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Rhodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL Associate Editors 
STUART KIMBALL HARRIS 


VOLUME 48 


1946 


The New England Botanical Club, Ine. 


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


9 1946 


Dodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL . Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. January, 1946. No. 565. 


CONTENTS: 
Some Mosses from Windsor, Nova Scotia. Herbert Habeeb. ... 
Cypripedium Calceolus, var. parviflorum. M. L. Fernald. ...... 


Contributions from the Gray Herbarium of Harvard University— 
No. CLX. Technical Studies on North American Plants. 
000 locutus CEST SERA QUAL U-. c1: ec E d 


The New England Botanical Club, Ine. 


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


RHODORA.—a monthly journal of botany, devoted primarily to the flora of the 
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EDIBLE WILD PLANTS OF EASTERN NORTH AMERICA 
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Gray Herbarium of Harvard University, Cambridge 38, Mass. 


Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. January, 1946. No. 565. 


SOME MOSSES FROM WINDSOR, NOVA SCOTIA 
HERBERT HABEEB 


Tut following is a list of noteworthy mosses gathered in the 
immediate vicinity of Windsor, N.S. The writer collected 250 
numbers of moss comprising 100 species. The collections were 
made in the spring of 1942. 

In company with Prof. J. S. Erskine, many an atiis free 
from duties was spent roaming the fields and woods neighboring 
on King’s Collegiate School at Windsor, N. S. Prof. Erskine 
devoted himself to observing birds and flowering plants, while the 
writer took on the mosses. 

The first noteworthy collector of Nova Scotia mosses was A. 
Menzies, who collected the type of Dicranum fulvum Hook. near 
Halifax; others were J. Fowler, A. H. MacKay, J. Macoun and 
G. E. Nichols. Of present day writers, E. B. Bartram! has 
published an article on Nova Scotia mosses; while Margaret S. 
Brown? has put out quite a complete list of Nova Scotia bryo- 
phytes. 

Of the mosses here listed many are new to Nova Scotia, and 
some may be said to be coastal plain elements. The nomencla- 
ture used in this article is according to Grout. The writer is 
grateful to Dr. A. J. Grout for the helping hand often lent in the 
determination of critical specimens. 


AULACOMNIUM PALUSTRE (Web. & Mohr) Schwaegr. var. IM- 
BRICATUM Bry. Eur. On ground, and humus. 

1 Bartram, E. B. Some Nova Scotia Mosses. RHODORA, 24: 121-124, 1922. 

2 Brown, M. S. Liverworts and Mosses of Nova Scotia. Proc. N. S. Inst. of Sc. 


19: 161-198. 1936. 
3 Grout, A. J. List of Mosses of North America North of Mexico. The Bryologist, 


43:117-131. 1940. 


2 Rhodora [JANUARY 


Plants referable to this variety are not uncommon in the 
Acadian region, as the writer has also a number of specimens from 
New Brunswick. Yet, it should be noted that some plants seem 
to posses leaves of both the variety and species. 


DICRANUM SCOPARIUM Hedw. var. ORTHOPHYLLUM Brid. On 
ground. ‘This is the form with the straight, strict leaves. 

HERUM FUSCESCENS Turn. Common on ground in the 
woods. 

DrrRICHUM PALLIDUM (Hedw.) Hampe. Two collections from 
Windsor, off the ground. 

PLEURIDIUM SUBULATUM (Hedw.) Lindb. On rather sterile 
ground. This moss was determined by Dr. Grout. 

GRIMMIA ALPICOLA Hedw. On rocks, and boulders. 


The writer has a suspicion that Grimmia alpicola is nothing 
more than a short-capsuled, muticous-leaved form of Grimmia 
apocarpa Hedw. 


AMBLYSTEGIUM JURATZKANUM Schimp. On a rock. 

BRACHYTHECIUM ACUTUM (Mitt.) Sull. On ground, and rotten 
wood. 

BracHYTHECIUM Besti Grout. On ground in woods. 


This moss is rather common in the Acadian region. 


BRACHYTHECIUM CAMPESTRE Bry. Eur. On ground. 

BRACHYTHECIUM FLEXICAULE Ren. & Card. On partly burnt 
wood, and ground. 

BRACHYTHECIUM POPULEUM (Hedw.) Bry. Eur. On rocks, 
ground. 


In both Nova Scotia and New Brunswick, Brachythecium 
populeum breaks down into two forms. One form is typical with 
a percurrent costa in the leaves, while the other has a costa ex- 
tending 24 the length of the leaf. 


BRACHYTHECIUM VELUTINUM (Hedw.) Bry. Eur. On ground, 
debris, and lower trunk of a tree. 

BryHniA Hurren Bartr. Was mixed with Brachythecium 
rivulare on debris in a damp situation. The determination was 
made by Dr. Grout, who mentions it as one of only a few collec- 
tions. 

AMBLYSTEGIUM VARIUM (Hedw.) Lindb. var. ovatum Grout. 
Concerning this specimen Dr. Grout wrote “costa even shorter 
than in type". , 

DREPANOCLADUS ADUNCUS (Hedw.) Warnst. var. KNEIFFI 
(Bry. Eur.) Warnst. Two collections were made from the water 


1946] Habeeb,—Some Mosses from Windsor, N. 8. 3 


of a swamp. The one of May, 1942, is a large form. The other of 
April, 1942, is a small form. 
DREPANOCLADUS ADUNCUS var. CAPILLIFOLIUS (Warnst.) 
Wynne. In water of a swamp. 
EURHYNCHIUM HIANS (Hedw.) J. & S. A half dozen numbers 
collected from the ground, rotten log in a swamp, wood debris, etc. 
L3 


A variable moss; a couple of specimens seem to approach the 
European Eurhynchium Swartz (Turn.) Hobk. in leaf outline. 


EURHYNCHIUM STRIGOSUM (Hoffm.) Bry. Eur. var. RoBUSTUM 
Roll. On ground. 

HYGROAMBLYSTEGIUM IRRIGUUM (Wils.) Loeske. Not uncom- 
mon on debris in damp situations and alongside of brooks. 

