inundatis ad Plymouth, Oakes (Gu, Ny, 
US, YU). RHODE ISLAND: Kent Co., Warwick, Bailey, 27 July 1883 (us); 
Washington Co., South Kingston, Congdon (MO, NY, YU). NORTH CAROLINA: 
Columbus Co., between Ash and Old Dock just west of the Waccamaw River, 
Wilbur 2892 (micu); New Hanover Co., Wilmington, McCarthy, 1885 (Gu, NCU, 
US). SOUTH CAROLINA: Horry Co., near Ocean Drive, Schallert, 12 June 1933 
(NY). 

15. Sabatia bartramii sp. nov.’ Sabbatia chloroides var. coriacea 
Ell., Sk. Bot. S. C. & Ga. 1: 286. 1817. Type (cHanL!). S. chloroides 
var. flexuosa Ell., l.c., excluding synonym. Type (CHARL!) ? Sabbatia 
chloroides var. erecta Ell., l.c. There is no specimen representing this 
variety in Elliott’s herbarium. The original description is practically 
a translation of Walter’s description of Chironia decandra, which is cited 
in synonymy. ? Pleienta flexuosa [Ell.] Raf., Fl. Tell. 3: 30. 1837, 
without basionym. ? Sabbatia chloroides B stricta Griseb., Gen. et. Sp. 
Gent. 125. 1839. The description is again almost word-for-word the 
same as that of Chironia decandra Walt., which Grisebach cited in syn- 
onymy. Sabbatia dodecandra stricta (Griseb.) Mohr, Bull. Torrey Club 
24: 26. 1897. Sabbatia decandra sensu Harper, Bull. Torrey Bot. Club 
27:432. 1900. 

Perennial herb (25-)50-80(-100) em. high, arising from slender to 
thick rhizome 4-6 or more em. long, 3-5 mm. in diameter. Stems almost 
invariably solitary, erect and rather rigid, terete, more or less smooth 
and ridgeless or but finely nerved, hollow, 2-5 mm. in diameter. Branches 
usually restrieted to the upper third of the stem, typically alternate but 
occasionally opposite at one or two of the primary nodes; ascendent 
usually forming an angle with the stem of from 30-45 degrees, usually 
10-20(-40) em. long, typically unbranched and bearing but one flower, 
or if branched seldom with more than 2 flowers. Root-system of numerous 
clustered, fleshy roots 5-12 em. long, 1-2 mm. in diameter bearing but 
few, slender, fibrous lateral roots. Leaf-texture thick, succulent, drying 
rather thickly chartaceous or rarely the lowermost becoming thinly 
membranous upon drying, venation obscure excepting the midvein which 
is often elevated beneath, the apices often callous-mucronate. Leaves 


6 Sabatia bartramii sp. nov. Perennes a rhizomatibus. Folia crassa, succulenta, 
saepissime siccitate plus minusve rugosa; radicalia rosulata, spathulata vel oblanceo- 
lata; caulina lanceolata vel linearia, ultima supreme plerumque angustiora quam 
caulis diametro. Flores plerumque 10-12 partiti, calycis lobis setaceis vel subulatis, 
succulentis, saepe semiteretibus. Specimen typicum in savanna prope Pensacolam, 
in Florida, legerunt Webster et Wilbur, sub numero 3577, et in Herb. Universitatis 
Michiganensis conservatum, 


92 Rhodora [Vor. 57 


strongly dimorphie in appearance with an abrupt transition from the 
basal to the cauline; the basal strongly spreading, conspicuously rosulate, 
oblanceolate, somewhat oblong, or more typically broadly spatulate, 
strongly but gradually long-tapering to the almost petiolate base, obtuse 
or rarely acute, (2.5-)4-8.5(-13) em. long, (4-)12-17(-22) mm. wide; 
cauline strongly ascendent and becoming even closely appressed, the 
lower lanceolate but gradually, or even abruptly, reduced to linear 
or even very narrowly linear above, where their width is often about 
equal to, or less than, the diameter of the stem, (1.5-)2.5-5(-6.5) em. 
long, (1-)2-8(-15) mm. wide. Inflorescence of reduced cymules of 
commonly one and more rarely more than two flowers borne on slender, 
rigid, finely ridged pedicels (3-)4-8(-12) em. long, terminal or arranged 
on usually undivided lateral branches of one or more rarely of several 
nodes. Calyx-tube ridgeless, crateriform to campanulate, usually broadly 
so, occasionally, especially in smaller flowers, somewhat turbinate, (2-) 
3-4(-8) mm. long, usually straw-colored in strong contrast to the darkly 
chlorophyllose lobes. Calyx-lobes typically strongly subulate, (4—)8-12 
(-20) mm. long, often rather succulent, usually drying somewhat rugose- 
thickened, ellipsoidal below to nearly round above in cross-section, often 
somewhat revolute, erect, strongly ascendent or but weakly spreading, 
dark green in color. Corolla-tube cylindrical, (5-)6-8(-9) mm. long, 
usually about 2-3 times as long as the calyx-tube, apparently pale- 
yellow externally and darker within. Corolla-lobes (8-)10-12(-13) in 
number, (16-)22-32(-35) mm. long, (5-)7-10(-12) mm. wide, obovate- 
spatulate or rarely oblong to elliptie, usually broadly obtuse, deep rose- 
magenta, rose-pink, rose, or rarely white, with an irregularly but slightly 
toothed, yellow, oblong patch at the base of the lobe 3-5 mm. long, usually 
bordered by a dark red line. Filaments slender, (3-)4-6(-7) mm. long, 
pale yellow; anthers linear, slender, golden-yellow, 5-7 mm. long. Style 
4-6 mm. long; stigmatic lobes slender, 7-10 mm. long. Capsule ovoid, 
6-8 mm. long, 4-6 mm. in diameter. TYPE LOCALITY: savanna about 
9 miles west of Pensacola, Escambia Co., Florida. Type: Webster & 
Wilbur 3577 (MICH). DISTRIBUTION: Savannas or low pine barrens from 
southern Georgia and Alabama to southeastern Mississippi and nearly 
throughout Florida except for the extreme southern tip. Map 15. 


This very striking species is recognized at once by the strongly 
subulate calyx-lobes, which are thickened and semicircular in 
cross-section; the usually somewhat fleshy leaves; the obtuse, 
spatulate basal leaves contrasting strongly with the abruptly 
reduced cauline leaves, the uppermost of which are very nar- 
rowly linear being little, if any, broader than the diameter 
of the stem. These features are in very strong contrast with 
the thin, flat, linear to foliaceous calyx-lobes of the other southern 
species of the same area whose upper cauline leaves are always 
at least several times the diameter of the stem. Also the basal 


1955] Wilbur,— The North American Genus Sabatia 93 


rosette of those species is either absent or little developed or, 
if present, the cauline leaves are only very gradually reduced. 
The rather high percentage of misidentifications of specimens of 
this species in the past, or the inclusion of material of other 
species under its name, has been due largely to the key-differences 
as presented by Small. "The distinction first utilized by him was 
that of the length of the corolla-lobes. In this species, however, 
the length overlaps very considerably with the other species 
of the dodecandra-alliance commonly collected in the area. 
In the last mentioned work he added another key-distinction, 
that of the shape of the corolla-lobes, which also was poorly 
chosen, for that floral structure is also extremely variable. 


S. decandra, the name that has designated this species for 
more than fifty years rests upon a binomial published in Walter's 
"Flora Caroliniana" (1788). Walter's Flora provides very 
brief diagnoses of the six species of what was then called Chironia. 
Linnean binomials were applied to three of the entities and 
new ones provided for the remaining species. The characteriza- 
tion of them all is so brief that one is able to place them no more 
than tentatively even if quite familiar with the genus. Blake 
stated (Rhodora 17: 129. 1915.) that there are now but seven 
specimens of Sabatia in Walter's herbarium, located at the 
British Museum. 

A print has been obtained from the Gray Herbarium of the 
page bearing all the specimens of Sabatia still in Walter's col- 
lection. Although the photographie print was not such that 
absolute determination of all seven existing specimens in the 
genus could be made, it was more than sufficient to prove con- 
clusively that at present there is among the fragments no material 
representing the species that we have been calling S. decandra. 

The seven fragments are tentatively identified from the photo- 
graph as follows: 3 specimens of S. calycina (Lam.) Heller (= 
Chironia dichotoma Walt.); 2 specimens of S. difformis (L.) 
Druce (= Chironia lanceolata Walt.); 1 specimen of Sabatia 
dodecandra var. foliosa (Fern.) Wilbur (called Chironia dode- 
candra L. in the Flora); and what appears to be a specimen 
of S. stellaris Pursh (probably passing in Walter's Flora as 
Chironia campanulata L., a species often confused with the annual 
even by present day authors). 


94 Rhodora [Vor. 57 


In the absence of an authentic specimen of Chiron?a decandra, 
one must rely heavily upon the original description which is as 
follows: 

decandra 6. flor. decemfidis colore dodecandrae, foliis 

linearibus, caule rigido erecto. 
This description, as Harper has admitted (Bull. Torrey Club 27: 
432. 1900.), is to say the least “rather brief." However, Harper 
concluded that since “there is no known plant in the southeastern 
states which answers it exactly, and as his name for the species 
is cited in synonymy by both Elliott and Grisebach, no hesitation 
is felt in taking it up here." I do not believe that Walter's 
description is sufficient to limit it to the species to which Harper 
applied it. The characterization of a 10-parted corolla colored 
like S. dodecandra with linear leaves and an erect, rigid stem 
fits S. gentianoides Ell. equally as well as S. decandra in the 
sense of Harper. There are occasionally encountered specimens 
of S. dodecandra that match the few features mentioned and 
certainly S. kennedyana, which formerly was not known from 
outside of Nova Scotia and southern New England but is now 
known to be part of the flora of southeastern North Carolina 
and northeastern South Carolina, might often be briefly described 
by those same few words. 

The species to which Harper applied Walter's epithet is not 
as yet represented by a single herbarium specimen from South 
Carolina. In fact I have seen no specimens from north of the 
Ogeechee River in Georgia and only one specimen from north 
of the Altamaha River. Harper reports (Bull. Torrey Club 
37: 595. 1910.) having seen this species from a train window in 
Hampton County, South Carolina at two different locations. 
The nearest location represented by a herbarium specimen 
of the plant called S. decandra by recent authors is almost 150 
miles from Walter's plantation and even Harper's sight records 
are nearly 100 miles from the area delimited by Walter. 

The preface of “Flora Caroliniana" informs one that almost 
all of the species described were to be found within a radius 
of fifty miles of Walter's plantation on the banks of the Santee 
River near the town of St. Stephen. Fraser (1789) adds to 
our knowledge of the source of materials included in the Flora 
in the following manner: 


1955] Wilbur,—The North American Genus Sabatia 95 


The botanical description of many of the plants which I found are contained 

in the Flora Caroliniana; with the author of which, the late Mr. Walter, I 
became acquainted soon after my arrival in Carolina. He had collected, 
when I went into that country, plants which afforded him six hundred and 
forty descriptions. I increased his work, by the specimens I produced to 
him, to one thousand and sixty; amongst which are upwards to two hundred 
new species, and thirty new genera; of all which, as well as the other plants in 
the Flora, I have now dried specimens in my possession, and many valuable 
living plants. Many of the most valuable specimens and living plants I 
collected are still remaining in my hands undescribed. 
Fraser by his own account spent nineteen months on this trip 
and collected over thirty thousand specimens. During his 
travels he ranged “from the south boundaries of Georgia to 
the northward of Carolina." Upon these journeys he would 
have had ample opportunity to encounter the numerous species 
that have puzzled botanists in the past since those species at 
present are not known from Walter’s neighborhood or even from 
South Carolina. Even so it would appear that most of Fraser's 
additions to the flora came from the limited area prescribed 
by Walter. 

Lacking authentic material of Chironia decandra and in view 
of the extremely brief and inconclusive original description, 
it is now impossible to determine the identity of the name. 
The description is so generalized that it is equally applicable 
to at least two species known from the area from which most of 
the specimens in the Flora are stated to have been found. 

For the reasons stated above I feel that the assignment of 
Walter's epithet to the pluripetalous southern perennial with 
subulate calyx-lobes is not justified by the available evidence. 
I am naming this plant Sabatia bartramii. William Bartram 
prepared an unmistakable sketch of the upper portion of this 
species which has recently been reproduced (Amer. Phil. Soc. 
Trans. 33: Plate XXIV. 1943.). It is quite possible that the 
nomen nudum mentioned by Bartram in his “Travels” was this 
species. More than sixty miles south of the Altamaha River, 
according to his own estimate, he describes crossing a land of 
“high open forest of stately pines, flowering plains, and extensive 
green savannas, chequered with the incarnate Chironia pulcher- 
rima, and Asclepias fragrans . . ." 


REPRESENTATIVE SPECIMENS:—GEORGIA: Appling Co., near Baxley, Bilt- 
more Herb. 14965b (us); Baker Co., near Bethany, Eyles 7247 (cu, GH); 
Brantley Co., 3 mi. e. of Nahunta, Wilbur & Webster 2746 (micu); Bryan Co., 


96 Rhodora [Vor. 57 


about 2 mi. w. of Pembroke, McKay, 14 Aug. 1930 (micH); Calhoun Co., 
5 mi. e. of Arlington, Thorne 5444 (cu, GA); Charlton Co., near Saddlebag 
Pond, Wright 871 (cv); Clinch Co., 4 mi. e. of Homerville, Eyles 6331 (Ga); 
Cook Co., 0.5 mi. se. of Sparks, Wilbur & Webster 2691 (micu); Miller Co., 
in sw. corner of county just north of Donalsonville, Duncan 6755 (MICH); 
Sumter Co., without exact locality, Harper 461 (F, GH, MO, NY, US); Ware Co., 
5 mi. se. of Waycross, Wilbur & Webster 2737 (MICH). FLORIDA: Alachua Co., 
Waldo, West, 14 June 1927 (rtas); Baker Co., 9 mi. s. of Macclenny, West & 
Arnold, 12 July 1946 (rras); Bay Co., 8 mi. n. of Lynn Haven, Webster & 
Wilbur 3623 (micu); Broward Co., Ft. Lauderdale, Eaton 798 (F, GH); Clay 
Co., 2 mi. n. of Goldhead Branch State Park, West & Arnold, 15 July 1947 
(FLAS); Collier Co., near Fort Shackleford, Small $340 (DUKE, GH, NY, TENN, 
us); Dade Co., Humbugus Prairie, Small, Mosier, & Small 6885 (xv); DeSoto 
Co., without exact locality, Schallert, 29 July 1940 (vanRk); Dixie Co., s. of 
Oldtown, West & Arnold, 11 Aug. 1937 (FLAS); Duval Co., Baldwin, Nash 
2250 (F, aH, MICH, MO, NY, US); Escambia Co., 9 mi. w. of Pensacola, Webster 
& Wilbur 3577 (wricg); Flagler Co., near Andalusia, Arnold, 25 June 1942 
(FLAS); Franklin Co., near Apalachicola, Biltmore Distrib. Chapman Herb. 
3305b (an, Ny, Us); Gulf Co., 7 mi. n. of Port St. Joe, Small & West, 8 Aug. 
1935 (FLAS); Hardee Co., near Limestone, Kirk, 8 July 1942 (rLas); Hernando 
Co., Bayport, Jones 76 (cu, us); Lee Co., about 8 mi. se. of Fort Myers, 
Standley 447 (F, GH, Mo, Us); Levy Co., 14 mi. se. of Chiefland, Webster & 
Wilbur 3640 (micH); Manatee Co., Manatee River, Rugel 155 (Mo, us); 
Nassau Co., 0.5 mi. s. of Callahan, Wright 873a (cv); Orange Co., without 
exact locality, Fredholm 5374 (Gu, us); Osceola Co., Kissimmee, Eaton 1062 
(F, GH); Palm Beach Co., 1 mi. n. of Lake Park, Hannon, 12 June 1948 (FLAS); 
Pasco Co., south of Denham, Hood 3572 (ras); Polk Co., Polk City, McFarlin 
5979 (MICH, TEX); St. Johns Co., St. Augustine, Garber, July 1876 (us); 
Sarasota Co., Myakka, Barrett 45 (us); Volusia Co., near Seville, Curtiss 6843 
(GH, KSC, MO, NA, NY, US); Wakulla Co., 1.5 mi. s. of Sopchoppy, Webster & 
Wilbur 3629 (micu); Washington Co., 4 mi. w. of Chipley, Webster & Wilbur 
3611 (MICH). ALABAMA: Baldwin Co., 1 mi. n. of Stapleton, Webster & 
Wilbur 3522 (micu); Mobile Co., Crichton, Sawada, 18 Aug. 1933 (NY); 
Monroe Co., Claiborne, Blanton 53 (GH, MT, NO, NY, OKLA, US). MISSISSIPPI: 
Jackson Co., about 4 mi. e. of Moss Point, Webster & Wilbur 3466 (mic). 


II. Section PsrupocHIRONIA Griseb., Gen. et Sp. Gent. 125. 1839. 


Subg. Plurimaria Raf., Med. Fl. 2: 76. 1830, in part, not Plurimaria 
Raf. (as genus), Fl. Tell. 3: 31. 1837. 
Pleienta Raf., Fl. Tell. 3: 30. 1837, in part, an illegitimate name 
since its type is the same as that of Sabatia. 
Lapithea Griseb., Prodr. 9: 48. 1845. 
Subg. Pseudochironia (Griseb.) Blake, RHoporaA 17: 56. 1915. 
Annuals. Floral parts plurimerous, 7-12-parted. Flowers sessile or 
very nearly so and borne either solitarily or more typically in compact, 
capituliform eymules. Anthers about half-twisted laterally even prior 
to pollen-discharge and never becoming revolute. TYPE SPECIES: Sabatia 
gentianoides Ell. 


The two species comprising this group are very strongly 
differentiated from the other eastern American species belonging 


1955] Wilbur,— The North American Genus Sabatia 97 


to the subtribe Erythraeinae with strongly bilobed, linear stig- 
matic branches. Their distinctive features are such that 
Grisebach, after first placing the only species known to him 
in his section Pseudochironia coordinate with all the other species, 
finally six years later decided upon generic status calling it 
Lapithea. Bentham (1873) maintained this group as the genus 
Lapithea as did Gilg (1895). Small treated the group as of 
generic rank in all of his work. Blake (1915) discussed the 
differences and concluded that ‘‘the group is better treated as 
of subgeneric value." Subgeneric status still seems to express 
best the phylogeny and relationship of the group without be- 
littling the distinctive features that so distinguish the two 
species from all others in the genus. The two groups of species 
are here considered to be of sectional rank. 

The two species are both very easily distinguished from the 
species comprising the rest of the genus and also from each other. 
S. capitata, known only from the ancient Appalachian highlands, 
is believed to be more primitive than the Coastal Plain $. 
gentianoides which possesses greatly reduced and modified foliage 
in comparison to that of the supposedly more primitive species. 


KEY Tro THE SPECIES OF SECTION PSEUDOCHIRONIA 


A. Basal and cauline leaves not strongly contrasted in either size or shape; 
cauline leaves oblong, elliptic, or rarely weakly lanceolate, usually broader 
than 1 em. and never less than 5 mm. wide, several to many times the 
diameter of the stem in width; calyx-lobes linear, their margins parallel 
for most of their length, erect................ 020 .00005 16. S. capitata. 

A. Leaves strongly dimorphic, the basal strongly contrasting to those of the 
stem in both size and shape; cauline leaves very narrowly linear, less than 
5 mm. wide, usually equaling or less than the diameter of the stem, 20—60 
times longer than broad; calyx-lobes subulate, the tips often recurved or 
^2 2c» PM cis UMEN 17. S. gentianoides. 


16. Sabatia capitata (Raf.) Blake, RHopora 17: 54. 1915. Pleienta 
capitata Raf., Fl. Tell. 3:30. 1837. Sabbatia Boykinii A. Gray in Chapm., 
Fl. S. U. S. 354. 1860. S. gentianoides B. Boykinii (A. Gray) Wood, 
Am. Bot. & Flor. 266. 1870. Lapithea Boykinii (A. Gray) Small, 
Fl. SE. U. S. 929. 1903. Lapithea capitata (Raf.) Small, Man. SE. 
Fl. 1051. 1933. 

Erect annual (15-)30-45(-70) em. high; stem 1-4 mm. in diameter, 
rigid, hollow, more or less terete to slightly 4-angled, smooth or often, 
upon closer examination, with few, very slightly raised lines extending 
between the internodes. Branches usually present except in depauperate 
plants, either opposite or alternate but in well-developed plants probably 
more commonly opposite, often restricted to the upper third of the stem 


98 Rhodora [Vor. 57 


S. DODECANDRA "m 
€ VAR. DODECANDRA > ` 
© VAR. FOLIOSA S. KENNEDYANA 


S.BARTRAMII 


oS CAPITATA 
\ @ S.GENTIANOIDES 
Mars 13-16. Map 13, upper left; map 14. upper right; map 15, lower left; map 16, 
lower right. 


but occasionally branching nearly to the base, varying greatly in length 
ranging from (1-)3-8(-30) em. long, rather strongly ascendent forming 
an angle of about 20-40 degrees with the stem. Root-system of several 
to numerous fibrous roots usually about 3-8 em. long, the larger of which 
are often somewhat woody and up to 2 mm. in diameter. Leaves weakly 
if at all dimorphic, the basal not strongly contrasting with the cauline 
in either shape or size, thin, not at all succulent, drying thin, smooth 
and brittly chartaceous, venation rather conspicuous with 3-5 nerves at 
least somewhat elevated beneath, the midvein especially so; the basal, 
when present and well-developed, obovate to broadly spatulate, sometimes 
rosulate, obtuse, tapering to the almost petiolate base, not strikingly 
contrasted to those above, 2.3-5 cm. long, 1-2 em. wide, the cauline 
broadly, or more rarely narrowly, oblong, elliptie, or weakly lanceolate, 
usually about 2-4 times longer than broad, about 5-15 times as wide 
as the diameter of the stem, weakly ascending, strongly spreading or 
even somewhat reflexed, (2.5-)3.5-5(-6.5) em. long, (7-)10-20(-25) mm. 


1955] Wilbur,— The North American Genus Sabatia 99 


wide, broadly obtuse or sometimes even acute, typically the base some- 
what clasping. Inflorescence cymose, terminating the main stem or the 
lateral branches. Flowers sessile or very nearly so, borne singly or in 
few to several (2-5 or very rarely more)-flowered compact clusters. Each 
flower closely associated with 2 subtending oblong to elliptic bracts about 
1-2 em.long. Calyx-tube broadly campanulate, 3-6 mm. long, smooth, 
unnerved, membranous. Calyx-lobes linear, 4-10 mm. long, and about 
1-2 mm. wide, erect or strongly ascendent, often strikingly unequal, 
7-12 in number, about 1-2 times the length of the calyx-tube and about 
half the length of the corolla. Corolla-tube about 5-7 mm. long or 
about 1-2 mm. longer than the calyx-tube, probably pale greenish-yellow. 
Corolla-lobes about 12-25 mm. long, 5-13 mm. wide, elliptical, obovate 
or spatulate, acute to more typically rounded-obtuse, pale rose-pink, 
pink, or rarely white with an unlobed, pale yellow semicircle at the base 
about 1-2 mm. high. Filaments slender, (2-)3-4 mm. long; anthers 
stout, slightly twisted laterally even prior to pollen discharge, bright 
yellow, (2.5-)3-5(-6) mm. long. Style 3-5 mm. long; stigmatic branches 
spatulate, 6-9 mm. long. Capsule apparently ovoid. TYPE LOCALITY: 
“Unaka and Cherokis Mts. TYPE: unknown. DISTRIBUTION: Open 
hardwoods on hillsides and ridges in southwestern North Carolina (?), 
northeastern Georgia, southeastern Tennessee, and northeastern and 
central Alabama. Map 16. 


This species, strikingly different from all others, has by far 
the most restricted range of any in the genus. It is also the 
only species which is completely limited to the southern Ap- 
palachian uplands, a biological reservoir from which spread 
much of the ancestral flora of eastern North America. 

This species was first discovered in the Unaka and Cherokee 
Mountains along the southern portion of the boundary between 
North Carolina and Tennessee and was very fully described by 
Rafinesque in the “Flora Telluriana" (1837). However, like 
most of that author's work, it was apparently not given serious 
attention and the identity of his name was not made known for 
over seventy-five years. "The species was first effectively brought 
to the attention of “reputable botanists" nearly twenty-five 
years after Rafinesque's description when it was published as 
S. Boykinii in Chapman's Flora (1860), and attributed to Gray. 
This was based upon material in Torrey's herbarium sent by 
Dr. Boykin from “middle Georgia." There it was seen by Gray 
and according to the annotation was apparently intended as 
a joint publication of Torrey and Gray. The remaining syn- 
onymy of this little collected species involves merely various 
combinations with either of these basionyms. 


100 Rhodora [Vor. 57 


REPRESENTATIVE SPECIMENS:—NORTH CAROLINA: County unknown but 
probably Cherokee Co., "from Cherokee" collector and date unknown (Mo). 
GEORGIA: Bartow Co., 3.8 mi. n., 14? w. of Allatoona Dam, Duncan 8613 (aA); 
Catoosa Co., Catoosa Springs, Biltmore Herb. 4512b (us); Walker Co., Pigeon 
Mt., Wilson 172 (Ny, vs); Whitfield Co., base of Dick's Ridge, Wilson 142 
(Ny, vs); County unknown, Boykin (aH, NY, Us, TYPE and ISoTYPES of S. 
boykinii). TENNESSEE: Marion Co., Cumberland Mts. at Whiteside, White, 
22 July 1895 (us). ALABAMA: Blount Co., without exact locality, Rugel, 
Oct. 1843 (Ny); Cherokee Co., Lookout Mt., Freeman, 11 July 1905 (ksc); 
Clay Co., Millerville, Pollard & Mazon 173 (Ny, vs); DeKalb Co., Val- 
ley Head, Biltmore Herb. 4512e (us); Etowah Co., near Coosa River, Mohr, 
July 1880 (vs); Jackson Co., Eliza, Graves 1191 (mo); Marshall Co., 
3.2 mi. n. of Boaz, Hubricht B1668 (rus, Mo); Shelby Co., Calera, Everts 
(Ny); Talladega Co., s. of Childersburg, Hood 285 (Frias); Tuscaloosa Co., 
without exact locality, Smith, July (us). 


17. Sabatia gentianoides Ell, Sk. Bot. S. C. & Ga. 1: 286. 1817. 
Pleienta gentianoides [Ell.] Raf., Fl. Tell. 3: 30. 1837, without basionym. 
Lapithea gentianoides (Ell) Griseb., Prodr. 9: 48. 1845. Sabbatia 
oligophylla Featherman, Rep. Bot. Surv. S. & Cent. La. 72. 1871. 

Erect annual (15-)30-50(-65) em. high; stem 1-3 mm. in diameter, 
rigid, hollow, more or less terete but with fine and irregular internodal 
ridges. Very commonly unbranched below the terminal inflorescence, 
but when branched, the branches usually restricted to the upper quarter 
of the stem, occasionally branched in the lower half or even more rarely 
with several stems arising from the base; commonly either opposite or 
alternate but more typically alternate, usually about 5-8 em. long or 
rarely up to 20 cm., strongly ascendent forming an angle of about 20-40 
degrees with the stem.  Root-system of several to numerous slender, 
fibrous roots 3-8 em. long and less than 1 mm. in diameter. Leaves 
thick and somewhat succulent, drying rather thickly chartaceous and 
rugose, strikingly dimorphic, venation obscure except for the midvein; 
those of basal rosette wide-spreading, oblong to orbicular-spatulate, 
typically long-persistent, (1—)2-3(-3.5) em. long, (4-)8-12(-15) mm. 
wide, usually about 2-3 times as long as wide, obtuse, rather strongly 
tapering to the base and therefore appearing petiolate; the cauline very 
narrowly linear, 20-60 times longer than broad, width equaling or less 
than the diameter of the stem, strongly ascendent and sometimes even 
appressed especially in the lower portion of the leaf, (15-)40-80(-100) 
mm. long, 1-3 mm. wide, acute and slightly callous-tipped, sessile or clasp- 
ing the stem. Flowers sessile or very nearly so, borne singly at the 
principal apex or at that of the lateral branches, or more typically in 
more robustly developed plants, especially at the apex of the principal 
axis, in few to several (2-5 or rarely even 7) flowered compact clusters. 
Each flower closely associated with 2 subtending, narrowly linear bracts, 
(1-)1.5-4(-7) em. long. Calyx-tube broadly campanulate, (3-)4-6(-8) 
mm. long, smooth and unnerved, membranous, light green to pale yellow- 
ish-green. Calyx-lobes conspicuously subulate, (3-)5-10(-17) mm. long, 
spreading-ascending, arching outward especially at the tip, often unequal 
in size, 7-12 in number, 1-2 times as long as the calyx-tube, and usually 


1955] Wilbur,—' The North American Genus Sabatia 101 


about half or less than half the length of the corolla. Corolla-tube 6-10 
mm. long, about 1.5-2 times as long as the calyx-tube, pale greenish- 
yellow, cylindrical. Corolla-lobes (12-)18-24(-30) mm. long, (4-)6-8(-11) 
mm. wide, elliptical, spatulate to oblanceolate, acute to obtuse, pink 
to deep rose with an unlobed greenish-yellow area at the base of the 
lobe 2-3 mm. high. Filaments slender, short, often about half as long as the 
anthers, 1-3 mm. long, pale yellow; anthers stout, slightly twisted laterally 
even prior to pollen discharge, golden-yellow, (3-)4-6(-7) mm. long. Style 
5-8 mm. long; stigmatic branches spatulate, 4—7 mm. long, 1.5-3 mm. wide. 
Capsule ovoid, 7-10 mm. high, about 6 mm. wide at the base. TYPE LOCAL- 
ITY: “in Bullock Co., Georgia." Type: Mr. Abbott s.n. (CHARL!). DISTRIBU- 
TION: Pinelands and savannas along the Coastal Plain from North Caro- 
lina south into Florida and west into eastern Texas. Map 16. 


This species is unmistakable once the distinctive characteristics 
of the plant are understood. "The combination of a pluripetalous 
corolla and an annual habit alone set this species off from all 
the members of Section Husabatia. These two features together 
with sessile to sub-sessile flowers arranged in capitate to sub- 
capitate clusters, and the laterally half-twisted anthers (as 
opposed to revolute) are shared by both S. gentianoides and 
S. capitata which together form section Pseudochironia. S. 
gentianoides is not only clearly distinguished morophologically 
by its strongly dimorphic basal and cauline leaves but also 
geographically by its restriction to the Coastal Plain which 
contrasts sharply with the southern Appalachian endemic, 
S. capitata. | Elliott/s species is one of the most easily recogniza- 
ble and nomenclatorially least misunderstood of all the species. 
In spite of this it has been surprisingly often misidentified; 
ten to fifteen per cent of the specimens that I have seen were 
misnamed. This is at least in part due to the incomplete 
ranges published in Small’s Flora (1903) “Ga. to Fla." and also 
in his Manual (1933) “Fla. to Tex. and Ga." The species is 
known to range from Texas to Florida and northward throughout 
eastern North Carolina. 


That S. oligophylla is a synonym of S. gentianoides is strongly 
indicated by the original description and conclusively shown 
by a water-color plate drawn for Professor Featherman and 
preserved at the Gray Herbarium. Featherman thought that 
it differed from S. gentianoides for that species “which it seems 
to resemble, has no bracts.” S. gentianoides does, of course, 
possess bracts at the base of each flower. 


102 Rhodora [Vor. 57 


REPRESENTATIVE SPECIMENS:—NORTH CAROLINA: Bladen Co., on Rt. 211, 
2.1 mi. ne. of Columbus Co. line, Fox & Whitford 1867 (Ncs); Brunswick Co., 
2.5 mi. s. of Grissettown, Fox & Godfrey 2836 (GH, MICH, NCS, NCU); Carteret 
Co., Newport, Godfrey 5789 (aH, vs); Columbus Co., 1 mi. se. of Delco, Fox 
& Godfrey 2872 (ncs, smu); Duplin Co., 8 mi. w. of Richlands, Godfrey 5784 
(aH, Us) Johnston Co., s. of Selma, Dlans, 28 July 1933 (ncu); Nash Co., 
between Bailey and Middlesex, Oosting 35574 (pUKE); New Hanover Co., 
near Wilmington, Abbe & Spalteholz, 3 Aug. 1927 (cu, ncu); Northampton 
Co., near Jackson, Rhoades, Aug. 1935 (aH); Onslow Co., near Folkstone, 
Alexander, 25 July 1923 (xcv); Pender Co., Burgaw, Godfrey 5924 (an, NCS, 
us); Wake Co., just w. of Raleigh, Fox 1828 (DUKE, aH, Ncs). Also represented 
(F, NY) by specimens reputedly from Statesville, Iredell Co., made by M. E. 
Hyams. These and other Coastal Plain plants known only from the Pied- 
mont by his collections are suspected of having been gathered in the Coastal 
Plain and distributed with his printed Statesville labels. SOUTH CAROLINA: 
Georgetown Co., 12 mi. nw. of Georgetown, Godfrey & Tryon 745 (DUKE, F, 
GH, MO, NY, TENN, Us); Horry Co., 6 mi. e. of Loris, Wilbur 2902 (micH). 
GEORGIA: Bullock Co., without exact locality, Abbot Tyre of S. gentianoides 
(CHARL); Calhoun Co., 3 mi. e. of Cordrays Pond, Thorne 6476 (cv); Charlton 
Co., 8 mi. s. of Folkston, Wright 863 (cu); Dodge Co., Eastman, Biltmore 
Herb. 4509c (Ny); Sumter Co., without exact locality, Harper 1121 (aH, Mo, 
Ny, US); Worth Co., 3.5 mi. w. of Sylvester, Thorne 6350 (F, GA, MT, US). 
FLORIDA: Baker Co., without exact locality, Curtiss 2233 (CU, F, MO, NY, US, 
yu); Bradford Co., w. of Starke, Bratley & Murrill, 5 July 1940 (FLas); Clay 
Co., Doctor's Inlet, West & Arnold, 29 June 1939 (DUKE, FLAS, MO, US); 
Escambia Co., near Ala. state line, Porter, 21 July 1938 (rr4s); Franklin Co., 
near Apalachicola, Biltmore Dist. Chapman Herb. 4509a (Gu, Ny, us); Gulf 
Co., Wewahitchka, Chapman (Mo); Indian River Co., 5 mi. n. of Vero Beach, 
Woodson & Schery 126 (mo); Jackson Co., w. of Marianna, Porter, 21 July 
1938 (FLAS); Liberty Co., near Bristol, Arnold, 23 July 1940 (rLAs); Nassau 
Co., Boulogne, Hume & West, 14 July 1933 (FLas); Okaloosa Co., about 3 mi. 
e. of Crestview, Webster & Wilbur 3589 (micn); St. Johns Co., near St. Augus- 
tine, Garber, July 1876 (BRU, F, NY, US, YU); Walton Co., Argyle, Curtiss 6484 
(CU, GH, MO, NY, SMU, US). ALABAMA: Baldwin Co., 8 mi. s. of Foley, Webster 
& Wilbur 3532 (mic); Butler Co., near Bolling, Smith, 28 Aug. 1885 (F, Mo, 
us); Mobile Co., about 12 mi. sw. of Mobile, Webster & Wilbur 3478 (micuH). 
MISSISSIPPI: Forest Co., about 8 mi. se. of Hattiesburg, Webster & Wilbur 
3379 (micH); Hancock Co., Bay St. Louis, Langlois, 12 Sept. 1883 (PENN); 
Harrison Co., Biloxi, Tracy 7007 (cv, F, GH, Mo, US); Jackson Co., 2 mi. e. 
of Ocean Springs, Webster & Wilbur 3450 (micu); Pearl River Co., Picayune, 
Reed, 2 Aug. 1934 (NA); Perry Co., 9 mi. n. of Beaumont, Webster & Wilbur 
8420 (MICH). LOUISIANA: Allen Parish, Oakdale, Bomhard, 22 June 1933 
(No); Calcasieu Parish, 3 mi. sw. of Vinton, Webster & Wilbur 3212 (micn): 
Grant Parish, 4 mi. s. of Pollock, Webster & Wilbur 3258 (micu); Natchitoches 
Parish, about 2 mi. s. of Kisatchie, Correll 9796 (puxe); Orleans Parish, New 
Orleans, Carpenter (Mo); Rapides Parish, Alexandria, Hale (BRU, F); St. Landry 
Parish, Opelousas, Carpenter (an, Nv); St. Tammany Parish, 1 mi. n. of Abita 
Springs, Pennell 4150 (PENN); Vernon Parish, 2 mi. w. of Leander, Webster & 
Wilbur 3241 (mica). Texas: Hardin Co., Sour Lake, Tharp, 29 July 1929 
(tex); Harris Co., Houston, Hall 509 (BRU, F, GH, MO, NY); Henderson Co., 
without exact locality, T'harp 2882 (vs); Houston Co., Grapeland, Palmer 
14405 (Mo, vs); Jasper Co., Jasper, Fisher 32117 (F); Jefferson Co., Beaumont, 
Hooks, 14 July 1931 (rEx); Liberty Co., 21 mi. se. of Cleveland, Webster & 


1955] Wilbur,—The North American Genus Sabatia 103 


Wilbur 3171 (wmicH); Orange Co., Orange, Letterman, 11 Aug. 1880 (wo); 
Rusk Co., without exact locality, Vinzent 39 (mo); Smith Co., without exact 
locality, Tharp, 5 July 1924 (rex); Trinity Co., Big Thicket, Murray, 1 June 
1938 (NA); Tyler Co., 2 mi. n. of Warren, Webster & Wilbur 3204 (Micn). 


DOUBTFUL AND EXCLUDED SPECIES 


CHIRONIA DECANDRA Walt., Fl. Car. 95. 1788. Nomen dubium. See 
discussion under Sabatia bartramii Wilbur. 

CHIRONIA PULCHERRIMA Bartram, Bartr. Trav. 19. 1791. Nomen nudum. 

PLEIENTA FASCICULATA Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. 

PLEIENTA QUINQUENERVIA Raf., New Fl. 4: 91. 1838. Perhaps a 
synonym of S. calycina (Lam.) Heller. 

PLEIENTA RIGIDA Raf., Herb. Raf. 80. 1833. 

SABBATIA ALBEOLA Raf., Cat. Bot. Gard. Transyl. Univ. 15. 1824. 
Nomen nudum. 

SABBATIA AMENA Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. Probably 
a misprint for S. amoena (= S. stellaris Pursh). 

SABBATIA ANCEPS Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. 

SABBATIA AUSTRALIS Cham. & Schlect., Linnaea 1: 194. 1826.  — Zygo- 
stigma australe (Cham. & Schlect.) Griseb., Gen. et Sp. Gent. 50. 1839. 

SABBATIA CENTAURIUM (L.) Raf., Casket 1827: 316. f.29-1827. Pub- 
lication not seen. Merrill (1949) states that this name was based on 
Gentiana Centaurium L. (= Centaurium umbellatum Gilib.). 

SABBATIA DECANDRA (Walt.) Harper, Bull. Torrey Club 27: 432. 1900. 
Nomen dubium. See discussion under S. bartramii Wilbur. 

SABBATIA DECLINATA Raf., Herb. Raf. 69. 1833. Nomen nudum. 
Probably a synonym of S. stellaris Pursh. 

SABBATIA LINGULATA Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. 

SABBATIA NIVEA Raf., Med. Fl. 2: 77. 1830. Briefly discussed under 
S. quadrangula Wilbur. 

SABBATIA OBTUSIF. [obtusifolia] Raf., Fl. Tell.3:30. 1837. Nomen nudum. 

SABBATIA PETIOLATA Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. 

SABBATIA PULCHELA Raf., Herb. Raf. 69. 1833. Publication not seen. 
Merrill (1949) states that it was listed from “Long Island, New Jersey, 
Chesapeake Bay" and “= Sabatia gracilis Salisb. = S. campanulata 
(Linn.) Britton." However, if it is not the Centaurium listed below, 
it is more likely a synonym of Sabatia stellaris Pursh. 

SABBATIA PULCHELLA (Sw.) Spreng., Syst. ed. 16. 581. 1825. = Cen- 
taurium pulchellum (Sw.) Druce. 

SABBATIA PUMILA Raf., Herb. Raf. 69. 1833. Publication not seen. 
Merrill (1949) provides the following information: *nom. Long Island, 
New Jersey, Chesapeake Bay." Perhaps, judging from the localities, 
a synonym for S. stellaris Pursh. 

SABBATIA SIMPLEX Bertol., Misc. Bot. 10: 27. (1842-1863). Publica- 
tion not seen. A. Gray (Syn. Fl. 2: 116. 1878.) equates this name 
with Rhexia stricta Pursh. 

SABBATIA STRICTA Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. 

SABBATIA TENUIFOLIA Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. 


104 Rhodora (Vor. 57 


SABBATIA UMBELLATA Raf., Fl. Tell. 3: 30. 1837. Nomen nudum. 
SABBATIA VERTICILLARIS Spreng., Syst. ed. 16. 580. 1825. Described 
from South America. Grisebach (Prodr. 9: 50. 1845.) equates this 
name with Schuebleria stricta Mart., a synonym of Curtia gentianoides 
Cham. & Schlecht. 
LITERATURE CITED 

Apanson, M. 1763. Familles des plantes. Paris. 2 vols. cecxxv, 189; 640 pp. 

BENTHAM, G., & Hooker, J. D. 1862-1883. Genera plantarum. London. 
3 vols. (Gentianaceae by Bentham 2(2): 799-820. 1873.) 

BLAKE, S. F. 1915. Notes on the genus Sabatia. Rnmopoma 17: 50-57. 
pl. 112. 

Brirron, N. L., & Miispaucu, C. F. 1920. The Bahama flora. New 
York. viii, 695 pp. 

CHAPMAN, A. W. 1860. Flora of the southern United States. Ivison, 
Phinney and Company. New York. xxxvii, 621 pp. 

Dram, C. C. 1940. Flora of Indiana. Indiana Dept. of Conservation, 
Division of Forestry. Indianapolis. 1236 pp. 

Dow, G. 1838. A general history of dichlamydeous plants. J. G. and F. 
Rivington. London. 4 vols. (Sabatia in 4: 206-207.) 

ELLIOTT, S. 1816-1824. A sketch of the botany of South Carolina and 
Georgia. J. R. Schenck. Charleston. 2 vols. 666; viii, 741 pp. 
(Sabatia in 1: 282-287. 1817.) 

FERNALD, M. L. 1950. Gray’s manual of botany. ed. 8. The American 
Book Company. New York. lxiv. 1632 pp. 

Fraser, J. 1789. A short history of the Agrostis Cornucopiae;—and also 
some account of a journey to the Cherokee Nation in search of new 
plants. London. $8 pp. 

Gita, E. 1895. Gentianaceae in Die natürlichen Pflanzenfamilien. 4(2): 
50-108. 

Gurason, H. A. 1952. The new Britton and Brown illustrated flora of the 
northeastern United States and adjacent Canada. Lancaster. 3 vols. 
(Sabatia in 3: 56—58.) 

Gray, A. 1848. A manual of the botany of the northern United States. 
J. Munroe and Company. Boston. lxxii, 710 pp. 

. 1878. The synoptical flora of North America. ed. 1. (Sabatia 
in 2(1): 113-116.) 

GmisEBACH, A. H. R. 1839. Genera et species gentianearum. Stuttgart. 

viii, 364 pp. 
. 1845. Gentianaceae in Prodromus systematis naturalis regni 
vegetabilis. 9: 38-141. 

Gronovius, J. F. 1739. Flora Virginica. Leiden. 

Lansouw, J., & STAFLEU, F. A. 1952. The herbaria of the world. Regnum 
Vegetabile 2: 1-167. 

Linnagus, C. 1753. Species plantarum. Stockholm. 2 vols. 

Merrit, E. 1949. Index Rafinesquianus. Arnold Arboretum. ix, 296 pp. 

Purs, F. 1814. Flora americae septentrionalis. White, Cochrane and 
Company. London. 2vols. xxxvi, 75lpp. (Sabatia in 1: 137, 138.) 

SMALL, J. K. 1903. Flora of the southeastern United States. New York. 
xii, 1370 pp. 

. 1933. Manual of the southeastern Flora. New York. xxii, 

1554 pp. 


Volume 57, no. 674, including pages 37—72, was issued 11 March, 1955. 


Douora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS ( 

RALPH CARLETON BEAN Associate Editors 
RICHARD ALDEN HOWARD ( 

CARROLL EMORY WOOD, JR. 


Vol. 57 April, 1955 No. 676 
CONTENTS: 

Evidence for the Hybrid Origin of Drosera anglica. Carroll E. 
Wood, Jr: o oo is 052,9, od ee eee sa cs cn cs oun 105 

Epipactis Helleborine in Illinois. Julian A. Steyermark......... 131 


Northeastward Range Extensions for Two Species of Hypericum 
in Nova Scotia. Daud Eraka.. osos ke. 132 


an A EE EP ORITUR os 132 


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EVIDENCE FOR THE HYBRID ORIGIN OF 
DROSERA ANGLICA 


CARROLL E. Woop, JR. 


AMPHIPLOIDY às à mechanism whereby new species of plants 
come suddenly into being has been demonstrated in a number 
of groups of plants and new examples continue to come to light. 
These include both artificial species unknown in nature and an 
increasing number of wild and cultivated species for which an 
origin in the past by hybridization followed by polyploidy has 
been either proved or postulated. (See Stebbins, 1950, Chap. 
8. That this process continues to operate in nature has been 
shown by three wild species which are known to have originated 
by amphiploidy within historic time (Spartina | Townsendii, 
Tragopogon mirus and T. miscellus). From the nature of 
amphiploidy it would be expected that such species might 
spring into existence more or less simultaneously at two or more 
different points, thus being polytopic in origin. This indeed 
appears to be the situation with Tragopogon mirus, for which 
Ownbey (1950) and Ownbey & McCollum (1954) have presented 
evidence of independent origin at three separate localities. 
Similarly, an independent origin presumably is true for each of 
five colonies of T. miscellus reported by the same authors. It 
may also be expected that a widespread and successful species 
of amphiploid origin might be synthesized again and again in 
the course of time as long as the diploid ancestors continue in 
existence and hybrids between them are genetically and eco- 
logically possible. Ordinarily, it would be difficult to detect 
such an event occurring under natural conditions unless the 
diploids and the original amphiploid occupy different areas 


106 Rhodora [Vor. 57 


at the time of resynthesis, thus making it possible to distinguish 
the new amphiploid from the old. It is with such an event, 
however, that this report 1s concerned: the finding of sterile 
hybrids between two diploid sundews, Drosera linearis and D. 
rotundifolia, and the chance finding of a small colony of a fertile 
hybrid of the same parentage which, from all of the evidence 
available, appears to be a spontaneous resynthesis of the cir- 
cumpolar Drosera anglica, the only known tetraploid of this 
genus in the northern hemisphere. 

It may be remarked both for the historical interest attached 
and for its bearing upon the origin of Drosera anglica that a 
hybrid origin for this species was suggested by Winge (1917) 
in the classical paper in which he proposed the origin of polyploid 
series of species by hybridization followed by chromosomal 
doubling. As Stebbins (1950) has pointed out, he apparently 
was at the time unaware of Digby’s work (1912) on Primula 
kewensis, the only concrete supporting example. Winge cited, 
however, the equally classical work of Rosenberg (1904, 1909) 
on a triploid hybrid in Drosera which had focused attention on 
polyploidy and on chromosomal behavior in hybrids. 

Rosenberg reported that in Drosera rotundifolia, with 10 pairs 
of chromosomes, and Drosera anglica, with 20 pairs of chromo- 
somes, meiosis is normal with the chromosomes pairing and 
disjoining regularly. However, in D. X obovata, the hybrid 
between them, he found that the 30 chromosomes formed 10 
bivalents which disjoined normally while the remaining 10 
chromosomes remained unpaired and were distributed irregu- 
larly. The interpretation suggested was that the 10 chromo- 
somes from D. rotundifolia paired with 10 of those from D. 
anglica, the other 10 having no homologues and remaining 
unpaired. (Shimamura later [1941] reported identical behavior 
in Japanese material of these three plants.) Recounting these 
and other observations of Rosenberg, Winge wrote (p. 244): 
"I could therefore imagine that D. longifolia [D. anglica] was 
produced by hybridization between two species or forms with 
x = 10, the chromosomes of which had become added together 
in the zygote, and then suffered division. On further crossing 
with D. rotundifolia only half would then be capable of conjuga- 
tion with those of the latter." 


1955] Wood, Jr.,—' The Hybrid Origin of Drosera anglica — 107 


Although it now appears that Drosera anglica is indeed an 
amphiploid of which D. rotundifolia is one of the progenitors, 
it has been pointed out by various authors (e.g., Sharp, 1934; 
Darlington, 1937; Dobzhansky, 1941) that in this particular 
example Rosenberg’s interpretation of chromosomal behavior 
is open to doubt, it being impossible to determine whether this 
is a case of allosyndesis, as suggested, with pairing only between 
“rotundifolia” and “anglica” chromosomes or whether the 20 
“anglica? chromosomes are forming 10 pairs with each other 
or whether some intermediate degree of autosyndesis is present 
with some “rotundifolia”? chromosomes and some “anglica” 
chromosomes pairing with other members of their own sets. 
On the basis of the behavior of the chromosomes in the sterile 
D. linearis X D. rotundifolia I should also wish to re-examine 
this interpretation, as well as to question the regular formation 
of 20 bivalents in D. anglica. It is to be noted, however, that 
Winge’s suggestion based on Rosenberg’s interpretation fits 
in with all of the other evidence which suggests a hybrid origin 
for Drosera anglica. 


A STERILE DIPLOID HYBRID 


In August, 1951, during the course of a field trip from the 
University of Michigan Biological Station to beach pools along 
the shore of Lake Huron at Hammond Bay, Presque Isle County, 
near the tip of the southern peninsula of Michigan, a clump of a 
Drosera with obovate-spatulate leaves was collected by Mr. 
Clare B. Kenaga from the midst of a large colony of Drosera 
linearis. In leaf-shape this plant resembled Drosera intermedia, 
a widespread species of wet and very acid habitats, but the 
leaves were much larger, the color a yellowish green with bright 
red tentacles, in contrast to the rather dull, dark red of D. 
intermedia in northern Michigan, and the plants were quite 
sterile, the ovaries containing only aborted ovules. The sterility 
of the plants, the leaf shape, and the occurrence of D. linearis 
in the pools and D. rotundifolia on hummocks nearby suggested, 
instead, a hybrid between these two species. Two summers 
later when I was again at the Biological Station, I had the 
opportunity of revisiting Hammond Bay and of studying other 
areas along the northern shores of the southern peninsula of 


108 Rhodora [Vor. 57 


20 


| 
Myr m 


24 


21 22 23 

FIGURES 1-24. Leaves of Drosera. Tracings from herbarium specimens, tentacles 
and leaf-bases omitted. X 4. 1. Drosera rotundifolia, Hammond Bay, Mich., 
Wood 8236, et al. (au). 2. D. linearis, Hammond Bay, Mich., Wood 8075, 8316 (an). 
3. Sterile hybrid (D. linearis X D. rotundifolia), Hammond Bay, Mich., Wood 8315, 
Wood 8327, et al. (au). 4. Sterile hybrid, Lake Orion, Mich., Sutton 440 (MICH). 
5. Sterile hybrid, Raspberry Island, Isle Royale, Mich., McFarlin 2238 (MicH). 
6. Sterile hybrid, Lake Orion, Mich., Billington 66 (micH). 7. Sterile hybrid, Lake 
Orion, Mich., Billington 66 (Ny). 8. Sterile hybrid, Emmet Co., Mich., Wood 8122 
(aH). 9. Sterile hybrid, Presque Isle, Mich., Wood 8307 & McVaugh (au). 10. 
Fertile hybrid (D. linearis X D. rotundifolia Amphiploid), Presque Isle, Mich., 
Wood 8308 & McVaugh (aH). 11. D. anglica, Keeweenaw Co., Mich., Hermann 
8208 (Ny). 12. D. anglica, Ireland, Prager, 1896 (au). 13. D. anglica, Gaspé Co., 
Quebec, Collins, Fernald & Pease (Pl. Exsic. Gray. 218) (NY). 14. D.anglica, Sweden, 
H. Svenonius, 1932 (wv). 15. D. anglica, Japan, Uno 21043 (au). 16. D. anglica, 
Marquette Co., Mich., Dachnowski, 1906 (wicm). 17. D. anglica, Glacier Park, 


1955] Wood, Jr.,— The Hybrid Origin of Drosera anglica — 109 


Michigan. The sterile plants are of sporadic occurrence around 
the northern Great Lakes in company with Drosera linearis and 
D. rotundifolia and are indeed hybrids between these two species. 

MonrnoLocy. ‘The parental species, the common and wide- 
spread Drosera rotundifolia! and the sometimes abundant but 
very much more local Drosera linearis, differ from one another 


D. ROTUNDIFOLIA 


STERILE HYBRID 


D. LINEARIS 


Leaves spreading or horizontal. 


Petioles compressed and winged, 
moderately to densely hairy 
with flat, twisted hairs. 


Leaf blades suborbicular to 
orbicular-reniform, broader than 
long, (4-)6-10 mm. long, (5-) 
7-11 mm. wide, bright yellowish 
green, the tentacles bright red. 


Leaf base slightly winged, the 
stipules adnate 2-3 mm. above 
the base, of fimbriate segments. 


Inflorescences (4-)7-16 (-21) cm. 
long, the peduncle (3.2-)6-15 
cm. long. 


Flowers per inflorescence (3-) 
4-13(-15). 


Calyx green; petals white; 
pollen white. 


Capsules oblanceolate-elliptic in 
outline, 4.5-5.5 mm. long, 2.5 
mm. diameter. 


Seeds light brown, shiny with a 
metallic luster, sigmoid-fusi- 
form, finely longitudinally stri- 
ate, 1-1.5 mm. long. 


Leaves ascending or spreading. 


Petioles somewhat compressed 
(intermediate), sparsely to mod- 
erately hairy. 


Leaf blades obovate-spatulate to 
elongate-spatulate, longer than 
broad, 11-17(-20) mm. long, 
4-6(-7) mm. wide, bright yel- 
lowish green, the  tentacles 
bright red. 


Leaf base with a narrow wing- 
like margin and numerous 
fimbriate, crinkled segments. 


Inflorescences | (2.5-)6-14(-19) 
em. long, the peduncle (2.3-) 
4-13.5 cm. long. 


Flowers (2-)4-8(-11). 

Calyx blackish; petals white, 
pink tinged without; pollen 
orange. 


Ovaries fail to develop into fruit. 


Seeds fail to develop. Ovules 
aborting. 


Leaves ascending or erect. 


Petioles terete, glabrous or with 
a few scattered hairs. 


Leaf blades exactly linear, the 
tip rounded, 20-50 mm. long, 
2-2.5 mm. wide, dull green or 
reddish, the tentacles dull red. 


Leaf base dilate-winged, the 
stipules adnate 2-4 mm. above 
the base, of numerous crinkled, 
fimbriate, setaceous segments. 


Inflorescences (3-)5-11(-12) cm. 
long, the peduncle (2.7-) 
4-8(-8.7) cm. long. 


Flowers (1-)4-6(-7). 


Calyx blackish; petals white, 
tinged pink at tips without; 
pollen orange. 


Capsules elliptic in outline, 
5-5.5 mm. long, 2.5-3.5 mm. 
wide. 


Seeds shining black, oblong- 
obovate in outline or rhom- 
boidal, minutely and densely 
pebbled, 0.5-0.8 mm. long. 


! See Wynne (1944) for range-maps of the North American species of Drosera. 


Montana, Mains 6053 (MICH). 
(NY). 19. D. anglica, Alaska, Anderson 764 (NY). 
D. rotundifolia), Sweden, Bergström, 1915 (em). 


Japan, Suzuki 45 (GĦ). 


18. D. anglica, Kauai, Hawaiian Ids., Rock, 1909 
20. D. x obovata (D. anglica x 
21. D. x obovata, Ozegahara, 
22. D. X obovata, Finland, Hallström 1202 (wv). 23. D. 


x obovata, Sweden, Asplund, 1923 (wv). 24. D. X obovata, Newfoundland, 
Fernald, Long & Fogg 1759 (au). 


110 Rhodora [Vor. 57 


in à number of conspicuous features, including habitat and 
habit, leaf-shape, cross-sectional shape and pubescence of 
petioles, stipules, the number of flowers per inflorescence, color 
of anthers and color, shape and surface-markings of seeds. 
There are also other less conspicuous differences. In most 
respects the sterile, spatulate-leaved plants from Hammond 
Bay are morphologically intermediate between the two species, 
but in a few characteristics resemble one or the other species 
more strongly. In pollen color and color of the calyx the 


27 


?6 


FIGURES 25-32.  Cross-sections of petioles of Drosera. X 20. Cross-sectional 
shape of petioles offers interesting possibilities as an additional distinguishing character- 
istic in fresh material of Drosera, as will be noted from the five species and one 
hybrid shown here. Fresh specimens or material preserved in liquid must be 
used, however, for the tissues shrink so badly in drying that it is difficult to re-expand 
petioles from herbarium specimens. Vascular bundles are shown diagrammatically, 
the xylem in black, the phloem in white. 

25. Drosera rotundifolia, middle of petiole; Emmet Co., Mich., Wood 8076 (anm). 
26. D. linearis X D. rotundifolia, middle of petiole; Hammond Bay, Mich., Wood 
8237 et al. (au). 27. D. brevifolia, middle of petiole; Carteret Co., N. C., Wood 
8237 (aH). 28. D. capillaris, middle of petiole; Pender Co., N. C., Wood 8236 (an). 
29. D. rotundifolia, petiole near the base of the blade. 30. D. linearis, middle of 
petiole; Emmet Co., Mich., Wood 8120 (an). 31. D. capillaris, petiole near base 
of the blade; compare with fig. 29. 32. D. intermedia, middle of petiole; Cheboygan 
Co., Mich, Wood 8130 (ax). 


plants are similar to D. linearis but in the yellow-green of the 
leaf blades and the bright red of the tentacles and in the elongate 
shape of the aborted ovules they approach D. rotundifolia. 
A comparison of all three plants is given in the accompanying 
table? from which the intermediacy of the sterile plants is, I 
believe, evident. (See also fig. 1-9, 25, 26, 30.) 

The only other species which could be involved as a parent 
of the hybrid is Drosera intermedia Hayne, which it superficially 

2 Since there is abundant evidence of ecological variation in Drosera, the com- 
parisons drawn in the table are based upon material from northern Michigan, rather 


than upon specimens from throughout the range of the parental species, so that a 
more uniform sample is obtained for this purpose. 


1955] Wood, Jr.,—' The Hybrid Origin of Drosera anglica — 111 


resembles in leaf-shape. This species in northern Michigan, 
however, is a plant of very acid and wet habitats, particularly 
Sphagnum bogs where it occupies the wettest depressions in the 
Sphagnum mat. It occasionally crosses with D. rotundifolia, 
which grows on the drier Sphagnum, producing a sterile hybrid 
with much more rotund leaves than the sterile plant of the beach 
pools. I have never found Drosera intermedia growing anywhere 
in the vicinity of D. linearis. Drosera intermedia is also ruled 
out by its completely glabrous petioles, D-shaped in cross- 
section (compare figs. 25, 26, 30 & 32), small, spatulate leaf 
blades, free stipules and inflorescences which emerge horizontally 
before curving upward. 

On morphological grounds the sterile plant of the Hammond 
Bay beach pools is certainly a hybrid between Drosera linearis 
and D. rotundifolia. It is now known from a number of other 
localities and the description given below is drawn from all of 
these collections. Since I believe that the amphiploid of this 
hybrid is Drosera anglica, nothing but confusion would result 
from a “specific” name for this putative hybrid. It is accord- 
ingly designated only by formula. 


Drosera linearis Goldie X D. rotundifolia L. 


Very short-stemmed rosette plant intermediate between the parental 
species and occurring with them. Principal leaves ascending or spreading, 
(2-)2.5-6(-8) em. long, the petioles (1.3-)2-4(-5.5) em. long, slightly 
compressed and winged, sparsely to moderately hairy with crinkled or 
twisted white hairs to 2 mm. long; base of petiole flattened, with a narrow 
wing-like margin, the stipules pink, adnate 2-4 mm. above the base, 
with several flat fimbriate segments and few to many twisted flat hairs 
to 5mm. long. Leaf blades obovate-spatulate to oblanceolate-spatulate, 
the apex rounded, 11-23(-30) mm. long, 4-6(-8) mm. wide, bright 
yellowish-green, the tentacles bright red (as in D. rotundifolia), to 5 mm. 
long. Inflorescences 1-3(-4), seapose, including the peduncle (2.5-) 
6-14(—20) em. tall, the first usually the largest; peduncle (2.3-)4-13(-15) 
em. long, glabrous or with scattered hairs near the base. Flowers (2-) 
4-8(-11) per inflorescence, the pedicels 1-2.5 mm. long. Calyx dark 
blackish-green (as in D. linearis), 4-5 mm. long, the lobes oblong, obtuse 
to acute, 3-3.5 mm. long, 1-1.5 mm. wide, the margins glandular-denticu- 
late. Petals white, obovate-cuneate or narrowly obovate, 4—4.5 mm. 
long, 2.5 mm. wide. Stamens 3.5-4 mm. long, the pollen orange (as in 
D. linearis). Ovary failing to develop into a fruit; ovules fusiform, 
aborting; styles slender, clavate, bifurcate to’ the base, the branches 
1.5-2 mm. long. Somatic chromosomes 20; meiosis irregular. 


112 Rhodora [Vor. 57 


SPECIMENS EXAMINED.’ Canada. Ontario. Bruce Co.: edges of pools 
in marly bog-marsh area near Lake Huron, Oliphant, Annabel Twp., Cain, 
Hagenah & Thompson, 6 July 1947 (BLH); grassy bog, Sauble Beach, Arotkov 
9099 (us). , 

United States. Michigan. [Charlevoix Co.:] border of a (marl) lake near 
Charlevoix, C. F. Wheeler, 26 Aug. 1894 (au). Emmet Co.: marly flats with 
scattered Thuja and Larix behind dunes along Sturgeon Bay, Sec. 5, R5W, 
T38N, Bliss Twp., Wood 8122 (aH). Keeweenaw Co.: transition zone 
between sedges and sphagnum, Raspbery Island bog, Isle Royale, R. M. 
Linn, 9 & 15 Aug. 1954 (aH); bog on Raspberry Island, Rock Harbor, Isle 
Royale, J. B. McFarlin 2238, 22 July 1930 (micu); Isle Royale, University 
Party, 1868 (MicH) (probably this plant—one plant with a plant of D. inter- 
media). Oakland Co.: [all of the following are from the same locality]: 
marsh, edge of Lake Orion, B. F. Chandler, 30 July 1914 (BLH, vs); boggy 
ground, Marl Lake, near Orion, Chandler, 19 Sept. 1915 (MICH); wet open 
ground (boggy ground) on the margin of Marl Lake, Orion Twp., Billington, 
9 & 13 July 1916 (Ny), 9 July & 13 Aug. 1916 (wrcn), 9 July 1916 (wrcn); 
borders of Marl Lake near Orion, O. A. Farwell 4289, 9 July 1916 (BLH, GH), 
428914 (au), 4371, 13 Aug. 1916 (GH); Marl Lake, north of Orion, Sutton 
440, 30 July 1914 (urcH); Marl Lake, near Rochester, Ballard, July 1897 
(anm). Presque Isle Co.: on edge of open beach pool at Hammond Bay, Sec. 
22, R3E, T36N, Kenaga 14, 6 Aug. 1951 (umBs); boggy beach pool, Evans 
property, Hammond Bay near old Coast Guard Station, F. K. & A. G. Sparrow, 
19 July 1952 (uwBs); open sand and small hummocks along seeping edges 
of shore and sandy, marly beach pools in front of Thujas along low shore 
of Hammond Bay, Lake Huron, w. of Coast Guard Station, Sec. 22-23, 
R3E, T36N, Ocqueoe Twp., Wood 8077, 23 June 1953, 8315, 22 Aug. 1953, 
Wood 8237, E. Clover & M. Fulford, 24 July 1953 (aH, MICH, NY, UMBS, US); 
rare, with D. rotundifolia and D. linearis, both of which are abundant, marly 
flats back of Lake Huron beach, small bay, ca. 3 mi. n. of Presque Isle P. O., 
between Sec. 21 & 27, R8E, T34N, McVaugh 10892, 11 July 1949 (wicH), 
Wood 8807 & McVaugh, 22 Aug. 1953 (an). 


Ecotoey. In northern Michigan Drosera rotundifolia and 
D. linearis usually occupy very different habitats. Drosera 
rotundifolia is characteristically a plant of Sphagnum bogs but 
it also occurs on other mosses, on wet, rotting logs, or on moist 
or wet sands, usually in acid habitats with a pH in the neighbor- 
hood of 4 or 5. The very different D. linearis, in contrast, is 
always (insofar as I can determine) a plant of constantly wet, 
open, circumneutral (pH 6.5-7.5) or slightly alkaline habitats, 
such as marl bogs and the seeping beach flats of the northern 
Great Lakes, especially Lake Huron. In these areas the plant 
often occurs by the thousands, literally covering the ground. 
Although the respective habitats of the two species are, for the 

3'The abbreviations used to designate herbaria are those of Lanjouw and Stafleu 
in Index Herbariorum, Pt. I., The Herbaria of the World, ed. 2, Utrecht, 1954, with 


the exception of the University of Michigan Biological Station, Cheboygan, Michigan 
(UMBS). 


1955] Wood, Jr.,— The Hybrid Origin of Drosera anglica — 113 


most part, quite separate, at some points they come together 
and it is in these areas that hybrids have been found. 

A description of one such area, that at Hammond Bay, Presque 
Isle County, Michigan, is given below. At this locality, back 
from the gently curving shore of Hammond Bay south of 
the Ocqueoc River, the land drops abruptly from the level of 
the fossil beaches of glacial Lake Algonquin to a Thuja bog 
which stretches toward Lake Huron. At the edge of the bog, 
some 40—50 yards from the lake shore, is a low, stabilized dune 
with a dirt road running along the top to several summer cottages. 
On the shoreward side of the road the sand slopes gently to 
a wet depression, the inner edge of a beach pool, a distance of 
some 25 feet. Characteristic plants of the stabilized upper 
portion of the slope are Pinus Strobus, Larix laricina, Thuja 
occidentalis, Juniperus horizontalis, Shepherdia canadensis, An- 
dropogon scoparius, Calamovilfa longifolia, Arctostaphylos Uva- 
ursi, Linnaea borealis var. americana, and Campanula rotundifolia. 
The lower portion of the slope, some 10 feet in width, is more 
open and is quite moist or even wet with constant seepage from 
the Thuja bog. Here are found scattered shrubs of Myrica 
Gale with Sarracenia purpurea, Primula intercedens, Lobelia 
Kalmii and numerous plants of Drosera rotundifolia. In the 
very shallow water of the wet depression, 8 feet wide at the 
foot of the slope, occur Rhynchospora capillacea, Pogonia ophio- 
glossoides, Sarracenia purpurea, Utricularia cornuta, U. inter- 
media, many plants of Drosera linearis and scattered individuals 
of the obovate-spatulate-leaved hybrids. A low, somewhat 
irregular ridge, from 6 inches to a foot high and from two to 
five feet wide, rises from the wet depression paralleling the 
shore. Beyond this the shallow beach pool, broken by hum- 
mocks, reaches about 30 feet to end abruptly at a wide, sandy 
beach which slopes gradually into Lake Huron. The low, 
irregular ridge provides conditions suitable for the growth of 
both Drosera linearis and D. rotundifolia and it is here that the 
greatest numbers of hybrid plants (75 or more individual plants 
or clumps of various sizes) were found. Also on this ridge were 
a few small Thujas and an occasional plant of Myrica Gale, 
along with Cladium mariscoides, Scirpus hudsonianus, Lilium 
philadelphicum, Pogonia ophioglossoides, Sarracenia purpurea, 


114 Rhodora (Vou. 57 


Gentiana procera, Lobelia Kalmii and Solidago ohionis. The 
beach pool proper is dotted with scattered plants of Sarracenia 
purpurea, but none of the sundews grows there. The con- 
spicuous plants of the beach pool are Cladium, Rhynchospora 
and the two species of Utricularia. The sandy beach is occupied 
by the common shore plants of the region, including Equisetum 
variegatum, Calamagrostis inexpansa, Scirpus americanus, Juncus 
balticus var. littoralis, Triglochin palustris, Salix interior, Cakile 
edentula var. lacustris, Parnassia glauca, Hypericum Kalmianum 
and Lobelia Kalmii, the distribution of these seemingly dependent 
upon the varying wetnesses of the beach. 

The principal area in which the hybrid sundews are found 
parallels the shore for about 250 feet. Beyond this on either 
side conditions change and the two parental species are separated 
again, although at another area a quarter of a mile away similar 
conditions obtain and here again hybrids are found. 

In each locality in northern Michigan where I have collected 
the hybrids ecological conditions similar to those described 
above occur. A seeping shore with open, marly flats and scat- 
tered Thujas or larches with hummocks at their bases are in- 
dications that hybrids may be looked for (but not always found, 
of course). Drosera linearis will be confined to the constantly 
wet soils or very shallow water, while D. rotundifolia may occur 
on the drier hummocks or at the bases of the trees. Hybrids 
have been found growing with either parent; at one locality 
(Emmet Co.), after a great deal of search I found only two 
hybrids, one in the midst of a colony of D. rotundifolia, the other 
among plants of D. linearis some distance away. I assume 
that the cross has occurred in both directions. Generally, 
however, the hybrid seems to be best developed in the wetter 
locations which favor Drosera linearis. 

The hybrid was formerly abundant at Lake Orion, in southern 
Michigan, where both Drosera rotundifolia and D. linearis (the 
latter here near its southern limit) occurred. Conditions there 
seem to have been very favorable for its growth, for many of the 
specimens from this locality are large and vigorous, including 
one, the largest I have seen, with leaves 8 cm. long and an 11- 
flowered inflorescence 20 cm. tall. Accompanied by Dr. Rogers 
MeVaugh and Dr. W. H. Wagner, Jr., I visited Lake Orion in 


1955] Wood, Jr.,— The Hybrid Origin of Drosera anglica 115 


August of 1953 in search of the plant, only to find that the 
lake level has been raised several feet artificially, apparently 
eliminating Drosera linearis and the hybrid quite effectively. 
Drosera rotundifolia still survives on sphagnum hummocks 
along the shrubby border at several places. 

CyroLoav. While, on the basis of occurrence and morphology 
alone, the spatulate-leaved plants may be presumed to be 
hybrids, high sterility is an additional indication of their hybrid 
nature. In an attempt to determine possible cytological causes 
for this sterility meiosis was studied in both parental species 
and in the hybrid from material collected at Hammond Bay 
on June 23, 1953. Mr. Robert Linn has also sent buds from 
plants which prove to be this hybrid growing in a bog on Rasp- 
berry Island, Isle Royale, Michigan. Various other collections 
of Drosera rotundifolia and D. linearis, as well as of D. intermedia, 
from northern Michigan have been studied. In the course 
of this investigation it has subsequently been possible to examine 
meiotie material of Drosera brevifolia, D. capillaris, D. filiformis 
and D. filiformis var. Tracyi from the southeastern United States, 
so that I have had the opportunity of studying all of the species 
occurring in North America, with the exception of one of the 
most crucial, the tetraploid Drosera anglica. The chromosome 
numbers now known in the genus are given in the accompanying 
table. 

Cytological material from northern Michigan was fixed in a 
modified Carnoy’s fixative (4 chloroform: 3 alcohol: 1 acetic 
acid) and that from the Southeast in Newcomer’s fixative 
(1953). Both gave excellent results. Anthers were squashed 
in propriono-carmine, drawings made with the aid of a camera 
lucida, and the slides made permanent by Bradley’s vapor 
transfer technique (Bradley, 1948). 

In all material of Drosera linearis, D. rotundifolia and D. 
intermedia examined ten bivalents are formed in the first division 
of meiosis. One pair of chromosomes is associated with the 
nucleolus in each of these species. (This is also true of D. 
linearis X D. rotundifolia and the various other diploid species 
reported here.) Chromosomes pair and disjoin with no irregu- 
larities (fig. 33-36). In the sterile hybrid, although in many 
instances the 20 chromosomes seem to be at least loosely asso- 


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117 


Wood, Jr.,—The Hybrid Origin of Drosera anglica 


1955] 


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118 Rhodora [Vor. 57 


1955] Wood, Jr.,—' The Hybrid Origin of Drosera anglica — 119 


ciated into 10 pairs in Prophase I, some pairs either fall apart 
or fail to move onto the spindle so that at Metaphase I only 
3-7 bivalents are present. (fig. 37-45). One, two or three 
trivalents may occur (fig. 41—44) and an occasional quadrivalent 
is seen (fig. 45). 

For example, of 28 metaphases (the largest number analyzable 
from any one hybrid plant), 3 pairs of chromosomes were present 
on the spindle in 3 cells, 4 pairs in 6 cells, 5 pairs in 11 cells, 6 
in 5 cells and 7 in 3 cells. Twelve cells showed one trivalent 
and five had two. Only a single quadrivalent was seen in this 
plant (fig. 45). The number of unpaired chromosomes varied 
from 2 to 10 and the number of bivalents not on the spindle 
from one to four. Figure 41 shows a cell with three bivalents 
and one trivalent on the spindle, while four bivalents lie off the 
spindle. 

Chromosomal distribution is consequently quite erratic. Some 
chromosomes lag and divide tardily and there is evidence of 
bridges and fragments (fig. 46-48). Following meiosis con- 
joined nuclei, micronuclei and other irregularities are often 
seen. It is not surprising that the hybrids of Drosera linearis 
and D. rotundifolia are sterile. 

In the Droseraceae the four microspores from each pollen 
mother cell remain associated and develop into a united tetrad 
of pollen grains, generally tetrahedral in arrangement (fig. 49—50). 
It is thus possible to see the products of each pollen mother 
cell and the extent to which meiotic irregularities result in the 
formation of abnormal microspores or pollen grains. Fresh 
Drosera pollen mounted in lactophenol with a little cotton blue 
absorbs the stain readily making it fairly easy to determine 


FIGURES 33—53.  Meiosis in Drosera. x 800. 33. Drosera linearis, metaphase I; 
Hammond Bay, Mich., Wood 8075. 34. D. rotundifolia, metaphase I; Hammond 
Bay, Mich., Wood 8076. 35. D. capillaris, metaphase I, Pender Co., N. C., Wood, 
8236. 30. D. brevifolia, diakinesis; Carteret Co., N. C., Wood, 8237. 37-45. D. 
linearis X D. rotundifolia (2n, sterile), metaphase I, Hammond Bay, Mich., Wood 
8077, 8237. 46-47. D. linearis X D. rotundifolia (sterile), metaphase II, showing 
tardily disjoining bivalents and dividing univalents, Wood 8077, 8237. 48. D. linearis 
x D. rotundifolia (sterile), metaphase II, Wood 8077, 8237. 49. Three pollen 
tetrads from D. rotundifolia, Wood 8305 & McVaugh (au). 50. Three pollen tetrads 
from D. linearis, Wood 8075 (au). 51. Four pollen ''tetrads" from D. linearis x D. 
rotundifolia (sterile), Wood 8077, with 4, 5, 9 and 8 microspores, respectively. 52. 
Two pollen tetrads from fertile hybrid, D. linearis X D. rotundifolia Amphiploid, 
Wood 8308 & McVaugh. 53. Pollen ‘‘tetrad’’ with 5 grains from same fertile hybrid. 
Fig. 49—53, x 250. 


120 Rhodora [Vor. 57 


the number of associated pollen grains and something of the 
condition of the protoplasmic contents. 

In hybrids from Hammond Bay approximately half of the 
grains are associated in tetrads, although the four are sometimes 
of different sizes. The remaining 'tetrads" consist of from 
2 to 9 mierospores or grains of various sizes, in most of these 
(about 40 per cent of the total) consisting of 5 or 6 (fig. 51). 
In no instances were any grains with evenly distributed and 
staining protoplasmie contents observed in the hybrids. Most 
grains (even those united in regular tetrads) lack contents alto- 
gether and the remainder show only deeply stained and clumped 
protoplasm. Apparently pollen sterility approaches 100 per cent. 
This is in great contrast to both of the parental species which 
are very nearly as fertile as the hybrid is sterile. 

In spite of the lack of sexual reproduction, at Hammond Bay 
and presumably at Lake Orion the hybrid reproduced itself 
vegetatively by adventitious plantlets from leaf blades, scapes, 
or even flowers, the carpels of which may be converted into a 
rosette of leaves similar to that described by Fernald as Drosera 
rotundifolia var. comosa. This kind of vegetative reproduction 
was seen at Hammond Bay in July and August and was shown 
by experimentally detached leaves which produced adventitious 
plantlets from the adaxial surfaces of the blades. Such repro- 
duetive methods are shared (at least potentially) by most 
species of the genus.* Vegetative reproduction may thus be of 
considerable importance in allowing the persistence and even 
spread of a sterile plant over a period of years. It greatly 
inereases the chances for the formation of restitution nuclei 
or other phenomena which could lead to polyploidy and the 
produetion of a fertile amphiploid such as the plant to be dis- 
cussed next. 

A FERTILE HYBRID 


On July 11, 1949, Dr. Rogers McVaugh had collected two 
plants of the sterile hybrid growing with abundant Drosera 


4 In his Flore Laurentienne (1935, p. 273), Marie-Victorin noted vegetative repro- 
duction in D. rotundifolia, saying, “At the maturation of the fruits, the leaves de- 
taching themselves are disseminated by water or wind and they are from this moment 
capable of giving rise to adventitious buds which develop into new plants. The 
process recalls the classic case of the budding of Bryophyllum."  [Translation.] 
See Behre (1929) for descriptions of modes of vegetative reproduction or Lloyd (1942) 
for a general review in English. 


1955] Wood, Jr.. —The Hybrid Origin of Drosera anglica 121 


linearis and D. rotundifolia on beach flats along the shore of 
Lake Huron about 3 miles north of Presque Isle Post Office, 
Presque Isle County, Michigan (McVaugh 10892). Since it 
was of considerable interest to obtain some idea of how wide- 
spread and frequent hybridization between these two species 
may be, Dr. McVaugh kindly showed me this locality on August 
22, 1953. "The ecological situation was similar to that in which 
hybrids have been found elsewhere, with marly flats becoming 
a cobble beach, a seeping shoreline along a Thuja bog and scat- 
tered plants of Thuja and Larix with ridges and hummocks of 
herbaceous vegetation. Drosera linearis was abundant on the 
wet flats and D. rotundifolia grew among the herbaceous plants 
and on mossy hummocks at the bases of Thuja or Larix. Both 
species of sundew were in mature fruit and at the point of 
shedding their seeds. Careful search produced only a single 
hybrid, a plant which had flowered but which was very definitely 
sterile. However, on the lower portion of a mossy hummock 
at the base of a small Thuja and in close proximity to both 
Drosera linearis and D. rotundifolia, we found a colony of ten 
or so plants of various sizes which were quite identical in their 
vegetative characteristics with both the sterile hybrid collected 
here and those seen elsewhere. Three of these plants, although 
rather small, had flowered and, unlike the other hybrids, had 
matured fruit and were shedding seeds. (Wood 8308 & McVaugh, 
GH.) 

These fruiting plants are quite remarkable. The capsule 
is well developed, oblong in outline, 7-7.5 mm. long and 3.5 mm. 
wide, thus larger than that of either Drosera linearis or D. 
rotundifolia in this area. The scapes were 6 to 11 cm. long 
and bore 3 to 7 flowers. It is the seeds, however, which are 
most notable. These are precisely what might be expected 
in a fertile hybrid between D. linearis and D. rotundifolia, with 
the black coloration of the seeds of the former, the spindle-shape 
and size of those of the latter and surface markings which are 
quite intermediate. Most remarkable of all, they are a perfect 
match for the seeds of the cireumpolar tetraploid Drosera anglica, 
the ‘only species of the northern hemisphere, other than D. 
rotundifolia, with fusiform seeds. This, in a group in which 
seed characteristics have proved to be among the most distinctive 


122 Rhodora [Vor. 57 


and stable, can not be regarded as mere coincidence. (See 
Wynne, 1944, text and figures, and the same illustrations in 
The New Britton & Brown.) Figures 60-63 show a comparison 
of the cells from the outer seed coats of Drosera rotundifolia, 
D. linearis, the fertile plants from Presque Isle and D. anglica 
from Montana. 

Although this small colony of exceptional fertile plants might 
conceivably represent a recent introduction of Drosera anglica 
from some locality to the north (the nearest in Michigan in 
the upper peninsula at Marquette, some 180 miles west-north- 
westward),? the circumstances of its occurrence suggest that 
these are plants of D. linearis—D. rotundifolia parentage which 
have somehow acquired fertility. In any event, the identical 
vegetative characteristics of these fertile plants which produce 
“anglica”? seeds and of the sterile offspring of D. rotundifolia 
and D. linearis need explanation, for some tie-up with Drosera 
anglica is indicated, whether this is an introduction to the area 
or whether it is a spontaneous development. I can see no reason 
to think that this is anything other than a fertile hybrid. 

Assuming that these are indeed derived from D. linearis and 
D. rotundifolia, the very production of seeds by the fertile 
plants, in contrast to the sterile hybrids with their erratic 
chromosomal behavior and inviable pollen, suggest that these 
exceptional plants may be amphiploids on the same chromosomal 
level as Drosera anglica, a tetraploid with 20 pairs of chromo- 
somes. 

It was not possible to determine directly the chromosome 
number of the fertile plants, but this was attempted by a com- 
parison of the areas and volumes of epidermal guard cells on 
the principle that the cells of a recent amphiploid, having twice 
the number of chromosomes of a diploid should have twice the 
nuclear volume and roughly twice the cellular volume. Leaves 
preserved in 70 per cent alcohol were available from Presque 
Isle plants of Drosera rotundifolia, D. linearis, the sterile hybrid 
and the fertile hybrid, but boiled-up herbarium material had 
to be used for Drosera anglica. Whole leaves were mounted 

5 I have no evidence of Drosera anglica in the lower peninsula of Michigan, although 
Wynne (1944) shows two localities on her map. One of these is certainly Lake Orion 


and represents sterile D. linearis X D. rotundifolia and I suspect that the other, 
farther north in the part of the state with which I am most familiar, does also. 


1955] Wood, Jr.,—' The Hybrid Origin of Drosera anglica 123 


in water, the epidermis on the backs of the leaves studied, 
and pairs of guard cells (chosen at random from the central 
portion of the blade) drawn with camera lucida at a final magni- 
fication of 1100 diameters. The guard cells of the two diploid 
species and their sterile hybrid are all smaller than those of 
either the fertile hybrid or D. anglica (fig. 54—59). Areas of 


lit | 


FIGURES 54—59. Epidermis from lower side of leaf blades of Drosera. X 125. 
54. Drosera rotundifolia, Presque Isle, Mich., Wood 8305 & McVaugh. 55. D. 
linearis, Presque Isle, Mich., Wood 8306 & McVaugh. 56. D. linearis X D. rotundi- 
folia (sterile), Presque Isle, Mich., Wood 8307 & McVaugh. 57. D. linearis X D. 
rotundifolia (fertile), Presque Isle, Mich., Wood 8308 & McVaugh 58. D. anglica, 
Sweden, Svenonius, 1932 (Ny). 59. D. anglica, Montana, Mains 6053 (MICH). 

Fiaures 60—63. Portions of outer seed coats responsible for surface sculpturing 
of seeds of Drosera. x 75. 60. D. rotundifolia, Wood 8305 & McVaugh. 61. D. 
linearis, Wood 8306 & McVaugh. 62. D. linearis X D. rotundifolia (fertile), Wood 
8308 & McVaugh. 63. D. anglica, Mains 6053. 


the individual cells of each of the plants were determined by 
means of a planimeter, the average of five circumferences being 
used for each cell. In attempting to obtain a rough approxima- 
tion of the volumes, the diameter of each guard cell was used 
as its depth and the volume calculated as though the guard 
cell were a flat-topped, straight-sided figure. Areas and volumes 
were calculated only for purposes of comparison; no attempt 
was made to convert these to the actual volumes of the cells. 
The results are given in the adjoining table. 


124 Rhodora [Vor. 57 


A statistically significant difference exists between the mean 
areas of the guard cells of D. linearis and D. rotundifolia. Their 
sterile hybrid lies midway between, as might be expected. The 
mean area of the guard cells of the fertile hybrid is 1.5 times 
that of the sterile hybrid. There is no significant difference 
between the means of the fertile hybrid and those of D. anglica 
from either Montana or Sweden. 

From the volume relationships it is quite evident that two 
distinet groups of plants, corresponding to the known diploids 
and tetraploids, are represented. The fertile plant from Presque 
Isle fits in with the latter group and the mean volume of its 
guard cells is 1.9 times that of the sterile plant. No statistically 
significant difference exists between the means of the volumes 
of Drosera rotundifolia, D. linearis, or their sterile hybrid, or 
between those of Drosera anglica from Europe, that from America 
and the fertile hybrid, but the difference between the two groups 
is highly significant and not due to chance. 

Since in this genus the petals are persistent, shriveling together 
above the capsule and protecting the dried stamens and pollen 
grains adhering to the stigmas, it is often possible to recover 
pollen tetrads from plants with mature fruits. While the 
shape and confluence of the grains makes the determination 
of the volumes impossible, those of the fertile hybrid are con- 
spicuously larger than those of either putative parental species 
or the sterile hybrid. (Compare fig. 52-53 with fig. 49—51.) 
It may be remarked in this connection that Erdtmann (1943) 
gives the width of a tetrad of Drosera anglica as 72 microns, 
the equatorial diameter of a single grain about 50 microns, and 
the width of a tetrad of D. rotundifolia as about 55 microns, 
the diameter of a single grain about 34 microns. I have not 
followed this line of evidence further. 

Although these indications are not as precise as is desirable, 
it seems highly probable, on the basis of its fertility, guard cell 
area and volume, and pollen size, that the fertile hybrid from 
Presque Isle is at the tetraploid or near tetraploid level, repre- 
senting a spontaneous amphiploid of Drosera linearis and D. 
rotundifolia. 

Such a plant might be expected to be fairly fertile and could 
propagate itself sexually, setting seeds by self-pollination. The 


Wood, Jr.,—The Hybrid Origin of Drosera anglica 125 


1955] 


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126 Rhodora [Vor. 57 


flowers of Drosera are usually open for cross pollination only 
in full sunlight from morning to mid-day or early afternoon, 
after which the petals close bringing the anthers into contact 
with the stigmas, insuring pollination. A number of unde- 
veloped ovules were noted among the seeds of the fertile Presque 
Isle hybrids; these might indicate either a degree of sterility or 
merely lack of fertilization of some ovules. Of 70 pollen *'tet- 
rads” recovered from these plants, 69 had four grains and a 
single one five (fig. 52, 53). It was not, of course, possible to 
determine whether or not the contents had been functional. 


DROSERA ANGLICA 


There is no question of the close morphological similarity 
of the fertile and sterile hybrids. Lacking the flowering scapes 
or without a compound microscope it would be quite impossible 
to distinguish between them. It is also quite impossible to 
distinguish between the distinctive seeds of the fertile plant 
and those of Drosera anglica. If the fertile plant is Drosera 
anglica at a station considerably outside of its known range, 
the identical features of this plant and the sterile hybrid indicate 
that D. anglica was derived from the sterile hybrid of D. linearis 
and D. rotundifolia by amphiploidy. If the fertile plant is 
regarded as a spontaneous amphiploid or near amphiploid 
derived from the sterile hybrid it is still tied to D. anglica in 
the same way by its seeds and other morphological features. 
The indications are that the latter is the case and that D. anglica 
must have first originated in this same manner at some unknown 
time in the past. "This should be regarded, then, as an instance 
of the spontaneous resynthesis of a widespread and successful 
species. 

The morphological similarities of the sterile and fertile hybrids 
to Drosera anglica are striking. Fernald and Wynne both 
identified the sterile hybrid as D. anglica and a specimen of the 
sterile plant from Lake Orion is illustrated as D. anglica in The 
New Britton & Brown (Gleason, 1952). In studying specimens 
of D. anglica I unhesitatingly identified a series of plants in 
bud from a bog on Raspberry Island, Isle Royale, Michigan, 
(McFarlin 2238) (fig. 5) as this species. However, when Mr. 
Robert Linn sent me cytological material from seemingly iden- 


1955] Wood, Jr.,— The Hybrid Origin of Drosera anglica 127 


tical plants from the same bog the plant proved to be the sterile 
diploid hybrid! 

In all of the characteristics which I have been able to compare, 
the hybrids and Drosera anglica match well, with the exception 
of the leaves of many specimens. The hybrids from the lower 
peninsula of Michigan tend to have obovate-spatulate leaves, 
(fig. 3-9), in contrast to the narrower linear-spatulate or very 
narrowly obovate-cuneate leaves of Drosera anglica over much 
of its range (fig. 11-19). Some leaves are a perfect match, 
(e.g., compare fig. 5 and the leaves at either end of fig. 3 with 
fig. 11-19), but most leaves of the hybrid resemble a great deal 
more those of Drosera X obovata, the sterile triploid hybrid of 
D. anglica and D. rotundifolia (fig. 20-24). Several possibilities 
suggest themselves by way of explanation. Ecological differ- 
ences may well be responsible, at least in part. Shimamura 
(1941) who studied D. anglica, D. rotundifolia and D. X obovata 
in Japan found that, although tetraploid D. anglica and triploid 
D. X obovata were separable in the field when collected, they 
became morphologically indistinguishable when cultivated in 
Tokyo. It may also be noted that the specimens of D. X obovata 
figured by him are very narrow-leaved and would easily pass 
as many European or American specimens of D. anglica. (Com- 
pare fig. 21 with fig. 11-19.) "Thus the difference between the 
Isle Royale hybrids and those from farther south (fig. 5 vs. 
fig. 3-4, 6-9) may be merely an ecological difference. 

Another possibility which warrants careful investigation but 
which must, for the present, await further evidence is that 
Drosera anglica is not a genomic allopolyploid but a segmental 
allopolyploid which resembles D. linearis more strongly than 
it does D. rotundifolia. In view of the partial chromosomal 
pairing shown by the sterile hybrid it would seem quite likely 
that this type of polyploidy might come about in the course of 
stabilization of a newly formed allopolyploid derived from it. 

As was indicated in the introduction, the evidence from 
cytology for the origin of Drosera anglica is both scanty and in 
need of re-examination. The work of Rosenberg (1904, 1909) 
and Shimamura (1941) should be repeated with more modern 
cytological techniques. Additional evidence could come from 
the hybrid of D. anglica and D. linearis, if it can be found. 


128 Rhodora [Vor. 57 


(This plant should occur in nature where the two species grow 
in close proximity, as they sometimes do. It should be difficult 
to distinguish from its parents in the field except by its failure 
to set seeds.) If Drosera anglica originated as suggested, some 
multivalent formation and other chromosomal irregularities 
might be expected both in this species and in its hybrids with 
D. linearis and D. rotundifolia. Neither Rosenberg nor Shima- 
mura reported anything of the kind in either D. anglica or 
D. X obovata, perhaps because both used sectioned material 
in which multivalents are much more difficult to see than in 
smears and squashes. It is worth noting that in one paper 
Rosenberg (1904) illustrated a metaphase of D. anglica with a 
lagging univalent, although he later maintained that 20 pairs 
of chromosomes were regularly formed in D. anglica and 10 
pairs with 10 unpaired chromosomes in D. X obovata. 

As for this latter plant, the question of autosyndesis versus 
allosyndesis remains unsettled. The chromosomes of the two 
parental species (D. anglica and D. rotundifolia) are not morpho- 
logically distinguishable from one another and the chromosome 
number of the one parent is a straight multiple of the other. 
There is nothing to suggest, however, that D. rotundifolia and 
other species with 10 pairs of chromosomes (all of which have 
been referred to as diploids in this paper) are in reality polyploids 
with a basic chromosome number of 5, a situation which might 
well lead to autosyndesis in D. X obovata. The section Rossolis, 
to which all of the plants under discussion belong, seems to be 
a polyploid series with 10 as the base number of chromosomes. 
(See table of chromosome numbers.) The presence of only 
a single pair of chromosomes in association with the nucleolus 
in each of the 10-paired species points in the same direction, al- 
though this is not conclusive evidence (Stebbins, 1950, p. 464). 

In spite of the equivocal nature of the cytological evidence 
from the pairing relationships of the chromosomes of D. X 
obovata, the hypothesis based upon it (that D. anglica has derived 
half of its chromosomes from D. rotundifolia) is worth con- 
sidering and is highly suggestive, especially in view of the evi- 
dence from the fertile hybrid from Presque Isle County, Michi- 
gan. If it be assumed that half of the chromosomes of D. 
anglica have indeed come from D. rotundifolia, it becomes 


1955] Wood, Jr.,— The Hybrid Origin of Drosera anglica — 129 


necessary to find a second species with the proper combination 
of morphological and physiological characteristics to produce 
D. anglica. Of all of the species known from the northern 
hemisphere only D. linearis can supply these and only D. linearis 
is not ruled out by one or more conspicuous morphological 
features which would certainly be evident in a hybrid. (See 
especially Diels, 1906, for characteristics of other species.) 
Moreover, insofar as I can determine, both D. linearis and D. 
anglica are unique among northern hemisphere species in growing 
in marly habitats, although the latter plant appears to have 
a greater ecological amplitude and certainly is not confined to 
these. 

Thus, although the question must remain unsettled for the 
present and many more ramifications of the problem continue 
to develop, the evidence from the sterile Drosera linearis X 
D. rotundifolia, that from the fertile hybrid with its “anglica” 
seeds, the close morphological correspondences, the cytological 
evidence and the distributional data all indicate strongly that 
Drosera anglica is an amphiploid which originated through the 
hybridization of D. linearis and D. rotundifolia at some time 
in the past and which may continue to appear from time to time 
in the future. 

ACKNOWLEDGMENTS 

This work was begun at the University of Michigan Biological Station, 
continued at the University of North Carolina at Chapel Hill and brought 
to its present state at the Arnold Arboretum and the Gray Herbarium. 
At all of these places many people have kindly helped in one way or 
another with the problem and I am indebted to them all. Among those 
to whom I am very grateful are Elzada U. Clover, Margaret Fulford, 
Agnes Kugel, Edgar Anderson, Carter Broad, R. B. Channell, Delzie 
Demaree, F. C. Gates, Rogers McVaugh, Howard F. L. Rock, Reed C. 
Rollins, E. G. Voss, and W. H. Wagner, Jr. Mr. Robert M. Linn, of Duke 
University, went to great trouble, for which I am most appreciative, to 
obtain crucial cytological material from Isle Royale during the summer 
of 1954. During the course of this study I have had the opportunity 
of examining all of the specimens of Drosera at the Cranbrook Institute 
of Science, Duke University, the Gray Herbarium, the University of 
Michigan, the University of Michigan Biological Station, the New York 
Botanical Garden, the University of North Carolina, the University of 
Pennsylvania, and the University of Wisconsin. The curators of these 
herbaria have been most cooperative in allowing me to study material 
from the collections in their care.—THE ARNOLD ARBORETUM OF HARVARD 
UNIVERSITY. 


130 Rhodora [Vor. 57 


LITERATURE CITED 


BEHRE, K. 1929. Physiologische und Zytologische Untersuchungen über 
Drosera. Planta 7: 208-306. 

BRADLEY, MURIEL V. 1948. A method for making aceto-carmine squashes 
permanent without removal of the cover slip. Stain Tech. 23: 41-44. 

Daunt, A. O. 1937. Chromosome studies in sundew (Drosera). Bull. 
Marine Lab. Woods Hole 73: 368. 

DARLINGTON, C. D. 1937. Recent advances in cytology, ed. 2. Philadel- 
phia. 

DrkLs, L. 1906. Droseraceae in Engler, Das Pflanzenreich 4 (112): 1-136. 

Diasy, L. 1912. The cytology of Primula kewensis and of other related 
Primula hybrids. Ann. Bot. 26: 357-388. 

DoszHansky, TH. 1941. Genetics and the origin of species, ed. 2. New 
York. 

ERDTMANN, G. 1943. An introduction to pollen analysis. Waltham, Mass. 

GLEASON, H. A. 1952. The new Britton & Brown illustrated flora of the 
northeastern United States and adjacent Canada. New York. 

Hxrrz, E. 1926. Der Nachweis der Chromosomen: Vergleichende Studien 
über ihre Zahl, Grosse und Form im Pflanzenreich. I. Zeit. für Bot. 
18: 625-681. 

Levine, M. 1916. Somatic and reduction division in certain species of 
Drosera. Mem. N. Y. Bot. Gard. 6: 125-147. 

Luoyp, F. E. 1942. The carnivorous plants. Waltham, Mass. 

ManrE-VicroRIN, FRÈRE. 1935. Flore Laurentienne. Montreal. 

NaAKAJIMA. 1933. Japanese Jour. Genetics 9: 1 ff. [not seen.] 

NARASIMHACHAR, S. G. 1949. A contribution to the embryology of Drosera 
Burmannii Vahl. Proc. Ind. Acad. Sci. (Sec. B) 29: 98-104. 

Newcomer, E. H. 1953. A new cytological and histological fixing fluid. 
Science 118: 161. 

OwNBEY, M. 1950. Natural hybridization and amphiploidy in the genus 
Tragopogon. Amer. Jour. Bot. 37: 487-499. 

OwNsBEY, M. and G. D. McCotitum. 1954. The chromosomes of Trago- 
pogon. Rhodora 56: 7-21. 

RosENBERG, O. 1908. Das Verhalten der Chromosomen in einer hybriden 
Pflanze. Ber. der Deutsch. Bot. Gesellsch. 21: 110-119. PI. 7. 

1904. Uber die Tetradenteilung eines Drosera-Bastardes. Ber. 
der Deutsch. Bot. Gesellsch. 22: 47—53. Pl. 4. 
1909. Cytologische und morphologische studien an Drosera 

longifolia X rotundifolia. Kungl. Svenska Vetenskapsakademiens Handl. 
43 (11): 1-65, 4 pl. 

SHARP, L. W. 1934. Introduction to cytology, ed. 3. New York. 

SHIMAMURA, T. 1941. Cytological study of Drosera obovata Mert. & 
Koch with special reference to its hybridity. Bot. Mag. (Tokyo) 55: 
553-558. 

STEBBINS, G. L. 1950. Variation and evolution in plants. New York. 

Wince, O. 1917. The chromosomes, their numbers and general importance. 
Compt. Rend. Trav. Lab. Carlsberg. 13: 131-275. 

Wynne, Frances E. 1944. Drosera in eastern North America. Bull. 
Torrey Bot. Club 71: 166-174. 


1955] Steyermark,—Epipactis Helleborine in Illinois 131 


ErreAcTIS  HELLEBORINE IN ILLINOIs.—The last general 
account of this orchid appeared in Rhodora 53: 240—242, 1951, 
in which the records of its occurrence in Michigan were discussed 
by Drew and Giles. The concluding sentence in their article 
is “These facts suggest that this orchid may be reported from 
other adjacent mid-western states within the next few years." 
Illinois can now be added to the list. 

During the summer of 1954 while botanizing a steep, wooded 
slope along a creek in the Biltmore Estates subdivision north 
of Barrington, northeastern Illinois, I located eight plants of 
Epipactis Helleborine in bud. I made daily observations and on 
July 21 some of the plants began to bloom. They were found 
about half-way up a natural forested slope covered by Acer 
saccharum, Tilia americana, and Quercus rubra, with an under- 
story of Staphylea trifolia and Ostrya virginiana. Some of the 
herbaceous plant associates were Asarum canadense, Uvularia 
grandiflora, Smilacina racemosa, Polygonatum canaliculatum, 
Hydrophyllum virginianum, and Smilax ecirrhata. The area is 
part of an undisturbed section of woodland that harbors many 
species unusual in this section, such as Elymus riparius, Botry- 
chium multifidum var. intermedium, Liparis liliifolia, Panax 
quinquefolius, Lithospermum latifolium, Fraxinus nigra, Aster 
Shortit, as well as both species of Hepatica. So far as I have 
been able to ascertain, the woodland is a virgin climax stand, 
and it certainly has not been disturbed within memory. The 
soil in which the orchids grew was neutral. 

During the fifteen years of my residence in this section, this 
is the first trip I had ever taken to the particular section of the 
ravine in which the Epipactis occurred, so that I can only state 
that the summer of 1954 is the first year this orchid has come 
under my observation. It is very likely that it has been in 
this location for a number of years, and has merely been un- 
observed. 

The data for this collection is as follows:—north-facing, 
wooded slopes of ravine along creek tributary to Grassy Lake, 
on south side of Eton Drive, Biltmore Estates subdivision, 
5 miles north of Barrington, Lake Co., Illinois, July 21, 1954, 
Sleyermark 76351 (Herb. Chi. Nat. Hist. Mus.).—JULIAN A. 
STEYERMARK, CHICAGO NATURAL HISTORY MUSEUM AND MISSOURI BOTANI- 
CAL GARDEN. 


132 Rhodora [Vor. 57 


NORTHEASTWARD RANGE EXTENSIONS FOR Two SPECIES OF 
Hypericum IN Nova Scotia. Hypericum mutilum L., var. 
parviflorum (Willd.) Fern., known (Roland 1947) the length 
of the Nova Scotian mainland, is newly recorded for the Flora 
of Cape Breton Island: INVERNESS co.: Judique, occasional 
on brook bank, 13 Aug. 1951, Smith, Schofield, Sampson & Bent 
5014 (DAO, AC ADIA). 

Hypericum dissimulatum Bickn., discovered in Yarmouth, 
Digby, Lunenburg and Halifax Counties (Third Lake, Windsor 
Junction) by the Gray Herbarium Expedition to Nova Scotia, 
is now recorded for prerou co.: Barney River, 28 Aug. 1935, 
Groh (DAO). In the new Gray's Manual, Fernald (1950) 
says that this species may be a hybrid of H. canadense with 
either H. boreale or H. mutilum. In its fastigiately crowded 
dense cymes and its small apiculate capsules, this specimen 
resembles M. boreale; in its linear-oblong leaves, acute bracts 
and acute sepals, /7. canadense. The capsules are 2.5-3 mm. 
long, smaller than those of either putative parental species, 
approaching the size of those of H. mutilum, but as they show 
no developing seeds, this may be accounted for by hybridity. 
There is no suggestion of the rounded leaves and blunt sepals 
of H. mutilum, var. parviflorum. Thus, this may well be inter- 
preted as a specimen of the hybrid H. boreale X canadense.— 
Davip ÉRSKINE, TORONTO, ONTARIO, CANADA. 

CROOKED OAKS ON CAPE Cop AND ManTHA'S. VINEYARD.— 
While the late Dr. Frank G. Speck and the writer were studying 
the ethno-biological relations of Wampanoag Indians on Cape 
Cod and Martha’s Vineyard (Jour. Wash. Acad. Sci. 38 (8): 
257-205. 1948.), our attention was called to many oak trees 
in the region of our study which had erooked trunks. In some 
sections of North America such growth forms are explained as 
having been produced by Indians who broke the trunk of sap- 
lings in order to mark boundaries. In this case, however, 
Dr. Speck found no evidence that such an explanation could 
be attributed to the native tradition of the Wampanoag on 
Cape Cod and Martha’s Vineyard.—Rauen W. DEXTER, DE- 
PARTMENT OF BIOLOGY, KENT STATE UNIVERSITY, KENT, OHIO. 


Volume 57, no. 675, including pages 73-104, was issued 28 March, 1955. 


Dodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


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Vol. 57 May, 1955 No. 677 
CONTENTS: 
Echinodorus in the American Tropics. Norman C. Fassett........ 133 


Two New Carices from Southeastern United States. F. J. 
Hermann; ee ee, Tu. 156 


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Vol. 57 May, 1955 No. 677 


ECHINODORUS IN THE AMERICAN TROPICS 
NonMaN C. FASSETT ! 
I. INTRODUCTION 


Tuais study includes all the species of Echinodorus in the New 
World north of the Tropic of Capricorn; all are tropical with the 
exception of one that is confined to the United States. The 
temperate zone of South America has been excluded because of 
the several species described from Argentina, Uruguay and 
southern Brazil, that are not well represented in the available 
collections.? 

The first extensive treatment of Echinodorus as a genus dis- 
tinguished from Alisma and Sagittaria was by Micheli, in 
DeCandolle, Monographiae Phanerogamarum vol. 3: 44-60. 
1881. It was a splendid work; the descriptions of species were 
not only long and detailed, but covered just the points that seem 
most diagnostie today. Further reference to Micheli's acumen 
will be made in the discussion of E. andrieuxt. Micheli, to an 
extent not followed by later workers, made good use of the pel- 
lucid markings present in the leaves of many species. These 
can be seen only with transmitted light. The present writer 
uses a binocular microscope placed over a hole in the table-top, 
with an electric light in the drawer below. 

1 Professor Fassett died September 16, 1954. Fora tribute to him and an account 
of some of his botanical activities see RHopora 56: 233-242, 1954. Eds. 

2 Material has been borrowed from the United States National Museum, the New 
York Botanical Garden, the Chicago Natural History Museum, the Gray Herbarium, 
the Academy of Natural Sciences of Philadelphia, and the Missouri Botanical Garden. 
Locations of cited specimens are indicated by the abbreviations recommended in the 


recently published Index Herbariorum. Maps 1-3, 7, and 13c are on a base map 
copyrighted by McKnight & McKnight, Bloomington, Illinois. 


134 Rhodora [Vor. 57 


Unfortunately, many of the collections cited by Micheli are 
not represented in the herbaria available to the present writer. 
A few fragments of types turn up, but most of them consist of a 
bud or two and are of no value; some have mature nutlets and 
are very helpful. There are a few of Micheli's names that I 
have not been able to account for. 


II. SUBGENERA AND SECTIONS 


The dwarf plants passing as Echinodorus tenellus, or E. par- 
vulus, have been the basis of the genus Helianthium.? The 
segregation seems to have been on the following set of differences: 
Echinodorus: (1) style apical; (2) fruit-heads echinate; (3) 
achenes flat; (4) achenes beaked; (5) stamens 12-30, or rarely 
fewer; (6) carpels numerous.  Helianthium: (1) style not apical; 
(2) fruit-heads not echinate; (3) achenes very turgid; (4) achenes 
beakless or obscurely beaked; (5) stamens 6 or 9; (6) carpels 
relatively few. 

Another character not listed in the literature appears in the 
anthers; they are versatile in Echinodorus (sensu stricto) and 
basifixed in Helianthiwm. 

Some of these characters do not hold, and those that do hold 
seem insufficient for generic segregation. The style is nearly 
apical in two hitherto undescribed South American representa- 
tives of Helianthium (Figs. 71 and 72), and strongly lateral in 
Echinodorus cordifolius (E. radicans)—see Fig. 77 in Gray’s 
Manual, ed. 8, or figure on page 96, vol. 1, Britton and Brown’s 
Illustrated Flora, ed. 2; fruiting heads are scarcely echinate in 
E. cordifolius. Two species of the Helianthtium group have 
beaked achenes (Figs. 71 and 72), as does Echinodorus nymphaei- 
folius, which is also put into Helianthium on the basis of its small 
number of carpels and stamens. The dozen or so carpels and 
6 or 9 stamens with basifixed anthers in Helianthiwm, contrasted 
with about 100 carpels and usually 12-30 stamens with versatile 
anthers in Echinodorus (sensu stricto), appear to be consistent, but 
scarcely sufficient for generic separation. 

The genus Echinodorus may be subdivided as follows: 

a. Carpels many in a dense head; stamens 12-30; anthers versatile. 
Subgenus 1. Hchinodorus. 


3 Britton, Man. ed. 2, 54. 1905; Small, N. Am, Fl. 17, pt. 1: 45. 1909; Britton & 
Brown, Ill. Fl. ed. 2, 1: 95. 1913. 


1955] Fassett,—Echinodorus in the American Tropies 135 


a. Carpels 20 or fewer in a loose head; stamens 6 or 9; anthers basifixed. 


b. 
b. 


Subgenus 2. Helianthium. 
Ribs of achenes crested; blades of emersed leaves cordate; inflores- 
cence compound. ...................020.. Section Nymphaeifolii. 
Ribs of achenes not crested; blades of all leaves linear-lanceolate to 
elliptic; inflorescence of one or few whorls.........Seetion Tenelli. 


Subgenus Ecuinoporus.  Échinodorus Britton, Man. ed. 2: 54. 1905; 
Small, N. Am. Fl. 17, pt. 1: 45. 1909; Small, Man. S. E. Fl. 21. 1933. 
Type species, Alisma rostratum Nutt. 
Subgenus Helianthium (Engelm.) stat. nov. Helianthium Engelm. in 
Britton, l.c.; Small l.c. 
Section Nymphaeifolii, sect. nov., carpellorum costis alatis; 
laminis foliorum cordatis. Type species Alisma nymphaei- 
folium Griseb. 
Section Tenelli, sect. nov., carpellorum costis non alatis; laminis 
foliorum lanceolatis vel ellipticis. Type species Alisma tenellum 
Mart. 


The nutlets of Sect. Tenelli frequently have what may be 
termed a beaklet; it is à small umbo, of the same texture as the 
rest of the pericarp, directly below the style (Fig. 67) or some- 
times some distance below it (Fig. 70). Sometimes the style 
appears to come directly from the beaklet (Fig. 69). When a 
longer beak is present, it seems to be of stylar origin (Figs. 71, 72). 


III. Taxonomic TREATMENT 


a. Carpels many in a dense head; stamens 12-30; anthers versatile. 


Subgenus Echinodorus. 


b. Sepals 12-20-nerved, thin, withering, not accrescent, reflexed or loosely 


ascending in fruit; pellucid markings of leaves, if present, not reticulate. 
c. Blades of leaves cordate or truncate at base (Figs. 1-13) (elliptic on 
some dwarfed individuals, Figs. 8 and 9, and linear on some submersed 
leaves); petiole not winged. 
d. Flowers on pedicels several times as long as the fruiting heads. 

e. Blades with pellucid lines or dots (pellucid markings occasionally 
obscure or absent, especially on dwarfed or submersed mem- 
branous leaves); leaves glabrous except in species no. 8. 

f. Nutlets about 24 as broad as long, the dorsal margin rounded 
at summit (Fig. 22), and facial glands nearly round; little- 
known plant of Mexico..................005. 1. E. virgatus. 

f. Nutlet twice as long as broad, the dorsal margin sloping down- 
ward; facial glands mostly.longer than wide (Figs. 23 and 51). 
g. Sepals with papillose ridges; scape erect when young but 

soon procumbent; nutlet with summit of keel often crested 

and beak ascending (Figs. 23, 24)......... 2. E. cordifolius. 
g. Sepals with smooth veins; scape erect at maturity; nutlet 

with keel entire and beak erect or nearly so (Figs. 25-34). 


136 Rhodora IVoL. 57 


h. Leaves glabrous, with pellucid lines often several mm. 
long (Fig. 55); facial glands of nutlet often entering the 

base of the long (0.5-2 mm.) beak (Figs. 25, 29) 
3. E. Berteroi. 
h. Leaves with stellate hairs on lower surface near base of 
blade; pellucid dots abundant (Fig. 56) but lines absent 
or rare (except in varieties (Figs. 57, 58) in Paraguay and 
Argentina); facial glands of nutlet rounded at both ends 
and scarcely approaching to within one gland-length of the 

short. (0.2 mm.) and stout beak (Figs. 30, 31) 

4. E. grandiflorus. 
e. Leaves without pellucid markings, with stellate hairs on lower 
surface near base of blade and summit of petiole (hairs often 

rubbing off and leaving papilla-like bases). (Fig. 11). 

i. Inflorescence erect, often compound, with pedicels reaching 3 
em. in length... 5. E. muricatus. 

i. Seape creeping, simple, sometimes 2 m. long, rooting at nodes 
and sometimes proliferous, with erect pedicels sometimes 
reaching 7 em. in length... nonna ananuna 6. E. fluitans. 

d. Flowers nearly sessile... sees 7. E. bracteatus. 
c. Blades tapered or subtruneate at base (Figs. 14-21), petioles mostly 
winged. 
j. Petioles not winged (Fig. 13); little-known plant of Cuba. 
8. E. ovalis. 
j. Petioles winged (Figs. 14-21). 
k. Leaves veined from the base of the blade or sometimes slightly 
pseudopenninerved (Figs. 14, 15, 17-20); flowers pedicelled. 

l. Inflorescence racemose; pedicels 2-4-angled, scarcely ribbed or 
fluted; body of nutlet 2-2.5 times as long as wide (Figs. 41-43), 
with beak (0.5)-0.8-1 mm. long, the face with one gland (rarely 2) 

9. E. Andrieuzi. 

l. Inflorescence paniculate on larger plants; pedicels (when dry) 
with about. 10 winged angles; body of nutlet more than half as 
wide as long (Figs. 48, 49), with short stout beak and no glands 

10. E. paniculatus. 
k. Leaves pseudopenninerved (Figs. 16, 21); flowers sessile or nearly 

So. 

m. Leaves many times as long as wide, acuminate (Fig. 21); rachis 
of inflorescence with green wings about 2 mm. wide; fruit 
glandless (Fig. 50)... 0000000000000 00 ce eee 11. E. trialatus. 

m. Leaves about 3 times as long as wide, acute (Fig. 16); rachis of 
inflorescence not winged; fruit with several facial glands 


(Fig. 51). ce 12. E. Grisebachii. 
b. Sepals about 30-nerved, thick and brittle, enlarging in fruit to cover the 
fruiting head; pellucid markings reticulate (Fig. 64)....13. Æ. tunicatus. 


a. Carpels 20 or fewer in a loose head; stamens 6 or 9; anthers basifixed. 
Subgenus Helianthium. 
n. Ribs of nutlet crested (Fig. 53); blades of emersed leaves ovate and deeply 
eordate.... sese re Section NV ymphaeifolii. 
14. E. nymphaeifolius. 
n. Ribs of nutlets not crested (Figs. 66-72); blades of all leaves linear- 
lanceolate to elliptie (Figs. 73-76)... 000.000.0002. 0005. Section T'enelli. 


1955] Fassett,—Echinodorus in the American Tropics 137 


o. Summit of nutlet rounded above the beaklet (Figs. 66-70); stylar beak 
nearly absent or up to 0.4 mm. long; anthers 0.2-1.0 mm. long. 

p. Facial ribs 3 on each side (Figs. 66, 67, 70) or suppressed (Fig. 68) 

15. E. tenellus. 

p. Facial ribs 4 on each side (Fig. 69).............. 16. E. isthmicus. 

o. Summit of nutlet horizontal or sloping downward from the beaklet 

(Figs. 71, 72); stylar beak 0.6-0.8 mm. long; anther 0.4-0.6 mm. long. 

q. Nutlet with 4 distinct facial ribs (Fig. 72). ...17. E. quadricostatus. 

q. Nutlet with facial ribs nearly or quite obsolete (Fig. 71). 
18. E. magdalenensis. 


1. Ecuinoporus viRGATUS (Hook. & Arn.) Micheli in DC. Monogr. 
Phan. 3: 54. 1881; Buch. in Engler, Pflanzenr. 4, pt. 15: 32. 1903; Small, 
N. Am. Fl. 17, pt. 1: 47. 1909. Alisma virgata Hook. & Arn., Bot. Beech. 
Voy. 311. 1839. 

Tepic, Mexico. 

Our Fig. 22 shows the very distinctive nutlet, from a fragment 
of the Type (Mo). “Mexico” is the only locality given in the 
original description, but on page 275 is the statement, “When no 
habitat is mentioned, the specimens are understood to have been 
collected at Tepic.” Fig. 1 is traced from a drawing of the TYPE 
(NY); I have not seen an actual leaf to examine the pellucid lines, 
which Micheli describes as being elongate. The drawing of the 
type shows about 10 flowers at a node, on pedicels 0.5-1 em. long. 


2. ÉCHINODORUS CORDIFOLIUS (L.) Griseb. in Goett. Abh. 7: 257, repr. 
109. 1857, at least as to Linnean basionym, not as to other synonyms; 
Fernald, Rhodora 49: 107. 1947, and in Gray's Manual, ed. 8. 85. 1950. 
Alisma cordifolia L. Sp. Pl. 343. 1753. Sagittaria radicans Nutt., Trans. 
Am. Phil. Soe. 5: 159. 1837. Echinodorus radicans Engelm. in Gray, 
Man. 460. 1848, and ed. 2. 438. 1856; Robinson & Fernald in Gray's 
Manual, ed. 7. 84. 1908; Fassett, Man. Aquat. Pl. 93. 1940; Muenscher, 


Aquat. Pl. U.S. 84. 1944. 


In transferring the Linnean Alisma cordifolia to Echinodorus, 
Grisebach confused it with another species, perhaps the Brazilian 
E. macrophyllus (Kunth) Micheli. Grisebach also refers to Vell. 
Ie. Fl. Flumin. 10, t. 31. 1827, which illustrates an erect com- 
pound inflorescence quite unlike that of Æ. cordifolius as here 
treated. But, as Fernald demonstrates, the name rests on 
Alisma cordifolia L. from Virginia. 

Leaves erect, long-petioled, the blade broadly ovate, shallowly cordate 
or truncate at base and rounded at apex, 6-20 em. long, 4-8 em. wide; 


pellucid markings nearly round or elongate, seldom exceeding 1 mm. but 
rarely reaching 3 mm. in length, mostly 1 mm. or more apart (Fig. 54), 


138 Rhodora \Vou. 57 


tay 
++ 


HHeseessd 


* 
* 
* 
* 
As 
* 
* 
* 
* 
H 
' 
-~ 
+t.. 


À” . 


+ 

+ 
+ 
+ 


ts. 
E 


Map 1. Echinodorus cordifolius. 


very rarely obscure or lacking; scape at first erect, soon prostrate, usually 
with long-triangular bracts but sometimes with small leaves at the nodes: 
flowers 5-15 at a node, on pedicels 2-6 em. long; sepals with papillose 
ridges; nutlets about 2.5 mm. long and 0.8-1.2 mm. wide, with ascending 
beaks only about 0.2 mm. long (rarely —0.8 mm.) so that the head of 
nutlets appears smooth to the naked eye; nutlet with an often irregularly 
crested dorsal keel (Figs. 23, 24); facial ribs 3-4, often abruptly bent, the 
two toward the dorsal keel most strongly winged toward the summit; 
glands 1 or 2, short and rounded, not approaching to within one gland- 
length of the base of the beak. 

Atlantie Coastal Plain from southern Texas to northern Florida and 
the District of Columbia; up the Mississippi Valley to central Tennessee, 
southern Indiana, central Illinois, northeastern and southwestern Mis- 
souri, southeastern Kansas, eastern Oklahoma and northeastern Texas 
(Map 1). District of Columbia: flats of Potomac at Chain Bridge, 1 
August 1900, Steele (Ny). Virginia: Franklin, Southampton Co., 22-29 
July 1893, Heller 1026 (Pu, Mo, F, Us, Ny). North Carolina: Wilmington, 
2 August 1900, Williamson (Ny, pH); Edenton, Chowan Co., 10 July 
1938, Godfrey 5353 (us). South Carolina: mud bank, Santee River, near 
Jamestown, Berkeley Co., 27 June 1939, Godfrey & Tryon 118 (Mo, vs, 
ny); St. Johns, Porcher (Ny). Georgia: margin of pool in Flint River 
bottoms, Dooly Co., 3 September 1900, Harper 564 (us, Ny). Florida: 


1955] “assett, —Echinodorus in the American Tropics 139 


Aspalaga, October 1897, Chapman (mo); Apalachicola, Chapman (NY). 
Illinois: Menard Co., 1861, Bebb (F); Menard Co., August 1864, Hall 
(F, PH); lakes by Cahokia, 23 August 1878, Eggert (Mo); Bluffs Lake, St. 
Clair Co., 27 July 1878, Eggert (mo); swamps opposite St. Louis, August 
1863, Engelmann (Mo); Flat Lake, Calhoun Co., 19 October 1920, Metcalf 
1117 (vs). Indiana: in the dried up bottom of the Pitcher pond about 
6 mi. northwest of Mt. Vernon, Posey Co., 8 October 1916, Deam 22303 
(Ny). Kentucky: in water at Swan Pond, Wicliffe, 18 August 1923, 
McFarland & Anderson 168 (wo, vs). Tennessee: Nashville, August, 
Gattinger 2741 (wo, us, Ny). Alabama: Prattville, 22 July 1874, Smith 
(vs); exposed open places, Mobile, 30 May 1884, Mohr (vs). Mississippi: 
Jackson, 19 July 1925, Cook (vs); ereek bottom, Natchez Trace Parkway, 
24 June 1948, McDougall 1683 (vs). Missouri: low wet woods, near 
Torch, Ripley Co., 12 July 1933, Palmer & Steyermark 41598 (Mo); 
Little Creve Coeur Lake, 15 September 1891, Douglass (wo); Poplar 
Bluffs, Butler Co., September 1897, Russell (mo); Kennett, 18 September 
1893, Bush (mo); wet mud around margin of Goose Lake, Papinsville, 
Bates Co., 1 October 1938, Steyermark 9986 (F, Mo); low woods bordering 
Stulz Lake along Marmaton River, Nevada, Vernon Co., 29 September 
1938, Steyermark 9800 (r, Mo); ditches along road, Quilin, Butler Co., 27 
May 1939, Steyermark 26640 (v); Kings Lake, Lincoln Co., 7 August 
1927, Kellogg 816 (mo); Pike Co., 1860, Peck (r); Pence Lake, Vernon Co., 
1 August 1919, McAtee 3042 (vs). Arkansas: moist soil in bottoms, 
Little Rock, 7 June 1885, Haase (pH); Bayou Bartholomew, Lincoln Co., 
Yorktown, 10 September 1936, Demaree 13760 (mo, us); Fulton, 18 
September 1900, Bush 931 (mo); low swampy woods, Corning, Clay Co., 
24 June 1914, Palmer 6056 (mo, r); sandy bogs along Yellow Creek, near 
MeNab, Hempstead Co., 27 October 1924, Palmer 26722 (mo); Dryden, 
Craighead Co., 22 August 1913, Emig 55 (M0); wet spring ditches, Lake 
City, Craighead, 25 June 1929, Demaree 6940 (F, us); Lake Waponoca, 
Crittenden Co., 17 November 1910, McAtee 1884 (us). Louisiana: 
single plant at spring in pine barrens, Alexandria, 27 May 1899, Ball 482 
(mo); Hamburg, 13 September 1912, McAtee 2215 (us); Baton Rouge, 
July 1914, Griffins (NY); swamp east of Tallulah, 29 June 1946, Fassett 
26692 (wis). Kansas: Cowley Co., 1898, White (mo); Wilson Co., 1895, 
Haller 983 (mo, us, Ny). Oklahoma: Verdigris River, 21 August 1895, 
Blankinship (Mo, vs); E. Dartlesville, Washington Co., 31 August 1927, 
Stratton 505 (mo); Heavener, 20 June 1936, de Gruchy 119 (Ny). Texas: 
swamps, Dallas, 5 June 1902, Reverchon 4043 (Mo, Ny); Santa Maria, 
1889, Nealley 505 (r); water’s edge, Riviera, Brooks Co., 9 April 1933, 
Clover 825 (NY); Houston, October 1842, Lindheimer (mo); San Antonio, 
Bexar Co., 30 June 1911, Clemens 124 (mo); Los Fresnas, 10 July 1923, 
Runyon 446 (us); in marshy pond, Tarrant Co., 10 August 1925, Ruth 
1349 (us). 

3. Ecnivoponvs Berteroi (Spreng.) n. comb. Alisma Berteroi Spreng., 
Syst. 2: 163. 1825. A. Berteroanum Balbis in R. & S., Syst. Veg. 7: 1605. 
1830; Kunth, Enum. 3: 152. 1841. A. cordifolius Kunth, Enum. 3: 152. 


140 Rhodora (VoL. 57 


1841, not L. A. Sprenglii Kunth, Enum. 3: 152. 1841. Eehinodorus 
cordifolius 8. Berteroanus Griseb. in Goett. Abh. 7: 257, reprint 109. 1857. 
E. rostratus Engelm., Gray's Man. 460. 1848. 

Emersed leaves erect, long-petioled, the largest (reaching perhaps 20 
em. in length) broadly oval and shallowly cordate (Fig. 2) or broadly 
ovate (Fig. 3), the smaller becoming narrower in proportion, down to the 
small elliptie or lanceolate leaves on dwarfed plants (Figs. 4, 5, 7); pellucid 
lines usually very clear, the longest commonly reaching about 2.5 mm. in 
length (a maximum of 11 mm. observed), extending nearly from veinlet 
to veinlet (Fig. 55), rarely locally absent; submersed leaves, when present, 
usually ribbon-like and membranous; scape erect, the lower branches 
compound; head of nutlets appearing echinate due to the spreading 
beaks; nutlets (Figs. 25-29) broadly keeled, with 2 winged ribs alternating 
with 3 non-winged ribs; facial gland commonly single, close to the beak, 
usually with an attenuate tip entering its base. 

West Indies and southern Mexico, north to Ohio, Illinois, Missouri, 
Kansas, Texas and California (Maps 2a & 2b). Reported from Central 
America in Gray's Manual, ed. 8, and in North American Flora, probably 
erroneously. 


The positive identification of Alisma cordifolia as the creeping 
plant also known as Echinodorus radicans (no. 2 of the present 
treatment) was followed by the adoption of the name Æ. rostratus 
(Nutt.) Engelm. for the erect species sometimes called E. cordi- 
folius (Fernald, Rhodora 49: 108. 1947). But there are at least 
two older names for E. rostratus, both based on material from 
Guadeloupe. The description of Alisma Berteroi was brief: “A. 
foliis oblongis utrinque attenuatis 3 nervis, scapo 3 gono, capsulis 
cuspidatis." Only the last two words are helpful, referring to 
the beaked fruit (Figs. 25-29). The leaves were certainly from 
a dwarfed plant, like our Figs. 5 & 9. The description of 
A. Berteroanum is ample and clear, and the leaves are stated to 
be 5- or 7-nerved. Kunth, substituting the name A. Sprenglii 
for A. Berteroi, suggested that it was but a form of A. Bertero- 
anum. The more ample material now at our disposal demon- 
strates the polymorphism of the leaves of this species (Figs. 2-9) 
as quite sufficient to include both A. Berteroi and A. Berteroanum. 

The northern and southern phases of this species differ mainly 
in the length of beak on the nutlet; much material in the region 
of overlap, in Texas, Oklahoma and California, is intermediate. 
a. Beak of nutlet 1.2-2 mm. long, 14-24 as long as the body (Figs. 25-27); 

anthers 0.8-1.2 mm. long.................. 3a. E. Berteroi var. Berterot. 


a. Beak of nutlet 0.5-0.8 (-1) mm. long, 13-15 as long as the body (Figs. 
28, 29); anthers 0.5-0.8 mm.long......... 3b. E. Berteroi var. lanceolatus. 


1955] Fassett,—Echinodorus in the American Tropics 141 


3a. E. Berreror (Spreng.) Fassett, var. BERTEROI. Alisma Berteroi 
Spreng., l.c. From southern California, Kansas, Oklahoma and Texas, 
where many individuals are more or less intermediate with the northern 
variety, var. Berteroi ranges southward to central Mexico and northern 
Yueatan. I have not seen material from the mainland of Florida, but 
it is on the Keys. It appears to be local in Cuba, but abundant through- 
out the Bahamas, Hispaniola, Puerto Rico and the Lesser Antilles. Just 
as it approaches but is not known to reach continental Florida, so it is 
represented by several collections from Curacao but not from continental 
South America (Map 2a). 

Florida: Key West, June 1839, Ritch (us); dry ponds, Lower Mate- 
cumbe, 4 February 1892, Simpson 434 (us, Ny); Key West, Blodgett (Ny). 
Kansas: Cowley Co., 1898, White (mo, vs). Oklahoma: creek bed, 
covered with water to a depth of about 4’’, also on wet banks, 2 mi. sw. 
of Cache, 26 May 1936, de Gruchy 13 (mo); pond, 6 mi. n. of Oklahoma 
City, 30 May 1939, Waterfall 1182 (ny). Texas: on banks of Lake 
Como, Tarrant Co., 12 October 1919, Ruth 203 (eH, F); dry sink in prairie, 
Caldwell Co., 13 July 1943, Barkley 13135 (vs); in paludosis prope las 
Nuces, April 1834, Berlander 2422 (us); San Antonio, 2 October 1900, 
Bush 1222 (mo); San Antonio, 23 June 1911, Clemens 120 (mo); water's 
edge, La Joya, Hidalgo Co., 14 March 1933, Clover 639 (Nv); land on 
which water has stood near old river channel, Brownsville, Cameron Co., 
1-5 August 1921, Ferris & Duncan 3178 (Mo, Ny); water-hole, Corpus 
Christi, September 1884, Havard (us); Comanche Spring, New Braunfels, 
August 1850, Lindheimer 1233 (F, NY); swamp, Pleasanton Rd. 9 mi. s. 
of San Antonio, 9 July 1931, Metz 198 (Ny); shallow pools, Silsbee, Hardin 
Co., 13 June 1916, Palmer 10174 (mo); shallow pools, Uvalde, 14 October 
1916, Palmer 11031a (wo); Lubbock, Reed 3169 (vs); Curry Creek, 
Blanco Co., 1884, Reverchon 1772 (mo); Lavaca River, Jackson Co., 27 
August 1941, Tharp 15 (Mo); Demmit Co., 24 June 1941, Tharp (mo); 
Brownsville, 26 October 1927, Rose & Russell 24243 (us); Austin, August 
1921, Schulz 906 (Ny); San Marcos, Spring, 1897, Stanfield (ny). Cali- 
fornia: San Diego Co., August 1897, Allen (NY); Lakeside, San Diego Co., 
23 (or 30?) June 1906, Brandegee (Mo, F); lake margin, Laguna Cañon, 
Laguna Beach region, 26 July 1916, Crawford (mo); Sweetwater, July 
1884, Orcutt (Mo, F); drying lake bed, Stanford University lake, Santa 
Clara Co., 14 July 1946, Rose 46218 (mo). Baja California: along stream 
below Santiago, 6 May 1931, Wiggins 5663 (us, Ny, GH); Sonora: Canyon 
Estrella, Dist. Alamos, 1 October 1933, Gentry 421M (rF); Agua Caliente, 
Alamos, 10 November 1933, Gentry 896M (us). Chihuahua: water 
hole near Guadalupe, 11-12 October 1852, Thurber 805 (au). Coahuila: 
Torreon, 13-20 October 1898, Palmer 466 (F, GH, MO, US, Ny). Tamau- 
lipas: Matamoros, 2 September 1939, LeSueur 53 (F); El Mulato, 16 
August 1030, Bartlett 10984 (F, GH). Michoacan: swamps, Zamora, 
5000 ft. alt., 21 May 1901, Pringle $484 (F, GH, PH, MO, US, NY); LaHuerta, 
District of Apatzingan, 29-30 December 1890, Maury 5310 (Ny). Guer- 
rero: Iguala, 25 October 1900, Pringle 9277 (Gu, vs). Yucatan: Progreso, 
5 March 1899, Millspaugh 1692bis (F, vs); in fresh water pool, Progreso, 


142 Rhodora [Vor. 57 


11-15 August 1932, Steere 3093 (Ny). Bahamas: Long Cay, 7-17 Decem- 
ber 1905, Brace 4081 (F, Us, Ny); Pompey Bay, Acklin’s Island, 21 Decem- 
ber to 6 January 1906, Brace 4424 (r, Ny); water holes, Stopper Hill, 
Crocked Island, 9-23 January 1906, Brace 4829 (r); sink holes near 
Georgetown, Great Exuma, 22-28 February 1905, Britton & Millspaugh 


1955] Fassett,—Echinodorus in the American Tropics 143 


3104 (F, GH, US, NY); Cat Island, 1-6 March 1907, Britton & Millspaugh 
5798 (r); Watling's Island, Graham's Harbor to Cockburn Town, 15 
March 1907, Britton & Millspaugh 6206 (r, Ny); Grand Turk Island, 
The Wells and vicinity, 16 March 1911, Millspaugh & Millspaugh 9338 
(F£); black loam, swampy, Fresh Creek settlement, Andros, 31 March 
1905, Wight 256 (F, Gu, Ny). Cuba: Wright, without locality (Gx); 
Barreras, Playa de Tarara, Habana, 28 August 1930, Léon 14661 (GĦ); 
Playa de Marianao, Prov. Habana, 22 February 1910, Britton & Wilson 
4556 (NY); little lagoon on a serpentine hill, Guyana, Havana, 1 June 
1911, Léon 2585 (Ny); in water, serpentine hill, Cuabal de Figueras, ne. 
of Canasi, Matauzas, 31 May 1928, Leon 13381 (Ny). Jamaica: in shaded 
swamps, Salt Ponds, 27 December 1915, Harris 13312 (F, GH, NY, US). 
Haiti: shallow pond and swamp area between Terrier Rouge and Fort 
Liberté, northeastern alluvial plain, 26 June 1941, Bartlett 17472 (au, vs); 
moist place, Presqu’ile do Nord-Ouest, Port-de-Paix, Etang-Portalus, 
13 March 1928, Ekman 9683 (vs); among cat-tails, common, Etang 
Saumatre, 7 April 1920, Leonard 3516 (GH, Ny, vs); muddy bank of 
stream, scarce, La Source, Pikmi, Gonave Island, 7 July 1920, Leonard 
5188 (us, Ny); Ennery, Dept. de l'Artibonite, alt. 325-900 m., 3 February 
1926, Leonard 9449 (us); in bed of Mole River, Mole St. Nicolas, 13-19 
February 1929, Leonard & Leonard 13139 (us, GH, NY); muddy roadside, 
west of Trois Rivières, Port de Paix, 12 May 1929, Leonard & Leonard 
15630 (us); sea level, Bayeux, near Port Margot, 3 August 1903, Nash 60 
(r, vv). Dominican Republic: Peñon, Prov. Barahona, September 
1911, Fuertes 1132 (F, GH, Ny, vs); Sto. Domingo in paludosis ad Rio 
Muñoz, 100 m., 15 June 1887, Eggers 2477 (Ny); sitio húmedo, cerca de 
la playa, Cerea de Gaspar Hernandez, 5 April 1950, Jimenez 1958 (vs). 
Puerto Rico: river swamp, Coamo Reservoir, 8 February 1922, Britton, 
Britton & Brown 5949 (Ny); along stream, Coamo Springs, 23 March 
1906, Britton & Cowell 1340 (Ny); Guayanilla, 10 March 1913, Britton 
& Shafer 1793 (Ny, us); borders of Lake Guanica, Guanica, 11-12 March 
1913, Britton & Shafer 1868 (Nv, vs); in paludosis, Coamo, 21 December 
1885, Sintenis 3200 (aH, vs); Guanica, in arenosis litoralibus ad lagunas, 
20 January 1886, Sintenis 3375 (us, Mo); Guanica, in litore ad la Plata, 
24 February 1886, Sintenis 3844 (F, GH, PH, NY, US); Palo Seco, 3 February 
1916, Stevenson. 3837 (vs); marsh, Guanica Lagoon, 17 March 1937, 


FIG. l-21. LEAVES OF SUBGENUS ECHINODORUS (X 4). FIG. 1: E. virgatus. 
Mexico, Beechey (from drawing of type in NY). Fig. 2-5. E. Berteroi var. Berteroi. 
2: Puerto Rico, Sentenis 3200 (US). 3: Guadeloupe, Duss 3652 (US). 4, 5: Haiti, 
Bartlett 17472 (US). Fig. 6—9: E. Berteroi var. lanceolatus. 6: Illinois, Engelmann 
(MO). 7-9: Illinois, Engelmann (Type of E. rostratus var. lanceolatus, in MO). 
Fig. 10. ŒE. grandiflorus var. grandiflorus, Panama, Woodson & Schery 754 (US). 
Fig. 11. ŒE. muricatus, Brazil, Tate 61 (NY). Fig. 12. XE. bracteatus var. bracteatus, 
Ecuador, Hitchcock 20084 (US). Fig. 13. ŒE. ovalis, Cuba, Wright (US). Fig. 
14, 15. E. Andrieuri. 14: Mexico, Pringle 8256 (US). 15: Mexico, Mexia 1065 
(NY). Fig. 16. E. Grisebachii, Cuba, Wright 4198 (isotype in US). Fig. 17-20. 
E. paniculatus var. paniculatus. 17: Venezuela, Fernandez 71 (US). Fig. 18: 
Brazil, Krukoff 2030 (US). 19: Brazil, Glaziou 14288 (US). 20: Venezuela, Chardon 
26 (US). Fig. 21. E. trialatus, Colombia, Cuatrecasas 5283 (type in US) 


| 
| 
| 


144 Rhodora [Vor. 57 


2002F "i n 
NORTH AMERICA - Mem D 
Q 800 490 690 MILES mS ed i 
——S——*—— NS A 24 4 
ca. aata one. tn j peaa Ae 
— NA m ? 2g wor 
copraiont av vr f 
MEKMONT a MeKNIGHT, BLOOMINGTON. ILLINOIS 


Map 2a. Echinodorus Berteroi var. Berteroi. 


Vélez (Nv). Virgin Islands: King's Hill gut, St. Croix, 17 March 
1897, Ricksecker 255 (¥, Mo, vs); Catherine's Rest, 14 February 1896, 
Ricksecker 275 (F, GH, MO, Ny, Us). Antigua: in small colonies in estate 
ponds and in sluggish streams, Elliots, 15 February 1938, Box 1370 (vs); 
ponds and the lower reaches of sluggish streams, Judges, 1 November 
1937, Box 1243 (us). Guadeloupe: Capesterre Marie-Galante, 1895, 
Duss 3652 (F, GH, MO, NY, PH, US); Grande-Terre, 5 March 1937, Stelé 
1555 (us); Gosier, March 1937, Questel 4584 (us). Barbados: Ealin 
Grove pond, 13 August 1906, Dash 163 (us, Ny). Curacao: Water hole, 
Mt. Pleasant, 20-27 March, 1913, Britton & Shafer 3115 (F, Ny, vs). 

3b. E. Berreror (Spreng.) Fassett, var. lanceolatus (Engelm.) n. 
comb. Æ. rostratus var. lanceolatus Engelm. ex Wats. & Coult. in Gray’s 
Man. ed. 6. 556. 1891. Æ. cordifolius var. lanceolatus Mackenzie & Bush, 
Man. Fl. Jackson Co., Missouri 10. 1902: Robinson & Fernald in Gray's 
Man. ed. 7. 84. 1908. E. cordifolius f. lanceolatus Fernald, Rhodora 38: 
73. 1936. E. rostratus f. lanceolatus Fernald, Rhodora 49: 108. 1947, 
and in Gray’s Man. ed. 8. 85. 1950. Alisma rostratum Nutt., Trans. 
Am. Phil. Soe. 5: 159. 1837. Echinodorus rostratus Engelm., Gray's 
Man. 460. 1848; Wats. & Coult. in Gray's Man. ed. 6. 556. 1891; Fernald, 
Rhodora 49: 108. 1947, and in Gray’s Man. ed. 8. 85. 1950. K. cordi- 
folius Mackenzie & Bush, Man. Fl. Jackson Co., Missouri 10. 1902; 


1955] Fassett,—Echinodorus in the American Tropics 145 


Map 2b. Echinodorus Berteroi var. lanceolatus. 


Britton & Brown, Ill. Fl. 1: 86. 1896; Small, Fl. S.E. U.S. 42. 1903; 
Robinson & Fernald in Gray's Man. ed. 7. 84. 1908; Small, N. Am. FI. 
17, pt. 1: 47. 1909; Small, Man. S.E. Fl. 22. 1933; Fassett, Man. Aquat. 
PI. 93. 1940; Muenscher, Aquat. Pl. U.S. 82. 1944. 

Centering in the lower Missouri and central Mississippi River valleys, 
north to central Illinois and southern South Dakota, thence southward 
to intergrade with var. Berteroi in Texas and in outlying districts west- 
ward in California (Map 2b). 

Ohio: pond 14 mi. west of Vause Station, Liberty Twp., Ross Co., 23 
July 1936, Bartley & Pontius 91 (Ny). Illinois: Oquawka, Henderson 
Co., 29 July 1892, Patterson (Fr); Peoria Co., Stewart (F); Peoria, Brendell 
(r, vs); Beardstown, August 1928, Turner 784 (F); Menard Co., 1861, 
Bebb (F); Menard Co., various collections of Hall (F, Mo, Us, NY); margin 
of ponds, American Bottom, September 1846, Engelmann (mo)—marked 
"Janceolat." by Engelmann, and to be taken as Type of £F. rostratus var. 
lanceolatus Engelm.); Mitchell, Madison Co., 7 September 1927, Steyer- 
mark 871 (Mo); Mason Co., 23 August 1845, Mead (mo); St. Clair Co., 
12 June 1879, Eggert (F, Mo); East St. Louis, 9 August 1897, Eggert (F, 
us, NY). Iowa: mud flats, infrequent, Fremont Co., 15 August 1898, 
Fitzpatrick (F, Mo, Ny); Hamburg, Hitchcock (mo, Ny); California Junction, 
Harrison Co., October 1904, Shimek (us) & 25 August 1908, Shimek (mo). 
Missouri: Independence, 22 August 1882, Bush 10 (mo); Atherton, Jackson 
Co., 9 July 1896, Bush 437 (mo, us); Clay Co., 9 August 1895, Bush 598 
(mo); St. Louis, August 1845 & August 1848, Engelmann (Ny); Sheffield, 


146 Ithodora [VoL. 57 


28 September 1895, Mackenzie 556 (Mo, Ny); Wolf Island, Mississippi 
Co., 11 July 1933, Palmer & Steyermark 41508-A (mo). South Dakota: 
Yankton, September 1892, Thornber (mo); water hole, Vermillion, Clay 
Co., 18 August 1927, Over 17422 (us). Nebraska: Omaha, August 1899, 
Williamson (pH); in ditch along road, flooded early in growing season, 
Holt Co., 20 miles south of O'Neill, 20 August 1941, Tolstead 41464 (mo); 
Dodge Co., 18 July 1883, Pepoon (wo). Kansas: Claflin, 19 August 
1929, Benke 5099 (F); ponds, Crawford Co., 1896, Hitchcock 853 (Mo, Us, 
Ny); Ellis Co., 28 July 1885, Kellerman (us, Mo); Manhattan, 29 July 
1892, Clothier (us); Argentine, 23 September 1895, Mackenzie (Ny). 
Oklahoma: Verdigris, 5 October 1894, Bush 605 (Ny, Mo); shallow water, 
Fish Hatchery Pond, Medicine Park, 28 May 1936, de Gruchy 52 (Mo, NY); 
Sapulpa, 1 October 1894, Bush 606 (M0); pond west of Daugherty, Murray 
Co., 2 June 1945, Waterfall 6044 (M0); wet sandy shore of Crystal Lake, 
2 miles north of Norman, Cleveland Co., 9 October 1936, Hopkins 775 
(mo); Cache Creek, 2 miles southwest of Cache, 26 May 1936, de Gruchy 
13 (NY); Durant fish hatchery, Durant, 29 May 1936, de Gruchy 66 (NY). 
Texas: bases submerged in water of small pool near Barton Springs 
Creek, 23 July 1943, Barkley 15 (mo, Ny); common in ponds, Columbia, 
14 October 1900, Bush 1497 (mo); Dallas, 2 July 1872, Hall 622 (F, mo, 
Us, NY); Dallas Co., 1926, Hynes 15 (mo); Comanche Spring, New Braun- 
fels, September 1849, Lindheimer 1232 (Mo, rH, vs); Corpus Christi, 
May 1913, Orcutt 5800 (mo); ditches, Wharton, Wharton Co., 24 September 
1914, Palmer 6630 (r, Mo); shallow pools, Brazoria, Brazoria Co., 3 May 
1916, Palmer 9666 (mo); Lake Como, Tarrant Co., 7 October 1920, Ruth 
203 (F, Mo); Camp Barkley, Taylor Co., 1943, T'olstead 7346 (mo). Cali- 
fornia: Ramona, October 1903, Brandegee (us); Lakeside, San Diego Co., 
30 June 1906, Brandegee (us); Laguna Canyon, 26 July 1916, Crawford 
(us); Tranquillity, Fresno Co., 10 August 1937, Hoover 2656 (vs); La- 
guna Canon, Orange Co., alt. 350 ft., 3 September 1918, Johnston 2151 
(us); Sweetwater, July 1884, Orcutt 321 (us); shores of Elsinore Lake, 
San Diego Co., 3 November 1891, Parish 2246 (F, Mo); Laguna Lakes, 
15 July 1919, Streit (pH); Moreno Dam, San Diego Co., alt. 3200 ft., 
7 October 1938, Wiggins 9204 (us). 

There are two sheets that would extend the range considerably, 
but whose authenticity is to be doubted. One is labelled Dela- 
ware: borders of ponds, Canterbury, July, Wm. M. Canby (NY). 
Mr. Bayard Long, of the Academy of Natural Sciences of Phila- 
delphia, writes me that this is without doubt a case of mixing of 
labels, and that E. rostratus (E. Berteroi) does not to his knowl- 
edge appear anywhere in the region. Canby did collect Æ. 
parvulus at Canterbury and his material appears in several 
herbaria. Actually, the label on the sheet in question bears the 
name Echinodorus parvulus; "parvulus" has been crossed out 
and “cordifolius” written above it, in a hand other than the 


1955] Fassett,— Echinodorus in the American Tropics 147 


original collector’s. The other sheet is labelled Minnesota: 
borders of ponds in mud, Fort Snelling, July 1888, Dr. W. H. 
Forwood (US). Dr. Gerald B. Ownbey of the University of 
Minnesota writes me that none of the many collectors in the 
Fort Snelling area have taken this species, unknown north of 
southern Iowa. He quotes Dr. John Moore who thinks Dr. 
Forwood was an army doctor who did some traveling on the 
plains. In view of these facts, and of the extensive collecting 
that has been done in this part of Minnesota and southward 
along the Mississippi River, it seems likely that Dr. Forwood's 
plant did not come from Fort Snelling. 

Although the northern phase of E. Bertero?, like the southern, 
generally has leaves of the: broadly cordate type (Fig. 6), the 
earliest name applied in the varietal category is that of Engel- 
mann, applied to dwarfed plants with lanceolate leaves (Figs. 
8, 9 are from the type). And so we have another case where a 
slightly atypical plant, named and distinguished from the com- 
mon phase, becomes the rvP& of the common phase from which 
it was originally distinguished. The type of E. rostratus var. 
lanceolatus was only a small and narrow-leaved plant such as 
one might piek up on almost any pond-shore where the species 
abounds. 

Alisma rostratum Nutt. was described from “the ponds of the 
Verdigris river of Arkansas" (now in Oklahoma). The type is 
at the Academy of Natural Sciences of Philadelphia, and through 
the kindness of Mr. Walter M. Benner I have before me two 
achenes from this collection. Fig. 28 illustrates one of them; 
while it belongs with the northern short-beaked variety, it 1s in 
the region of overlap and shows a little more tendency toward 
the long-beaked variety than do plants from regions farther north. 

Introgression between E. Berteroi and E. cordifolius is suggested 
by several individuals. Missouri: Stoddard Co.; woods border- 
ing Swan Pond, T 28 N, R 10 E, sect. 35, 4 mi. south of Advance, 
28 August 1948, Steyermark 66159 (r). Leaves with pellucid 
lines 2.5 mm. long, and achenes with a suberect beak and alter- 
nate ribs winged, as in Æ. Berteroi; scape creeping, sepals papil- 
lose, and glands of the achene well below the beak and rounded 
as in E. cordifolius. Illinois: lakes by Cahokia, 23 August 1878, 
Eggert (PH). Leaves with pellucid lines reaching 3 mm. as in 
E. Berteroi; scape creeping as in E. cordifolius; achenes short- 


148 Rhodora [Vor. 57 


beaked as in Æ. cordifolius but sometimes with facial glands 
tapered into the beak as in E. Bertero?; papillose condition of 
sepals intermediate. Texas: Lavaca Run, Jackson Co., 27 
August 1941, Tharp 15 (mo), and Gillespie Co., Jermy (mo). 
Pellucid lines closely spaced and reaching 4 mm. in length, and 
sepals scarcely papillose, as in Æ. Berteroi; scape creeping as in 
E. cordifolius. 

4. ECHINODORUS GRANDIFLORUS (Cham. & Schlecht.) Micheli in 
DC., Monogr. Phan. 3: 57. 1881. Alisma grandiflorum Cham. & 
Schlecht., Linnaea 2: 152. 1827; Kunth, Enum. 3: 153. 1841; R. & S., Syst. 
Veg. 7: 1606. 1830; Seub. in Mart., Fl. Bras. 3, pt. 1: 108. 1847. Alisma 
floribundum Seub. in Mart., Fl. Bras. 3, pt. 1: 109. 1847. Echinodorus 
floribundus Seub. in Warm., Symb. fasc. 13: 113. 1872. E. grandiflorus 
x floribundus Micheli in DC., Monogr.'Phan. 3: 58. 1881. FE. grandi- 
Jlorus var. floribundus Hauman, Anal. Mus. Nae. Buenos Aires 27: 311. 
1915, in part at least. K. grandiflorus var. longiscapus (Arech.) Hau- 
man, l.c., in part. Æ. muricatus Woodson & Schery, Fl. Panama, Ann. 
Mo. Bot. Gard. 30: 101. 1943, probably not E. macrophyllus B. muri- 
catus Griseb. 

Leaves erect, long-petioled except in individuals from dry soil; blades 
shallowly cordate, about as wide as long, often very large and reaching 
55 em. in length; pellucid markings usually very clear (Figs. 56-58); 
inflorescence simple or branching; flowers verticillate on pedicels that 
may reach 4 em. in length as the fruits mature; nutlets (Figs. 30-31) 
winged on the back, with 3 or 4 ribs that are sometimes slightly winged 
toward the summit, usually 2 facial glands that are elongate and rounded 
at both ends and are placed well below the summit of the nutlet, and a 
short stout beak. 


VARIETIES OF ÉCHINODORUS GRANDIFLORUS 
a. Blades of leaves with pellucid dots, short lines very rare or absent, petals 
white or pink... naaa aaau var. grandiflorus. 
a. Blades of leaves with numerous pellucid lines as well as dots. 
b. Petals yellow; summit of petiole and base of blade with stellate hairs 
var. aureus. 
b. Petals white; leaves glabrous... ... illis var. ovatus. 


4a. E. GRANDIFLORUS (Cham & Schlecht.) Micheli, var. GRANDIFLORUS. 
Alisma grandiflorum Cham. & Schlecht., l.c. Leaves with tubercle- 
based stellate hairs about the summit of the petiole and base of the blade, 
the hairs sometimes wearing off so that the persistent bases give a muri- 
cate appearance; pellucid markings nearly all dots (Fig. 56) with lines 
very rare or absent; petals white or rarely pink. 

Cuba and Guatemala to Paraguay and southern Brazil (Map 3). 
This range appears to be bicentrie, with a long gap between Colombia 
and Bolivia. I have not found any positive differences between the 
var. grandiflorus of the two parts of the range. 


Map 3. Echinodorus grandiflorus var. grandiflorus. 


Cuba: Prov. Pinar del Rio, Arroyo Mantua, S. Francisco ad stagnum, 
flor. albi, 29 May 1920, Ekman 10974 (xy). Guatemala: Rio de Los 
Esclavos, Dept. Santa Rosa, August 1892, Heyde & Lux 4074 (GH, NY, 
us); vicinity of Quirigua, Dept. Izabal, alt. 75-225 m., 15-31 May 1922, 
Standley 23858 (GH, NY, vs); in a slough, Quirigua, 17 May 1937, Muensch- 
er 12065 (Fr); edge of small stream, herb 1 m. tall, petals white, near 
El Molino, Dept. Santa Rosa, alt. about 600 m., 26 November 1940, 
Standley 78399 (r); Los Amates, Dept. Izabal, alt. 90 m., 20 February 
1907, Kellerman 6615 (v); Lago Retana, between Ovejero and Progreso, 
alt. 600 m., 26 November 1939, Steyermark 31985 (F); in water of swamp, 
valley of Río Chiquimula, 114 mi. northeast of Chiquimula, alt. 400 m., 
21 October 1939, Steyermark 30120 (r). El Salvador: edge of river, 
plants 1 m. high, petals white, vicinity of Ahuachapan, Dept. Ahuacha- 
pan, alt. about 700-1100 m., 16-25 January 1947, Standley & Padilla 
2518 (F); herb 3-4 ft., fls. white, open swamp, vicinity of Tepetitán, 
Dept. San Vicente, alt. about 400 m., 6 March 1922, Standley 21427 
(GH, NY, vs); Zacatecoluca, March 1922, Calderon 301 (GH, Ny, US); 
herb 3-4 ft., fls. white, swamp along river, vicinity of Nahulingo, Dept. 
Sonsonate, alt. about 220 m., Standley 22019 (GH, Ny, vs); herb 3-5 ft., 
fls. white, in swamp, abundant, vicinity of Ateos, Dept. La Libertad, 
17 April 1922, Standley 23346 (us). Honduras: in marsh, acaulescent 
herb 3-4 ft., inflorescence much branched, vicinity of Tela, Dept. At- 
lántida, at sea level, 14 December 1927-15 March 1928, Standley 54522 
(r, vs); Tela, 1923, van Severén 38 (vs); corolla white, in marsh, near 


150 Rhodora (VoL, 57 


25 BERTERO 27 
var BERT EROJ 


3T 29 
BRAC TEATUÉ 


28 
BERTEROI 29 
varLANCEOL ATUS 


924 | so% y j 
M3 TRALATUS of 
CRANOF LORUS GRISBACH|| 9» NYMPHAEIFOLIUS 


1955] Fassett,—Echinodorus in the American Tropics 151 


Tela, 8 September 1034, Yuncker 4985 (r, Mo); fls. blancas, en sabana, 
margenes del Rio Jalán, 3 kms. al norte de Guaimaca, Dept. Morazán, 
13 June 1950, Molina 3065 (v); El Banco, Dept. Comayagua, alt. 640 
m., 13 March 1945, Valerio 2341 (v); in marsh, herb 1 m., petals white, 
vicinity of Juticalpa, Dept. Olancho, 380-480 m., 5-16 March 1949, 
Standley 18057 (r). Nicaragua: in ditch, rlants 1 m. high, fls. white, 
vicinity of La Libertad, about 500-700 m., Dept. Chontales, 29 May- 
1 June 1947, Standley 8977 (F). Panamá: swamp, Boquete, Prov. Chiri- 
qui, 3800 ft., 21 May 1938, Davidson 691 (F, GH); flowers lavender pink, 
lower portion of valley and marshes along R. Antón, El Valle de Antón, 
about 500 m., 2 February 1935, Hunter & Allen 381 (F, vs); inflorescence 
1.5-2 m. tall, marshy borders of streams, El Valle de Antón and vicinity, 
500-700 m., 23-27 July 1935, Seibert 490 (an, Mo, NY); flowers white, 
vicinity of Boquete, alt. 1200-1500 m., 24-26 July 1940, Woodson & 
Schery 754 (GH, MO, US); in bog, fiowers white, between Las Margaritas 
and El Valle, 15 July, 8 August 1938, Woodson, Allen & Seibert 1736 
(Gu, Ny). Colombia: loco uliginoso, 1100 m., Timbs, Dept. Valle, 3 
January 1937, von Sneidern 1135 (Ny). Bolivia: Buer a Vista, Dept. Santa 
Cruz, flor. blanca, 29 October 1916, Steinbach 2868 (v). Brazil: peren- 
nial herb 1.5 m. high, white flower, road to Sio Miguel near km. 11, 
swampy land near corriga, alt. 650 m., State of Minas Geraes, 28 Decem- 
ber 1929, Mexia 4179 (F, GH, MO, PH, Us, NY); Cambuquira, 25 December 
1935, Barreto 927 (Fr); Minas Geraes, 1840, Claussen, cited by Micheli 
as var. floribundus (Gu); lccis paludosis, Pinhaes, Paraná, 885 m., 14 
October 1914, Dusén 1132a (GH); Blumenau, Sta. Ca'arina, January 
1888, Ule 540—leaf with pellucid lines and dots of var. ovatus but pubes- 
cence of var. grandiflorus (us); Minas Geraes, 1845, Widgren (us); Minas 
Geraes, 1865, Regnell 418 (us); Prov. Caera, August-Nc vember 1838, 
Gardner 1860, cited by Micheli urder var. floribundus (Ny). Paraguay: 
Pileomayo River, Morong 853 (mo, vs). 

Fig. 22-53. NUTLETS OF SUBGENUS ECHINODORUS ( X 10). Fig. 22: E. virgatus, 
Mexico, Beechey (fragment of type, in MO). Fig. 23, 24: E. cordifolius. Tennessee, 
Gattinger 2741 (US). Fig. 25-27. E. Berteroi var. Berteroi. 25: Curacao, Britton 
& Shafer 3115 (US). 26: Puerto Rico, Sintenis 3375 (US). 27: Mexico, Pringle 8484 
(US). Fig. 28, 29. ŒE. Berteroi var. lanceolatus. 28: Illinois, Engelmann (type of 
E. rostratus var. lanceolatus in MO). 29: Oklahoma, Nuttall (type of Alisma 
rostrata, in PH). Fig. 30, 31. ŒE. grandiflorus var. grandiflorus. 30: Honduras 
Standley 53610 (US). 31: Uruguay, Bartlett 2182 (unident fled variety in US). 
Fig. 32, 33. E. muricatus. 32: Brazil, Tate 61 (US). 33: Brazil, Tate 61 
(NY). Fig. 34. Unidentified species, perhaps E. muricatus, Bolivia, Rusby 1419 
(NY). Fig. 35, 36. E. macrophyllus, Brazil, Pereira 3602 (US). Fig. 37, 39. 
E. bracteatus var. bracteatus, Ecuador, Hitchcock 29984 (NY); fig. 39: var. 
efenestratus, Ecuador, Rimbach 90 (type in F). Fig. 40. ŒE. ovalis, Cuba, Wright 
(immature nutlet in US). Fig. 41—43. E. Andrieurii. 41: Mexico, Rose, Standley 
& Russell 14100 (NY). 42: Mexico, Mexia 1065 (US). 43: Mexico, Hinton 2667 
(US). Fig. 44—49. E. paniculatus. 44: Bolivia, Kuntze (NY). 45: Venezuela, Chardon 
26 (US). 46: Bolivia, Cárdenas 4466 (US). 47: Venezuela, Fernandez 71 (US). 48: 
Bolivia, Cárdenas 4466 (US). 49: Ecuador, Hitchcock 20278 (US). Fig. 50. E. 
trialatus, Colombia, Cuatrecasas 4283 (type in US). Fig. 51. E. Grisebachii, Cuba, 
Wright 3198 (US). Fig. 52. E. tunicatus, Panama, Maxon 7095 (US). Fig. 53. 
E. nymphaeifolius, Mexico, Mell 2090 (US). 


152 Rhodora [Vor. 57 


4b. E. GnANDIFLORUS. (Cham. & Schlecht.) Micheli, var. aureus, 
Fassett, n. var., foliorum laminis punctis pellucidis lineisque brevibus 
(0.2-0.5 mm. longis) instructis; floribus flavis; fructus ignotus.—Pubes- 
cence of leaves as in var. grandiflorus; pellucid dots intermixed with 
lines 0.2-0.5 mm. long; flowers vellow.—Cuba: in bog, 2 ft. or more, 
flowers yellow, near Rincón, Prov. Havana, 20 January 1905, van He- 
mann 440 (TYPE in F; GH, NY). In absence of fruit, the exact relation- 
ships of this plant cannot be determined with certainty. 


Throughout the northern part of its continental range, K. 
grandiflorus is constant in its diagnostic characters, responding, 
apparently, only to ecological factors in such features as size of 
blades and length of petioles. Southward, in southern Brazil, 
Argentina, Uruguay, and Paraguay, many plants are glabrous, 
with pellucid lines, mostly 1 mm. or less long, replacing many of 
the dots in the blades (Fig. 57): they probably represent Æ. 
grandiflorus var. ovatus Micheli in DC., Monogr. Phan. 3: 
58. 1881. In about the same region appear other plants with 
pedicels up to 4 em. in length, and pellucid lines to 2 mm. long 
(Fig. 58), often suggesting those of KE. Berterot. Several taxa 
have been described from the area, including K. Sellowianus 
Buch., Pflanzenr. 4, fam. 15: 30. 1903, and E. longiscapus 
Arechavaleta, Anal. Mus. Nac. Montevideo 4, pt. 1: 67, pl. 2. 
19083. The treatment of Argentinean Alismataceae by Hauman, 
Anal. Mus. Nac. Buenos Aires 27: 311. 1915, appears to have 
the advantage of being based on familiarity with the plants in 
the field, but Hauman does not express complete certainty as to 
the relationships of the species named in this paragraph. 


5. ECHINODORUS MuRIcATUS Griseb., Bonplandia 6: 11. 1858. Æ. 
macrophyllus 8 muricatus Micheli in DC., Monogr. Phan. 3: 50. 1881. 
Not E. muricatus Woodson & Schery, Fl. Panama, Ann. Mo. Bot. Gard. 
30: 101. 1943. 

Leaves erect, long-petioled, the blades cordate at base, a little longer 
than wide (Fig. 11), sometimes reaching about 50 em. in length, with- 
out pellueid lines or dots (veins rarely pellucid); pubescence stellate 
mostly about the summit of petiole and base of blade, often wearing off 
and leaving papilla-like bases; inflorescence a meter or more high, com- 
pound and ample; pedicels 1—3 em. long, subtended by bracts or some- 
times by reduced cordate leaves; fruit with body 2.5-3 mm. long and 
beak 0.3-1 mm. long, keeled, with 4-5 scarcely winged ribs on each 
face, mostly with 1—3 glands (Fig. 32) or rarely glandless (Figs. 33-34). 

Panama to Ecuador and British Guiana (Map 4). Colombia: wet 
swamp, alt. 130-140 m., Puerto Berrio, Dept. Antioquia, 14 July 1917, 


1955] Fassett, —Echinodorus in the American Tropics 153 


Pennell & Rusby 57 (Ny); Puerto Berrio, 11-13 January 1918, Pennell 
3734 (GH, Ny). Venezuela: La Rubiera cerea de Calabozo, Guarico, 
1925, Grisel 1—mixed with E. paniculatus (Ny, vs). British Guiana: 
in water, 4 feet high, Limao, Mount Roraima, 21 September 1927, Tate 
61 (Ny); Berbice, June 1889, Jenman 5162 (Ny); near Neue Amsterdam, 
April 1889, Jenman 5080 (NY). 


I have seen no Echinodorus with leaves actually muricate, but 
there are 3 species with stellate hairs that eventually drop off to 
leave just the tuberculate bases: these are E. grandiflorus, E. 
bracteatus and E. muricatus. The type of E. muricatus is from 
Panama; I have not seen this type, nor any other material of 


P id n = SP = 
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2 AA A BR 
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C: l 4! . 
pae Sm QC 
EE." T @ at 
CAEN 
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Map 4. Echinodorus muricatus. 


E. muricatus from Panama. But Micheli, who seems to have 
examined the type, and whose observations concerning pellucid 
markings appear to be very reliable, lists Æ. macrophyllus B 
muricatus under “Folia punctis lineisve pellucidis destituta," 
thus excluding the possibility of the type of E. muricatus be- 
longing with E. grandiflorus or E. bracteatus. 

E. muricatus was treated by Micheli as a variety of the more 
southern FL. macrophyllus (Kunth) Micheli. Both species have 
leaves without pellucid markings (Fig. 59) but those of Æ. 
macrophyllus are glabrous. The achene of Alisma macrophyllum 
was described by Kunth (Enum. Pl. 3: 51. 1841) as being 2-ribbed 
on each side. The only good fruiting material of E. macrophyllus 
available to the writer is Pereira 3602 from São Paulo (v8); it 
has fruits mostly 3-ribbed (Fig. 35) or 4-ribbed (Fig. 36). This 
plant also shows variation in the number of glands, not mentioned 
by Kunth, but given by Micheli as usually 3 or 4. 

Collections from near Lake Rogagua, Bolivia, H. H. Rusby 
1419 (flowers & fruit —Nv) and 1415 (leaves & fruit—ny) are 


154 Rhodor: (VoL. 57 


5 BGRANDFLORUS 


59 MACROPHYLLUS 


N: 


60 BRACTEATU 64 TUNCATUS 


57 GRANDFLORUS 6! OALIS 6% NYMPHAEIFOLUS 


FIG. 54-65. PORTIONS OF LEAVES OF SUBGENUS ECHINODORUS, AS SEEN BY TRANS- 
MITTED LIGHT (X about 7). Fig. 54. E. cordifolius. Tennessee, Gattinger 2741 
(US). Fig. 55. E. Berteroi. Santo Domingo, Eggers 2477 (US). Fig. 56. E. 
grandiflorus var. grandiflorus. Honduras, Standley 53610 (US). Fig. 57. E. 
grandiflorus var. ovatus. Argentina, Venturi 2416 (US). Fig. 58. ŒE. grandiflorus, 
unidentified variety. Bolivia, Cárdenas 4465 (US). Fig. 59. E. macrophyllus, 
Brazil, Pereira 3602 (US). Fig. 60. ŒE. bracteatus var. bracteatus, anama, Standley 
26724 (US). Fig. 61. FE. ovalis, Cuba, Wright, (US). Fig. 62. E. trialatus, 
Colombia, Cuatrecasas 7301 (US). Fig. 63. E. Grisebachii, Cuba, Wright 3198 (US). 
Fig. 64. E. tunicatus, Costa Rica, Lankester 947 (US). Fig. 65. E. nymphaeci- 
folius, Mexico, Mell 2090 (US). 