HYLOCOMIUM BREVIROSTRE (Beauv.) Bry. Eur. On rocks, 
brookside, and swampy ground. 

HYPNUM cuRVIFOLIUM Hedw. One collection made in April 
from the ground. 

LEPTODICTYUM RIPARIUM (Hedw.) Warnst. This is mixed 
with its forma longifolium (Schultz) Grout; from turf and sticks, 
edge of à wet situation. 

PLAGIOTHECIUM LAETUM Bry. Eur. On rocks, rotten wood. 

ORTHOTRICHUM SORDIDUM Lesq. & James. On tree trunks, 
ironwood and maple. 

ORTHOTRICHUM SPECIOSUM Nees. On old willow log. 


The writer has collected good Orthotrichum elegans Hook. & 
Grev. in New Brunswick, but this collection from Winsdor seems 
to be closer to O. speciosum in its larger size and shaggier ap- 
pearance. Some leaves are as long as 6 mm. 


PoLYTRICHUM COMMUNE Hedw. var. PERIGONIALE (Mx.) Bry. 
Eur. On ground, rotten logs, etc. 


More common than the species in the Acadian region. 


POLYTRICHUM JUNIPERINUM Hedw. var. ALPESTRE Bry. Eur. 
This form with the slender strict habit and shorter capsules is 
found around Windsor in boggy situations. 

PoLYTRICHUM FORMOSUM Hedw. On ground, May. 

AsrToOMUM MUHLENBERGIANUM (Sw.) Grout. On ground in 
rather sterile pasture. 

PHASCUM CUSPIDATUM [Schreb.] Hedw. On ground in sterile 
pasture. This is typical and not the var. americanum Ren. & 
Card. 

WEIsIA MicrosTomaA (Hedw.) C. Müell. On ground, sterile 
pasture. Weisia microstoma when in fruit is easily separated 
from Weisia viridula Hedw. 


4 Rhodora [JANUARY 


Astomum Muhlenbergianum, Phascum cuspidatum and Weisia 
microstoma were found growing in association with each other, 
and sometimes with Pottia truncata (Hedw.) Furnr. These three 
species may have a coastal-plain distribution, at least in eastern 
Canada. For the writer has not as yet found them in central 
New Brunswick. 


GRAND FALLS, 
New Brunswick 


CYPRIPEDIUM CALCEOLUS L., var. parviflorum (Salisb.), comb. 
nov. C. Calceolus, B. in part, L. Sp. Pl. 951 (1753). C. parvi- 
florum Salisb. in Trans. Linn. Soc. i. 77, t. 2, fig. 2 (1791). C. 
flavescens DC., B. C. parviflorum (Salisb.) DC. in Redouté, Lili- 
aceae, 1. pl. 20 (1802). C. luteum Raf., var. parviflorum (Salisb.) 
Raf. Med. Bot. i. 142 (1828). C. hirsutum Mill, var. parvi- 
florum (Salisb.) Rolfe in Orchid Rev. xv. 184 (1907). C. bulbo- 
sum Mill, var. parviflorum (Salisb.) Farwell in Fifteenth Rep. 
Mich. Acad. Sci. 170 (1913). Calceolus parviflorus (Salisb.) 
Nieuwl. in Am. Midl. Nat. iii. 118 (1913). 


The large-flowered Cypripedium Calceolus, var. pubescens 
(Willd.) Correll in Bot. Mus. Lfts. (Harvard), vii. 14 (1938) is 
relatively southern, characteristic of rich and dry to merely damp 
woodland from Georgia to Missouri, thence north to the Northern 
States, reaching its northeastern limit in central Maine. The 
smaller-flowered var. parviflorum, on the other hand, is relatively 
northern, extending from the southern part of the Labrador 
Peninsula to northern British Columbia, thence south to New- 
foundland, Prince Edward Island, New Brunswick, New England, 
New Jersey, upland of Maryland, mountain-region to Georgia 
and Tennessee, Ohio, Indiana, Illinois, Missouri, New Mexico, 
Utah and Washington, mostly in calcareous bogs or mossy 
swamps or mossy woods or on wet rocks, slopes or shores. The 
differences between the two are clearly brought out by Mr. 
Albert M. Fuller in his masterly Studies on the Flora of Wis- 
consin I. The Orchids, Orchidaceae—Bull. Pub. Mus. Milwau- 
kee, xiv!. 64-70 (1933), a treatment meriting careful considera- 
tion.—M. L. FERNALD. 


1946] Fernald,—Studies on North American Plants 5 


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


TECHNICAL STUDIES ON NORTH AMERICAN PLANTS 
M. L. FERNALD 


(Plates 993-1020)! 


I. Some SPECIES IN RAFINESQUE’S "HERBARIUM 
RAFINESQUIANUM” 


(PnATES 993 and 994) 


Dr. Merrill has asked me about the identities of some of the 
species published in the rare work of Rafinesque, his Herbarium 
Rafinesquianum (1833). Unlike too many of the publications 
of that highly variable and temperamental genius, this little book 
is carefully written, with logical discussions and with diagnoses of 
genera and species actually in hand, specimens of which were 
offered for sale. What a pity that we cannot now buy the series! 
Here are many well described novelties from many parts of 
North America, many of which have clear priority over the de- 
scriptions of others. For the most part their identification can 
safely be made only by those intimately familiar with the areas 
concerned: Texas, Oregon, etc.; but in checking on the region I 
best know it has been found that several of our long-familiar 
specific names must lapse, while some in other sections of the 
world are obviously later homonyms. In the following memo- 
randa only those names about which I feel no doubt are noted; 
others of them must be considered by specialists on other floras. 

SPIRANTHES LACERA (Raf.) Raf. Herb. Raf. 44 (1833). Neottia 
lacera Raf. Fl. Ludovic. 171 (1817), nomen, and in Am. Month. 
Mag. Crit. Rev. ii. 206 (1818) with full description. Neottia 
gracilis, var. B. secunda Bigel. Fl. Bost. ed. 2: 322 (1824). 