1955] Fassett,—Echinodorus in the American Tropics 155 


stellate-pubescent like E. muricatus and may be conspecific with 
it. The nutlets are consistently glandless (Fig. 34) with some- 
what branching facial ribs. 


6. EcuiNoponus fluitans Fassett, n. sp., planta aquatica, foliorum 
petiolis longis, laminis 14 em. longis 8 em. latis, ad apicem obtusis, ad 
basim cordatis-subtruncatis; petiolis ad apicem et laminis ad basim 
tuberculosis-muricatis aut cum pilis stellatis; scapis horizontalibus, usque 
ad 1 m. longis; nodis cum radicibus et circa 5 floribus erectis eum pedicellis 
5-8 cm. longis; fructus ignotus.— Colombia: aquatic, growing in about 


Map 5. Echinodorus bracteatus. 


30 em. of water. Flowers white, showy; stem to 2 m. long, bearing 
several inflorescences at intervals, frequently proliferous from these. 
In ponds near Riohacha, Dept. Magdalena, 24 November 1944, Oscar 
Haught 4450 (TYPE in vs).—Seen only from Colombia. 


In habit this most closely resembles Æ. cordifolius, with its 
clusters of long-pedicelled flowers from a prostrate axis. "The 
leaf, however, is quite different, with blades nearly twice as long 
as wide, more pointed at apex and less deeply cordate at base, 
and quite without pellucid markings. 


7. ECHINODORUS BRACTEATUS Micheli in DC., Monogr. Phan, 3: 
59. 1881. 

Leaves erect, long-petioled except on plants from drier places, the 
blades about 24 as broad as long, cordate at base, obtuse at tip, mostly 
about 30 em. long but sometimes more than 50 cm. long (Fig. 12); in- 
florescence branched except in dwarfed plants, sometimes nearly 3 m. 
tall; seape and branches sharply but narrowly 3-winged; flowers sessile 
or nearly so, usually slightly exceeded by the narrow long-acuminate 
bracts; nutlets (Figs. 37-39) rather wide, short-beaked, broadly keeled 


156 Rhodora [Vor. 57 


on the back, with about 5 facial ribs of which usually one is widened 
and wing-like toward the summit; facial gland single or absent. 
Panama to Ecuador (Map 5). 


(To be continued) 


TWO NEW CARICES FROM SOUTHEASTERN 
UNITED STATES 


F. J. HERMANN 


IN southern and central Florida occurs a perplexing sedge 
that keys out to Series Fetae of Section Ovales in Mackenzie’s 
monograph (N. Amer. Flora 18: 118. 1931),—a series with which 
it really has no close affinity. Actually, in all characteristics 
except its ovate-orbicular rather than obovate perigynium, 
it is most closely related to Carex alata Torr. & Gray (Series 
Alatae). But from this species it further differs in lacking 
aristate or long-acuminate scales, the lower being obtuse and the 
upper acute to short-acuminate, and in its longer, more slender 
perigynium beak. The perigynium is usually broadest near 
the middle (occasionally near the base or top) and is less thin 
and flat than that of C. alata. "The shape of the perigynium 
is more suggestive of that of C. brevior (Dewey) Mackenz., but 
from this it is readily distinguished by the narrower, stipitate 
achene, the larger perigynium with much larger corky, crescent- 
shaped areas bordering the achene, and the lack of ventrally 
hyaline leaf-sheaths. This sedge may be known as: 


Carex vexans sp. nov. (Ovales). Caespitosa; culmi folia plus minusve 
superantes; folia ad basim culmi maxima ex parte aggregata, vaginis non 
artis, modo sub ore albidis; squamae lanceolatae, obtusae vel acutae vel 
interdum breviacuminatae; perigynia modice concavo-convexa, crasse mem- 
branacea vel subcoriacea, ovato-orbicularia, manifeste spongiosa, in rostrum 
1-1.5 mm. longum abrupte contracta; achaenia elliptico-oblonga stipitata. 

Cespitose; culms 2.5-7 cm. high, equaling to considerably exceeding the 
leaves, bluntly triangular, scabrous below the heads, otherwise smooth; 
sterile shoots conspicuously developed; lowest leaves bladeless; leaves with 
well-developed blades 2 to 5 to a fertile culm, mostly on the lower half of the 
culm, the blades flat, linear, 4-15 em. long, 2.5-4 mm. wide, scabrous on the 
margins and midrib toward attenuate apex, the sheaths rather loose, white- 
hyaline only below the mouth, rupturing tardily and not becoming fibrillose, 
the ligule from slightly longer to twice as long as wide, 2.5-4 mm. long, 2-3 
mm. wide, blunt; head oblong to ovoid-oblong, 1.5-3 cm. long, 1-1.5 em. wide, 
the spikes 3 to 6, green to glaucous-green, gynaecandrous, aggregated or the 


1955] Hermann,— Two New Carices 157 


lowermost somewhat separate, ovoid-oblong to subglobose, 8-12 mm. long, 
6-9 mm. wide, rounded or abruptly tapering at the base, truncately obtuse 
to rounded-obtuse at the apex, the perigynia ascending, their beaks spreading 
at maturity; bracts setaceous, often lacking except below the lowest spikes; 
scales lanceolate, obtuse to acute or occasionally short-acuminate, shorter 
and much narrower than the perigynia, more or less tawny-hyaline, the 
center green and l-nerved; perigynia shallowly concavo-convex, 4—5.2 mm. 
long, 3-4 mm. wide, thick-membranaceous to subcoriaceous, greenish stram- 
ineous or in age deep brown, glossy, the body ovate-orbicular, broadest near 
the middle or occasionally toward the base or top, conspicuously corky, 
strongly winged to the base, minutely serrulate above the middle, finely 
many-nerved dorsally, few-nerved ventrally, rounded or abruptly tapering 
at the sessile base, abruptly tapering into a beak 1-1.5 mm. long, the beak 
flat, serrulate, dorsally cleft, rather shallowly bidentate; achenes lenticular, 
elliptic-oblong, 1.5-2 mm. long, 1 mm. wide, on a slender stipe 0.75-1 mm. 
long, apiculate, stramineous to brown; stigmas two, slender, 1.5-2 mm. long, 
reddish brown. 


FLORIDA : Collier County—open places, edge of hammock, Deep Lake, 
J. B. McFarlin 4651, March 31, 1931, and marginal woods, Collier West 
Pasture, Sec. 25, T 475, R28E, R. O. Woodbury & R. S. Rummell SE-71, 
June 15, 1954; Hendry County—wet border of roadside ditch along State 
Road 25 about 11.8 miles west of Lewiston, C. C. Deam 58,634, Feb. 2, 
1938 and C. C. Deam 61,177, March 19, 1941 (ryPE—U. S. National 
Herbarium); Lake County—Eustis, G. V. Nash, May 28 to June 15, 1895. 


In 1938 Drs. Rogers MceVaugh and J. H. Pyron collected an 
anomalous sedge near Chatsworth, Gilmer County, Georgia, 
which bears a superficial resemblance to Carex communis Bailey, 
particularly to var. wheeler? Bailey because of its small, few- 
flowered, predominantly sessile staminate spikes. It is at once 
set off from this, however, by its large pistillate scales, these 
being 115 to 2 times the length of, and wider than, the perigynia 
and so partially concealing them. The pistillate scales differ 
further from those of C. communis in being long-acuminate and 
pale-green rather than obtuse or acute (occasionally short- 
acuminate) and reddish purple. The predominantly pale 
staminate scales also differ markedly in having the midrib 
excurrent as a short awn. The short ligule, generally wider 
than long or, exceptionally, as long as wide, is suggestive of 
C. pensylvanica Lam., rather than of C. communis, but the 
lack of stolons and much less fibrillose base of the plant clearly 
set it off from this. 

It was hoped that a postponement of publication of this 
sedge might result in additional localities turning up for it, 


158 Rhodora (Vou. 57 


but to date it remains unique; so it may be a very local species. 
It is here proposed as: 


Carex amplisquama sp. nov. (Montanae). Dense caespitosa, estoloni- 
fera; culmi tenues basibus aliquantum fibrillosis; foliorum ligula plerumque 
latior quam longa; spica terminalis mascula vulgo parva paucifloraque, 
squamis elliptico-oblongis, obtusis, cuspidatis vel aristulatis; spicarum femin- 
earum squamae ovato-lanceolatae, longe acuminatae vel aristatae perigynia 
multo longiores, pallidovirides; perigynia obovoidea 2.75-3 mm. longa. 

Densely cespitose, without long horizontal stolons; culms 1.3-5 dm. high, 
slender, rather sharply triangular and scabrous above, equaling or somewhat 
exceeding the leaves, mostly phyllopodie, somewhat fibrillose and strongly 
purplish-red at the base, the dried leaves of the previous year conspicuous; 
leaves several to a culm, clustered at the base, the blades flat, linear, 5-25 
em. long, 1.5-4 mm. wide, scabrous on the margins and veins toward the 
acuminate apex, the sheaths shallowly concave to slightly convex at the 
mouth, the lower breaking and becoming fibrillose, the ligule wider than 
long (1-1.75 X 1.25-2.5 mm.) or exceptionally slightly longer than wide; 
inflorescence 2-4 (rarely 6) em. long; staminate spike solitary, linear, sessile 
to short-peduncled (the peduncle occasionally up to 1 em. long), generally 
few-flowered and small, 6-12 mm. long, 0.75-2.25 mm. wide, the scales 
elliptic-oblong, obtuse (the lowermost generally retuse) and cuspidate or 
more frequently the midrib excurrent as a short awn, hyaline and greenish 
to stramineous or brown-tinged except for the prominent green midrib; 
pistillate spikes 2 or 3, ovate-orbicular, separate, sessile or the lowermost 
subsessile, 4-6 mm. long, 4-5 mm. wide, containing 2-4 ascending perigynia ; 
lowest bract foliaceous, very short-sheathed, usually equaling or exceeding the 
inflorescence; upper bracts reduced or squamiform; scales ovate-lanceolate, 
long-acuminate, frequently awned, much exceeding the perigynia, greenish- 
hyaline to somewhat brown-tinged except for the broad green midrib; peri- 
gynia obovoid, 2.75-3 mm. long, 1.35-1.5 mm. wide, 2-keeled, puberulent, 
light-green, closely enveloping the achene, obtusely trigonous, abruptly 
contracted into a spongy base and into a bidentate beak 0.5-0.7 mm. long; 
achenes orbicular-obovoid, trigonous with convex sides, yellowish green, 2 
mm. long, 1.5-1.75 mm. wide, abruptly tapering at the base to a stipe 0.5 mm. 
long, the apex short-apiculate; style short, deciduous; stigmas 3, long and 
slender. 

GEORGIA: Gilmer County—dry rocky roadside, 8 miles east of Chats- 
worth, J. H. Pyron and Rogers McVaugh 2951, May 15, 1938 (TYPE 
U. S. National Herbarium).—pLANT INDUSTRY STATION, BELTSVILLE, 
MARYLAND, 


1955] Voigt, —Southern Illinois Flora 159 


SOUTHERN ILLINOIS FLora: RECENT AppitTions.—Field col- 
lections during the past summer (1954) have resulted in the 
addition of several new specimens to the herbarium of Southern 
Illinois University and constitute additions to the known flora 
of Illinois, based on the listings of Jones’ Flora of Illinois (1950). 


A new cup-grass, Eriochloa contracta Hitche., was found in 
considerable abundance on the roadside and low ground near 
the levee of the Pine Hills recreation area of Union County, 
Ill. The 8th edition of Gray’s Manual gives the range of this 
species as follows: La. to N.M., Kansas; adv. in Mo. and Va. 
The specimens were collected Oct. 13, 1954. Specimens have 
been deposited at the herbaria of the University of Illinois, 
The Illinois State Museum, and the U. S. National Herbarium. 
The determination was verified by staff members of the U. S. 
National Herbarium. 

Early in the fall of 1954, à conversation with Dr. L. S. Minck- 
ler, Silviculturist, U. S. Forestry Service, Carbondale Research 
Center, revealed that he had seen what he thought was an 
American Holly tree, [lex opaca Ait., in Union County. On 
Sept. 25, he brought a specimen (not flowering or fruiting) 
to me for my opinion as to its identity. As well as one can 
tell in the absence of flowers or fruits it does seem to be Ilex 
opaca. Dr. Minckler suggests that its location is such that 
it must be naturally occuring in southern Illinois. The place 
of collection is near Alto Pass in Ramberger Hollow. -The 
American Holly is listed in Gray's Manual of Botany, 8th 
edition, for southern Illinois and the neighboring states of 
Kentucky, Missouri, southern. Ohio and southeastward. The 
specimen is deposited in the herbarium of southern Illinois 
University. 

An exploration of land abandoned following coal stripping 
operations in Perry County, on the land of the Truax-Traer 
Coal company about 5 miles south of Pinckneyville, Illinois, has 
resulted in the discovery of a colony of smooth-leaved Silphium 
terebinthinaceum Jacq. The upper surface of the leaf was more 
or less completely smooth except for a narrow fringe around 
the edge of the leaf. The leaves were typically rough-hairy be- 
neath. Correspondence with Dr. E. Lucy Braun for whom 
the smooth-leaved variety was named and subsequent checking 


160 Rhodora [VoL. 57 


of the material by Dr. Braun and by Dr. Julian Steyermark 
reveals that Steyermark considers our material referable to 
Silphium terebinthinaceum Jacq. var. Lucy-Erauniae Steyermark. 
The previously known stations for this taxon are in Adams 
County, Ohio, the area of the type; Rowan County, Ky., Pendle- 
ton County, Ky; Oktibbeha County, Mississippi, and from 
Lord's Park near Elgin, Illinois.! Our specimens were some- 
what smaller in stature than is usual for typical S. terebinthina- 
ceum. The plants were in bud at the time of collection which 
was August 4, 1954. 

One of the most interesting plants discovered the past season 
was Trichachne insularis (L.) Nees, found one-half mile south 
of the village of Cambria in Williamson County. This plant 
was not listed in Gray's Manual, 8th ed., and the range as 
given in Hitcheock’s Manual of the Grasses of the United States 
(1950) is: Florida, Alabama (Mobile), southern Texas, southern 
Arizona; Mexico; West Indies to Argentina. A small clump 
was found by the roadside and in my excitement I pulled all 
of the material for specimens. Unfortunately we shall never 
know if it would have survived there. Judging from the range 
it is very doubtful that it could have survived our winters. 
Owing to the situation then it would not be correct to make 
it an addition to our flora. Its occurrence here is nevertheless 
interesting. The plants were growing well and had produced 
seed. It will be interesting to watch the location for possible 
germination of seed next year. The specimens were deposited 
in the herbaria of the University of Illinois, Illinois State Museum 
and U. S. National Museum. Workers at the last mentioned 
institution verified the determination. The date of collection 


was Oct. 13, 1954. Jon W. VOIGT, so. ILL. UNIVERSITY, CAR- 
BONDALE, ILL. 


Volume 57, no. 676, including pages 105—182, was issued 15 April, 1955. 


1 Steyermark, Julian A., 1951. <A glabrous variety of Silphium terebinthinaceum, 
Ruopora 53: 133-35. 1951. 


Hovova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL ` 
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RALPH CARLETON BEAN 
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CARROLL EMORY WOOD, JR. 


Associate Editors 


Vol. 57 June, 1955 No. 678 


CONTENTS: 


Preliminary Studies in the Genus Dentaria in Eastern North 
America. F. H. Montgometi. -aooiie AN Aa 161 


Echinodorus in the American Tropics (continued). Norman C. 


The New England Botanical Club, Ine. 


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Rbodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 57 June, 1955 No. 678 


PRELIMINARY STUDIES IN THE GENUS DENTARIA 
IN EASTERN NORTH AMERICA* 


F. H. MoNTGOMERY 


THE taxonomy of the eastern North American species of the 
genus Dentaria (Cruciferae) has been a problem to botanists 
for many years. So closely may characteristics of some speci- 
mens in one species resemble those of another species, that the 
task of separating them seems hopeless. Considering the two 
most widely distributed species, Wiegand and Eames (1926) 
remarked “Many plants of Dentaria differ in various ways from 
typical D. diphylla and D. laciniata. These have been a constant 
source of difficulty to local botanists. Field studies through 
many seasons have led the authors to the belief that these 
plants are all of hybrid origin, with D. diphylla and D. laciniata 
as parents." 

Coulter, as far back as 1875, suggested, “These several species 
(D. maxima, heterophylla, laciniata, and multifida) should cer- 
tainly be reduced to one, and some such name as D. heterophylla 
be given to it." Indeed, had he been aware of the variability 
of D. diphylla, he might well have included it with the other 
species. 

The fact that hybrids between nearly all species have been 
suggested also gives an indication of the complex taxonomic 
status of the species. Haberer and House (1923) described three 
hybrids, D. diphylla X maxima Haberer, D. laciniata X maxima 
Haberer, and D. laciniata X diphylla House. The second 
hybrid was considered to be similar to D. incisifolia Eames, 


* Field work in this study was assisted by a Grant from the Research Council of 
Ontario. 


162 Rhodora [Vor. 57 


and the third to correspond to D. anomala Eames. Dr. E. 
Lucy Braun on herbarium specimens and in personal corre- 
spondence has suggested the hybrids D. laciniata X heterophylla, 
and D. multifida X heterophylla. The presence of the parent 
species in the areas where the hybrids were found, as well as 
appearances, was the basis for her determinations. 

Up to the present, the differentiation of species and hybrids 
has been based chiefly on morphology. The data presented 
in this paper are based not only on a morphological study of 
specimens from numerous herbaria and in the field, but also 
include information obtained from cytological, embryological, 
pollen and fertilization studies. 


MATERIALS AND METHODS 
Herbarium material was examined from the following sources: 


Dr. E. Lucy Braun (private collection), Cincinnati, Ohio; Clemson College, 
Clemson, 8. C.; Chicago Museum of Natural History, Chicago, Ill.; Cornell 
University, Ithaca, N. Y.; Division of Botany and Plant Pathology, Ottawa, 
Ont.; Gray Herbarium, Harvard University, Cambridge Mass.; MeMaster 
University, Hamilton, Ont.; Montreal Botanic Garden, Montreal, Que.; 
Missouri Botanical Garden, St. Louis, Mo.; National Herbarium, Washington, 
D. C.; National Museum of Canada, Ottawa, Ont.; New York Botanical 
Garden, New York, N. Y.; North Carolina State College, Raleigh, N. C.; 
University of Cincinnati, Cincinnati, Ohio; University of Georgia, Athens, 
Ga.; University of Iowa, Ames, Iowa; University of Michigan, Ann Arbor, 
Mich.; University of North Carolina, Chapel Hill, N. C.; University of 
Tennessee, Nashville, Tenn.; University of Toronto, Toronto, Ont.; Uni- 
versity of Wisconsin, Madison, Wisc.; University of Western Ontario, London, 
Ont. 


Rhizomes were collected by the writer from Ontario, and 
the states of Alabama, Georgia, New York, Tennessee, and 
Virginia. Dr. Wilbur H. Duncan of the University of Georgia 
very kindly eontributed further materials from Alabama and 
Georgia. Root-tips from these rhizomes were fixed in acetic- 
aleohol, 1:3, stained by the Feulgen technique, mounted in 
aceto-carmine and squashed under the coverslip. Chromosome 
counts and photographs were made before the slides were made 
permanent. 

Pollen for determining the percentage of mature pollen grains 
was obtained for all species, and from areas throughout the 
range of each species. Following the technique of Downs (1943), 
pollen was mounted in polyvinyl alcohol. To give a stain so 


1955] Montgomery,—Studies in the Genus Dentaria 163 


that the pollen grains could be more easily seen, we dissolved 
acid fuchsin in the mounting medium. Mature pollen grains 
stained a deep pink; aborted or empty grains did not take the 
stain. 

Taxonomic NOTES 


O. E. Schulz (1903) monographed the genus Cardamine, and 
in his monograph considered the genus Dentaria to be a section 
cf Cardamine. The combining of the two genera was not with- 
out precedent, for Crantz (1769), Robert Brown (1812), and 
Wood (1870) listed the species of Dentaria under Cardamine. 
Recent floras in America, as well as in Europe, however, have 
continued to separate the two genera. Until both have been 
studied more completely by modern taxonomical methods, no 
conclusive evidence as to the relationship between them can be 
established. 

The first species of Dentaria to be described in eastern North 
America was D. laciniata by Willdenow (1800), apparently from 
letters and collections sent to him by Muhlenberg. The main 
diagnostic characteristic was the three, three-parted leaves. In 
1803, Michaux described as D. concatenata, a species having 
tuberous segments united by a narrow connective, and with 
three, three-parted leaves. This has been made synonymous 
with D. laciniata. This leaf character and the concatenate 
root-stocks are still accepted as the outstanding features of the 
species. The rhizomal leaf (if present) and the flowering stem 
arise from the end of the rhizome segment of the previous year. 
The involucral leaves may be two instead of the typical three. 
They may be glabrous or pubescent. The peduncles and 
pedicels most frequently do have pubescence. The trichomes 
are comparatively long and more or less perpendicular to the 
leaf, peduncle and pedicel surface. In this respect, they resemble 
the trichomes of D. maxima. 

Although all species are polymorphic, in no other has varieties 
been described. When the leaves are two, they may be opposite, 
var. opposita Farwell (1931). Typically the leaves are in whorls 
but they may be alternate as in var. alterna Farwell (l.c.). As 
in the original description the leaflets are basically three-parted, 
but the lateral leaflets may be again divided, giving a five-parted 
appearance. Several varietal names have been given to varia- 


164 Rhodora [Vor. 57 


tions in those specimens having simply three-parted leaflets, 
e.g. var. furcata Small, (1903), var. integra (O. E. Schulz) Fernald, 
(1908), var. latifolia Farwell (l.c.). Variety coalescens Fern. 
(1938) has the leaves merely three-cleft or pinnatifid. Schulz 
(1903) described a variety with the pubescence also covering 
the silique, var. lasiocarpa. 

Dentaria laciniata has a wide distribution (fig. 4). It extends 
farther westward and southward than any other species, but 
no records have been seen for Maine and the Maritime Provinces 
of Canada. Only in southern Ontario has it been found north 
of the Great Lakes. 

Michaux (1803) described D. diphylla as the Dentaria with 
dentate roots, with two stem-leaves approximate and three- 
parted. The rather stout, continuous, dentate rhizomes, on or 
close to the surface of the ground, are the best distinguishing 
feature of the species. The basal or rhizome-leaf, and the 
flowering stem arise from the same point at the end of the 
rhizome, but the new underground growth forms so quickly 
in the spring that these may appear to originate from the surface 
of the rhizome. In all other species the above ground parts 
appear to come from the constricted portion of the rhizome— 
actually the tip of the former year’s growth. The two, three, 
or more involucral leaves may be opposite, sub-opposite, or 
alternate. These leaves are three-parted. The very short, 
forward-pointing trichomes on the margins of the leaflets may 
be used to distinguish this species from D. laciniata, maxima 
and multifida; however, D. heterophylla has similar marginal 
trichomes. The peduncles and pedicels are nearly always 
glabrous. The number, size, and color of the flowers varies 
considerably, and no constant character has been noted. 

Dentaria diphylla covers most of the range of the genus (fig. 3) 
but does not extend as far west as D. laciniata. It is, however, 
found farther northward and eastward than D. laciniata. 

Dentaria multifida was catalogued by Muhlenberg (1813) but 
was first described by Nuttall (1818). It is the most delicate 
of our species. It has very small, concatenate rhizomes with 
the segments rather firmly united. The leaves are basically 
similar to D. laciniata, being three-parted. "The primary divi- 
sions are further divided into linear segments. For this reason 


1955] Montgomery,—Studies in the Genus Dentaria 165 


DISTRIBUTION OF EASTERN NORTH AMERICAN SPECIES OF Dentaria. Fig. 1. D. 
multifida; Fig. 2. D. heterophylla; Fig. 3. D. diphylla; Fig. 4. D. laciniata; Fig. 5. D. 
maxima; Fig. 6. Composite map of distribution areas. 


Leavenworth (1824) described D. dissecta from a specimen from 
Cherokee Co., Tenn. This has been placed in synonymy. No 
pubescent specimens have been seen. Some specimens of D. 
multifida may approach D. laciniata in appearance, and vice- 
versa. Schulz (l.c.) treated D. multifida as a variety of D. lacini- 
ata and said that it was bound with the main species through 
a continuous series of intermediates. 

D. multifida has a restricted distribution (fig. 1). Its area 
of greatest frequency is Tennessee, northern Georgia and northern 
Alabama. 


166 Rhodora [Vor. 57 


Nuttall’s description of D. heterophylla (1818) might well 
apply to D. diphylla, except for the concatenate rhizome. In 
D. heterophylla the segments are short, slender and rather firmly 
united by a narrow connective. The size of the segments 
distinguishes them from those of D. laciniata. In examining 
specimens it was frequently necessary to observe the rhizomes 
in order to separate D. heterophylla and D. diphylla. 

The area of D. heterophylla (fig. 2) 1 much more restricted 
than that of diphylla and (aciniata, but wider than that of 
multifida. It has migrated into southern Indiana, Ohio and 
Pennsylvania. 


Dentaria maxima was rather thoroughly described by Nuttall 
(le.). His description of the rhizomes requires some clarifica- 
tion. Nuttall described them as “concatenate” but this would 
not distinguish them from those of D. laciniata, D. heterophylla 
and D. multifida. The rhizomes of Dentaria mazima closely 
resemble those of D. diphylla, but are distinguished by alternately 
enlarged and only slightly constricted regions. Also, as to the 
size and number of leaves and leaflets, the specimens that Nuttall 
described must have been rather singular. “Stem often near 
2 ft. high,—leaves 5 to 7." In the specimens examined, stems 
seldom exceeded 1 ft. in height, and most frequently there 
were 3 alternate leaves. However specimens with one, two, 
or several leaves are to be found. The leaflets are usually 
more sharply toothed than those of D. diphylla. The leaf 
characters vary, and in some areas they approach those of 
diphylla. ln other localities, e.g. Taughannoek Falls and 
Enfield Glen, New York, and Milton and Kitchener, Ontario, 
the leaves are difficult to distinguish from those of laciniata. 
Nuttall mentioned the slightly asperate margin of the leaf. 
The trichomes there were found to be longer and more nearly 
perpendieular to the margin of the leaf than the trichomes of 
diphylla. ‘They are superficially similar to those of laciniata. 
The peduncles are most frequently glabrous, but pubescent 
specimens have been seen. 

D. maxima occurs entirely within the area of the Wisconsin 
glaciation (fig. 5), and its main distribution is in New York 
State, and the Provinces of Ontario and Quebec. 


Rhodora Plate 1206 


CHROMOSOMES OF DenTARIA Species (X 2000). Fig. 7. D. diphylla, 2n = 96; 
Fig. 8. D. multifida, 2n. = + 112; Fig. 9. D. heterophylla, 2n. = + 128; Fig. 10. D. 
laciniata, 2n. = + 240; Fig. 11. D. marima, 2n = + 208; Fig. 12. D. multifida from 
Rome, Ga., 2n — 64. 


1955] Montgomery,—Studies in the Genus Dentaria 167 


CyTOLOGY 

The chromosomes in all species are small and numerous. 
This latter fact made it difficult always to obtain complete 
separation of the chromosomes. Their size prohibited a study 
of chromosome morphology, but most of them seemed to have 
a median centromere. Some had a sub-terminal centromere 
which was rather elongated and often made it uncertain whether 
one should count one or two chromosomes. 

Table 1 gives the chromosome numbers for the species studied. 


TABLE 1. CHROMOSOME NUMBERS FOR EASTERN NORTH AMERICAN 
SPECIES OF DENTARIA 


No. of cell 2n 
Species Source of material counts no. 
D. diphylla Kitchener, Guelph, Ont. 21 96 
(Fig. 7) 
D. heterophylla Markham, Va.; Rome, Ga.; Winston 
(Fig. 9) Co. Ala.; Johnston City, Tenn.; 42 +128 
D. laciniata Kitchener, Guelph, Ont. 15 +240 
(Fig. 10) 
D. maxima Guelph, Milton, Ont.; Enfield Glen, 
(Fig. 11) N. Y.; Taughannock Falls, N. Y. 15 +208 
D. multifida Vilanow, Ga.; Chattanooga, Tenn. 10 +112 
(Fig. 8) 
(Fig. 12) Rome, Ga. 5 64 


Chromosome studies by other workers have shown that most 
species of Dentaria studied from other parts of the world are 
also polyploids. Those species for which chromosome counts 
have been made are as follows: 


TABLE 2. CHROMOSOME NUMBERS OF SPECIES OUTSIDE OF 
EASTERN NoRTH AMERICA 


Species 2n Chromosome No. Author 

D. polyphylla 48 Schwarzenbach, 1922 

D. pentaphylla 48 Schwarzenbach, 1922 

D. pinnata 48 Schwarzenbach, 1922 

D. bulbifera ca 96 Manton, 1932 

D. californica 32 Manton, 1932 

D. savensis ca 80 Manton, 1932 

D. macrophylla 16 Morinaga & Fukushima, 1931 


With the exception of D. californica these are European and 
Asiatic species. 

The basic chromosome number has been given as x = 8 by 
Darlington and Janaki Ammal (1945). This number is certainly 


168 Rhodora [Vor. 57 


basic to the American species studied here, but the 2n number 
exceeds those found so far in other parts of the world. 

It is the writer's opinion that the chromosome numbers given 
in Table 1 are not all that may be obtained for specimens in 
our area. The fact that one specimen of D. multifida from Rome, 
Ga., had 2n = 64 would give credence to this opinion; also the 
origin of our present species would indicate that other chromo- 
some numbers may be expected, if different populations are 
examined. 

Some verification of chromosome number relationships was 
sought in stomata number per sq. mm. This method was first 
suggested by K. and H. J. Sax (1937). Babcock and Stebbins 
(1938) used stomatal size as a criterion to indicate chromosome 
number in Crepis. In Dentaria, neither method proved to be 
completely satisfactory. However, in three species, D. diphylla, 
D. laciniata, and D. maxima, stomatal counts per sq. mm. 
could be used with some degree of accuracy. D. diphylla 
averaged 92 stomata, D. laciniata 52 stomata, and D. maxima 
65 stomata per sq. mm. It was not possible to distinguish 
D. heterophylla and D. multifida by this technique. 


STERILITY IN THE GENUS DENTARIA 

Sterility in some European species, particularly Dentaria 
bulbifera, has been the subject of several papers, Ernst, (1918), 
Gams, (1922), Schwartzenbach, (1922), Leopold (1928), and 
Fritsch, (1922). Eames (1903) remarked on the sterility of 
D. diphylla, and the new edition of Gray's Manual notes that 
siliques are rarely matured in D. diphylla and D. maxima. 
While studying herbarium specimens, observation of those 
specimens that had fruiting material showed that although 
pods were developed, few, if any, developed mature seed. Studies 
in the field led to the same conclusion. Pods may develop to a 
good size, but when the contents are examined, they will most 
frequently be found to contain aborted ovules. Efforts to get ap- 
parently mature seeds of D. diphylla, D. laciniata and D. maxima 
from Ontario to germinate were entirely unsuccessful. 

Causes of sterility were sought in pollen and ovules. Anthers 
of all species were observed for aborted pollen. In all, 70 speci- 
mens from throughout the area of the genus were examined, 


1955] Montgomery,—Studies in the Genus Dentaria 169 


and with few exceptions the percentage of normal appearing 
pollen was high. Even in those specimens that were considered 
by the collector to be hybrids, good pollen was usually the rule. 
It was evident that sterility could not be attributed to poor 
pollen. 

This led to the questions: Were the flowers being pollinated? 
Is self- or cross-pollination required for production of seeds? 
To obtain some answers to these questions 50 flowers from 
each of the species D. diphylla, D. maxima, and D. laciniata 
were self-pollinated, and 50 flowers of each were cross-pollinated 
with pollen from what were considered to be different clones. 
Not one seed was obtained from these pollinations. This 
would confirm observations that these three species are highly 
sterile. 

Sections of ovules of the same three species were studied, 
and it was observed that shortly after the eight-celled stage 
was reached in the embryo sac, disintegration of the cells of the 
embryo sac took place and the ovules finally aborted and died. 
'The eastern North American species of Dentaria proved to be 
another example of a polyploid series being sexually sterile. 
Undoubtedly they reproduce mostly, if not entirely, apomictically 
by vegetative reproduction, for which the nature of the rhizome 
seems admirably suited. In this respect they resemble D. 
bulbifera as discussed by Ernst (l.c.), D. glandulosa by Schulz 
(l.e.). 

DISCUSSION 

This study of the eastern North American species of Dentaria 
indicates that there is great difficulty in determining the species 
by morphology alone. Numerous plants with intermediate 
characters appear and there is an intergradation of characters 
in most of the species. Cytological investigations indicate 
that the species in the area being considered have high poly- 
ploidy, 2n = 64, 96, 112, 128, 208, and 240, and these numbers 
support the separation of the species. Normal pollen, with 
few exceptions, throughout the range of the genus is high. 
It has been found, in at least three species, that disintegration 
of the contents of the embryo sacs takes place shortly after 
they have reached the 8-celled stage. The same three species 
are sexually sterile and reproduce vegetatively by rapid develop- 


170 Rhodora [Vor. 57 


ment of the rhizomes, partioulariy in in the early spring and during 
the time of flowering. 

It is apparent that we have in this group of plants, an agamie 
complex similar to that described by Babcock and Stebbins 
(l.c.) in western American members of the genus Crepis. How- 
ever, we have not yet discovered the diploid ancestors of this 
eastern North American complex. Probably they have been 
overlooked, or, they may no longer be in existence. As Stebbins 
(1941) has pointed out, “The systematic structure of an agamic 
complex can be understood only when its diploid sexual members 
are all known, and the limits of their variability have been 
determined." We are, therefore, compelled to discuss our 
representatives from the data that are available. 

It is well known that in genera such as Taraxacum, Hieracium, 
and Crepis certain groups are highly polymorphic. This poly- 
morphism may be attributed to hybridization, polyploidy, and 
apomixis. Stebbins (l.c.) has pictured the development of the 
agamic complex, and Dentaria seems to fit well into this picture. 

The diploid species, which were well defined systematically, 
by hybridization and polyploidy produced many new forms 
which because of their adaptability were able to invade habitats 
unsuited to their diploid parents. They may even have crowded 
out the diploid parents, if they ever existed on this continent. 
The numerous variations that were produced were finally reduced 
by the process of natural selection to the few remnants which 
represent our present species, and the many intermediates 
that cause so much taxonomic difficult y. 

Gustafson (1947) showed that hybridization and polyploidy 
contribute to the production of apomicts. Hybridization may 
bring together genes responsible for sexual sterility, and poly- 
ploidy builds up the numbers of these genes, and may also 
alter the physiological condition of the cells so that the sterility 
faetors may have full expression. In our highly polyploid 
members of the genus Dentaria, there has been a great accumula- 
tion of these sterility factors, and the genus has reached the 
stage of evolution where there is complete sexual sterility. The 
genus, therefore, depends for its existence on vegetative repro- 
duction. According to Stebbins this is the last stage in the 
evolution of apomicts. Having lost their sexuality, they can 
produce nothing new and eventually die out. 


1955] Montgomery,—Studies in the Genus Dentaria 171 


The distribution of Dentaria maxima north of the southern 
limits of the glaciation is an interesting phytogeographical 
problem. That it is a hybrid between two of our species, D. 
diphylla and D. laciniata, as suggested by Wiegand and Eames, 
is excluded for several reasons. "These two species are usually 
found in dry mesophytic woodlands, but D. mazima has been 
found most frequently in low, moist, wooded areas or on moist, 
wooded hillsides. The chromosome number, 2n = + 208 does 
not indicate a hybrid origin with these two species as parents. 
The sexual sterility of D. diphylla and D. laciniata also argues 
against its being a direct hybrid between these two species; and 
for similar reasons it seems highly improbable that it can be a 
hybrid between any species of Dentaria now known in eastern 
America. 

It would seem then, that D. maxima is a member of the agamic 
complex that originated before complete sexual sterility was 
effected in the members of the complex. It may have been 
derived during some interglacial period of the Pleistocene 
glaciation. During the Wisconsin glaciation the species survived 
near the border of the ice sheet, and with the return of temperate 
conditions found the glaciated soils a more suitable habitat 
than the Tertiary soils to the south. 


CONCLUSIONS 


1. The species of the genus Dentaria in eastern North America 
are highly polymorphie, and sharp morphological lines of 
distinction between species cannot be drawn. 

2. The species D. diphylla, D. multifida, D. heterophylla, D. max- 
ima, and D. laciniata can be distinguished cytologically on 
the basis of chromosome numbers. All species are high 
polyploids. 

3. Sexual sterility is known to occur in three species, D. diphylla, 
D. maxima and D. laciniata. Examination of specimens of D. 
heterophylla and D. multifida indicates that sterility also 
occurs in these species. Sterility has been shown to be 
the result of incomplete development in the embryo sac. 

4. The genus is considered to be of ancient origin, and the poly- 
morphic character of the species is the result of hybridization, 
polyploidy and apomixis. 


172 Rhodora [Vor. 57 


5. Dentaria maxima occurs only in the glaciated regions of 
eastern North Ameriea, and is thought to have originated 
during some interglacial period. After the Wisconsin glacia- 
tion it migrated north into the glaciated areas rather than 
south into the tertiary soils. 


KEY ro THE EASTERN NORTH AMERICAN SPECIES OF DENTARIA 


a. Rhizome not segmented or with alternately enlarged and slightly 

constricted regions; rachis glabrous (rarely pubescent). 
b. Diameter of rhizome constant along its entire length; cauline leaves 
mostly 2, opposite or alternate; trichomes on margins of leaflets 0.1 
mm. long and appressed. ..............00 000000200 D. diphylla. 
b. Rhizome with alternately enlarged and slightly constricted regions; 
number of cauline leaves and arrangement variable but most fre- 
quently 3 and alternate; trichomes on margins of leaflets 0.2-0.3 mm. 
long and spreading... ..........0.0 eee D. maxima. 
a. Rhizome definitely formed of segments united by fragile connectives. 
c. Rachis pubescent (rarely glabrous); trichomes of rachis and margins 
of leaflets 0.2-0.3 mm. long and spreading............ D. laciniata. 

e. Rachis glabrous (rarely pubescent). 

d. Cauline leaves usually 2, opposite or alternate; leaflets 3; trichomes 
on margins of leaflets 0.1 mm. long and appressed. . D. heterophylla. 
d. Cauline leaves most frequently 2; leaflets dissected into 3-7 
linear segments 1-3 mm. broad; margins of leaflets without tri- 
Chomes... 0... 0.6 ees D. multifida. 


ACKNOWLEDGMENTS 


My sincere thanks are extended to the Curators of all the 
herbaria from which study material was received; to Dr. L. O. 
Gaiser for her advice in the preparation of the manuscript; to 
Dr. Reed C. Rollins for his encouragement during the progress 
of the work, and to many others who have contributed materials 
and information without which the study could not have been 
completed.—DEPARTMENT OF BOTANY, ONTARIO AGRICULTURAL COLLEGE, 
GUELPH, ONTARIO. 

LITERATURE CITED 

Bascock, E. B. and G. L. STEBBINS, JR. 1938. The American species 
of Crepis. Carneg. Inst. Wash. Pub. No. 504. 

Brown, Rosperr 1812. Genera et species plantarum. In Aiton Hortus 
Kewensis. Ed. 2. 4:101. ` 

COULTER, J. 1875. Botany Bulletin 1: 8. 

Crantz, H. J. 1769. Classis Cruciformium. p. 66. 

Daruneton, C. D. and E. K. JanaKı AMMar, 1945. Chromosome 
atlas of cultivated plants. George Allen & Unwin, Ltd. London. 


1955] Montgomery,—Studies in the Genus Dentaria 173 


Downs, W. G. 1943. Polyvinyl alcohol—a medium for mounting and 
clearing biological specimens. Science 97: 539-40. 

Eames, E. H. 1903. The Dentarias of Connecticut. RHODORA 5: 202. 

Ernst, A. 1918. Bastardierung als Ursache der Apogamie im Pflanzen- 
reich. Jena. 

FARWELL, O. A. 1931. Botanical gleanings in Michigan, VII. Amer. 
Mid. Nat. 12: 58. 

FERNALD, M. L. 1908. Notes on some plants of northeastern America. 
Ruopona 10: 84. 

. 1938. Noteworthy plants of southeastern Virginia. Rmo- 
DORA 40: 421. 

. 1950. Gray’s manual of botany, ed. 8. American Book 
Co. N. Y. 

Fritscu, K. 1922. Ist Cardamine bulbifera als Abkómmling eines 
Bastardes aufzufassen? Ber. Deutsch. Bot. Ges. 40: 193-195. 

Gams, H. 1922. Noch einamal über die Herkunft von Cardamine 
bulbifera (L.) Crantz und Bermerkungen über sonstige halb- und 
ganzwaisen. Ber. Deutsch. Bot. Ges. 40: 362-367. 

GUSTAFSSON, AKE. 1947. Apomixis in higher plants. Lunds Univer- 
sitets Ársskr. N. F. 43: Nr. 2; 70-349. 

HovsEe, H. D. 1923. Report of the State Botanist for 1921. N. Y. 
State Mus. Bull. 243-244: 44. 

LEAVENWoRTH, M. C. 1824. Notice of four new species collected in 
Alabama. Amer. Jour. Sci. 7: 62. 

LroroLp, W. 1928. Beiträge zur Kenntnis der Gattung Cardamine 
mit besonderer Berücksichtigung der hybridenfrage in der Sektion 
Dentaria. Acad. d. Wiss. in Wien Mat. Nat. Klasse. 101: 325-360. 

MicHaUx, A. 1803. Flora Boreali-Americana 2: 30. 

MUHLENBERG, H. 1813. Catalogus Plantarum Americae Septentrion- 
alis, 63: 60. 

NUTTALL, T. 1818. The genera of North American plants and a 
catalogue of the species to year 1817. 2: 66. 

Sax, K. and H. J. Sax. 1937. Stomata size and distribution in diploid 
and polyploid plants. Jour. Arnold Arboretum, 18: 164-172. 

ScuurLZ, O. E. 1903. Monographie der Gattung Cardamine. Bot. 
Jahrb. 32: 280-623. 

ScHWARTZENBACH, F. 1922. Untersuchungen über die sterilitàt von 
Cardamine bulbifera (L.) Crantz. Flora 115: 393-514. 

SMALL, J. K. 1903. Manual of the southeastern flora. Published 
by the author. New York. 

STEBBINS, G. L. Jr. 1941. Apomixis in Angiosperms. Bot. Rev. 7: 
507—542. 

WiEGAND, K. M. & A. J. Eames, 1926. The flora of the Cayuga Lake 
Basin, New York. Cornell Univ. Agri. Expt. Sta. Mem. 92. 

WinrpENow, C. L. 1800. Species Plantarum. III: 479. 

Woop, A. 1870. American Botanist & Florist. Part 4, p. 37. A.S. 
Barnes & Co., N. Y. 


174 Rhodora [Vor. 57 


ECHINODORUS IN THE AMERICAN TROPICS 


NonMAN C. FASSETT 


(Continued from p. 156) 


7a. E. BRACTEATUS Micheli var. BRAcTEATUS. Blades with stellate 
tubereulate-based hairs toward the base, and with pellucid dots and 
very short lines (Fig. 60). 

Panama: around El Paraiso, Canal Zone, alt. 30-100 m., 24 January 
1911, Pittier 2531 (GH, Ny, vs); Gatun, November 1859, Hayes 130 
(GH); open swamp, between France Field, Canal Zone, and Catival, 
Prov. Colón, 9 January 1924, Standley 30444 (vs); wet forest, near 
the big swamp east of the Río Tecumen, Prov. Panamá, 11 December 


Map 6. Echinodorus ovalis. 


1923, Standley 26724 (vs); Mindi, 28 February 1905, Cowell 177 (NY); 
Colón, Monkshill, 5 June 1874, Kuntze (Ny). Ecuador: in water of 
swamp and mud, Prov. Guayas, Oil Camp between Guayaquil and 
Salinas, alt. 0-100 m., 21-24 June 1923, Hitchcock 20084 (vs, Ny; sheet 
in GH is the variety below); in paludosis, Balao, January 1892, Eggers 
14216 (us); moist sandy creek bed, alt. 40 m., 20 km. southweast of 
Guayaquil, 29 August 1938, Worth, Morrison & Horton 8935 (vs). 

7b. E. BRACTEATUS Micheli, var. efenestratus Fassett, n. var., foli- 
orum laminis glabris, punctis lineisve pellucidis destitutis.—Leaf blades 
glabrous and without pellucid markings.—Ecuador: coast plain, on 
marshy ground, near water. Stem 214 m. high. Flowers white. A. 
Rimbach 90 (Tyrer in F); coast plain, in swampy ground or open places. 
Perennial herb, 215 m. high. Leaf blade as much as 40 em. long and 
30 em. broad. Flowers white. July. A. Rimbach 287 (us); swamp, 
in water, and in mud along edge, Prov. Guayas, Oil Camp between 
Guayaquil and Salinas, 21-24 June 1923, Hitchcock 20084 (an). 

8. EcuiNoboRUS ovaris Wright in Sauvalle, Fl. Cub. 564. 1871; 
Buch. in Engler, Pflanzenr. 4, fam. 15; 31. 1903; Small, N. Am. FI. 17: 
pt. 1: 48. 1909. 

I have seen but two collections of this little-known Cuban species. 
The pellucid marking of the blades (Fig. 61) is much like that of E. cordi- 


1955] Fassett,—Echinodorus in the American Tropics 175 


folius (Fig. 54), but the pellucid lines are more abundant and the venation 
appears to be a little finer and more complex. The scape, also, is much 
like that of E. cordifolius and looks as if it might become prostrate as in 
that species. The sepals of E. ovalis, however, have smooth ribs quite 
unlike the muricate ribs of Æ. cordifolius. An immature nutlet is shown 
in Fig. 40; it looks as if it would be short beaked, with several ribs and 
facial glands. 

Western Cuba (Map 6). Cuba: Wright, without data (vs); Wright 
3713, without other data (aH); pond west of Batabanó, Habana, 8 Novem- 
ber 1928, León 13707 (Gu). E. ovalis is listed by Buchenau from Pinar 
del Rio and Hato Salado, and these localities have also been plotted on 
Map 6. 

9. ECHINODORUS ANpRIEUXI (Hook. & Arn.) Small, N. Am. Fl. 17, 
pt. 1: 46. 1909. Alisma Andrieuxi Hook. & Arn., Bot. Beech. Voy. 
311. 1839. Echinodorus ellipticus y, ovata Micheli in DC., Monogr. 
Phan. 3: 52. 1881, not E. ellipticus var. ovatus Hauman, Anal. Mus. 
Nac. Buenos Aires 27: 309. 1925. E. ellipticus f. ovata F. J. Meyer, 
Bot. Centralbl. Beihefte 49 (1): 331. 1932. 

Leaves erect, long-petioled; blades narrowly elliptic (Fig. 14), to 
ovate (Fig. 15), mostly tapered to truncate at base, usually 15-20 em. 
long but scmetimes reaching 30 cm. or more, usually with very obscure 
pellucid lines; petiole 2-winged; inflorescence simple and wand-like or 
rarely with short weak branches at base, with many rather compact 
verticels; bracts slightly connate at base, long-acuminate at tip; pedicels 
from nearly lacking to 2 em. in length; nutlets with body about l% as 
wide as long, 4-7-ribbed, with usually one facial gland (Figs. 41-43); 
beak about L4 as long as the body. 

Western Mexico to El Salvador (Map 7), and Niearagua according 
to Micheli. Sinaloa: Bellavista, Mazatlán, March 1931, Gonzales 
6846 (r, vs); Río Piaxtla, San Ignacio, alt. 20 m., 13 March 1918, Nar- 
vaez 199 (us); Mazatlán, 6 April 1910, Rose 14100 (Ny, vs); moist field, 
Villa Union, 2 April 1910, Rose 13963 (vs); Balboa, January 1923, Ortega 
4994 (vs). Nayarit: Tepic, Beechey—TrYPE (drawing in Ny; fragment 
in Mo); Tepic, 5 January—6 February 1892. Palmer 1844 (F, GH, NY, 
vs); Tepic, 12 February 1927, Jones (Mo). Jalisco: in rain pools, forms 
dense clumps, 1 m., fl. white, alt. 20 m., Tuxpan, Palapar Redondo, 
6 November 1926, Mexia 1065 (F, GH, Mo, NY, US). Michoacán: Tacupa, 
Huetamo, 22 March 1934, Hinton 5817 (Gu, Ny). Morelos: swamps 
near Cuernavaca, 5000 ft., 29 September 1899, Pringle 8256 (F, GH, 
MO, PH, NY, US); Tetecala, 7 October 1890, Maury 4922 (Ny). Guerrero: 
swamp, Acapulco, February 1895, Palmer 530 (us). Oaxaca: Tem- 
ascaltepec, 21 November 1932, Hinton 2667 (Gu, us); Juquila, 24 Decem- 
ber 1921, Conzatti 4512 (us); Tehuantepec, Andrieux 91 (drawing in 
ny; fragment in Mo). Chiapas: in marshy lakeside, 2 m. high, Aca- 
coyagna, 28 September 1947, Matuda 17014 (r). Guatemala: Dept. 
Chiquimula, valley of Río Chiquimula, 114 mi. northeast of Chiquimula, 
alt. 400 m., 21 October 1939, Steyermark 30150 (F); Dept. Jutiapa, in 


176 Rhodora [Vor. 57 


Map 7. Echinodorus Andrieuxi. 


marsh on plain, between Jutiapa and La Calera, southeast of Jutiapa, 
alt. about 850 m., 2 November 1940, Standley 76103 (v); Dept. Jutiapa, 
open marsh, between Jutiapa and La Burrere, northeast of Jutiapa, 
alt. 800-850 m., 1 November 1940, Standley 75999 (r). British Hon- 
duras: Corozal district, Gentle 373 (vs). El Salvador: Dept. Santa 
Ana, shallow open pool, south of Texistepeque, 14 km. north of Santa 
Ana, alt. 1266 ft., 13 October 1950, Fassett 28281 (F). 


In the herbarium of the New York Botanical Garden are two 
beautifully detailed pencil drawings of specimens in the Kew 
Herbarium. One is Andrieux 91, “circa Tehuantepec, in ditione 
Oaxacana, Septembr." The other is the Beechey collection from 
Mexico. Both are marked as rype. “When no habitat is 
mentioned, the specimens are understood to have been collected 
at Tepic" (Hook. & Arn., p. 275), so the Beechey collection 
presumably eame from there. Following the description of 
Alisma Andrieuxi is a citation of Andrieux 91, which would at 
first sight appear to be the type. But the next paragraph, 
stating that “the same species was found by M. Andrieux about 
Tehuantepec of Oaxaca," distinctly implies that the species is 
primarily based on the Beechey collection from Tepic, and that 
M. Andrieux also collected the same species. The Beechey 
collection should therefore be taken as type. 


1955] Fassett,—Echinodorus in the American Tropics 177 


Micheli places E. Andriewxi under E. ellipticus as y, ovata, 
citing collections from Mexico, Nicaragua and Brazil, but I have 
seen nothing from South America that appears conspecific with 
the Mexican plants. Macbride, Field Mus. Pub. Bot. Ser. 11: 
4. 1931, states that E. palaefolius (Nees. & Mart.) Macbr. (= E. 
ellipticus (Mart.) Micheli) occurs in Mexico, probably basing this 
statement on that of Micheli. Actually, E. ellipticus and £E. 
palaefolius are both described as having leaves with deeply 
cordate blades and angled petioles, while Æ. ellipticus y, ovata 
(= E. Andrieuzt) is distinguished by having blades decurrent on 
the petiole. Its relationships are, then, rather with the Æ. 
paniculatus—E. subalatus complex. 

Micheli described the leaves of W. ellipticus, including Æ. 
Andrieuxi, as having “lineis pellucidis longis, raris, saepe fere 
inconspicuis." He was quite correct, but I find myself wondering 
just how he managed to see the lines. My own apparatus is 
without doubt better than any he had: my work-table has a 
21% inch square hole cut in the top, with a 60-watt bulb in the 
drawer below, shining up through the herbarium sheet into a 
modern binocular microscope. Only by waiting until after dark 
and turning off all overhead lights was I able to see dimly the 
pellucid lines in E. Andrieuri. Micheli was right—they are 
therein most individuals. He must have been a very painstaking 
and thorough observer. Recently I have had opportunity to 
examine living plants in El Salvador; the pellucid lines are 
nearly invisible. 


10. ECHINODORUS PANICULATUS Micheli in DC., Monogr. Phan. 3: 
51. 1881. 

Blades varying from elliptic (Fig. 18) or ovate (Fig. 17) to linear- 
lanceolate (Figs. 19, 20), broadly cuneate or truncate to narrowly cuneate 
at base, acute at tip, sometimes reaching 30 em. in length; petiole with 
a distinct wing on each side; inflorescence sometimes simple, but often 
with branches 20 cm. long; flowers long-pedicelled; nutlets (Figs. 44— 
49) very flat, mostly more than half as wide as long, with several narrow 
wing-like ribs, broadly keeled dorsally and ventrally, without facial 
glands, the beak short and stout or sometimes nearly absent. 

10a. E. PANICULATUS Micheli var. PANICULATUS. Leaves ovate to 
linear-lanceolate, with pellucid lines very obscure or absent, and without 
pubescence. Colombia and Venezuela to Peru, Paraguay and southern 
Brazil (Map 8). Colombia: Santa Marta, 1899-1901, Smith 2326 (F, 
GH, MO, NY, PH, US); Baranquilla, along shores of the Magdalena River, 
June, 1927, Elias 202 (us); La Rubiera near Calabozo, Guárico, Grisol 1— 


178 Rhodora [Vor. 57 


Map 8. Echinodorus paniculatus. 


mixed with E. muricatus (Ny, vs). Venezuela: Sanare, 1358 m., Lara, 
May 1930, Saer 489 (r); Hato de Matapalo, in swamp, Llanos del Alto- 
Apure, alt. 150 m., 7 May 1911, Jahn 206a (us); Dist. Fed., Laguna 
Conejo Blanco, El Valle, 900 m., 17 August 1941, Fernandez 71 (us). 
Ecuador: dried up pond, Province Guayas, Milagro, alt. 50 m., 30 June— 
2 July 1923, Hitchcock 20278 (GH, Ny, vs). Peru: Alto Río Huallaga, 
alt. 360-900 m., Dept. San Martin, Williams 5815 (r, vs). Bolivia: 
Santa Cruz, Chiquitos, Cardenas 4466 (us); pampas near Lake Rogagua, 
alt. 1000 ft., 4 November 1921, Rusby 1649 (ny); Potero Largo in from 
Velasco, 200 m., July 1892, Kuntze (NY). Brazil: Prov. Piaui, Gardner 
2741 (coryPE—Us); Province de Rio-Janeiro, Glaziou 13221 (r; photo 
ex Museo Botanico Berolinense, r, GH, us); in open field, low land, flooded 
in rainy season, region of River Mearin, State of Maranhao, 9 March 
1933, Krukoff 2030 (GH, Ny, us); Minas Geraes, 16 December 1864, 
Regnell 418 (us); Matto Grosso, 1891, 92, Moore 969 (Ny). Paraguay: 
Pilcomayo River, Morong 1039 (xv, pH). 


The nutlets are unmistakable (Figs. 44-49) and one can be 


reasonably sure of the conspecificity of fruiting individuals, 
despite variation in leaf-width and type of inflorescence. Micheli 


1955] Fassett,—Echinodorus in the American Tropics 179 


described the nutlets as being multiglandular, but he had only 
young fruits which sometimes have a gummy appearance that 
falsely promises later development of glands. 

I have placed in this group collections of heterogeneous aspect, 
and cannot be certain that a single entity is represented. There 
is variation in stamen number, from 9 to 20 or more. Two 
sheets show leaves slightly pseudopenninerved, while the rest 
are nerved from the base of the blade. Some plants have ob- 
scure pellucid lines in the leaves, and others no lines at all. 

Of course, with such uncertainty concerning the taxonomic 
status of the sheets cited above, application of the name cannot 
be at all certain. I find great difficulty in placing some names 
used by Martius in R. & S. Syst. Veg., and Martius himself 
expresses some doubts as to such names as Alisma ellipticum, 
A. intermedium and A. subalatum. Similar difficulties appear 
with Micheli's treatment of Echinodorus Martii, E. intermedius, 
E. paniculatus, E. ellipticus and E. subalatus. 


10b. E. PANICULATUS Micheli, var. dubius Fassett, n. var., foliis laminis 
ovatis, ad basim obtusis vel truncatis, coriaceis, punctis pellucidis in- 
structis; petiolis ad apicem et laminis ad basim cum pilis stellatis in- 
structis; scapis paniculatis; fructus ignotus.—Colombia: Guanabanal, 
Dept. El Valle, 2 June 1922, Killip 6218 (TYPE in US; GH, NY, PH). 


This individual, of uncertain status, is appended to E. pani- 
culatus because its winged petiole and scarcely cordate blades 
will key it to that species. It could as well be placed with Æ. 
grandiflorus on account of its leathery blade with pin-point pel- 
lucid dots and stellate hairs. There are no fruits in the ample 
paniculate inflorescence. If future collectors do not show it to 
be a population in southern Colombia, with fruiting individuals 
to indicate its true relationships, it may then be considered as a 
hybrid of the two species whose characters it shares. 


11. EcuiNoponmvs trialatus. Fassett, n. sp., foliorum petiolis alatis, 
laminis acuminatis subaequantibus, venis laminarum pseudopinnatis; 
scapi internodis late 3-alatis; pedicellis 3-6 mm. longis; fructibus subro- 
stratis, costis filiformibus alatis ad apicem.—Leaves with blades about 
as long as the winged petioles and tapering into them (Fig. 21), long- 
tapered at tip, with the upper pair of veins paralleling the midrib and 
leaving it at some distance from the base; pellucid markings lacking 
(Fig. 62); inflorescence simple, its rachis with 3 green wings about 2 mm. 
wide; flowers subsessile or on pedicels up to 6 mm. long; fruit short- 


180 Rhodora [Vor. 57 


Map 9. kchinodorus trialatus. 


beaked (Fig. 50), with about 5 ribs winged toward the summit, and 
without facial glands. 

Lowlands of Colombia (Map 9). Colombia: Río Casanare, Bar- 
ranco de Atahuarpa, alt. 120 m., 30 October 1938, Cuatrecasas 4283 
(ryPE—Us); Sabanas de Santa Isabel, Meta, January 1937, Gamia 
5117 (vs); hierba sumergida, Comisaría del Vaupés, San José del Guaviare, 
en matas de monte, 240 m. alt., 4 November 1939, Cuatrecasas 7391 
(us); desiecated pool, alt. 200-250 m., Chinu, Dept. Bolivar, Pennell 
4096 (NY). 


nad c 


Map 10. Echinodorus Grisebachii. 


12. E. Grisepacuiut Small, N. Am. Fl. 17, pt. 1: 46. 1909. Æ. inter- 
medius *var.," Griseb. Cat. Pl. Cub. 218. 1866. EF. intermedius Micheli 
in DC., Monogr. Phan. 3: 54. 1881, in part; not Alisma intermedium 
Mart. 

Leaves rather small, the winged petioles only slightly exceeding the 
blades which are 7-10 em. long, ovate-oblong, with the upper pair of 
veins paralleling the midrib for some distance (Fig. 16), abundantly 
furnished with pellucid dots and lines of varying lengths (Fig. 63); scape 
simple or slightly branched, with close verticels of nearly sessile flowers; 
nutlet short-beaked (Fig. 51), more broadly winged on the dorsal edge 
than the ventral, with few ribs of which the outermost is wing-like, 
and with several glands. 


1955] ^ Fassett,—Echinodorus in the American Tropics 181 


Cuba and Costa Rica (Map 10). Cuba: Prov. Pinar del Rio, vicinity 
of Coloma, 8 March 1911, Britton & Cowan 9849 (vs); Wright 3198, 
without further data (rype number—vs). Costa Rica: Colorado de 
Coto, 20 February 1936, Valerio 1389 (F). 

The name E. intermedius was applied to this plant by Micheli, who 
thought the same species grew in Brazil. Grisebach, transferring Alisma 
intermedium Mart. to Echinodorus, added ‘‘var.: lineolae folii pellucidae 
conformes; Wr. 3198"; this forms the basis for E. Grisebachii Small. 

13. EcnuiNoponRus TUNICATUS Small, N. Am. Fl. 17, pt. 1: 48. 1909. 
E. longipetalus sensu Woodson & Schery, Ann. Mo. Bot. Gard. 30: 101. 
1943, not Micheli. 


Map 11. Echinodorus tunicatus. 


Leaves with blades 15-30 cm. long, 10-20 em. wide, deeply cordate 
at base, glabrous or sparsely muricate to somewhat pubescent with 
spreading flattened hairs toward the junction with the terete or costate 
petiole, the pellucid lines forming a network that is unrelated to the 
pattern of veins (Fig. 64); sepals pale yellow, firm, thick and brittle, 
about 30-ribbed or sometimes nearly smooth, enlarging and closely 
eupped about the fruiting head and about each other, the whole forming 
a flattened-globose head 1-1.5 em. in diameter, the calyx much depressed 
at base about the pedicel; petals 5-6 mm. long (fide Woodson & Schery); 
stamens about 30; achenes wedge-shaped, 3 mm. long, 1 mm. wide at 
summit and 0.3-0.4 mm. wide at base, the corky summit narrowed to 
an erect or ascending beak that is nearly 1 mm. long (Fig. 52); facial 
ribs 3-4; glands several. 

Panama and Costa Rica (Map 11). Costa Rica: Limón Province, 
palm swamp between Río Reventazón and Río Parismina on Castilla 
Farm, 10 m., 2 April 1930, Dodge & Neverman 7185 (Mo); from near sea- 
level creeks, Reventazón, Lankester 947 (us). Panama: in low hollow, 
Marraganti and vicinity, alt. 10-100 ft., 3-9 April 1908, Williams 991 
(TYPE in Ny); Agricultural Experiment Station at Matías Hernandez, 
Prov. Panamá, 25 December 1914, Pittier 6894 (Gu, Ny, vs); Old Experi- 
ment Station, 3 miles east of Panama City, 13 June 1923, Maxon 7095 
(cH, vs); in mud, acaulescent, common, near the big swamp east of the 
Río Yecumen, Prov. Panamá, 11 December 1923, Standley 26525 (vs); 


182 Rhodora [Vor. 57 


between Las Sabanas and Matías Hernández, Prov. Panamá, 21 January 
1924, Standley 31898 (us); Juan Díaz, Prov. Panamá, 11 January 1924, 
Standley 30506 (vs). 

Woodson & Schery, l.c., include this under the Brazilian 
E. longipetalus Micheli. But E. longipetalus, while sharing the 
accrescent sepals and pellucid network of E. tunicatus, is a 
very different species with narrow elliptic-oblanceolate leaves, 
sepals not cupped together, and petals 3-3.5 em. long. 

A third species with accrescent sepals, also from Brazil, is Æ. 
brevipedicellatus (Kuntze) Buch. It has the general aspect of 
E. longipetalus but its much narrower leaves are obviously 
different. Buchenau separated these two species on fruiting 
characters; I have not seen mature fruit. Æ. brevipedicellatus is 
keyed by Buchenau under "lineae et puncta pellucida desunt"; 
actually it has a pellucid network like that of Æ. tunicatus and 
E. longipetalus, obscured by the thickness of the leaf, but be- 
coming visible in a piece of leaf that has been boiled in alcohol. 

From Peru, Schunke 279 (US), determined as FE. intermedius 
by Standley, has a well-marked pellucid network, but the sepals 
are thin, about l6-nerved, and not accrescent; it is without 
doubt an undescribed species. 

14. EcHINODORUS NYMPHAEIFOLIUs (Griseb.) Buch. Bot. Jahrb. 2: 
483. 1882. Alisma nymphaeifolium Griseb. Cat. Pl. Cub. 218. 1866; 
Micheli in DC. Monogr. Phan. 3: 36. 1881. Helianthium nymphaei- 
folium Small, N. Am. Fl. 17, pt. 1: 45. 1909. 

Submersed leaves ribbon-like, thin and flaccid, 15-20 em. long, about 
1 em. wide; emersed leaves long-petioled, the blades oval and deeply 
cordate, 3-12 em. long and 1.5-9 em. wide; pellucid markings scattered 
lines 0.2-0.8 mm. long (Fig. 65); inflorescence 2-4 times compound, 
forming an erect ovate or conie panicle; nutlets 1.4 mm. long, 1 mm. 
wide, with a beak 0.2 mm. long, flat-sided, with a broad crested keel, 
and with erested ribs and 1 or 2 long glands on each face (Fig. 53). 

Cuba and Yucatan (Map 12). Cuba: without locality, Wright 3196 (an, 
vs, Mo); Los Palacios, Province of Pinar del Rio, 3, 4 January 1912, Shafer 
11664 (us); El Sábalo, Finca Salanalamar, near sea level, 22 December 1937, 
Killip 32286 (vs); La Gloria, Camaguey, 3 February 1909, Shafer 260 (v); 
Columbia, Isle of Pines, 19, 21 March 1916, Britton, Britton & Wilson 15703 
(us, F). Mexico: Ciudad del Carmen, Campeche, 15 April 1933, Mell 2090 
(us); Tuxpefia, Campeche, 30 January 1932, Lundell 1270 (wo, vs, 
GH, NY, F). British Honduras: Maskall Pine Ridge, January 1934, 
Gentle 1109 (F, GH, MO, NY). 

«I am indebted to my collegue, Prof. R. I. Evans, for help in this and other like 
problems. 


1955] Fassett,—Echinodorus in the American Tropics 183 


15. EcHINODORUS TENELLUS (Mart.) Buch. Abh. Nat. Ver. Bremen 
2: 21. 1868; Micheli in DC. Monogr. Phan. 3: 47. 1881; Buch. in Pflanz- 
enreich 4, pt. 15: 1903. Alisma tenellum Mart. in Roem. & Schultes, 
L., Syst. 7, pt. 2: 1600. 1830; Kunth, Enum. 3: 149. 1841. 

Robinson, Rhodora 5: 85-89. 1903, points out a seeming inaccuracy 
in the illustration of this species in the Flora Brasiliensis: the carpels 
are shown in a single ring rather than in a head as they should be in the 
genus Echinodorus. However, some plants do give a definite impression 
of a ring of carpels, until closer examination shows them to be borne in 
a head, but being few in number they have been squeezed into a ring- 
like arrangement. 


Map 12. Echinodorus nymphaeifolius. 


VARIETIES OF E. TENELLUS 
a. Nutlet 0.9-1.1 mm. long; stylar beak absent or nearly so (Figs. 66-68); 
anther 0.2-0.6 mm. long; umbel usually single and terminating the scape. 
b. Blade of leaves tapered at each end so that the sides are slightly concave 
toward the tip (Fig. 75); anther 0.2-0.6 mm. long. 
c. Facial ribs well-developed (Fig. 66)..... 15a. E. tenellus var. tenellus. 
c. Facial ribs nearly or quite suppressed (Fig. 68). 
15b. E. tenellus var. ecostatus. 
b. Blade of leaves narrowed from the middle to an acute or obtuse base and 
tip (Figs. 73, 74); anther 0.2 (-0.3) mm. long. 
15e. E. tenellus var. parvulus. 
a. Nutlet 1.4-1.8 mm. long; stylar beak 0.2-0.5 mm. long (Fig. 76); anther 
0.5-1.0 mm. long; umbels often 2, the terminal 3-6 cm. above the lower. 
15d. E. tenellus var. latifolius. 


Two phases of E. tenellus appear in southern Brazil, the type 
region for this species. The smaller plant has a nearly beakless 
nutlet about 1 mm. long (Fig. 66) and leaves 1-3 mm. wide and 
long-tapered to both ends (Fig. 75); it is illustrated in Fl. Bras. 
3, pt. 1: pl. 13, fig. 2, plant to the right. "The larger plant has a 
nutlet about 1.5 mm. long and a well-developed beak (Fig. 70), 
and leaves often with an elliptic blade (Fig. 76); this is illustrated 
in Fl. Bras., l.c., plant to the left. 


184 Rhodora [Vor. 57 


In the original description of Alisma tenellum but one collection 
is cited, so this can be considered as TYPE: In palmetis udis 
depressis Provinciae Minas geraés Brasiliae: Martius. The de- 
scription says ‘‘capsulis obsolete mucronulatis" and “Folia . . 
utrinque angustata . . . 14-1 pol, 34-114 lin. lata," so it is 
presumably the phase with beakless nutlets and narrow leaves, 
to be taken up in this treatment as var. tenellus. 

In the Flora Brasiliensis some 5 collections are cited and the 
description is expanded to include forma latifolia with leaves up 
to 4 lines long. This is presumably what is illustrated by the 
drawing to the left on plate 13 referred to above. The only 
nutlet shown in detail is definitely beaked, in contradistinction 
to the original description, and is also presumably forma latifolia. 
Differing in leaves, fruit, anthers and inflorescence, forma lati- 
folia will be taken up as a variety in the present treatment. 


15a. E. TENELLUS var. TENELLUS. Alisma tenellum Mart. l.c.; Seubert 
in Mart. Fl. Bras. 3, pt. 1: 105, t. 13, fig. 2, plant to right. 1848. A. 
ranunculoides L. var. brasiliense A. de St. Hilaire, Voy. Distr. Diam. 
2: 432. 1833. 

South America, mostly in the Amazon basin, but occurring from 
Venezuela to southern Brazil (Map 13a). Venezuela: Laguna de la 
Culebra, near S. Carlos, Cojedes, in half-dried places, 6 April 1925, 
Pittier 11707 (Ny); Anzoátegui, Estanque de El Baño de El Tigrito, 26 
January 1944, Pittier 15168 (vs); Apure, Cunavicha, 13 February 1941, 
Chardon 252 (us). Colombia: low muddy bank of the Rio Muco, 150 
m. alt., ca. 80 km. northwest of San José de Ocuné, Comisaria del Vichada, 
13 January 1944, Herman 10934 (an, vs); La Serranía entre los ríos 
Ariari y Meta, Llano Grande, 320 m. alt., 26 November 1939, Cuatre- 
casas 7878 (vs). Brazil: Santarem, Prov. Pará, July 1850, Spruce 
(GH, MO, NY); Minas Geraes, ex herb. Mart., probably fragments of 
TYPE (MO); Piauf, Gardner 2740, cited in Fl. Bras. (us); Lagoa Salgadinho, 
Sud-Piauí, 1913, Luetzelburg 378 (wv); Ceará, low sandy flats about 
Açude Joao Lopez, Fortaleza, 10 September 1935, Drouet 2453 (aH); 
Lagoinha, Campo Grande, Matto Grosso, 4 September (or 9 April?) 
1936, Archer & Gehrt 36327 (vs); Habira do Campo, Prov. de Minas, 
1887, Glaziou 17299 (us). Peru: Prope Tarapoto, 1855-56, Spruce 
4491 (GH, NY). Bolivia: Reis, alt. 1500 ft., February 1886, Rusby 555 
(GH, MO, NY, US). 

15b. E. TENELLUS var. ecostatus. Fassett, n. var., achaeneis 1.0-1.1 
mm. longis, obsolete mucronatis, cum costulis obscuris aut nullus (Fig. 68). 

This appears to be an extreme derived from var. tenellus, occurring 
at and beyond its northern limits. Var. ecostatus grades into var. tenellus, 
and on some individuals the nutlets are variable as to development of 
ribs, 


1955] Fassett,— Echinodorus in the American Tropics 185 


` 
D 
an ; - Se TR ee eee ee un 
i 
J S 


Map 13a. Echinodorus tenellus var. tenellus. 


Panama to Venezuela (Map 13b). Panama: on clay in dried up bed 
of pond, Peronome and vicinity, 23 February—22 March 1908, Wil- 
liams 246 (us; Ny). Venezuela: in drying lagoon near Ospino, Por- 
tuguesa, 26 December 1925, Pittier 12018 (us—ryrE; Ny); Culebra 
Lagoon near S. Carlos, Cojedes, in half dried places, 6 April 1925, Pit- 
tier 11707 (us). 

15e. E. TENELLUS var. parvulus (Engelm.) n. comb. KE. parvulus 
Engelm. in Gray, Man. ed. 2; 438. 1856; Rand, Rhodora 5: 83-85. 1903; 
Robinson, Rhodora 5: 85-89. 1903. E. subulatus Engelm. in Gray, 


vinqetesdsealuectsuvucwUrAdanss neminem Emme eme emeret tnmen tnm mmm dta qa m mA ao ar m Rar atr m dre Siu mE D quim 


?- . 
c Samos. 


[7] 


Map 13b. Echinodorus tenellus var. ecostatus. 


Man. ed. 1; 460. 1848, not Alisma subulatum L., Helianthium tenellum 
Britton, Man. ed. 2; 54. 1904. H. parvulum Small, N. Am. Fl. 17, 
pt.1:45. 1909. 

Rare and local, from southern Mexico to Texas, Florida and Cuba, 
northward to Missouri and on the Coastal Plain to Massachusetts 
(Map 13c). Massachusetts: Winter Pond, Winchester, 7 August 1870, 
Morong (F), 8 August 1876, Morong (mo), 17 August 1876, Morong (ny, 


186 Rhodora [Vor. 57 


us); 28 August 1878, Faxon (GH, MO, NY, US); September 1878, Morong 
(r), 27 September 1916, Ware 6524 (us), 10 September 1902, Churchill 
(mo); Mt. Auburn, Cambridge, August 1868, James (au), August 1869, 
James (Mo, NY, US); September 1869, James (Mo). New York: Maple 
Grove, Long Island, 4 July 1904, Bicknell (Ny); Queens, Long Island, 
16 September 1921, Ferguson 934 (Ny); Queens, Long Island, 25 October 
1928, Ferguson 7378 (Ny, us). New Jersey: Nullys Pond, Delanco, 
19 September 1918, Pennell 9917 (wv); Delanco, Burlington Co., 17 
August 1907, Van Pelt (au), 11 August 1908, Van Pelt (Ny). Dela- 
ware: Canterbury, August 1874, Canby (Gu, us, F). South Carolina: 
Santee Canal, July, Ravanel (aH). Georgia: Open Pond, Decatur Pond, 
12 August 1901, Harper 1202 (an, Mo, Ny, vs). Florida: Quincy, Chap- 
man (Ny); Tallahassee, 31 May 1925, Harper (Ny, vs); Dunnellon, 
Marion Co., 11 June 1900, Curtiss 6658 (GH, Mo, NY, US); Tampa, April, 
Curtiss 2739 (GH, MO, NY, US); Fort Meade, 20 March 1880, Smith (us). 
Illinois: East St. Louis, 4 August 1892, Eggert (Ny); St. Clair Co., 11 
August 1892, Eggert (Mo). Missouri: margin of ponds on the hills north 
of St. Louis, August 1845, Engelmann (mo); margins of ponds back 
of St. Louis, August 1845, Engelmann (F); St. Louis, August 1845, Engel- 
mann (NY); margins of ponds in the hills west of town, St. Louis, Septem- 
ber 1845, Engelmann (wo); in ponds northwest of St. Louis, August 
1848, Engelmann (wo); St. Louis, August 1848. Engelmann (NY); 
common along open sedgy shoreline, Adobesee Pond (sink-hole pond 
now ruined by artificial disturbance), 9 mi. southeast of West Plains, 
Howell Co., 4 September 1949, Sleyermark 69124a (rF). Mississippi: 
borders of ponds near Osyka, July 1899, Cocks (Tulane). Texas: Horse- 
shoe Lake, Jackson Co., 9 August 1920, Drushel 4143 (Mo, vs). Cuba: 
Rancho Boyeros, Provincia de Habana, 22 July 1904, Baker & Wilson 
375 (Ny); Mazorra, 23 November 1904, Baker & Abarca 4210 (am). 
Mexico: Vera Cruz, 1855, Müller 2142 (NY). 


Most of the range of this variety is on the Atlantic Coastal 
Plain and on low ground, geologically youthful, around the Gulf 
of Mexico (outlined with a broken line on Map 13c). From the 
Coastal Plain it extends its range northeastward to Winter Pond, 
Winchester, Massachusetts, whose assemblage of rare or isolated 
southern plants has long been a subject of comment.’ In the 
Mississippi Valley, also, its range extends north of the Coastal 
Plain to the region of St. Louis. The recent collection by Dr. 
Steyermark from southern Missouri in the heart of the Ozark 
Plateau (cross-hatched on Map 13c) may be the clue to the 
ancestral area from which the North American phase of Echino- 
dorus tenellus invaded the Coastal Plain. 


5 Rand, Rhodora 5: 83. 1903; Sears, Rhodora 10: 42. 1908; Fernald, Rhodora 10: 
143. 1908; Rhodora 13: 151. 1911. 


f 
ttres 


/ 
-- 9. 
M: 


ts.. 
^ eu 
t--e. 


Echinodorus tenellus var. parvulus. 


Map 18c. 


188 Rhodora [Vor. 57 


GP FE NENNEN SM 


7 Y b" ` 
mep 7 Fo a 
M Y Net fe Hu 
UAE | = 359... 
Meet Ul T o- 


Map 13d. Echinodorus tenellus var. latifolius. 


E. TENELLUS var. PARVULUS f. Randii Fassett, n.f., plantae submersae, 
steriles aut fertiles; foliis linearibus, non petiolatis, membranaceis, 1-3 
mm. latis.—Massachusetts: Winter Pond, Winchester, Oct. 10, 1901, 
E. L. Rand (rve& in Gray Herbarium). 


While it has been collected only at Winter Pond, the submerged 
form must surely be expected wherever the species grows. LE. 
tenellus commonly grows on wet shores or at the water's edge; 
the leaves are more or less clearly petioled with blades narrowed 
to each end, of firm texture, feather-veined, narrowly callous- 
margined and callous-tipped. The submersed forms (see also 
var. latifolius) have leaves becoming linear, of thin membranous 
texture with lacunate tissue especially near the midrib, and more 
obseurely veined. 

(To be concluded) 


Volume 57, no. 677, including pages 133-160, was issued 9 June, 1956. 


Douora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN ` Associate Editors 
RICHARD ALDEN HOWARD 

CARROLL EMORY WOOD, JR. 


Vol. 57 July, 1955 No. 679 


CONTENTS: 


Distribution and Abundance of Shortia galacifolia. P. A. Davies 189 


Echinodorus in the American Tropics (concluded). Norman C. 
kasset i. ene es a Gacvicte oe PEE 202 


The New England Botanical Club, Ine. 


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Vol. 67 July, 1955 No. 679 


DISTRIBUTION AND ABUNDANCE OF 
SHORTIA GALACIFOLIA 


P. A. DAVIES 


From 1839, when Asa Gray found an unnamed specimen 
from the Carolinas in André Michaux’s herbarium in Paris and 
named it Shortia galacifolia,! until 1877 when George M. Hyams 
rediscovered a small station of it on a hillside near a small stream 
which flowed into the Catawba River about five miles north 
of Marion in McDowell County, North Carolina,? the prime 
efforts of botanists and collectors was its rediscovery. Following 
Hyams rediscovery of Shortia, the effort shifted to determining 
its distribution and abundance. In 1879, Asa Gray and party, 
including Charles S. Sargent of the Arnold Arboretum, sought 
for additional plots without success on the west side of the 
Linville Mountains at North Cove and on the Iron Mountain 
Range, both in the region of Hyams' discovery. No more 
extensive excursions or discoveries were made in this area until 
recently. 

Nine years after Hyams’ discovery, Charles S. Sargent, 
with his guide, Frank E. Boynton of Highlands, North Carolina, 
while seeking Michaux's Magnolia cordata at the headwater 
of the Keowee Hiver, found Shortia galacifolia near the con- 
fluence of the Horsepasture and Toxaway Rivers in Oconee 
County, South Carolina. Desiring additional specimens and 
information on its distribution and abundance, Sargent requested 

1 Gray, Asa, Amer. Jour. Sci. & Arts 42: 48, 1842, 

? Letter, J. W., Congdon to Asa Gray, Oct. 18, 1878 (Gray Herb.). 


3 Redfield, J. H., Bull. Torrey Bot. Club 6: 331-336, 1879. 
* Sargent, C. S., Amer. Jour. Sci. & Arts, Ser. III, 32: 466-473, 1886. 


190 Rhodora [Vor. 57 


Boynton to return to the forks of the river, their collecting 
ground of the previous month, and explore the region. This 
Boynton did and found large quantities of Shortia on Bearcamp 
Creek, a lower tributary of the Horsepasture River? Three 
years later (1889) Boynton found acres of it in the locale of 
the junction of the Whitewater River and Devilfork Creek and 
in the lower Jocassee Valley, all in Oconee County, South 
Carolina.’ In the spring of 1890, T. G. Harbison found Shortia 
in great abundance on the Horsepasture River between its 
junction with the Toxaway River and Bearcamp Creek.” 

The acres of Shortia that F. E. Boynton discovered along 
the Whitewater River in Jocassee Valley were easily accessible 
by roads and trails. Now that an abundant supply had been 
found, more than enough to meet the demands of botanists 
for herbarium specimens and nurserymen to supply the publie 
with living plants, the scientifie interest brought on by nearly 
fifty years of intensive search for it was lessened. Almost an 
equal amount of time was to pass before interest was again 
awakened for additional knowledge on its distribution and 
abundance. 

Charles F. Jenkins sought information on the distribution 
of Shortia by assembling data from herbarium sheets in thirty- 
one of the leading herbaria in the United States. In all he 
found ninety-eight sheets and gave the distribution as follows: 
“Forty-four were collected along the banks of the Whitewater 
River in Oconee County, South Carolina. Fourteen more are 
listed as coming from the Jocassee Valley, also in Oconee County. 
Thirteen were found in McDowell County, North Carolina, 
more definite locations not being given. Eight were found 
in the Toxaway Gorge in Transylvania County, North Carolina, 
three along Bearcamp Creek and two along the Horsepasture 
River, both located in Transylvania County. One each from 
the mountains of N. W. Carolina, four miles N. W. Salem, South 
Carolina, and Macon County, North Carolina. The remainder 
are specimens with labels giving no definite locality or coming 
from cultivated plants."'* 

5 Letter, F. E. Boynton to C. S. Sargent, Nov. 7, 1886 (Gray Herb.). 

5 Boynton, F. E., Garden & Forest, 2: 214—215, 1889. 


7 Clute, W. N., Amer. Bot. 32: 65-68, 1926. 
3 Jenkins, C. F., Arnoldia 2: 13-28, 1942, 


1955 Davies,— Distribution of Shortia galacifolia 191 


In the library of the Botany Department, University of 
North Carolina, is a distributional map of Shortia prepared 
by William C. Coker from both herbarium sheets and field 
records. For McDowell County, North Carolina, he gives 
stations on Tom's Creek, branches of John's Creek, and several 
places on the east side of the lower part of Bald Mountain. 
Dr. Coker includes on his map the single station reported by 
F. M. Crayton on the Linville River west of Table Rock Moun- 
tain in Burke County, North Carolina. On the Keowee River 
and its sources in Pickins County, South Carolina, he lists 
two locations on the east side of the Keowee River below the 
entrance of Big Eastatoe Creek; in Oconee County, South 
Carolina, four stations above the entrance of Big Eastatoe Creek, 
sixteen places on the upper sources of Little River, and several 
plots along the Whitewater; and in Transylvania County North 
Carolina, three records along the Toxaway River and two on 
Toxaway Creek.? 

The southernmost location is the one discovered by A. B. 
Massey in 1911 on the Keowee River below the entrance of 
Little River. In recording his discovery Massey relates, “This 
is a small station on the bank of the Keowee River at the foot 
of a wooded bluff. When I last visited the place, there were 
only a few plants; however, it may have increased in recent 
years."!" On November 3, 1952, D. E. Wade visited Massey's 
station and explored the area adjacent to it. He describes 
his observations: “In a rocky ravine there are at least six clumps 
of Shortia, the largest [Massey station] was approximately 3 by 
6 feet at an elevation of about 615 feet, while the highest was 
at about 680 feet. The entire ravine is cooled by a small stream 
which is fed by springs at various places. "The woody plant 
life is marked by hemlock, rhododendron, laurel, holly and 
beech. It looks as if the stand had been there many hundreds 
of years. Most of the clumps are definitely above flood level." 
In July 1953, D. C. Wade showed the writer the Massey station 
and the additional stands he found. The clumps were restricted 
and the plants were rather small but healthy. 

° Map, W. C. Coker, Botany Dept. University of North Carolina, copy sent to 
P. A. Davies, May 18, 1948. 


10 Map, A. B. Massey to P. A. Davies, June 30, 1952. 
11 Map and letter, D. E. Wade to P. A. Davies, Nov. 7, 1952. 


192 Rhodora [Vor. 57 


C. F. Moore of Brevard, North Carolina, located several 
stands on the Keowee River between Laurel Branch and 
Whitewater River, a single station near the source of Laurel 
Branch, a small area on the Bearwallow Creek about one-half 
mile from its mouth, and on subsequent visits with B. C. 
Olney observed many patches along the lower part of this 
creek. They also found Shortia on Rock House Creek, near 
the mouth of Laurel Fork Creek, on the Keowee River be- 
tween Laurel Branch and the Toxaway River, along the 
Whitewater River in Jocassee Valley, on eastern branches of the 
Horsepasture River and along a ridge between the Horsepasture 
River and Bearwallow Creek.” 

In April, 1917, W. W. Diehl saw Shortia in abundance and 
in full bloom on the northeast bank of the Whitewater River 
between the Thompson and Toxaway Rivers, and on three 
other occasions between 1921 and 1927 near the mouth of 
Bearwallow Creek. In the Gray Herbarium is a specimen 
collected by E. T. Wherry on July 12, 1936 along Bearwallow 
Creek and another by H. J. Oosting on July 15, 1936 in the 
Toxaway Gorge, Transylvania County, North Carolina. During 
March, 1944, A. E. Prince, J. A. Berly and O. L. Cartwright 
found Shortia in flower along the road near the Whitewater 
River just below the entrance of Devilfork Creek and later 
(1946) Prince found it plentiful on the right bank of the Toxaway 
River above its confluence with the Whitewater River.* The 
Rev. A. Rufus Morgan, Franklin, North Carolina, found Shortia 
on the Toxaway River above the North Carolina border and on 
the Whitewater River just above its junction with the Thompson 
River.’ W. H. Duncan reported a station in an open woods 
along a tributary of Middle Fork Creek in Oconee County, 
South Carolina.'® 

In March 1949, W. H. Duncan, Haskell Venard and G. W. 
McDowell discovered the first station in Georgia. It was a 
small colony (8 X 4 feet) near Reed Creek in Rabun County, 
and in August of the same year Duncan found a smaller colony 
near the original one.!? 

1? Map, C. F. Moore to P. A. Davies, June 30, 1952. Also, letter, Aug. 2, 1954. 

13 Letter, W. W. Diehl to P. A. Davies, Sept. 23, 1952. 

4 Prince, A. E., Rhodora, 49: 159-161, 1947. Also, map and letter, May 24, 1953. 


15 Map, A. Rufus Morgan to P. A. Davies, Aug. 8, 1951. 
15 Letter, W. H. Duncan to P. A. Davies, May 23, 1952. 


1955] Davies,— Distribution of Shortia galacifolia 193 


Several healthy and expanding patches of Shortia are present 
in Highlands, North Carolina and its environs. All available 
data indicate that these plantings were made from plants gathered 
by F. E. Boynton, T. G. Harbison and others along the sources 
of the Keowee River, particularly the Whitewater River. The 
writer has examined many streams and their smaller tributaries 
about Highlands that appear to offer suitable habitat without 
finding a single indication of Shortza. 

Many sources of reference state that Shortia galacifolia is 
rare. These statements have probably arisen from the fact 
that Asa Gray and his colleagues sought for it without success 
for thirty-eight years (1839-1877) and the patch that G. M. 
Hyams finally discovered near a tributary of the Catawba River 
was too small to supply the demands for herbarium specimens. 
The recently discovered patches along the sources of this river 
are not numerous. On the sources of the Keowee River most 
collectors have observed Shortia on the fringe of its range or 
have made distributional judgments from the colonies within 
easy reach by roads or trails along the Whitewater River or 
Bearwallow Creek. In certain places on the drainage areas of 
the Toxaway and Horsepasture Rivers it is one of the most 
abundant plants, and F. E. Boynton, C. F. Moore, B. C. Olney, 
and the writer have observed patches containing one or more 
acres. Until recently the areas containing an abundance of 
Shortia on the sources of the Toxaway and Horsepasture Rivers 
were accessible only by a few strenuous trails and nonserviceable 
roads. Over the Poinsett Lumber and Manufacturing Co. road 
to these areas, Shortia may now be seen in abundance. 

Shortia grows in various habitats: hanging from rocks near 
water falls (water dripping from its leaves) in sand at the edge 
of running water, on fallen decaying tree-trunks, in shady, 
deep, moist loam soils, and on dry hillsides. Its most favorable 
habitat is a cool, damp, shady stream bank, with moist circulating 
air, and a surface covering of two to six inches of loose organic 
debris with a fertile, moderately acid soil (pH 4.2—6.0) beneath. 
While Shortia is more frequently found beneath Rhododendron 
maximum it will grow equally as well under the shade of T'suga 


17 Duncan, W. H., Haskell Venard and G. W. McDowell, Rhodora 52: 229-232, 
1950. 


194 Rhodora [Vor. 57 


canadensis, Pinus Strobus, Pinus rigida, Liquidambar Styraciflua, 
Liriodendron Tulipifera and other acid tolerating trees and 
shrubs. Plants found in the most favorable habitats have 
large leaves with long petioles and large flowers on long pe- 
duncles; while those growing under less favorable conditions 
(dry hillsides) have smaller leaves on short petioles and small 
flowers on shorter peduncles. 

Fortunately for the preservation of Shortia, the largest per- 
centage of it is confined to large private holdings and government 
land. Along the Toxaway and Horsepasture Rivers and their 
tributaries in Oconee and Pickens Counties, South Carolina, 
the Poinsett Lumber and Mfg. Co. holds the areas of greatest 
concentration of Shortia, while in Transylvania County, North 
Carolina, it is the Duke Power Co. which controls such areas. 
The writer, his colleagues and students are indebted to Thomas 
Mitchell and Furman Chastain of the Poinsett Lumber and 
Mfg. Co. and C. F. Moore of the Duke Power Co. for permission 
to seek Shortia on the company lands. On the sources of the 
Catawba River in McDowell County, North Carolina more 
than one-half the SAortia plants are in the Pisgah National 
Forest. Both the public and private organizations have the 
means of controlling the destroyers of it, so Shortia is safe for 
posterity. 

The writer became interested in Shortia while preparing a 
paper on the life of Charles W. Short, first of the Kentucky 
born botanists and recipient of Asa Gray’s honor, for the Filson 
Club History Quarterly. Available references on Shortia did 
not present a unified picture of its distribution and little could 
be found about its abundance. In order to obtain more com- 
plete knowledge of its distribution and abundance, investigations 
were made which included extensive field excursions and the 
following of all significant leads through correspondence and 
conversations with persons who had collected Shortia or had 
heard of places where it could be found. 

Explorations were made by working up or down the rivers, 
creeks, or branches. Not all the small tributaries were followed 
to their sources. In several places the terrain was too rough 
or the thickets so impenetrable that no explorations were made 


18 Davies, P. A., Filson Club Hist. Quart., 19: 131—155, and 19: 208-249, 1945. 


1955] Davies,— Distribution of Shortia galacifolia 195 


for fear of bodily injuries. Most of the surveys were made 
in dry weather and at low water stages, so the explorers were 
able to wade the streams through most of the gorges and to 
escape the dense thickets. 

It was the author's original intention to publish a full account 
of the distribution of Shortia, giving exact plot locations.  Realiz- 
ing that unscrupulous collectors might destroy the easily acces- 
sible small patches, particularly on the fringes of the range, 
he decided to publish only general locations, and to deposit in 
the library of the Gray Herbarium data on exact location, 
including maps, and the correspondence accumulated during 
the study. In this way Shortia may be protected and the 
information on exact locations be made available to anyone 
interested in checking various plots or continuing the study of 
its distribution. 

The most southern extensive search by the writer and his 
colleagues was on the middle and eastern sources of Little 
River near the divide which separates its watershed from that 
of the Whitewater River. Along these small headwater streams 
many scattered patches were observed. A more complete 
survey of the smaller western branches would undoubtedly yield 
many new locations. 

Jocassee Valley, located in Oconee County, South Carolina, 
extends from above the confluence of the Whitewater and 
Thompson Rivers to below the Chapman Bridge over the Keowee 
River. Many streams enter the valley. The largest are the 
Whitewater, Thompson and Toxaway Rivers and Devilfork 
Creek. . The writer has carefully examined all parts of the valley. 
The acres of Shortia that Boynton discovered in 1889 have largely 
been destroyed by cultivation, grazing and road building. How- 
ever, along many of the streams and on some of the steeper 
slopes where grazing has not taken place, Shortia is still plentiful. 
There is enough of it in the valley to meet the demands of nursery- 
men for hundreds of years. The author knows of one ravine 
near the Keowee River where nurserymen have been gathering 
Shortia for more than 30 years without greatly depleting it. 

At the upper end of the valley, along the camp road on the 
south side of the Whitewater River, a few colonies of Shortia 
were observed. This river was explored all the way to the 


196 Rhodora [Vor. 57 


upper falls and no Shortia was found beyond the first road- 
bridge (elevation circa 1060 feet) above its confluence with the 
Thompson River. 

Following the Thompson Hiver from its junction with the 
Whitewater to approximately the North Carolina boundary 
numerous patches of Shortia were observed. This river was 
again examined above state highway 92 (Upper Whitewater 
Falls Road) without finding any indication of Shortia. Wright 
and Coley creeks, the two main eastern tributaries of the Thomp- 
son River in South Carolina were examined to their sources. 
As the elevation increased, Shortia became more abundant. 
Near the divide which separates the branches of Coley Creek 
from those of Mill Creek, the writer found on the Coley Creek 
side, a patch of Shortia on a moist, steep and shaded hillside. 
Below and near the patch were numerous young plants which 
were judged to be from one to three years old. In 1953, C. F. 
Moore and the writer found a similar condition below a patch 
and near the edge of Bearwallow Creek. This is a clear and 
encouraging indication that in certain places where conditions 
are favorable, Shortia is increasing rather than decreasing. 
No attempt was made to explore the tributaries of the Thompson 
River between the North Carolina boundary and state highway 
92. 

On Devilfork Creek, the large southern tributary of the 
Whitewater Hiver, scattered stations were recorded along its 
banks, but many were present higher up on the mountain along 
the numerous branches of the lower tributaries. As the head- 
waters of the southern branches of this creek are separated 
by narrow ridges from the northern branches of Little River, 
the upper stations on each drainage area are in close proximity. 

The Toxaway River ends in Jocassee Valley by joining the 
Whitewater to form the Keowee River and extends northward 
into its numerous branches above the old Lake Toxaway site 
in Transylvania County, North Carolina. Along its course 
many streams flow into it, but the majority are small and un- 
named. On its banks and along its numerous tributaries Shortia 
is plentiful, some in small isolated patches, others in large 
areas extending up the mountainside. No Shortia was found 
one fourth mile above the entrance of Bearwallow Creek, although 


1955] Davies,— Distribution of Shortia galacifolia 197 


the river was examined into its sources. Only the larger creeks 
and branches of the Toxaway River which appear on topographic 
maps will be considered in this paper although a great many 
of the smaller branches contain an abundance of it. 

The center of distribution of Shortia for the sources of the 
Keowee River basin appears to be along the Toxaway River 
below the entrance of Horsepasture (Green) River and near 
that of the Devil's Hole Creek. This position is approximately 
halfway between the stations to the south on the northern 
branches of Little River, along Devilfork Creek, on the White- 
water River and those to the north on Bearwallow Creek. In an 
east-west direction it is between the station on the upper part 
of Laurel Fork Creek and the Thompson River. It is interesting 
correlation that it was near this center of distribution, at the 
forks of the Toxaway and Horsepasture Rivers, that C. S. 
Sargent believed André Michaux, in the inclement weather of 
December 1788, collected the type specimen and that he almost 
a hundred years later (September, 1886) rediscovered it for this 
part of the Carolinas. 

Several large and many small streams enter the Toxaway 
River from its junction with the Whitewater to the south and 
the Horsepasture to the north. All except Devils Hole Creek 
are unnamed and Shortia was found, varying in abundance, 
on nearly all of them. As one passes through the Toxaway 
watershed over the Poinsett Lumber and Mfg. Co.’s new road 
to the confluence of the Toxaway and Horsepasture Rivers, 
Shortia may be observed in abundance along the branches of 
Devil's Hole Creek and other streams in this area. Because 
of the recent cutting of trees along the upper branches, resulting 
in the drying of the surface debris, and the covering of the 
leaves with dust from the dirt road, a large part of these stands 
may not survive. 

On the Horsepasture River between its junetion with the 
Toxaway and Bearcamp Creek (main tributary of the Horse- 
pasture in South Carolina), the great amount of Shortia that 
T. G. Harbison observed in 1890 was still there when the area 
was examined in March of 1951 by the author accompanied 
by L. M. Brown, and in June of the same year with Arland 
Hotchkiss. Both sides of the river were checked and the larger 


198 Rhodora [Vor. 57 


amount was found on the more gently, less exposed, southwestern 
slopes. Rocks and soil dumped on the lower side of a new 
logging road built around the mountain in the latter part of 
1951 buried many large patches. Above the road Shortia can 
still be found in considerable quantities. 

Along the Horsepasture River above the entrance of Bearcamp 
Creek, the amount of this plant diminishes as one moves north- 
ward. The farthest north Shortia was found on this river or 
its tributaries was in North Carolina, at an elevation of about 
1600 feet, between the approach of the Duke Power Road to 
the river and Windy Falls. In this area C. F. Moore and B. C. 
Olney found an unusual station, and as Moore describes it, 
“This interesting location was high on a ridge between the 
Horsepasture River and Bearwallow Creek, but nearer the 
Horsepasture. Shortta was growing on the dry hillside far 
removed from any stream. It was not in a cove but on a well 
exposed crest. There appeared to be several acres of it."'!? 

The lower part of Bearcamp Creek and its main north branch 
were explored well above the North Carolina boundary, and 
the large quantity of Shortia that F. E. Boynton discovered 
along the lower part of this creek in the fall of 1886 still remains. 
It was found hanging from the moist wall of the gorges, on decay- 
ing fallen trees and in the loose, fertile acid soil (pH 5.2) along 
the stream. Along Mill Creek, a southern branch of Bearcamp 
Creek, Shortia was frequently found, and in certain places 
extending up the mountain. As its sources near the divide 
(circa 1800 ft.), which separates the drainage areas of the Horse- 
pasture River from those of the Thompson River, were reached, 
Shortia became less abundant. 

Laurel Fork Creek, the lower main tributary of the Toxaway 
River above its junction with the Horsepasture River, entends 
into Pickens County, South Carolina. On this eastern branch, 
Shortia is growing near the lower bridge crossing, about one- 
fourth mile above its mouth. C. F. Moore and B. €. Olney 
previously recorded this location. At the old millsite, farther 
up the creek, it was found in abundance extending up the moun- 
tain sides. Numerous stations were observed along the creek 
and its branches in the two miles explored above the mill-site. 
In March of this year C. F. Moore and J. L. Kapp found it in 


19 Letter, C, F, Moore to P. A. Davies, June 30, 1952. 


1955] Davies,— Distribution of Shortia galacifolia 199 


abundance further up this creek to an elevation of about 1600 
feet." On Jackies Branch, a northern tributary of Laurel Fork 
Creek, Shortia was found near its mouth. No attempt was made 
to explore the headwaters of this small stream. 

Three creeks joining the river just below the North Carolina 
border were next examined. Cobb and Bear Creek rising in 
Transylvania County, North Carolina, flow southeast into 
Oconee County, South Carolina, while the third creek (unnamed) 
arises in Pickens County, South Carolina, and flows almost due 
west to the river. Cobb Creek, the smaller and more northerly 
branch on the west side of the river was explored along the lower 
two-thirds of its course. This creek had very little water, so 
conditions were not favorable. A few small patches were found 
near its mouth. Bear Creek was followed from its confluence 
with the river to the Duke Power Co. road. Habitats were 
favorable and many stations of Shortia were observed. No 
examinations were made above the Duke Power Co. road. The 
eastern unnamed branch was examined but a short distance and 
several small patches were found. 

In the lower part of Transylvania County, three streams enter 
the Toxaway River at the Cane Brake: Rock and Toxaway 
Creeks from the east and a small unnamed one from the west. 
Along the lower and more gentle terrain of Rock (Rockhouse) 
Creek no Shortia was observed. At the narrows about three- 
fourths mile above its mouth, it was found but not in abundance. 
No further inspection of this creek was attempted. In 1948 
C. F. Moore reported a station higher up near its sources. 
Toxaway Creek was explored as far as the falls. Again Shortia 
was found along its banks. Frozen Creek, a northern branch 
of Toxaway Creek, was explored above Frozen Lake without 
suecess. The farmers in this area had never seen or heard of it. 
The unnamed creek from the west was examined about one-half 
way up its course. Eight stations were counted. 

Studies were made on the lower one-fourth mile of Auger 
Fork Creek. No Shortia was found except near the river. 
The small unnamed creek just above the Auger Fork Creek was 
examined near the road crossing. No Shortia was found at this 
place. 

Bearwallow Creek is a main western division of the Toxaway 


20 Letter C. F. Moore to P. A, Davies Aug. 2, 1954. 


200 Rhodora [Vor. 57 


River below the falls. With the exception of the Whitewater 
River area, more persons have observed and collected Shortia 
along the lower part of this creek than in any other part of the 
Keowee drainage basin. The writer has been along the lower 
part of this creek five times and has observed patches of Shortia 
each time. The fourth time, the creek was reached from a 
logging road and trail from F. O. Fisher’s place between Rosman 
and Cashiers on highway 64. No Shortia was found along this 
creek until an elevation of about 1600 feet was reached. The 
stations along Bearwallow Creek and the one observed on the 
Toxaway River about one-fourth mile above the entrance of 
this creek mark the northern station of Shortia observed by 
the writer in the Keowee drainage area. 

The amount of Shortva on the drainage areas of the Catawba 
River is exceedingly small when compared to that on the tribu- 
taries of the Keowee River. Except for one small station re- 
ported by F. M. Crayton on the Linville River, west Table Rock 
Mountain in Burke County, all locations are near streams with 
sources in the lower part of the Bald Mountain chain. 

Although several streams, most of which are small and not 
indicated on available maps, rise from the lower part of the 
Bald Mountain chain, only three have received names: Toms 
Creek on the west, Johns Creek (Fish Hatchery Creek) on the 
southeast, and McGegers Branch to the east. W. C. Coker 
reports a station on Toms Creek. Milton Bowman and the 
writer in June 1952 located Coker’s station and found one above 
and another below it. Three small stations were observed on 
a small unnamed branch of Toms Creek flowing from the south- 
ern tip of Bald Mountain. Unfortunately no record was made 
of the exact location of Hyam's station, although he stated it 
was on a mountain side near a small stream that flowed into 
the Catawba River about five miles north of Marion. The 
area at the southern tip of the Bald Mountain chain fits all 
known data as to the location of Hyam’s station. Here the road 
from Marion to Turkey and North Coves runs close to the moun- 
tain, crossing several streams, and is about five miles north of 
Marion. 

Johns Creek has three main branches, all with sources in the 
Pisgah National Forest. On the right branch no Shortia was 


1955] Davies,— Distribution of Shortia galacifolia 201 


found, probably due to heavy grazing. Along the left branch 
a small pateh was observed on a hillside just above the old 
sawmill site. Many stations were found on its sources in the 
national forest. The middle branch, frequently called the fish 
hatchery branch, has all its sources in the national forest. Above 
the hatehery and along the main tributary, Shortia is growing 
almost to the top of the ridge. 

The stations reported by W. C. Coker on the east side of 
the Bald Mountain, were checked and found to be in good 
condition. An examination of MeGegers Branch and other 
streams between it and Johns Creek resulted in several small 
patches. No explorations were made above MeGegers Branch 
on the east side of the mountains. Limekiln, Cox and Arm- 
strong Creeks on the west side of the Bald Mountain chain were 
checked at various places without success. 

Two attempts were made to reach the Crayton station on 
the Linville River below the Table Rock Mountains. On April 
12, 1952, Milton Bowman and the writer descended the gorge 
between Hawksbill and Table Rock Mountains. Narrowness 
of the gorge and high water prevented us from reaching the 
area directly below Table Rock Mountain although the river was 
explored for some distance. No Shortia was found. Later the 
river was ascended from Lake James to a place below Chimney 
Gap before heavy rains forced a retreat. The part examined 
gave no indication of Shortia. A group of senior scouts ascending 
the river were given leaves and flowers of Shortia and instructed 
where to look. They were particularly asked to search the area 
directly below Table Rock Mountain. Although their main 
interest was to ascend the river to the Linville Falls, they did 
spend some time looking for Shortia. Their results were nega- 
tive. From the writer's experience along this river, he is led 
to believe that the Crayton station is simply an outpost, and 
that if other patches are present they are few in number.— 
DEPARTMENT OF BIOLOGY, UNIVERSITY OF LOUISVILLE, LOUISYILLE, KEN- 
TUCKY. 


202 Rhodora [Vor. 57 


ECHINODORUS IN THE AMERICAN TROPICS 


NonMAN C. FaAssETT 
(Continued from p. 188) 


15d. E. rENELLUs, var. latifolius (Seubert) Fassett, n. comb. Alisma 
tenellum forma latifolia Seubert in Mart. Fl. Bras. 3, pt. 1: 195, t. 13, 
fig. 2, nutlet and plant to left. 1848. 

Southern Brazil to Venezuela, West Indies, and northern Central 
America (Map 13d) Guatemala: Lake Zotz, Petén, I8 May 1933, 
Lundell 8293 (r); Dept. Jalapa, Potrero Carrillo, 13 miles northeast of 
Jalapa, alt. 1500-1700 m., 12 December 1939, Steyermark 33098 (r); 
Dept. Alta Verapaz, bogs near Santa Cruz, along road to San Cristóbal, 
alt. about 1350 m., 16 April 1941, Steyermark 42754 (v); Dept. Alta 
Verapaz, large swamp just east of Yactic, alt. 1300 m., 20 February 
1942, Steyermark 43977 (r). Honduras: in bog, in pine forest, El Achote, 
near Siguatepeque, Dept. of Comayagua, alt. 1500 m., 18 February 
1928, Standley 56099 (us, F). Jamaica: growing in swamp, Cornwall, 
St. Elizabeth, I8 July 1915, Harriss 12095 (Gu, F, MO, NY, US); Cornwall 
swamp, near Locovia, 1915, Perkins (NY). Dominican Republic: Prov. 
de la Vega, prope Tarabacoa in paludosis 550 m. alt., June 1912, Fuertes 
1666 (v, GH, NY, US); Llano Costero, Prov. Sto. Domingo, Cuenca, edge 
of laguna, 6 January 1929, Ekman 11025 (vs). Porto Rico: Laguna 
Yeguada near Vega Baja, 24 March 1922, Britton, Britton & Brown 
6773 (r, NY, US); muddy marsh, vicinity of San Juan, 11 & 12 February 
1914, Britton & Cowell 1483 (Gn, Ny, US); Martin Pena, 12 February 1914, 
Johnston 1314 (Ny). Venezuela: Tovar, 1854-55, Fendler 1358 (am); 
La Trinidad de Maracay, at an altitude of 440 meters, State of Aragua, 
January-February 1913, Pittier 5844 (Ny, vs); in morass, El Limón, 
near Maracay, Aragua, 19 January 1922, Pittier 10104 (GH, NY, Us); 
Anzoátegui, headwater of Río Guanipa, 28 December 1940, Pittier 
14637 (vs). British Guiana: herb, on edge of drying pond, basin of 
Rupununi River, lat. about 2° 52’, 25 & 26 October 1937, Smith 2285 
(x, Mo, Ny, vs). Bolivia: Reyes, alt. 1000 ft., October 1921, Rusby 
1540A (Ny, vs). Brazil: Lagoa Santo, Minas Geraes, 3 August 1936, 
Archer. 3612 (us); Omolas, Matto Grosso, 6 August 1908, Hoehne 30 
(us). Paraguay: in regione lacus Ypacaray, May 1913, Hassler 12654 
(F, GH, MO, US); common in a dry lagoa, Tapytá, April 1931, Jórgensen 
4722 (NY, US, F, MO); fluminis Apa, February 1902, Hassler 8453 (GH, MO). 


This is often the largest phase of E. tenellus, some of the 
plants having a scape 35 em. long, with leaves 20 em. long and 
12 mm. wide (Fig. 76). A collection from Honduras: El Achote, 
hills above the plains of Siguatepeque, Dept. Comayagua, 
Y uncker, Dawson & Youse 6521 (F), has a slight development 
of a fourth lateral rib on some achenes. This may represent a 


1955] Fassett,—Echinodorus in the American Tropics 203 


di 
E, H 
. 
° 
-- * 
/ n 
b: 
"e. 3 
Ns f 
N, 
Maey f N 
Ca DN f 
c* J AULAM 
AMENS r2. ` zs 


Map 14. Echinodorus isthmicus. 


slight introgression from populations to the southeastward in 
Costa Rica, where that character is well-developed. 


E. TENELLUS var. LATIFOLIUS f. apanecae Fassett, n.f., plantae in 
aqua vadosa; foliis linearibus longis-oblanceolatis, 12-22 cm. longis, 
flaccidis, membranaceis, ad basim attenuatis.—El Salvador: in about 
1 foot of very cold water, Lagunita las Ninfas, Apaneca, alt. 5430 ft., 
28 January 1951, Fassett 28704 (GH, wis); shallow water, mud bottom, 
Laguna Verde, Apaneca, alt. 5100 ft., 28 December 1950, Fassett 28505 
(TYPE in Chicago Natural History Museum, GH); same data, no. 28508 
(F); same locality, 30 December 1950, Fassett 28525 (F). 


When the fertile terrestrial form is not found nearby the 
identification of submersed forms is more difficult. Since Æ. 
tenellus var. latifolius is the only one of this group that has been 
collected in Central America, these plants presumably belong to 


Map 15. Echinodorus magdalenensis, dots; E. quadricostatus, triangle. 


204 Rhodora [Vor. 57 


that variety. "The leaves are much larger than in the terrestrial 
forms. A few leaves of no. 28508 are a little more like the 
terrestrial ones, and show the callous margin and tip charac- 
teristic of E. tenellus. 

16. Ecurvoponas isthmicus Fassett, n. sp., foliis lanceolatis, 3-7 em. 
longis, 2-5 mm. latis; seapis 6-12 em. altis, saepi verticillatis; antheris 
0.7 mm. longis; achaeniis 1.6 mm. longis, 4-costatis utrinque, rostro nullo 
aut 0.2 mm. longo (Fig. 69). 

Costa Riea and Panama (Map 14). Costa Rica: Cienega da Agua 
Buena, Cafias Gordas, alt. 1100 m., February 1897, Pittier 11126 (us). 
Panama: wet meadows along Río Caldera, south of EI Boquete, alt. 
1250 m., | March 1918, Killip 3618 (Us—rYpPE). 

17. EcuiNopoRus quadricostatus Fassett, n. sp., laminis foliorum 
3.5-5 em. longis, 3-6 mm. latis; achaeniis 1.8 mm. longis, 4-costulatis 
utrinque (Fig. 72); rostro 0.8 mm. longo; antheris 0.6 mm. longis. 

Northern Peru (Map 15, triangle). Peru: Dept. Loreto, Balsapuerto, 
altitude about 220 meters, January 1933, Klug 2875 (GH—TYPE; Us, 
F); Dept. Loreto, Yurimaguas, lower Río Huallaga, alt. about 135 meters, 
22 August—9 September 1929, Killip & Smith 27963 (us); lower Río 
Huallaga, Dept. Loreto, October-November 1929, Williams 4698 (F). 

18. EcuiNoponvs magdalenensis Fassett, n. sp., laminis foliorum 
lanceolatis, 3-5 em. longis, 5-9 mm. latis; achaeniis 1.6-1.7 mm. longis, 
costis obscuris aut obsoletis (Fig. 71); rostro 0.6-0.7 mm. longo; antheris 
0.4—0.5 mm. longis. 

In Colombia, in the basin of Río Magdalena and on the nearby Pacific 
coast (Map 15, dots). Colombia: Department of Antioquia, Puerto 
Berrio, alt. 130-140 m., 11-13 January 1918, Pennell 3717 (am, US), 
Department of El Valle, Cuchilla, east of Zarzal, alt. 1200-1600 m., 
22 July 1922, Pennell, Killip & Hazen 8523 (vs—rvrE); Department 
of Valle, costa del Pacifico, Isla del Guayabel, en lo desembocadura 
río Cajambre, 0-5 m. alt., 11 & 12 February 1944, Cuatrecasas 16229 
(us). 


IX. DisrRIBUTION OF CHARACTERS IN SECTION TENELLI 


The morphological relationships as based on nutlets are shown 
in Figs. 66-72. Specific distinctions are shown by double lines 
and varietal by single lines; this is on a basis of how the char- 
acters act, not on how different they look. The rounded summit 
of the achene correlates with the beak poorly developed or 
absent, and plants with 3-ribbed nutlets are quite distinct from 
those with 4-ribbed nutlets, but the short beak grades imper- 
ceptibly into no beak at all and various intermediates are found 
between the ribbed achenes and the ribless ones. Nine different 
combinations of characters are theoretically possible, of which 
5 have been found. 


1955] Fassett,—Echinodorus in the American Tropics 205 
M M Bm N 
~ f ~ j 
M m 
> M / 
R U u $& 
c Pag ib. 4 im 
2 S LM ‘l iin d I~ 
ba | 3 iet PESA PN N 
3 x 68 | XUL IAR 
5 3 as à 
ix / S| E TENELLUS 69 S 
B S| ver ECOSTATUS E./STHMICUS S 
; m 
o 
£ A 
~ 
35 C 
SIS 
D 
Ñ |S 
x 
E. TENELLUS 
var. LATIFOL/US P 
E. QUADRI- a 
COSTATUS = 
x 
m 
Q |= 
SS 
ov 
NIE 
E. MAGDALENENSIS AQ 
3 RIBS NO RIBS 


FIG. 66-72. NUTLETS OF SECTION TENELLI (X 20). Fig. 66. ŒE. tenellus var. 
tenellus; Brazil, Gardner 2740, Fig. 67. E. tenellus var. parvulus; Massachusetts, 
Faxon. Fig. 68. E. tenellus var. ecostatus; Venezuela, Pittier 11707. Fig. 69. E. 
isthmicus; Panama, Killip 3618. Fig. 70. E. tenellus var. latifolius; Venezuela, 
Pittier 5844. Fig. 71. ŒE. magdalenensis; Colombia, Pennell 3717. Fig. 72. E. 


quadricostatus. Peru, Killip & Smith 27963. Specimens in United States National 
Herbarium. 


206 Rhodora [Vor. 57 


\ 
var. PARVULUS | 


16 
75 


var. TENELLUS 


var. LAT/FOL/IUS 
E TENELLUS 


FIG. 73-76, VARIETIES OF E. TENELLUS, NATURAL SIZE, LEAVES OR PLANTS, FROM 
PHOTOGRAPHIC PRINTS OF SPECIMENS IN Us. Fig. 73. Var. parvulus: Florida, Harper 
39. Fig. 74. Var. parvulus: Missouri, Eggert. Fig. 75. Var. tenellus: Venezuela, 
Chardon 252. Fig. 76. Var. latifolius: Dominican Republic, Ekman 11025. 


1955] Fassett,—Echinodorus in the American Tropics 207 


On Maps 16 & 17, characters are mapped, and on Map 18 the 
two previous maps are combined to give a map of the recognized 
taxa. 

Map 16. The long beak, which is correlated with a sloping 
summit of the nutlet, is in a relatively small area in the northern 
Andean region. The short beak and undeveloped beak both 
occur in a great area centering on the Amazon basin, but from 
this region of coéxistence they reach out in slightly different 
directions; the short beak follows the geologically ancient route 
through the Antilles to northern Central America, while the 
undeveloped beak runs from northern South America into the 
younger lands of the Isthmus of Panama. The undeveloped 
beak then reappears in western Cuba and Mexico, ranging thence 
northward on the Atlantic Coastal Plain to Massachusetts and 
in the Mississippi embayment to Missouri. 

Map 17. The species with 4 ribs on the nutlet have been 
found in Peru (altitude 135-220 meters) and in Costa Rica and 
Panama (1199-1250 meters). More widespread is the nutlet 
with 3 ribs. Lack of ribs is confined to the lowlands from 
Venezuela and Colombia to Panama; the ribless condition blends 
on the one hand with the 3-ribbed phase to produce Æ. tenellus 
var. ecostatus (Fig. 68) and combines on the other hand with the 
4-ribbed long-beaked Æ. quadricostatus type to produce E. 
magdalenensis (Fig. 71). 

From the morphological relationships shown in Figs. 66-72, 
and the geographical relationships shown in Maps 16-18, we 
may speculate concerning phylogenetie relationships. Phy- 
logeny is, as is so often the case, more susceptible to speculation 
than to proof. If we hypothesize as ancestral types E. tenellus 
var. tenellus (Fig. 66) mainly of the Amazon basin, and Æ. 
quadricostatus (Fig. 72) of the headwaters of the Amazon, we 
may then see FH. tenellus var. latifolius (Fig. 70) as deriving its 
beak from E. quadricostatus. Var. latifolius then spreads 
through the range of var. tenellus and beyond it through the 
Antilles to Central America. As for E. tenellus var. ecostatus 
and E. magdalenensis, these occupy contiguous areas and are 
both characterized by partial or complete suppression of ribs; 
when there is some suggestion of ribs, Æ. tenellus var. ecostatus 
shows 3 and E. magdalenensis shows 4. It seems unlikely that 


208 Rhodora [Vor. 57 


Map 16. Distribution of beak types. 


= LI 
ena eme! ae 


Map 17. Distribution of rib types. 


1955] Fassett,—Echinodorus in the American Tropics 209 


| E. tenellus vA 
à use] var. parvulus owe 


ara M. CSS 5 


3 ae “sor 
v DE té ———— dcs 
SUP ES 
» j 
E. isfhmicus <, ` ; 
— 


E. tenellus v. ecostatus” 
E magdalenensis ^ 


E. quadricostatus NANSA 


` H 
Sue — 


Map 18. Composite of maps 16 and 17 to show distribution of taxa. 


the same habit of rib suppression should have originated inde- 
pendently in two species in adjacent areas. The greater con- 
stancy of rib suppression in E. magdalenensis suggests that that 
species arose first, perhaps by mutation from an KE. quadricostatus 
stock, and that its character has subsequently been transmitted 
to an E. tenellus stock in the geologically young lowlands of 
Venezuela and adjacent Panama. Æ. isthmicus may be composed 
of another recombination of the characters of E. tenellus and 
E. magdalenensis. 

Another hypothesis is suggested by my colleague Dr. Jonathan 
Sauer, involving £. magdalenensis (Fig. 71) as a primitive type, 
with a genetic make-up involving 4 ribs but with the factor for 
ribs suppressed; this would involve the origin of E. quadricostatus 
(Fig. 72) from E. magdalenensis and an £E. tenellus stock to intro- 
duce expression of the factor for ribs, and of E. isthmicus (Fig. 69) 
and E. tenellus var. ecostatus (Fig. 68) from different combinations 
of characters of E. magdalenensis and E. tenellus. 


210 Rhodora [Vor. 57 


V. Krys FOR IDENTIFICATION IN LIMITED AREAS 


ECHINODORUS IN THE UNITED STATES 
a. Slender plants rarely 10 em. high; carpels 20 or fewer in a loose head; 

stamens 6 or 9; anthers basifixed; nutlets beakless (Fig. 67). 

E. tenellus var. parvulus. 
a. Robust plants with ovate or cordate leaves (except for dwarfed individuals 
on drying shores); carpels many in dense heads; stamens 9-30; anthers 

versatile; nutlets beaked (23-29). 

b. Sepals with smooth veins; profile of fruiting head appearing echinate to 
the naked eye, due to the long (0.5-1.8 mm.) beaks on the nutlets; each 
side of nutlet with 5 arching ribs of which the 2 wing-like ones are alter- 
nate with the others; glands of nutlet acuminate at the upper end and 
entering the base of the beak (Figs. 25-29); pellucid lines of leaves 
mostly less than 1 mm. apart and often several mm. long (Fig. 55); 
scape erect with whorled and usually compound branches. 

c. Beak of nutlet 1.2-2.0 mm. long, 15-24 as long as the body (Figs. 
25-27); anthers 0.8-1.2 mm. long......... E. Berteroi var. Berteroi. 
c. Beak of nutlet 0.5-0.8(-1.0) mm. long, 14-14 as long as the body 
(Figs. 28, 29); anthers 0.5-0.8 mm. long. . E. Berteroi var. lanceolatus. 

b. Sepals with papillose ridges; profile of fruiting heads appearing nearly 
smooth due to the short. (0.2-0.8 mm.) beaks on the nutlets; each side of 
nutlet with 3-4 abruptly curved and sometimes joining ribs of which the 
1 or 2 toward the dorsal edge are wing-like toward the tip (Figs. 23, 24); 
glands of nutlet rounded at both ends and not closely approaching the 
beak; pellucid lines of leaves mostly 1 mm. or more apart and rarely 
exceeding 1 mm. in length (Fig. 54); scape erect when young but soon 
procumbent, with verticillate peduncles at the nodes. .... E. cordifolius. 


ECHINODORUS IN THE WEST INDIES 
a. Carpels 20 or fewer in a loose head; stamens 6-9; anthers basifixed. 
b. Leaves narrowly lanceolate or elliptic (Fig. 76); ribs of nutlets not crested 
(Fig. 700).........ssssssssee ee E. tenellus var. latifolius. 
b. Leaves cordate; ribs of nutlets crested (Fig. 53)... . E. nymphaeifolius. 
a. Carpels many in a dense head; stamens 9-30; anthers versatile. 
c. Veins of leaf radiating from base of the blade (Figs. 2, 8, 10); petiole 
terete or fluted, not 2-winged. 

d. Beak more than !$ as long as the body of the nutlet (Figs. 25-27), 
with tapering apex of glands entering its base; pellucid lines of leaves 
crowded, often several mm. long (Fig. 55)... EK. Berteroi var. Berterot. 

d. Beak shorter, the rounded ends of the glands not reaching its base 
(Figs. 30, 31, 40); leaves with more scattered pellucid short lines 
(Figs. 57-58) or dots (Fig. 56). 

e. Blades with stellate hairs toward the base, about as wide as long 
(Fig. 10). 
Jf. Blades with pellucid dots and few or no very short lines (Fig. 56), 
petals white... 0.00.00 0.0.000.. E. grandiflorus var. grandiflorus. 
f. Blades with pellucid lines 0.2-0.5 mm. long; flowers yellow. 
E. grandiflorus var. aureus. 
e. Blades glabrous, about twice as long as wide (Fig. 13).... E. ovalis. 
c. Upper pair of veins paralleling the midrib for some distance (Fig. 16); 
petiole winged. .......... VEMM ... E. Grisebachüt. 


1955] Fassett,— Echinodorus in the American Tropics 211 


ECHINODORUS IN MEXICO 
a. Carpels 10 or fewer in a loose head; stamens 6-9; anthers basifixed. 
b. Leaves narrowly lanceolate or elliptie (Fig. 73, 74); ribs of nutlet not 
crested (EIE Of) ues oe: oce cd E. tenellus var. parvulus. 
b. Leaves cordate; ribs of nutlet crested (Fig. 53)...... E. nymphaeifolius. 
a. Carpels many in a dense head; stamens 9-30; anthers versatile. 
c. Petiole terete or fluted, not winged (Figs. 1—5), blades with clear pellucid 
dots or lines. 
d. Leaves glabrous, with pellucid lines (Fig. 55). 
e. Nutlets about 24 as wide as long, the summit rounded on one side 
(Fig. 22), with very short beak and round facial glands... . E. virgatus. 
e. Nutlet twice as long as wide, tapered to the very long beak (Figs. 
25-27), the facial glands tapered at apex to enter the base of the 
Denke qus coe DU rs eee E. Berteroi var. Berteroi. 
d. Leaves with stellate hairs and pellucid dots (Fig. 56). 
E. grandiflorus var. grandiflorus. 
c. Petiole winged (Figs. 14, 15); blades with very obscure pellucid lines or 
TARE NERO Te es nnno PETERS E. Andrieuxi. 
ECHINODORUS IN CENTRAL AMERICA 
a. Carpels 20 or fewer in a loose head; stamens 6-9; anthers basifixed. 
b. Leaves narrowly lanceolate or elliptic (Figs. 75, 76); ribs of nutlet not 
crested (Figs. 68-70). 
c. Facial ribs 3 on each side (Fig. 70), or absent (Fig. 68). 
d. Facial ribs nearly or quite suppressed (Fig. 68); nutlets 0.9-1.1 mm. 


long; stylar beak absent or nearly so...... E. tenellus var. ecostatus. 

d. Facial ribs well developed (Fig. 70); nutlets 1.4-1.8 mm. long; 
stylar beak 0.2-0.5 mm. long............ E. tenellus var. latifolius. 

c. Facial ribs 4 on each side (Fig. 69)........0...00.0.0000. E. isthmicus. 
b. Leaves cordate; ribs of nutlets crested (Fig. 53)...... E. nymphaeifolius. 


a. Carpels many in a dense head; stamens 9-30; anthers versatile. 
e. Sepals 12-20-nerved, thin, withering, not accrescent, reflexed or loosely 
ascending in fruit, pellucid markings of leaves, if present, not reticulate. 
f. Leaves with blades cordate at base (Figs. 10-12); petioles terete or 
fluted, not winged; base of blades and summit of petioles with stellate 

hairs or muricate (Fig. 11). 

g. Flowers pedicelled; blades with few or no pellucid lines, sometimes 
with pellucid dots. 

h. Blades with copious pellucid dots (Fig. 56) 
E. grandiflorus var. grandiflorus. 
h. Blades without pellucid markings................ E. muricatus. 

g. Flowers sessile; blades with very short pellucid lines (Fig. 60). 

E. bracteatus var. bracteatus. 
f. Leaves glabrous, with blades tapered (Figs. 14, 16) or subcordate 

(Fig. 15) at base; petioles 2-winged. 

i. Veins radiating from the base of the blade (Figs. 14, 15); leaves 
with pellucid lines very obscure or absent; nutlets long-beaked, 
with 1 facial gland (Figs. 41-43). .................. E. Andrieuzi. 

i. Upper pair of veins paralleling the midrib for some distance (Fig. 
16); leaves with well-marked pellucid lines (Fig. 63); nutlets short- 
beaked, with several facial glands (Fig. 51)......... E. Grisebachit. 

e. Sepals about 30-nerved, thick and brittle, enlarging in fruit to cover the 
fruiting head; pellucid markings reticulate (Fig. 64)....... E. tunicatus. 


212 Rhodora [Vor. 57 


ECHINODORUS IN TROPICAL SOUTH AMERICA 


a. Slender plants rarely 10 cm. high; carpels 20 or fewer in a loose head; 
stamens 6 or 9; anthers basifixed. 
b. Summit of nutlet rounded above the beaklet (Figs. 66, 68, 70); stylar 
beak 0.5 mm. or less long; facial ribs of nutlet present or absent. 
c. Nutlet 0.9-1.1 mm. long; stylar beak nearly absent; anther 0.2-0.6 
mm. long. 
d. Facial ribs well developed (Fig. 66)........ E. tenellus var. tenellus. 
d. Facial ribs nearly or quite suppressed (Fig. 68). 
E. tenellus var. ecostatus. 
c. Nutlet 1.4-1.8 mm. long; stylar beak 0.2-0.5 mm. long (Fig. 70); 
anther 0.5-1.0 mm. long. ................. E. tenellus var. latifolius. 
b. Summit of nutlet horizontal or sloping downward from the beaklet 
(Fig. 71); stylar beak 0.6-0.7 mm. long; facial ribs of nutlet nearly or 
quite obsolete... E. magdalenensis. 
a. Robust plants usually 50 em. or more high; carpels many in dense heads; 
stamens 9-30; anthers versatile. 
€. Leaves with blades more or less cordate at base (Figs. 10-12); petiole 
terete or fluted but not 2-winged; leaves mostly stellate-pubescent or 
muricate about summit of petiole and base of blade. 
f. Flowers long-pedicelled. 
g. Blades with pellucid dots (Fig. 56). . E. grandiflorus var. grandiflorus. 
g. Blades without pellucid markings. 
h. Inflorescence erect, often compound, with pedicels reaching 3 em. 
in length. 22... cc Ie E. muricatus. 
h. Seape creeping, simple, sometimes 2 m. long, rooting at nodes and 
sometimes proliferous, with erect pedicels sometimes reaching 
TOT. ec cc tet e sehe er e ere esee as E. fluitans. 
f. Flowers nearly sessile. 
i. Leaves with very short pellucid lines (Fig. 60) and stellate hairs. 
E. bracteatus var. bracteatus. 
i. Leaves glabrous and without pellucid markings. 
E. bracteatus var. efenestratus. 
e. Blades tapered (Figs. 18-21) or truncate (Fig. 17) at base; petioles 
2-winged. 
j. Veins radiating from the base of the blade (Figs. 17-20); internodes of 
inflorescence sharply keeled but not winged.......... E. paniculatus. 
j. Upper pair of veins running parallel to the midrib for some distance 
(Fig. 21); internodes of inflorescence with 3 herbaceous wings about 
2mm. wide... ... 2.6... ra E. trialatus. 


Volume 57, no. 678, including pages 161—188, was issued 11 July, 1955. 


odora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chiet 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS | 

RALPH CARLETON BEAN Associate Editors 
RICHARD ALDEN HOWARD ( 

CARROLL EMORY WOOD, JR. - 


Vol. 57 August, 1955 No. 680 


CONTENTS: 


Chromosome Numbers in the Genus Sesbania (Leguminosae): 
Evidence for a Conservative Treatment. B. L. Turner...... 213 


Cytotaxonomic Observations on North American Ferns. Warren 
H. Wagner, Jr... aie epee es Ore TE no s n MERE om 219 


The New England Botanical Club, Ince. 


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Vol. 57 August, 1955 No. 680 


CHROMOSOME NUMBERS IN THE GENUS 
SESBANIA (LEGUMINOSAE): 
EVIDENCE FOR A CONSERVATIVE TREATMENT 


B. L. TURNER 


Sesbania is à genus with approximately 50 species occurring 
in the warmer areas of both hemispheres, particularly in wet 
habitats. In many parts of the world some of the species are 
used extensively as green manures for soil improvement. In 
Texas the native species have become troublesome weeds in 
irrigated rice fields. 

The genus has been treated in various ways by taxonomists 
ever since its initial description in 1777. Pollard (1897), Ryd- 
berg (1923), and Jacobs (1941) have reviewed some of the 
pertinent literature, hence only a brief summary of its taxonomic 
history will be given here. 

Bentham and Hooker (1865) recognized the genus as having 
three distinct subgenera or sections: (1) Eusesbania, (2) Dauben- 
tonia, and (3) Glottidium. Taubert (1891) treated Sesbania 
in Engler and Prantl’s Pflanzenfamilien in the same fashion, 
recognizing these three subgenera. A similar treatment was 
followed by most workers until Small (1903), in treating the 
four species found in the United States, recognized the sub- 
genera as distinct genera, thus re-establishing the names Sesban 
(= Sesbania proper—composing the subgenus Eusesbania as 
treated above), Agati (included by most workers in Eusesbania), 
Daubentonia, and the monotypic genus Glottidtum. 

Small’s treatment was based primarily on fruit differences 
among the various taxa: Sesbania (including Agati) with linear, 


214 Rhodora [Vor. 57 


many-seeded, non-winged legumes; Daubentonia with thickened, 
several-seeded, four-winged legumes; Glottidium with thin, two- 
seeded legumes, the seeds remaining in a dry, bladdery, bag- 
like endocarp at maturity. 

Phillips and Hutchinson (1921) concluded, in a revision of the 
African species of Sesbania (23 in number), that the genus was 
best treated semi-conservatively, recognizing in Sesbania the 
subgenera Eusesbania (including Agati) and Daubentonia, 
but agreeing with Small in his treatment of Glottidium as a 
distinet genus. Again, the principal reason for recognition of 
the latter taxon was given as legume morphology. 

In spite of Phillips and Hutchinson’s contribution, Rydberg 
(1924) maintained the four genera, Sesbania (as Sesban), Agati, 
Daubentonia, Glottidium, and, in addition, placed the species 
Sesbania longifolia (Cav.) DC. in a newly created, monotypic 
genus, Daubentoniopsis. 

It is obvious that taxonomic workers have not exhibited any 
degree of unanimity in the recognition of genera or subgenera 
when evidence has been based on external morphology alone. 
As a result, Senn (1938), on the basis of three reported chromo- 
some numbers in the genus, hinted at the possible validity for at 
least some of the segregated genera, stating: “Sesbania with 
n numbers 6, 7, and 16 seems to be in need of thorough study 
as regards the constitution of the genus . . . The occurrence 
of three unrelated chromosome numbers in Sesbania probably 
means that some of these species belong in different genera." 

Senn counted 2n = 12 for the single species of Sesbania 
investigated in his study. Unfortunately, as pointed out by 
Jacobs (1941) and more recently by Rao (1946), Haque (1946), 
and Sampath (1947), the chromosome numbers of n = 16 and 
n = 7 reported by Kawakami (1930), and Krishnaswami and 
Ayyangar (1935) for the other two species, on which Senn’s 
statement was based, were found to be erroneous. When re- 
investigated, the correct number for both the species was found 
to be 2n = 24. Jacobs added two more species counts in his 
study, reporting the number 2n = 12 in both instances. 

Heretofore, the following counts had been established for the 
genus: 


1 Darlington and Janaki-Ammal (1945) list a species, Sesbania australis, as having 
2n = 30, and give Kreuter (1930) as the authority. This is undoubtedly an error 


1955] Turner,— The Genus Sesbania 215 


Fiaures 1-3. Camera lucida drawings of the meiotic chromosomes in Sesbania 
Spp. 1. S. exaltata, la. Univalents indicated by arrows. 1b. Univalents paired 
but without chiasmata. 2. S. drummondii. 3. S. vesicaria. ( X 2000). 


SPECIES 2n AUTHORITY 
Eusesbania 
S. sesban (= S. aegyptiaca) 12  Haque; Jacobs; Rao; Sampath 
S. speciosa 12 Jacobs; Sampath 
S. punctata 12 Frahm-Leliveld (1953) 
S. bispinosa (= S. aculeata) 12, 24 Haque; Jacobs; Rao; Sampath 
S. exaltata (= S. macrocarpa) 12 Atchison (1949); Turner 
(Present paper) 
S. marginata 12  Castronova (1945) 
S. sericea 24 Frahm-Leliveld 
Agati 
S. grandiflora 24 Haque; Jacobs; Rao; Sampath; 
Tjio (1948) 
Daubentonia 
S. punicea 12 Covas and Schnack (1946) 
S. tetraptera 12 Senn 


since reference to the publication cited shows that there is no Sesbania mentioned 
in Kreuter's paper; the report is for Carmichaelia australis, which is properly listed 
by D. & J. on page 163. 


216 Rhodora [Vor. 57 


From the above tabulation, the basic number, x = 6, can be 
inferred for the genus. The species counted include two of the 
usually accepted subgenera, Eusesbania and Daubentonia, as 
well as S. grandiflora, which is sometimes separated from Euses- 
bania and placed in the monotypic subgenus or genus, Agati. 

In the present paper, chromosome numbers are reported 
for three species native to the United States, including S. vesi- 
caria, the monotypic member of the subgenus Glottidium. 
The counts were made from P.M.C. smears. Buds were col- 
lected in 4 chloroform : 3 absolute alcohol : 1 glacial acetic acid 
and allowed to remain for several hours. Young anthers were 
subsequently removed and squashed in acetocarmine. Attempts 
to obtain satisfactory root-tip squashes were unsuccessful. 

Below are listed the sources of materials and corresponding 
n chromosome numbers of the species examined.” 

SPECIES SOURCE n 
5usesbania 
Texas. Travis Co.: Austin 
S. exaltata (Raf.) Cory (Grown from seed collected 6(51121) 
in Austin, Texas). Aug. 17, 
1954. Turner 3655. 
Daubentonia 
Texas. Galveston Co.: 2 mi. 
S. drummondii (Rydb.) Cory N.W. of Texas City. Aug. 8, 6 
1953. Turner 3149. 
Glottidium 
Texas. Travis Co.: Austin. 
S. vesicaria (Jacq.) Ell. (Grown from seed collected 6 
in Giddings, Texas) Aug. 17, 
1953. "Turner 3656. 

Smears from à number of plants of S. exaltata consistently 
showed five bivalents and two univalents at first metaphase 
(Fig. 1). Similar meiotic configurations were reported for 
S. sesban and S. bispinosa by Jacobs. All three of these species 
belong to the subgenus Eusesbania. S. drummondii and S. 
vesicaria both showed 6 bivalents at metaphase. Chromosome 
morphology appeared similar in the three species examined. 


DISCUSSION 
Rollins (1953) has briefly discussed the value and limitations 
of chromosome numbers in the circumscription of plant taxa. 


2 Voucher specimens have been deposited in The University of Texas Herbarium, 
Austin, Texas. 


1955] 'Turner,— The Genus Sesbania 217 


He points out that chromosome numbers are valuable evidence 
for taxonomic purposes in some cases and of little or no im- 
portance in others. The genera in the Leguminosae are notable 
for their constancy. It was only natural that Senn raised the 
question of possible generic validity for those taxa thought to 
have different numbers. Re-examination of these species and 
counts of additional members of all proposed generic segregates, 
except the recently proposed Daubentoniopsis, shows the base 
number to be x = 6. 

The constancy of chromosome numbers in Sesbania does not 
necessarily mean the segregate taxa are not “good” genera; 
on the other hand it does indicate that there is no cytologic 
evidence to justify their segregation. The author agrees with 
Rollins when he states, *' chromosomes provide essentially 
the same kind of evidence to be derived from other parts of the 
plant."  Sesbania, then, has another character that links the 
subgenera together in a single taxon, giving support to such 
legume workers as Bentham and Taubert who considered external 
morphological features alone in their world-wide treatments 
of this genus. It appears that where generic segregation has 
been proposed for these taxa by recent American workers there 
has been a tendency to place excessive weight on the characters 
of the mature pod. From the standpoint of total morphology 
in the various subgenera, the sum of their resemblances far 
exceeds their differences. 

It is hoped that future genetical work and comparative 
studies (embryological and anatomical) will be forthcoming 
so that a more complete synthesis of information will be available 
from which to draw taxonomic conclusions. Until such addi- 
tional work is completed it seems best to treat the proposed 
segregates as subgeneric taxa in Sesbanza. 


SUMMARY 


Chromosome counts of n = 6 for three species of Sesbanza are 
reported: S. exaltata, S. drummondii, S. vesicaria. These species 
belong to the respective subgenera Eusesbania, Daubentonia, 
and Glottidium. Previous chromosome reports for the genus 
have been reviewed. From established counts, a base number 
of r = 6 may be inferred for the genus. Until more evidence 
is forthcoming, it has been concluded that the genus Sesbania 


218 Rhodora (Vou. 57 


is best treated as containing the subgenera, Daubentonia and 
Glottidium, as well as the other generic segregates that have 


been proposed by various authors.—'rHE PLANT RESEARCH INSTITUTE, 
THE UNIVERSITY OF TEXAS, AUSTIN, TEXAS, AND THE CLAYTON FOUNDATION 
FOR RESEARCH. 


LITERATURE CITED 


Arcuison, E. 1949. Chromosome numbers and geographical relationships 
of miscellaneous Leguminosae. J. Elisha Mitchell Sci. Soc. 65: 118-130. 

Bentuam, G. and J. D. Hooker. 1865. Sesbania. Gen. Pl. 1: 502. 

CasrRONOVA, A. 1945. Estudio cariologico de doce especies de Leguminosas 
Argentinas. Darwiniana 7: 38-58. 

Covas, G. and B. ScuNAck. 1946. Numero de chromosomas en autofitas 
de la region de Cuyo (Republica Argentina). Rev. Argent. Agron. 
13: 153-166. 

DanLiNGTON, C. D. and E. K. JaNAKr-AMMaL. 1945. Chromosome Atlas 
of Cultivated Plants. London. 

FrauM-LELIVELD, J. A. 1953. Some chromosome numbers in tropical 
leguminous plants. Euphytica 2: 46—48. 

HaqvE, A. 1946. Chromosome numbers in Sesbania spp. Cur. Sci. 15: 78. 

Jacoss, K. T. 1941. Cytological studies in the genus Sesbania.  Bibliogra- 
phia Genetica 13: 225-297. 

Kawakaml, J. 1930. Chromosome numbers in Leguminosae. Bot. Mag. 
Tokyo 44: 319-328. 

Krevter, E. 1930. Beitrag zu karyologisch—systematischen studien an 
galegeen. Planta 11: 1-44. 

KRrIsHNASWAMI, N. and G. N. RANGASWAMI AYYANGAR. 1935. Chromo- 
some numbers in Sesbania grandiflora Pers. Cur. Sci. 3: 488. 

Puiuurrs, E. P. and J. Hurcuinson. 1921. A revision of the African species 
of Sesbania. Bothalia 1: 40-64. 

Potuarp, C. L. 1897. Studies in the flora of the central gulf region. I. 
Bull. Torr. Bot. Club. 24: 151-154. 

Rao, Y. S. 1946. Chromosome numbers in Sesbania. Cur. Sci. 15: 78. 

Rorrivs, R. C. 1953. Cytogenetical approaches to the study of genera. 
Chronica Botanica 14: 133-139. (Part of a symposium on PLANT 
GENERA, their nature and definition.) 

RYDBERG, P. A. 1923. Genera of North American Fabaceae I. Tribe Galegeae. 
Am. Jour. Bot. 10: 485-498. 

RYDBERG, P.A. 1924. Sesbanianae. N. Amer. Fl. 24: 202-209. 

SAMPATH, S. 1947. Chromosome numbers in Sesbania spp. Cur. Sei. 
16: 30-31. 

SENN, H. A. 1938. Chromosome number relationships in the Leguminosae. 
Bibliographia Genetica 12: 175-336. 

SMALL, J. K. 1903. Flora of the Southeastern United States. New York. 

Tuto, J. H. 1948. The somatic chromosomes of some tropical plants. 
Hereditas 34: 135-146. 


1955] Wagner, Jr.,——North American Ferns 219 


CYTOTAXONOMIC OBSERVATIONS ON 
NORTH AMERICAN FERNS! 


WARREN H. WAGNER, JR. 


DURING the last five years several contributions to our knowl- 
edge of the cytotaxonomy of North American ferns have been 
made. A list by Britton (1953) of thirty chromosome determina- 
tions of ferns collected in southern Ontario was followed by a 
report on the cytology of the Dryopteris spinulosa complex 
by Manton and Walker (1953), and later by a study of the 
Appalachian Aspleniums by the writer (1954). The present 
paper records à number of additional records based largely 
on plants of Michigan (Table I). 

It seems desirable at this time to make a résumé (Table II) of 
our present knowledge of the chromosomes of North American 
ferns, and to point out, at least in some cases, where changes or 
modifications are suggested by the new evidence. The results 
thus far have yielded a number of taxonomic facts which may be 
expected to modify some of the concepts of relationship which 
have traditionally prevailed. A “progress report" pertaining 
to North American ferns is especially appropriate at this time 
since two important new revisions of northeastern American 
ferns have recently appeared, namely those of Gray's Manual 
of Botany, 8th ed. (Fernald, 1950), and the new Britton and 
Brown Illustrated Flora (Morton, zn Gleason, 1952), two revi- 
sions which differ from each other quite significantly in a number 
of taxonomic interpretations and which thus strongly point up 
the need for new information. While groups of species in 
certain of our genera, such as Dryopteris sens. str. and Aspleniwm, 
form natural species complexes of considerable appeal to the 
cytotaxonomist, there are single, isolated species also, such as 
Onoclea sensibilis and Pteretis pensylvanica, which warrant at- 
tention because of their taxonomic interest at the generic and 
familial levels of classification. Of our sizeable species-groups, 
the knowledge of our native Dryopteris, Asplenium, and Botry- 
chium species is being rapidly filled in. Notably absent, how- 
ever, are data on such important and problematic indigenous 
assemblages as Cheilanthes and Woodsia. Of single, isolated 


1 This study was made possible by a Summer Faculty Research Fellowship awarded 
by the Horace H. Rackham School of Graduate Studies, University of Michigan. 


220 Rhodora [Vor. 57 


species, there are certain ones which particularly invite study, 
notably such plants as Lygodium palmatum, Pityrogramma 
triangularis and its forms, and Trichomanes boschianum. That 
Schizaea pusilla, a disjunct species which occurs along the 
eastern seaboard, may have interesting cytological conditions 
is suggested by Selling’s finding (1944) that its spores exist in 
two size groups. 

Several of the most important types of evidence that have 
emerged from the study of chromosomes of ferns should be briefly 
mentioned. First, the data suggest that a given basic number 
will tend to be constant in large, related assemblages, and 
that “aneuploidy tends to characterize the relation between 
genera or groups of genera rather than between species." (Man- 
ton, 1950). For example, the Asplentum group apparently 
has a base number which is uniformly 36, while the two genera 
Dryopteris and Polystichum have 41, Athyrium has 40, and 
Cystopteris 42. Therefore, the determination of chromosome 
numbers may affect decisions concerning the validity of certain 
genera, such as Gymnocarpium which Morton maintained as 
distinct from Dryopteris but which Fernald did not. The 
general problem of generic interpretation in the assemblage 
formerly construed as belonging to one large genus *Dryopteris" 
has assumed special significance since Holttum's proposal that 
some of its elements may be so distinct as to represent a separate 
family, the Thelypteridaceae. This problem will be discussed 
further below. Another type of evidence possible from cyto- 
logical studies of ferns bears on whether or not a given taxon 
is of hybrid origin. Such endemic plants as Cystopteris fragilis 
var. simulans, Botrychium minganense, and B. oneidense have 
certain seemingly intermediate morphological traits which lead 
one to suspect that they may have arisen by hybridization 
between other species. In sterile hybrids the behavior of the 
chromosomes at the time of spore production may sometimes 
lead to knowledge of their parents. In fertile hybrids, the 
chromosome complement may be double that of the parents, 
ie. allopolyploids. A third type of evidence associated with 
chromosomes relates to the evaluation of subspecies or varieties. 
'Thus Britton (op. cit.) has shown that the rare native American 
variety of the hart's-tongue fern, Phyllitis scolopendrium var. 


1955] Wagner, Jr.,—North American Ferns 221 


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Fig. 1, Botrychium multifidum, Cheboygan Co., Mich. (Wiegand); Fig. 2, B. 
multifidum, Alpena Co., Mich.; Fig. 3, B. oneidense, Jackson Co., Mich.; Fig. 4, B. 
oneidense, Saginaw Co., Mich.; Fig. 5, B. dissectum f. dissectum, Saginaw Co., 
Mich.; Fig. 6, B. dissectum f. obliquum, Saginaw Co., Mich.; Fig. 7, B. lunaria var. 
onondagense, Emmet Co., Mich. (Voss); Fig. 8, B. simplex var. tenebrosum, Wash- 
tenaw Co., Mich.; Fig. 9, B. simplex var. simplex, Washtenaw Co., Mich.; Fig. 10, 
B. pumicola, Klamath Co., Oregon (R. D. Brown); Fig. 11, B. matricariaefolium, 
Livingston Co., Mich.; Fig. 12, B. lanceolatum, Emmet Co., Mich. (Voss); Fig. 13, 
B. virginianum, Washtenaw Co., Mich. 


americanum, possesses the tetraploid condition, in contrast 
to the typical European plant which is a diploid. 

I am indebted to a number of people for their interest and 
assistance in this project. In particular, I wish to acknowledge 
the expert help of Mr. Dale J. Hagenah, of the Cranbrook 
Institute of Science, who has contributed numerous living 
specimens for study, and Mr. Walter F. Kleinschmidt and the 
Staff of the Botanical Gardens of the University of Michigan, 
who have cultivated the living plants. Thanks are due also 
to those who supplied material, either fixed cytological specimens 
or living plants, unavailable in the immediate vicinity of Ann 
Arbor, including Mr. Donald F. M. Brown, Mr. Richard M. 


222 Rhodora [Vor. 57 


Brown, Dr. Edward G. Voss, Miss Florence M. Wiegand, 
Dr. Édgar T. Wherry, and Dr. Carroll E. Wood. 

Meruops: Leaflets or other fertile parts were collected in the 
field or in the greenhouse and put in Newcomer's solution (New- 
comer, 1953) prior to squashing in aceto-orcein. Some of the 
earlier collections were fixed in a modification of Carnoy's 
solution (4 parts chloroform, 3 parts alcohol, 1 part glacial 
acetic acid). The results were essentially the same but the 
material seemed to last in a condition suitable for squashing 
for a longer period in Newcomer’s solution. For members 
of the genus Botrychium it was desirable to crush and macerate 
masses of sporangia in the aceto-orcein on a microscope slide, 
and then to remove the large pieces of debris before adding the 
cover-slip. In addition, it was found effective to dry the outer 
surfaces of the Botrychium sporangia first before placing them 
in the stain, as this reduced the tendeney toward precipitation. 
In all cases the top of the cover-slip was tapped with the point 
of a dissecting needle in order to spread out the sporocytes for 
pressing. The slide was heated nearly to boiling, and then 
pressed repeatedly with the forefinger on the surface of the 
cover-slip. In selecting cells to study, those which appeared 
to be intact, i.e., with unfragmented margins, were chosen. 
For each collection several perfect cells were carefully drawn 
with camera lucida at approximately 1450 times magnification 
using an oil-immersion objective, and a 15X widefield ocular. 
Using this technique, the size of the flattened cell cannot, of 
course, be controlled. However, those cells in whieh the ehromo- 
somes were most spread out were usually selected, although 
sometimes only slightly flattened cells showed chromosomes 
clearly and were used (cf. figs. 17 and 18). The voucher speci- 
mens are deposited in the Department of Botany, University 
of Michigan. They were made of the same plants or, in some 
cases, of plants from the same clone or colony, by mounting 
à leaf or leaves on herbarium sheets and pasting a eamera lucida 
drawing of the chromosomes on the same sheet. 

OBSERVATIONS: The genus Botrychium was the focus of con- 
siderable attention in this study because it was suspected that 
two of its species (B. minganense and B. oneidense) might be of 
hybrid origin. Although a number of species and varieties 


1955] Wagner, Jr.,——North American Ferns 223 


are reported here for the first time, there still remain several 
taxa to be investigated. One of the primary difficulties in 
the study of Botrychium is in determining the time of meiosis 
in a given locality and, because of this, collections from a number 
of places turned out to be too far advanced in sporogenesis 
to be of any use in revealing chromosome numbers. Quite 
clearly in Botrychium the monoploid number is usually 45. 
Botrychium virginianum so far appears to be the only exceptional 
species, having an apparent base number of 46. This species 
has been found by Britton, in material from southern Ontario, 
and confirmed by the author from one locality in Michigan, 
to have a chromosome number of n = 92 (fig. 13). The chromo- 
somes themselves appear to be conspicuously smaller than those 
of the other species of Botrychium examined. These cytological 
differences are interesting in view of the morphological features 
that characterize this species, such as the thin-textured blade, 
the partially exposed leaf primordium (the so-called ‘‘bud’’), 
and the small sporangia. It will be profitable to determine 
whether the other members of the subgenus Osmundopteris, 
which, according to Clausen (1938), includes the native B. 
virginianum and four species of other parts of the world, will 
also reveal the base number 46. 

In the subgenus Sceptridium, only two varieties of B. dissectum 
have been previously reported upon (Britton, 1953). The 
question of the related taxon, B. oneidense (Gilbert) House, 
has been an especially vexing one taxonomically since its original 
description by Gilbert around the turn of the century as a 
variety of “Botrychium ternatum.” The latter epithet was 
used in the past to apply jointly to both B. dissectum and to 
B. multifidum, taxa now recognized as wholly distinct species 
by current authors. However, the plant in question, B. onei- 
dense, is at present in the rather paradoxical situation of being 
interpreted by one author (Fernald, 1950) as a form of B. dis- 
sectum (B. dissectum forma oneidense (Gilbert) Clute), and by 
another (Morton, 1952) as a variety of the other species, B. 
multifidum (B. multifidum var. oneidense (Gilb.) Farwell). How- 
ever, Fernald (op. cit., p. 21) describes the plant in question 
as being “embarrassingly transitional" between B. dissectum 
and B. multifidum; and Morton (p. 18) calls it "intermediate 


224 Rhodora [VoL. 57 


TABLE I. CHROMOSOME NUMBERS‘ 


Ophioglossaceae 

Botrychium multifidum (Gmel.) Rupr. n 
Levi Bur's farm, Cheboygan Co., Mich., June 26 (F. M. Wiegand)...45 
Williams Lake Rd., Oakland Co., Mich., June 19..........isuuuu.. 45 
Sharon Hollow, Washtenaw Co., Mich., June 22.................. 45 


Lime sinks near Leer, Alpena Co., Mich., July 3 (D. J. Hagenah)... .45 
B. oneidense (Gilbert) House 


Waterloo-Munith Rd., Jackson Co., Mich., July 18................ 45 

Jackson Rd., s. of Chelsea, Washtenaw Co., Mich., July 30......... 45 

S of Frankenmuth, Saginaw Co., Mich., July 31...........isiuuuuu. 45 
B. dissectum Spreng. f. obliquum (Muhl.) Fern. 

U. of Mich. Arboretum, Ann Arbor, Washtenaw Co., Mich., July....45 

S of Frankenmuth, Saginaw Co., Mich., July 31...........issusuu. 45 
B. dissectum Spreng. f. dissectum 

S of Frankenmuth, Saginaw Co., Mich., July 381............0..... 45 
B. lunaria (L.) Sw. var. onondagense (Underw.) Clute 

Mackinaw City, Emmet Co., Mich., May 15 (E. G. Voss). ..... ca. 90 
B. simplex E. Hitch. var. tenebrosum (A. A. Eat.) Clausen 

Mud Lake, Washtenaw Co., Mich., May 10...................... 45 
B. simplex E. Hitch. (typical) 


Silver Hill Rd., near hdqtrs., Pinkney Area, Washt. Co., May 14....45 

Oakwood Rd., near Sashabow Rd., Oakland Co., Mich., June 19.ea. 45 
B. pumicola Coville 

Llao Rock, Crater Lake, Klamath Co., Oregon, June 19 (R. D. Brown) 45 
B. matricariaefolium A. Br. 


2 mi. w. of Howell, Livingston Co., Mich., May 7.............. ca. 90 

Silver Hill Rd., near hdqtrs., Pinkney Area, Washt. C., May 14..ca. 90 

Maple River Twp., Emmet Co., Mich., May 15 (E. G. Voss). . ..ea. 90 
B. lanceolatum (Gmel.) Angst. ssp. angustisegmentum (P. & M.) Clausen 

Maple River Twp., Emmett Co., Mich., May 15 (E. G. Voss).......45 
B. virginianum (L.) Sw. 

Hankerd Rd., Pinkney Area, Washtenaw Co., Mich., May 15....... 92 


“Polypodiaceae” 
Pteridium aquilinum (L.) Kuhn var. pseudocaudatum (Clute) Heller 
1.5 mi. s. of Lillington on Route 15A, Harnett Co., N. C., gh 


= , 


July 28 (C. E. Wood)... sss 0 
Onoclea sensibilis L. 

Jackson Rd., s. of Chelsea, Washtenaw Co., Mich., July 29......... 37 
Cystopteris bulbifera (L.) Bernh. 

Paint Creek, Ross Co., Ohio, gh, Sept. 5 (D. F. M. Brown)......... 42 

Oxford 40, Oakland Co., Mich., June 17...........sss esses sss 42 


C, fragilis (L.) Bernh. 

Lake Michigamme, Marquette Co., Mich., gh, April (D. J. Hagenah) 84 

Grand Ledge, Eaton Co., Mich., gh, May 16 (D. F. M. Brown)..... 84 
C. dickieana Sim 

Mt. Bohemia, Keweenaw Co., Mich., gh, July 19 (D. J. Hagenah)...84 
C. fragilis var. tennesseensis (Shaver) MeGregor 

Fayette, Delta Co., Mich., gh, July 19, 20 (D. J. Hagenah)........ 126 

Trout Lake, Chippewa Co., Mich., gh, Aug. 13 (D. J. Hagenah)....126 
C. fragilis var. simulans (Weatherby) MeGregor 

Catoctin Furnace, Frederick Co., Md., gh, June 17 (E. T. Wherry) ca. 84 


Bernville, Berks Co., Penn., gh, July 26 (E. T. Wherry)............ 84 
Thelypteris (Phegopteris) hexagonoptera (Michx.) Weatherby 

Sharon Hollow, Washtenaw Co., Mich., June 22.................. 30 

Pine Lake, Hope Twp., Berry Co., Mich., June 25 (D. J. Hagenah)..30 

Jackson Rd., s. of Chelsea, Washtenaw Co., Mich., July 4.......... 30 


* Where chromosome determinations were made from materials grown in the 
greenhouse, the date of meiosis is probably not significant, so those cases where green- 
house plants were used are indicated by “gh”, 


1955] Wagner, Jr.,——North American Ferns 225 


TABLE I.—Continued 
T. noveboracensis (L.) Nieuwl. 


Waterloo-Munith Rd., Jackson Co., Mich., June 25................ 27 
Podunk, Barry Co., Mich., June 29 (D. J. Hagenah)............... 27 
Jackson Rd., s. of Chelsea, Washtenaw Co., Mich., July 4.......... 27 
Proud bake, Oakland Co. Mich July Ira 7 te ee, 27 


T. palustris Schott. var. pubescens (Lawson) Fern. 
M92, near Green Lake, Waterloo Area, Washt. Co., Mich., July 16. .35 
Athyrium filix-femina (L.) Roth var. michauxii Mett. 


Sharon Hollow, Washtenaw Co., Mich., July 2.................... 40 
A. thelypteroides (Michx.) Desv. 

Sharon Hollow, Washtenaw Co., Mich., July 2.........0.....0.00.02, 40 

Jackson Rd., s. of Chelsea, Washtenaw Co., Mich., July 4.......... 40 
A. pyenocarpon (Spreng.) Tidestr. 

Sharon Hollow, Washtenaw Co., Mich., August 3................. 40 
Woodwardia (Anchistea) virginica (L.) Sm. 

Mud Lake, Washtenaw Co., Mich., July 10....................... 35 

Near Stockbridge, Ingham Co., Mich., July 18.................... 35 

Obon Midland’ Co.) Mich. July 31.022 2.0.27 fs i ne 35 


W. (Lorinseria) areolata (L.) Moore 
Eastern North Carolina (plant in U. of N. C. greenhouse), Aug. 15 


COR IDEA Colere DY ae mad ucc c x I-A ccm 35 
Asplenium ruta-muraria L. var. cryptolepis (Fern.) Wherry 

Sunfish Creek, Pike Co., Ohio, gh, May 23 (D. F. M. Brown).......72 
between B. multifidum . . . and B. dissectum var. obliquum 


and possibly a hybrid between them." Obviously it was es- 
pecially desirable to determine whether or not there was cyto- 
logical evidence of hybridity in the presumed intermediate. 

To this end, materials of B. multifidum were studied from 
four different counties of Michigan, of B. oneidense from three 
counties, and of B. dissectum from two counties. For all three 
of the basic taxa under discussion the chromosome number 
proved to be » = 45, as determined in meiosis, There was no 
evidence of hybridity, either in irregular pairing or polyploidy, 
in the materials of B. oneidense which were unquestionably 
typical of this taxon (cf. figs. 1-6). For this reason, and because 
of numerous field observations which Mr. Hagenah and the 
writer have made of the members of this complex in the field, 
I believe that B. oneidense merits interpretation as a distinct 
species, and that the current concepts of this entity (a) as a 
variety or form of B. dissectum, (b) as a variety or form of 
B. multifidum, or (c) as a hybrid between them, may be erroneous. 
'The hybrid hypothesis especially does not seem to be supported 
by cytological evidence. Furthermore, all three entities have 
been found to co-exist in the same locality without evidence 
of intergradation and the presumed “intermediate”? does not 
seem really to be intermediate. It is hoped that in the near 


226 Rhodora [Vor. 57 


future a detailed analysis of the problem of B. oneidense can be 
presented. 

In the subgenus Kubotrychium, only the European form of 
Botrychium lunaria has been reported (Manton, op. cit., n = 45). 
In North America, the major taxonomie problem involving 
this subgenus appears to the writer to be the plant described 
as B. minganense Victorin, which was treated by subsequent 
authors as merely a variety or form of B. lunaria. Although 
its author described it as a distinet species, Clausen (1938) 
reduced it to varietal status under B. lunaria, stating (p. 68) 
that “collections from Michigan, Alberta, and various places 
in the Rocky mountains indicate that, at least in those areas, 
a complete transition occurs." Morton (1952) also interprets 
B. minganense as a variety of B. lunaria, and writes (p. 16) 
that it is “scarcely distinguishable from the typical variety" 
of B. lunaria; Fernald (1950, p. 22) treats it simply as a form. 

Our interest in B. minganense has become heightened by a 
number of field studies which indicate that the Michigan rep- 
resentative is clearly a distinct entity, capable of living sym- 
patrically with typical P. lunaria and entirely without transition. 
In some respects it appears to be intermediate between P. 
lunaria and B. matricariaefolium. Although we are still unable 
to report a chromosome count in B. minganense, its spores have 
been reported to be larger than those of B. lunaria, suggesting 
that it may be a polyploid. However, the latter possibility is 
complicated now by the fact that the single population of B. 
lunaria which has been studied turned out to be a polyploid 
itself, with approximately 90 pairs of chromosomes at meiosis. 
It should be emphasized in this case that the material, from 
Emmet Co., Mich., represents the endemic variety, B. lunaria 
var. onondagense, and material wholly typical of B. lunaria from 
North America has not yet been examined. The problem of 
chromosome numbers in the B. lunaria group is therefore recom- 
mended to workers in various places in the area of occurrence of 
these taxa, roughly from Labrador to Alaska and California, 
east to northern Wisconsin and Michigan. B. matricariaefolium 
has yielded approximately 90 pairs of chromosomes in the three 
populations examined, so it is polyploid like B. lunaria var. 
onondagense. 


1955] Wagner, Jr.,——North American Ferns 227 


The two extreme varieties of Botrychium simplex which occur 
most commonly in Michigan have such different habitats and 
aspects that 1t was suspected that they might prove to be different 
cytologically. Botrychium simplex var. tenebrosum is normally 
a plant of deep shady woods in swamps, whereas B. simplex 
of the typical variety occurs usually in open fields on hillsides 
in relatively dry habitats. Cytologically, however, the two 
varieties proved to be indistinguishable (figs. 8 and 9). The 
closely related B. pumicola, a rare and local species known only 
from a couple of localities in Oregon and California, was found 
also to be like typical diploid Eubotrychiums in its cytology 
(fig. 10). 

Eubotrychium sect. Lanceolatae Clausen comprises a single 
species, B. lanceolatum, which is represented in the northeastern 
United States primarily by its variety angustisegmentum. Speci- 
mens of the latter from Emmet Co., Michigan, showed chromo- 
some complements which were indistinguishable from those 
of members of Subgenus Sceptridium and Sect. Lunariae (fig. 12). 

ONocLEA: The genus Onoclea is monotypic and its single 
specles, O. sensibilis, occurs both in eastern North America 
and in eastern Asia. In morphology it is a unique genus and 
the generic relationships are not clear, although, as pointed 
out by Britton (op. cit), it has been placed in close proximity 
to Pteretis. This interpretation rests largely on the fact that 
the sporangia are produced on much-modified fertile fronds. 
Britton has found » — 37 in Canadian plants, with a possible 
error in interpretation of not more than one bivalent. The 
writer determined 37 pairs in clear preparations of material 
from Washtenaw Co., Mich., thus adding more certainty to this 
number. The large-sized chromosomes of Onoclea are distine- 
tive, and Fig. 15 shows a cell which is only slightly flattened and 
in which the chromosomes themselves have apparently not 
been flattened at all. Britton points out that since the chromo- 
some number of Pteretis pensylvanica is 40, the plants are prob- 
ably not closely related. Research on the comparative mor- 
phology and anatomy of these two genera is strongly recommend- 
ed, to see especially whether they might have acquired their 
similar-appearing fertile fronds through parallel evolution from 
unrelated ancestors in which the fronds are monomorphic. 