S. MONTANA Raf. Herb. Raf. 45 (1833). S. ovalis Lindl. Gen. 
Sp. Orch. Pl. 466 (1840). S. cernua, var. parviflora Chapm. FI. 
So. U. S., ed. 3: 448 (1897). Gyrostachys parviflora (Chapm.) 
Small, Fl. Se. U. S. 318 (1903). 5S. parviflora (Chapm.) Ames, 
Orchidaceae, fasc. 1: 137 (1905). Ibidium ovale (Lindl.) House 

1The cost of preparing and engraving the plates met in part through grants from 


the AMERICAN PHILOSOPHICAL Society and from the DEPARTMENT OF BIOLOGY OF 
HARVARD UNIVERSITY. 


6 Rhodora [JANUARY 


in Muhlenbergia, i. 128 (1906). J. parviflorum (Chapm.) Jen- 
nings in Ann. Carneg. Mus. iii. 485 (1906). Triorchis ovalis 
(Lindl) Nieuwland in Am. Midl. Nat. ii. 123 (1913). S. 
Smallii Schlechter in Beih. Bot. Centralbl. xxxvii?. 358 (1920). 

S. TUBEROSA Raf. Herb. Raf. 45 (1833). S. Beckii Lindl. 
Gen. Sp. Orch. Pl. 472 (1840), at least as to deser. S. simplex 
Gray, Man. ed. 5: 506 (1867), not Griseb. Gyrostachys simplex 
(Gray) Ktze. Rev. Gen. ii. 664 (1891). S. Gray? Ames in Rnuo- 
DORA, vi. 44 (1904). Gyrostachys Grayi (Ames) Britton, Man. 
ed. 2: 300 (1905). ZIbidium Beckii (Lindl.) House in Muhlen- 
bergia, i. 128 (1906). Gyrostachys Beckii (Lindl.) W. Stone, PI. 
So. N. J. 375 (1912). Triorchis Grayi (Ames) Nieuwland in Am. 
Midl. Nat. iii. 123 (1913). Triorchis Beckii (Lindl. House in 
Am. Midl. Nat. iv. 206 (1920). 


Unfortunately, the original description of Neottia lacera Raf. 
in Am. Month. Mag. Crit. Rev. ii. 206 (1818) did not get into 
Index Kewensis, although the other species, described in the 
same column of the identical page and directly preceding it, was 
there entered; and this first specific name of the pair, Neottia 
plantaginea Raf. l. c., was taken up by Torrey as the basis of 
Spiranthes plantaginea (Raf.) Torr. (1843) and, since Torrey had 
taken it up and it, therefore, got into reputable literature, it has 
been the nomenclatural basis of binomials by Britton, House and 
Nieuwland. But Neottia lacera, described with it and again in 
Herbarium Rafinesquianum, has been quite ignored, as have the 
other eight names under Spiranthes in the latter work. Never- 
theless Spiranthes lacera, based on Neottia lacera (1818), was 
several years earlier than Neottia gracilis Bigelow, Fl. Bost. ed. 
2: 322 (1824), the nomenclatural basis of S. gracilis (Bigelow) 
Beck, Bot. 333 (1833). Bigelow’s original account was as follows: 


*NEOTTIA GRACILIS. Slender Neottia. 

N. foliis radicalibus ovatis; scapo vaginato, floribus spiraliter secundis; 
labello obovato, crispo. 

Leaves radical, ovate; scape sheathing; flowers in a spiral row; lip 
obovate, curled. 

Root fascicled. Leaves radical, on short petioles, ovate, acute, nerved, 
caducous. Scape erect, slender, eight to twelve inches high with a few 
sheathing scales or leaf[ljlets. Flowers white in a twisted spike. Bractes 
closely applied to the germ, ovate, acuminate. Germs obovate. Petals 
linear, crystalline, parallel, the three upper ones cohering. Lip obovate- 
spatulate, curled, its base swelling with the lateral petals connected before 
it. Anther parallel to the style.—In dry, hilly woods.—July.—Perennial. 

The leaves falling off frequently cause the plant to appear leafless at 
the time of flowering. 


1946] Fernald,—Studies on North American Plants 7 


Variety 8. secunda. Spike unilateral, hardly twisted; flowers more 
slender. Perhaps a different species.—In Conway, New-Hampshire.— 
July. 


Rafinesque's original description in the American Monthly 
Magazine and Critical Review was 


24. Neottia lacera Raf. Smooth radical leaves oblong obtuse flat, 
scapes vaginated, sheaths acute: spike slender, flowers one sided spiral 
nodding, bracteas longer than the ovary, labellum canaliculated re- 
flexed obtuse laciniated.—Obs. Detected in 1816, in the swampy 
woods, near Glen’s Fall’s, Lake George, and the Luzerne mountains, 
blossoming in July and August, flowers white, scape slender about one 
foot high, root palmated. 


Rafinesque’s second account, in his Herbarium Rafinesquianum 
(1833), was briefer but contained the synonym “N fi. e. S]. 
gracilis, Beck, 1833", which nomenclaturally rested on Neottia 
gracilis Bigel. and, incidentally, indicated the priority in 1833 
of Beck over Herbarium Rafinesquianum. From this it would 
be natural to infer that Spiranthes gracilis must give way to S. 
lacera; but in this case Rafinesque ‘builded better than he knew", 
just as Jacob Bigelow did when he separated as “Perhaps a dif- 
ferent species” the more slender-flowered plant of the White 
Mountains. 

Spiranthes gracilis, as generally interpreted, consists of two 
quite different species: one relatively southern, the true S. 
gracilis, i. e. Neottia gracilis Bigelow; the other relatively north- 
ern, the Neottia lacera Raf. (1818), or N. gracilis, var. B. secunda 
Bigelow (1824), the S. lacera (Raf.) Raf. (1833). Study of all 
the material in the Gray Herbarium, the Ames Herbarium, the 
herbarium of the New England Botanical Club and that of the 
New York State Museum, which has been kindly placed at my 
disposal by Dr. House, brings out several striking differences. 
Some of these are shown in PLATES 993 and 994. They may be 
briefly stated as follows!: 


1 SPIRANTHES LACERA (as S. gracilis) is beautifully illustrated in that remarkably 
accurate study of orchids, by Albert M. Fuller, with photographs by George L. Waite, 
Studies on the Flora of Wisconsin Part 1: The Orchids; Orchidaceae— Bull. Pub. Mus. 
Milwaukee, xiv. no. 1, pl. 36 (1933). This plate well displays S. lacera. Although 
Mr. Fuller conservatively followed long-established usage, he obviously saw two 
elements in his S. gracilis, saying (p. 113): ‘‘While the flowers usually occur in spirals 
on the raceme, plants with distinctly 1—sided (secund) racemes appear to be plentiful 
in northern Wisconsin". If the Milwaukee Museum has more such careful ‘‘Studies’’ 
we shall all welcome them. 


8 Rhodora [JANUARY 


SPIRANTHES LACERA (PLATE 993): basal leaves usually present at flowering 
time, though often wilted, submembranaceous, semi-translucent, the veins 
and veinlets clearly evident, the veinlets simple or subsimple, forming an 
obvious loose mesh; spike secund or with 1-few spirals, the flowers distant in 
few elongate series; sepals and petals narrowly lanceolate to lance-linear, 
forming a slender tube 1-1.75 (rarely -2.25), averaging 1.4 mm., in diameter, 
much longer than thick and not strongly ringent; dilated summit of lip droop- 
ing, with a broad white border.—Very dry to moist acid open (rarely shaded) 
soil, Magdalen Islands to Manitoba, south to Nova Scotia, New England, 
Long Island, more rarely to southeastern Virginia, upland to North Carolina 
(up to 3300 ft.) and Tennessee, southern Ontario, Michigan, northern Illinois, 
Wisconsin and Minnesota. Flowering from June 15 to September 11 (AvER- 
AGE of 135 collections Auaust 5). 

S. GRACILIS (PLATE 994): basal leaves rarely present at flowering time 
(though occasionally on non-flowering younger plants), thick, opaque, the 
veinlets barely visible by strong transmitted light, more branched and form- 
ing an obscure but relatively fine mesh; spike strongly spiraling, the approxi- 
mate flowers in many short secund series; tube of perianth more ringent, the 
bases of the broader sepals and petals forming a tube 1.5-2.5, averaging 2 mm., 
thick; white border of summit of lip narrower.—Dry to moist open soil, or in 
open woods and thickets, Florida to Texas, north to southwestern Maine, 
southeastern New Hampshire, central Vermont, southeastern, central and 
western New York, Ohio, Indiana, Illinois, Missouri and Oklahoma. Flower- 
ing late July to October (avERAGE of 110 collections SEPTEMBER 2). 

All material seen from Quebec, New Brunswick, ‘Nova Scotia, 
northern and central Maine and New Hampshire, northern Ver- 
mont, northeastern New York, Ontario, Michigan, Wisconsin, 
Minnesota and Manitoba belongs to S. lacera. In eastern New 
York that plant abounds northward, being the only one of the two 
species found in the Adirondack region and near Lake George, 
from Clinton and St. Lawrence Counties to Saratoga, Fulton and 
Oneida Counties, while in Cattaraugus County to the west it 
occurs at an altitude of 2000 feet. It is the plant of shores and 
slopes near Lake George (our ria. 2) and is clearly the plant de- 
scribed by Rafinesque from there. 

Spiranthes gracilis, of wide southern range, spreads northward 
at low altitudes to Cumberland County, Maine, Strafford and 
Hillsboro Counties, New Hampshire, southeastern Addison 
County, Vermont, and in eastern New York from Long Island 
northward to Albany County. In southern New England and 
southern and central New York both species occur. It is, there- 
fore, specially illuminating to note the collector's data when they 
have placed them both on one sheet or under the same label. 
Thus, the late Charles W. Jenks, getting them both in Bedford, 
Massachusetts, and calling them both S. gracilis, noted them on 
his label and sheet as a, b and c; a being flowering S. lacera col- 


lected July 26, b fruiting material of the same collected August 24, 


Rhodora Plate 993 


Photo. B. G. Schubert 


SPIRANTHES LACERA: FIG, 1, two plants, X 1, from type-region of S. gracilis, var. secunda; 
FIG, 2, two spikes, X 1, from type-region of species: FIG. 3, profile, X 6, of portion of spike; 
ric. 4, face-view, X 6, of same flowers; FIG. 5, lip, X 10; FIG. 6, venation of basal leaf, X 10, 


by transmitted light. 


Rhodora Plate 994 


Photo. B. G. Schubert 


SPIRANTHES GRACILIS: FIG. 1, spike, X 1. from type-region; FIG. 2, profile, 6, of flowers, 
and FIG. 3, face-view, X 6, of flowers; FIG. 4. lip, X 10; Fic. 5, venation of basal leaf, X 10, 
by transmitted light. 


1946] Fernald,—Studies on North American Plants 9 


and c young flowering material (the upper half of the spike in 
bud) of S. gracilis collected August 25. Another sheet from 
Massachusetts, from the herbarium of H. M. Ballou, has the two 
under one label, the slender-flowered S. lacera dated July 18, 
the flowering S. gracilis marked *Aug.". One other mixed sheet 
brings out the difference in flowering period. This is one of the 
late R. W. Woodward's beautiful sheets from Franklin, Con- 
necticut, the label bearing the notes: “plants with basal leaves 
July 14", these being S. lacera in anthesis; *plants without basal 
leaves Aug. 11", these being S. gracilis, so young that the re- 
curving budded tips have not straightened up. 