228 Rhodora [Vor. 57 


Preripium: Typical plants of Pteridium aquilinum (L.) Kuhn 
from Europe were determined by Manton to have a chromosome 
number of n = 52. More recently Britton has reported that 
the eastern American var. latiusculum (Desv.) Underw. has this 
number likewise; and in the present study a plant of P. aquilinum 
var. pseudocaudatum (Clute) Heller collected by Dr. Carroll 
E. Wood, Jr. near Lillington, N. C., was found also to have 
n — 52 (fig. 14). 

CvsroPTERIS: Some years ago, Mr. Weatherby (1935) dis- 
cussed the forms of the Cystopteris fragilis complex in eastern 
North Ameriea, and he showed that there are a number of 
distinct varieties, including a taxon he described as forma 
simulans Weath., with leaf cutting like C. bulbifera. Since 
that time two rather important findings have been made: (1) 
A peculiar *bulblet-bearing" plant which evidently combines 
characters of C. fragilis and C. bulbifera has been reported from 
a very wide area of Kansas to Pennsylvania and the southern 
shore of Lake Superior (the last localities discovered by D. J. 
Hagenah and as yet not reported in detail). Bulblet-bearing 
plants were named C. tennesseensis by Shaver (1950) and plants 
somewhat like them had been reported earlier by Wherry (1944) 
and Wagner (1944). Since then, McGregor (1950) has reduced 
C. tennesseensis to varietal status under C. fragilis, and he 
points out that it differs from var. simulans (Weath.) McGregor 
primarily in having minute white glands and narrower pinnae. 
It is now known that both forms have *'bulblets" along the 
rachis and occasionally along the costae of the pinnae, although 
var. simulans commonly lacks them. All the authors concerned 
with the relationships of these plants in recent years have 
suggested hybridity, especially on the basis of the “bulblets” 
(which, however, are morphologically quite different from those 
of C. bulbifera)?. (2) The second recent discovery involves an 
overlooked American fern, Cystopteris dickieana Sim. It has 
now been recorded from a number of localities where it had 
previously been undetected because of the microscopic nature 


2 The first mention of bulblets in Cystopteris fragilis that I have found is by Pinker- 
ton, M. Elizabeth, 1933, Ferns and Fern Allies of Missouri, Ann. Missouri Bot. 
Gard, 20: 45-78, who writes (p. 59) under C, fragilis, “An exceedingly variable species. 
Hybridization with C. bulbifera might account for some of the aberrations. Forms 
bearing bulblets have been included under this group as that character does not 
seem to be a constant feature for C. bulbifera alone." 


220 


Fig. 14, Pteridium aquilinum var. pseudocaudatum, Harnett Co., N. C. Note: 
this figure is drawn at approximately two-thirds the magnification of the others, and 
is not to scale. (C. E. Wood); Fig. 15, Onoclea sensibilis, Washtenaw Co., Mich.; 
Fig. 16, Cystopteris bulbifera, Ross Co., Ohio (D. F. M. Brown); Fig. 17, C. fragilis, 
Eaton Co., Mich. (D. F. M. Brown); Fig. 18. C. dickieana, Keweenaw Co., Mich. 
(D. J. Hagenah); Fig. 19, C. fragilis var. simulans, Berks Co., Penn. (E. T. Wherry); 
Fig. 20, C. fragilis var. tennesseensis, Delta Co., Mich. (D. J. Hagenah); Fig. 21, 
Thelypteris hexagonoptera, Washtenaw Co., Mich.; Fig. 22, T. hexagonoptera, 
Barry Co., Mich. (D. J. Hagenah); Fig. 23, 'T. noveboracensis, Jackson Co., Mich.; 
Fig. 24, T. novaboracensis, Oakland Co., Mich.; Fig. 25, 'T. palustris var. pubescens, 
Washtenaw Co., Mich.; Fig. 26, Athyrium filix-femina var. michauxii, Washtenaw 
Co., Mich.; Fig. 27, A. thely pteroides, Washtenaw Co., Mich.; Fig. 28, A. pycnocarpon, 
Washtenaw Co., Mich.; Fig. 29, Woodwardia virginica, Washtenaw Co., Mich. 
(somatic division); Fig. 30, the same (meiotic division); Fig. 31, W. virginica, Ingham 
Co., Mich.; Fig. 32, W. areolata, U. of North Carolina greenhouse (C. E. Wood); 
Fig. 33, Asplenium ruta-muraria, Pike Co., Ohio (D. F. M. Brown). 


of its distinction from typical C. fragilis. It differs from the 
latter species in its spores which are rugose (like those of Woodsza, 
for example) while those of C. fragilis are echinate (Alston, 
1950). The cytological data to be presented here should be 
expanded considerably before we may consider our under- 
standing of North American Cystopteris to be complete from 
this standpoint. 

However, one point emerges clearly in connection with the 


230 Rhodora [Vor. 57 


presumed hybrids of Cystopteris which now favors McGregor’s 
varietal interpretation of the taxa in the “hybrid” group. Cyto- 
logical studies of C. fragilis in Europe and in North America 
(the latter including the reports of n = 84 by Britton and the 
present writer, fig. 17) show that there is so far no evidence 
of a low number of 42, such as that found in C. bulbifera in 
material from southern Ontario and from Michigan and Ohio 
(fig. 16). The European typical C. fragilis not only occurs 
in tetraploid races, but in hexaploid as well. In North America 
only the tetraploids have been found. A fertile hybrid of 
Cystopteris bulbifera and C. fragilis, whether allopolyploid or 
apogamous, would probably be expected to show at least the 
hexaploid eondition of 126 pairs at meiosis. combining the 84 
pairs of C. fragilis with the 42 of C. bulbifera. But such was 
not the case in plants that show the “hybrid” characteristics 
and which conform to MeGregor's concept of var. simulans 
collected by Dr. Wherry at Bernville, Berks Co., Pa. (fig. 19) 
where the number is n = 84. In plants like the foregoing from 
Catoctin Furnace, Frederick Co., Md., the precise number 
has not been yet determined but it is certainly not more than 
two bivalents different from 84 pairs. Accordingly, at least 
two populations do not readily conform to a hybrid hypothesis 
based on current knowledge of the cytology of the two putative 
parents as they occur in North America.’ On the other hand, 
similar *bulblet-bearing" materials from Fayette, Delta Co., 
and from Trout Lake, Chippewa Co., Michigan, which fit 
MeGregor's interpretation of var. tennesseensis in having many 
delicate glands on the leaf surfaces have yielded the hexaploid 
number of chromosomes, showing clearly in a number of prepara- 
tions 126 pairs at meiosis. The latter are the first reports of 
hexaploids in the genus as it occurs in North America. The 
fact that the "hybrids" show two numbers now complicates 

s The possibility should be recognized, however, that there might exist now or 
might have existed at some time in the past more primitive populations of Cystopteris 
fragilis in which the chromosome number was diploid. Thus var, simulans, it might 
be argued, could have arisen from the hybridization of C. bulbifera and a diploid 
C. fragilis, and var. tennesseensis arose from C. bulbifera and a tetraploid C. fragilis. 
(The fact that hexaploids are known in European C. fragilis indicates that almost 
surely there were at one time, at least, diploids in that species.) The wide ranges 
now known in the "hybrids" make it reasonable that they could have originated in 


ancient times. The point is that we do not—at present—have direct evidence of 
diploid C. fragilis in any known North American populations. 


1955] Wagner, Jr.,—North American Ferns 231 


the picture, but McGregor’s interpretation of the "hybrid" 
plants as comprising two taxa seems to be substantiated. 

Of C. dickieana, only one population has thus far been studied 
in North America and this, like the European plants of the 
same taxon studied by Manton (op. cit., pp. 117-120), yielded 
n = 84 (fig. 18). In the writer's opinion the taxonomic treat- 
ment of this plant as a species should be questioned, since up 
to now I am not aware that it has been shown to differ in more 
than a single reliable character from C. fragilis. It is even 
conceivable that the difference between rugose and echinate 
spores in this case depends on merely one or a few genes. 

DRYOPTERIS sens. lat.: One of the major taxonomic problems 
in filieinean classification in general is the proper disposition 
of the generic groups formerly associated with Dryopteris in the 
broad sense, as illustrated locally by Fernald's treatment (1950, 
pp. 31-37) which includes widely divergent plants. He states, 
however, that Dryopteris (s.l. is a 'large and very complex 
world-wide genus, its sections by some considered to form 
genera." Morton (1952, pp. 48-55) considered the eastern 
North American species as belonging to three genera, viz. 
Thelypteris (syn. Lastrea of Copeland, 1947), Gymnocarpium, 
and Dryopteris s.s., and he has given his reasons for this interpre- 
tation elsewhere (Morton, 1950, pp. 214-218). The most 
radical treatment of the generic groups is that of R. E. Holttum 
(1946, pp. 130-133) mentioned above, in which the Thelypteris 
group is interpreted as a distinct family, Thelypteridaceae, 
demarcated from the Dryopteris s. s. group which he deals with 
as a subfamily (Dryopteridoideae) of his Dennstaedtiaceae. 
Such a seemingly “violent” interpretation will no doubt surprise 
many taxonomists versed in traditional fern classification, but 
the cytological evidence accumulated thus far appears indeed 
to warrant a sharp division of some sort between these generic 
groups. Manton has recently concluded (1954, pp. 16-17) that 
“there is very clear evidence that Copeland’s Aspidiaceae is 
polyphyletic at least to the extent that the Thelypteris group 
of genera with » — 31, 34, 35, and 36, are wholly discordant, 
the rest of the ‘family’ being all based on 40, 41, or 42. Holt- 
tum's new family of 'Thelypteridacea is therefore strongly 
supported." With this in mind, the comments in the following 


232 Rhodora [Vor. 57 


discussion will be divided into two parts, the first concerning 
the Thelypteris group and the second the Dryopteris group. 

The chromosome number of the North American representa- 
tive of the type species of T'helypteris (T. palustris var. pubescens 
(Lawson) Fernald) has been reported as » — 35 by Britton in 
Canadian material, and is confirmed by the writer on material 
from Washtenaw Co., Michigan (fig. 25). The two new num- 
bers to be reported here in the Thelypteris group are of special 
interest, including one which establishes the base number of the 
segregate Phegopteris for the first time, and another which is 
lower than any previously reported. 

The *broad beech fern," Thelypteris (Phegopteris) hexagonop- 
tera (Michx.) Slosson is an endemic North American species 
ranging from Quebec and Minnesota south to Florida and 
Texas. It is closely related to T. phegopteris (L.) Slosson, the 
"northern" or “narrow beech fern," a circumboreal species 
which has been found to be apogamous and to possess 90 pairs 
of chromosomes at meiosis in British plants studied by Manton 
and in Canadian plants studied by Britton. Both of the species 
under discussion have also been placed in a separate genus, 
Phegopteris. With regard to the widespread northern species, 
Manton wrote (1950, p. 184) that '"The possible origin of the 
species . . . is at present unknown, since the chromosome 
number (90) is unlike that of any of the genera with which 
Phegopteris has from time to time been classed, and there is 
therefore no clue to its nearest affinities.” It was found that 
the majority of sporangia of this species show only 8 spore 
mother cells, instead of the usual 16, and that both generations 
possess 90 chromosomes, thus indicating the apogamous type 
of life-cycle. 

It is therefore extremely interesting to report that three 
populations of the endemic American species, T. hexagonoptera, 
were found to possess only 30 pairs of chromosomes at meiosis 
(figs. 21, 22). This new number suggests (a) that 30 is probably 
the original, basic number for this pair of species (i. e., for 
Phegopteris as a segregate genus or subgenus), (b) that T. hexa- 
gonoptera very likely has the more primitive, sexual type of 
life-cycle, and (c) that there is some justification from the 
cytological evidence for upholding Phegopteris as a distinct genus. 


1955] Wagner, Jr..—North American Ferns 233 


The chromosomes of the “New York fern," Thelypteris nove- 
boracensis (L.) Nieuwland were examined because Britton (op. 
cit., p. 577) indicated that it was a difficult species to study 
and reported the number of » = 29 + 2 only for interest and 
as a tentative determination. Because also of the divergence 
of this tentative number from the previously reported ones 
of 31, 34, 35, and 36, it seemed desirable to examine several 
populations, and the results that emerged in this investigation 
(figs. 23, 24) turned out to be even more divergent than had 
been expected. The chromosome number of T. noveboracensis 
appears to be unique—n = 27. The chromosome number 
range now known in the Thelypteris group—from 27 to 36— 
indicates either that within this assemblage the generalization 
that chromosome numbers are more or less uniform for related 
groups at the level of genus does not apply here, or that the 
Thelypteris group is polyphyletic, and its members are perhaps 
less closely related phylogenetically than had been supposed. 

In the group of Dryopteris sens. str., it is now well known that 
the basic number of chromosomes of n = 41 is quite different 
from the numbers known in the Thelypteris group with the 
possible exception of Thelypteris (Gymnocarpium) dryopteris 
and T. (G.) robertianum. The primary cytotaxonomic interest 
of Dryopteris in North America concerns the degree of hybridiza- 
tion which has played a role in the evolution of the species. 
It should be stressed that to a considerable extent authors have 
seemingly underestimated the degree to which hybridization 
has occurred within this genus in eastern North America: 
practically no large *woodfern" swamp is lacking in numerous 
intermediate individuals which the critical plant collector will 
inevitably notice, and which often provide “headaches”? in 
identification. Furthermore, these plants which are certainly 
of hybrid origin are apomictic, and are capable over a period 
of years of forming populations by vegetative reproduction. 
Spread of the plants is accomplished by small buds which develop, 
several at a time, at the bases of the withering petioles in the 
autumn, and these are capable of forming successive ‘‘families”’ 
of plants around old individuals. As it stands, there is no 
really thorough-going source for determination of these inter- 
mediate plants in the herbarium, nor, in fact, has it been until 


234 Rhodora [Vor. 57 


very recently that convincing cytological evidence for their 
hybridity has appeared. The attention of pteridologists is 
called to the important work of Manton and Walker (1953), 
not yet completed, which is of major interest to students of 
this group. 

ATHYRIUM: In tropical areas the genus Athyrium often con- 
stitutes a bewildering assemblage taxonomically; in North 
America, however, there are but few species and the major 
problems are at the subspecific level. Such taxonomic work 
as has been done on this group has been largely of a subjective 
nature and little detailed knowledge of the plants concerned 
is available. Intensive investigations are much needed, es- 
pecially among the indigenous forms of the Athyrium filix- 
femina complex, which includes at least several distinctive 
subspecies or varieties, and a large number of forms. Until 
now, only typical A. filix-femina from Britain has been reported 
(n = 40), and A. filix-femina var. michauxii Mett. (syn. A. 
angustum Small) from southern Ontario and Michigan (with the 
same number). 

Of other North American indigenous species of Athyrium 
there are no previous records. Athyrium thelypteroides (Michx.) 
Desv. from two localities in Washtenaw Co., Michigan, yielded 
n = 40. The most distinctive of the native species, A. pycno- 
carpon (Spreng.) Tidestr., of which Morton (1952, p. 43) wrote 
“This species is of uncertain generic relationships,” also showed 
n = 40. There thus appears to be no special evidence from 
chromosome number to support placing A. pycnocarpon in a 
separate genus (e.g., as Diplazium pycnocarpon Broun, or Homo- 
losorus Small). It is interesting to observe that meiosis occurred 
fully a month later (early August) in A. pycnocarpon than it 
did in the two other species which were found growing with 
it in the same locality. In A. pyenocarpon there were not even 
crosiers of fertile fronds evident above the ground level at the 
time fully formed fertile fronds of A. filix-femina and A. thely- 
pteroides were collected in early July. 

Woopwarpia: Britton (op cit., p. 577) wrote of the Virginia 
chainfern, Woodwardia (Anchistea) virginica, that the chromo- 
some number was n = 36, and that “this chromosome number 
may indicate an affinity between this genus and Asplenium.” 


1955] Wagner, Jr.,—North American Ferns 235 


(Such a relationship was suggested earlier by Bower and other 
authors on what seemed to be morphological resemblances.) 
Because of this interesting apparent correlation, and the ready 
availability of this species in southern Michigan, three popula- 
tions were studied. Meiotic figures were observed in all three, 
and in one population mitotic figures of archesporial cells were 
also studied. The results differ from those of Britton: in all 
three cases there was no doubt that the gametie number is 35. 
(In a couple of sporocytes, two unpaired univalents were clearly 
observed, suggesting an occasional early separation of one of 
the bivalents at the time of metaphase, but otherwise in the 
numerous cells examined there was no question concerning the 
number.) Britton indicated in his count, however, a possible 
error of + 1. The present number of 35 does not so evidently 
suggest a relationship of Woodwardia to Asplenium. 

Toward the end of this investigation, Dr. Wood kindly sent 
materials of the very different-appearing species, Woodwardia 
(Lorinseria) areolata, from living plants in the University of 
North Carolina greenhouse. It had been expected that this 
species might perhaps prove to have a different chromosome 
number, especially in view of the fact that the number » — 34 
is now known in Old World species of Blechnum and Wood- 
wardia. However, it turned out that W. areolata has the same 
number as W. virginica, n = 35 (fig. 32), and there is, there- 
fore, no evidence from this standpoint that our two native 
species belong to different genera. 

ASPLENIUM: One of the groups of leptosporangiate ferns which 
has been most thoroughly investigated is the Asplenium group 
characterized by a basic number of 36. Among its species 
apogamy is shown by Asplenium monanthes, intraspecific euploidy 
by A. trichomanes, and allopolyploidy by A. ebenoides and others. 
In the Appalachian spleenworts, three species that show mor- 
phological intermediacy between other species have been found 
to have 72 pairs at meiosis, supporting a hypothesis of allo- 
polyploidy for these taxa (namely, the Alabama population of 
A. ebenoides, A. pinnatifidum, and A. bradleyi). Aneuploidy 
in the Asplenium group was suggested for the first time by 
Britton’s report of n = 35 for Camptosorus rhizophyllus in south- 
ern Ontario. However, four populations tested by the author 


236 Rhodora [Vor. 57 


TABLE II. SUMMARY or CHROMOSOME NUMBERS REPORTED FOR 
NORTH AMERICAN FERNS® 


Ophioglossaceae n 
Botrychium multifidum............... 45 (Wagner, 1955) 
B. oneidense.......0000.0 0020.00.00 0 0200 45 (Wagner, 1955) 
B. dissectum f. obliquum.............. 45 (Britton, '53; Wagner, '55) 
B. dissectum f. dissectum.............. 45 (Britton, '53; Wagner, '55) 
B. lunaria var. onondagense........ca. 90 (Wagner, 755) 
B. simplex var. tenebrosum............ 45 (Wagner, '55) 
B. simplex var. simplex............... 45 (Wagner, '55) 
B. pumicola... noaa auauua aaan 45 (Wagner, '55) 
B. matricariaefolium............... ca. 90 (Wagner, '55) 
B. lanceolatum var. angustisegmentum. .45 (Wagner, '55) 
B. virginianum... 0.00.0... 000000005 92 (Britton, 53; Wagner, 755) 
Osmundaceae 
Osmunda regalis var. spectabilis........ 22 (Britton, '53) 
O. einnamomea......eee ees 22 (Britton, '53) 
“Polypodiaceae” 
Dennstaedtia punetilobula......... 34 + 1 (Britton, '53) 
Pteridium aquilinum var. latiusculum.. 52 (Britton, 753) 
P. aquilinum var. pseudocaudatum..... 52 (Wagner, 755) 
Pellaea atropurpurea..............05.. 87 (Manton, '50) 
P. glabella... 0.00. 116 + 2 (Britton, '53) 
Adiantum pedatum................0.. 29 (Britton, '53) 
Pteretis pensylvanica. ...... liess. 40 + 1 (Britton, 53) 
Onoclea sensibilis................... '..97 (Britton, '53; Wagner, 755) 
Woodsia ilvensis.................41 + 1 (Britton, '53) 
Cystopteris bulbifera.................. 42 (Britton, '53; Wagner, '55) 
C. fragilis; i... iiis sisse 84 (Britton, '53; Wagner, '55) 
C. fragilis var. tennesseensis..........126 (Wagner, '55) 
C. fragilis var. simulans............... 84 (Wagner, '55) 
C. dickieana... 0.0.00 .00000 0000. eee 84 (Wagner, '55) 
Polystichum acrostichoides. ........ sss. 41 (Britton, '53) 
Dryopteris spinulosa...............0.. 82 (Manton & Walker, '53) 
D. intermedia. .................. 82 + 1 (Britton, 753) 
= oe e 41 (Manton & Walker, '53) 
D. fruetuosa... 2.2.2... eee triploid (Manton & Walker, '53) 
D. boottii.... nananana aaa unpaired (Britton, 753) 
JPN triploid (Manton & Walker, '53) 
D. marginalis... naana aana aaa aaa aaa 41 (Britton, '53; Manton & 
Walker, '53) 
D. goldiana....... llle 41 (Manton & Walker, '53) 
D. cristata... naaa ee ee ee 82 (Manton & Walker, '53) 
D. clintoniana...........02.. 00.0060. 123 (Manton & Walker, '53) 
Thelypteris phegopteris............... 90 (Britton, 53) 
T. hexagonoptera.... isses 30 (Wagner, '55) 
T. noveboracensis............0005 29 + 2 (Britton, 93) 
MENU 27 (Wagner, '55) 
T. palustris var. pubescens............ 35 (Britton, '53; Wagner, '55) 


Athyrium filix-foemina var. michauxii...40 (Britton, '53; Wagner, '55) 


5 While this paper was in press, I. Manton and W. A. Sledge's paper entitled 
“Observations on the Cytology and Taxonomy of the Pteridophyte Flora of Ceylon” 
(Phil. Trans. Roy. Soc. London, Ser. B., Biol. Sci. 238: 127—185, 1954) has appeared, 
including the following additional cytological records for North America: Pityro- 
gramma triangularis (California), n = 30; Adiantum pedatum (Vancouver), n = 29; 
and Woodwardia chamissoi Brack. (California), n — 34. 


1955] Wagner, Jr.,—North American Ferns 237 


TABLE II.—Continued 


A. thelypteroides. ...............uieee 40 (Wagner, '55) 
A- DyChODBTDOR ....2....7.--..... 2.23 40 (Wagner, '55) 
Woodwardia (Anchistea) virginica..36 + 1 (Britton, '53) 

VETERES a a a 35 (Wagner, 755) 


W. (Lorinseria) areolata............... 35 (Wagner, '55) 
Asplenium ruta-muraria............... 72 (Wagner, '55 
AMIS VITIC Crean PET EE rr 36 (Britton, '53) 
A. trichomanes................... 36, 72 (Britton, '53) 
A Diatyneurong 9 unge le a a 36 (Wagner, '54) 
'ASMDrAGOICONNENNG E S e iiclree ees 72 (Wagner, '54) 
'AcmmoniBDumee A Lour. eere et tes 36 (Wagner, '54) 
'A-Spinnatitidum s v.a oerte 72 (Wagner, '54) 
"AeErrüdelli eee oio aa triploid (Wagner, '54) 
A. ebenoides.............. (72 univalents) (Wagner, '54) 
— — — —— (Alabama population)...... 72 (Wagner, '54) 
Phyllitisscolopendrium var. american 72 + 2 (Britton, '53) 
Camptosorus rhizophyllus............. 35 (Britton, '53) 
:-—— JM EDD 36 (Wagner, '54) 
Polypodium virginianum.............. 37 (Manton, '50) 


PE E 74 (Britton, '53) 


from Ohio, Virginia, and Indiana, showed the usual aspleniaceous 
number of 36. It should be considered possible that Britton’s 
report was based on an aberrant population; this is a vegeta- 
tively reproducing species, and one which might be capable 
of continued reproduction under abnormal chromosomal con- 
ditions. One of the most interesting of the recent findings, 
mentioned earlier in this paper, was Britton's determination 
of the chromosome number of the rare limestone-inhabiting 
American variety of Phyllitis scolopendrium as tetraploid, in 
contrast to the common European representative which is a 
diploid. 

It was worthwhile to find out whether the ill-distinguished 
American representative of Asplenium ruta-muraria (separated 
by Fernald, without justification in this writer’s opinion, as a 
wholly distinct species, A. cryptolepis) might also prove to be 
distinct cytologically from its European counterpart. The 
results were negative. Ohio material of this plant showed n = 72, 
as do the European plants (fig. 33). 

From the taxonomic viewpoint, it should be emphasized 
here that the recent reduction by Morton (1952, p. 38) of A. 
trudellii to varietal status under A. pinnatifidum has been found 
to be untenable. This conclusion is based not only on grounds 
that previously supported its treatment as a hybrid taxon, 
but on its cytology which also indicated its hybrid nature. 


238 Rhodora [Vor. 57 


There is a triploid chromosome complement present with ap- 
proximately 36 pairs and 36 univalents at meiosis (Wagner, 
1954, pp. 111-113). 

I am more and more inclined to believe that our eustom of 
treating hybrid ferns as binomial taxa (e.g., Asplenium X ebe- 
noides, A. X gravesit, Dryopteris X boottii, etc.) can be justified 
on the grounds that follow: (1) Evidently hybridization has 
played a role in the formation of sexual, allopolyploid species, 
as well as apogamous species, and these species behave in nature 
as independent populations with their own separate ranges and 
ecology, (2) Where the hybrids are sterile, vegetative reproduc- 
tion may enable them to form large clones of importance in the 
plant community, not unlike those of the dandelion in cultivated 
places, and there is some evidence from work now being carried 
out in this laboratory that forms that lack the capacity to 
reproduce vegetatively may reproduce (in a manner as yet 
unexplained) by what appears to be ordinary spore-production, 
in spite of the fact that the spores are normally abortive and 
meiosis is highly irregular. 


CONCLUSIONS 


Forty-eight populations of North American ferns, including 27 species 
and varieties, were examined cytologically in relation to taxonomic 
interpretation. The following cytotaxonomic observations are discussed: 

1. The subgenus Osmundopteris of Botrychium, as represented by B. 
virginianum, seems to be the most distinctive, cytologically, of the sub- 
genera, having an apparent basic number of 46 chromosomes and chromo- 
somes of small size. 

2. The remaining two subgenera of Botrychiwm, namely Sceptridium 
and Kubotrychium appear to be similar to one another cytologically and 
to possess a base number of 45 chromosomes. Polyploidy has now been 
observed in Eubotrychium. 

3. There is no evidence of hybrid origin of B. oneidense (n = 45). On 
the contrary, this taxon appears on the basis of chromosomal and other 
information to behave like a normal species, distinct from both B. dis- 
sectum (n = 45) and B. multifidum (n = 45), and not like a variety of 
either of them. 

4. Inthe group of B. lunaria, B. minganense appears from field studies to 
be a taxon specifically distinet from B. lunaria, although in some respects 
it appears to be intermediate between the latter species and B. matri- 
cariaefolium. | However, cytological data on B. minganense are still not 
available. B. lunaria var. onondagense has proved to be a polyploid with 
n = ca. 90, in contrast to typical B. lunaria of Europe, n = 45. The 
related B. matricariaefolium is also polyploid (n = ca. 90). 


1955] Wagner, Jr.,——North American Ferns 239 


5. Two well-marked varieties of Botrychium simplex, the typical 
variety and var. tenebrosum, show no distinctions in chromosome number 
from each other, and the clcsely related B. pumicola is also indistinguish- 
able (all three with n = 45). 

6. Eubotrychium Sect. Lanceolatae as represented by B. lanceolatum 
ssp. angustisegmentum (n = 45) resembles the species of Sect. Lunariae. 

7. Further research on the relationship of Onoclea sensibilis (n = 37) 
and Pteretis pensylvanica (n = 40) is suggested by the differences in their 
chromosome numbers. 

8. Pteridium aquilinum var. pseudocaudatum (n = 52) is no different 
from the other varieties of the bracken thus far stucied. 

9. A hypothesis of hybridity to explain the ‘‘bulblet-bearing” popula- 
tions of Cystopteris fragilis is not directly supported by presently known 
cytological facts, since C. fragilis in Europe and in the United States has 
not been found to have less than 84 pairs of chromosomes, while C. 
bulbifera has 42. ‘“Bulblet-bearing”’ plants of C. fragilis var. simulans 
show 84 pairs, thus offering no support to an allopolyploid hypothesis. 
On the other hand, the ''bulblet-bearing" C. fragilis var. tennesseensis 
yielded n = 126, justifying the recent taxonomic separation of these two 
varieties. 

10. The American Cystopteris dickieana was found to have n = 84 
chromosomes like its European counterpart. The only difference (exine 
sculpture) thus far demonstrated for this taxon does not seem by itself 
to justify its treatment as a species. 

11. The chromosome number of Thelypteris (Phegopteris) hexagonoptera 
indicates that the base number of the segregate genus Phegopteris (for- 
merly in question because the related T. phegopteris hasn = 90 chromo- 
somes and is apogamous) is n = 30. 

12. Thelypteris noveboracensis with n = 27 is the lowest-numbered form 
yet reported for the Thelypteris group. The latter group now is known 
to have a range in chromosome number from 27 to 36, suggesting that 
either the chromosome numbers in this group tend to fluctuate more 
readily than in other fern groups, or the group is actually polyphyletic. 

13. The species of Hudryopteris are well distinguished by their basic 
chromosome number (n = 41) from the thelypteroid ferns, and the evi- 
dence that has been accumulated in the past several years shows clearly 
that hybridization has played a strong role in the evolution of many of 
the North American populations. 

14. There is no reason especially from chromosome number to warrant 
the placing of Athyrium pycnocarpon in a separate genus from A. thelyp- 
teroides or A. filix-femina since all have n = 40. 

15. Both species of Woodwardia of the northeastern United States 
possess n = 35 chromosomes, in contrast to Old World species of Blechnum 
and Woodwardia. 

16. The ill-defined eastern North American variety of Asplenium ruta- 
muraria with n = 72, is indistinguishable in chromosome number from 
the typical European form. 


240 Rhodora (Vor. 57 


17. Asplenium X trudellii should be considered a hybrid taxon, rather 
than as a variety of A. pinnatifidum, because of its triploid chromosome 
complement and its meiotic behavior. 

18. Our taxonomic custom of treating hybrid ferns as binomials seems 
justified to this writer in view of their capacity to form populations and 
behave as typical sexual or asexual plant species. 


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Ciausen, R. T. 1938. A monograph of the Ophioglossaceae. Torr. 
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CoprELAND, E. B. 1947. Genera Filicum. Waltham, Mass. 

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Horrruwm, R. E. 1946. A revised classification of leptosporangiate 
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Manton, I., and S. WALKER. 1953. Cytology of the Dryopteris 
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———————, 1952. Pteridophyta, in Gleason, H. A., The New Britton 
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SuavER, J. M. 1950. A new fern, Cystopteris tennesseensis sp. nov., 
from Tennessee. Tennessee Acad. Sci. Jour. 25 (2): 106-113. 

Waaner, W. H. 1944. Another occurrence of the apparent hybrid 
Cystopteris. Am. Fern Jour. 34: 125-127. 

———————, 1954. Reticulate evolution in Appalachian Aspleniums. 
Evolution 8: 103-118. 

WEATHERBY, C. A. 1935. A new variety of Cystopteris fragilis and 
some old ones. Rhodora 37: 374-378. 

Wuerry, E. T. 1944. Cystopteris Bluff. Am. Fern Jour. 34: 92-94. 


Volume 57, no. 679, including pages 189—212, was issued 2 August, 1955. 


Dodota 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chiet 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN > Associate Editors 
RICHARD ALDEN HOWARD 

CARROLL EMORY WOOD, JR. 


Vol. 57 September, 1955 No. 681 


CONTENTS: 
The Auriculate-leaved Species of Lesquerella (Cruciferae). 


ECA A Ata a E A erroe a 241 


Plants New to Illinois and to the Chicago Region. Julian A. 
Steyermark and Floyd A. Swink........ eee 265 


The New England Botanical Club, Ine. 


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JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 57 September, 1955 No. 681 


THE AURICULATE-LEAVED SPECIES OF 
LESQUERELLA (CRUCIFERAE) 


REED C. ROLLINS 


AN intensive study of the most easterly species of the genus 
Lesquerella has raised a number of problems regarding the 
taxonomy of those with auriculate cauline leaves. With the 
possible exception of L. lasiocarpa, these are all more closely 
related to each other than to other species of the genus and should 
be formed into a separate section or subgenus. There is now 
definite proof that the flat-podded L. Lescurii is genetically 
very closely related to the very unlike globose-podded L. den- 
sipila. They cross freely in the field (Rollins, 1954) and can 
be readily crossed in the laboratory, producing highly viable 
seeds. Second generation offspring retain the high fertility of 
the Fi, so that there appears to be little or no bar to free gene 
exchange between them. The total evidence (to be presented 
elsewhere) shows without question that L. Lescurit, though 
singular in many of its characteristics, should not form a mono- 
typie section as maintained by Payson (1922) in his monograph 
of the genus. "The earlier treatment of Watson (1888) in which 
L. Lescurii is associated with L. auriculata and L. grandiflora 
is preferable. 

The differentiation of Lesquerella in the Central Basin of 
Tennessee resulting in such morphologically divergent species as 
L. Lescurii, L. densipila and L. perforata has posed questions as 
to their origin and nearest relatives within the genus. Where did 
these Tennessee Lesquerellas come from and by what migratory 
route did they get there? The genus in large part is southwestern 


242 Rhodora [Vor. 57 


and Mexican in distribution with a heavy concentration of 
species in Texas. However, there are a goodly number in the 
Rocky Mountain area and several species in South America. 
But for the relatives of the Tennessee species, we need to look 
no farther west than Texas and Oklahoma. There, among the 
three possible species, L. auriculata appears to be the most 
likely progenitor of the more easterly species. Not only does 
it agree in many characteristics with the Tennessee and Alabama 
species under consideration but the chromosome number of 
n =8 is the same. L. grandiflora, the other possible Texas 
relative, has n = 9. 

The discovery of a new species, L. lyrata, in northwestern 
Alabama is significant in connection with the problems posed 
above. This species, lying geographically and morphologically 
between L. densipila of Tennessee and L. auriculata of Oklahoma 
and Texas, fills a gap as nearly as one could ask for it. L. 
lyrata provides the proper evolutional step from L. auriculata 
to L. densipila as well as a possible remnant of a more or less 
continuous distribution pattern that must have connected the 
Tennessee and southwest species in a past era. It is interesting 
that L. lyrata should only now be found and furthermore that 
L. gracilis! should be discovered in Mississippi for the first time. 

Lesquerella as a whole is under study, and it is hoped that a 
monograph of this interesting genus may eventually be presented. 
'The present paper deals with only the auriculate-leaved species. 
The herbarium citations follow the symbols of Index Herbariorum 
(1954). I am indebted to Dr. L. O. Gaiser and Mrs. Winslow 
Briggs for making the chromosome counts given at the end 
of each description in those instances where it is known. Dr. 
Elsie Quarterman of Vanderbilt University has kindly assisted 
by providing information about the Central Basin of Tennessee 
and by participating in the field work, for which I am grateful. 


1 Last April, in company with Mr. George R. Cooley and Dr. James D. Ray, Jr., 
I examined a large population of L. gracilis (Hook.) Wats. growing on heavy black 
prairie land just north of Okolona in Chickasaw County, Mississippi. Here the 
plants were as thoroughly at home as on the black-land prairies of Texas. Thanks 
to the efforts of Dr. Ray, Mr. Cooley, and Mr. L. J. Brass, there are now three known 
stations for L. gracilis in Mississippi, the southernmost in Lowndes County and the 
northernmost in Pontotoc County, all on the “prairie strip." To my knowledge, 
the genus Lesquerella has not been previously reported from either Mississippi or 
Alabama. 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella 243 


KEY TO THE SPECIES 
A. Pedicels recurved; siliques slightly compressed to flattened contrary to 
septum; filaments not dilated at base; cauline leaves nonauriculate to 
obscurely auriculate, not clasping the stem........... 1. L. lasiocarpa. 
A. Pedicels ascending; siliques globose, subglobose, pyriform, or flattened 
parallel to septum; filaments dilated at base; cauline leaves markedly 
auriculate and clasping the stem. 

B. Siliques strongly flattened parallel to septum; valves pubescent with a 
mixture of large bulbous-based trichomes and an understory of small 
branched trichomes; flowers yellow.................. 2. L. Lescurii. 

B. Siliques globose, subglobose or pyriform, not flattened; valves glabrous 
or uniformly pubescent with simple or forked trichomes; flowers 
yellow or white. 

C. Flowers yellow; siliques globose or subglobose; septum complete. 
D. Siliques densely pubescent with minute simple or forked trichomes; 


styles pubescent; plants of Tennessee. .......... 3. L. densipila. 
D. Siliques glabrous; styles glabrous; plants of Alabama, Oklahoma, 
and Texas. 


E. Ovules 4-8 in each silique, septum dense, opaque; stigma scarcely 
expanded; siliques less than 3 mm. high, depressed globose 
D T.I LII LUC MI LL 4. L. lyrata. 

E. Ovules 12-20 in each silique; septum less dense, translucent; 
stigma markedly expanded; siliques 4-8 mm. high, globose 
to elliptical. 

F. Lower stems hirsute with spreading simple trichomes; lower 
cauline leaves markedly auriculate; petals 4-5 mm. wide; 
infructescences dense. .............2+0+5: 7. L. auriculata. 

F. Lower stems pubescent with appressed branched trichomes; 
lower cauline leaves not auriculate; petals 6-9 mm. wide; 
infructescences lax, greatly elongated... ... 8. L. grandiflora. 

C. Flowers white; siliques pyriform or depressed globose to somewhat 
didymous; septum perforate. 
G. Siliques densely hirsute, slightly didymous to depressed globose; 


valves glabrous on interior; styles hirsute........ 5. L. stonensis. 
G. Siliques glabrous to very sparsely hirsute, pyriform; valves densely 
pubescent on interior; styles glabrous. .......... 6. L. perforata. 


1. Lesquerella lasiocarpa (Hook. ex Gray) Watson, Proc. Amer. Acad. 
23: 251, 1888. 


Annuals, biennials or perennials; stems several to numerous, slender, 
decumbent to procumbent, pubescent with branched trichomes or a 
combination of branched and simple spreading trichomes, 1-5 dm. long; 
basal leaves petiolate, oblanceolate in outline, variable, sinuate dentate 
to somewhat lobed or incised, 3-10 em. long, 1-3 cm. wide, densely pu- 
bescent with small branched trichomes or a mixture of these and large 
trichomes; cauline leaves sessile, narrowed toward base, varying from non- 
auriculate to barely auriculate or with definite auricles present, sinuate 
dentate to incised, obovate elliptical or oblong in outline, densely pubescent, 
1-4 em. long, 0.5-2 cm. wide; infructescence lax; pedicels slender, re- 
curved in fruit, not expanded toward summit, densely pubescent, 1.5- 


244 Rhodora [Vor. 57 


2.5 em. long; sepals green, pubescent with small branched trichomes 
to hirsute, linear-oblong, nearly acuminate, 4-5 mm. long, 1.5-2 mm. 
wide; petals broadly obovate, 6-8 mm. long, 4.5-5.5 mm. wide, light 
yellow, often drying purplish, blade nearly orbicular, limb very short; 
anthers sagittate, 2-3 mm. long; filaments not dilated at base, barely 
exceeding anthers in length; glands formed into horn-like projections 
at the bases of the filaments; siliques sessile, definitely flattened contrary 
to the partition to only slightly flattened, orbicular in outline to elliptical 
or cordate; densely pubescent with a mixture of small branched and 
large simple trichomes or with the simple trichomes absent, 5-9 mm. 
long, 4-9 mm. wide; style 1-1.5 mm. long; stigma expanded; replum 
oblong to nearly elliptical; septum transparent; ovules 8-14 in each 
locule; funiculi attached to septum at base; seeds nearly orbicular, slightly 
longer than broad, margined, ca. 1.5 mm. long; cotyledons accumbent. 


KEY TO THE VARIETIES 
Siliques hirsute with large simple trichomes as well as being pubescent with 
an understory of small branched trichomes. 

Plants annual; caudex not noticeably thickened; cauline leaves scarcely 
cuneate at base, barely auriculate to definitely so; low elevations in 
Texas and coastal eastern and northeastern Mexico. 

Siliques orbicular to elliptical in outline; leaf-lobes acute to acuminate 
Meme e ee aae a la. var. lasiocarpa. 
Siliques cordate in outline; leaf lobes rounded to obtuse, plants of Mexico 
eet er ee oe ths epee wetness were neu ereseentas le. var. ampla. 
Plants biennial or perennial; caudex thick; cauline leaves cuneate at base, 
nonauriculate; mountains and foothills of northeastern Mexico......... 
eee e meae e ee ee ee eee ees 1d. var. heterochroma. 
Siliques pubescent with small branched trichomes only. .1b. var. Berlandieri. 


la. L. lasiocarpa var. lasiocarpa. 


Vesicaria lasiocarpa Hook. ex. Gray, Smithson. Contrib. 5: 13. 1853. 
Synthlipsis Berlandieri var. hispida Watson, Proc. Am. Acad. 17: 321. 


1882. 

Alyssum lasiocarpa Kuntze, Rev. Gen. Pl. 2: 931. 1891. Type seen at 
Kew, but there is no indication of place of collection on the type sheet. 
Gray, l.c., says, "between Bexar and Trinity River, May, 1828, Berlandier 
(in herb Hook.).” 


Texas: Rio Frio north of Dilley, Frio Co., Painter, Lucas & Barkley 14206 
(MO, TEX); 3 mi. south of Dilley, La Salle Co., Painter & Barkley 14305 (Ex); 
Rockport, Arkansas Co., Tharp s.n. (au); Corpus Christi, Nueces Co., Tracy 
9348 in part (NY); same locality, E. J. Palmer 11215 (mo, TEX); same locality, 
Heller 1405 (an, MO); Bishop, Nueces Co., Young s.n. (TEX); Alice, Jim Wells 
Co., E. J. Palmer 11259 (mo, TEX); San Diego, Duval Co., Croft s.n. (GĦ, NY); 
Apache Ranch, ca. 40 mi. s.w. Catarina, Webb Co., Blair et al. 48-500 (TEX); 
6 mi. n. Laredo, Webb Co., K. M. & M. C. Wiegand 750 (au); 6 mi. w. of 
Aguilares, Webb Co., Tharp et al. 51-1693 (rex); King Ranch, Kleberg Co., 
M. C. Johnston 5410 (rex); Kingsville, Kleberg Co., High 57 (mo); 41% mi. e. 
of Hebbronville, Jim Hogg Co., M. C. Johnston 54124 (OKLA, TEX); Donna, 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella 245 


Hidalgo Co., Clover 624 (Ny); 3 mi. n. of Edinburg, Hidalgo Co., Painter & 
Barkley 14448 (TEX); s. of Raymondville, Willacy Co., Gentry 52-658 (TEX); 
w. of Holly Beach, Cameron Co., M. C. Johnston 54201 (TEx); w. of Browns- 
ville, Cameron Co., Hansen 321 (an, MO, NY, TEX). 

Mexico: Hacienda el Carrizo, Nuevo León, Pringle 10236 (an, NY); 23 mi. 
n. of Sabinas Hidalgo, Nuevo León, Webster & Barkley 14591 (Tex); Sierra 
de San Carlos, Tamaulipas, Berlandier 3101 (am): Laredo, Tamaulipas, 
Berlandier 157 (GH); 16 mi. s. of Nuevo Laredo, Tamaulipas, Heard 2 (TEX); 
San Lorenzo de Leguna, 70 mi. s.w. from Parras, Coahuila, E. Palmer 26 
(an, type of Synthlipsis Berlandieri var. hispida). 


Ib. L. lasiocarpa var. Berlandieri (Gray) Payson, Ann. Mo. Bot. 
Gard. 8: 139. 1922. 
Synthlipsis Berlandieri Gray, Bot. Mex. Bound. Surv. 34. 1859. 


Texas: Corpus Christi, Nueces Co., Tracy 9348 (GH, MO, Nx in part); Sanz 
Ranch, Willacy Co., M. C. Johnston 54547 (Tex); San Juan, Hidalgo Co., 
Parks 18003 (aH). Mexico: Matamoros, Tamaulipas, April, 1836, Berlandier 
3017 (GH, type; MO); same location, Berlandier 710, 778, 1517, 2127, 2198, 
3102 (an); 


1c. L. lasiocarpa var. ampla Rollins, var. nov. 

Herba annua; foliis obtusis amplis; siliquis cordatis. 

Mexico: vicinity of Victoria, Tamaulipas, Feb. 1 to Aptil 9, 1907, E. 
Palmer 41 (an, type; Nv, isotype); vicinity of Pueblo Vieja, Vera Cruz, 2 
kilo. s. of Tampico, E. Palmer 366 (au). 

1d. L. lasiocarpa var. heterochroma (Watson) Rollins, comb. nov. 

Synthlipsis heterochroma Watson, Proc. Amer. Acad. 17: 321. 1882. 


Mexico. Nuevo León: Guajuco, 27 mi. s.e. from Monterrey, March, 
1880, E. Palmer 33 (anu, type); Villa de Santiago, Leavenworth 120 (am); 
Linares, J. C. & E. M. Frye 2522a (an); 12 mi. e. of Monterrey, Barkley 14354 
(TEX). 

Lesquerella lasiocarpa is tremendously variable. Not only 
can one find a great range in the size of individual plants but 
every part of the plant seems to run through a rather wide 
range of variation. For example, this species has been com- 
monly thought of as having siliques that are rather markedly 
flattened contrary to the partition, the resulting shape of the 
replum being narrowly oblong. However, the actual amount 
of flattening varies all the way from the rather strongly flattened 
type to siliques that are just short of being round. In the 
latter type, the replum is broadly elliptical to nearly round. 
Other characters are not noticeably correlated with the degree 
of flattening. Many of the plants collected are obviously 


246 Rhodora [Vor. 57 


annuals, but some, as in the type of Watson's Synthlipsis hetero- 
chroma and other specimens from Nuevo León, seem to be 
perennials. Unfortunately, I have had no field experience 
with this partieular species and can only attempt to interpret 
the specimens. 

Although quite definitely a Lesquerella, this species has certain 
features in common with the monotypic genus Synthlipsis and 
suggests a possible ancient connection between the two genera. 
If this view gains further support as evidence concerning the 
phylogeny of the Cruciferae is accumulated, it would point to a 
southern, perhaps Mexican, origin for the genus Lesquerella. 
This idea is in immediate conflict with a widely held view that 
Lesquerella is related to and probably originated from either 
Vesicaria or Alyssum of the Old World. 

The varieties of L. lasiocarpa are not especially marked except 
with relatively minor characters. Variety Berlandieri lacks 
large simple trichomes on the fruits and has a reduced number 
elsewhere on the plants. I am not sure that this is a natural taxon 
for it seems to occur within the range of var. lasiocarpa and has 
even been collected at the same location. Variety ampla is 
more southerly in its range than the other varieties and marks 
a trend toward cordate-shaped fruits and large blunt leaves. 
The simple trichomes on the fruits are not so pronounced as 
in var. lasiocarpa where they are frequently large, stiff and with 
a bulbous base. Payson did not recognize var. ampla as separable 
from var. lasiocarpa although he had available to him the same 
collections that I have studied. However, the Gray Herbarium 
specimen, which he did not see, is the one with mature fruits 
where the cordate shape is strongly evident. 

In var. heterochroma appears a trend of development not seen 
in other varieties, that of a perennial caespitose habit. The 
specimens so far collected all have a comparatively thick caudex. 
A note by one collector (Barkley) indicates that the plants were 
taken near a brook, suggesting a moist habitat. All collections 
appear to have come from above one thousand feet in elevation. 
The other varieties are low elevation plants occurring mainly 
on the Gulf coastal plain. 

Although four varieties are recognized in L. lasiocarpa, the 
present taxonomic treatment leaves considerable to be desired. 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella — 247 


Not only is some explanation for the tremendous variation 
needed but there are trends of differentiation observable, for 
example that toward globose siliques, that require a more 
adequate explanation than can be surmised from specimens 
alone. A genetical and cytological study of this species is 
definitely needed and would contribute profitably to an under- 
standing of the role of L. lasiocarpa in the phylogeny of the 
genus as a whole. 

The cauline leaves of some plants of L. lasiocarpa are non- 
auriculate; in others they are slightly auriculate; and in some, 
definite auricles are present. The questions follow naturally 
as to whether L. lasiocarpa should be included with the definitely 
auriculate species as in the present treatment and whether it is 
more closely related to them than to other species of the genus. 
There is little to go on at present to provide answers to these 
questions except to say that L. lasiocarpa is as nearly in place 
with the auriculate species as with any group of species in the 
genus. 


2. Lesquerella Lescurii (Gray) Watson, Proc. Amer. Acad. 23: 250. 
1888. 

Vesicaria ? Lescurii Gray, Manual Bot. ed. 2, 38. 1856. 

Alyssum Lescurii Gray, Manual Bot. ed. 5, 72. 1867. 

Annual; stems several to numerous, erect or decumbent, simple or 
usually branched, hirsute below, densely pubescent with branched tri- 
chomes above and below, 1-3 dm. long; basal leaves petiolate, lyrate, 
deeply lobed, 3-7 em. long, 5-20 mm. wide, hirsute below with mostly 
simple trichomes, coarsely pubescent above with a mixture of large simple 
and smaller branched trichomes, lateral lobes remote; cauline leaves 
sessile, auriculate, dentate, broadly oblong to ovate, coarsely pubescent 
with large simple and smaller branched trichomes, upper leaves often 
lacking the simple trichomes 0.5-2 cm. long, 3-10 mm. wide; pedicels 
divaricate, nearly straight, densely pubescent with coarse branched 
trichomes, 8-15 mm. long, not expanded at summit; sepals coarsely 
pubescent, yellowish at anthesis, broadly oblong, 3-4 mm. long, 1.5-2 
mm. wide, outer pair slightly saccate, inner pair flat at base; petals yellow, 
obovate, rounded to slightly emarginate at apex, gradually narrowed 
to base with no sharp differentiation into blade and claw, 5-7 mm. long, 
3.5-4.5 mm. wide; glandular tissue in a continuous mold subtending the 
filaments, surrounding the attachment point of single stamens; filaments 
dilated at base, 2.5-3.5 mm. long; anthers oblong, ca. 1 mm. long; siliques 
sessile, strongly flattened parallel to septum, orbicular to slightly louger 
than broad, 4-6 mm. long, 3-4 mm. wide, valves coarsely and densely 
pubescent on the exterior with large simple bulbous-based trichomes 


248 Rhodora [Vor. 57 


and an understory of small branched trichomes, sparsely pubescent with 
minute branched trichomes on the interior; septum complete, dense; 
replum thick, glabrous; style glabrous, 1.5-2 mm. long; stigma greater 
in diameter than the style in dried material, the same diameter as the 
style on the growing plant; ovules 2-4 in each locule; funieuli attached 
to septum at base only; seeds flattened, prominently margined, nearly 
orbicular in outline to slightly longer than broad, 2-2.8 mm. long, 1.5-2 
mm. broad; cotyledons accumbent. N = 8. Plate 1207. 

L. Lescurii is limited in distribution to central Tennessee. The follow- 
ing specimens, supplementing those cited earlier (Rollins, 1952), show a 
considerably wider range than was formerly known. 


Tennessee. Summer County: 4.5 mi. e. of Gallatin, Deam 61336 (DEAM). 
Wilson County: 2 mi. n. of Green Hill, Rollins 5309 (au). Rutherford County: 
Stewart's Creek, 14 mi. e. of Smyrna, Rollins 55174 (cu). Williamson County: 
near Arrington Creek, 14 mi. s.e. of Arrington, Rollins 55116 (GH); near 
Arrington Creek, ca. 7 mi. n. of College Grove, Rollins 65117 (Gu); roadside, 
1 mi. n. of Kirkland, Rollins 5517 (au); near Mill Creek, 3 mi. n. of Nolensville, 
Rollins 65110 (au); 2.5 mi. n. of Triune, Rollins 55111 (GH). Cheatham 
County: 3 mi w. of Ashland City, Rollins 55151 (Gx); flood plain of Cumber- 
land River, 10 mi. n.w. of Ashland City, Rollins 55156 (an); near Pleasant 
View, Quarterman & Stauffer 4948 (au). Davidson County: flood plain of 
South Harpeth River near Linton, Rollins 53134 (an). 


Of the fifty-four mounted specimens of L. Lescurii assembled 
from various herbaria for study, and excepting my own speci- 
mens, all but two were collected within the city of Nashville. 
Actually, the species is abundant in many localities in the six 
counties of the northern portion of the Central [Nashville] Basin 
of Tennessee. It seems strange that documentation for the 
wider range now known is not available in herbaria. 

L. Lescurii appears early in the season, sometimes beginning 
to flower in early March, but most often it is at its flowering 
peak in April. Whole hillsides or open fields may be covered 
by a dense stand of the plants. This species seems to thrive 
equally on the flood plains of the river valleys and on the thin 
soil of open cedar glade-like areas of the hill slopes. 

The branched trichomes present on the interior of the valves 
have not been seen by previous students. This feature is rare 
in the Cruciferae, but is now known in several species of Les- 
querella including L. perforata where it was first noticed in the 
genus (Rollins, 1952). The long bulbous trichomes of the 
exterior valve surfaces are distinetive as may be seen in plate 
1207. These trichomes are attached to a pedicel of tissue that 
projects above the valve wall. An understory of small branched 


Rhodore Plate 1207 


PLATE 1207. Lesquerella Lescurii. "art of an infructescence, X6 (Rollins 55174). Photo by 
Frank White. 


Rhodora Plate 1208 


PLATE 1208. Lesquerella densipila. Part of an infructescence, X6 (Rollins & Quarterman 
65146). Photo by Frank White. 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella — 249 


trichomes is present on young fruits, but these are often shed 
as the siliques mature. 

There is considerable variation from plant to plant in any 
given population of L. Lescurii and trends of variation in dif- 
ferent populations are sometimes recognizable. The intra- 
population variation is most frequently quantitative and may 
often be associated with local environmental factors, such as 
depth of soil, exposure and moisture availability. If conditions 
for prolonged growth are not present, a given plant may be 
small in stature and cease flowering after the setting of a few 
fruits. However, if factors are favorable, growth continues 
and the fruiting racemes become considerably elongated. The 
largest plants are found on the deep soils of the flood plains. 

The taxonomic history of L. Lescurii reflects the doubts of 
Gray as to its rightful generic position. He was undoubtedly 
led to refer this species to Vesicaria in the beginning because 
other species of Lesquerella were being referred there by Hooker 
and other European botanists. The flattened siliques must 
have finally influenced him to move it into Alyssum, but it 
was equally out of place there as Watson finally showed. L. 
Lescurii has now been crossed artificially with L. densipila 
and L. perforata and it hybridizes naturally with L. densipila. 


3. Lesquerella densipila Rollins, Rhodora 54: 186. 1952. 


Annual; stems several to numerous, erect or the outer decumbent at 
base, simple or branched, purplish below, 1-4 dm. high, hirsute below 
with spreading simple trichomes, rachis of inflorescence and upper portion 
of stems hirsute with smaller less spreading and frequently branched 
trichomes; basal leaves petiolate, lyrately pinnatifid to pinnately lobed, 
obtuse, 4-8 em. long, 1-1.8 em. wide, terminal lobe comparatively large, 
lateral lobes deeurrent on rachis, hirsute on upper surface with mostly 
simple trichomes, lower surface with a mixture of large simple and smaller 
branched trichomes; cauline leaves sessile, auriculate, broadly ovate to 
oblong, 1-3 em. long, 0.5-1.5 em. broad, lower broadly obtuse, upper 
smaller and tending toward acuteness, dentate to nearly lobed, hirsute 
on both surfaces with mostly simple spreading trichomes; inflorescence 
racemose, 1-2 dm. long; fruiting pedicels divaricately ascending, straight, 
expanded at summit, 1-2 em. long, pubescent with a mixture of simple 
and branched trichomes; sepals yellowish, nonsaccate, sparsely to generally 
covered with appressed branched trichomes, often with spreading single 
trichomes in addition, oblong, alternating members flat and boat-shaped, 
narrowed toward apex but remaining rounded, 2.5-4 mm. long, 1.5-2 mm. 
wide; petals yellow, broadly obovate, not markedly differentiated into 


250 Rhodora [Vor. 57 


blade and claw, 6-8 mm. long, 4-5 mm. wide; filaments strongly dilated 
at base, attached to anthers just below middle, anthers nearly versatile, 
not sagittate, ca. 1.5 mm. long; glandular tissue in a thin continuous 
mold beneath stamens, forming projections between single and paired 
stamens and an abbreviated ring around the base of the filament of the 
single stamens; siliques subglobose to slightly broader than long, un- 
compressed, 3-4 mm. in diameter, densely pubescent with minute simple 
or forked spreading trichomes; styles 2-3 mm. long, pubescent sometimes, 
glabrous above, slightly expanded into a capitate stigma; 2-4 ovules 
in each loeulus, funieuli free except at their very base; septum entire; 
replum nearly orbieular, pubescent; seeds margined, flattened, orbicular 
to slightly longer than broad, brown, 2-2.5 mm. in diameter; cotyledons 
accumbent; radicle short. N = 8. Plate 1208. 


Tennessee. Rutherford County: 4 mi s.e. of Murfreesboro, Rollins & 
De Selm 55122 (au); near West Fork of Stone's River, 1.3 mi. s.w. of Bethel, 
Rollins & De Selm 55124 (an); near West Fork of Stone's River where state 
route 102 crosses, Rollins & Bold 55134 (anu); floodplain of the Harpeth River, 
1 mi. n. of Eagleville, Rollins 5526 (au). Bedford County: roadside near 
Alexander Creek, 2 mi. n. of the North Fork of Duck River, Rollins 5537 (Gu). 
Williamson County: 1 mi. n. of College Grove, R. C. & D. Rollins 5215 (Gu); 
L5 mi. n. of Kirkland, between Triune and College Grove, Rollins 5315 (an); 
same locality, Rollins 5518 and 55112 (au); Kirkland, Rollins 53138 (am); 
3 mi. s.e. of Kirkland, Rollins 5519 (au); 6 mi. n. w. of College Grove near 
the Harpeth River, Rollins 5514 and 55114 (au). Marshall County: Duck 
River bottom, north of Verona, Sharp, Felix and Adams 11187 (Gu, type; 
ut, isotype); 1 mi. n. of Chapel Hill, Rollins 5319 (au); near Duck River, 3 
mi. s. of Chapel Hill, R. C. & D. Rollins 5217 (au); Lo mi. s. of Duck River, 
ca. 5 mi. s. of Chapel Hill, Rollins 5321 (aH). Maury County: flood plain 
of the Duck River, 10 mi. n.w. of Lewisburg, Rollins 5539 (Gu); open field 
and cedar glade, 4 mi. w. of Columbia, Rollins 55108 (GH); same location, 
Rollins and Quarterman 55146 (GH). 


Though originally thought to be extremely local in its occur- 
rence, L. densipila now proves to have a range in area approach- 
ing that of L. Lescurii. Recent collections from Rutherford, 
Bedford and Maury Counties provide the evidence for a con- 
siderably enlarged geographic area as compared to that known 
at the time the species was described in 1952. The species 
is confined to the Central Basin of Tennessee with its predomi- 
nance of limestone soils. 

The populations of L. densipila usually consist of a large 
number of individuals often numbered in the tens of thousands. 
These may be in open glade-like areas or along the river and 
stream bottoms which are subjected to spring flooding each 
year. Asin L. Lescurii, there is considerable variation between 
individuals and between populations. These are principally 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella 251 


single character variations which can be seen from plant to 
plant, but there are few correlated differences between indi- 
viduals of the same populations. "Thus, independent assortment 
of genetic characters seems to be the rule. L. dens?pila is largely 
self incompatible, thereby insuring a continuing intermixing 
of the genes of any given population. Where different popula- 
tions are involved, there are some minor correlated character- 
istics. However, L. densipila 1s an easily recognized species, 
and the characters that make it distinguishable from other 
species are present in every population. The exception to 
this comes in the hybrid populations that result from the natural 
crossing of L. densipila and L. Lescurii and of L. densipila and 
L. stonensis. 

Lesquerella densipila and L. Lescurii come together at the 
junction of Arrington Creek, where the latter is found in abun- 
dance, and the Harpeth River, where L. densipila is abundantly 
found. From the entrance of Arrington Creek on down the 
Harpeth River, there are numerous populations of hybrids 
that have resulted from the crossing of these species. In these, 
the variation is very wide and the characters, for the most part, 
segregate independently. The fruit shape on most of the 
plants is somewhere between that of being extremely flattened 
which is characteristic of L. Lescuri? and the globose type that 
is found in L. densipila. Occasional hybrid plants have flattened 
siliques, and in occasional ones the siliques are globose. The 
latter may be hirsute with very long trichomes, and the flattened 
type may lack simple trichomes altogether thus combining 
in different ways these characters of both species. In like 
manner, many other characteristics are combined in a complex 
series. Data supporting these conclusions have been given 
previously (Rollins, 1954). 

Natural hybridization between £L. densipila and L. stonensis 
has not been previously demonstrated, and the facts are at 
present being worked out.  Lesquerella densipila var. maxima 
was based on hybrid plants from a population on Stones River 
whose variation was recognized, but was inexplicable at the 
time of publication. Now sufficient field work has been done 
to show that L. densipila is present on the West Fork of Stones 
River and some of the smaller tributaries to it. L. stonensis 


252 Rhodora [Vor. 57 


is so far known only from the East Fork of Stones River. These 
two species meet at the junction of the east and west forks of 
the river, and from that point down river hybrid populations 
are produced which apparently quickly incorporate the genetic 
makeup of stray plants of either species that come within range 
of cross pollination. The hybrids of L. densipila X L. stonensis 
have previously been referred to under the name L. dens?pila var. 


maxima. 
4. Lesquerella lyrata Rollins, sp. nov. 


Annual, stems one to several, usually simple, erect, outer decumbent 
at base, 1-3 dm. long, densely hirsute below with simple spreading tri- 
chomes, upwards becoming a mixture of large simple and smaller branched 
trichomes with the small branched trichomes predominating above; 
basal leaves petioled, lyrate, 2-7 em. long, 6-15 mm. wide, hirsute with 
simple trichomes or with a mixture of small branched and large simple 
trichomes, terminal lobe large and orbicular to elliptical; cauline leaves 
sessile, auriculate, clasping, ovate to broadly oblong, obtuse, nearly 
entire to coarsely dentate, 5-20 mm. long, 4-10 mm. wide, densely to 
sparsely hirsute with simple or simple and branched trichomes; inflores- 
cences dense; pedicels slender, straight, divaricately ascending, densely 
pubescent, 1-1.5 em. long; sepals pubescent, spreading and yellowish 
at anthesis, oblong, 3-4 mm. long, 1.2-1.5 mm. wide, outer pair very 
slightly saccate at base; petals yellow, broadly obovate, 5-7 mm. long, 
3.5-4 mm. wide, slightly rounded, truncate or shallowly retuse at apex, 
limb short; glandular tissue subtending paired stamens, surrounding 
single stamens; paired stamens ca. 4 mm. long, single stamens ca. 3 mm. 
long; filaments dilated at base, those of paired stamens 3-3.5 mm. long, 
those of single stamens 2-2.5 mm. long; siliques depressed globose to 
subglobose, often slightly depressed at base of style and slightly didymous, 
glabrous, sessile, 2.5-3 mm. high, 3-4 mm. broad; styles slender, glabrous 
1-1.5 mm. long; stigma unexpanded, nearly same diameter as style; 
replum orbicular to slightly broader than high; ovules 2-4 in each locule, 
funiculi free; seeds flattened, oval to nearly orbicular in outline, margined, 
brown, variable in size, 1.5-2.5 mm. on longest dimension, larger seeds 
in capsules with 2 per locule; cotyledons aecumbent. N — 8. Plate 1209. 


Herba annua; caulibus erectis vel decumbentibus simplicibus vel rare 
ramosis hirsutis 1-3 dm. longis; foliis radicalibus lyratis petiolatis hirsutis 
2-7 cm. longis, 6-15 mm. latis; foliis caulinis sessilibus auriculatis oblongis 
vel ovatis obtusis pubescentibus dentatis vel integris 5-20 mm. longis, 4-10 
mm. latis; pedicellis ascendentibus divaricatis pubescentibus, 1-1.5 em. 
longis; sepalis flavis oblongis pubescentibus 3-4 mm. longis; petalis luteis 
late obovatis 5-7 mm. longis, 3.5-4 mm. latis; siliquis subglobosis depressis 
glabris 2.5-3 mm. altis, 3-4 mm. latis; stylis tenuibus glabris 1-1.5 mm. longis; 
loculis 2-4-ovulatis; seminibus marginatis brunneis compressis; cotyledonibus 
accumbentibus. 

Alabama. Franklin County: cedar glade, roadside and field, near Richard- 
son's Crossing, 7 miles east of Russellville, April 16, 1955, Reed C. Rollins, 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella 253 


George R. Cooley and L. J. Brass 5599 (an, type); same location, April 5, 1955, 
Rollins 5548 (au); same location, April 27, 1955, Rollins 55188 (Gu) ; bottom land 
of creek, 4 miles north of Richardson's Crossing, about 9 miles east of Russell- 
ville, April 27, 1955, Rollins 55187 (an). Franklin-Moulton County line: road- 
side near cedar covered hill, 1.5 miles east of Newburg, April 5, 1955, Rollins 
5547 (aH). 

Lesquerella has not heretofore been known from Alabama. 
However, I was led deliberately to look for it there because of 
the close parallel in the over all distribution pattern of the most 
easterly auriculate species of Lesquerella and of the genus Leaven- 
worthia. The latter has been known from several stations 
in northwestern Alabama for some time. It seemed probable 
that if Lesquerella did occur between Tennessee and Texas, 
it would be on limestone or soils of limestone origin. The 
Leavenworthia stations in Moulton and Franklin Counties were 
picked as target areas for the search.  Lesquerella was first 
found on April 5th near the Moulton-Franklin County line, 
as I proceeded westward on Alabama State Route 24. Here 
there was a small population of a few dozen plants extending 
along the roadside for about fifty yards. The glabrous fruits 
immediately indicated this Lesquerella was not the same species 
as any of those known from Tennessee, although the auriculate 
cauline leaves left no doubt about its being related to them. 
However, there remained the possibility that it was an outlying 
station of L. auriculata a species found chiefly in Oklahoma, 
but extending into Texas. My enthusiasm was aroused and 
I began to search in all directions to find additional populations. 
As the area of my search grew larger, I moved gradually west 
beyond the small village of Newburg, but no new populations 
were encountered until Richardson’s Crossing was reached. 
At this location, marked by a lone roadside building housing the 
Richardson family and a general store, Lesquerella was found 
in abundance. This population, occurring with Leavenworthia, 
Sedum and similar cedar glade inhabiting plants, extended over 
a sizeable area of the glade itself, spilling up onto the roadside 
and over much of an adjacent old cotton field. Here was 
adequate material for field study, a mass collection and cyto- 
logical fixations. On April 16th I had the pleasure of showing 
this station to Mr. George R. Cooley and Mr. L. J. Brass, at 
which time the type series was collected. 


254 Rhodora [Vor. 57 


Lesquerella lyrata is most nearly related to L. densipila on 
the one hand and to L. auriculata on the other. The siliques 
are on the average smaller than in either of these species, being 
much smaller than in L. auriculata. Both the latter and L. 
lyrata have glabrous siliques, but they are fundamentally sepa- 
rated by ovule number. In L. auriculata there are from 12 to 20 
ovules in each silique, while in L. lyrata there are only 4 to 8. 
The basal leaves are quite different in the two. Those of L. auri- 
culata are often spatulate and nearly entire but there is variation 
to a sinuate dentate leaf similar to that of L. grandiflora. How- 
ever in all of the material I have seen, the basal leaves of L. 
lyrata are definitely lyrate with a large terminal obtuse lobe 
and much reduced more or less acute lobes or teeth along the 
margins. 

Aside from ovule number, perhaps the most noticeable dif- 
ferences between L. auriculata and L. lyrata are in silique size 
and shape. In the former, the siliques are longer than broad, 
elliptical in outline, with a rounded apex, while in the latter 
they are broader than long and often with a slight depression 
at the base of the style. As to size, the fruit of L. lyrata measures 
2.5-3 mm. high, whereas in L. auriculata, it ranges from 4-6(-8) 
mm. high. The stigma is rather strongly expanded in L. auri- 
culata being at least twice the diameter of the style. In L. 
lyrata, the stigma is very slightly expanded beyond the style 
diameter itself. 

The siliques of L. lyrata are only slightly smaller than in 
L. densipila and they are of the same general shape although 
perhaps more markedly depressed globose to nearly didymous 
than in the most extreme populations of the latter. Also, 
the ovule number runs about the same in the two species, so 
far as extremes are concerned. No data have been assembled 
on the point but I have the impression that 7 or 8 ovules per 
silique occur most frequently in L. densipila while the most 
frequent number in L. lyrata is 4. The one very marked dif- 
ference between these species is the indument on the exterior of 
the siliques and on the styles. In L. lyrata, both are perfectly 
glabrous. In L. densipila, the exteriors of the siliques are densely 
pilose with minute trichomes and the styles are hirsute. 

All of the known facts of morphology point to a closer relation- 
ship between L. denszpila and L. lyrata than between the latter 


Rhodor: Plate 1209 


PLATE 1209. Lesquerella lyrata. Part of an infructescence, X6 (Rollins, Cooley & Brass 5599). 
Photo by Frank White. 


Rhodora Plate 1210 


PLATE 1210. Lesquerella  stonensis. Part of an infructescence, X6 (Rollins, 55176). 
Photo by Frank White. 


LÀ 


1955] Rollins, —Auriculate-leaved Species of Lesquerella 255 


and L. auriculata. This presumed affinity has not been tested 
experimentally. The chromosome numbers of all three are 
the same, n — 8. 


5. Lesquerella stonensis Rollins, sp. nov. 


Annual, stems several from the base, simple or usually branched, 2-4 
dm. long, erect, the outer usually decumbent at base, densely hirsute 
with simple trichomes below, densely pubescent above with simple or a 
combination of simple, forked and more lightly branched trichomes; 
basal leaves petioled, lyrately lobed to pinnatifid, obtuse, 3-6 cm. long, 
8-15 mm. wide, densely hirsute, simple trichomes present on upper surface, 
a mixture of long simple and shorter forked or branched trichomes present 
on lower surface, lobes variable, terminal lobe large and ovate to nearly 
orbicular, entire to sinuate dentate, lateral lobes triangular to broadly 
oblong, obtuse, decurrent on the leaf rachis; cauline leaves sessile, auric- 
ulate, clasping, broadly oblong to ovate dentate, 1-5 em. long, 5-15 mm. 
wide, lower cauline leaves densely hirsute with predominantly simple 
trichomes on upper surface, a mixture of simple, forked and branched 
trichomes on lower surface, upper cauline leaves densely pubescent with 
predominantly forked or branched trichomes; infructescence 8-15 cm. 
long, rachis densely pubescent with mostly branched or forked trichomes; 
pedicels straight or nearly so, divaricately ascending, densely pubescent, 
1-2.5 em. long; sepals oblong to nearly ovate, narrowed toward apex, 
pubescent, 4.5-5.5 mm. long, 1.5-2 mm. wide, outer pair slightly saccate, 
inner pair narrower and nonsaccate; petals white with a yellowish short 
claw, obovate, rounded to emarginate at apex, 7-9 mm. long, 5-6 mm. 
wide; filaments dilated at base, those of the paired stamens 4—4.5 mm. 
long, those of single stamens 3-3.5 mm. long; glandular tissue continuous, 
surrounding insertion point of single stamens, subtending insertion point 
of paired stamens; siliques depressed globose to slightly didymous, densely 
hirsute with simple trichomes, 4-5 mm. broad, 3-4 mm. high; edge of 
replum hirsute, not raised above valve margins; valves glabrous on 
interior; styles hirsute at least below, slender, ca. 2 mm. long; stigma 
expanded; replum orbicular to slightly wider than long, rounded at apex; 
septum usually perforate, sometimes entire, perforation variable in size; 
funiculi not attached to septum; ovules 4-6 in each locule; seeds dark 
brown, flattened, oval in outline, margined, 1.8-2 mm. long, ca. 1.5 mm. 
wide; cotyledons aceumbent. Plate 1210. 


Herba annua, caulibus ramosis vel rare simplicibus erectis vel deeumbentibus 
hirsutis 2-4 dm. altis; foliis radicalibus petiolatis lyratis hirsutis 3-6 cm. 
longis, 8-15 mm. latis; foliis caulinis sessilibus auriculatis dentatis oblongis 
vel ovatis 1-5 em. longis, 5-15 mm. latis; pedicellis divaricatis pubescentibus 
1-2.5 em. longis; sepalis oblongis vel ovatis pubescentibus 4.5-5.5 mm. longis, 
1.5-2 mm. latis; petalis albis obovatis 7-9 mm. longis, 5-6 mm. latis; siliquis 
subglobosis depressis dense hirsutis 4-5 mm. latis, 3-4 mm. altis; stylis hirsutis 
ca. 2 mm. longis; loculis 4—6-ovulatis; seminibus marginatis, brunneis com- 
pressis 1.8-2 mm. longis, ca. 1.5 mm. latis; cotyledonibus accumbentibus. 


256 Rhodora [Vor. 57 


Tennessee. Rutherford County: field ana flood plain, East Fork of 
Stones River, Walterhill, April 26, 1955, Reed C. Rollins 55176 (au, type); 
same location, April 1, 1955, Rollins 5507 (am); pasture and hilltop, near 
East Fork of Stones River, 415 mi. n.w. of Walterhill, April 1, 1955, Rollins 
5508 (an); east of Old Jefferson near point where state route 102 crosses East 
Fork of Stones River, April 22, 1955, Reed C. Rollins and Harold Bold 55138 
(au); pasture near East Fork of Stones River, 1 mi. s.w. of Lascassas, April 
26, 1955, Rollins 55177 (Gu). 

The variability of populations of Lesquerella densipila var. 
mazima noticeable in the field first suggested that these were of 
hybrid origin. In publishing this variety (Rollins, 1952) I 
called attention to the divergence in trichome length on different 
plants of the same population. Later, in working up data 
compiled in seeking to discover species that might have figured 
in the origin of var. maxima, it became clear that while L. 
densipila was probably involved as one parental species, there 
was no known species in the Central Basin of Tennessee that 
could have played the role of the other parent. There were 
characters such as fruit shape, trichome size and trichome 
distribution that could not have arisen from L. densipila or any 
other known species. In the spring of 1955, working on the 
hypothesis that the plants of var. maxima were of hybrid origin 
and that an unknown Lesquerella was involved in producing 
these hybrid plants, I sought to find the unknown. L. stonensis 
is that heretofore unknown species. 

The situation with regard to the hybrid origin of var. maxima 
is now fairly clear, though a complete study has not been made, 
up to the present. Lesquerella stonensis exists as a pure species 
at various localities along the East Fork of Stones River. L. 
densipila exists as a pure species on the West Fork of Stones River 
and to the south and west of this area in the Central Basin. Both 
species occupy flood plain habitats and presumably their most 
important direction of movement is downstream, the seeds 
being carried during the annual spring floods. "The two species 
come together where the two forks of Stones River meet and 
hybridization takes place there. From the junction of the 
East and West Forks downstream, all of the populations exam- 
ined showed evidences of their hybrid origin. 

Lesquerella stonensis is a white flowered species, most closely 
related, as shown by most of its characteristics, to the yellow 
flowered L. densipila. It differs not only in flower color from 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella — 257 


Fig. A-G.  Lesquerella perforata. A—habit sketch X14. B—flower X3. C— 
replum and perforate septum X4. D—F—trichomes of the stems and leaves X125. 
G—silique X4. Drawings by B. Tugendhat. 


258 Rhodora [Vor. 57 


L. densipila but also in the size and shape of the siliques and in 
the length of the trichomes on the valves of the siliques. In 
L. stonensis, the larger fruits are depressed globose to slightly 
didymous while the smaller fruits of L. densipila are globose 
tosubglobose. On the valve surfaces, a dense covering of minute 
simple or forked trichomes is present in L. densipila, but in L. 
stonensis the simple trichomes are much longer and form a less 
dense indument. By comparing plate 1210, L. stonensis, and plate 
1208, L. densipila, which are reproduced at the same scale of 
magnification, the differences of indument between these species 
can readily be seen. Most of the siliques of L. stonensis examined 
possessed a perforate septum. Sometimes the perforation was 
small. However, in many instances it was larger, ranging 
upward in size to about one-fourth of the total area of the 
septum. 


6. Lesquerella perforata Rollins, Rhodora 54: 190. 1952. 


Annual; stems several to many, outer usually decumbent at base, 
inner erect, simple or branched, 1 dm. high, densely hirsute below with 
large spreading mostly simple trichomes, pubescent above with less- 
spreading mostly branched trichomes; basal leaves lyrately lobed, petiolate, 
2-5 em. long, 5-15 mm. wide, lobes variable, terminal lobe orbicular to 
ovate, entire or dentate, obtuse to more pointed, lateral lobes broadly 
oblong, entire or shallowly toothed, becoming remote toward petiole; 
hirsute on both surfaces, with mostly simple trichomes, marginal tri- 
chomes smaller and branched; cauline leaves sessile, aurieulate, broadly 
oblong to nearly ovate, sagittate, dentate, 8-20 mm. long, 4-8 mm. wide, 
hirsute above with simple trichomes, below with a mixture of simple 
and branched trichomes; pedicels straight, divaricately ascending, 
scarcely swollen at apex, 6-12 mm. long, uniformly pubescent with 
branched trichomes or with a mixture of simple and branched trichomes; 
sepals oblong, pubescent with a mixture of large and small branched 
trichomes, 3.5-5 mm. long, 1.5-2 mm. wide; petals white to pale lavender 
with a yellowish claw, sometimes tinged with light purple when dry, 
unguiculate, obovate to very broadly spatulate, emarginate to nearly 
entire, 7-9 mm. long, 5-6 mm. wide; filaments dilated at base, those 
of paired stamens 4—4.5 mm. long, anthers ca. 1.3-1.5 mm. long; glandular 
tissue subtending all filaments and nearly surrounding those of the 
single stamens, with projections between single and paired stamens; 
siliques inflated, variable in shape, broadly obovoid to subpyriform, very 
slightly stipitate, sparsely hirsute with large simple or forked trichomes 
to nearly glabrous on the exterior, densely pubescent with small den- 
dritically branched trichomes on the interior, 4-6 mm. long, widest 
above the middle, 4 mm. wide; septum nearly obsolete, represented by 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella 259 


only a narrow band of tissue around the inner margin of the replum; 
styles 1.5-.5 mm. long, unexpanded or only very slightly expanded at 
apex; ovules 2-6 in each loculus, funiculi nearly free, seeds slightly longer 
than broad to nearly orbicular, flattened, margined, 1.5-2.5 mm. long, 
1.2-2 mm. broad; cotyledons aceumbent. N = 8. Plate 1211. 


Tennessee, Wilson County: open field near Spring Creek, 5 mi. n. of 
Lebanon, 1952, R. C. & D. Rollins 5207 (an); same general locality, 1953, Rol- 
lins 5304 (GH); same locality, 1955, Rollins & Bold 55189 (an); near Lebanon, 
Sharp 83 (TENN); western edge of Lebanon, R. C. & D. Rollins 5208 (an); 
near Barton Creek, 4% mi. w. of Lebanon, Rollins 5306, 53145 (GH); 3 mi. w. 
of Lebanon, Rollins & Bold 55141 (an). 

Aside from the perforate septum and the very dense covering 
of dendritic trichomes on the interior of the valves which I 
have emphasized before, l.c., one of the striking features of this 
species is the papery quality of the siliques. As may be seen 
in plate 1211, the valves are somewhat wrinkled and unusually 
strongly veined. ‘These are not raised veins, but they do have 
greater density than the papery valve itself. The replum margin 
stands above the level of the adjacent valves which flare slightly 
outward on each side of it. "Thus, the edge of the replum to- 
gether with the valve margins forms a definite ridge that nearly 
encircles the silique. 

Mass collections of fruiting racemes have been available 
for study, and one of the very noticeable things about the plants 
of a population of L. perforata is the variation in the shape of 
the silique. In some plants, the siliques are definitely pyriform, 
in others nearly the shape of an inverted triangle, and in still 
others the siliques are much depressed and almost didymous. 
Also, there appears to be a difference in ovule number between 
two populations. In ten plants from the population near 
Spring Creek represented by No. 55139, there was an average 
of 3.1 ovules per locule. Ten plants of the population three 
miles west of Lebanon, No. 55141, had an average of 5.0 ovules 
per locule. The significance of these and other differences 
between populations is not understood at present, but I shall 
not be surprised if further study eventually shows that this and 
other species of Lesquerella of the Central Basin are in the 
process of fractionating, evolutionally speaking, to produce a 
new array of forms that may eventually become quite distinct. 

Lesquerella perforata is apparently local in its occurrence, 
but I suggest this with considerable caution. In three different 


260 Rhodora [Vor. 57 


years, I have searched the general area around Lebanon for 
populations of Lesquerella hoping to discover a broader range for 
L. perforata. On each occasion, the species was found in great 
abundance at the known stations, but these are all within six 
miles of Lebanon. Attempts to find it farther afield were not 
successful. However, my experience with Lesquerella elsewhere 
in the Central Basin of Tennessee leads me to be cautious about 
stating a definite range for a given species. At first, L. densipila 
was thought to be from a very limited area, but now, as a result 
of recent field work, it is known to be quite widespread in the 
Central Basin. 


7. Lesquerella auriculata (Engelm. & Gray) Watson, Proc. Am. Acad. 
23:250. 1888. 


Vesicaria auriculata Engelm. & Gray, Bost. Journ. Nat. Hist. 5: 240. 
1845. 

Alyssum auriculatum (Engelm. & Gray) Kuntze, Rev. Gen. Pl. 2: 931. 
1891. 