Not only did Jacob Bigelow think that Spiranthes lacera (his 
Neottia gracilis, var. B. secunda) was “perhaps a different species"; 
Asa Gray, having material of it, probably from northern New 
York, was puzzled by it. His specimen resided for nearly a 
century in the Gray Herbarium, unnoticed in a pocket, pasted 
on a sheet of typical S. gracilis, but with a folded manuscript 
discussing its details and a significant '((?)" after the unsatis- 
factory name. Furthermore, I find that in both the Gray Her- 
barium and that of the New England Botanical Club a specialist 
on the Orchidaceae has recently separated out strikingly charac- 
teristic sheets of S. lacera and has annotated them as the very 
different S. Beckii; at least they did not seem to him to be 5. 
gracilis! 

There can be hardly a doubt of the identity of Spiranthes 
montana Raf. with the beautifully distinct S. ovalis Lindl. 
Rafinesque's description was brief but clear: 


8. Sp. montana, Raf. Caule basi folioso, fol. radic. obl. cuneatis- 
obtusis, caulinis lanc obt. spicis obl. dense spir. bract. obt. acum. fl. 
mediocris, labellum obl. obt. erosum.—Cumberland mts. pedal. disc. 
1823. 


This species (as S. ovalis) has been collected by my companions 
and me several times in Virginia. Its cuneate-oblong or oblan- 
ceate, obtuse or acutish lower leaves, its well developed cauline 
leaf or leaves and the thick and short spike, tapering when young 
but rounded at summit at maturity, are characteristic, as is the 
lip. Although local, the species is scattered in rich, preferably 
calcareous woods from Virginia across the Cumberland Mts. and 
Plateau of Kentucky to bluffs of southern Indiana and to Mis- 


10 Rhodora [JANUARY 


souri, south to northern Florida, Alabama, Mississippi, Louisiana 
and eastern Texas. Dr. E. Lucy Braun records it from three 
counties of Kentucky in the Cumberland area and specimens 
from the Cumberland Mts. of Tennessee are well known. 

As to the identity of Spiranthes tuberosa Raf. and the white- 
lipped S. Beckii Lindl. there is certainly no doubt. Here is the 
original diagnosis: 

10. Sp. tuberosa, Raf. rad. tuberosa monorchis, caule filif. aphyllo, 
vaginis setaceis, spic. gracilis vix spiralis secunda, bract. brevis acutis, 


fl. parvis, labellum cuneato acuto.—Disc. by M. Durand in New Jersey, 
pedal. 


This, with “rad. tuberosa monorchis", ete., is surely the charac- 
teristic species with a single tuberoid, described by Gray as 5. 
simplex: “Root a solitary oblong or spindle-shaped tuber; no leaves 
at flowering time; scape 5'-9' high, bearing a small narrow 
(rarely l-sided) spike of very short flowers (perianth 1"—1Y4" 
long) . . . —E. Mass. (Nantucket, Dr. Robbins), New Jersey 
(C. F. Austin, &c.) and Delaware, Wm. M. Canby". 


HABENARIA MARITIMA Raf., Herb. Raf. 74 (1833) antedates 
by nearly 60 years the Californian H. maritima Greene, Pittonia, 
ii. 298 (1892), basis of Piperia maritima (Greene) Rydb. in Bull. 
Torr. Bot. Cl. xxviii. 641 (1901). 

With only limited understanding of the Californian endemies 
I refrain from renaming the latter. By Ames it is treated as H. 
elegans (Lindl.) Bolander, var. maritima (Greene) Ames, Orchi- 
daceae, iv. 113 (1910). It is noteworthy, however, that both 
Jepson and Abrams, with intimate field-acquaintance of both, 
maintain them as distinct. 

Rafinesque's Habenaria maritima was obviously one of the 
numerous fluctuations of H. clavellata (Michx.) Spreng. (1826), 
which was based on Orchis clavellata Michx. Fl. Bor.-Am. ii. 155 
(1803) from Carolina and which, in spite of the slightly 3-lobed 
tip of the lip was described by Michaux “cornu longitudine 
ovarii, clavato; labello ovali, integro". Rafinesque’s description 
was quite as definite: 

4 Habenaria maritima, Raf. Caule angulato, folia unica longa 
cuneata lane. ceteris subulatis, Spica brevis paucifl. 5-8 fl. bract. lanc. 
ovar. eq. calcar recurvo clavato labello oblongo truncato.—On the Sea 


Islands of New Jersey in swamps, semipedal, flowers small greenish 
white. 


1946] Fernald,—Studies on North American Plants 11 


Those who consider Habenaria maritima Greene a good species 
need a name for it. 


CALIPOGON PARVIFLORUM Raf. Atl. Journ. i. 148 (1832). 


Several years earlier than Calopogon parviflorus Lindl. Gen. 
Sp. Orch. Pl. 424 (1840), that species considered identical with 
Ophrys barbata Walt. Fl. Carol. 221 (1788), the basis of CALo- 
POGON BARBATUS (Walt.) Ames, Orchidaceae, ii. 227 (1908). 
Rafinesque’s plant was evidently of this species, as indicated by 
his "stem one leaved 3-5 flore", for C. barbatus has 1-7 flowers, 
its var. multiflorus (Lindl. Correll in Bot. Mus. Lfts. vii. 71 
(1940), based on C. multiflorus Lindl. l. c. 425 (1840), having 
more. The only other species to consider for Rafinesque’s 
species of “Fl. and Louis", with ‘3-5 . . . flowers spicate, min- 
ute, bracts subulate, labellum undulate", is C. pallidus Chapm. 
(1860). Originally described “Scape 10-20-flowered", that 
species, at the northern limit of its range, in southeastern Vir- 
ginia and North Carolina, may, in the smaller specimens, have 
as few as 3-6 flowers. Until authentic material of Rafinesque's 
species is found, it is wisest to let C. pallidus stand; his specific 
epithet, however, clearly antedates that of Lindley. 


GooDYERA OBLONGIFOLIA Raf. Herb. Raf. 76 (1833). Spiran- 
thes decipiens Hook. Fl. Bor.-Am. ii. 203, t. 204 (1839). G. 
Menziesit Lindl. Gen. Sp. Orch. Pl. 492 (1840). Orchioides 
decipiens and O. Menziesii Ktze. Rev. Gen. ii. 675 (1891). 
Peramium M enziesii (Lindl. Morong in Mem. Torr. Bot. Cl. v. 
124 (1894). Peramium decipiens (Hook.) Piper in Contrib. U. 
S. Nat. Herb. xi. 208 (1906). G. decipiens (Hook.) F. T. Hub- 
bard in Standardized Pl. Names, 328 (1923). 


Rafinesque's description of Goodyera oblongifolia from the 
mountains of Oregon seems unequivocal: 

10 Goodyera seu T'ussaca oblongifolia, Raf. Fol. radie petiol oblongis 
ovatis acutis 5 nervis non reticulatis, subtus glaucis, caule gracile vaginato, 
spica laxiflora, fl. remotis hirsutis, "bract. lanc. acut. ovar. eq. ovarium 
tereto.—Oregon mts. subpedal, fl. white small. 

Hooker said “Scape 8 inches to a foot high”; Lindley’s descrip- 
tion of Goodyera Menziesii, “Hab. in Americae septentrionalis 
orå occidentali, Menzies, Douglas”, began: 


“G. subacaulis, foliis oblongis venosis unicoloribus petiolis longioribus, 
spicá laxá . . . , bracteis ovario aequalibus". 


12 Rhodora [JANUARY 


There is little difference (except in finish) between this account 
of the habit of G. Menziesii and Rafinesque’s definition of the 
earlier G. oblongifolia. 

CORALLORHIZA MONTANA Raf., Herb. Raf. 75 (1833) should be 
added to the synonymy of C. odontorhiza (Willd.) Nutt. (1818), 
which was Cymbidium odontorhizon Willd. (1805). Rafinesque's 
species from the “‘Wasioto mts, and hills, autumnal” had the 
“labello involuto truncato . . . , capsulis pendulis globosis, . . . 
flowers small yellowish, with red spots on the lip". The small 
flowers, pendulous, globose capsule and autumnal flowering seem 
to settle the identity. 


DENTARIA GRANDIFLORA Raf. Herb. Raf. 47 (1833). D. 
macrocarpa Nutt. ex Torr. & Gray, Fl. i. 88 (1838). Cardamine 
pulcherrima Greene in Erythea, i. 148 (1893). D. macrocarpa, 
var. pulcherrima (Greene) Robinson in Gray, Syn. Fl. il. 154 
(1895). D. tenella Pursh, var. pulcherrima (Greene) Detling 
in Am. Journ. Bot. xxiii. 273 (1936). 


Rafinesque's description was clearly of the largest-flowered 
extreme which Greene later described as Cardamine pulcherrima. 
Rafinesque's account of his plant from Oregon follows: 


2. Dentaria grandiflora, Raf, Caule flexuoso apice diphyllo, fol. oppos. 
petiolis alatis, trifoliatis, foliolis sessilib. ovato-lanceol, ineq. serratis, 
racemo brevis umbellato grandifloro.— Pedal. fl. incarnate one inch long. 
petals entire equal to stam. 


SANGUISORBA' STIPULATA Raf., Herb. Raf. 47 (1833). SS. 
canadensis L. Q. latifolia Hook. Fl. Bor.-Am. i. 198 (1834). S. 
sitchensis C. A. Meyer, Fl. Ochot. 34 (1856). Poterium sitchense 
(C. A. Meyer) S. Watson, Bibl. Index, i. 303 (1878). S. lati- 
folia (Hook.) Coville in Contrib. U. S. Nat. Herb. iii. 339 (1896). 


Unfortunately the long-familiar name Sanguisorba sitchensis 
for the northwestern species must lapse. Rafinesque’s descrip- 
tion of his plant from Oregon was clear: 


3. Sanguisorba stipulata, Raf. Caule apice panicul. angul. nudo, fol. 
amplis, foliolis stipulatis petiolatis alternis, cordatis ovatis grandident. 
spicis parvis obl. bract. ovatis acum.—Foliolis twice as large as in S. 
officinalis, 3 inches long. 


Compare the description by Abrams of Sanguisorba sitchensis: 


[11 


. Stipules rounded, coarsely toothed; leaflets oblong-ovate, 2-7 cm. 
long, rounded at the apex, cordate at the base, coarsely serrate, petiol- 
ulate", etc. 


1946] Fernald,—Studies on North American Plants "ES 


EXPLANATION OF PLATES 993 AND 994 


PLATE 993, SPIRANTHES LACERA (Raf.) Raf.: FIG. 1, two plants, X 1, from 
Randolph, Coós County, New Hampshire, Pease, no. 31,500; ria. 2, spike, X 1, 
from the type-region, Northwest Bay, Lake George, Warren County, New 
York, House, no. 30,076; rra. 3, profile of portion of spike, X 6, from no. 
31,500; FIG. 4, face-view, X 6, of portion of spike, from no. 31,500; rra. 5, lip, 
X 10, from no. 31,500; ric. 6, venation of basal leaf, X 10, by transmitted 
light, from Willoughby, Vermont, September 4, 1896, G. G. Kennedy. 

PLATE 994, S. aRACILIS (Bigelow) Beck: ria. 1, spike, X 1, from Wellesley, 
Massachusetts, August 17, 1945, F. W. Hunnewell; ric. 2, portion of spike, 
X 6, from same collection; FIG. 3, face-view of portion of spike, X 6, from 
Winchester, Massachusetts, August 16, 1945, Ernest Rouleau; Fria. 4, lip, X 10, 
from last specimen; FIG. 5, venation of dried-out old basal leaf, X 10, by trans- 
mitted light, from East Hartford, Connecticut, Weatherby, no. 1434. 