Annual; stems several to many, erect or decumbent at base, simple, 
0.5-2 dm. high, rather stout, hirsute with long simple trichomes and with 
an understory of smaller branched trichomes; basal leaves with a short 
petiole, lyrate to sinuate dentate, sometimes nearly entire, usually obtuse, 
rarely somewhat acute, 2-5 em. long, 8-15 mm. wide, hirsute on margins, 
midrib, and much of upper surface with simple trichomes, upper and 
lower surfaces often pubescent with smaller branched trichomes; cauline 
leaves auriculate, sessile, entire to dentate, usually overlapping on the 
stem, oblong to sagittate, 1-4 em. long, 3-10 mm. wide; pedicels divar- 
icately ascending to divergent at about 45°, nearly straight, hirsute with 
spreading simple and branched trichomes, 7-15 mm. long, not expanded 
at summit; sepals narrowly oblong, hirsute with spreading simple and 
branched trichomes, 4-6 mm. long, 1.5-2 mm. wide, outer pair slightly 
saccate, inner pair flat at base; petals yellow, obovate, entire to slightly 
emarginate at apex, 7-10 mm. long, 4-5 mm. wide; filaments abruptly 
dilated at base; siliques globose to longer than broad, glabrous, 4-6 
(-8) mm. long, 4-6 mm. wide, nearly sessile, valves glabrous within; 
septum entire or rarely with a small perforation; styles glabrous, 1.5-2 
mm. long; stigma expanded; ovules (4-) 6-8 (-10) in each locule; seeds 
nearly orbicular, flattened, margined, ca. 2 mm. in diameter; cotyledons 
accumbent. N = 8. 


Oklahoma: 2 mi. so, Choteau, Mayes Co., D. M. Moore 55-11 (au); 
6 mi. so. Ponco City, Kay Co., Waterfall & McCoy 11402 (OKLA, TEX); 1 mi. 
w. Perry, Noble Co., Harding 296 & 334 (okLA); 4 mi. e. Stillwater, Payne 
Co., Waterfall 9907 (OKLA, TEX); edge of Enid, Garfield Co., Gephardt 16 & 
160 (vs); near Kingfisher, Kingfisher Co., Stevens 188 (GH, MO, NY, OKLA, US); 


Rhodora Plate 1211 


PLATE 1211. Lesquerella perforata. Part of an infructescence, X5 (Rollins & Bold 55139). 
Photo by Frank White. 


Rhodora Plate 1212 


E DI M MM 
2 


ol "rta e 
oye we Fre mt 


Li 


T 


PLATE 1212. Lesquerella grandiflora. Part of an infructescence, X5 (Rollins 5561). 
Photo by Frank White, 


1955]  Rollins,—Auriculate-leaved Species of Lesquerella 261 


U. 8. Highway 66 e. of S. Canadian R. bridge, Canadian Co., Waterfall 3609 
(MO, OKLA); open plains, Grady Co., Goodman 2090 (GH, MO, NY); 3 mi. so. 
Union, Grady Co., Rollins 53126 (an); 8 mi. so. Watonga, Blaine Co., Goodman 
2382 (GH, MO, NY); near Hinton, Caddo Co., Demaree 11919 (GH); 3 mi. n. 
Lawton, Comanche Co., Rollins 53123 (au); 5 mi. n. Granite, Green Co., 
Bull 72 (mo). 


Texas: Big Sandy, Upshur Co., Reverchon 2967 (mo); Terrell, Kaufman Co., 
Reverchon 3717 (GH, MO, NY, US); s.w. Dawson, Navarro Co., Reverchon s.n. 
(M0); near San Felipe on the Brazos, Austin Co., March, 1844, E. Lindheimer 
217 (GH, type; GH, MO, isotypes); Brazos Prairie, Feb., 1846, Lindheimer s.n. 
(NY). 

The rarity and the curiously disrupted distribution of Les- 
querella auriculata in Texas contrast with the abundance and 
the more or less integrated distributional area of this species 
in Oklahoma. Adding to the puzzling known distribution is 
the fact that there are no recent collections from Texas. "Those 
of Reverchon go back to 1902 and 1903, and there are no more 
recently collected specimens in the herbaria from which I have 
received material. In 1953, I made a special search in Texas 
for L. auriculata especially in Kaufman and Navarro Counties 
and in the area around San Felipe, the type station. Near San 
Felipe, the related L. grandiflora was present in abundance, 
but I could not find L. auriculata. In fact, by the time the 
San Felipe area had been searched as thoroughly as time would 
permit, I was almost ready to conclude that L. auriculata had 
been based on an unusual specimen of L. grandiflora. However, 
I have studied the type and a series of isotypes (four sheets of 
specimens of the type series in all), and there is no doubt but 
that they belong to the same species as the material from farther 
north in Texas and that of Oklahoma. 

It is difficult to point up the differences between L. auriculata 
and L. grandiflora in such a way as to be convincing as to the 
specific nature of each. The smaller stature of L. auriculata 
is perhaps significant, but there are populations of L. grandi- 
flora with equally small plants. The basal leaves are often 
lyrate to nearly entire and broadly oblanceolate in L. auriculata; 
and if all populations possessed basal leaves of this type, it 
would be considered a distinctive feature. However, there 
are à number of collections that show evenly dentate and more 
or less acute basal leaves similar to those of L. grandiflora. 
Characters such as the nature of the pubescence, flower size, 


262 Rhodora [Vor. 57 


and the density of the infructescence appear to be the most 
constant for purposes of identification. In spite of the rela- 
tively few rather unsatisfactory morphological differences, there 
seems to be no doubt that two species, not a single one, are 
present. This view is reinforced by the evidence from chromo- 
some numbers. In two of the Oklahoma populations of 
L. auriculata (Rollins 53123 & 53126) where counts were made, 
n = 8 was found. On the other hand, n = 9 has been found 
in three populations of L. grandiflora. This aneuploid relation- 
ship between L. auriculata and L. grandiflora is almost certain 
to provide a genetic basis for breeding incompatibility between 
the two species. 

The different populations of L. auriculata are by no means 
identical with each other, and there is considerable variation 
within populations as well. In most plants of L. auriculata, 
the siliques are globose or nearly so, but there are occasional 
plants in which they are longer than broad, approaching the 
elliptical in outline. Such plants were found in a population 
three miles north of Lawton, Oklahoma (Rollins 53123), and 
I have seen specimens of this type collected by others. The 
ovule number is higher than usual in these more elongate fruits; 
for example, in the plants of my collection there were as many 
as ten ovules in each locule. The most frequent number is 
between six and eight. There is also a trend toward a reduced 
ovule number indicated by plants from the most northeasterly 
known station, Mayes County, Oklahoma (Moore 55-11). In 
this collection, I have examined fruits with only four and five 
ovules per locule. This particular collection is rather unusual 
in other respects, such as the deep cutting of the basal leaves 
and the rather markedly dentate cauline leaves. 


8. Lesquerella grandiflora (Hook.) Watson, Proc. Am. Acad. 23: 250. 
1888. 


Vesicaria grandiflora Hook., Bot. Mag. 63: pl. 3464. 1836. 

V. grandiflora var. pinnatifida Gray, Bost. Jour. Nat. Hist. 6: 146. 
1850. 

V. brevistyla Torr. & Gray, Fl. N. Am. 1: 102. 1838. 

Alyssum grandiflorum (Hook.) Kuntze, Rev. Gen. Pl. 2: 931. 1891. 


Annual; stems several to numerous, erect to decumbent at base, simple 
or branched, 2-7 dm. long, densely pubescent with branched usually 


1955] Rollins, —Auriculate-leaved Species of Lesquerella 263 


five-parted trichomes; basal leaves petiolate, oblanceolate, irregularly 
dentate to bipinnatifid, acute to obtuse, varying greatly in size from 
5-15 em. long, densely pubescent with branched, mostly five-parted 
trichomes, rays of trichomes often erect; cauline leaves oblong to lanceo- 
late, acute to obtuse, usually overlapping, dentate to conspicuously 
toothed, mid-rib conspicuous, densely pubescent with branched trichomes, 
1-4 cm. long, sessile, upper definitely auriculate, lower often narrowed 
toward base and usually lacking auricles; pedicels ascending to divaricate, 
straight to slightly curved upward, densely pubescent, 1-2 cm. long; 
sepals oblong, densely pubescent, nonsaccate, 5-7 mm. long, 2-3 mm. 
wide; petals yellow, broadly obovate to nearly orbicular in outline, rounded 
to slightly emarginate at apex, usually narrowed abruptly to a short 
claw, 8-11 mm. long, 6-9 mm. wide; anthers sagittate, 2.5-3 mm. long; 
filaments dilated at base, 3-4 mm. long; glandular tissue projecting, 
subtending paired stamens, nearly surrounding base of single stamens; 
siliques globose to slightly longer than broad, glabrous, 4-6 mm. high, 
4-5 mm. wide, with a very short gynophore to nearly sessile; valves 
glabrous within; styles 1-1.5 mm. long; stigmas expanded; funiculi 
slender, 1-1.5 mm. long, attached to septum by lower half; seeds margined, 
orbicular, flattened, 2.5-3 mm. in diameter; cotyledons accumbent. 
N =9. Plate 1212. 


Texas: Houston, Harris Co., Biltmore Herb. 14807 (mo); San Felipe, 
Austin Co., 1835 Drummond 20 bis (x, type; NY, type of Vesicaria brevistyla; 
GH, isotypes); 3 mi. s.e. of San Felipe, Austin Co., Rollins 5352 (au); Herring- 
ton, Brazos Co., E. J. Palmer 13440 (Mo, vs); 12 mi. s.w. Caldwell, Burleson 
Co., Cory 51660 (us); Columbus, Colorado Co., Howell 356 (us); Bastrop, 
Bastrop Co., Duval s.n. (us); Victoria, Victoria Co., Tracy 9193 (GH, MO, NY, 
us); prairies east of Victoria, Feb., 1845, Lindheimer 300 (an, type of Vesicaria 
grandiflora var. pinnatifida; wo, isotypes); Austin, Travis Co., Hall 23 (an, 
MO, NY, US); Caldwell Co., McBryde s.n. (Ny); 11 mi. n.w. of Westhoff, Gonzales 
Co., Rollins 5363 (an); 6 mi. s. of Lulling, Gonzales Co., Rollins 5556 (an); 
Ottine, Gonzales Co., Cory 5691 (GH); 4 mi. n. Cuero, Dewitt Co., Rollins 5561 
(GH); Goliad, Goliad Co., E. J. Palmer 9136 (Mo); 3 mi. s. Seguin, Guadalupe 
Co., Cory 54056 (OKLA); 14 mi. w. Stockdale, Wilson Co., Rollins 5364 (au); 
3 mi. w. of La Vernia, Wilson Co., Rollins 5365 (an); 4 mi. s.e. Floresville, 
Wilson Co., Rollins 5567 (aH); Riviera, Kleberg Co., Harrison s.n. (us); 
Kingsville, Kleberg Co., Sinclair s.n. (GH, Mo); Llano, Llano Co., Reverchon 
1488 (GH, MO, NY, US); 16 mi. s. of Llano, Rollins 5575 (an); Edge Falls, 
Rendall Co., H. B. Parks s.n. (au); 6 mi. w. Thelma, Bexar Co., Johnson & 
Webster 578 (GH, OKLA, US); near Moore, Frio Co., E. J. Palmer 33861 (an, 
MO, NY); Carrizo Springs, Dimmit Co., E. J. Palmer 33750 (NY). 


In stature and vegetative characters, Lesquerella grandiflora 
is quite variable. Individual plants appear to respond sharply 
to shading, moisture, and similar factors of the environment. 
At several locations, I have observed populations that extended 
from sunny to shady habitats. In the sun, the plants were 
short with comparatively small leaves and shortened internodes. 


264 Rhodora [Vor. 57 


Plants in the shade or growing in a dense growth of other herbs 
were tall with large thin leaves and elongated internodes. Her- 
barium specimens need to be interpreted with these points in 
mind. Vesicaria grandiflora var. pinnatifida of Gray appears 
to be a shade form, and the deeply cut leaves found on the type 
specimen are usual in shade grown plants. 

The populations of L. grandiflora that I have seen were all on 
sandy, loose, well-drained soils. These habitats are relatively 
dry and depend on current rains for a moisture supply. The 
most frequent occurrence of the species appears to be in the 
south central part of Texas just east of the Edwards Plateau. 
However, it 1s extremely abundant in the valley of the Llano 
River in Llano County which is well within the Plateau area. 
In general, L. grandiflora occurs south and west of L. auriculata, 
but the two species evidently come together at San Felipe in 
Austin County. There are no recent records of L. auriculata 
at the latter locality, but the species was founded on material 
from that location. 

The problem of distinguishing L. auriculata and L. grandiflora 
has been dealt with above. There is a sharp break between 
petiolate basal leaves and markedly auriculate cauline leaves 
in L. auriculata. However, in L. grandiflora, the transition is a 
gradual one with the lower cauline leaves being narrowed to the 
base and devoid of auricles in most instances. Proceeding up 
the stem, the leaves become less narrowed at base, and the 
auricles become more prominent. Another point of difference 
that should perhaps be stressed again is in the infructescence. 
In L. auriculata, the infructescence is dense and rather short 
while in L. grandiflora, the infructescence tends to elongate 
greatly and becomes rather loose as a result.—GRAY HERBARIUM 
OF HARVARD UNIVERSITY. 


LITERATURE CITED 


Lanjouw, J. AND F. A. STAFLEU. 1954. Index Herbariorum, part I. The 
Herbaria of the World, Ed. 2. Regn. Veg. 2: 1-179. 

Payson, E. B. 1922. A Monograph of the Genus Lesquerella. Ann. Mo. 
Bot. Gard. 8: 103-235. 

Rouurns, Reep C. 1952. Some Cruciferae of the Nashville Basin, Ten- 
nessee. RHODORA 54: 182-192. 

— — — ——. 1954. Interspecific Hybridization and Its Role in Plant Evo- 
lution. VIII Cong. Int. Bot. Rapp. & Comm. Sect. 10: 172-180. 

WarsON, SERENO. 1888. Contributions to American Botany. Proc. Am. 
Acad. 23: 249 287. 


1955] | Steyermark and Swink,— Plants New to Illinois 265 


PLANTS NEW TO ILLINOIS AND TO THE 
CHICAGO REGION 


JULIAN A. STEYERMARK AND FLOYD A. SWINK 


THE last report by the authors appeared in Ruopora 54: 
208-213. 1952. As on previous occasions, the authors have 
either collected or have found these new records in the her- 
barium of the Chicago Natural History Museum. All specimens 
colleeted have been deposited in the herbarium of that institution. 


1. ApDITIONS TO THE FLORA OF ILLINOIS 


TRILLIUM ERECTUM L. This species is not given by Jones in the second 
edition of his Flora of Illinois. During the spring of 1954 while botanizing 
a tract of rich woodland on the property of Mr. Richard Babcock in north- 
eastern Illinois, the senior author saw numerous plants belonging to this 
species growing with hundreds of Trillium grandiflorum. Mr. Babcock 
was certain that neither of the two species had been introduced, since he 
and his mother before him have maintained the property in its natural 
condition. Associated with the rich mesophytic woodland flora were 
Hepatica acutiloba, H. americana, Mitella diphylla, Asclepias exaltata, and 
many other native species. My field notes for the Trillium erectum col- 
lection read: “ovary deep purple, petals spreading from base, maroon- 
purple, peduncle ascending to spreading, above the leaves.” The data 
for this collection are as follows: rich wooded slopes along headwaters of 
tributary of Boone Creek, T 44 N, R 7 E, sect. 11, on property of Mr. 
Richard Babcock, 4 mi. east of Woodstock, Me Henry Co., May 9, 1954, 
Steyermark 76094. 

This locality is close to the Ringwood station, cited by Jones (p. 100) 
in the second edition of his Flora of Illinois for Trillium cernuum, in 
which he includes var. macranthum Wieg. The senior author has ex- 
amined the Ringwood specimen collected by Dr. Geo. Vasey, identified 
by him as T. cernwum and later verified by Stanley J. Smith, but it is not 
possible to judge in the dried condition of preservation two of the critical 
characters used in delimiting 7. cernuum, i. e., color of the ovary and 
position of the petals in anthesis, whether spreading from the base (as in 
T. erectum) or from the middle (as in T. cernuum). The senior author 
has determined this Vasey specimen as T. erectum, rather than as T. 
cernuum, on the basis of the field observations on plants from the Wood- 
stock locality, which is near the Ringwood station of Vasey's collection. 
The E. J. Hill collection from Wolf Lake, Chicago, cited under T. cernuum, 
has not been examined. It is probable, however, that both T. cernuwm 
and T. erectum will have to be admitted to the Illinois flora. 

RUMEX ACETOSELLA L., forma INTEGRIFOLIUS (Wallr.) G. Beck. The 
data for this collection are: in shaded ground of Col. Fabyan Forest 
Preserve south of Geneva, Kane Co., June 8, 1952, Swink 1198. 


266 Rhodora [Vor. 57 


FnaGARIA vesca L., forma ALBA (Ehrh.) Rydb. A colony of about 
fifty plants was found by Mrs. Charles W. Miller growing wild on her 
property in the Biltmore Estates subdivision of Lake County. In this 
colony the fruit was whitish with white superfieial achenes; the calyx 
lobes were spreading on all specimens noted. In general the plants are 
of a paler green color than in the common redfruited variety with which 
they were associated. "The data for this collection are as follows: upland 
oak-hickory open woodland on property of Chas. W. Miller, Kimberley 
Road, Biltmore Estates subdivision, 5 mi. north of Barrington, Lake 
Co., June 26, 1952, Steyermark 73627. 

Baprista MINOR Lehm. Mr. Karl Bartel, a keen local ornithologist and 
amateur botanist, discovered this species growing in a natural dry prairie 
associated with typical dry prairie species, such as Eryngium yuccifolium, 
Silphium laciniatum, and Silphium terebinthinaceum. As only one plant 
was noted and the station is well out of the known range (Mo. and Kans. 
to Tex.) of the species, it is suspected that the plant has been introduced 
at this locality. There is no indication at this time, however, to suggest 
how it could have been introduced from the southwest, as the nearest 
railroad is some distance away. The data for this collection are: in a 
dry prairie near 105th St. and Central Ave. near Chicago Ridge, Cook 
Co., August 2, 1953, Swink & Bartel 2521. 

Viana sinensis (L.) Endl. The authors found this Cowpea as an 
adventive in a state forest area. The data for the collection are as follows: 
sandy soil in Quercus stellata-Q. marilandica-Q. velutina forest, Mason 
State Forest, 5 mi. north of Forest City, Mason Co., August 13, 1949, 
Steyermark & Swink 68854. 

ViTIS RIPARIA Michx., var. svyRTICOLA (Fern. & Wieg.) Fern. Several 
collections which may be referred to this variety are the following: Jo 
Daviess Co.: 14 mi. south of Council Hill, Lansing 4145; Kankakee Co.: 
Altorf Island, 9 mi. northwest of Kankakee, Steyermark 68633; Kankakee, 
E. J. Hill; Du Page Co.: Naperville, Umbach; Cook Co.: Graceland, 
Chicago, F. C. Gates 252; Camp Harrison, between 154th and 159th streets, 
T 36 N, R 14 E, NE \ sect. 13, 114 mi. west of Calumet City, Steyermark 
68303; Winnetka, Sherff 1917; La Salle Co.: Starved Rock, Greenman, 
Lansing & Dixon 5. 


2. ADDITIONS TO THE FLORA OF THE CHICAGO REGION 


Bromus squarrosus L. This was collected in sandy soil along a path 
near the Baltimore & Ohio Railroad west of Lake Street in the Miller 
section of Gary, Lake Co., Indiana, June 21, 1952, Swink 1362. 

TRIODIA FLAVA (L.) Smyth, forma cuprea (Jacq.) Fosberg. An abun- 
dant grass known from several stations: along U. S. 12 near Lakeside, 
Berrien Co., Michigan, Swink 1904; sandy soil of a forest preserve parking 
lot on the south side of Ridge Road east of Thornton, Cook Co., Illinois, 
September 25, 1954, Swink 2762. 

ELYMUS INTERRUPTUS Buckl. This was collected in woods near South 
Lake of the Grand Marais lake chain near Stevensville, Berrien Co., 
Michigan, June 15, 1952, Swink 1299. 


1955] Steyermark and Swink,— Plants New to Illinois 267 


SPOROBOLUS VAGINIFLORUS (Torr. Wood, var. INAEQUALIS Fern. 
This collection was found in gravelly soil along Whitcomb St. near 17th 
St. in Gary, Lake Co., Indiana, September 21, 1952, Swink 1911. 

MUHLENBERGIA ASPERIFOLIA (Nees & Meyen) Parodi. Although 
Deam found this once in Lake Co., Indiana, it has not been reported 
previously for the Illinois portion of the Chicago region. The data for 
this record follow: along Douglas Park r.r. tracks property, just east of 
52nd Avenue, Cicero, Cook Co., Illinois, July 29, 1953, Swink 2356. 

PANICUM MILIACEUM L. This was collected along a roadside in the 
Lakewood Estates subdivision north of Dundee, Kane Co., Illinois, 
July 24, 1954, Swink 2751. 

LuzurA LUZULOIDES (Lam.) Dandy & Wilmott. This is collected in 
woods of Covenent Harbor near Lake Geneva, Walworth Co., Wisconsin, 
July 4, 1953, Swink 2272. 

ALLIUM STELLATUM Fraser. Found in a dry prairie near the ski slide 
in Fox River Grove, Mc Henry Co., Illinois, August 3, 1952, Swink 1636. 
This record applies only to the Illinois area, as it was found once in the 
dune area of Indiana (see Deam's Flora of Indiana, p. 1034, for details). 

NYMPHAEA ODORATA Ait. Collected in shallow water of the boggy 
area of South Lake of the Grand Marais lake chain near Stevensville, 
Berrien Co., Michigan, June 21, 1953, Swink 2207. 

JEFFERSONIA DIPHYLLA (L.) Pers. Collected in rich woods near 
Butternut Springs, 3 miles east of Wheeler, Porter Co., Indiana, April 
19, 1953, Swink 1949. This new record applies only to the Indiana area, 
as the plant has been collected in Cook Co., Illinois. 

PopoPHYLLUM PELTATUM L., forma APHYLLUM Plitt. This form was 
found in rich woods of Pilcher Arboretum east of Joliet, Will Co., Illinois, 
May 16, 1953, Swink 2016. 

SISYMBRIUM LOESELU L. This was collected in a barnyard north of 
Helm Road and east of Route 25, about 3 miles northeast of Carpenters- 
ville, Kane Co., Illinois, June 27, 1953, Swink 2221. 

FILIPENDULA RUBRA (Hill) Robins. This rare and local species, 
previously known in the Chicago region only from Du Page County, 
Illinois, has recently been discovered by Mr. Karl Bartel in Will and 
Cook counties, Illinois. The data for these collections are: marshy 
ground along a roadside south of an old scout camp three miles southeast 
of Custer Park, Will Co., Illinois, August 2, 1953, Swink & Bartel 2425; 
marshy ground along 107th Street east of Kean Avenue north of Palos 
Park, Cook Co., Illinois, August 2, 1953, Swink & Bartel 2505. 

Rusus Laciniatus Willd. Collected along the east side of the Des 
Plaines River in Schuth's Grove just north of Cermak Road near North 
Riverside, Cook Co., Illinois, December 6, 1953, Swink 2664. 

Cassia NICTITANS L. Collected in sandy soil south of Ridge Road, 114 
miles east of Thornton, Cook Co., Ilinois, August 14, 1954, Swink 2755. 
This record applies only to the Illinois area, as the plant has been found 
in the Indiana dunes. 

CROTALARIA SAGITTALIS L. Collected in sandy soil south of Ridge 


268 Rhodora [Vor. 57 


Road, 114 miles east of Thornton, Cook Co., Illinois, August 14, 1954, 
Swink 2763. 

DESMODIUM GLUTINOSUM (Muhl.) Wood, forma CcHANDONNETH (Lunell) 
Schub. Collected in woods near Long Lake near Black Hawk Beach, 
Porter Co., Indiana, July 14, 1952, Swink 1467. 

LESPEDEZA LEPTOSTACHYA Engelm. Collected in a dry gravelly prairie 
along the south side of Shoe Factory Road west of Sutton Road north 
of Bartlett, Cook Co., Illinois, Swink 2757. Jones (2nd edition, p. 179, 
Flora of Illinois) records this species from McHenry and Winnebago 
counties in northern Illinois. 

EUPHORBIA VERMICULATA Raf. Collected in sandy soil near Lake 
Michigan in Bridgman, Berrien Co., Michigan, September 20, 1952, 
Swink 1898. 

ViBURNUM OPULUS L. Collected in woods of Petrifying Springs Park 
east of Somers, Kenosha Co., Wisconsin, July 26, 1953, Swink 2289. 

ASTER FURCATUS Burgess. Jones (2nd edition, p. 273, Flora of Illinois) 
records this species only from Starved Rock, La Salle Co. Two collections 
from the Chieago region are known, in addition, as follows: upland slopes 
near creek, just east of Kimberley Road, Biltmore Estates subdivision, 
6 miles northeast of Barrington, Lake Co., Illinois, August 14, 1948, 
Steyermark 65964; wooded north-facing slopes along creek, same locality, 
September 16, 1947, Steyermark 65017. Other collections of this species 
in the Chicago Natural History Museum Herbarium from Illinois are: 
Winnebago Co. (E. W. Fell 531109); Peoria Co. (Brendel, Stewart); 
Tazewell Co. (Stewart); Ogle Co. (Patterson); Henderson Co. (Patterson); 
Woodford Co. (McDonald). 

CENTAUREA VOCHINENSIS Bernh. The authors found this species 
abundant locally in open grassy places along both sides of Butterfield 
Road (Route 55), near Lambert Road south of Wheaton, Du Page Co., 
Illinois, July 25, 1954, Steyermark & Swink 76353; Swink & Steyermark 
2758. 

HIERACIUM VuLGATUM Fries. Collected in woods of Covenant Harbor 
near Lake Geneva, Walworth Co., Wisconsin, July 4, 1953, Swink 2287.— 
CHICAGO NATURAL HISTORY MUSEUM AND COLLEGE OF PHARMACY, UNIVER- 
SITY OF ILLINOIS. 


Volume 57, no. 680, including pages 218—240, was issued 12 August, 1955. 


Dodova 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS | 

RALPH CARLETON BEAN Associate Editors 
RICHARD ALDEN HOWARD ( 

CARROLL EMORY WOOD, JR. 


Vol. 57 October, 1955 No. 682 

CONTENTS: 1 
The Vegetation of Sanibel Island, Lee County, Florida. 

George R. Cooley.. aee suas CREER MO as os e's Renan 269 
Non-validity of Nuttallian Names in Fraser's Catalogue. 

Loyd H. Shinnera’. oc. ccc oc ca oo We es er irre 290 
The Archer Method for Mounting Herbarium Specimens. 

Reed C. Rollins... 1. cage eae cody eeadhee ces 294 
Pinus rigida Miller in Quebec. Ernest JKOWIeON iis co LoT d 299 


The New England Botanical Club, Ine. 


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GRAY HERBARIUM of Harvard University, 
Cambridge 38, Mass., U. S. A. 


Rhodor: Plate 1213 


Prare 1213. Aerial photograph of east-central portion of Sanibel Island, showing vegeta- 
tional features. 


Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 57 October, 1955 No. 682 


THE VEGETATION OF SANIBEL ISLAND 
LEE COUNTY, FLORIDA 


iEORGE R. COOLEY 


INTRODUCTION 


SANIBEL ISLAND lies in the Gulf of Mexico, a few miles off 
the southwestern coast of peninsular Florida near the mouth 
of the Caloosahatchee River about fifteen miles from the city 
of Fort Myers. Lying as it does, less than three degrees north 
of the Tropic of Cancer, Sanibel Island has many characteristics 
of tropical and sub-tropical insular areas. Furthermore, being 
only a few miles from the mainland, the island has the appearance 
of many sections of South Florida. Yet it is not exactly the 
same as any region of the tropies or of Florida, historically, 
geographically, geologically, or botanically. Occupied by white 
people for little more than seventy years, influenced by rivers, 
ocean currents, and mighty storms; made up almost entirely 
of shells and shell material;and the home of plants of the tropical, 
sub-tropical, and temperate zones, it is unlike, in many respects, 
any area to be found in the southern United States. 


History OF SANIBEL 


It is said that on his trip to the Gulf of Mexico and Mississippi 
River in 1539, De Soto stopped at Sanibel Island. Captain 
Bernard Romans, ‘‘Draugh™ Mathem* Navigat"" and ‘Nat- 
uralist & Botanist" in 1774, when describing the region around 
Charlotte Harbour north of Sanibel, said, “This nook in the 
land forms what the Spaniards call Ensenada de Carlos, i.e. 
Charle’s Bay: the piece of coast that trends E. and W. is the 
beach of an island called Sanybel, This place is further remarkable 


270 Rhodora (Vor. 57 


for a great number of pine-trees without tops, standing at the 
bottom of the Bay, like which there is no spot in the whole 
extent of this coast. 'The northernmost entrance is likewise 
remarkable for a singular hummock, or grove of pine-trees, 
standing very near the beach, and the only one of its form and 
kind in all these parts." No other records seem to be extant 
which would help us in an understanding of the flora of this 
island before its discovery by the white man four hundred years 
ago, or for that matter at any time up to the present. 

At the time of De Soto's exploration and for three hundred 
years thereafter a tribe of Muspah Indians apparently made 
their homes on Sanibel Island and nearby Captiva Island. 
Great kitchen middens, mostly shells, give evidence of long 
tenure by these Indians. Mr. Julien C. Yonge of the Library 
of Florida History found in early records that in 18831 a New 
York company set up a colonial development at the eastern 
end of the island; there were also comments of a traveler to 
the effect that two years thereafter no evidence of this develop- 
ment remained except the framework of the largest house. 
About 1883 an agricultural and horticultural development 
started which was to change the face of the island for a period 
of over forty years. All the arable land on the island was 
cleared and cultivated (Map C). Citrus fruits and garden vege- 
tables, principally tomatoes, eggplants, and squashes, were raised 
in considerable quantities. Boats came from the terminus of the 
railroad on the mainland a score of miles to the north to collect 
the produce. It is reported that one gardener in 1901 received 
$1,100 for the tomatoes grown on one acre of ground. In 
time, competition from the mainland lessened the prosperity 
of Sanibel farmers so that when a disastrous hurricane struck 
the island in 1926, half the population left. Those who remained 
have served winter visitors, fishermen, and shell collectors, 
for Sanibel has become known as one of the three great shell 
collecting beaches of the world. The replacement vegetation 
of the previously cultivated land is now again steadily progressing 
toward a climax. 

GEOGRAPHY 


All of the islands off the west coast of Florida lie north and 
south like reefs along the shoreline of the peninsula except 


1955] | Cooley,— The Vegetation of Sanibel Island 271 


\@tHARLOTTE HA 


SANIBEL 
ISLAND 


ESTERO 
ISLAND 


Mar A. The Charlotte Harbor area of southwestern Florida, showing the location 
of Sanibel Island 


272 Rhodora [Vor. 57 


Sanibel which lies east and west, at right angles to the others, 
taking the full brunt of storms from the south and being sub- 
jected to the constant flow of a northward moving gulf stream. 
The island is about 10.75 statute miles long and 2.6 statute 
miles across at its widest point; it lies at 26° 30’ N. latitude and 
82° 10° W. longitude (Map A.) 

Nowhere is the land more than fourteen feet above the level 
of the Gulf of Mexico. The elevation of most marsh areas 
is six feet or less; the grasslands, six to eight feet, and the ham- 
mocks and former cultivated areas from six to fourteen feet 
above sea level. 

From the aerial photograph (Plate 1214, fig. 2) one can obtain 
a general idea of the geographical features and vegetation of the 
island. At the rounded point to the east is a 70-foot lighthouse; 
west of it can be seen a road bordered by casuarina trees. At 
this point there is the ferry landing, a restaurant, the post office 
and the home of the postmistress. On both sides of this road, 
which crosses the narrow end of the island, can be seen cleared 
areas representing an incipient real estate development. The 
dark space toward the lighthouse is a rather dense button man- 
grove thicket; the black spots represent red mangroves. The 
rest of the dense growth consists of button mangrove (Con- 
ocarpus erecta) and its associates, with a cactus thicket at the 
side toward the lighthouse. In the photograph the dark mass 
in the upper left indicates the presence of red mangrove swamps, 
which form matted growths along most of the inner shore on 
Pine Island Sound. The lighter-colored sweeps of gray are 
button mangrove swamps where airplants grow in abundance. 
A road extends east and west through the middle of the island. 
This is now a well-maintained macadam highway connecting 
by a concrete bridge at the west end of the island with neigh- 
boring Captiva. Along this road can again be seen rows of 
casuarinas planted about 1912 to 1914. South of the main road 
the grasslands are extensive, changing into Spartina marshes 
where the land is lower between the ancient shell ridges. Over 
the part of the island not shown the Spartina marshes are very 
much larger, being fully one-half mile in breadth and far-reach- 
ing in extent. A natural drainage area is shown by the ir- 
regular dark line through the center of the grasslands. The 


1955] Cooley,— The Vegetation of Sanibel Island 273 


straight white line indicates the road leading to the island’s 
only general store and a community of winter dwellings. From 
this road toward the west, on the north side of the main highway, 
one can see formerly cultivated fields. A large part of the land 
on both sides of the main road from this section across the entire 
island is where the farming of the 1883-1928 period was done. 
The design at the lower left shows a winter resort. There are 


SAN/BEL /SLAND 


PLANT HABITATS 


L7 7] Grassland 
and prairie 
Spartina marsh 
um = í palmetto 
Salt flat 

à Mangrove 
(ang swamp 
Palmetto jungle and mized woods 


V Cactus thicket 


Mar B. The principal habitats and vegetational groupings on Sanibel Island. 


four such regions on the Gulf side of the island. Plate 1213 is 
an aerial photograph of the east central portion of the island. 
It shows vegetational features similar to those described above. 


ECOLOGICAL FEATURES 


The beach on the Gulf side ends abruptly at a ridge of shells 
washed up by a previous great storm. The island, other than 
the mangrove swamps and inner salt marshes, is made up of 
ridges more or less parallel to the Gulf shore. In ages past these 
ridges were formed in the same way as the present outer ridge 
which was created by the storm of October, 1953. Great quanti- 
ties of shells were washed up onto the beach and over the beach- 
strand vegetation, burying all but the tallest of the shrubs and 
trees (Plate 1214, Fig. 1). Four and five feet of shells covered 
the strand for varying widths of fifty to one hundred-fifty feet 
forming a new ridge. Back of the new ridge is a marsh of 
Spartina and its associates growing up through broken shells. 
Successive ridges and low places produce prairie growths and 


274 Rhodora [Vor. 57 


marsh vegetation, then transition vegetation and hammocks 
as the distance lengthens from the beach (Map B). 

The inner north shore facing the quiet waters of Pine Island 
Sound is free from the effeets of waves and storms and the 
vegetation consists of a great mangrove swamp. Back of the 
mangroves are tidal marshes, back of these on slightly higher 
ground are button mangroves bordering on hammocks and 
mixed woods as the elevation increases. 


SANIBEL /SLAND 


AREAS FORMERLY UNDER 
CULTIVATION 


Mar C. Former cultivated areas of Sanibel Island. 


Among the red mangroves organic muck has formed to the 
depth of several feet; in the Spartina marshes a foot or more of 
humus has accumulated; but all other soils are made up of de- 
composing shell material and varying amounts of decaying 
vegetable matter. 

Evidence of human occupation and disturbance in the form 
of ditehes old and new, abandoned homesites, fence-rows, and 
former groves, give character to much of the higher ground. 
(See Map C showing formerly cultivated land, as defined by an 
old resident.) 

Research discloses meager records of the flora of Sanibel 
and few specimens indeed are in herbaria of America. Botanists 
have neglected this interesting place. At the suggestion of 
Dr. Richard A. Howard the writer was prompted to undertake 
a floristic study of Sanibel and to that end made a winter's stay 
on the island and two subsequent visits to it. 


1955] Cooley,— The Vegetation of Sanibel Island 275 


A. S. Hitchcock visited Sanibel in 1900, and three specimens 
collected by him at that time have now been located at the 
Gray Herbarium. He was probably the instigator of S. M. 
Tracy’s trip to Sanibel in 1901, when the new species Eragrostis 
tracyi Hitchcock was collected, but no notes and only few speci- 
mens of this trip can now be found. George A. Orric collected 
on neighboring Captiva Island in 1915, but it is not known 
that he made any study of Sanibel. On January 24, 1924 
Roland M. Harper spent a few hours on the island and, as stated 
in a letter to the author, “took what notes I could on the vegeta- 
tion, and a few pictures, but do not think I collected any plants." 
James B. McFarlin did some collecting on Sanibel November 
4, 1934, incidental to his study of the flora of Polk County, 
central Florida. In 1936, J. R. Swallen collected five grasses; 
and in 1951, R. Bruce Ledin spent two days on Sanibel collecting 
52 numbers for the Buswell Herbarium at the University of 
Miami. Ledin collected plants of Sanibel that were typical - 
of the Florida Keys of Monroe County. He was amazed at 
the number of identical species found in both places. About 
35 of 52 species collected were considered to be essentially West 
Indian plants. Specimens in the herbarium of the University 
of Florida (from Sanibel) are probably less than a dozen, accord- 
ing to Miss Lilian E. Arnold, Curator. John H. Davis and the 
writer spent a day on Sanibel in 1951, but did little collecting. 

Of all these visits few comments remain for our understanding 
of the flora of Sanibel. In 1927 in “Natural Resources of South 
Florida" Harper said of the West Coast islands, “the vegetation 
comprises that of beaches and dunes, cactus thickets, salt 
flats, palm savannas, mangrove swamps, and a little tropical 
hammock. There is a little truck-farming on some of the islands, 
such as Sanibel.” 

This study purports to provide a general understanding of 
the ecology of the Island and its present vegetation to the end 
that comparisons in the future may be accurately made. 


VEGETATIONAL FEATURES 


Even the casual visitor to Sanibel is impressed by the number 
and ubiquity of the cabbage palm (Sabal palmetto). Throughout 
the island it is found in almost all habitats, giving its name to 


216 Rhodora [Vor. 57 


some, as cabbage palm prairie (Plate 1215, Fig. 3) to the grass- 
lands, and cabbage palm jungle to the hammocks. <A quarter 
of a mile from the ferry landing adjoining the oldest settlement 
area on the island 1s a stand of Sabal palmetto reminding one of a 
mature reforestation plantation so tall, so straight, and so 
dominant are the trees of this species. 

Clinging to the cabbage palms and often growing rampant 
over them are Rhus radicans and less frequently Parthenocissus 
quinquefolia. Living on them epiphytically are Ficus aurea, 
Vittaria lineata, and Phlebodium aureum and that strange crypto- 
gam of the sub-tropics, Psilotum nudum, with slender, branching, 
delicate stems is sometimes found at the base of the cabbage 
palms. 

Another tree which is characteristic of the modern Sanibel 
landscape is the Australian pine (Casuarina equisetifolia) which 
lines many of the roads and beaches. Brought to the island 
about forty-five years ago, Casuarina has become well established. 
By the distribution of vast numbers of seeds it has extended 
its territory far beyond the roadsides and beach strands where 
it was originally planted. At one place in the center of the 
island it has multiplied to the point of forming a wood where 
it tends to become dominant. On Wulfert Road is a row of 
Casuarina cunninghamiana about 30 to 40 years old. This 
species 1s apparently not becoming naturalized. 

From the main black-top road which extends through the 
center of the island for its entire length, mangroves are noticeable 
at only one point. However, a boat ride along the inner shore 
for almost the entire length of the island reveals a mangrove 
swamp of eight square miles. Dominant at the water’s edge 
is the red mangrove (Rhizophora mangle), its interlacing and 
arching trunks, branches, and roots forming a maze through 
which progress can be made only with persistence and a sharp 
machete. Wherever the soil has been built up above the mean 
tide level, black mangroves (Avicennia nitida) appear with trunk 
diameters up to 4” at breast height. Deeper in the swamp 
(hence rarely seen from a passing boat) and only where the soil 
is generally above the tide are the white mangroves (Laguncu- 
laria racemosa), attaining a size no greater than large shrubs. 

Between the red mangrove swamps and the hammocks and 
cabbage palm savannas toward the middle of the island are 


1955] Cooley,—The Vegetation of Sanibel Island 271 


interesting belts of button mangroves (Conocarpus erecta, Plate 
1215, Fig. 4). Here flourish air plants in some profusion; not 
much Spanish moss (Tillandsia usneoides) but an abundance of 
Epidendrum tampense, Tillandsia fasciculata, T. balbisiana, T. 
utriculata, and T. tenuifolia. At four places cactus thickets 
have been formed among the button mangroves; wicked places 
for the botanist, but happy homes for gopher snakes which 
grow to lengths of seven and eight feet. 

As the button mangroves become numerous, extending toward 
the center of the island (Plate 1216, Fig. 6), they are joined by 
an interesting transition group of a score or more of hammock 
trees and shrubs, including the following: 


Ardisia escallonioides Randia aculeata 
Baccharis glomeruliflora Phytolacca rigida 
Chiococca alba Rivina humilis 

Dodonaea jamaicensis Zanthoxylum fagara 
Eugenia anthera Capparis cynophallophora 
Eugenia myrtoides Rapanea guianensis 
Eugenia azillaris Sideroxylon foetidissimum 
Forestiera porulosa Bumelia angustifolia 
Jacquinia keyensis Psychotria nervosa 


Pithecellobium unguis-cati 


On the larger of these trees, Spanish moss grows more abundantly 
than other air plants. Cabbage palms are present, acting as 
hosts to the shoestring fern (Vittaria lineata), the golden polypody 
(Phlebodium aureum), and young growths of the strangling fig 
(Ficus aurea) (Plate 1216, Fig. 5). They also support a vigorous 
growth of poison ivy (Rhus radicans) and Virginia creeper 
(Parthenocissus quinquefolia). 

The writer should mention here Sanibel’s second worst pest 
(mosquitoes are first), poisonivy. Apparently it will not tolerate 
salt water and is therefore not found in the mangrove swamps 
or the salt marshes. The beaches and the black-top road do 
not provide happy homes for Rhus radicans either, but every 
other habitat is congenial to it. On a cabbage palm was found 
a poison ivy plant with a stem 9 cm. in diameter at breast 
height. One branch 13 dm. long bore 6,130 blossoms. Often 
leaves are found 10 cm. wide and 20 cm. long. On the cabbage 
palms some poison ivy plants grow 10 meters high. In a cabbage 
palm thicket six hundred seventy-two poison ivy plants or sprouts 
were counted in a quadrat twenty by twenty feet. 


278 Rhodora [Vor. 55 


In the buttonwood swamps and the hammocks and near Indian 
shell mounds has grown the tree cotton (Gossypium hirsutum). 
For a long time, despite the fact that commercial cotton is not 
raised within two or three hundred miles of Sanibel, the De- 
partment of Agriculture has sent four men to the island several 
times a year to rid it of wild cotton and thus deprive the boll 
weevil of food and a breeding place. "The species has persisted 
but is now almost extinct on Sanibel as only three young plants 
were found in à two-day search in February (1954). 

Hammocks on Sanibel vary greatly. As indicated above, 
one hammock is a stand of pure Sabal palmetto; many are Sabal 
palmetto jungles where trees, shrubs, vines and herbs compete; 
some in the center of the island are distinguished by Quercus 
virginiana; and some are mixed forests. The gumbo limbo 
tree (Bursera simaruba), although frequent and widespread 
through the higher elevations of the island, is nowhere abundant. 
Because of its red bark it is often mistakenly called mahogany 
by natives and visitors alike. In a mixed wood on Wulfert 
Road stands a slash pine (Pinus elliottii var. densa) with several 
younger trees near by. Millions of slash pines grow on the 
mainland and yet Sanibel can boast of but one small colony. 

Several square miles of Spartina marshes lie between the 
principal hammocks and the coastal shell ridge. Here the 
dominant is Spartina bakeri, accompanied by Cladium jamai- 
cense, Andropogon glomeratus, Bacopa monnieri, and Sesuvium 
portulacastrum (Plate 1217, Fig. 7). In the marshes the presence 
of ancient shell ridges is evidenced by plants which prefer slightly 
drier ground. Luxuriant growths appear of Baccharis halimi- 
folia, Sporobolus domingensis, Aristida patula, Fimbristylis 
castanea, Flaveria linearis, Kosteletzkya virginica, Mikania batati- 
folia, Rhus radicans, Melothria pendula, Melanthera deltoides, 
and on still higher ground Sabal palmetto and its prairie associates 
(Plate 1217, Fig. 8). 

The grass lands between the coastal shell ridge and the Spar- 
tina marshes are largely dry open areas in which the following 
species grow: 


Bouteloua hirsuta Piriqueta caroliniana 
Sporobolus virginicus Pluchea purpurascens 
Aristida patula Rhynchosia michauzxii 


Chloris petraea Samolus ebracteatus 


1955] Cooley,—The Vegetation of Sanibel Island 279 


Bidens pilosa Sida carpinifolia 
Cirsium horridulum Waltheria americana 
Coreopsis leavenworthit Conyza canadensis 
Flaveria floridana Vigna repens 
Flaveria linearis Lochnera rosea 
Gaura angustifolia Crotalaria striata 
Opuntia austrina Cassytha filiformis 
Physalis elliottii Cenchrus pauciflorus 


The shrubby areas also present are composed of: 


Forestiera porulosa Lantana involucrata 
Baccharis halimifolia Lantana ovatifolia 

M yrica cerifera Sophora tomentosa 
Ernodea littoralis Trichostema suffrutescens 
Eugenia axillaris Capraria biflora 


Jacquinia keyensis 


Charles Torrey Simpson said “To the naturalist . . . the 
seashore is the most fascinating place in the world." On 
Sanibel’s seashore the coastal shell ridge—a pile of countless 
shells—influences the botanist by some powerful and irresistible 
charm. To be sure, the hand of man has contributed to the 
flora of the coastal ridge by his plantings of Casuarina equiseti- 
folia to bind the soil and break the wind, but the hand of nature 
has brought to the beach strand other plants entrancing in 
their characteristics and different in their forms and habits. 

Coccoloba uvifera, the sea grape, changes in appearance many 
times a year. Its shiny, bright new leaves in March replace 
the dull *platter" leaves which fall to cover the earth beneath; 
then 15 cm. spikes of greenish yellow flowers extending from the 
axils of pairs of near-opposite leaves attract the bees and insects; 
clusters of large fruit of deepening color tempt the local house- 
wives to make sea-grape jelly; then the aging leaves grow dull 
again and await the rebirth of another spring. 

A pan-tropical half-shrub, Scaevola plumieri (Fig. 9) tolerates 
the salt spray and saline soil of the upper beach. Its succulent 
leaves retain a full green color throughout the year; its springtime 
white flowers have five white involute petals arranged palmately 
in a plane. 

The active waters of the beach give Rhizophora mangle no 
chance for a footing, and the dry character of the ridge excludes 
Avicennia nitida and Laguncularia racemosa. Conocarpus erecta, 


1 "In lower Florida wilds,” page 276. New York. 1920. 


280 Rhodora [Vor. 57 


the button mangrove, has more success along the coastal shell 
ridge, generally near the marshes or in man-made ditches (Plate 
1218, Fig. 10). "The cinereous form of Conocarpus erecta grows at 
four stations, none of them near the coast. On the neighboring 
island of Captiva Ambrosia hispida forms runners exceeding 
fifty feet, but on Sanibel the plant is rare and small. Another 
soil-binder, however, is by no means so rare; Ipomoea pes-caprae, 
the railroad vine, climbs over both the ground and the vegeta- 
tion. 


Fig. 9. Flower of Scaevola plumieri, X 4. 


Most abundant of the shrubby growths of the coastal shell 
ridge is Suriana maritima, the bay cedar, in the West Indies 
often called tassel plant, with small yellow flowers and alternate 
entire leaves. With it are scattered plants of Uniola paniculata 
with culms two meters in height. These sea oats serve to 
bind the coastal soil but are never dominant on Sanibel. 

I collected what appeared to be two species of Oenothera occur 
on the beach, one with yellow flowers and one with reddish 
flowers. Both are Oenothera humifusa, whose flowers are yellow 
when new blown and reddish when older, the change taking place 
even from morning to afternoon. 

The creeping plants of the marshes, Sesuvium, Portulaca, and 
the salt-tolerant grasses of the wet areas are not found on the 
coastal ridge, for it is a dry habitat where rain filters quickly 


1955] Cooley,— The Vegetation of Sanibel Island 281 


through the crushed shell aggregate. It is to be expected that 
Calonyction should clamber over the shrubs of the coastal shell 
ridge, but Calonyction tuba grows only on the low ridge at the 
inner side of the island where storms are not severe and Calonyc- 
tion aculeata lives mostly one hundred fifty yards and more back 
from the gulf beach. 

Common on the coastal ridge is Baccharis halimifolia and less 
frequent is Baccharis dioica. The ordinarily entire leaves of 
the latter are here sometimes indented, possibly from hybridiza- 
tion. It has the distinctive habit of extending its tap root 
horizontally just beneath the ground at right angles to its stem. 

From one end of the island to the other on twelve miles of outer 
shell ridge many native, adventive, and waif species make their 
homes. Solidago sempervirens var. mexicana is present but not 
abundant. Bidens pilosa, with its bright yellow-centered flower 
heads, is much more abundant, and its barbed achenes, known 
as Spanish needles, cling tenaciously to everyone who brushes 
past the plant. Mention should be made of the chaff flower 
(Achyranthes ramosissima). This tropical herb of the amaranth 
family has a spreading manner of growth among the herbs and 
low shrubs. Cakile (Plate 1221, Fig. 17) is widespread through 
the sub-tropics, but only one species of it (Cakile fusiformis) 
is found on Sanibel and at only one spot on disturbed ground 
near a real estate development. 

There is only one extensive broad beach area (Plate 1219, 
Fig. 13), and that toward the western end of the island. This 
beach is up to two hundred yards wide, and is composed of 
both quartz sand and shell fragments, but unlike typical sandy 
beaches drifts little. Such vegetation as manages to exist in 
the broiling sun and salt air includes Sesuvium portulacastrum 
(Plate 1219, Fig. 12), Suriana maritima (Plate 1220, Fig. 14), 
Scaevola plumieri, Euphorbia buxifolia, Oenothera humifusa, Iva 
imbricata, Atriplex arenaria, Uniola paniculata, Sporobolus virgini- 
cus, Spartina patens, and Ipomoea pes-caprae. The rolling beach 
grades downward into marshes with a varied border of Dalbergia 
ecastophyllum, Coccoloba wvifera, Suriana maritima, Conocarpus 
erecta, Casuarina equisetifolia, and other shrubs. 

The white panicles of Yucca alozfolia do not last long but for 
a short time offer a spectacle to those ‘“‘shellers” whose eyes are 


282 Rhodora [Vor. 57 


raised from the beach. No other Yucca grows on Sanibel. 
Chloris petraea, Lochnera rosea, Rhus radicans, Cenchrus pauci- 
florus, Sporobolus virginicus, Euphorbia spp., Heliotropium parvi- 
florum, H. polyphyllum, H. curassavicum, Conyza canadensis, and 
Asclepias verticillata are a few of the vascular plants crowding 
the slopes and crest of the coastal shell ridge. 

Eragrostis tracyi was described as a new species from a col- 
lection made on Sanibel in May, 1901. It was not collected 
again on Sanibel until April 13, 1954, but has never been found 
elsewhere. It is common on disturbed ground, roadsides, 
lawns, around homes, and in cleared areas. Diligent search 
over grasslands and savannas produced no plants. The habitat 
it occupies leads one to believe that Eragrostis tracyi has been 
introduced onto Sanibel. But from where did it come? 


CHECK LIST 


Nomenclature provides a problem for the taxonomists of the 
Southern States. No manual exists to which a botanist can 
go for names of southern plants in keeping with recent taxonomic 
studies and judgment, and in conformity with the International 
Code of Nomenclature. The author searched the monographs 
and writings of the last twenty years and has used the names 
which in large part conform to the best present-day usage. 

Mr. Jason R. Swallen, of the United States National Museum, 
identified the grasses collected, and Professor Erdman West and 
Miss Lillian E. Arnold of the University of Florida helped with 
the determination of all other specimens. Specimens have been 
deposited in the United States National Museum, the Gray 
Herbarium, the herbarium of the New York Botanical Garden, 
the herbarium of the University of Florida, and the Cooley 
Herbarium of Rensselaerville, New York. 

In the check list which follows seven habitat columns are 
used after Lemon (ms). The column headed Mangrove Swamps 
for convenience includes red mangrove swamp plants; Spartina 
Marshes include salt marsh plants; and Grasslands and Savannas 
combine both wet and dry areas. Frequent ancient shell 
ridges throughout the island increase the elevation slightly at 
many points, providing congenial habitats for plants of dry 
preference in the midst of damp and wet areas. No effort has 


1955] Cooley,—The Vegetation of Sanibel Island 283 


been made to define these habitats precisely; rather general 
ecological names have been used. 


DEFINITIONS 


Dominant (D) Prevailing, tending to drive out all other plants and to 
resist invasion by them. 

Abundant (A) Teeming, in copious supply. 

Common (C) General, not local, so frequent as to be found easily. 

Frequent (F) Often to be met with, but not general. 

Infrequent (I) Isolated, sparse, occurring at considerable distances. 

Rare (R) Seldom met with, so infrequent as to be unexpected. 


fields 
Coastal 
shell ridge 
marshes 
Grasslands 
& savannas 
Mangrove 
swamps 
Palmetto 
jungles 


Old 
= Spartina 


Chara zeylanica Klein in Willd. 
Psilotum nudum L. 
Acrostichum danaeaefolium Langsd. & Fisch. 
Blechnum serrulatum Rich. F 
Nephrolepis exaltata (L.) Schott I 
Phlebodium aureum L. 
Polypodium polypodioides (L.) Watt 
Pteridium aquilinum (L.) Kuhn, var. caudatum 
(L.) Sadebeck I 
Pteridium aquilinum (L ) Kuhn, var. pseudo- 
caudatum (Clute) Heller I 
Vittaria lineata Sw. F F 
Typha angustifolia L. 
Typha domingensis Pers. 
Pinus elliottii Engelm., var. densa Little & Dorm. I 
Najas guadalupensis (Spreng.) Morong 
Andropogon glomeratus (Walt.) BSP. F 
Andropogon virginicus L. F 
Aristida patula Chapm. ex Nash 
Aristida purpurascens Poir. 
Bouteloua hirsuta Lag. 
Cenchrus echinatus L. F 
Cenchrus incertus M. A. Curtis C 
Cenchrus pauciflorus Benth. C 
Chloris petraea Swartz C 
Cladium jamaicense Crantz 
Cynodon dactylon (L.) Pers. I 
Dactyloctenium aegyptium (L.) Beauv. 
Digitaria sanguinalis (L.) Scop. 
Distichlis spicata (L.) Greene 
Eleusine indica (L.) Gaertn. 
Eragrostis ciliaris (L.) R. Br. I 
Eragrostis elliottii S. Wats. 
Eragrostis pilosa (L.) Beauv. 
Eragrostis tracyi Hitchc. 
Leptochloa dubia (H. B. K.) Nees. 
Muhlenbergia capillaris (Lam.) Trin. 
Panicum adspersum Trin. 


h 
> 
- 
[anl 


Q7 


M M ge 


aami 
aa 
Qu “pry u 


Q gH 


gy ORE 


284 Rhodor: 


Panicum agrostoides Spreng. 

Panicum albomarginatum Nash 
Panicum amarulum Hitche. and Chase 
Panicum bartowense Scribn. & Merr. 
Panicum dichotomiflorum Michx. 
Panicum neuranthum Griseb. 

Panicum virgatum L. 

Paspalum ciliatifolium Michx. 
Paspalum vaginatum Swartz 
Pennisetum glaucum (L.) R. Br. 
Rhynchelytrum roseum (Nees) 8. & H. 
Setaria geniculata (Lam.) Beauv. 
Setaria glauca (L.) Beauv. 


Setaria macrosperma (Scribn. & Merr.) Schum. 


Sorgum halepense (L.) Pers. 

Spartina bakeri Merr. 

Spartina patens (Ait.) Muhl. 

Sporobolus domingensis (Trin.) Kunth 

Sporobolus juncea (Michx.) Kunth 

Sporobolus virginicus (L.) Kunth 

Triplasis purpurea (Walt.) Chapm. 

Uniola paniculata L. 

Cyperus globulosus Aubl. 

Cyperus ligularis L. 

Cyperus martindalei Britton 

Cyperus planifolius L. C. Rich. 

Cyperus polystachyos Rottb., var. texensis 
(Torr.) Fern. 

Cyperus pseudovegetus L. 

Cyperus rotundus L. 

Cyperus strigosus L. 

Dichromena colorata (L.) Hitchc. 

Eleocharis atropurpurea (Retz.) Kunth 

Eleocharis geniculata (L.) R. & 8. 

Fimbristylis castanea (Michx.) Vahl 

Scirpus americanus Pers. 

Scirpus cyperinus (L.) Kunth, var. eriophorum 
Michx. 

Cocos nucifera L. 

Sabal palmetto (Walt.) Torr. 

Serenoa repens (Bartr.) Small 

Lachnocaulon anceps (Walt.) Morong 

Tillandsia balbisiana Schult. 

Tillandsia circinata Schlech. 

Tillandsia fasciculata Sw. 

Tillandsia recurvata L. 

Tillandsia tenuifolia L. 

Tillandsia usneoides L, 

Tillandsia utriculata L. 

Commelina erecta L. 

Juncus megacephalus M. A. Curtis 

Sansevieria guineensis Willd. 

Smilax auriculata Walt. 


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a) 
Se 
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Sa OF 
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C 
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I 
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I 
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C 
€ Y 
F I 
C F 
I 
F 
Escape 


Spartina 
marshes 


F 


Grasslands 
& savannas 


Mangrove 
swamps 


amamama 


[Vor. 57 
© 
E 
Zé T2 
EEEE 
AB AE 


F 


F 


Rhodor:z Plate 1214 


PLATE 1214. Fic. 1, Recently formed shell ridge. Fie, 2, Aerial photograph 
showing ancient shell ridges paralleling the south coast. 


5 


121 


Plate 


Rhodora 


4, Ad- 


Fia. 


, Sanibel’s most abundant tree, Sabal palmetto. 


jacent communities of Spartina and Conocarpus. 


1215. Fia. 


PLATE 


Rhodor: Plate 1216 


PLATE 1216. Fra. 5, Strangling fig around cabbage palm at edge of mixed woods. 
Fic. 6, Typical button mangrove thicket, with Tillandsia in center. 


Rhodor: Plate 1217 


PLATE 1217. Fia. 7, Bacopa monnieri dominant in low places of Spartina marshes. 
Fic. 8, Spartina marsh, showing cabbage palm “islands,” on the slightly higher ground 
of ancient shell ridges. 


Rhodora Plate 1218 


PLATE 1218. Fia. 10, Thicket with Conocarpus, Sabal, and Yucca in transition zone 
between grassland and Conocarpus swamps. Fia. 11, Gumbo limbo at edge of Pal- 
metto jungle. 


Rhodor: Plate 1219 


PLATE 1219. Fig, 12, Sesuvium portulacastrum dominant on a low beach area 
with straggling Conocarpus in the background. Fic. 13, The broad beach at the 
western end of the island. 


Rhodor: Plate 1220 


PLATE 1220. Fic. 14, The stretch of broad beach at the western end of the island, 
showing Suriana maritima and Uniola paniculata, | F1G. 15, Agave sisalana persisting 
in an old field, after cultivation. 


Rhodor: Plate 1221 


PLaArEk 1221. Fra. 16, Solanum gracile, a common weed of disturbed areas. Fia. 17, 
Cakile fusiformis, the only species of this genus found on Sanibel. Fia. 18, Borrichia 
Jrutescens dominating a salt marsh, with buttonwood in background. 


1955] Cooley, 


Yucca aloifolia L. 

Agave americana L. 

Agave decipiens Baker 

Agave sisalana Perrine 

Hymenocallis keyensis Small 

Sisyrinchium atlanticum Bickn. 

Epidendrum tampense Lindl. 

Zeuxine strateumatica (L.) Schltr. 

Casuarina cunninghamiana Miq. 

Casuarina equisetifolia Forst. 

Salix longipes Anders. 

M yrica cerifera L. 

Batis maritima L. 

Quercus virginiana Mill. 

Ficus aurea Nutt. 

Parietaria nummularia Small 

Coccoloba uvifera (L.) Jacq. 

Atriplex arenaria Nutt. 

Chenopodium album L. 

Chenopodium ambrosioides L., var. 
anthelminticum (L.) Gray 

Salicornia perennis Mill. 

Suaeda linearis (Ell.) Moq. 

Achyranthes ramosissima (Mart.) Standley 

Amaranthus hybridus L. 

Froelichia floridana (Nutt.) Moq. 

Iresine celosia L. 

Philoxerus vermicularis (L.) R. Br. 

Boerhaavia paniculata L. C. Rich. 

Phytolacca americana L. 

Rivina humilis L. 

Sesuvium maritimum (Walt.) BSP. 

Sesuvium portulacastrum L. 

Portulaca oleracea L. 

Portulaca phaeosperma Urban 

Portulaca pilosa L. 

Annona glabra L. 

Cassytha filiformis L. 

Argemone leiocarpa Greene 

Argemone mexicana L. 

Capparis cynophallophora L. 

Cakile fusiformis Greene 

Lepidium virginicum L. 

Chrysobalanus icaco L. 

Chrysobalanus oblongifolius Michx. 

Abrus precatorius L. 

Acacia farnesiana (L.) Willd. 

Caesalpinia bonduc L. 

Caesalpinia crista L. 

Canavalia obtusifolia DC. 

Cassia tora L. 

Centrosema virginianum (L.) Benth. 

Chamaecrista aspera (Muhl.) Greene 


a 


- 


30 


bei ee d 


^ Coastal 
` shell ridge 


F 


x 


krj mi om 


Spartina 
marshes 


foo 


The Vegetation of Sanibel Island 


& savannas 
Mangrove 


Grasslands 
swamps 


-— 


— 


F 


Palmetto 
jungles 
Mixed 
woods 


C 


F 


285 


uud a 


-— 


286 Rhodora [Vor. 57 


Coastal 
shell ridge 
Spartina 
marshes 
Grasslands 
& savannas 
Mangrove 
swamps 
Palmetto 
jungles 
Mixed 
woods 


Crotalaria incana L. 
Crotalaria linaria (Small) Fern, & Schub. F 
Crotalaria rotundifolia (Walt.) Poiret F 
Crotalaria striata DC. F 
Dalbergia ecastophyllum (L.) Britt. I 
Desmodium tortuosum (Sw.) DC. F I 
Erythrina herbacea (L.) Small I 
Galactia parvifolia Rich. F F 
3alactia volubilis (L.) Britton C 
Neptunia lutea (Leavenw.) Benth, I 
Piscidia piscipula (L.) Sarg. I 
Pithecellobium unguis-cati (L.) Benth. I I 
Rhynchosia lewtonii (Vail) Small I F 
Rhynchosia michauxii Vail I F 
Rhynchosia minima (L.) DC, I 
Sesbania macrocarpa Muhl. I 
Sophora tomentosa L. I F F 
Tamarindus indica L. F 
Vicia acutifolia Ell. C 
Vigna repens (L.) Kuntze F 
Oxalis dillenii Jacq. FE 
Zanthoxylum clava-herculis L. I 
Zanthoxylum fagara (L.) Sarg. F F I 
Suriana maritima L. A 
Bursera simaruba (L.) Sarg. F I F 
Asemeia cumulicola Small F 
Asemeia grandiflora (Walt.) Blake, var. 

leiodes Blake F 
Polygala incarnata L. F 
Cnidoscolus stimulosus (Michx.) Engelmann 

& Gray F 
Croton floridanus Ferguson I I 
Euphorbia ammannioides H.B.K. F 
Euphorbia buxifolia L. C 
Euphorbia heterophylla L. C 
Euphorbia hirta L. C I 1 
Euphorbia hyssopifolia L. I 
Euphorbia maculata L, F 
Euphorbia pilulifera L. F 
Buphorbia trichotoma H.B.K. F 
Phyllanthus abnormis Baill. F 
Ricinus communis L. 
Stillingia aquatica Chapm. 
Stillingia sylvatica L. 
Rhus radicans L. 
Schinus terebinthifolius Raddi I 
Maytenus phyllanthoides Benth. I 
Cardiospermum halicacabum L. I 
Dodonaea jamaicensis DC. I I F 
Parthenocissus quinquefolia (L.) Planch. F F 
Gossypium hirsutum L, I 
Kosteletzkya virginica (L.) Presl F I 
Kosteletzkya virginica (L.) Presl, var. 

althaeifolia (Chapm.) A. Gray R 


1955] Cooley,— The Vegetation of Sanibel Island 


Sida carpinifolia L. f. 
Waltheria americana L. 


Piriqueta caroliniana (Walt.) Urban, var. 


glabra (DC.) Urban 
Carica papaya L. 
Mentzelia floridana Nutt. 
Acanthocereus floridanus Small 
Opuntia austrina Small 
Opuntia dillenii (Ker.) Haw. 
Opuntia flcus-indica (L.) Mill. 
Opuntia keyensis Britton 
Ammannia latifolia L. 
Lythrum lanceolatum Ell. 
Rhizophora mangle L. 
Conocarpus erecta L. 


Conocarpus erecta L., var. sericea Forst, 


ex DC. 
Laguncularia racemosa (L.) Gaertn. 
Eugenia anthera Small 
Eugenia axillaris (Sw.) Willd. 
Eugenia buxifolia (Sw.) Willd. 
Psidium guajava Raddi 
Saura angustifolia Michx., var. eatonii 
(Small) Munz. 
Ludwigia microcarpa Michx. 
Oenothera humifusa Nutt. 
Oenothera laciniata Hill 
Eryngium baldwinii Spreng. 
Hydrocotyle umbellata L. 
Ptilimnium capillaceum (Michx.) Raf. 
Jacquinia keyensis Mez. 
Ardisia escallonioides S. & C. 
Rapanea guianensis Aubl. 
Samolus ebracteatus H.B.K. 
Samolus floribundus H.B.K. 
Bumelia angustifolia Nutt. 
Sideroxylon foetidissimum Jacq. 
Forestiera porulosa (Michx.) Poir. 
Ligustrum ovalifolium Hassk. 
!ynoctonum mitreola (L.) Britton 
Polypremum procumbens L. 
Eustoma exaltatum (L.) Griseb. 
Sabatia campanulata (L.) Torr. 
Lochnera rosea (L.) Reichenb. 


Rhabdadenia biflora (Jacq.) Muell-Arg. 


Urechites lutea (L.) Britton 
Asclepias verticillata L. 
Cryptostegia grandiflora R. Br. 
Cynanchum palustre (Pursh) Heller 
Cynanchum scoparium Nutt. 
Sarcostemma clausum (Jacq.) R. Br. 
Calonyction aculeatum (L.) House 
Calonyction tuba (Schlecht.) Colla 
Cuscuta campestris Yuncker 


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& savannas 
Mangrove 
swamps 
Palmetto 
jungles 


= Grasslands 
Mixed 


— 


Qu 


ous 


287 


woods 


— 


288 Rhodora 


Ipomoea cathartica Poir. 

Ipomoea pes-caprae (L.) Sweet 

Ipomoea sagittata Cav. 

Heliotropium curassavicum L. 

Heliotropium parviflorum L. 

Heliotropium polyphyllum Lehm, 

Avicennia nitida Jacq. 

Lantana camara L. 

Lantana involucrata L. 

Lantana ovatifolia L. 

Lippia nodiflora L. 

Verbena scabra Vahl 

Trichostema suffrutescens Kearn. 

Physalis elliottii Kunze 

Physalis pubescens L. 

Solanum gracile Link. 

Bacopa monnieri (L.) Pennell 

Buchnera floridana Gandoger 

Capraria biflora L. 

Ernodia littoralis Sw. 

Gerardia maritima Raf. 

Scoparia dulcis L. 

Dicliptera assurgens Juss. 

Thunbergia fragrans Roxb. 

Chiococca alba (L.) A. Hitchc. 

Chiococca pinetorum Britton 

Galium hispidulum Michx. 

Galium tinctorium L., var. floridanum 
Wiegand 

Genipa clusiaefolia (Jacq.) Griseb. 

Houstonia nigricans (Lam.) Fern. 

Houstonia procumbens (Walt.) Standley 

Psychotria undata Jacq. 

Randia aculeata L. 

Sambucus simpsonii Rehder 

Melothria pendula L. 

Lobelia feayana A, Gray 

Scaevola plumieri Vahl 

Ambrosia artemisiifolia L., var. elatior 
(L.) Descourtils 

Ambrosia hispida Pursh 

Aster subulatus Michx. 

Baccharis angustifolia Michx. 

Baccharis dioica Vahl 

Baccharis glomerulifolia Pers. 

Baccharis halimifolia L. 

Berlandiera subacaulis Nutt. 

Bidens pilosa L., var. radiata Schultz-Bip. 

Borrichia frutescens (L.) DC. 

Cirsium horridulum Michx., forma elliottii 
(T. & G.) Fern. 

Conyza canadensis (L.) Britton, var. pusilla 
(Nutt.) Cronq. 


Old 
fields 


a-aa 


awas 


- 


à 


Coastal 
shell ridge 


F 


Spartina 
marshes 


Grasslands 
& savannas 


F 


C 


= Mangrove 
swamps 


C 


[Vor. 57 
e 

En 

vo yu 
Éu E 
-g Rale] 
Gu — 

RE 
I 


I I 
I 

I C 

R 

F F 

I 


1955] Cooley,—The Vegetation of Sanibel Island 289 


© as o 
-j o g a &H A s 
gE £2 38 BETE gu 
S Sm KE 39 aR Ew RB 
Ta © Sa 89 ag ag s 
=D Š A Ln mw 3 3 o 
od OF GE 64 SE ES SE 
Coreopsis leavenworthii T. & G. (o EO F 
Emilia sonchifolia (L.) DC. I 
Erechtites hieracifolia (L.) Raf. R F 
Erigeron quercifolius Lam. C F 
Eupatorium capillifolium (Lam.) Small C 
Eupatorium mikanioides Chapm. F F 
Flaveria floridana J. R. Johnston F C 
Flaveria linearis Lag. F [6 
Gnaphalium purpureum L, C 
Iva frutescens L. I 
Iva imbricata Walt. C 
Melanthera deltoidea Michx. C I F 
Mikania batatifolia DC. F C Fr 
Pectis linearifolia Urban F F 
Pectis prostrata Cav. I 
Pluchea purpurascens (Sw.) DC. F 
Pluchea rosea Godfrey F F 
Solidago sempervirens L., var. mexicana 
(L.) Fern. I Ko 
Sonchus oleraceus L. F 


Verbesina virginica L., var. laciniata A. Gray C 


290 Rhodora [Vor. 57 


NON-VALIDITY OF NUTTALLIAN NAMES 
IN FRASER'S CATALOGUE 


Luoyp H. SHINNERS 


In 1890 Edward L. Greene reprinted in Pittonia (vol. 2, pp. 
116-119) “A Catalogue of New and Interesting Plants Col- 
lected in Upper Louisiana, and principally on the River Mis- 
sourie, North America, [by T. Nuttall,] for sale at Messrs. 
Fraser’s Nursery For Curious American Plants, Sloane Square, 
King’s Road, Chelsea, London,” dated 1813. The reprint was 
made from a copy at the Philadelphia Academy, in which Nuttall 
had inserted his name after the first part of the title, indicated 
above by brackets. Under present rules of nomenclature, 
commercial catalogues and newspapers are not to be accepted 
as media for effective publication of scientific names after 1 
January 1953, but the regulation is unfortunately not retroactive. 
Following Greene's resurrection of Fraser's Catalogue, a number 
of names from it have come into circulation, credited to Nuttall. 
But an examination of the listings in the Catalogue, Pursh's 
Flora Americae Septentrionalis (1814), and  Nuttall's The 
Genera of North American Plants (1818), shows that Nuttall 
himself did not accept most of the names in the Catalogue, and 
could scarcely have been its author. 

Of the 89 names in Fraser's Catalogue, 71 are starred as new: 
69 species (one not named beyond genus), 2 varieties. In his 
Genera, Nuttall mentions only 32 of the new species and one 
of the new varieties. Three are credited to other sources than 
the Catalogue: Allium stellatum “Sims, Bot. Mag." ; Bartonia 
decapetala “Bot. Mag." (listed as synonym of B. ornata 
Nuttall, the only species, the genus starred as new); Oenothera 
macrocarpa "PH." (i. e., Pursh; listed as synonym of O. alaía 
n. sp. by Nuttall). Ten names are starred as new in the Genera, 
without mention of previous publication, though all had appeared 
in the Catalogue: Calymenia pilosa, Cactus viviparus, C. feroz, 
C. fragilis, Cheiranthus asper (spelled asperus in the Catalogue), 
Elaeagnus argentea (spelled argenteus in the Catalogue), Gaura 
coccinea, Oenothera caespitosa, O. serrulata, Rudbeckia purpurea 
var. serotina. Fourteen names appear in the Genera credited 
to “T. N. in Fras. Catal; two are misquoted (Glycyrrhiza 


1955] Shinners,—Nuttallian Names in Fraser's Catalogue — 291 


lepidota, which in the Catalogue is Liquiritia lepidota; Artemisia 
nutans, as synonym of A. cernua n. sp., is A. cernua in the 
Catalogue); two (in addition to Artemisia nutans) are placed 
in synonymy (Lactuca oblongifolia under L. integrifolia n. sp., 
with L. pulchella Pursh as an additional synonym; Seseli lucidum 
under S. divaricatum Pursh); ten are accepted and starred as 
new though with reference to previous publication in the Cata- 
logue (Amorpha canescens, A. nana, Calymenia angustifolia, 
Dalea aurea, Lilium andinum, Malva coccinea, Phalangium 
esculentum, Psoralea esculenta, P. incana, Troximon glaucum). 
Six more are credited in the Genera only to “Fras. Catal.’’; 
one is neither described nor starred as new (Erigonum flavum), 
the remaining five are both (Astragalus gracilis, Oenothera 
albicaulis, Penstemon grandiflorum, P. cristatum, P. erianthera). 


Eighteen new names appear in Fraser's Catalogue with some 
descriptive remarks. In addition there is a brief joint descrip- 
tion for two species of Sideranthus. Only four are acknowledged 
by Nuttall to have been his: Amorpha nana, Lilium andinum, 
Malva coccinea, Phalangium esculentum. Two are credited by 
Nuttall only to “Fras. Catal.": Eriogonum flavum (not described 
in the Genera), Penstemon grandiflorum. One is credited to 
Pursh without reference to the Catalogue, though first described 
there: Oenothera macrocarpa (as synonym of O. alata n. sp.). 
Ten are not mentioned by Nuttall in his Genera: Astragalus 
crassicarpus, Dalea enneandra, Hyssopus anethiodorus, Rud- 
beckia columnifera, Sideranthus integrifolius, S. pinnatifidus, 
Thuraria herbacea, Virgilia grandiflora, Vitis campestris, Yucca 
glauca. The first, second, fourth, and tenth of these are in 
current use, attributed to Nuttall, though he quite plainly 
disclaimed them. Three more are described as new in 1818 
without reference to the Catalogue: Cactus viviparus, Oenothera 
caespitosa (spelled cespitosa in the Catalogue), Rudbeckia pur- 
purea var. serotina. 

It must be concluded from this record that Nuttall was not 
the author of Fraser’s Catalogue, and did not consider it an 
accepted medium for publication of new names. His autograph 
on the Philadelphia copy was intended to mean not “A Cata- 
logue, by T. Nuttall, of Plants Collected in North America," 
but “A Catalogue of Plants, Collected in North America by 


292 Rhodora [VoL. 57 


T. Nuttall.” He evidently supplied manuscript names with 
some of the plants which he sent to the Frasers. References 
in his Genera to their Catalogue were an effort to salvage some 
(not all) of these names. He was resentful of Pursh's having 
anticipated him in publishing them, as some of his remarks 
indicate. Under Amorpha canescens he says, “Mr. Pursh places 
his usual mark of v.v. to this species, although he had never 
seen a flowering specimen except in my herbarium." In a 
footnote under Bartonia ornata he writes, “Mr. Pursh, before 
he had perused the notes which I had made from the living plant 
on the Missouri, with an intention of rendering them publie, 
had not then, by his own acknowledgment, anything like ma- 
terials for publishing this genus, . . . This unfortunate want 
of fidelity, prevented me from communicating to Mr. F. Pursh, 
many of the plants which now appear in this work. . .. It 
was surely not honorable in Frederick Pursh, whom I still 
esteem as an able botanist, to snatch from me the little imaginary 
credit due to enthusiastie researches made at the most imminent 
risk of personal safety!” 

It would be unjust to accuse Pursh of totally unethical be- 
havior. Nuttall was a shy and secretive man, and it is altogether 
likely that he did not confide to Pursh his plan to publish a 
North American flora when the German botanist visited him 
and examined his herbarium. Pursh must have guessed that 
at least some of the names in Fraser's Catalogue had originated 
with Nuttall, but he did not know which ones, nor did he know 
precisely Nuttall's intentions concerning them. Communica- 
tion was slow and diffieult, and Nuttall himself, in his frequent 
travels, not easily accessible. 


But the fine etiquette of the matter, as seen at this late date, 
cannot be our basis for determining the validity of the names 
published in the Catalogue. These were anonymous as pub- 
lished. We do not know whether the Catalogue was prepared 
by one of the Frasers or by a clerk in their employ. Since the 
names were not avowedly accepted by any author, they were not 
validly published. (Cf. Jacques Rousseau, “One Plant Name 
in Search of an Author," Taxon 4: 40-42, 1955.) It might be 
argued that we must at least keep the names claimed later by 
Nuttall. We should then have to keep those not claimed by 


1955] Shinners,—Nuttallian Names in Fraser's Catalogue — 293 


him also. Nuttall himself did not retain all the names which 
he acknowledged to be his. We cannot very well allow that he 
could legitimize some parts of the Catalogue and not others. 
His inconsistent treatment gives us sufficient legal grounds, 
if anonymity be not enough, to disregard all names in the Cata- 
logue. 

Nor can it be held that usage gives any support to retention 
of the names. The inconsistencies of later authors are strikingly 
shown in the eighth edition of Gray's Manual. “Yucca glauca 
Nutt." was an anonymous name explicitly disclaimed by Nuttall, 
who does not mention it in his Genera, but lists Y. angustzfolia 
Pursh. ‘Psoralea esculenta Pursh” first appeared in Fraser's 
Catalogue as a nomen subnudum, but unlike Yucca glauca, was 
claimed by Nuttall in 1818. The same is true of Amorpha 
nana (A. microphylla Pursh). “Dalea enneandra Nutt." is an 
identical case with that of Yucca glauca; Nuttall in 1818 called it 
D. laxiflora Pursh, and did not mention the binomial later er- 
roneously attributed to him. Astragalus caryocarpus Ker is in- 
consistently (but correctly!) adopted in place of A. crassicarpus, 
another anonymous name never accepted by Nuttall. *''Glycyr- 
rhiza lepidota (Nutt.) Pursh should strictly be credited to Pursh 
alone: if dated from 1813, it may be considered a misprint 
corrected by Nuttall in 1818 from Liquiritia to Glycyrrhiza. 
“Sphaeralcea coccinea (Pursh) Rydb.” is correct, but again 
inconsistent, since the basonym (Malva coccinea) first appeared 
in Fraser's Catalogue aecompanied by a description, and was 
subsequently claimed by Nuttall as his. Mamillaria vivipara 
(Nutt.) Haw. has basonym dated from 1818, not 1813. ‘Pen- 
stemon grandiflorus Nutt." (credited by him only to Fraser’s 
Catalogue, not to himself) must give way to P. Bradburi? Pursh. 
Ratibida columnifera (Nutt.) Woot. & Standl. (basonym never 
acknowledged by Nuttall) returns to its long familiar name, 
R. columnaris (Pursh) Rafinesque. 

! Correctly under then existing rules of nomenclature, which outlawed names 
based on mixtures. As since revised, Article 76 of the International Code requires 
the adoption of A. carnosus Pursh for this species: though based in large part on the 
later-named Sophora sericea Nutt., there is no doubt but that the name carnosus 
was intended to describe the fruit of Astragalus caryocarpus (also named later). 


Nuttall by implication, and Torrey & Gray explicitly, typifled Pursh's epithet with 
the Astragalus element, for which it must now be revived. 


294 Rhodora [Vor. 57 


THE ARCHER METHOD FOR MOUNTING 
HERBARIUM SPECIMENS 


REED C. ROLLINS 


SINCE the publication by W. Andrew Archer! in 1950 of an 
article entitled, “New Plastic Aid in Mounting Herbarium 
Specimens", there has been an increasing interest in the use 
of plastic for this purpose. In presenting his formula five years 
ago, Dr. Archer invited prospective users to further improve it 
and to modify methods of procedure to suit particular needs. 
We have been using the Archer Method with some slight modifi- 
cations at the Gray Herbarium with excellent results for several 
years. The frequent calls for information concerning our 
present formula and procedure have prompted me to put these 
into print for the convenience of those interested. 


ORIGINAL ARCHER FORMULA 


‘Toluene—800 cc. 

Methanol—200 ec. 

Ethel cellulose (Dow Chemical Ethocel?) 
standard 7 cps.—250 grams. 

Dow Resin 276 V-2—75 grams. 


New York BOTANICAL GARDEN MODIFIED FORMULA 


Toluene—720 cc. 
Methanol—180 ce. 
Ethocel—250 grams 
Dow Resin—75 grams 


Gray HERBARIUM MODIFIED FORMULA 


Toluene—880 cc. 
Methanol—220 cc. 
Ethocel, standard 10 cps. 
Dow Resin—75 grams 


1750 ec. (560—650 grams). 


First mix methanol and toluene; then add and dissolve the 
resin completely. Finally mix in the ethocel adding it gradually. 
The resulting stiff coarse mixture should be stirred slowly until 
the ethocel is partly dissolved. "There will be many bubbles 


! Rhodora 52: 298-299. 1950. 

? Ethocel and Dow Resin are obtainable from the Dow Chemical Company, Mid- 
land, Michigan. At present the Ethocel may not be purchased in less than 50 pound 
lots at a cost of about $40.00. The Dow Resin is sold in a 5 gallon container. This 
may seem expensive, but fifty pounds of Ethocel and five gallons of Dow Resin to- 
gether with appropriate amounts of the solvents have provided the plastic for mount- 
ing over 40,000 specimens. 


1955] Rollins,——Mounting Herbarium Specimens 295 


and large particles present at this stage, but the solution will 
clear in about 24 hours. 

It will be quickly noted that the users at both New York 
and at Gray found the original formula less viscous than desirable. 
In the New York modification, less solvent is used, and the 
proportions of Toluene and Methanol are maintained. In 
working toward greater viscosity, my own procedure was to 
obtain Ethocel of standard 10 eps. (to replace standard 7 cps.) 
and to increase the proportion of ethel cellulose. The first 
combination that came close to meeting with satisfaction was 
based on the use of 1000 cc. of solvent. However, this proved 
to be difficult to use in the dispensers; hence, another 100 ce. 
of the solvent was added to the formula. It should be understood 
that the formula was arrived at by trial and error, and I feel sure 
that several different combinations of the basie materials orig- 
inally put together for this purpose by Archer would be very 
satisfactory. 