II. DIFFICULTIES IN NORTH AMERICAN Sarrx 
(Plates 995-1006) 


1. MUHLENBERG'S NORDAMERIKANISCHEN WEIDEN ANTE- 
DATED BY Micuaux.—The first decade of the 19th century and 
the two decades immediately preceding it were of the greatest 
significance in making known the more generalized flora of eastern 
North America. Not appreciating the ultimate significance of 
the exact date of issue (like too many editors of so-called learned 
societies today), editors brought out scattered or independent 
papers under a blanketing title-page with one arbitrary date for 
the whole series. Something has been done to clarify the dates 
of actual publieation of numerous debatable works and much 
more remains to be done, especially since the over-nice suscepti- 
bilities of librarians and book-binders have long led to the dis- 
carding of or the shifting of the positions of the original covers 
(and, of course, the trimming off of all carefully made marginal 
memoranda). In the intricate genus Saliz one paper has been 
outstanding as containing the original descriptions and drawings 
of leaves of the commoner species of the eastern United States. 
This is the brief article entitled Uber die Nordamerikanischen 
Weiden von Hrn. Pred. MUHLENBERG mit Anmerkungen des Hrn. 
Prof. WiLLDENOW, which was article no. XIV (pp. 233-242, tab. 
VI) in Der Gesellschaft Naturforschender Freunde zu Berlin 
Neue Schriften, iv, with the general title-page dated 1803. 

Since Michaux, in his Flora Boreali-Americana, ii. 225, 226 
(1803), also published as new five species of Saliz from Canada 
and the eastern United States the exact dates of issue of the two 


14 Rhodora [JANUARY 


nearly contemporaneous treatments have to be settled. To be 
sure, it has long, following Willdenow who had an editorial finger 
in the Muhlenberg paper, been assumed to need no investigation. 
In fact, so dominating was the influence of Willdenow and his 
remarkable and compendious Species Plantarum that the dictum 
emanating from those sources has rarely been challenged. Thus, 
the Muhlenberg treatment of American willows was reprinted 
by Konig and Sims in their Annals of Botany, ii. 62-69, pl. 5 
(1805)! with the title: On North American Willows, by the Rev. 
Mr. MuHLENBERG, with Notes of Professor WILLDENOW. Further- 
more, although Sims had been regularly citing Michaux, Flor. 
Bor.-Am. in his articles in Curtis's Botanical Magazine, begin- 
ning on December 1, 1803, so that it appears that he knew that 
remarkable work, he and Konig in 1805 contrived to overlook 
the five species of Salix published by Michaux, for, as an ex- 
planation of their reprinting of the Muhlenberg paper they 
wrote: “of all the species of these regions, we know but one 
through Mr. von Wangenheim [S. conzfera, an abnormal plant] 
and another through Mr. Aiton [S. tristis]." Nevertheless, the 
five species of Michaux can hardly be waved aside; surely not if 
the sketchy accounts by Marshall are satisfactory for the estab- 
lishment of three of our species! 

My attention was drawn to this technical matter through 
noting, while studying Schneider's various papers on American 
willows, that in Journ. Arn. Arb. ii. 189 (1921) Schneider, without 
a word of explanation, reduced outright to S. cordata Muhlenberg 
(our PLATES 995 and 996) the utterly different S. cordata Michx. 
(our PLATES 997-1006); and, furthermore, that in vol. i. 158 
(1920) he had reduced to S. adenophylla Hook. (our PLATE 997) 
the amazingly different S. syrticola Fernald (our PLATES 1001 
and 1002). I was naturally surprised at what has been called 
"this Schneid. treatment" of these species, for I had studied 
Michaux’s willows as well as Hooker's type of S. adenophylla 
(PLATE 997) and I knew that S. cordata Michx. has quite different 
aments from those of S. cordata Muhl.; in fact, that it is the best 
kind of S. adenophylla! And I also knew the several fundamental 
characters which distinguish S. syrticola from the others. It is, 


1 The title-page says 1806, but the late Mr. James Britten pointed out in Journ. 
Bot. xl. 419 (1902) that the pages including the Muhlenberg reprint were issued 
“1 June, 1805.” 


1946] Fernald,—Studies on North American Plants 15 


consequently, evident that the application of Schneider’s apology 
should be éxtended quite to the Atlantic coast, for he gave it too 
occidental a bearing when he wrote, “As to the [§] Cordatae, I am 
not yet well enough acquainted with some of the western forms of 
this group to be able to draw a sharp line between them and the 
[8] Adenophyllae" (Journ. Arn. Arb. i. 148 (1920)). 

Returning to the question of dates, Dr. Schubert, in RHODORA, 
xliv. 149 (1942), has clearly shown that Michaux's Flora Boreali- 
Americana was on sale in March, 1803. It was very soon being 
cited: for instance, Sims in Curtis’s Botanical Magazine, xix, 
under plates 703, etc.; plate 703, of Iris virginica (with definite 
citation of “Michaux Flor. Bor.-Amer. 1. 22"), being engraved in 
time for publication on December 1, 1803. Nearer home, in 
Paris, in the 780-page (therefore not written in a day) vol. vi. 
(1804) of Lamarck's Encyclopédie Méthodique, Poiret added at 
the end of his treatment of Salix the note (p. 661): “Michaux, 
dans sa Flore de l'Amérique septentrionale, cite les espéces sui- 
vantes", followed by transcripts from Michaux and more de- 
tailed descriptions by Poiret. The actual placing on sale of 
Michaux's Flora in March, 1803, can hardly be questioned. 

As to the Muhlenberg paper on Nordamerikanischen Weiden 
I have again asked the aid of Dr. Schubert. The following items 
are most important. The Vorrede of vol. iv. of Der Gesellschaft 
Naturforschender Freunde zu Berlin Neue Schriften, with the 
title-page dated 1803, certainly was prepared before the volume 
was actually published. "This Foreword bears the definite date 
"Berlin, den 3ten Mai 1803", two months after Michaux's 
Flora was on sale. This concerns vol. iv. of the Neue Schriften. 