In a recent letter, Dr. Archer notes that the Gray Herbarium 
Formula, using 625 gms. of ethocel, has proven to be too thick 
for some dispensers. He is now using the following formula: 
toluene 880 cc.; methanol 220 cc.; ethocel (standard 10 cps.) 
575 gms.; resin 75 gms. I feel sure that the use of a weight 
measure for the ethocel is at the base of the inconsistencies 
experienced by different persons in making up the formula. For 
example, the weight of 1750 cc. of one lot of ethocel was 659.7 
gms., while the weight of 1750 cc. of another lot was 560.5 gms. 
It appears that volume measure is more reliable than weight 
measure. At least, we have always used and gotten consistent 
results with a volumetric measure in handling the ethocel. It 
may be necessary to make minor adjustments in the formula 
with each new lot of ethocel. However, I do not believe the 
proportions are at all critical and they may be shifted somewhat 
without materially devaluing the final product. 

We find it possible to mount large woody materials, even pine 
cones, by use of the above formula. The viscosity is such that 
it will hold well on the sides of a branch or stem and not produce 
a weak point in the strand by thinning at the vertical part. 
Our aim is to put the plastic in a strap-like form over the stem 
or a portion of the leaf etc., in such a way as to have each end 


296 Rhodora [Vor. 57 


of the plastie strap firmly anchored on the mounting sheet. 
On large leaves, holes are punched with a paper punch a centi- 
meter or two from the edge at intervals around the leaf. The 
plastic strap is spread from the mounting paper to the hole 
through which it reaches the paper beneath the leaf. Thus, 
a large leaf can be as firmly strapped to the mounting sheet as 
can stems and other parts of the plant. This procedure essen- 
tially eliminates the need or the desirability of smearing the 
specimen with plastic on the side placed downward on the 
mounting sheet. The mounting job, then, is essentially one 
of strapping the specimen to the sheet with plastie straps squirted 
from a pressure oil can. Occasionally on especially bulky 
materials or where a specimen will not lie flat, plastic is used 
beneath the projecting part of the specimen to provide a kind 
of seat upon the paper. It is important to have a strap stretch 
from paper at one end to paper at the other. The bonding 
of the plastie with the specimen itself is often imperfect and 
cannot be relied upon to hold. This is not true with the plastic 
to paper bonding which is exceptionally strong. 

Some users of plastic as a mounting medium have experienced 
difficulty from having stacked the sheets before the plastic was 
fully dry. The sheets stuck together and produced real dif- 
fieulty. Actually, the plastic dries quite rapidly, and the sheets 
can usually be safely stacked in about 40 minutes to an hour. 
Also, some users require a large amount of table space for their 
mounting operation because the specimens are laid out on the 
sheets side by side on tables. After a group of specimens are 
laid out and weights put into position, the mounter applies the 
plastic. While these are drying, a second group of specimens 
is laid out, ete. When the first group is dry, the sheets of 
specimens are then stacked. We have eliminated the need 
for a large amount of table space and made the time factor in 
drying unimportant by two simple procedures. Furthermore, 
the mounter is permitted to sit down while working. 

The sheet of mounting paper is placed upon a double-faced 
corrugated cardboard slightly larger in size than the sheet. 
The label is affixed to the sheet; the specimen is placed and 
weighted; and finally the plastic is applied. The mounter 


3 For weights, we have found broken up type-plates to be ideal. These lead sheets 


1955] Rollins, —Mounting Herbarium Specimens 297 


then puts four small wooden blocks or, other objects of similar 
nature, on the cardboard near the four corners. The cardboard 
carrying the mounted specimen with the plastic still wet is then 
stacked. Each successive cardboard in the stack is held above 
the wet plastie of the sheet below, and the stack may be left until 
the plastic is dry without inconvenience. The wooden blocks are 
of two or three lengths so that four blocks as much as an inch 
long may be selected if the thickness of the specimens requires 
it. The cardboards are easily obtained from paper box manu- 
facturers. For us, the double-faced corrugates commonly used 
in plant presses serve very well. 

Dispensing the plastic provides no great problem. We find 
the “pistol grip” pressure oil can of the type used for spraying the 
springs of automobiles very satisfactory. This type of dis- 
penser, called to my attention by Dr. David D. Keck, can be 
used continuously for several months without cleaning. At 
first, we were careful to cover the end of the spout while not in 
use, thinking the plastie would harden in it. However, this 
proved to be an unnecessary precaution, for the plastic seals 
the end of the spout very quickly; and the short plug formed 
is easily squeezed out even after several days have elapsed. 
If the plastic hardens in the spout, the nozzle can be soaked 
in the solvent. The spout of the dispenser is made of brass, 
and the copper present reacts with the plastic to make the 
plastie greenish in color. If the dispenser has not been used 
for several days, the plastic in the spout will be quite colored. 
For those who find this objectionable, it is a small waste to 
squeeze that amount away before starting the mounting of 
specimens. The barrel of the dispenser should not be brass, 
for the whole of the contents will then become colored. When- 
ever the dispenser becomes fouled after long usage, it can be 
cleaned by disassembling it and submerging the parts in the 
solvent for several days. 

Our mounters find it worthwhile to wipe the end of the spout 
of the dispenser frequently to keep the flow of plastic smooth. 
This produces a neater strap and prevents unnecessary threading 
of the plastic itself. In fact, they find it convenient to work 


can be readily broken into many different sizes. Of course, any relatively heavy 
bits of metal will do very well. 


298 Rhodora [Vor. 57 


with the dispenser in one hand and a small cloth in the other 
wiping the nozzle after every few straps extruded. 

The plastic once made up appears to have excellent keeping 
qualities if kept in an airtight container. Once dried after use 
in mounting, it is tough and durable and not easily broken even 
by a direct hit with a metal object. There is some contraction 
of the plastie on drying, and this tends to pull the specimen 
tight against the paper giving the full value of support to the 
specimen that the paper can provide. The fact that the plastic 
does contract makes it inadvisable to use long straps, especially 
if several are put parallel to one another. Several long parallel 
straps will contract enough to pull the mounting paper into 
wrinkles. We use straps mostly below an inch in length although 
occasional longer ones are required and can be safely used. 
The first tendency of the mounter in using the plastic is to put 
long straps over several culms of grass or similar specimens. 
This should be avoided not only because it tends to form wrinkles 
in the sheet, but also produces unsightly mounted specimens as 
well. While the strap should be kept relatively short, its thick- 
ness may be increased by passing the stream of plastic back and 
forth several times. 

Some feel it is too early to evaluate completely the Archer 
Method from the point of view of what may happen to the plastic 
straps after forty or fifty years in the herbarium. However, 
if the plastic holds after five years, it seems to me that it might 
be expected to hold indefinitely. One thing we do know is 
that the plastic does not appear to become brittle with the passage 
of time. And there are apparently no a priori grounds upon 
which to predict any appreciable change in the chemical or the 
physical nature of the plastic straps unless they are subjected to 
a powerful solvent. The latter seems highly unlikely in a well 
kept herbarium. 

The chief advantage of the Archer Method is the speed-up 
possible in the mounting. In rough calculations, we have found 
that a mounter can prepare and mount three or four specimens 
using this technique in the time required to mount one specimen 

4 Paradichlorobenzene has been reported to soften the plastic even after it has dried 
thoroughly. Our own tests with a high concentration of p d b at room temperature 


did not produce any noticeable affect. However, no tests were made under varying 
conditions of temperature and moisture. 


1955] Rollins —Mounting Herbarium Specimens 299 


using paste and gummed cloth strips. Where heavy specimens 
are sewed, the time saved is even greater, for sewing is not 
required. A second advantage comes from the possibility of 
repairing specimens in the herbarium itself. A dispenser filled 
with plastic can be kept handy for remounting loose specimens 
or broken parts by the botanist while he is actually working 
in the herbarium. This eliminates the necessity of having 
the specimen out of the herbarium in the hands of the mounter 
and of having to refile it in the herbarium at a later time. Per- 
sonally, I believe the use of plastic makes a better looking 
mounted specimen. However, all botanists may not agree 
on this point.—GRAY HERBARIUM OF HARVARD UNIVERSITY. 


Pinus RIGIDA MILLER IN QuEBEC.—Owing to the lack of 
authentic records, the presence of Pinus rigida Miller in the 
province of Quebec had always been in doubt. During a field 
trip in the region of Saint-Chrysostóme, Chateauguay County, 
made on September 4th, 1954, Mr. Lionel Cinq-Mars and the 
author have found a natural stand of Pinus rigida at Cairnside. 
The mixed stand covers an area of about a square mile. The 
trees growing in the forest attain around thirty-five feet in 
height and about eight inches in diameter. When they grow 
on the bare Potsdam sandstones, they are stunted and the 
lower branches are heavily covered with cones. Pinus resinosa 
Ait. and Pinus Strobus L. are also present in the area.—ERNEST 
ROULEAU, HERBIER MARIE-VICTORIN, UNIVERSITÉ DE MONTRÉAL. 


Volume 57, no. 681, including pages 241—268, was issued 4 October, 1956. 


Hovova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS 

RALPH CARLETON BEAN Associate Editors 
RICHARD ALDEN HOWARD 

CARROLL EMORY WOOD, JR. 


Vol. 57 November, 1955 No. 683 


CONTENTS: 


Contributions to the Flora of Nova Scotia: V. Results of Explora- 
tion in Cumberland County. W. B. Schofield.............. 301 


Plants New to Missouri. FE. J. Palmer and J. A. Steyermark..... 310 


Contributions to the Flora of Southern Illinois. Robert H. 
Mohlenbrock oa pret sk RES MEE Oe ates IS 319 


An Additional Species of the Lichen Genus Buellia from the West 
Indies. «Henry Imahaug ecm. sooo NLIS I 323 


The New England Botanical Club, Ine. 


8 and 10 West King St.. Lancaster, Pa. 
Botanical Museum, Oxford St., Cambridge 38, Mass. 


RHODORA.—A monthly journal of botany, devoted primarily to the 
flora of the Gray's Manual Range and regions floristically related. 
Price, $4.00 per year, net, postpaid, in funds payable at par in 
United States currency in Boston; single copies (if available) of not 
more than 24 pages and with 1 plate, 40 cents, numbers of more 
than 24 pages or with more than 1 plate mostly at higher prices 
(see 3rd cover-page). Back volumes can besupplied at $4.00. Some 
single numbers from these volumes can be supplied only at advanced 
prices (see 3rd cover-page). Somewhat reduced rates for complete sets 
can be obtained on application to Dr. Hill. Notes and short scien- 
tific papers, relating directly or indirectly to the plants of North 
America, will be considered for publication to the extent that the 
limited space of the journal permits. Illustrations can be used 
only if the cost of engraver's blocks is met through the author or his 
institution. Forms may be closed five weeks in advance of publica- 
tion. Extracted reprints, if ordered in advance, will be furnished at 
cost. 


Address manuscripts and proofs to Reed C. Rollins, 
Gray Herbarium, 22 Divinity Ave., Cambridge 38, Mass. 


Subscriptions (making all remittances payable to RHODORA) to 
Dr. A. F. Hill, 8 W. King St., Lancaster, Pa., or, preferably, Botanical 
Museum, Oxford St., Cambridge 38, Mass. 


Entered as second-class matter March 9, 1929, at the post office at 
Lancaster, Pa., under the Act of March 3, 1879. 


INTELLIGENCER PRINTING COMPANY 


Specialists in Scientific and Technical Publications 
EIGHT WEST KING ST., LANCASTER, PA. 


CARD-INDEX OF NEW GENERA, SPECIES AND 
VARIETIES OF AMERICAN PLANTS 


For all students of American Plants the Gray Herbarium Card-index 
of Botanical Names is indispensable. It is a work of reference essen- 
tial to scientific libraries and academies and all centers of botanical 
activity. It includes genera and species from 1885 to date. "The sub- 
divisions of species from 1885 to date are now included and from 1753 
to 1886 are in the process of being inserted. Issued quarterly, at $25.50 
per thousand cards. 


GRAY HERBARIUM of Harvard University, 
Cambridge 38, Mass., U. S. A. 


TRbooora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 57 November, 1955 No. 683 


CONTRIBUTIONS TO THE FLORA OF NOVA SCOTIA: 


V. RESULTS OF EXPLORATION IN 
CUMBERLAND COUNTY 


W. B. ScHOFIELD 


CAPE BLoMIDON, Kings County, has long been known as a 
botanically rich area containing a number of interesting montane 
species. Saxifraga Aizoén Jacq. (var. neogaea Butters) was re- 
ported from the area many years ago (Lawson, 1884), but no 
substantiating specimen seems to have survived.! A. E. Roland 
(1938, 1947) reported Draba arabisans Michx. and Arabis 
Drummondii Gray which are locally quite abundant on an ex- 
posed boulder slope. Recently J. S. Erskine (Smith & Erskine, 
1954) has collected Lycopodium Selago L., Poa glaucantha 
Gaudin, and Trisetum spicatum (L.) Richter, var. pilosiglume 
Fernald from Amethyst Cove, a short distance from the Cape; 
these are local on the cooler cliff shelves. 

A relationship of the flora of Cape Blomidon to that of north- 
ern Cape Breton Island has been implied on the basis of rather 
inadequate botanical evidence, but much evidence is accumulat- 
ing as the floras of both areas become better known. The rela- 
tion of Blomidon’s flora to that of Cumberland County has been 
practically unknown, but the few collections from that county 
had suggested a positive relationship (Smith & Erskine, 1954). 

Intensive botanical exploration in the northern portion of 
Cape Breton Island has stimulated interest in the montane 

1A recent re-collection of this species from Cape Blomidon indicates that it is still 
present there: scattered plants on dry soil, shelves and pockets of lower portion of cliff, 


about 2 miles south of Cape Split, Cape Blomidon, Kings County, W. B. Schofield 
and D. H. Webster 5873. 


302 Rhodora [Vor. 57 


element of Nova Scotia’s flora. It was one of the aims of the 
writer to accumulate evidence from Cumberland County either 
supporting or refuting the implied relationships of its flora to 
more northerly areas. This exploration, carried on during the 
summers of 1953 and 1954, was done under the sponsorship of 
the Nova Scotia Research Foundation. The results have been 
most gratifying. 

Unquestionably the most interesting association of montane 
species known in peninsular Nova Scotia is to be found in the 
vicinity of Cape d'Or, Cumberland County. The montane 
aspect of the west-facing cliff-top is quite remarkable. There 
the strong winds have so blasted the area behind the precipice 
that a narrow treeless border has been left wherein one finds 
the following association of species: Saxifraga Aizoón Jacq., 
var. neogaea Butters; Astragalus Robbinsii (Oakes) Gray; Oxy- 
tropis johannensis Fern.; Chrysanthemum Leucanthemum L., 
var. pinnatifidum LeCoq & LaMotte; Festuca rubra L.; Sedum 
Rosea (L.) Scop.; Plantago juncoides Lam, var. decipiens (Barné- 
oud) Fern.; Poa pratensis L. (sensu lato); Poa compressa L.; 
Oenothera biennis L.; Cardamine parviflora L., var. arenicola 
(Britt.) Schulz; Antennaria canadensis Greene; Campanula 
rotundifolia L.; Trisetum spicatum (L.) Richter, var. pilosiglume 
Fern.; Cirsium arvense (L.) Scop.; Draba arabisans Michx.; 
Achillea lanulosa Nutt.; Solidago bicolor L., and Agropyron 
trachycaulum (Link) Malte, var. novae-angliae (Scribn.) Fern. 

As can be noted by the list, highly competitive weedy species 
are producing a marked change in the vegetation of the cliff- 
top. Doubtless the rare species will continue to persist in the 
more exposed areas where they tend to flourish. 

Behind this narrow band is the tangled barrier of alder, 
which forms another distinct border to the cliff-margin vegeta- 
tion. In areas among these alders are found small open patches 
of Potentilla fruticosa L., and Heracleum maximum Bartr. 

Other areas that harbor interesting species include a high 
cliff at New Prospect, near Parrsboro, where Draba arabisans 
Michx.; Carex rosea Schkuhr; C. convoluta Mackenz.; Muhlen- 
bergia mexicana (L.) Trin.; Milium effusum L.; and Festuca 
obtusa Biehler, are found in abundance (although not montane, 
these species are extremely local in the province). At Advocate, 


1955] Schofield,—' The Flora of Nova Scotia 303 


above the salt marsh, on the eastern bank of Burke Brook, 
Montia lamprosperma Cham. abounds near a cold springy 
area and Stellaria humifusa Rottb., carpets large patches of 
the marsh near the brook. 

The cliffs of Isle Haute, the largest of Nova Scotia's Bay 
of Fundy islands, also hold several interesting species, among 
which are Draba arabisans Michx.; Arabis Drummondii Gray; 
Poa glaucantha Gaudin and Lycopodium Selago L. (the final 
species being exceedingly rare). 

The deeper brook valleys yielded rather sparse collections 
of montane species. McAlese Brook, New Prospect, was by 
far the richest, possessing on the moist slope near its waterfall 
Carex atratiformis Britt., and on the cliff above Lycopodium 
Selago L., and Dryopteris fragrans (L.) Schott., var. remotiuscula 
Komarov. 

The flora of Cumberland County cannot be considered com- 
pletely known on the basis of these two summers' collections, 
but these certainly point out the importance of intensive ex- 
ploration in one area. In the following list are four species 
new to the province and numerous records of very local species. 

All collections mentioned are from Cumberland County unless 
noted otherwise; all numbers given without the name of a col- 
lector are those of the writer. Specimens have been deposited 
at the Acadia University Herbarium. 

Special acknowledgement is due the sponsoring institution: 
Nova Scotia Research Foundation, without whose support 
much of this exploration could not have been done. The 
writer is also most grateful for the considerable aid and helpful 
advice received from Dr. E. C. Smith of Acadia University and 
J. S. Erskine of Wolfville, Nova Scotia. 

Lycopodium Selago L. Cliff-top, south side, Isle Haute (J. S. Erskine & 
W. B. Schofield, JSE 53.038); locally abundant in rock crevices of river bank, 
West Moose River (3170); local colony on moist cliff facing McAlese Brook, 
New Prospect (3234). The species, widely distributed, but local, in northern 
Cape Breton Island, is very rare in peninsular Nova Scotia. 

Sparganium minimum Fries. Very abundant in marginal water of Wig- 
more Lake (4245). Known from a few local stations in the province; the 
above colony covered an area of about fifteen square yards. 

Potamogeton spp. Several interesting species of this genus were collected 
and are to be treated later by D. H. Webster in a paper devoted to the genus. 


Najas flexilis (Willd.) Rostk. & Schmidt. This species is now known to 
be fairly widespread (cf. Smith & Schofield, 1952). "The following collections 


304 Rhodora [Vor. 57 


from Cumberland County mark its wide distribution there: rare in marginal 
shallows of Leak Lake (3557); marginal shallows of Lake Killarney (4165); 
on sand of shallow water, Newville Lake (3606); in wrack of Mattatall Lake 
(3996); small riverside pond, Head of River Hebert (W. B. Schofield & D. H. 
Webster 5632). 

Sagittaria graminea Michx. Common on gravelly shore of Newville Lake 
(3562); wet margin of Dewar’s Lake (4073); rare in mud of lake margin, 
Lake Killarney (4156); margins of Big Lake, Victoria (5394). 'The above 
collections do not support the suggestion of Roland (1947), “apparently local 
and rare in the northern and central parts of the peninsula." Although it 
often does not flower, the stiffly arching, thick, underwater leaves readily 
distinguish it from any other known Nova Scotian species of Sagittaria. 

Festuca obtusa Biehler. Abundant on hardwood slope, New Prospect 
(3453); very abundant on rich hardwood slope about one mile east of Refugee 
Cove (5191). A collection from Kings County is also of interest: occasional 
among small trees at base of high cliff, Cape Blomidon (Schofield & Webster 
4230). Previously known only from Five Mile River, Hants County (Fernald, 
1921). 

Poa alsodes A. Gray. Damp margin of MeGahey Brook (W. B. Schofield 
& P. A. Bentley 4295). This very slender species of Poa is readily distin- 
guished in the field from any other Nova Scotian species, for the very capillary 
rachillas and culms, that often tumble over, are quite different from the far 
coarser aspect of most species. Known from a few locations in N. S.; in the 
above location it grew abundantly on the damp shaded flood plain of the 
brook. 

P. glaucantha Gaudin. The following collections are all from Isle Haute: 
occasional in cliff crevices, usually damp, near Pigeon Point, North Side 
(3760); rock crevices, North Side, Boar's Head (3835); abundant on upper 
slopes and crevices, central North Side (3838); rare in cliff crevices, Wrack 
Cove (3886); rock crevices, Western Slope (3904). Previously known from 
a number of stations in northern Cape Breton (Smith & Schofield, 1952) 
and from Cape Blomidon, Kings County (Smith & Erskine, 1954). 

Schizachne purpurascens (Torr.) Swallen. Very rare in rock crevices, West 
Moose River (3760). Hitherto, in peninsular Nova Scotia, this species was 
known from only Moore's Falls, near Kentville, Kings County. Two further 
collections mark its presence on North Mountain, Kings County: rare in 
spruce woods, Cape Split (8310A); occasional clumps on grassy slopes above 
boulder scree, Cape Blomidon (Schofield & Webster 4550). The species 
is locally abundant in northern Cape Breton Island. 

Sphenopholis intermedia Rydb. Damp cliff gully, Moose Island, Colchester 
County (4041); seepy, mucky slope near Indian Springs Brook, Cape Blomidon, 
Kings County (Schofield & Webster 5233). Dore and Roland (1942) re- 
marked that this grass was found “where its roots were in contact with lime- 
stone or gypsum." The basalt of the above stations (plus inclusions of 
gypsum found in the sandstone) would probably produce a similar basic 
soil. Very local in Nova Scotia and previously unknown from the north- 
central counties. 

Trisetum spicatum (L.) Richter, var. pilosiglume Fern. Frequent on exposed 
cliff headlands, Cape d’Or (Schofield & Bentley 4798). Locally abundant 
in northern Cape Breton Island and known in peninsular Nova Scotia from 
the Cape Blomidon area. 

Muhlenbergia mexicana (L.) Trin. Common at moist base of cliff and in 


1955] Schofield,—' The Flora of Nova Scotia 305 


crevices, New Prospect (3449); banks of Wallace River, Wentworth (5248). 
Known previously from Kings, Hants, and Halifax Counties (Smith & Erskine, 
1954). The station in Wentworth is most interesting, in that it is the first 
to be discovered on a river not emptying into the Minas Basin. 

Milium effusum L. Occasional under hardwoods of slope at base of cliff, 
New Prospect (3142); damp woodland margin of MeGahey Brook (Schofield 
& Bentley 4785). A species fairly frequent in the rich hardwood stands 
of northern Cape Breton Island, it was previously known from the mainland 
of the province from Cape Blomidon, Kings County, and Five Mile River, 
Hants County. In the hardwood forests of Cape Chignecto the species 
flourishes. 

Eleocharis nitida Fernald. Rare on road to Cape d'Or (Schofield & Bentley 
4817). Other collections of this rare species: occasional in moist soil over 
basalt, Elliott Lake, Annapolis County (W. B. Schofield & J. S. Erskine 3113); 
frequent on woods road, Cape Split, Kings County (3308, 3319); damp wood- 
land roadside, Economy Mountain, Five Islands, Colchester County (4949); 
damp pockets in burned-over area, north east end, Seatari Island, Cape Breton 
County (E. C. Smith, W. B. Schofield, D. H. Webster, L. Slipp & J. Taylor 8596). 
The above collections were all found in association with soils derived from 
volcanic rock; this agrees with Fernald’s remark about this species (Fernald, 
1922). 

E. ovata (Roth) R. & S. var. ovata. In small tufts, margin of pond, True- 
manville (4200). This is a re-collection of the typical variety where Fernald 
(1950A) collected var. Heuseri Uechtritz and, apparently, the species, for 
he reports both from N. S. in the 8th edition of Gray's Manual (Fernald, 
1950B). A collection of J. S. Erskine from Sandy Cove, Digby County 
(Erskine 52.1268) is also the species. On the margin of a dried-up pond 
behind the barrier beach at Black Point, Halifax County, the species abounds 
(E. C. Smith, W. B. Schofield, D. H. Webster & P. A. Bentley 12644). 

E. ovata var. Heuseri Uechtritz. This is very abundant at Truemanville, 
being far more abundant than the species (4200a). A collection from the 
shore of Earltown Lakes (E. C. Smith, D. H. Webster & P. A. Bentley 11749) 
is also this variety. Previously known from Truemanville (Fernald 1950A). 
It is interesting to note that both of these taxa are found on soils derived 
from basic rock. Even the collections from Black Point are found on a small 
local area of Carboniferous limestone. 

Scirpus cespitosus L., var callosus Bigel. A single clump in rock crevice, 
Moose River (3251). This species, although exceedingly common on the 
Atlantic slope of Nova Scotia, seems to be very rare and local in the north- 
central counties. 

S. hudsonianus (Michx.) Fern. Abundant in wet cliff crevices, West Moose 
River (3174). This species is also rare in the north-central counties, being 
represented by only one collection (in the vicinity of River Hebert) on Roland's 
distribution map of the species (Roland, 1947). 

Rhynchospora fusca (L.) Ait. Abundant on boggy margin of Leak Lake 
(8385); common on moist margin of swamp, west end, Dewar's Lake (4045). 
Commonest in the south-western counties of the province, these mark further 
collections in the central portion. 

Rhynchospora capitellata (Michx.) Vahl. Abundant on the swampy margin 
of Dewar’s Lake (4064). This is also best known from the south-western 
counties. The above collection marks its extension into the northern part 
of the province. In common with all collections made outside the south-west- 


306 Rhodora [VoL. 57 


ern counties, the above material was very much slenderer than that common 
to those counties. 

Cladium mariscoides (Muhl.) Torr. Abundant on margin of Leak Lake 
(3481); marginal water of Mattatall Lake (4007); common on beach of Dewar’s 
Lake (4041); damp swamp of Big Lake (4249). This species is much com- 
moner in the province than previous records indicate. 

Carex rosea Schkuhr. Abundant under hardwoods of slope near base of 
cliff, New Prospect (3/43). Known from a few local stations in the province. 

Carex Mackenziei Krecz. Wet quaking areas of salt marsh, Five Islands, 
Colchester County (4951); mucky area of salt marsh, Advocate, Cumberland 
County (5103). The distribution of this species is poorly known, but as 
Roland (1947) remarks, it is "probably general." 

Carex pedunculata Muhl. Rich hardwood slope about one mile east of 
Refugee Cove (5190). Sterile plants of this species were noted on Moose 
Island, Colchester County, but were not collected. This species was pre- 
viously reported from only two stations on North Mountain: Cape Blomidon, 
Kings County and north of Annapolis, Annapolis County. Another collection: 
from dryish open woods, top of North Mountain, Arlington, Kings County 
(D. S. Erskine 906). A collection of J. S. Erskine indicates its presence in 
Hants County: swamp, Oulton's Ridge, near Windsor (J. S. Erskine, June 8, 
1947). 

C. atratiformis Britt. Local on moist cliff facing waterfall, McAlese Brook, 
New Prospect (3226). Known only from northern Cape Breton Island, 
the above collection marks its first report from peninsular Nova Scotia. 

C. capillaris L. var capillaris. Tiny plants forming rounded cushions 
on seepy exposed slope at cliff-top, Cape d'Or (Schofield & Bentley 4804, 
Schofield 5168). "Var. major Blytt is known from a number of stations in 
northern Cape Breton Island. The typical variety is readily distinguished 
from this by its possession of darker green leaves, shorter stature and in its 
formation of very dense tussocks rather than single erect clumps. New to 
Nova Scotia. 

C. comosa Boott. Roadside behind dyke, Advocate (5155); abundant 
clumps in swamp, Truemanville (4792). Reported only from the Annapolis 
Valley, the above collections indicate its presence in north-central Nova 
Scotia. 

C. Tuckermanii Boott. Local in meadow swale, Wallace River, Wentworth 
(56352). Known from Sweet’s Corner, Hants County (Smith & Erskine, 
1954). 

Juncus Vaseyt Engelm. Abundant cespitose clumps in cranberry bog, 
Linden (5400). New to Nova Scotia; this fills in a range gap, for it is known 
from adjacent New Brunswick. 

Luzula parviflora (Ehrh.) Desv., var. melanocarpa (Michx.) Buchenau. 
Wet wooded margin of McGahey Brook (Schofield & Bentley 4783); damp 
margin of Mill Brook (Schofield & Bentley 4825); rare on banks of Soldier 
Brook (5202). Reported once from peninsular Nova Scotia (Erskine, 1951), 
this from Three Sisters, Cumberland County. The above collections show 
that it is widespread (but never abundant) in the Cape Chignecto area. 

Trillium erectum L., forma albiflorum R. Hoffm. Occasional among typical 
plants, alder thicket, North Side, Isle Haute (J. S. Erskine & W. B. Schofield 
JSE 53.041). 'This form was previously reported from North Mountain, 
Annapolis County (Roland, 1947). 

Malaxis brachypoda (Gray) Fern. Rare in wet area beside trickle near 


1955] Schofield,— The Flora of Nova Scotia 307 


Indian Flats, Isle Haute (3773). Previously unknown from the province, 
the above collection fills in a gap in the range of this species, found locally 
in the neighbouring provinces and states. 

Liparis Loeselii (L.) Richard. Rare in damp area in field near road, Isle 
Haute (3830); springy area, roadside near Folly Lake (3981); abundant in 
railroad ditch, Pineo Lake, Conn's Mills (2462). Its presence in the northern 
part of the province had not previously been suspected. 

Geocaulon lividum (Richards.) Fern. Among heaths of bog, Spicer's Cove 
(Schofield & Bentley 4838). This is the second collection from peninsular 
Nova Scotia. A collection from among heaths of a moist heath bog, Auburn, 
Kings County (3097) is very close to the previously known station at King- 
ston. The species is widespread, but local, in Cape Breton Island. 

Polygonum hydropiperoides Michx., var. hydropiperoides. In water of 
River Hebert, near south end of Newville Lake (3559). This species was 
previously known only from south-western Nova Scotia. 

Stellaria humifusa Rottb. Brackish marsh near Cape d'Or (3948); forming 
mats near upper part of salt marsh, Advocate (5125). Previously reported 
as mainly from eastern Nova Scotia and Cape Breton Island, the above 
collections indicate its presence in the north-central portion of the province. 
A collection from the margin of a salt marsh, Five Islands, Colchester County 
(4950) indicates its presence in the adjacent county as well. 

Montia lamprosperma Cham. Abundant on cold trickle margin, upper 
salt marsh, east side of Burke Brook, Advocate (3618). This species is known 
from three other widely separated stations (Brier Island, Digby Co.; Port 
Hawkesbury, Inverness Co.; and Northwest Arm, Halifax Co.). The above 
collection falls within the expected range of the species. 

Ceratophyllum demersum L. On sludge-bottomed margin of Newville 
Lake (3607); in water of River Hebert, Newville (Schofield & Webster 5645) ; 
in wrack of Pineo Lake, Conn’s Mills (5463). Reported from two other 
stations in the province, both in Kings County. 

Draba arabisans Michx. Abundant in crevices of high cliff facing road, 
New Prospect (3150); cliff crevices, South Side, Isle Haute (J. S. Erskine & 
W. B. Schofield JSE 53.035, Schofield 3696); rare in cliff crevices and on ex- 
posed cliff top, Cape d’Or (Schofield & Bentley 4805) ; small moist rock outcrop 
on rich hardwood slope, one mile east of Refugee Cove (5199). Hitherto 
known very locally from Cape Blomidon, Kings County and from Cape Breton 
Island. It is relatively common on the cliffs of Isle Haute. 

Cardamine parviflora L., var. arenicola (Britt.) Schulz. Among rocks 
near boat-house, East End, Isle Haute (3659); talus on cliff, North Side, 
Pigeon Point, Isle Haute (3786); seepy area of gully on cliff, Cape d’Or (6182). 
This taxon is doubtless much commoner than was suspected by the single 
reported station near Halifax (Roland, 1947). The following collections help 
to support this suggestion: very rare under damp shade of overhanging 
boulder, near P. Jack Cove, Brier Island, Digby County (1625); abundant 
in moist humus pockets of boulder talus, Cape Blomidon, Kings County 
(3084). 

Arabis hirsuta (L.) Scop., var. pycnocarpa (M. Hopkins) Rollins. Small 
moist outcrop on rich hardwood slope about one mile east of Refugee Cove 
(5200). This species has been reported from Indian Brook, Victoria County 
(Smith & Erskine 1954). The following collection indicates its presence in 
Colchester County: talus slope, north-west side, Moose Island (4962). At 
the latter locality it was fairly abundant. 


308 Rhodora [VoL. 57 


Arabis Drummondii Gray. Talus slopes near Western Slope, Isle Haute 
(J. S. Erskine & W. B. Schofield JSE 53.036, Schofield 3692); one plant on 
talus slope, central North Side, Isle Haute (3839). Another species known 
previously from Cape Breton Island and Cape Blomidon, Kings County. 

Sarracenia purpurea L., forma heterophylla (Eaton) Fern. Peat bog, 
Spicer’s Cove (Schofield & Bentley 4834). An interesting colour form known 
from a few local stations (Smith & Schofield, 1952). 

Saxifraga Aizoón Jacq., var. neogaea Butters. Locally abundant on shel- 
tered cliff shelves, Cape d'Or (Schofield & Bentley 4802, Schofield 5164). 
Previously known from Cape Breton Island and Cape Blomidon, Kings 
County. 

Potentilla fruticosa L. Exposed cliff-top headlands, Cape d'Or (Schofield 
& Bentley 4811); cliff crevices above sea-stack near Refugee Cove (Schofield 
& Bentley 4819). Common at both the extreme northern and south-western 
ends of the province, the above collections note its local abundance in the 
north-central portion. 

Astragalus Robbinsii (Oakes) Gray, var. Robbinsii. Depressed clumps on 
exposed cliff headlands, Cape d’Or (Schofield & Bentley 4800); depressed 
tussocks on talus above sea stack near Refugee Cove (Schofield & Bentley 
4818). The treatment of Barneby in the New Britton & Brown has been 
followed. The above specimens differ only superficially from the description 
in the above-mentioned manual, and have therefore been included under 
the typical variety. These collections are of particular interest since they 
represent the rediscovery of a taxon thought to be extinct from its type area 
on "dry calcareous ledges, Winooski R., Vt." (Fernald, 1950B). It is rela- 
tively common and easily accessible at Cape d’Or. New to Canada. 

Gzylropis johannensis Fern. Frequent in cliff crevices and on exposed 
cliff headlands, Cape d'Or (Schofield & Bentley 4799, Schofield 5183). This 
species has been known for many years from remote St. Paul Island, Victoria 
County. The above collection marks its first report from peninsular Nova 
Scotia. 

Geranium  Bicknellii (Britt.) Fern. Abundant on talus overgrown with 
poison ivy, base of cliff, New Prospect (3134). This species has been reported 
from very few localities. 

Impatiens pallida Nutt. Luxuriant on slope, North Side, Isle Haute 
(3828). A rather uncommon species in the province, the above collection 
indicates its local occurrence in Cumberland County. 

Elatine minima (Nutt.) Fisch. & Meyer. Occasional in shallow water 
of Leak Lake margin (3396); abundant on mucky margin of Lake Killarney 
(4180). This species appears to be widely distributed in Nova Scotia, but has 
been overlooked (cf. Smith & Schofield, 1952). 

Myriophyllum tenellum Bigel. Abundant in marginal water of Leak Lake 
(3482); abundant in marginal water of Newville Lake (3601). Previously 
unknown from the north-central counties, but now found to be relatively 
frequent in lakes throughout the province. 

Conioselinum chinense (L.) BSP. Seen, but not collected, on the banks 
of West Moose River; dry headland, Cape d'Or (Schofield & Bentley 4810). 
A rather rare species in the province, previously unknown from the north- 
central counties. 

Bartonia paniculata (Michx.) Robinson, var. iodandra (Robinson) Fern. 
Damp margin of swamp, west end of Dewar’s Lake (4048). The plants 
were unusually etiolated, possibly due to their submergence during at least 


1955] Schofield,—The Flora of Nova Scotia 309 


part of their growth period. This collection notes the appearance of this 
taxon in the north-central portion of the province, where it has been unknown. 

Lindernia dubia (L.) Pennell. Damp gravel pit near Shinimecas Bridge 
(4218) (these tiny plants—up to 3 cm.—were growing among Ludwigia 
palustris and were flowering cleistogamously); damp, muddy bank of River 
Philip, near Oxford (5371). Known from only two other localities in the 
province. 

Littorella americana Fern. In marginal shallows of Folly Lake (3999). 
All plants were sterile. This marks its first report from the north-central 
portion of Nova Scotia and its second collection from the peninsula. It is 
also of local occurrence in Cape Breton Island (cf. Smith & Schofield, 1952). 

Galium boreale L., var. intermedium DC. A single colony on hill-top 
pasture, New Prospect (3262); dry field, Cross Roads (Schofield & Bentley 
4700). Known previously from Cape Blomidon, Kings County. 

Campanula aparinoides Pursh. Abundant on banks of Parrsboro River 
(3506); common in moist area near Frog Pond, Isle Haute (3722). This 
species appears to be more common than was previously suspected (cf. also 
Smith & Erskine, 1954). 

Lobelia spicata Lam. Locally abundant in dry field near shore, Linden 
(5404). Previously known from Blomidon, Kings County. 

Megalodonta Beckii (Torr.) Greene. In wrack of Mattatall Lake (3998). 
This appears to be the third report of this species from the province.—PERRY 
BIOLOGICAL LABORATORIES, ACADIA UNIVERSITY, WOLFVILLE, NOVA SCOTIA. 


LITERATURE CITED 


BARNEBY, R. C. in Gleason, Henry A. 1952. Astragalus in the New Britton & 
Brown Illustrated Flora of the Northeastern United States and Adjacent 
Canada, Vol. 2, N. Y. Botanical Garden. 

Dorr, W. G. & A. E. RoraANp. 1942. Grasses of Nova Scotia. Trans. 
N. S. Inst. Sci. 20: 179-288. 

ERsKINE, D. S. 1951. Species newly or rarely reported from Nova Scotia 
and Cape Breton Island. Rnopona 53: 264-270. 

ERSKINE, J. S. 1953. Additions and extensions to the Flora of Nova Scotia. 
RHODORA 55: 17-20. 

FERNALD, M. L. 1921. The Gray Herbarium Expedition to Nova Scotia, 
1920. RnuoponA 23: 89-111, 130-152, 153-171, 184-195, 223-240, 
257-278, 284-301. 

FERNALD, M. L. 1922. Notes on the Flora of Western Nova Scotia. HRHo- 
DORA 24: 157-164, 165-180, 201—208. 

FERNALD, M. L. 1950A. Three additions to the Flora of Nova Scotia. 
Ruopona 52: 18-19. 

FERNALD, M. L. 1950B. Gray's Manual of Botany, Eighth Edition, 
American Book Company, New York. 

Lawson, G. 1884. Notice of New and Rare Plants. Trans. N. S. Inst. 
Sci. 6: 68-75 

Perry, L. M. 1931. Vascular flora of St. Paul Island, Nova Scotia. Rno- 
DORA 33: 105-126. 

Ronawp, A. E. 1938. Notes on the flora of Cape Breton Island. Can. 
Field Nat. 52: 104-106. 

RoLANp, A. E. 1947. Flora of Nova Scotia. Trans. N. S. Inst. Sci. 21: 
95-642. 


310 Rhodora [VoL. 57 


RornaAwp, A. E. & W. G. Dore. 1942. Notes on the Flora of Nova Scotia 
III. RHODORA 44: 334—338. 

Scuorretp, W. B. & E. C. Surrg.. 1953. Contributions to the Flora of 
Nova Scotia III: Some interesting white forms. Can. Field Nat. 67: 93- 
94. 

SurrH, E. C. « W. B. Scuoriretp. 1952. Contributions to the Flora of 
Nova Scotia I & II. Ruopora 54: 220-228. 

SurrH, E. C. & J. S. EnskiNE. 1954. Contributions to the Flora of Nova 
Scotia IV. RHODORA 56: 242-252. 


PLANTS NEW TO MISSOURI 


E. J. PALMER AND J. A. STEYERMARK 


SINCE the publication in 1935 of the Catalogue of the Flowering 
Plants of Missouri (Ann. Mo. Bot. Gard. 22: 375-758), work 
has continued on the further botanical exploration of the state 
of Missouri and a large number of plants, including several 
genera not previously known in the state, have been added. 
The junior author of the Catalogue and of this paper has been 
particularly active and has made many collecting trips into 
nearly all parts of the state, resulting in some surprising dis- 
coveries. The senior author, since returning to Missouri in 1948, 
has devoted what time he could to an intensive exploration 
of several of the southwestern counties, with a few excursions 
into other sections. 

The resulting new discoveries have been reported in Rhodora 
from time to time in several short papers contributed by the 
junior author and one by the senior author, as well as in a joint 
paper on new fern discoveries in the American Fern Journal 
42: 61-66. 1952. William B. Drew also reported the dis- 
covery of four new records for the state (RHODORA 44: 248. 
1942), George B. Van Schaack reported Calamagrostis insparata 
new to the state (RHoporA, 56: 43. 1954), and C. L. Kucera 
published his findings of Lyonia ligustrina in Missouri (RHODORA, 
55: 155. 1953). The large number of additional plants now 
known in the state, as well as many changes in nomenclature 
and the interpretation of species necessary to bring it into 
conformity with the eighth edition of Gray's Manual, have 
made a revision of the Catalogue desirable, and it is hoped 
that such a revision can be published in the near future. The 
present paper is a further report of progress and a review of 


1955] Palmer and Steyermark,—Plants New to Missouri 311 


what has been done up to this time in increasing our knowledge 
of the state flora. 


Hysrrix PATULA Moench var. BiGELoviaANA (Fern. Deam. This 
variety has not been reported previously from Missouri. It was found 
on wooded north-facing ravine slopes tributary to Mill Creek ditch of 
Missouri River valley, T59N, R38W, sect. 14, 4 mi. south of Oregon, 
Holt Co., July 20, 1952, Steyermark 73829. 

LOLIUM MULTIFLORUM Lam. var. pimMinutuM Mutel. Not previously 
reported from Missouri, this variety was found in open grassy woodland 
near lake, Lewis and Clark State Park, T55N, R37W, sect. 33 and 28, 
16-1 mi. southwest of Armour, Buchanan Co., August 20, 1950, Steyermark 
70116. 

MUHLENBERGIA MEXICANA (L.) Trin. forma AMBiGUA (Torr.) Fern. 
The record for this form is based uopn the following specimen: gravelly 
open places along South Fork of Spring Creek along highway A, T23N, 
RSW, south V4 sect. 33, 8 mi. south of West Plains, Howell Co., September 
3, 1949, Steyermark 69076. 

PANICUM CONSANGUINEUM Kunth. This species, primarily of Atlantic 
and Gulf Coastal Plain distribution from northern Florida to eastern 
Texas north to southeastern Virginia, Tennessee, and Arkansas, has not 
been reported previously from Missouri. It is based upon the following 
collection: wooded oak-hickory sandy knoll, T26N, R14E, sect. 34, 444 
mi. southeast of Sikeston, New Madrid Co., May 18, 1950, Steyermark 
69667. This station is in the southeastern corner of Missouri, where 
occur many species the affinities of which align them with the flora of the 
Mississippi Embayment of the Gulf Coastal Plain. 

PasPALUM LAEVE Michx. (typical). The typical form of this species 
with the spikelets 2-2.5 mm. broad and with the leaf-blades and sheaths 
glabrous or nearly so has not been reported previously from Missouri. 
It is now known by the following collections: marshy springy ground at 
base of slopes of ravines bordering east-facing escarpment on Crowley 
Ridge, just south of Stephens Cemetery, T24N, RIOE, sect. 30, 115 mi. 
south of Pyletown, Stoddard Co., August 29, 1948, Steyermark 66187; 
knolls and depressions, T22N, RAE, sect. 35, 4 mi. south of Naylor, 
Ripley Co., October 20, 1948, Steyermark 66943; wet sedgy swales, wet 
woods, and swampy thickets along highway 25, 3 mi. west of Arbor, 
Cape Girardeau Co., September 22, 1946, Steyermark 64158. 

SCIRPUS POLYPHYLLUS Vahl. This addition to the state flora is based 
upon the following collections: swampy meadow along Bee Fork, on 
property of Joe Goforth, T32N, R2W, sect. 23, 414-434 mi. east of 
Bunker, Reynolds Co., July 7, 1951, Steyermark 72041; swampy meadow 
along Bee Fork, T32N, R2W, sect. 22, on property of Mr. Reese, 4 mi. 
southeast of Bunker, Reynolds Co., July 7, 1951, Steyermark 72002. 

Carex ABDITA Bickn. Whether or not this taxon can be maintained 
as distinet from C. umbellata Schkuhr is questionable. It perhaps should 
be considered only as a variety of C. umbellata. On the basis of the 


312 Rhodora [VoL. 57 


characters used for separating the two taxa, the following Missouri 
collection, with perigynia 2.2-2.8 mm. long with short beaks 0.5-0.6 mm. 
long and acute pistillate scales below becoming more attenuate in the 
upper ones, may be cited: rocky ground east of highway K, T33N, R33W, 
sect. 27, 2 mi. northwest of Liberal, Barton Co., April 16, 1949, Steyermark 
67176. 

Carex Buxpaumit Wahlenb. forma piLvTIoR Kükenth. This form, in 
which the pistillate scales are whitish or pale brown, has not been reported 
previously from the state. It was found in a ealeareous swampy meadow 
associated with other species of which the ranges in the United States are 
northward or northeastward: swampy meadow along West Fork of Black 
River, on property of D. C. Miner, T33N, R3W, sect. 23 and NE sect. 
26, 3-314 mi. northwest of Greeley, Reynolds Co., July 6, 1951, Steyer- 
mark 71984. 

In this collection some of the terminal spikes are staminate at the base 
as well as at the summit, while others are staminate only in the upper 
half. Dr. F. J. Hermann, who has kindly verified the junior author's 
determination, writes of the collection as follows: “The scales of Carex 
Buxbaumii normally tend to bleach out with age, as they do in many 
other species found in exposed habitats, so that typical Buxbaumii must 
often become f. dilutior late in the season. In the sheet you sent me both 
terminal spikes are staminate at the base as well as at the apex, the peri- 
gynia being restricted to the central portion. However, I doubt that 
sexual aberration in the terminal spike of this species is anymore deserving 
of recognition than seale-color. I have f. heterostachya and the typical 
form on the same plant in a collection from northern Michigan." 

CAREX DEBILIS Michx. (typical). The typical form of this species has 
not been reported previously from Missouri. It is known from the fol- 
lowing collection: depressions in swamp dominated by Lindera melissae- 
folium, surrounded by sandy, wooded and cultivated knolls, T22N, R4E, 
SE part of sect. 35, 484 miles south of Naylor, Ripley Co., May 28, 1951, 
Steyermark 71228, In this collection the perigynia are 8-8.5 mm. long, 
but the pistillate scales are greenish-white. 

Carex DbiGITALIS Willd. var. Macropopa Fern. This variety, previ- 
ously unknown from Missouri, is represented by the following collections: 
north-facing slopes along Crooked Creek, T31N, R9E, sect. 16, 2-3 mi. 
southeast of Bessville, Bollinger Co., May 17, 1950, Steyermark 69613a; 
rich, north-facing ravine along tributary of Dry Fork of Charrette Creek, 
T46N, R2W, sect. 19, 4 mi. northwest of Hopewell, 7 mi. southwest of 
Warrenton, Warren Co., June 8, 1952, Steyermark 73327. 

CAREX STRICTA Lam. var. srricrror (Dewey) Carey. Dr. F. J. 
Hermann has kindly verified the junior author’s determination of this 
species, neither the typical form of the species nor any of its varieties 
having been previously identified correctly from the state. The following 
collections are given: swampy meadow along Bee Fork, T32N, R2W, sect. 
23, 4% mi. southeast of Bunker, Reynolds Co., May 29, 1951, Steyermark 
71303; swampy meadow along Bee Fork, on property of Joe Goforth, T32N, 


1955] Palmer and Steyermark, Plants New to Missouri — 313 


R2W, sect. 23, 415-434 mi. east of Bunker, July 7 , 1951 , Steyermark 72032, 
"in dense colonies; leaves dark green; perigynia appressed.” 

POPULUS NIGRA L. var. rTALICA. Muenchh. Several small specimens 
escaped from cultivation and spreading form the basis for the following 
collection: open ground, border of woods, 4 mi. northwest of Webb City, 
Jasper Co., Sept. 27, 1952, Palmer 55194. 

UrMus PUMILA L. This species, now becoming common in cultivation, 
seems to be well established in the rocky valley of Hickory Creek, in 
Newton County, where a number of specimens of different sizes were 
seen. At the McDonald County station only one tree was seen. Rocky 
waste ground along Hickory Creek, Neosho, Newton Co., Oct. 19, 1953, 
Palmer 57208; same locality, Nov. 3, 1953; same locality (in bloom), 
March 6, 1954, Palmer 57218; near Beaver Brook Spring, Anderson, Mc 
Donald Co., Oet. 19, 1954, Palmer 59240. 

PILEA PUMILA (L.) Gray. (typical). The typical form of this species 
is based upon the following collection: along Bookout Branch and 
ravines tributary to Spring Creek, T64N, RISW, SW 14 sect. 21 and 
NE 1% sect. 28, 44% mi. northeast of Green City, Sullivan Co., August 
25, 1950, Steyermark 70149. 

ad UM viscosum L. forma APETALUM (Dumort.) Mert. & Koch. 
'This form, not previously reported from the state, is represented by the 
following collection: open grassy ground on top of ridge in sect. 24, cherty 
limestone upper slopes above northwest-facing wooded bluffs along Flat 
Creek, T45N, R21W, 215 mi. south of Sedalia, Pettis Co., May 20, 1949, 
Steyermark 67973. 

NYMPHAEA ODORATA Ait. forma Rosea Guillon. This form of the 
water-lily, in which the petals are roseate, was found well established in 
an artificial lake in northern Missouri. It has not been previously re- 
ported from the state. According to Mr. Raymond Buster of Ethel, 
Missouri, the plants were purchased from Vaughan's Seed Store in 
Chicago and planted in the lake ten to fifteen years ago. These plants 
are established, growing in a colony in which the flowers open in the 
morning and close about 2 p.m. The petals, on several flowers examined, 
number about 28 and vary from obtuse to rounded at the apex; there are 
4 sepals present. The flowers are fragrant, the petioles are coiled at the 
base, and the leaves are all floating and purple beneath. The data for 
this collection are as follows: near shore of Ethel Lake, T59N, R17W, 8 
part of sect. 25 and N part of sect. 36, 34 mi. west of Ethel, Macon Co., 
Sept. 14, 1954, Steyermark 77311. 

DRABA VERNA L. var. BOERHAAVI Van Hall. This variety, not previ- 
ously reported from the state, is based upon the following collection: 
grassy, rocky, open places along road bordering limestone wooded banks 
along creek along highway 21 and 49, just northeast of Centerville, 
Reynolds Co., March 30, 1949, Steyermark 67107. 

SEDUM SARMENTOSUM Bunge. This is an evident escape from cultiva- 
tion and it may have been originally planted in the vicinity, but is now 
well established and locally abundant in a wild spot near one of the spring 


314 Rhodora [Vor. 57 


heads. The data for the collections cited are: on moist mossy rocks, 
Haddock Spring, Newton Co., Nov. 3, 1953, Palmer 57195; same locality, 
May 19, 1954, Palmer 57624. 

SPIRAEA PRUNIFOLIA Sieb. and Zucc. This species, not recorded in the 
Sth edition of Gray's Manual, has become thoroughly established in at 
least two places in Missouri, and is based upon the following collections: 
escaped from old cemetery in woods on ridge top and established as a shrub 
forming thickets on north-facing, steep, wooded bluffs with limestone at 
top along Missouri River, T44N, R10W, sect. 16, just north and north- 
west of Osage City, Cole Co., May 20, 1950, Steyermark 69747; planted 
at edge of cemetery, but escaping and thoroughly established along fence 
row, above ravines bordering Wyaconda River, T62N, R6W, sect. 18, 7 
mi. northwest of Canton, Lewis Co., July 27, 1952, Steyermark 74161. 

PYRUS MELANOCARPA (Michx.) Willd. This remarkable addition to the 
state flora of an essentially boreal and northerly distributed species was 
discovered, not in the northern section of Missouri, where it might have 
been expected, but in the southeastern corner of the state ordinarily 
occupied by species of an essentially Mississippi Embayment-Gulf and 
Atlantic Coastal Plain distribution. The data for the collection are as 
follows: in midst of dense alder thickets along spring branch east and 
southeast of Pleasant Valley Church, east of spring-fed creek, along flood 
plain and Crowley Ridge junction, on property of E. Walker, T25N, 
R111, sect. 6, 314 mi. southeast of Bloomfield, Stoddard Co., August 20, 
1954, Steyermark 76825, “plants 3 feet tall." 

RUBUS ALLEGHENIENSIS Porter var. pLAUSUS Bailey. Not previously 
reported for Missouri, this variety is based upon the following collection: 
in open ground of valley between ravines tributary to wooded slopes 
along north side of North River, *Miller's Hills,’ T59N, R9W, west 
part of sect. 2, 215 mi. southeast of Burksville, Shelby Co., June 5, 1951, 
Steyermark 71705. 

RosA MICRANTHA Sm. This species has not been recorded previously 
from the state. It is based upon the following collection: fallow pasture 
above wooded slopes following ravine along tributary to Salt River, 
TOON, R13W, NE 14 sect. 1 and sect. 36, 114 mi. northwest of Locust 
Hill, Knox Co., Sept. 18, 1950, Steyermark 70672. 

GERANIUM DISSECTUM L. This and the following record of Geranium 
molle were originally brought to the attention of the junior author by 
Mr. Oscar Petersen, poet-naturalist and amateur botanist of 8t. Louis Co. 
Both species are of spontaneous occurrence and have become well estab- 
lished on his property since 1951. The data for this collection are as 
follows: on property of Mr. Oscar Petersen, 267 Elm Ave., Glendale, St. 
Louis Co., June 7, 1952, Steyermark 73314. 

GERANIUM MOLLE L. Collected on property of Mr. Oscar Petersen, 
267 Elm Ave., Glendale, St. Louis Co., June 7, 1952, Steyermark 73315. 

EUPHORBIA GEYERI Engelm. This species, not previously known from 
a definite Missouri collection, was found near the Iowa line and is based 
upon the following data: natural sandy prairie on slopes of old river ter- 


1955] Palmer and Steyermark,— Plants New to Missouri — 315 


race on beach paralleling Des Moines River, T65N, R6W, sect. 9, 112 mi. 
southeast of St. Francisville, Clark Co., August 13, 1949, Steyermark 
68876, “plant prostrate; leaves pale beneath." 

ACER RUBRUM L. forma TOMENTOSUM (Desf.) Dansereau. This form, 
not previously reported from Missouri, is based upon the following col- 
lection: cherty upper slopes of ravine tributary to Little Niangua River, 
T38N, R18W, sect. 4, 444 mi. southeast of Barnumtown, Camden Co., 
July 6, 1952, Steyermark 73743. 

JUSSIAEA URUGUAYENSIS Camb. This South American plant has be- 
come quite abundant and grows in large colonies for some distance along 
the creek in association with several other introduced plants, including 
Phalaris arundinacea L., Vicia Cracca L., Myriophyllum brasiliense Camb., 
and Myosotis scorpioides L. The garden Forget-me-not is also very 
abundant along rocky margins for some distance up and down the creek. 
It had previously been recorded from the vicinity of St. Louis. The data 
for the Jussiaea collection are as follows: in shallow running water of 
Hickory Creek, Neosho, Newton Co., Oct. 12, 1953, Palmer 57068. 

GAYLUSSACIA BACCATA (Wang. K. Koch. Another remarkable dis- 
covery, this species can now be definitely added to the known flora of the 
state, where it was found in an unglaciated section of east-central 
Missouri north of the Missouri River, where Trillium nivale also occurs. 
It was growing with Vaccinium vacillans and was almost mistaken 
in the field for that species. The data for the collection are as follows: 
cherty upper slopes above north-facing limestone bluffs along West Fork 
of Cuivre River, T50N, R3W, sect. 32, 814 mi. northeast of Bellflower, 
Montgomery Co., Sept. 16, 1954, Steyermark 77419. 

DopEcATHEON Mrapia L. var. BRACHYCARPUM (Small) Fassett forma 
PALLIDUM Fassett. This form of the variety, without red at the base of 
the leaves, is based upon the following collection: north- and northeast- 
facing, limestone, wooded, small bluffs along Camp Ground Creek, 
T40N, R11W, sect. 6, 6 mi. northwest of Tavern, Maries Co., May 19. 
1950, Steyermark 69701. 

BARTONIA PANICULATA (Michx.) Muhl. (typical). This surprising dis- 
covery was made at the head of a sandy spring branch where [lex opaca 
and Fraxinus tomentosa occurred at the base of Crowley Ridge in the 
southeastern section of Missouri, noted for its prominence of species 
following a Mississippi Embayment-Atlantic and Gulf Coastal Plain 
distributional pattern. The locality was not far from the alder thickets 
where Pyrus melanocarpa (reported above) was discovered on the same 
day. Inthe Missouri collection the corolla lobes are creamy-white above, 
1 mm. wide, the anthers are yellow, and the filaments white below and 
lavender above. The data for this collection are as follows: on mossy 
ground at head of sandy spring branch, at base of sandy ravine near 
junction of Crowley Ridge and lowland, T25N, R11E, NW 14 sect. 6, 
31% mi. southeast of Bloomfield, on property of Mr. Martin, Stoddard 
Co., August 20, 1954, Steyermark 76784. 

CONVOLVULUS sEPIUM L. var. REPENS (L.) Gray. Although Missouri 


316 Rhodora [Vor. 57 


is in the general range stated in the 8th edition of Gray’s Manual for this 
variety, it has not been recorded previously for the state. The record 
for its occurrence is based upon the following collection: swales in bottoms 
of Missouri River valley, T59N, R38W, SW sect. 14 and N part of sect. 
23, 4-415 mi. south of Oregon, Holt Co., July 20, 1952, Steyermark 73846. 

PHLOX DIVARICATA L, var. LaPHAMrI Wood, forma candida Palmer & 
Steyermark, f. nov. A typo differt floribus albis.—Low, rich, alluvial 
woods, 1 mile west of Nashville, Barton County, Missouri, April 30, 1953, 
Ernest J. Palmer 55411, noLoryrE, in Palmer Herb., isotype in Herb. 
Chi. Nat. Hist. Mus.; terrace along alluvial bottoms of Grand River along 
route 36, 4 mi. southwest of Chillicothe, Livingston Co., May 1, 1950, 
Julian A. Steyermark 69584, PARATYPE, in Herb. Chi. Nat. Hist. Mus. 

It is necessary to give a name to the white-flowered form of Phlox 
divaricata var. Laphamii. Dr. Wherry lists this white-flowered form in 
Bartonia 12: 34. 1930, but as stated by him in a recent communication 
with the present authors, his reference to the albino form, without a 
diagnosis or type specimen indicated, is not validly published. 

LrrHosPERMUM CANESCENS (Michx.) Lehm. forma pallidum Palmer & 
Steyermark, f. nov. A typo recedit corollis pallido-luteis.—Along south- 
east side of highway 54, 14 mi. southwest of junction with highway 154, 
T53N, R4W, W part of sect. 25, 4 mi. WSW of Curryville, Pike Co., 
Missouri, April 30, 1952, Julian A. Steyermark 73223, HOLOTYPE, in Herb. 
Chi. Nat. Hist. Mus., isotype in Mo. Bot. Gard. Herb. 

This form of the common Lithospermum canescens has cream-colored to 
pale yellow, instead of orange flowers. About a dozen plants were found 
growing scattered among the orange-flowered ones. Individual plants 
transplanted to the garden of the junior author have maintained the pale 
color of the corolla. 

LITHOSPERMUM CAROLINIENSE (Walt. MaeMill. I. M. Johnston 
(Journ. Arn. Arb. 33: 339-340. 1952) combines L. croceum Fernald with 
L. caroliniense, noting that (p. 340) “the floral differences used by Fernald 
to distinguish L. croceum from L. caroliniense are those which distinguish 
the short- and long-styled flowers of the species." We are in agreement 
with Dr. Johnston in combining the two species, as the differences between 
the two taxa, as given in the 8th edition of Gray's Manual, intergrade. 
If, however, L. caroliniense is maintained as distinct, the following Missouri 
collection should be cited: sandy open banks of formerly original sandy 
prairie, along west side of road, T25N, R14E, sect. 10, 614 mi. southeast 
of Sikeston, New Madrid Co., May 18, 1950, Steyermark 69674. 

VERBENA CANADENSIS (L.) Britt. forma candidissima (Haage & Schmidt) 
Palmer & Steyermark, comb. nov. V. canadensis var. candidissima, in 
Royal Hort. Soe. Dict. Gardening 4: 2210. 1951. 

The white-flowered form of this species has not been recorded previously 
from Missouri. It is based upon the following collection: exposed lime- 
stone bluff along road, along Long Creek, just north of Oasis, T22N, 
R22 W, sect. 10, 11, and 3, Taney Co., April 28, 1949, Cora Steyermark 
s.n. In this collection the corolla lobes are whitish and the corolla tube 
is whitish with pale lilac suffused throughout. 


1955] Palmer and Steyermark,—Plants New to Missouri — 317 


VERONICA DIDYMA Ten. This species has not been reported previously 
from Missouri. It is based upon the following collection: near gas station 
one mile north of Kohler City, Jefferson Co., April 20, 1952, H. E. Ahles 
5848. 

GALIUM TRIFLORUM Michx. X G. CIRCAEZANS var. HYPOMALACUM 
Fern. The data for this collection are as follows: in wooded ravines 
tributary to Thorp Branch, T59N, RSW, NE sect. 12, 414 mi. southeast 
of Oregon, Holt Co., July 20, 1952, Steyermark 73804. 'The present 
collection, growing with G. circaezans (Steyermark 73804a), consists of 
plants having the leaves in 6's and elliptic-lanceolate as in G. triflorum, 
but with pubescent stems and the midrib of the lower leaf surface pilose 
as in G. circaezans var. hypomalacum. So far as known, this apparent 
natural hybrid has not been recorded previously in literature. 

HOUSTONIA PUsILLA Schoepf forma ALBIFLORA Standl. The form of 
the species with white corollas has not been recorded previously from 
Missouri. The following collection is representative: open ground in 
valley of Ottery Creek, along highway A, northeast of Redmondville, 
Iron Co., April 26, 1952, Steyermark 73101. In this collection the plants 
are of a much lighter green color than in typical purple-flowered H. 
pusilla (H. patens Ell.). The width and length of the white corollas vary 
considerably within a given colony of the plants. Other Missouri collec- 
tions which may be referred to this form are Steyermark 4599 from Pulaski 
Co. and 4592 from Phelps Co. Although the corollas in these last two 
collections vary from white to white with pale blue or lilac, they are gen- 
erally white throughout. 

SPECULARIA LAMPROSPERMA (McVaugh) Fern. X 5. LEPTOCARPA 
(Nutt.) Gray. In a colony of the two species named above, with S. 
lamprosperma predominating, were found a few plants with characters 
quite intermediate between the two and of evident hybrid origin. The 
data for this collection are: chert glades along Shoal Creek, 4 mi. south- 
west of Joplin, Newton Co., May 29, 1952, Palmer 54078. 

LoBELIA CARDINALIS L. forma Rosea St. John. Not previously re- 
corded from Missouri, this form of the cardinal flower was collected in a 
wet, calcareous meadow along Parker Branch of West Fork of Black 
River, T33N, R3W, west part of sect. 15, 44 mi. northwest of Marcoot, 
5 mi. northwest of Greeley, Reynolds Co., Sept. 24, 1951, Steyermark 
72729. In this collection the corolla tube is pink to rose-colored, the 
corolla lobes are pink on the outside, white on the inside, and the staminal 
tube 1s white. 

ASTER AZUREUS Lindl. forma raAEviCAULIS Fern. The following col- 
lection may be referred to this form, not previously reported from the 
state: upland roadside banks, T59N, R31W, sect. 4, 314 mi. southwest of 
Fairport, Dekalb Co., Sept. 27, 1951, Steyermark 72869. This collection 
represents an unusually floriferous specimen, which may actually be a 
hybrid between A. laevis and A. azureus, with both of which species the 
plants were growing. The upper surface of the leaves in this collection 
are scabrous as in A. azureus and the stems are glabrous as in A. laevis. 

ASTER CORDIFOLIUS L. var. MoRATUS Shinners. Not previously re- 


318 Rhodora [Vor. 57 


ported for the state, this variety is based upon the following collection: 
along base of La Motte sandstone bluffs along Terre Bleue Creek, T37N, 
R6E, south part of sect. 20 and north part of sect. 29, 2-214 mi. south 
of Thurman, 5-6 mi. northwest of Sprott, Ste. Genevieve Co., October 3, 
1950, Steyermark 71041. 

ASTER CORDIFOLIUS L. var. POLYCEPHALUS Porter. In the 8th edition 
of Gray's Manual, the range for this variety is given as “SW. Que. to 
Ind., s.to N.E. and Ga." In addition to a specimen from Boone County 
previously recorded in our catalogue, it may be eredited to Missouri on the 
basis of the following collection: limestone glade along highway 131, on 
south side of South Fork of Blackwater River, T46N, R28SW, NW 14 sect. 
26, 3 mi. north of Holden, Johnson Co., Sept. 25, 1951, Steyermark 72751. 

ASTER DUMOSUS L. var. STRICTIOR T. & G. Not previously reported 
from Missouri, this variety may be added to the state flora on the basis 
of the following collection: meadow along north side of highway 80, 4.9 
mi. southwest of West Plains, Howell Co., Sept. 25, 1949, Steyermark 
693306. 

ASTER OBLONGIFOLIUS Nutt. forma ROSEOLIGULATUS (Benke) Shinners. 
This form has not been recorded previously from the state. It is based 
upon the following collection: edge of limestone escarpment of SE-facing 
Chapel Bluff along Niangua River, T37N, RISW, sect. 26, 8-814 mi. 
southeast of Macks Creek, Camden Co., October 24, 1954, Steyermark 
78223. The rays in this collection varied from pink to rose-colored. 

ASTER VIMINEUS Lam. (typical). Although the var. subdumosus Wieg. 
is known from Missouri, the typical variety of the species has not previ- 
ously been reported from the state. It is based upon the following 
colleetion: alluvial lower part of north-facing limestone slopes along Salt 
River, Mark Twain State Park, T54N, RSW, sect. 9 and 16, 1-2 mi. 
southwest of Florida, Monroe Co., Sept. 25, 1948, Steyermark 66529. In 
this collection the ray florets vary from 16 to 20 in number, the lobes of 
the corollas of the disk florets are erect, about 2$ the length of the throat, 
and are 0.9-1 mm. long. 

HELIANTHUS ANNUUS L. var. NANUs fl. pl. Hort. This double-flowered 
variety with the stems averaging about 3 feet high is locally common in 
Holt Co., northwestern Missouri. It has not been previously reported 
for the state. The collection upon which it is based is as follows: common 
escape throughout this area in low ground in extensive swale, T62N, 
R40W, sect. 25, 334 mi. southeast of Craig, Holt Co., July 20, 1952, 
Steyermark 73788. The robust-stemmed (averaging 6 feet tall), double- 
flowered variety is referred to var. chrysanthemoides Cockerell (Am. Nat. 
49: 617. 1915), but the present collection is better referred to the dwarf 
type. 

LIATRIS SCABRA (Greene) K. Schum. This species has not been previ- 
ously reported from the state. It is based upon the following collection: 
ravine slopes tributary to river, along Little Black River, between Green- 
ville Ford and Pennington Ford, T24N, R3E, sect. 10, 15, 22, 23, 26, 24, 
and 25, 10-13 mi. northeast of Doniphan, Ripley Co., Sept. 1, 1946, 
Steyermark 63966, 


1955] Mohlenbrock,— Flora of Southern Illinois 319 


CICHORIUM INTYBUS L, forma ALBUM Neum. This form with the 
flowers entirely white has not been recorded previously from Missouri. 
It is based upon the following collection: along route 129, 2.3 mi. south 
of Green City, Sullivan Co., August 25, 1950, Steyermark 70127.—WEBB 
CITY, MISSOURI, CHICAGO NATURAL HISTORY MUSEUM. 


CONTRIBUTIONS TO THE FLORA OF SOUTHERN ILLINOIS.— 
Field work in southern Illinois the latter part of 1954 produced 
several species of plants new to Illinois. In addition, the 
ranges of other uncommon species have been extended by further 
collections. 

Areas of particular interest occur in Randolph County; here are 
found such rare species for Illinois as Pinus echinata, Asplenium 
bradleyi, Ranunculus harveyi, Talinum calycinum, Rhamnus 
caroliniana, Carex torta, Carex aquatilis var. altior, and Solidago 
buckleyi. 

Herbaria and their abbreviations for specimens cited in this 
paper are as follows: Southern Illinois University (SIU), Univer- 
sity of Ilinois (UI), Illinois State Museum (ISM), Illinois 
Natural History Survey (NHS), and that of the author (A). 


SPECIES NEW TO ILLINOIS 


CAREX DEBILIS Michx. The habitat in Illinois for this species is near 
the base of a densely shaded east-facing hillside. Its common associates 
at this station are Spigelia marilandica and Pedicularis canadensis. The 
species occurs in south-central and south-eastern Indiana, the nearest 
stations to ours being in Lawrence and Crawford Counties. COLLECTION 
DATA: rich woods near Cave-in-Rock State Park, Hardin County, June 
20, 1954, Mohlenbrock 4257 (A). 

Carex SWANIL (Fern.) Mackenz. The only report of this species from 
Illinois is based on a questionable specimen from Vermilion County, 
although it is not uncommon in western Indiana. It apparently is un- 
known from Missouri. Two plants at our station are growing at the base 
of a sweet gum (Liquidambar styraciflua). COLLECTION DATA: dry soil at 
the edge of a marsh, one mile north of Murphysboro, Jackson County, 
June 5, 1954, Mohlenbrock 3327. (X). 

TALINUM CALYCINUM Engelm. Although this species is known from 
eastern Missouri, it appears that the present collection represents the 
first from Illinois.2 A single plant was found growing along the edge of 
a sandstone bluff in Randolph County on August 15, 1954. A search for 
additional plants of this species a week later in the same area proved 

! Deam, CHARLES C. Flora of Indiana. 1940. 

2 Personal correspondence with Mr. Harry E. Ahles, University of Illinois Her- 
barium, 


320 Rhodora (Vou. 57 


successful as some one hundred specimens were seen, many in full flower. 
They were associated with /santhus brachiatus, Opuntia humifusa, and 
Polygonum tenue. This station in southwestern Illinois extends the range 
of the species to the east. COLLECTION DATA: thin soil on a dry sandstone 
bluff, “Castle Rock", near Leanderville, Randolph County, August 15, 
1954, Mohlenbrock 4550 (SIU, A). 

RANUNCULUS PARVIFLORUS L. This species, naturalized from Europe, 
is uncommon in the midwest. It is unreported from Indiana and is known 
in Missouri only from Butler and Dunklin Counties? It grows in an 
open wet place in a mesic woods. COLLECTION DATA: low woods, Saltpeter 
Cave, five miles south of Murphysboro, Jackson County, Mohlenbrock 
1936. 

PrinimNium CcosTrATUM (Ell) Raf. It had been thought that two 
species of Plilimnium occurred in Illinois—P. nuttallii (DC.) Britt. and 
P. capillaceum (Michx.) Raf.^ However, when the author was asked to 
collect specimens of these two species, the first Ptilimnium collected 
keyed out to P. costatum. Upon checking the herbarium specimens of 
Ptilimnium at Southern Illinois University, University of Illinois, Illinois 
State Museum, and the Illinois Natural History Survey (the last three 
by Mr. Harry Ahles), it was discovered that all sheets labelled P. capil- 
laceum were actually either P. costatum or P. nuttallii. In addition, some 
P. costatum specimens had been identified as Carum carvi. Further field 
work led only to the discovery of more P. costatum. So far, no authentic 
specimens of P. capillaceum have been located for Illinois. SPECIMENS 
EXAMINED: Union County: wet soil, Big Muddy River bottoms, July 29, 
1941, G. D. Fuller and R. Fisher 746 (UI, NHS, ISM); river bottom 
forest, Clear Creek, Sept. 12, 1940, Fuller 289. Jackson County: rich 
woods northeast of Howardton, Sept. 13, 1941, McCree 1197 (SIU, UI); 
Campbell Lake area near Elkville, July 17, 1941, McCree 929 (SIU, UI); 
wet soil, Carbondale, Sept. 13, 1941, G. D. Fuller and W. B. Welch 1197 
(ISM); six miles east of Elkville, low woods, MoAlenbrock 1912 (A). 
Pulaski County: without definite locality or date, Fricke. 

PTILIMNIUM NUTTALLII (DC.) Britt. Randolph County: roadside in 
low ground, two miles northwest of Sparta, July 13, 1950, G. S. Win- 
terringer 4848 (ISM). Jackson County: north of Makanda, August 2, 
1950, Bailey and Swayne 1114 (SIU). 

RuELLIA CAROLINIENSIS (Walt.) Steud. The discovery of this species 
in Illinois is not too surprising since it is found rather frequently in 
southern Indiana and Missouri. Its associates at our station include 
Scutellaria ovata and Monarda fistulosa. COLLECTION DATA: edge of woods, 
near Cave-in-Rock State Park, Hardin County, June 20, 1954, Mohlen- 
brock 4286 (A). 

VERONICA POLITA Fries. (Veronica didyma Tenore of Pennell?). "This 

3 PALMER, E. J. AND J. A. STEYERMARK. An Annotated Catalogue of the Flowering 
Plants of Missouri. Ann. Mo. Bot. Gard. Vol. 22. 1935. 

1 Jones, G. N. Flora of Illinois. 1950. 

* PENNELL, Francis W. The Scrophulariaceae of Eastern Temperate North 
America. 1935. 


1955] Mohlenbrock,— Flora of Southern Illinois 321 


species is adventive from Eurasia. In southern Illinois, it has been found 
on a semi-weedy lawn, growing with Veronica arvensis but flowering much 
earlier. COLLECTION DATA: lawn, Murphysboro, Jackson County, March 
10, 1955, Mohlenbrock 4984 (A). 


ADDITIONAL DISTRIBUTION RECORDS OF SOME ILLINOIS PLANTS 


LoPHOTOCARPUS CALYCINUS (Engelm.) J. G. Sm. forma Maximus (En- 
gelm.) Fern. The giant form of this species was found growing in a 
drainage ditch in the southwestern part of Murphysboro. The width of 
the leaves slightly surpassed three decimeters. The "typical" form is 
present in Lake Murphysboro. In addition to Jackson County, the 
species is known in Illinois from Pope, Piatt, Mason, Fulton, Peoria, and 
Henderson Counties. Only Pope and Jackson are in the southern part of 
the state. COLLECTION DATA: drainage ditch, Murphysboro, Jackson 
County, August 14, 1954, Mohlenbrock 4735 (A). 

PANICUM LANUGINOSUM Ell. var. IMpLicatuM (Seribn.) Fern. This 
taxon, found in dry soil at the edge of a woods in Jackson County, is 
generally more widespread to the north in Illinois. COLLECTION DATA: 
Giant City State Park, Jackson County, June 19, 1954, Mohlenbrock 
$206 (A). 

HEMICARPHA MICRANTHA (Vahl) Pax. Previous to its discovery in 
Jackson County, this tiny sedge was known in Illinois only from the 
northern and central counties (the most southern record being from Piatt 
County). This species was found at two stations along the Mississippi. 
COLLECTION DATA: in sand, Mississippi River cutoff, one mile south of 
Grand Tower, Jackson County, August 20, 1954, Mohlenbrock 4640 (UI, 
A): near Cora, Jackson County, August 24, 1954, Mohlenbrock 4648 
(UI, A). 

CAREX BRACHYGLOSSA Mackenz.. The Jackson County collection ex- 
tends the range of this primarily northern Illinois species to the south. coL- 
LECTION DATA: edge of Walker Hill Pond, Grand Tower, Jackson County, 
July 15, 1954, Mohlenbrock 4753 (A). 

CAREX SPARGANIOIDES Muhl. The habitat for this species is rich 
mesic woodlands. The Jackson County locality is about 175 miles south 
of the nearest station in Illinois for this species (Menard County). CoL- 
LECTION DATA: rich, moist woods, Lake Murphysboro area, Jackson 
County, June 12, 1954, Mohlenbrock 2679 (A). 

ALLIUM STELLATUM Fraser. This is one of the species characteristic of 
the hilltop prairies found along the southwestern border of the state. 
These prairies, atop limestone bluffs, are minute replicas of the prairies 
to the west. Other species which grow in this type of community include 
Bouteloua curtipendula, Andropogon scoparius, Andropogon  gerardi, 
Petalostemum purpureum, Petalostemum candidum, and Kuhnia eupa- 
torioides. The Allium is known in Illinois only from Jackson, McHenry 
and Union Counties. COLLECTION DATA: hilltop prairie north of the Pine 
Hills, Jackson County, August 6, 1954, Mohlenbrock 4760 (UI, A). 

HEXALECTRIS SPICATA (Walt. Barnh. A colony of seventeen plants 


322 Rhodora [VoL. 57 


of this rare orchid was discovered growing on an east-facing slope in an 
oak-hickory woods in Jackson County. Only two other locations are 
known for this species in Illinois (Randolph and Pope Counties). cor- 
LECTION DATA: dry wooded slope, Fountain Bluff, Jackson County, July 
20, 1954, Sanders, Voigt, and Mohlenbrock 4330 (SIU) 

RUMEX MARITIMUS L. var. FUEGINUS (Phil.) Dusen. This species, like 
Hemicarpha micrantha, was found growing in sand one mile south of 
Grand Tower. It now is known in Illinois from McHenry, Tazewell, 
Whiteside, and Jackson Counties. COLLECTION DATA: in sand, Missis- 
sippi River cutoff, one mile south of Grand Tower, Jackson County, 
August 20, 1954, Mohlenbrock 4641 (A). 

POTENTILLA PARADOXA Nutt. Specimens of this plant were found 
growing in sand along the Mississippi River in close association with 
Hemicarpha micrantha and Rumex maritimus var. fueginus. In addition 
to Jackson County, it is known in Illinois from Randolph and St. Clair 
Counties. COLLECTION DATA: Mississippi River cutoff, one mile south of 
Grand Tower, Jackson County, August 20, 1954, Mohlenbrock 4637 
(UI, A). 

GEUM VIRGINIANUM L. Since this species was first reported from Illi- 
nois in 1954,° it has been found in three additional counties. The habitat 
at each station is in an open oak-hickory woods. COLLECTION DATA: 
Pounds Hollow, Gallatin County, summer, 1954, Mohlenbrock, Voigt, and 
Sanders 1921 (SIU); Panther's Den, Williamson County, summer, 1954, 
Mohlenbrock, Voigt, and Sanders 1976 (SIU); Jackson Hollow, Pope 
County, July 15, 1954, Mohlenbrock 4311 (A). 

CORNUS ALTERNIFOLIA L. f. This dogwood is not too uncommon in 
northern Illinois, but the Pope County collection is the first reported from 
southern Illinois. It grows in abundance in a rich beech-maple forest at 
Belle Smith Springs. COLLECTION DATA: rich woods, Belle Smith Springs, 
Pope County, June 23, 1954, Mohlenbrock 3889 (A), 

ONOSMODIUM HISPIDISSIMUM Mackenz. This species is not common in 
southern Illinois, the present collection being the first from Jackson 
County. It grows on an outerop of limestone along the Mississippi River. 
COLLECTION DATA: atop a limestone bluff, Devil's Bake Oven, near Grand 
Tower, Jackson County, June 27, 1954, Mohlenbrock 3206 (A). 

LIATRIS CYLINDRACEA Michx. This is another of the hilltop prairie 
species of southwestern Illinois. The Jackson County station is the only 
extreme southern Illinois record. COLLECTION DATA: hilltop prairie north 
of the Pine Hills, Jackson County, Sept. 9, 1954, Mohlenbrock 4758 (A, UI). 

The author wishes to thank Mr. Harry E. Ahles of the University of 
Illinois Herbarium who supplied the distributional records for each species 
in Illinois.—RonzERT H. MOHLENBROCK, DEPARTMENT OF BOTANY, 
WASHINGTON UNIVERSITY, ST. LOUIS, MISSOURI. 


6 MoHLENBROCK, R. H. RHopona 56: 227-2298, 1954. 


1955] Imshaug,—The Lichen Genus Buellia 328 


AN ADDITIONAL SPECIES OF THE LICHEN GENUS 
BUELLIA FROM THE WEST INDIES 


Henry A. [MSHAUG! 


THE present author has recently (Imshaug 1955) published 
a revision of the West Indian species of Buellia. Since then 
he has studied the West Indian species of the closely related 
genus Rinodina. Only three saxicolous species of Rznodina 
have been reported from the West Indies and all three were 
described by Wainio from material collected in the Virgin Islands 
by F. Børgesen in 1906; R. antillarum from St. Thomas, R. 
boergesenii from St. Croix and R. pyxinoides from St. John. 
Each species was collected from only one locality, each on a 
different island. I have been able to examine these collections 
through the courtesy of Dr. M. Skytte Christiansen to whom 
I would like to express my sincere appreciation. 

All three collections represent the same species which is identi- 
cal with Buellia microphylla Malme. Wainio’s three names, how- 
ever, antedate B. microphylla. Of the three names only the 
specific epithet pyxinoides has not previously been used in 
Buellia. 

A description of the West Indian material follows: 


Buellia pyxinoides (Wain.) Imshaug, comb. nov., based on Rinodina 
pyxinoides Wain. Ann. Acad. Sci. Fenn. A. 6(7): 75. 1915. Rinodina 
boergesenii Wain. Ann. Acad. Sci. Fenn. A. 6(7): 76. 1915. Rinodina 
antillarum Wain. Ann. Acad. Sci. Fenn. A. 6(7): 77. 1915. Buellia 
microphylla Malme, Ark. Bot. 21A(14): 40. 1927. 

Type CoLLECTIONS: R. pyxinoides—Collected on non-caleareous rocks 
at Cruzbay, St. John (West Indies) by F. Bgrgesen on March 13, 1906. 
Holotype in C. R. boergesenii—Collected on non-caleareous rock at 
Hams Bluff, St. Croix (West Indies) by F. Børgesen on Feb. 6, 1906. 
Holotype in C. R. antillarum—Collected on non-calcareous rock at 
Magensbay Estate, 800 ft. elev., St. Thomas (West Indies) by F. Børgesen 
on Dec. 28, 1906. Holotype in C. B. microphylla—Collected “in rupe 
praerupta subumbrosa” in Paraguay (Paraguari, Santo Tomas) by G. O. 
Malme, no. 1508 B. Isotype seen in MO. 

HYPOTHALLUS composed of many black patches which tend to become 
continuous. THALLUSs initiated as small, round, flat areoles which expand 
to become convex and lobate or squamulose, rarely becoming contiguous 
or rimose-areolate; cinereous or sometimes + ochraceous. APOTHECIA at 
first immersed but soon emergent with superficial thalloid covering 


1 Department of Botany, University of Idaho, Moscow, Idaho. 


324 Rhodora [Vor. 57 


which soon disappears, round or occasionally crenate, 0.3-0.7 mm. 
across; disks black, plane, naked; margins rather thick and concolorous. 

HYPOTHECIUM + colorless; exciple + colorless inside but with thick, 
dark brown margin. HYMENIUM 70-75 y thick, colorless, not inspersed 
with oil drops; paraphyses enlarged at apices and fusco-capitate, forming 
a light brown epithecium. Spores 8 in ascus, mischoblastiomorph, 
fumose, (14) 15-21 (24) x (7) 8-11 (12) u. CHEMICAL REACTIONS: Thallus 
section KOH —; apothecial tissues KOH—. 

MATERIAL SEEN: ST. CROIX—Hams Bluff, Børgesen, 1906 (C). sr. 
THOMAS—Magensbay Estate, 800 ft. elev., Børgesen, 1906 (C). sr. JOHN— 
on maritime rocks, Cruzbay, Børgesen, 1906 (C). 


Wainio recognized the similarity between Rinodina pyxinoides 
and R. boergesenii. The latter species was separated only by 
apothecia with thinner margins and non-mamillate disks. These 
were the same characters Wainio used to separate Buellia gyrosa 
Wain. (Syn. B. trachyspora) from B. trachyspora Wain. Wainio 
did not compare Rinodina antillarum with R. pyxinoides but 
the only significant difference I could find was in the spore size. 
Since the spore size seems rather variable and since my measure- 
ments do not agree with those given in the original descriptions 
they are recorded here: 


Author’s 
measurements Original 
(in KOH) deseription 
R. pyxinoides (holotype) 17-24 x 10-12 u 16-21 x 8-9 u 
R. boergesenii (holotype) 17-21 x 9-1l à 13-17 x 7-9 u 
R. antillarum (holotype) (14) 15-17 x 7- 8u (10) 12-16 x 5-8 u 
B. microphylla (isotype in MO) 18-24 x 10-14 u 16-21 x 8-9 u 


LITERATURE CITED 


ImsnauG, H. A. 1955. The lichen genus Buellia in the West Indies. Far- 
lowia 4 (4): 473-512. 1955. 


Volume 57, no. 682, including pages 259—299, was issued 31 October, 1955. 


odora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
REED CLARK ROLLINS, Editor-in-Chief 


ALBERT FREDERICK HILL 

STUART KIMBALL HARRIS l 

RALPH CARLETON BEAN Associate Editors 
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CARROLL EMORY WOOD, JR. 


Vol. 57 December, 1955 No. 684 

CONTENTS: 

French’s Shooting Star in Southern Illinois. John W. Voigt and 

TUUS R S WANE T a a ae IIO a R bc 325 
Pedanticism Runs Amuck. H. A. Gleason... ........ eese. 332 
A Dark-hooded Variant of Asclepias amplexicaulis. Leonard J. 

IU rii RIA E MEME lr RESET BUENO 336 
Scotch Heather 90H slr Ker 1 0000 eile ni e 337 
Errata — 25 0 e dn nu M E c 338 
IndexitoBVolumebo7 m up n cM a 339 


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Vol. 57 December, 1955 No. 684 


FRENCH'S SHOOTING STAR IN SOUTHERN ILLINOIS 
JouN W. VoicT AND JuLIUS R. SWAYNE 


FnRENCH'S shooting star Dodecatheon frenchii (Vasey) Rydb.! 
is of local interest. It was first discovered and named for 
George Hazen French an early teacher of biology at Southern 
Illinois State Normal (now Southern Illinois University), at 
Carbondale, Illinois. A specimen bearing the date 1870 is 
located in the herbarium of Southern Illinois University. The 
label bears the following inscription, ‘‘Dodecatheon Frenchn 
V., this is thought to be a new species as it differs much from 
D. meadia." The word type is written on the sheet near the 
label. 'The species name, authority initial and the word type 
are written in pencil while the label is written in ink. The 
specimen is in a fruiting condition. A second sheet collected 
on May 6, 1871, also from “Fern Rocks" is probably of the 
same collection as that marked type at the Chicago Natural 
History Museum. According to Fassett (1944) the sheet at 
Chicago was designated the type presumably by McBride. 
The original description did not designate a type or date of 
collection. The earliest collection date previously stated in 
print was 1871. Thus the 1870 date on the type specimen 
establishes the date of collection of the type series as one year 
earlier than previously indicated. 

The last complete work on the genus in eastern North America 
was that of Fassett (1944). Before Fassett’s treatment French’s 
shooting star had been known at different times as a variety, 
subspecies, and species. On the basis of Fassett’s work French’s 


1The nomenclature is that of Jones 1950. Flora of Illinois, University of Notre 
Dame Press, Notre Dame, Ind. 


326 Rhodora (VoL. 57 


shooting star has been accepted by many as a variety of Mead’s 
shooting star (D. meadia L.). 

Dodecatheon frenchii has been described as a plant having 
"leaves abruptly narrowed to the petiole . . . extreme plants 
with broad oval subcordate blades. The inclusion of all plants 
with leaves tending to be abruptly narrowed to the petiole 
has led to various interpretations of range as being much wider 
than it is" (Fassett, 1944). Vasey gave the range as Pennsyl- 
vania to southern Illinois and Arkansas. Rydberg (1932) gave 
it as Illinois, Minnesota, Arkansas, and Pennsylvania. Fassett 
(1944) restricts the range to the Illinois Ozarks with a single 
specimen of doubtful determination being cited from Wisconsin. 
Recent intensive searches in southern Illinois for the plant 
recognized by Fassett as D. frenchii show that it is restricted 
to à belt of about 10 miles width across the State (Fig. 1). 

Dodecatheon frenchiíi has always been found in canyons of 
streams flowing primarily southward and under sandstone 
ledges which may face any direction. It is found most often 
under north and east facing bluffs. D. meadia is found in several 
central and northern counties of Illinois, in the Illinois Ozarks 
and southward. 

It has been reported that D. frenchii differs genetically from 
the widespread phases of D. meadia, but that special ecological 
conditions are required for a phenotypic expression of the varietal 
phase (Fassett, 1944). He also reported that D. frenchii is 
changed to the likeness of D. meadia when grown under increased 
amounts of light and a longer light period. D. meadia is reported 
to be replaced by, or to grade into, D. frenchii under cliffs and 
shaded places. This ecological behavior has not been noted 
by us. The plants have been found to be distinct. The dis- 
covery of D. frenchii in open well-lighted upland woods at 
Jackson Hollow stil maintaining its distinctive vegetative 
characteristics demonstrates that this plant may occasionally 
be found away from the cliff habitat though it exhibits a strong 
"preference" for it. It also shows that the leaf-shape which 
so characterizes this taxon may also be manifested in brighter 
light and that reduced light is not necessary for its phenotypic 
expression. These plants were growing some fifty yards from 
the cliff where other plants of D. frenchii were growing. 

Light readings in the habitat of D. meadia ranged from 5,000 


1955] Voigt and Swayne,—French’s Shooting Star 327 


to 6,000 foot candles when readings in the open recorded 10,000 
foot candles. The readings in the habitat of D. frenchii were 
usually about 14-25 per cent less than the light recorded in the 
habitat of D. meadia. The light meter was held directly over 
the plants in such a position as to receive the fullest amount 
of sunlight. In addition to these readings, which were taken 
from several locations, the light was measured in the upland 
habitat of the open woods where about 40 plants of D. french 
were found growing at Jackson Hollow. Over a five hour 
period seven readings were made. The average of these read- 


RANDOLPH | PERRY JFRANKLIN HAMILTON | WHITE 


JACKSON WILLIAMSON |SALINE GALLAT 
e 

vest: LLS stkricfle 

[] e| @@ 

gee 


UNION JOHNSON POPE 


HARDIN, 


ALEXAND ULAS AS 
i j 


Fig. 1. The southern sixteen counties of Illinois. The white area represents the 
Shawnee Hills Section. The area to the north is glaciated. Dots represent stations 
for the occurrence of D. frenchii. Horizontal hatch is the Salem Plateau section; 
vertical hatch is the Coastal Plain province. Map adapted from Leighton, Ekblaw, 
and Horberg. 


ings, where most of the plants were growing, was about 5,000 
foot candles. The readings were started at 11:00 a.m. and 
terminated at 4:00 p.m. The average of seven readings over 
the same period of time for a station 3 feet distant from the 
first, where a few more plants were growing, was about 6,500 
foot candles. It is realized that averages of factor intensities 
are ordinarily of little value, but here the amount of light and 
its duration is equivalent to and often exceeds that found in 
the habitats of D. meadia. Light readings taken under the 
bluff and down the hill some 50 yards away where most of the 
D. frenchii plants were growing, showed only 11 foot candles 
at the edge of the bluff. At 3:45 p.m. the reading here was 
only 8-10 foot candles. The plants growing under the bluff 


328 Rhodora [Vor. 57 


never receive direct sunlight and for only half of the day do they 
receive as much as 200 to 300 foot candles. 

The effect of increased lighting on D. french?i and decreased 
lighting on D. meadia was studied in the greenhouse. Several 
dozen resting buds of D. frenchii and D. meadia were collected 
on January 17, 1952 and planted. The natural light was sup- 
plemented by two fluorescent tubes (30 watt) which were placed 
over the plants at a distance of about 20 inches. "The total 
day length period was 15 hours. The greenhouse temperatures 
were set for 60 degrees at night and 80 degrees during the day. 
The plants appeared above ground in about a week, grew rapidly 
and initiated flower stalks at the end of the third week. "The 
buds opened into flowers during the sixth week at which time 
the plants were harvested. "The time for the development and 
appearance of flowers was close to that required in nature. No 
important differences of leaf shape were noted between these 
plants and any others of D. frenchii that had been seen. Both 
taxa were subjected to diminished light by placing the same 
number of resting buds of each under a cubicle of plastic material 
which reduced the light to about 25 per cent of full sunlight. 
The only major change was in the length of the leaves of D. 
meadia. They became longer and thinner but did not otherwise 
change shape or bear any likeness to D. frenchi?. 

The plants of D. frenchii and D. meadia were started again 
after 2 months of storage in the bottom of a refrigerator. They 
were once more subjected to increased lighting and started 
their vegetative growth promptly. This time they were grown 
under three mazda bulbs (300 watts each). The bulbs were 
approximately 3 feet from the plants, and timed for a 15 hour 
day. These plants also showed no change of leaf shape. A 
third trial was made, following the same procedure as in the 
second, and again negative results were obtained. 

Fassett's observation of intergradation of D. frenchii with 
D. meadia or the change of leaf shape of D. frenchii to the likeness 
of D. meadia is suspected to be due to the variation of the popu- 
lations of D. frenchii. A considerable number of plants in any 
population of D. frenchii will be found to vary some in leaf 
shape. However when plants that are growing side by side 
show leaf shape variation which does not approach the shape, 
color, or texture of leaves of D. meadia then one wonders what 


1955] Voigt and Swayne,—French’s Shooting Star 329 


environmental influences can be so subtle as to change the 
leaves of one D. frenchii plant and not another growing next to it. 

Other environmental factors were investigated. These in- 
cluded measurements of organic matter, moisture content, pH, 
evaporation, temperature and humidity. Only the last three 
factors showed any appreciable differences in the two habitats. 
The maximum difference in temperature for any 24 hour period 
was 12 degrees Fahrenheit, and the average over a 5 week period 
was 7.5 degrees. The lower temperature always prevailed 
in the habitat of D. frenchii as did the higher humidity and 
lower evaporation rate. 


MORPHOLOGICAL ANALYSIS 


The shape of the leaf in both taxa has been found to be dis- 
tinct and unchanged by a change in the lighting of the environ- 
ment. When width of leaves is plotted against the length of 
leaves on logarithm paper for both taxa the points for each fall 
mostly on separate straight lines. These slope determinations, 
figured by least squares, were 0.9 for D. frenchii and 0.88 for 
D. meadia. Measurements were made from 201 samples of 
which 126 were for D. frenchii and 75 were for D. meadia. The 
samples were random and from all known stations. A few points 
from this plotting do fall between the two slope lines indicating 


B/P B/P 


Fig. 2a. Polygonal graph showing blade/petiole, length/width relationships and 
angle of contraction of blade to petiole of leaves of D. meadia which seemed ‘‘inter- 
mediate" by logarithmic plotting of length vs width of leaves. 

Fig. 2b. Polygonal graph showing relationships of blade/petiole, length/width 
and angle of contraction of blade to petiole of D. frenchii, These measurements 
were selected from leaves of plants which were “intermediate” in logarithmic plotting 
of length vs. width of leaves. 


330 Rhodora [Vor. 57 


that some leaves of each taxon are intermediate as to length and 
width. The length-width relationship of the leaves however 
does not show the character that is most important in dis- 
tinguishing the two taxa, namely, the abrupt angle of contraction 
of blade of leaf to the petiole in D. frenchii. 

The angle at which the blade joins the petiole was calculated 
for both taxa from the individuals which seem to be intermediate 
by the logarithmic plotting of length vs width of leaves. The 
measurement of the angle of contraction of blade to petiole 
was done according to the method of Voigt (1952). The in- 
dividuals which seem to be intermediate as to leaf length and 
width were plotted on polygonal graphs (fig. 2a & b) and are 
seen to be distinctly different. 

The blade-petiole ratio (B/P) was always greater for D. 
meadia. The length-width (L/W) ratio was also more variable 
for D. meadia. The angle ( €) of contraction made by the base 
of the blade to the petiole was always greatest for D. frenchii. 
Polygonal projection of other individuals whose measurements 
fall more closely to the slope lines would only have these differ- 
ences more greatly emphasized. This method of graphing has 
the advantage of giving an objective comparison while portraying 
several characters (Davidson, 1947). When the graphs of the 


B/P 


Fig. 3a. Polygonal graph showing morphological comparison of leaves of D. 
meadia and D. frenchii found growing in the same habitat. The individuals were 
six feet apart. Measurements were made of blade/petiole, length/width, and angle 
of contraction of blade to petiole. Heavy line is for D. meadia. 

Fig. 3b. Polygonal graph as previously done. The plants were found in the 
same area fifteen feet apart. The heavy line is for D. meadia. 


1955] Voigt and Swayne,—French’s Shooting Star 331 


two entities are superimposed a high degree of crossing of the 
lines emphasizes remoteness of morphological similarity whereas 
a high degree of paralleling denotes a closer morphological 
similarity. 

Measurements made in the manner just described are shown 
(fig. 3a) for D. frenchii and D. meadia growing together in the 
same habitat. Single individuals are shown. In each case 
only one individual of D. meadia was found. The plants rarely 
occur in the same habitat (in two cases, the habitat was that 
of D. frenchii). The nearest plant of D. frenchii to the single 
individual of D. meadia was selected for comparison. The 
distance apart in fig. 3a was 6 feet, and in fig. 3b the distance 
was 15 feet. In both instances the distinctness is quite ap- 
parent. 

Colonies of D. frenchii are not usually recognized by the 
color of the corolla lobes because white corolla lobes are found 
in nearly every plant. A dark purple color as given by Ryd- 
berg (1932) and Jones (1950) has not been found in the corolla 
lobes of D. frenchit. 

Plants of D. frenchii transplanted into the habitat of D. meadia 
at Giant City State Park in Illinois have remained distinct 
through three seasons of growth. They flowered in the third 
year. Plants of D. meadia transplanted into the habitat of 
D. frenchii have remained distinct as to leaf shape and have 
even retained their darker color. The leaves did become thinner. 

A change of D. meadia into D. frenchii has not been observed. 
The vegetative morphological distinctness together with the 
lack of intergrading forms and the differences in ecological 
behavior of these presumed species are probably significant. 
However, a more complete investigation including genetic and 
cytological studies are required for a final determination of the 
taxonomie status of these taxa. 

Detailed collection data are on file at the herbarium of South- 
ern Illinois University. As indicated by the map (fig. 1) sta- 
tions for D. frenchii are known from Jackson, Union, Johnson 
amd Pope Counties.—SOUTHERN ILLINOIS UNIVERSITY, CARBON- 


DALE, ILLINOIS. 
LITERATURE CITED 


Davipson, J. F. 1947. The polygonal graph simultaneous portrayal of 
several variables in population analysis. Madrono, 9: 105-110. 


332 Rhodora [Vor. 57 


Fassett, N. C. 1944. Dodecatheon in eastern North America. American 
Midland Naturalist No. 2, 31: 455-486. 

Jones, G. N. 1950. Flora of Illinois. American Midland Naturalist, 
monograph No. 5., The University of Notre Dame Press, Notre Dame, 
Indiana. 

Leicuton, M. M., G. E. ExBLAWw, AND L. HonBERG. 1948. Physiographic 
Divisions of Illinois. The Journal of Geology, Vol. 56, No. 1. 

RYDBERG, P. A. 1932. Flora of the prairies and plains of central North 
America. New York. 

VoraT, J. W. 1952. A technique for morphological analysis in population 
studies. Rhodora 54: 217-220. 


PEDANTICISM RUNS AMUCK 
H. A. GLEASON 


IT is now seventeen years since the article* was published and 
for seventeen years I have considered a reply to it. At first I 
remained silent out of respect to my friend Alfred Rehder, 
who was unfortunately, and I suspect rather unwittingly con- 
cerned with it. 

The title of the paper is misleading. Botanists who may at 
this point fear that they have been asked or will now be asked 
to learn a fourth name for this beautiful, valuable, and widely 
distributed tree may be at ease. "The article leaves the name 
Pseudotsuga taxifolia unchanged and refers only to the author- 
citation for it, that is, to the botanists who are responsible 
for the name in the recommended form. Nevertheless it is 
significant that the authors betray in their title something of 
the growing tendency to convert the ordinary binomial system 
of nomenclature into a trinomial or quadrinomial system, in 
which the "authorities" constitute the third and fourth terms. 
Every teaching taxonomist still tells his students that the name 
of a species consists of two terms which together are sufficient 
to designate the species and at the same time show something 
of its place in the scheme of classification. The International 
Code, in the formation of which the authors played such a 
prominent part, stil affirms the binomial system. But this 
growing tendency is often apparent. 

Since the Kew Bulletin, as well as the original sources on 
which Sprague and Green base their conclusions, may not be 


* SPRAGUE, T. A., AND M. L. Green. The botanical name of the Douglas Fir. 
Kew Bulletin 1938: 79, 80. 1938. 


1955] Gleason,—Pedanticism Runs Amuck 333 


generally accessible to American readers, the statements of 
the authors may be briefly summarized. 

The first binomial given to the Douglas Fir was Pinus taxi- 
folia by Lambert in 1803. Later the tree received two other 
specific epithets, mucronata from Rafinesque and Douglasii from 
Lindley. All three of these have been transferred to Pseudotsuga, 
resulting in three binomials, of which two must be synonyms 
and only one, if any, can be valid. All three have been in 
general use in American botany and forestry. Under prevailing 
rules of nomenclature, the oldest of these specific epithets must 
be used, provided it was valid when published and after its 
transfer produces a valid binomial. 

The oldest epithet is actually invalid, the name Pinus taxifolia 
as used by Lambert being a later homonym of Pinus taxifolia 
as previously used by Salisbury. Under the International 
Code an invalid epithet can not be validly transferred. There- 
fore Pseudotsuga taxifolia (Lambert) Britton is also invalid. 
Apparently the next available epithet is mucronata and the 
valid name Pseudotsuga mucronata (Raf.) Sudworth. Sprague 
and Green, however, claim that the Douglas Fir was also named 
Abies taxifolia by Poiret in 1804 and that this name is validly 
published and transferable. They then proceed (on behalf of 
Rehder) to transfer this epithet, which is considerably older 
than those of Rafinesque and Lindley, to Pseudotsuga, giving 
us the same name with different authors, P. taxifolia (Poir.) 
Rehder. 

Sprague and Green are expert in all the intricacies of modern 
nomenclatural etiquette and their interpretation of the rules, 
as applied to this particular case, may be accepted without 
question. In brief, the applicable rules are three: 

1. An invalid epithet (taxifolia in Pinus taxifolia Lamb.) 
upon combination with another generic name (Pseudotsuga) 
produces an invalid binomial (Pseudotsuga taxifolia Britt.) 

2. A later homonym (Pseudotsuga taxifolia Rehder) is not 
invalidated by an older one (Pseudotsuga taxifolia Britton) 
if both are based on the same type. 

3. The invalidity of an epithet in one genus (taxifolia in 
Pinus, as used by Lambert) does not preclude its valid use in 
another genus, even for the same species, provided it is described 
there as new and not merely transferred. 


334 Rhodora [Vor. 57 


These rules are the ones invoked by Sprague and Green and I 
agree with them completely. The sole question to be deter- 
mined is whether Poiret described the Douglas Fir as a “new” 
species, thereby creating a valid and transferable specific epithet. 

Lambert’s description (1803) reads as follows: 


Tab. 33. 
27. Pinus taxifolia. 
Nootka Fir. 


Pinus taxifolia, foliis solitariis planis integerrimis, strobilis oblongis, antheris 
inflato-didymis. 

Habitat ad Americae borealis oras occidentales. 

Descriptio. 

Habitus P. canadensis, at folia angustiora et paululum longiora, integer- 
rima. Amenta mascula ovata, subsessilia, multiflora; antheris inflato- 
didymus, crista reflexa, minima. 

The figure was taken from a specimen in the Banksian Herbarium, brought 
home by Mr. Menzies, by whom it was discovered on the Northwest coast 
of America, and who has favored me with the following particulars respecting 
this species. 

In general habit this tree resembles P. canadensis, and attains considerable 
height and size. The leaves are also very like those of the species just men- 
tioned, but narrower and their edges are entire, whereas the others are visibly 
serrated. The inflorescentia is much larger than in P. canadensis and there 
are more antherae. As for the Cones, I can give no account of them, those 
which were brought by Mr. Menzies having been unfortunately mislaid. 
That gentleman however informs me that they differ in their form from the 
cones of P. canadensis, and that they are longer. 


During the single year after this publication no other specimen 
of the Douglas Fir was brought to Europe. In 1804 Poiret 
had occasion to write about trees and the only source of infor- 
mation on the Douglas Fir was Lambert's description and plate. 
It is conceivable that he borrowed the specimen from the Banks 
herbarium, but not probable, since England and France were 
not on good terms at that time. — Poiret's description now follows, 
and the reader is advised to compare it carefully, clause by 
clause, with that of Lambert, remembering that the order of 
presentation is considerably changed. 


15. Sapin à feuilles d'if. Abies taxifolia. Lambert. 
Abies foliis solitariis, planis, integerrimis; strobilis oblongis, antheris inflato- 
didymis. Lambert, Descript. of Pin. pag. 51. tab. 33. 
Cette espéce a de grands rapports avec le Pinus canadensis par son port, 
& méme par l'élévation de son trone. Ses rameaux sont un peu diffus, opposés 


1955] Gleason,—Pedanticism Runs Amuck 335 


ou alternes; ses feuilles sont plus étroites & plus longues, trés entieres, glabres 
à leurs deux faces, planes, solitaires. Les chatons males sont ovales, presque 
sessiles, trés entieres, trés chargés de fleurs; les antheres renflées & à deux 
longes; leur créte réfléchie & fort petite; on soup conne que les cones sont 
beaucoup plus longs que ceux du Pinus canadensis. 

Cet arbre croît sur les côtes occidentales de l'Amérique septentrionale. 


Careful comparison of the two descriptions will show that 
almost all information given by Poiret has been taken, usually 
by literal translation, from Lambert. There is a little additional 
matter, as “rameaux sont un peu diffus, opposés ou alternes,” 
or "feuilles plus étroites, planes, solitaires,” and this might 
have been taken from Lambert’s plate. The only significant 
difference is the substitution of the generic name Abzes for 
Pinus. It is not mere plagiarism; he cites Lambert’s work 
accurately. It is merely the transfer of a specific epithet to a 
new generic position. Transfer of an invalid specific epithet 
produces an invalid binomial. If Lambert’s taxifolia produced 
an invalid binomial when transferred to Pseudotsuga by Britton, 
then it also produces one when transferred to Abies. 

Abies taxifolia, as used by Poiret, can not possibly be regarded 
as a new" species. It is a transfer, if there ever was one. The 
basonym and its author are clearly stated, the description is 
a mere translation into French. 

Was the epithet taxifolia ever used elsewhere in a description 
of the Douglas Fir as a “new” species? No such use is known 
to exist in the few years between its first and invalid appearance 
in 1803 and the publication of mucronata by Rafinesque. If such 
use was made of it at any later time, it would have resulted 
merely in an invalid synonym. The use of taxifolia as the 
specific epithet for the Douglas Fir is definitely and finally 
excluded, and the name of the tree is once again Pseudotsuga 
mucronata, the specific epithet by Rafinesque, the combination 
by Sudworth. Under that name it was known for many years 
in most American literature.—GREENWICH, CONN. 


336 Rhodora [Vor. 57 


A DARK-HOODED VARIANT OF ASCLEPIAS AMPLEXICAULIS.— 
Asclepias amplexicaulis Sm. is a strikingly-formed Milkweed, 
with a solitary (rarely 2 or 3), long-peduncled, ample umbel, 
and crisped-margined, broad, glaucous leaves. It is familiar in 
dry, open situations in the eastern United States. Normally, 
its hoods are pinkish, or flesh-colored. Small,! however, describes 
the hoods as ‘‘maroon.’”’ It seems odd, therefore, that while 
this term is hardly apt for the usual hood color of the usual 
colonies of the species, it can, with tolerance, be applied to the 
dark-colored hoods of a certain population I encountered on 
August 1, 1954, in the upland pine-sand country of southwestern 
North Carolina, at Lake Osceola, Hendersonville, Henderson 
County. No other specimens of A. amplexicaulis were noted 
nearby, and the only other asclepiad seen in the vicinity was 
A. incarnata L., abundant on the lake shores a few rods away. 
A single specimen of this aberrancy was collected, later deposited 
in the New York Botanical Garden Herbarium. This specimen 
exhibits no deviation from typical A. amplexicaulis in any 
character other than hood coloration. Technically, this color 
approaches Ridgway's? chip of “Dahlia Carmine." For a more 
immediately practical concept, the color of the hoods of this 
variant quite fairly approximates the well-known, living flower 
color of Trillium erectum L. 

The literature on A. amplexicaulis seems to be free of reference 
to flower color variation, but I am apparently not the first to 
encounter the present deviation. Mr. Joseph Monachino, of 
the New York Botanical Garden, who, incidently, verified my 
specimen, wrote me that he once encountered a ''dark-purple 
hooded" plant in New Jersey. Similarly, Mr. Roy Latham, 
of Orient, New York, informs me he has seen dark-hooded plants 
on Long Island. It appears, therefore, that there exists a wide- 
ranging color form of A. amplexicaulis, wherein the usual an- 
thocyanic pigmentation of the hoods is grossly intensified, the 
flowers (crown) thus appearing dark maroon-purple in life, 
more vivid and showy than typical plants. The dark form has 
not been adequately collected, and, therefore, awaits acquisition 
of supporting material for its proper delineation. Therefore, 


! SMALL, J. K. 1933. Manual of the Southeastern Flora. p. 1070. 
2? Ripaway, R. 1912. Color Standards and Nomenclature. Pl. XXVI. 


1955] Parker,—Scotch Heather 337 


field workers are asked to watch for dark-hooded A. amplexi- 
caulis, and to collect specimens with proper notations. I would 
be very grateful to learn of any such observations and collections. 

The hood color of most herbarium specimens of A. amplexi- 
caulis is a meaningless strawy hue. Occasionally a dried speci- 
men turns up in which the hoods have turned a deep plum shade. 
My specimen turned this color on drying. Yet, with all the 
variables to consider, this can hardly be taken to indicate a 
pattern. It is especially deplorable, that there is rarely a 
collector’s note on flower color with these herbarium specimens. 
Because flower color is so useless in identifying dried Milkweeds, 
Carr® prepared an asclepiad key to circumvent this character. 
Happily, the color of my aberrant specimen will be preserved. 
The living plant was photographed in color, and a print was 
deposited with the specimen. In addition, for comparative 
purposes, a color photograph of a normal pinkish Long Island 
plant was included.—Lreonarp J. UTTAL, 1258 BEACH ROAD, 
RIVIERA BEACH, FLA. 


Scorch Hratuer.—lIn the town of Hartland, in the north- 
western part of Hartford County, Connecticut, at an elevation 
of 1200 feet, there is a good stand of Calluna vulgaris, which 
extends over an area of about an acre. Since it is rather unusual 
to find this shrub in such a location, which has very little pro- 
tection from the winter cold, the writer attempted to trace the 
history of the introduction of the plant to this part of Con- 
necticut. Mr. L. E. Pearson, a forester in Connecticut, first 
noticed it when looking over the woodland of the present owners, 
Dr. and Mrs. Henry A. Sturman. 

Most of the following information was obtained by the Stur- 
mans in conversation with local inhabitants of the area. The 
present Sturman farm was owned by one John Schwaller and his 
wife, who came to America from Alsace-Lorraine in the 1870s 
and settled on the property in Hartland. It is reported that 
the original seeds of the present stand of heather were sent in 
a letter from Mrs. Schwaller’s mother who told her daughter 
that the shrub would be valuable for winter forage for the cows. 


3 CARR, K. 1942. A Key to North American Asclepiads. Castanea 7: 1-7. 


338 Rhodora (Vor. 57 


The exact year the seeds were sent is not known but presumably 
at least 40 years ago, and possibly 50 or 60 years ago. It would 
probably be safe to assume that the stand has been in existence 
for 50 years. 

The present site is an old field which has commenced to 
grow up with gray birch, white pine and some juniper, as well 
as mountain laurel. It would appear that the pines offer some 
protection from the winter storms. However, the site being 
on the top of a rather exposed hill, does not appear to be a 
location in which the heather would thrive. The remarkable 
thing is that it has apparently continued to spread slowly for 
about half a century. 

Some of the plants show evidence of winter killing in the tops 
but the branches underneath seem to remain protected so they 
leaf out again each spring and come into full flower each summer. 

At this writing, the sixth of August, the plants are in full 
bloom and present a beautiful sight. In conversation with 
Dr. and Mrs. Sturman a few days ago they said some Scottish 
friends of theirs, now living in this country, actually had tears 
in their eyes when they viewed the shrubs in full bloom.—S. E. 
PARKER, PLEASANT VALLEY, CONNECTICUT. 


Volume 57, no. 688, including pages 301-324, was issued 16 December, 1955. 


ERRATA 


Page 139, last line; for cordifolius read cordifolium. 
Page 221, line 1; for americanum read americana. 
Page 278, line 31; for deltoides read deltoidea. 

Page 280, line 16; for occur read occurring. 

Page 281, line 5; for aculeata read aculeatum. 

Page 291, line 12; for Erigonum read Eriogonum. 
Page 291, line 14; for grandiflorum read grandiflorus. 
Page 294, line 18; for Ethel read Ethyl. 

Page 310, line 30; for insparata read insperata. 


1955] Index to Volume 57 339 


INDEX TO VOLUME 57 


New scientific names and combinations are printed in boldface type 


Abies taxifolia, 333, 334, 335 

Abrus precatorius, 285 

Acacia farnesiana, 285 

Acanthocereus floridanus, 287 

Acer rubrum f. tomentosum, 315; 
saccharum, 131 

Achillea lanulosa, 302 

Achyranthes ramosissima, 281, 285 

Acinos arvensis, 73 

Acrostichum danaeaefolium, 283 

Adiantum pedatum, 236 

Aesculus discolor, 39; glabra, 38, 39, 
40; hippocastanum, 38, 39, 40, 
Plate 1205, 41; octandra, 39, 40, 
Plate 1205; parviflora, 39, 40; 
sylvatica, 39, 40, Plate 1205, 41 

Aesculus, Studies in the Hippocas- 
tanaceae, I. Variation in the 
Mature Fruit of, 37 

Agave americana, 285; decipiens, 
285; sisalana, 285, Plate 1220 

Agropyron trachycaulum var. no- 
vae-angliae, 302 

Alisma Andrieuxi, 175, 176; Ber- 
teroanum, 139, 140; Berteroi, 139, 
140, 141; cordifolia, 137, 140; 
cordifolium 139; ellipticum, 179; 
floribundum, 148; grandiflorum, 
148; intermedium, 179, 180, 181; 
macrophyllum, 153; nymphaei- 
folium, 182; nymphaeifolius, 135; 
ranunculoides var. brasiliense, 
184; rostratum, 135, 144, 147; 
Sprenglii, 140; subalatum, 179; 
subulatum, 185; tenellum, 135, 
183, 184; f. latifolia, 202; virgata, 
137 

Allium stellatum, 267, 290, 321 

Alyssum auriculatum, 260; grandi- 
florum, 262; lasiocarpa, 244; Les- 
curii, 247 

Amaranthus hybridus, 285 

Ambrosia artemesiifolia var. elatior, 
288; hispida, 280, 288 

American Tropics, Echinodorus in 
the, 133, 174, 202 

Ammannia latifolia, 287 

Amorpha canescens, 291, 292; nana, 
291, 293 


Andropogon gerardi, 321; glomera- 
tus, 278, 283; scoparius, 113, 320; 
virginicus, 283 

Annona glabra, 285 

Antennaria canadensis, 302 

Arabis Drummondii, 301, 303, 308; 
hirsuta var. pycnocarpa, 307 

Archer Method for Mounting Her- 
barium Specimens, The, 294 

Arctostaphylos Uva-ursi, 113 

Ardisia escallonioides, 277, 287 

Argemone leiocarpa, 285; mexicana, 
285 

Aristida patula, 278, 283; purpuras- 
cens, 283 

Artemisia cernua, 291; nutans, 291 

Asarum canadense, 131 

Asclepias amplexicaulis, A Dark- 
hooded Variant of, 336 

Asclepias amplexicaulis, 336, 337; 
exaltata, 265; fragrans, 95; incar- 
nata, 336; verticillata, 282, 287 

Asemeia cumulicola, 286; grandi- 
flora var. leiodes, 286 

Asplenium bradleyi, 235, 237, 319; 
cryptolepis, 237; ebenoides, 235, 
237; X ebenoides, 238; X gravesii, 
238; monanthes, 235; montanum, 
237; pinnatifidum, 235, 237, 240; 
platyneuron, 237; rutamuraria, 
229, 237, 239, var. cryptolepis, 
225; trichomanes, 235, 237; tru- 
delli, 237; X trudelli, 240; viride, 
237 

Aster azureus, 317; f. levicaulis, 317; 
cordifolius var. moratus, 317, var. 
polycephalus, 318; dumosus var. 
strictior, 318; furcatus, 268; lae- 
vis, 317; oblongifolius f. roseo- 
ligulatus, 318; Shortii, 131; subu- 
latus, 288; vimineus, 318, var. 
subdumosus, 318 

Astragalus caryocarpus, 293; crassi- 
carpus, 291, 293; gracilis, 291; 
Robbinsii, 302, var. Robbinsii, 
308 

Athyrium angustum, 234; filix- 
femina, 234, 239, var. michauxii, 
225, 229, 234, 236; pycnocarpon, 


340 Rhodora 


225, 229, 234, 237, 239; thelyp- 

teroides, 225, 229, 234, 237, 239 
Atriplex arenaria, 281, 285 
Auriculate-leaved Species of Les- 

querella (Cruciferae), The, 241 
Avicennia nitida, 276, 279, 288 


Baccharis angustifolia, 288; dioica, 
281, 288; glomerulifolia, 277, 288; 
halimifolia, 278, 279, 281, 288 

Bacopa monnieri, 278, Plate 1217, 
288 

Baptisia minor, 266 

Bartonia decapetala, 290; ornata, 
290, 292; paniculata, 315, var. 
iodandra, 308 

Batis maritima, 285 

Berlandiera subacaulis, 288 

Bidens pilosa, 279, 281, var. radiata, 
288 

Blechnum serrulatum, 283 

Boerhaavia paniculata, 285 

Borrichia frutescens, Plate 1221, 
288 

Botrychium dissectum, 223, 225, 
238, f. dissectum, 221, 224, 236, 
f. obliquum, 221, 224, 225, 236, 
f. oneidense, 223; lanceolatum, 
221, 227; lanceolatum ssp. angus- 
tisegmentum, 224, 239, var. an- 
gustisegmentum, 227, 236; lun- 
aria, 226, 238, var. onondagense, 
221, 224, 226, 236, 238; matri- 
cariaefolium, 221, 224, 226, 236, 
238; minganense, 220, 222, 226, 
238; multifidum, 221, 223, 224, 
225, 236, 238, var. intermedium, 
131, var. oneidense, 223; onei- 
dense, 220, 221, 222, 223, 224, 
225, 226, 236, 238; pumicola, 221, 
224, 227, 236, 239; simplex, 224, 
227, 239, var. simplex, 221, 227, 
236, var. tenebrosum, 221, 224, 
227, 236, 239; ternatum, 223; 
virginianum, 221, 223, 234, 236, 
238 

Bouteloua curtipendula, 321; hir- 
suta, 278, 283 

Bromus squarrosus, 266 

Buchnera floridana, 288 

Buellia from the West Indies, An 
Additional Species of the Lichen 
Genus, 323 


[Vor. 57 


Buellia gyrosa, 324; microphylla, 
323, 324; pyxinoides, 323; tra- 
chyspora, 324 

Bumelia angustifolia, 277, 287 

Bursera simaruba, 278, 286 


Cactus ferox, 290; fragilis, 290; vivi- 
parus, 290, 291 

Caesalpinia bonduc, 285; crista, 285 

Cakile edentula var. lacustris, 114; 
fusiformis, 281, Plate 1221, 285 

Calamagrostis inexpansa, 114; in- 
sperata, 310 

Calamintha alpina, 76; arvensis, 76; 
cretica, 76; glabella, 73, var. 
angustifolia, 73; grandiflora, 75; 
montana, 76; Nepeta, 75, 76; 
officinalis, 73, 75, 76; parviflora, 
76; trichotoma, 75 

Calamovilfa longifolia, 113 

Calluna vulgaris, 337 

Calonyction aculeatum, 281, 287; 
tuba, 281, 287 

Calycera balsamitaefolia, 72 

Calycera balsamitaefolia in the 
United States, 72 

Calymenia angustifolia, 291; pilosa, 
290 

Campanula aparinoides, 309; ro- 
tundifolia, 113, 302 

Camptosorus rhizophyllus, 235, 237 

Canavalia obtusifolia, 285 

Cape Cod and Martha's Vineyard, 
Crooked Oaks on, 132 

Capparis cynophallophora, 277, 285 

Capraria biflora, 279, 288 

Cardamine parviflora var. arenicola, 
302, 307 

Cardiospermum halicacabum, 286 

Carex abdita, 311; alata, 156; 
amplisquama 156; aquatilis 
var. altior, 319; atratiformis, 303, 
306; brachyglossa, 321; brevior, 
156; Buxbaumii, 312, f. dilutior, 
312, f. heterostachya, 312; capil- 
laris var. capillaris, 306, var. 
major, 306; communis, 157, var. 
wheeleri, 157; comosa, 306; con- 
voluta, 302; debilis, 312, 319; 
digitalis var. macropoda, 312; 
Mackenziei, 306; pedunculata, 
306; pensylvanica, 157; rosea, 
302, 306; sparganioides, 321; 


1955] Index to Volume 57 341 


stricta var. strictior, 312; Swanii, 
319; torta, 319; Tuckermanii, 
306; umbellata, 311; vexans, 156 

Carica papaya, 287 

Carum carvi, 320 

Cassia nictitans, 207; tora, 285 

Cassytha filiformis, 279, 285 

Casuarina cunninghamiana, 276, 
285; equisetifolia, 276, 279, 281, 
285 

Cenchrus echinatus, 283; incertus, 
283; pauciflorus, 279, 282, 283 

Centaurea vochinensis, 268 

Centaurium, 4; pulchellum, 103; 
umbellatum, 103 

Centrosema virginianum, 285 

Cerastium viscosum f. apetalum, 
313 

Ceratophyllum demersum, 307 

Chamaecrista aspera, 285 

Chara zeylanica, 283 

Cheiranthus asper, 290 

Chenopodium album, 285; am- 
brosioides var. anthelminticum, 
285 

Chiococca alba, 277, 288; pine- 
torum, 288 

Chironia, 2, 3, 93; amoena, 58; 
angularis, 18, var. @ latifolia, 18, 
20, 31, 8 angustifolia, 20, 30, var. 
6 angustifolia, 29; calycosa, 79; 
campanulata, 52, 93, 6 gracilis, 
52; chloroides, 86; cymosa, 7, 11, 
23, 25, 27; decandra, 90, 91, 94, 
95, 103; dichotoma, 79, 93; dode- 
candra, 3, 5, 86, 93; gracilis, 52; 
lanceolata, 7, 10, 11, 25, 93; 
paniculata, 7, 11, 24, 25, B 
angustifolia, 11; pulcherrima, 95, 
103; stellaris, 58; stellata, 58; 
venosa, 7, 12 

Chlora, 2; dodecandra, 86 

Chloris petraea, 278, 282, 283 

Chrysanthemum Leucanthemum 
var. pinnatifidum, 302 

Chrysobalanus icaco, 285; oblongi- 
folius, 285 

Chromosome Numbers in the Genus 
Sesbania (Leguminosae): Evi- 
dence for a Conservative Treat- 
ment, 213 

Cichorium intybus f. album, 319 


Cirsium arvense, 302; horridulum, 
279; f. elliottii, 288 

Cladium jamaicense, 278, 283; 
mariscoides, 113, 306 

Clinopodium, 74, 77, 78; incanum, 
75; rugosum, 75; vulgare, 73, 75 

Cnidoscolus stimulosus 286 

Coccoloba uvifera, 279, 281, 285 

Cocos nucifera, 284 

Commelina erecta, 284 

Conioselinum chinense, 308 

Conocarpus erecta, 272, 277, 279, 
280, 281, 287, Plate 1215; var. 
sericea, 287 

Contributions to the Flora of Nova 
Scotia: V. Results of Explora- 
tion in Cumberland County, 301 

Contributions to the Flora of South- 
ern Illinois, 319 

Convolvulus sepium var. repens, 
315 

Conyza canadensis 279, 282, var. 
pusilla, 288 

Cooley, G. R., The Vegetation of 
Sanibel Island, Lee County, 
Florida, 269 

Coreopsis leavenworthii, 279, 289 

Cornus alternifolia, 322 

Cronquist, A., A New Variety of 
Solidago ulmifolia, 36 

Crooked Oaks on Cape Cod and 
Martha's Vineyard, 132 

Crotalaria incana, 286; linaria, 286; 
rotundifolia, 286; sagittalis, 267; 
striata, 279, 286 

Croton floridanus, 286 

Cryptostegia grandiflora, 287 

Cumberland County, Contributions 
to the Flora of Nova Scotia: 
V. Results of Exploration in 301 

Curtia gentianoides, 104 

Cuscuta campestris, 287 

Cynanchum palustre, 287; scopa- 
rium, 287 

Cynoctonum mitreola, 287 

Cynodon dactylon, 283 

Cyperus globulosus, 284; ligularis, 
284; martindalei, 284; planifolius, 
284; polystachyos var. texensis, 
284; pseudovegetus, 284; rotun- 
dus, 284; strigosus, 284 

Cystopteris bulbifera, 224, 228, 229, 
230, 236, 239; dickieana, 224, 


342 Rhodora 


228, 229, 231, 236, 239; fragilis, 
224, 228, 229, 230, 231, 236, 239, 
f. simulans, 228, var. simulans, 
220, 224, 228, 229, 230, 236, 239, 
var. tennesseensis, 224, 229, 230. 
236, 239; tennesseensis, 228 
Cytotaxonomie Observations on 
North American Ferns, 219 


Dactyloctenium aegyptium, 283 

Dalbergia ecastophyllum, 281, 286 

Dalea aurea, 291; enneandra, 291, 
293; laxiflora, 293 

Dark-hooded Variant of Asclepias 
amplexicaulis, A, 336 

Davies, P. A., Distribution and 
Abundance of Shortia galacifolia, 
189 

Dennstaedtia punctiloba, 236 

Dentaria anomala, 162; bulbifera, 
167, 168; californica, 167; con- 
catenata, 163; diphylla, 161, 164, 
165, 166, 167, Plate 1206, 168, 
169, 171, 172; X maxima, 161; 
dissecta, 165; glandulosa, 169; 
heterophylla, 161, 164, 165, 166, 
167, Plate 1206, 168, 171, 172; 
incisifolia, 161; laciniata, 161, 
163, 164, 165, 166, 167, Plate 
1206, 168, 169, 171, 172, var. 
alterna, 163, var. coalescens, 164, 
var. furcata, 164, var. integra, 
164, var. lasiocarpa, 164, var. 
latifolia, 164, var. opposita, 163; 
X diphylla, 161; heterophylla, 
162; X maxima, 161; macrophyl- 
la, 167; maxima, 161, 163, 164, 
165, 166, 167, Plate 1206, 168, 
169, 171, 172; multifida, 161, 164, 
165, 166, 167, Plate 1206, 168, 
171, 172; X heterophylla, 162; 
pentaphylla, 167; pinnata, 167; 
polyphylla, 167; savensis, 167 

Dentaria in Eastern North Ameriea, 
Preliminary Studies in the Genus, 
161 

Desmodium glutinosum f. chan- 
donnettii, 268; tortuosum, 286 

DeWolf, Jr., G. P., A Note on the 
Name Calamintha, 73 

Dexter, R. W., Crooked Oaks on 
Cape Cod and Martha’s Vine- 
yard, 132 


[Vor. 57 


Dichromena colorata, 284 

Dicliptera assurgens, 288 

Digitaria sanguinalis, 283 

Diplazium pyenocarpon, 234 

Distichlis spicata, 283 

Distribution and Abundance of 
Shortia galacifolia, 189 

Dodecatheon frenchii, 325, 326, 327, 
328, 329, 330, 331; meadia, 325, 
326, 327, 328, 329, 330, 331; var. 
brachyearpus f. pallidum, 315 

Dodonaea jamaicensis, 277, 286 

Draba arabisans, 301, 302, 303, 307; 
verna var. Boerhaavia, 313 

Drosera anglica, Evidence for the 
Hybrid Origin of, 105 

Drosera angliea, 106, 107, 108, 111, 
115, 117, 121, 122, 123, 124, 125, 
126, 127, 128, 129; binata, 116; 
brevifolia, 110, 115, 116, 118; 
Burmanni, 116; capensis, 117; 
capillaris, 110, 115, 116, 118; 
cistiflora, 116; filiformis, 115, 116, 
var. Tracyi, 115, 116; intermedia, 
107, 110, 111, 115, 116; linearis, 
106, 107, 108, 109, 110, 111, 112, 
113, 114, 115, 117, 118, 119, 121, 
122, 123, 124, 125, 126, 127, 128, 
129; longifolia, 106; X obovata, 
106, 117, 127, 128; pygmaea, 116; 
regia, 116; rotundifolia, 106, 107, 
108, 109, 110, 111, 112, 113, 114, 
115, 117, 118, 119, 120, 121, 122, 
123, 124, 125, 126, 127, 128, 129, 
var. comosa, 120; spathulata, 117 

Dryopteris boottii, 236; X bootii, 
238; clintoniana, 236; cristata, 
236; fragrans var. remotiuscula, 
303; fructuosa, 236; goldiana, 
236; intermedia, 236; marginalis, 
236; spinulosa, 219, 236, 237; 


Echinodorus Andrieuxii, 133, 136, 
142, 150, 175, 176, 177, 211; 
Berteroi, 136, 139, 146, 147, 
148, 152, 154; var. Berteroi, 140, 
141, 142, 144, 145, 150, 210, 211, 
var. lanceolatus, 140, 142, 144, 
145, 150, 210; bracteatus, 136, 
142, 150, 153, 154, 155; var. 
bracteatus, 174, 211, 212, var. 
efenestratus, 174, 212; brevi- 
pedicellatus, 182; cordifolius, 134, 


1955] Index to Volume 57 343 


135, 137, 138, 140, 144, 147, 148, 
150, 154, 155, 175, 210, f. lan- 
ceolatus, 144; var. lanceolatus, 
144, 8 Berteroanus, 140; ellipti- 
eus, 177, 179, f. ovata, 175, var. 
ovatus, 175, y ovata, 175, 176; 
floribundus, 148; fluitans, 136, 
155, 212; grandiflorus, 136, 142, 
148, 150, 152, 153, 154, 179, « 
floribundus, 148; var. aureus, 
148, 152, 210; var. floribundus, 
148; var. grandiflorus, 148, 149, 
210, 211, 212; var. longiscapus, 
148; var. ovatus, 148, 152; 
irisebachii, 136, 142, 150, 154, 
180, 181, 210, 211; intermedius, 
179, 180, 181, 182; isthmicus, 
137, 203, 204, 205, 209, 211; 
longipetalus, 181, 182; longi- 
scapus, 152; macrophyllus, 137, 
150, 153, 154; 8 muricatus, 148, 
152, 153; magdalenensis, 137, 
203, 204, 205, 207, 209, 212; 
Martii, 179; muricatus, 136, 142, 
148, 150, 152, 153, 155, 211, 212; 
nymphaeifolius, 134, 136, 150, 
154, 182, 183, 210, 211; ovalis, 
136, 142, 150, 154, 174, 175, 210; 
palacfolius, 177; paniculatus, 136, 
142, 150, 177, 178, 179, 212; var. 
dubius, 179; var. paniculatus, 
177; parvulus, 134, 146, 185; 
quadricostatus, 137, 203, 204, 
205, 207, 209; radicans, 134, 137, 
140; rostratus, 140, 144, 146; f. 
lanceolatus, 144; var. lanceolatus, 
144, 147; Sellowianus, 152; sub- 
alatus, 179, 185; tenellus, 134, 
137, 183, 186, 188, 202, 204, 206, 
209, var. ecostatus, 183, 184, 
185, 205, 207, 209, 211, 212, var. 
latifolius, 183, 188, 202, 203, 
205, 206, 207, 209, 210, 211, 212, 
var. latifolius f. apanecae, 203, 
var. parvulus, 183, 185, 187, 
205, 206, 209, 210, 211; var. 
parvulus f. Randii, 188, var. 
tenellus, 183, 184, 185, 205, 206, 
207, 209, 212; trialatus, 136, 
142, 150, 154, 179, 180, 212; 
tunicatus, 136, 150, 154, 181, 182, 
211; virgatus, 135, 137, 142, 150, 
211 


Echinodorus in the American Trop- 
ics, 133, 174, 202 

Elaeagnus argentea, 290 

Elatine minima, 308 

Eleocharis atropurpurea, 284; geni- 
culata, 284; nitida, 305; ovata 
var. Heuseri, 305, var. ovata, 
305 

Eleusine indica, 283 

Elymus interruptus, 266; riparius, 
131 

Emilia sonchifolia, 289 

Epidendrum tampense, 277, 285 

Epifagus virginiana, 71, 72 

Epipactis Helleborine in Illinois, 
131 

Equisetum variegatum, 114 

Eragrostis ciliaris, 283; elliottii, 
283; pilosa, 283; tracyi, 275, 282, 
283 

Erechtites hieracifolia, 289 

Erigeron quercifolius, 289 

Eriochloa contracta, 159 

Eriogonum flavum, 291 

Ernodia littoralis, 279, 288 

Erskine, D., Northeastward Range 
Extensions for Two Species of 
Hypericum in Nova Scotia, 132 

Eryngium baldwinii, 287; yucci- 
folium, 266 

Erythrina herbacea, 286 

Eugenia anthera, 277, 287; axil- 
laris, 277, 279, 287; buxifolia, 
287; myrtoides, 277 


Eupatorium eapillifolium 289; 
mikanioides, 289 
Euphorbia ammannioides 286; 


buxifolia, 281, 286; Geyeri, 314; 
heterophylla, 286; hirta, 286; 
hyssopifolia, 286; maculata, 286; 
pilulifera, 286; trichotoma, 286; 
vermiculata, 268 

Eustoma exaltatum, 287; maculata, 
58 


Fassett, N. C., Echinodorus in the 
American Tropies, 133, 174, 202 

Ferns, Cytotaxonomic Observations 
on North American, 219 

Festuca obtusa, 302, 304; rubra, 302 

Ficus aurea, 276, 277, 285 

Filipendula rubra, 267 

Fimbristylis castanea, 278, 284 


344 Rhodora 


Flaveria floridana, 279, 289; linearis, 
278, 279, 289 

Florida, The Vegetation of Sanibel 
Island, Lee County, 269 

Forestiera porulosa, 277, 279, 287 

Fragaria vesca f. alba, 266 

Frasera caroliniensis, 10; verticil- 
lata, 10 

Fraser's Catalogue, Non-validity of 
Nuttallian Names in, 290 

Fraxinus nigra, 131; tomentosa, 315 

French’s Shooting Star in Southern 
Illinois, 325 

Froelichia floridana, 285 


Galactia parvifolia, 286; volubilis, 
286 

Galium boreale var. intermedium, 
309; circaezans, 317, var. hypo- 
malacum, 317; hispidulum, 288; 
tinetorium var. floridanum, 288; 
triflorum 317; X G. cireaezans 
var. hypomalacum, 317 

Gaura angustifolia, 279, var. 
eatonii, 287; coccinea, 290 

Gaylussacia baceata, 315 

Genipa clusiaefolia, 288 

Gentiana, 2; calycina, 79, 81; Cen- 
taurium, 103; procera, 114 

Geocaulon lividum, 307 

Geranium Bicknellii, 308; dissec- 
tum, 314; molle, 314 

Gerardia maritima, 288 

Geum virginianum, 322 

Gleason, H. A., Pedanticism Runs 
Amuck, 332 

Glechoma hederacea, 74 

Glycyrrhiza lepidota, 291, 293 

Gnaphalium purpureum, 289 

Gossypium hirsutum, 278, 286 


Hardin, J. W., Studies in the 
Hippocastanaceae, I. Variation 
in the Mature Fruit of Aesculus, 
37 

Heather, Scotch, 337 

Helianthium, 134, 135, 136; nym- 
phaeifolium, 182; parvulum, 185; 
tenellum, 185 

Helianthus annuus var. nanus, 318 

Heliotropium curassavicum, 282, 
288; parviflorum, 282, 288; poly- 
phyllum, 282, 288 


[Vor. 57 


Hemicarpha micrantha, 321, 322 

Hepatica acutiloba, 132, 265; amer- 
icana, 132, 265 

Heracleum maximum, 302 

Herbarium Specimens, The Archer 
Method for Mounting, 294 

Hermann, F. J., Two New Carices 
from Southeastern United States, 
156 

Hexalectris spicata, 321 

Hieracium vulgatum, 268 

Hippocastanaceae, I. Variation 
within the Mature Fruit of Aescu- 
lus. Studies in the, 37 

Hodgdon, A. R., Ilex glabra and a 
New Station for Kalmia latifolia 
in New Hampshire, 34 

Houstonia nigricans, 288; procum- 
bens, 288; pusilla, 317; f. albi- 
flora, 317 

Hybrid Origin of Drosera Anglica, 
Evidence for, 105 

Hydrocotyle umbellata, 287 

Hydrophyllum virginianum, 131 

Hymenocallis keyensis, 285 

Hypericum boreale, 132; X cana- 
dense, 132; canadense, 132; dis- 
simulatum, 132; Kalmianum, 
114; mutilum var. parviflorum, 
132 

Hypericum in Nova Scotia, North- 
eastward Range Extensions for 
Two Species of, 132 

Hyptis capitata, 75; radiata, 75 

Hyssopus anethiodorus, 291 

Hystrix patula var. Bigeloviana, 
311 


Ilex glabra and a New Station for 
Kalmia latifolia in New Hamp- 
shire, 34 

Ilex glabra, 34; opaca, 159, 315 

Illinois and to the Chicago Region, 
Plants New to, 265; Contribu- 
tions to the Flora of Southern, 
319; Epipactis Helleborine in, 
131; Illinois Flora: Recent Addi- 
tions, Southern, 159; French’s 
Shooting Star in Southern, 325 

Impatiens pallida, 308 

Imshaug, H. A., An Additional 
Species of the Lichen Genus 
Buellia from the West Indies, 323 


1955] Index to Volume 57 345 


Ipomoea cathartica, 288; pes-cap- 
rae, 280, 281, 288; sagittata, 287 

Iresine celosia, 285 

Isanthus brachiatus, 320 

Iva frutescens, 289; imbricata, 281, 
289 


Jacquinia keyensis, 277, 279, 287 

Jeffersonia diphylla, 267 

Juncus balticus var. littoralis, 114; 
megacephalus, 284; vaseyi, 306 

Juniperus horizontalis, 113 

Jussiaea uruguayensis, 315 


Kalmia latifolia, 34, 35, 36 

Kalmia latifolia in New Hampshire, 
Ilex glabra and a New Station 
for, 34 

Kosteletzkya virginica, 278, 286; 
virginica var. althaeifolia, 286 

Kuhnia eupatorioides, 321 


Lachnocaulon anceps, 284 

Lactuca integrifolia, 291; oblongi- 
folia, 291; pulchella, 291 

Laguncularia racemosa, 276, 279, 
287 

Lantana camara, 288; involucrata, 
279, 288; ovatifolia, 279, 288 

Lapithea Boykinii, 97; capitata, 97; 
gentianoides, 100 

Larix laricina, 113 

Lepidium virginicum, 285 

Leptochloa dubia, 283 

Lespedeza leptostachya, 268 

Lesquerella auriculata, 241, 242, 
243, 253, 254, 255, 260, 261, 262, 
264; densipila, 241, 242, 243, 
Plate 1208, 249, 250, 251, 254, 
250, 258, 260, var. maxima, 251, 
252, 256; gracilis, 242; grandi- 
flora, 241, 242, 243, 254, Plate 
1212, 261, 262, 263, 264; lasio- 
carpa, 241, 243, 245, 246, 247, 
var. ampla, 244, 245, 246, var. 
Berlandieri, 244, 245, 246, var. 
heterochroma, 244, 245, 240, 
var. lasiocarpa, 244, 246; Les- 
curii, 241, 243, 247, 248, Plate 
1207, 249, 250, 251; lyrata, 242, 
243, 252, Plate 1209, 254; per- 
forata, 241, 243, 248, 249, 257, 
258, 259, Plate 1211, 260; ston- 


ensis, 243, 251, Plate 1210, 255, 
256, 258 

Lesquerella (Cruciferae), The Auric- 
ulate-leaved Species of, 241 

Liatris cylindracea, 322; scabra, 318 

Lichen Genus Buellia from the 
West Indies, An Additional 
Species of the, 323 

Ligustrum ovalifolium, 287 

Lilium andinum, 291; philadelphi- 
cum, 113 

Lindera melissaefolium, 312 

Lindernia dubia, 309 

Linnea borealis var. americana, 113 

Liparis liliifolia, 131; Loeselii. 307 

Lippia nodiflora, 288 

Liquidambar Styraciflua, 194, 319 

Liquiritia lepidota, 291 

Liriodendron Tulipifera, 194 

Lithospermum canescens f. palli- 
dum, 316; caroliniense, 316; 
croceum, 316; latifolium, 131 

Littorella americana, 309 

Lobelia Cardinalis f. rosea, 317; 
feayana, 288; Kalmii, 113, 114; 
spicata, 309 

Lochnera rosea, 279, 282, 287 

Lolium multiflorum var. diminu- 
tum, 311 

Lophotocarpus calycinus f. maxima, 
321 

Ludwigia microcarpa, 287; palus- 
tris, 309 

Luzula luzuloides, 267; parviflora 
var. melanocarpa, 306 

Lycopodium Selago, 301, 303 

Lygodium palmatum, 220 

Lyonia ligustrina, 310 

Lythrum lanceolatum, 287 


Magnolia cordata, 189 
Malaxis brachypoda, 306 
Malva coccinea, 291, 293 
Mamillaria vivipara, 293 
Martha’s Vineyard, Crooked Oaks 
on Cape Cod and, 132 
Maytenus phyllanthoides, 286 
Megalodonta Beckii, 309 
Melanthera deltoidea, 278, 289 
Melissa, 77 
Melothria pendula, 278, 288 
Mentzelia floridana, 287 
Mikania batatifolia, 278, 289 


346 Rhodora 


Milium effusum, 302, 305 

Missouri, Plants New to, 310 

Mitella diphylla, 265 

Mohlenbrock, R. H., Contributions 
to the Flora of Southern Illinois, 
319 

Monarda fistulosa, 320 

Montgomery, F. H., Preliminary 
Studies in the Genus Dentaria in 
Eastern North America, 161 

Montia lamprosperma, 303, 307 

Muhlenbergia asperifolia, — 267; 
capillaris, 283; mexicana, 302, 
304, f. ambigua, 311 

Myosotis scorpioides, 315 

Myrica cerifera, 279, 285; Gale, 113 

Myriophyllum brasiliense, 315; 
tenellum, 308 


Najas flexilis, 303; guadalupensis, 
283 

Nephrolepis exaltata, 283 

Neptunia lutea, 286 

Neurola, 4, 43; arkanzica, 44, 46 

New Hampshire, Ilex glabra and a 
New Station for Kalmia latifolia 
in, 34 

New Variety of Solidago ulmifolia, 
A, 36 

Non-validity of Nuttallian Names 
in Fraser’s Catalogue, 290 

North American Ferns, Cytotaxo- 
nomic Observations on, 219 

North American Genus Sabatia 
(Gentianaceae), A Revision of 
the, 1, 43, 78 

North America, Preliminary Studies 
in the Genus Dentaria in Eastern, 
161 

Northeastward Range Extensions 
for Two Species of Hypericum 
in Nova Scotia, 132 

Note on the Name Calamintha, A, 
73 

Nova Scotia, Northeastward Range 
Extensions for Two Species of 
Hypericum in, 132 

Nova Scotia: V. Results of Explor- 
ation in Cumberland County, 
Contributions to the Flora of, 301 

Nymphaea odorata, 267, f. rosea, 
313 


[Vor. 57 


Oenothera alata, 290, 291; albi- 
eaulis, 291; biennis, 302; caespi- 
tosa, 290, 291; humifusa, 280, 
281, 287; laciniata, 287; macro- 
carpa, 290, 291; serrulata, 290 

Onoclea sensibilis, 219, 224, 227, 
229, 236, 239 

Onosmodium hispidissimum, 322 

Opuntia austrina, 279, 287; dil- 
lenii, 287; ficus-indica, 287; humi- 
fusa, 320; keyensis, 287 

Osmunda cinnamomea, 236; regalis 
var. spectabilis, 236 

Ostrya virginiana, 131 

Oxalis dillenii, 286 

Oxytropis johannensis, 302, 308 


Palmer, E. J. and Steyermark, 
J. A., Plants New to Missouri, 310 

Palmer, 8. C., Epifagus virginiana, 
71 

Panax quinquefolius, 131 

Panicum adspersum, 283; agros- 
toides, 284; albomarginatum, 284; 
amarulum, 284; bartowense, 284; 
consanguineum, 311; dichotomi- 
florum, 284; lanuginosum var. 
implicatum, 321; miliaceum, 267; 
neuranthum, 284; virgatum, 284 

Parietaria nummularia, 285 

Parker, 8. E., Scotch Heather, 337 

Parnassia glauca, 114 

Parthenocissus quinquefolia, 276, 
277, 286 

Paspalum ciliatifolium, 284; laeve, 
311; vaginatum, 284 

Pectis linearifolia, 289; prostrata, 
289 

Pedanticism Runs Amuck, 332 

Pedicularis canadensis, 319 

Pellaea atropurpurea, 236; glabella, 
236 

Pennisetum glaucum, 284 

Penstemon Bradburii, 293; crista- 
tum, 291; erianthera, 291; grandi- 
florus, 291, 293 

Petalostemum candidum, 321; pur- 
pureum, 321 

Phalangium esculentum, 291 

Phalaris arundinacea, 315 

Philoxerus vermicularis, 285 

Phlebodium aureum, 276, 277, 283 


1955] Index to Volume 57 347 


Phlox divaricata var. Laphamii f. 
candida, 316 

Phyllanthus abnormis, 286 

Phyllitis scolopendrium, 237, var. 
americana, 221, 237 

Physalis elliottii, 279, 288; pu- 
bescens, 288 

Phytolaeca americana, 285; rigida, 
277 

Pilea pumila, 313 

Pinus canadensis, 334, 335; Doug- 
lasii, 333; echinata, 319; elliottii 
var. densa, 278, 283; mucronata, 
333; resinosa, 299; rigida, 194, 
299; Strobus, 34, 113, 194, 299; 
taxifolia, 333, 334 

Pinus rigida Miller in Quebec, 299 

Piriqueta caroliniana, 278, var. 
glabra, 287 

Piscidea piscipula, 286 

Pithecellobium unguis-cati, 277, 286 

Pityrogramma triangularis, 220, 236 

Plantago juncoides var. decipiens, 
302 

Plants New to Illinois and to the 
Chicago Region, 265 

Plants New to Missouri, 310 

Pleienta, 4, 78; capitata, 97; dode- 
candra, 86; fasciculata, 103; 
flexuosa, 91; gentianoides, 100; 
leucantha, 86; quinquenervia, 
103; rigida, 103 

Pluchea purpurascens, 278, 289; 
rosea, 289 

Poa alsodes, 304; compressa, 302; 
glaucantha, 301, 303, 304; pra- 
tensis, 302 

Podophyllum peltatum f. aphyllum, 
267 

Pogonia ophioglossoides, 113 

Polygala incarnata, 286 

Polygonatum canaliculatum, 131 

Polygonum hydropiperoides var. 
hydropiperoides, 307; tenue, 320 

Polypodium  polypodioides, 283; 
virginianum, 237 

Polypremum procumbens, 287 

Polystichum acrostichoides, 236 

Populus nigra var. italica, 313 

Portulaca oleracea, 285; phaeo- 
sperma, 285; pilosa, 285 

Potentilla fruticosa, 302, 308; para- 
doxa, 322 


Preliminary Studies in the Genus 
Dentaria in Eastern North Amer- 
ica, 161 

Primula intercedens, 113; kewensis, 
106 

Pseudotsuga mucronata, 333, 335; 
taxifolia, 332, 333 

Psidium guajava, 287 

Psilotum nudum, 276, 283 

Psoralea esculenta, 291, 293; in- 
cana, 291 

Psychotria nervosa, 277; undata, 
288 

Pteretis pensylvanica, 219, 227, 236, 
239 

Pteridium aquilinum, 228, var. 
caudatum, 283, var. latiusculum, 
228, 236, var. pseudocaudatum, 
224, 228, 229, 236, 239, 283 

Ptilimnium capillaceum, 287, 320; 
costatum, 320; nuttallii, 320 

Pycnanthemum incanum, 75 

Pyrus melanocarpa, 314, 315 


Quebec, Pinus rigida Miller in, 299 
Quercus rubra, 131; virginiana, 278, 
285 


Randia aculeata, 277, 288 

Ranunculus harveyi, 319; parvi- 
florus, 320 

Rapanea guianensis, 277, 287 

Ratibida columnaris, 293; columni- 
fera, 293 

Revision of the North American 
Genus Sabatia (Gentianaceae), 
A, 1, 43, 78 

Rhabdadenia biflora, 287 

Rhamnus caroliniana, 319 

Rhexia stricta, 103 

Rhizophora mangle, 276, 279, 287 

Rhododendron maximum, 193 

Rhus radicans, 276, 277, 278, 282, 
286 

Rhynchelytrum roseum, 284 

Rhynchosia lewtonii, 286; mi- 
chauxii, 278, 286; minima, 286 

Rhynchospora capillacea, 113; capi- 
tellata, 305; fusca, 305 

Ricinus communis, 286 

Rinodina antillarum, 323, 324; 
boergesenii, 323, 324; pyxinoides, 
323, 324 


348 Rhodora 


Rivina humilis, 277, 285 

Rollins, R. C., The Archer Method 
for Mounting Herbarium Speci- 
mens, 294; The Auriculate-leaved 
Species of Lesquerella (Cruci- 
ferae), 241 

Rosa micrantha, 314 

Rouleau, E., Pinus rigida Miller in 
Quebec, 299 

Rubus allegheniensis var. plausus, 
314; laciniatus, 267 

Rudbeckia columnifera, 291; pur- 
purea var. serotina, 290, 291 

Ruellia caroliniensis, 320 

Rumex acetosella f. integrifolius, 
265; maritimus var. fueginus, 
322 


Sabal palmetto, 275, 276, 278, Plate 
1215, 284 

Sabatia  alainii, 67; amoena f. 
albiflora, 58; angularis, 18, 20, 21, 
30, 31, 46; f. cleistantha, 19, 22; 
arenicola, 43, 44, 47, 49, 50, 51; 
bartramii, 79, 91, 95, 98, 103; 
Boykinii, 99; brachiata, 18, 20, 
27, 29, 30, 31, 32, 47, 50; f. 
candida, 23, 27, 30; brevifolia, 52, 
65, 68; calycina, 65, 78, 79, 93, 
103; campanulata, 50, 51, 52, 57, 
60, 61, 66, 81, 103, 287, var. 
gracilis, 57, var. grandiflora, 64; 
campestris, 20, 31, 32, 43, 44, 46, 
47, 49, 50, 51; capitata, 97, 98, 
101; carnosa, 50, 51; chloroides, 
83; concinna, 31, 32, 47; corym- 
bosa, 10, 26, 70; cubensis, 80; 
decandra, 83, 93, 94; difformis, 6, 
7, 8, 9, 10, 11, 12, 16, 21, 23, 24, 
25, 26, 27, 93; dodecandra, 5, 78, 
79, 82, 88, 84, 85, 86, 89, 90, 94, 
var. dodecandra, 86, 98, var. 
foliosa, 86, 87, 93, 98; elliottii, 
10, 25, 26, 68, 70; foliosa, 83, 84, 
85; formosa, 47; gentianoides, 3, 
94, 96, 97, 98, 100, 101; gracilis, 
54, 57, 60, 66, 81, 103, var. 
cubensis, 80; grandiflora, 52, 60, 
64, 65, 66, 67; harperi, 84, 85; 
kennedyana, 79, 88, 89, 90, 94, 
98, f. candida, 88, f. encycla, 88; 
lanceolata, 11; macrophylla, 7, 8, 
13, 14, 15, 16, 21, var. macro- 


[Vor. 57 


phylla, 17, 21, var. recurvans, 
17, 21; maculata, 62; nana, 61; 
nervosa, 44; obtusata, 84, 85, 87; 
oligophylla, 101; palmeri, 62; 
paniculata, 9, 24, 25, 26, 27, 30, 
69, 70, 8 Elliottii, 68, var. angusti- 
folia, 27, var. latifolia, 27; pur- 
pusii, 62, 63; quadrangula, 9, 
11, 18, 21, 22, 24, 25, 26, 27, 30; 
recurvans, 14, 15, 16; simulata, 
61; stellaris, 52, 56, 58, 59, 60, 61, 
62, 63, 65, 66, 67, 93, 103 

Sabatia (Gentianaceae), A Revision 
of the North American Genus, 
1, 43, 78 

Sabbatia Alainii, 64; albeola, 103; 
amena, 103; amoena, 58, 103; 
anceps, 103; angularis, 18, f. 
albiflora, 19, f. cleistantha, 19, 
var. albiflora, 19, var. elatior, 19, 
var. latifolia, 19, var. pauciflora, 
19; angustifolia, 29; australis, 
103; Boykinii, 97; brevifolia, 70; 
calycosa, 79; campanulata f. 
albina, 52, var. amoena, 58, var. 
gracilis, 52; campestris f. albi- 
flora, 44; centaurium, 103; chlo- 
roides, 86; chloroides 8 stricta, 91, 
var. coriacea, 91, var. erecta, 91, 
var. flexuosa, 91; concinna, 29; 
corymbosa, 7, 12, 70, var. angusti- 
folia, 7, 12; cubensis, 79; cymosa, 
7, 22; decandra, 91, 103; de- 
clinata, 103; dichotoma, 79; dif- 
formis, 7, 12; dodecandra, 86; 
dodecandra stricta, 91; elliotti, 
70; foliosa, 87; formosa, 44, 47; 
gentianoides 8  Boykinii, 97; 
gracilis, 52, 57, 6 stellaris, 58, 
var. cubensis, 79, var. grandi- 
flora, 64; grandifolia, 67; Harperi, 
87; lanceol., 7; lanceolata, 7, 12; 
lingulata, 103; maculata, 58; 
maritima, 58; nana, 58; nervosa, 
46; nivea, 26, 103; obtusifolia, 
103; oligophylla, 100; Palmeri, 
58; paniculata, 7, 23, 69, 70, a 
latifolia, 7, 8 angustifolia, 7; 
petiolata, 103; pulchella, 103; 
pumila, 103; Purpusii, 58; quad- 
rangula, 103; recurvans, 17; sim- 
plex, 103; simulata, 58; stellaris, 
56, 103, f. albiflora, 58, 8 pumila, 


1955] Index to Volume 57 349 


58; stricta, 103; tenuifolia, 103; 
Tracyi, 52, 56, 57; umbellata, 
104; verticillaris, 104 

Sagittaria graminea, 304; radicans, 
137 

Salicornia perennis, 285 

Salix interior, 114; longipes, 285 

Sambucus simpsonii, 288 

Samolus ebracteatus, 278, 287; 
floribundus, 287 

Sanibel Island, Lee County, Florida, 
'The Vegetation of, 269 

Sansevieria guineensis, 284 

Sarcostemma clausum, 287 

Sarracenia purpurea, 113, 114, var. 
heterophylla, 308 

Satureja Acinos, 73; arkansana, 73; 
Calamintha, 73; glabella, 73, var. 
angustifolia, 73; hortensis, 73; 
vulgare, 73 

Saxifraga Aizoón, 301, var. neogaea, 
301, 302, 308 

Scaevola plumieri, 279, 280, 281, 
288 ' 

Schinus terebinthifolius, 286 

Schizachne purpurascens, 304 

Schizaea pusilla, 220 

Schofield, W. B., Contributions to 
the Flora of Nova Scotia: V. 
Results of Exploration in Cum- 
berland County, 301 

Schuebleria stricta, 104 

Scirpus americanus, 114, 284; ces- 
pitosus var. callosus, 305; cyperi- 
nus var. eriophorum, 284; hud- 
sonianus, 113, 305; longii, 90; 
polyphyllus, 311 

Scoparia dulcis, 288 

Scotch Heather, 337 

Scutellaria ovata, 320 

Sedum Rosea, 302; sarmentosum, 
313 

Serenoa repens, 284 

Sesbania aculeata, 215; aegyptiaca, 
215; australis, 214; bispinosa, 
215, 216; drummondii, 215, 216, 
217; exaltata, 215, 216, 217; 
grandiflora, 215, 216; longifolia, 
214; macrocarpa, 215, 286; mar- 
ginata, 215; punctata, 215; pu- 
nicea, 215; sericea, 215; sesban, 
215, 216; speciosa, 215; tetrap- 
tera, 215; vesicaria, 215, 216, 217 


Sesbania (Leguminosae): Evidence 
for a Conservative Treatment, 
Chromosome Numbers in the 
Genus, 213 

Seseli divaricatum, 291; lucidum, 
291 

Sesuvium maritimum, 285; por- 
tulacastrum, 278, 281, Plate 1219, 
285 

Setaria geniculata, 284; glauca, 284; 
macrosperma, 284 

Shepherdia canadensis, 113 

Shinners, L. H., Non-validity of 
Nuttalian Names in Fraser’s 
Catalogue, 290 

Shooting Star in Southern Illinois, 
French’s, 325 

Shortia galacifolia, 189, 190, 191, 
192, 193, 194, 195, 196, 197, 198, 
199, 200, 201 

Shortia galacifolia, Distribution and 
Abundance of, 189 

Sida carpinifolia, 279, 287 

Sideranthus integrifolius, 291; pin- 
natifidus, 291 

Sideroxylon foetidissimum, 277, 287 

Silphium laciniatum, 266; tere- 
binthinaceum, 159, 160, 266, var. 
Lucy-Brauniae, 160 

Sisymbrium loeselii, 267 

Sisyrinchium atlanticum, 285 

Smilacina racemosa, 131 

Smilax auriculata, 284; ecirrhata, 
131 

Solanum gracile Plate 1221, 288 

Solidago bicolor, 302; buckleyi, 319; 
ohionis, 114; sempervirens var. 
mexicana, 281, 289; ulmifolia var. 
palmeri, 36; var. ulmifolia, 36 

Solidago ulmifolia, A New Variety 
of, 36 

Sonchus oleraceus, 289 

Sophora tomentosa, 279, 286 

Sorghum halepense, 284 

Sparganium minimum, 303 

Spartina bakeri, 278, 284; patens, 
281, 284; Townsendii, 105 

Specularia lamprosperma, 317; X 
S. leptocarpa, 317 

Sphaeralcea coccinea, 293 

Sphenopholis intermedia, 304 

Spigelia marilandica, 319 

Spiraea prunifolia, 314 


350 Rhodora 


Sporobolus domingensis, 278, 284; 
juncea, 284; vaginiflorus var. 
inequalis, 267; virginicus, 278, 
281, 282, 284 

Staphylea trifolia, 131 

Stellaria humifusa, 303, 307 

Steyermark, J. A. and Palmer, 
E. J., Plants New to Missouri, 310 

Steyermark, J. A. and Swink, F. A., 
Calycera balsamitaefolia in the 
United States, 72; Plants New 
to Illinois and to the Chicago 
Region, 265 

Steyermark, J. A., Epipactis Hel- 
leborine in Illinois, 131 

Stilingia aquatica, 286; sylvatica, 
286 

Studies in the Hippocastanaceae, I. 
Variation within the Mature 
Fruit of Aesculus, 37 

Suaeda linearis, 285 

Suriana maritima, 280, 281, Plate 
1220, 286 

Swayne, J. R., and Voigt, J. W., 
French's Shooting Star in South- 
ern Illinois, 325 

Swertia, 2; difformis, 7, 9, 10, 11, 25; 
elliottii, 10, 26; lanceolata, 10; 
quadrangula, 11 

Swink, F. A., and Steyermark, J. A., 
Calycera balsamitaefolia in the 
United States, 72; Plants New 
to Illinois and to the Chicago 
Region, 265 

Synthlipsis Berlandieri, 245, var. 
hispida, 244; heterochroma, 245, 
246 


Talinum calycinum, 319 

Tamarindus indica, 286 

Thelypteris dryopteris, 233; hex- 
agonoptera, 224, 229, 232, 236, 
239; noveboracensis, 224, 229, 
233, 236, 239; palustris var. 
pubescens, 224, 229, 232, 236; 
phegopteris, 232, 236, 239; ro- 
bertianum, 233 

Thuja occidentalis, 113 

Thunbergia fragrans, 288 

Thuraria herbacea, 291 

Tilia americana, 131 

Tillandsia balbisiana, 277, 284; 
circinata, 284; fasciculata, 277, 


[Vor. 57 


284; recurvata, 284; tenuifolia, 
277, 284; usneoides, 277, 284; 
utriculata, 277, 284 

Tragopogon mirus, 105; miscellus, 
105 

Trichachne insularis, 160 

Trichomanes boschianum, 220 

Trichostema suffrutescens, 279, 288 

Triglochin palustris, 114 

Triodia flava f. cuprea, 266 

Trillium cernuum, 265, var. ma- 
cranthum, 265; erectum, 265, 336, 
f. albiflorum, 306; grandiflorum, 
265; nivale, 315 

Triplasis purpurea, 284 

Trisetum spicatum var. pilosiglume, 
301, 302, 304 

Troximon glaucum, 291 

Tsuga canadensis, 194 

Turner, B. L., Chromosome Num- 
ber in the Genus Sesbania 
(Leguminosae): Evidence for a 
Conservative Treatment, 213 

Two New Carices from Southeast- 
ern United States, 156 

Typha angustifolia, 283; domin- 
gensis, 283 


Ulmus pumila, 313 

Uniola paniculata, 280, 281, Plate 
1220, 284 

United States, Calycera balsamitae- 
folia in the, 72; Two New Carices 
from Southeastern, 156 

Urechites lutea, 287 

Utricularia cornuta, 113; inter- 
media, 113 

Uttal, L. J., A Dark-hooded Variant 
of Asclepias amplexicaulis, 336 

Uvularia grandiflora, 131 


Vaccinium vacillans, 315 

Vegetation of Sanibel Island, Lee 
County, Florida, The, 269 

Verbena canadensis f. candidis- 
sima, 316; var. candidissima, 
316; scabra, 288 

Verbesina virginica var. laciniata, 
289 

Veronica arvensis, 321; didyma, 
317, 320; polita, 320 

Vesicaria auriculata, 260; brevi- 
styla, 262; grandifolia, 262, var. 


3 1753 00341 3793 


1955] Index to Volume 57 351 


pinnatifida, 262, 264; lasiocarpa, 
244 

Viburnum opulus, 268 

Vicia acutifolia, 286; Cracca, 315 

Vigna repens, 279, 286; sinensis, 266 

Virgilia grandiflora, 291 

Vitis campestris, 291; riparia var. 
syrticola, 266 

Vittaria lineata, 276, 277, 283 

Voigt, J. W., Southern Illinois 
Flora: Recent Additions, 159 

Voigt, J. W. and Swayne, J. R., 
French’s Shooting Star in South- 
ern Illinois 325 


Wagner, W. H. Jr., Cytotaxonomic 
Observations on North American 
Ferns, 219 

Waltheria americana, 279, 287 

West Indies, An Additional Species 


of the Lichen Genus Buellia 
from the, 323 

Wilbur, R. L., A Revision of the 
North American Genus Sabatia 
(Gentianaceae), 1, 43, 78 

Wood, C. E. Jr., Evidence for the 
Hybrid Origin of Drosera anglica, 
105 

Woodsia ilvensis, 236 

Woodwardia areolata, 225, 229, 
235, 237; chamissoi, 236; vir- 
ginica, 225, 229, 234, 235, 237 


Yucca aloifolia, 281, 285; angusti- 
folia, 293; glauca, 291, 293 


Zanthoxylum clava-herculis, 286; 
fagara, 277, 286 

Zeuxine strateumatica, 285 

Zygostigma australe, 103