In the Góttingische gelehrte Anzeigen, Bd. iii for 1803, p. 1493, 
issued “den 17. Sept. 1803", vol. iii of the Neue Schriften was 
reviewed; but it was not until Bd. i for 1804 of Gótt. Anz. p. 
255 (“16. Febr. 1804”) that the first notice of Neue Schriften, iv 
appeared, while Muhlenberg's paper on Nordamerikanischen 
Weiden and succeeding papers in vol. iv were not reviewed until 
the issue of May 12, 1804. Another line of evidence is found in 
the dates of sending or receiving the manuscripts of articles 
published in vol. iv of Neue Schriften. The manuscript of 
article no. VI, by Domeier, was sent from “London, in December 
1802" (p. 110); article no. XVIII, by Karsten, was dated “am 


16 Rhodora [JANUARY 


15ten Marz 1803” (p. 328); article XXIII was submitted by 
Trommsdorf from “Erfurt, im Febuar 1803” (p. 391); while 
article XXIV, by Bode, was submitted from “Berlin, den 26sten 
April 1803” (p. 394). These dates are consistent with the inter- 
pretation derived from the first notice of vol. iv in the Góttin- 
gische gelehrte Anzeigen, “16. Febr. 1804". It is clear, then, 
that Michaux's Flora Boreali-Americana was on sale in March, 
1803, but that Muhlenberg's paper on American Willows could 
not have been issued prior to the date of the foreword, May 3, 
1803, and was probably not available until late in 1803 or early 
1804. 

Fortunately, in 1903 I made a detailed study of Michaux's 
Herbarium in Paris and made notes on or photographs of all 
types which were within my limited understanding. These have 
subsequently been supplemented by some hundreds of sharply 
clear photographs taken under the supervision of Metman by the 
photographer for the Muséum National d’Histoire Naturelle, 
Cintract, so that a fair proportion of Michaux’s species are under- 
stood. Unhappily, however, in case of Salix only my memoranda 
and some very vague photographs of 1903 are available. These 
notes, nevertheless, were explicit, except for the one southern 
species which I then did not know. 


(To be continued) 


Volume 47, no. 564, including pages 393-425, plate 992, and title-page of 
volume, was issued 18 December, 1945. 


FEB 13 1946 


Dodota 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED cee eat 
ALBERT FREDERICK HILL Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. February, 1946. No. 566. 


CONTENTS: 


Ninth Report of the Committee on Plant Distribution. 
R. C. Bean, C. H: Knowlton and A. F. Hl. ............... 17 


Contribution from the Gray Herbarium of Harvard University— 
No. CLX. Technical Studies on North American Plants. 
E Nomad (continded). .......... «mtr o RE Seien 3T 27 


Lythrum alatum in Maine. John C. Parlin. .................... 40 


The New England Botanical Club, Ine. 


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


RHODORA.—4 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 
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vanced prices (see 3rd cover-page). Somewhat reduced rates for complete sets can 
be obtained on application to Dr. Hill. Notes and short scientifie papers, relating 
directly or indirectly to the plants of the northeastern states, will be considered for 
publication to the extent that the limited space of the journal permits. Forms may 
be closed five weeks in advance of publication. Authors (of more than two pages of 
print) will receive 15 copies of the issue in which their contributions appear, 
if they request them when returning proof. Extracted reprints, if ordered in ad- 
vance, will be furnished at cost. 

Address manuscripts and proofs to 
M. L. Fernald, 14 Hawthorn Street, Cambridge 38, Mass. 
Subscriptions (making all remittances payable to RHODORA) to 


Dr. A. F. Hill, 8 W. King St., Lancaster, Pa., or, preferably, Botanical M 
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. 


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EDIBLE WILD PLANTS OF EASTERN NORTH AMERICA 
by Merritt LYNDON FERNALD and ALFRED CHARLES KINSEY 
Practical discussion of edibility and directions for recognition and prepara- 
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Cornwall-on-Hudson, New York, or Librarian, Gray HERBARIUM, Cambridge 
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MEMOIRS OF THE GRAY HERBARIUM. A series of illustrated quarto 


papers issued at irregular intervals, sold separately 


No. I. A Monograph of the Genus Brickellia, by B. L. Robinson. 150 
pp. 96 fig. 1917. $3.00. 


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Section Axillares, by M. L. Fernald. 183 pp., 40 plates, 31 maps. 1932. 
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No. IV. The Myrtaceous Genus Syzygium Gaertner in Borneo, by E. D. 
Merrill and L. M. Perry. 68 pp. 1939. $1.50. 


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Gray Herbarium of Harvard University, Cambridge 38, Mass. 


Rhodora Plate 995 


ie. x » 
| weil a .— INN S 


Photo. B. G. Schubert 


SALIX RIGIDA: FIG. 1, leaf of TYPE, X 1, after Muhlenberg; ric. 2, fruiting branch, X 1; 
FIG, 3, portion of young pistillate ament, X 5; FIG. 4, fruiting ament, X 45; ria. 5, portion 
of fruiting ament, X 5. 


QTRbooora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. February, 1946. No. 566. 


NINTH REPORT OF THE COMMITTEE ON PLANT 
DISTRIBUTION 


The present report deals with the tribes Chlorideae, Festuceae 
and Hordeae of the Gramineae, taken in the order of the seventh 
edition of Gray's Manual. This report completes the treatment 
of the Gramineae which has continued through two previous 
reports with which it has been prepared more or less concomi- 
tantly; acknowledgments there made apply equally here. 


PRELIMINARY LISTS OF NEW ENGLAND 
PLANTS—XXXIV 


The sign + indicates that an herbarium specimen has been 
seen: the sign — that a reliable printed record has been found. 


Me. N. H. Vt. Mass. R. I. Conn. 
I. CHLORIDEAE 


Bouteloua curtipendula (Michx.) Torr. 
Bouteloua gracilis (HBK.) Lag. 
Bouteloua radicosa (Fourn.) Griffiths 
Bouteloua rigidiseta (Steud.) Hitchc. 


T m 
+ 
+ 
e 
Bouteloua simplex Lag. + 
+ 
+ 
-} 


++4++ 


Chloris cucullata Bisch. 

Chloris virgata Swartz 

Cynodon dactylon (L.) Pers. 

Dactyloctenium aegyptium (L.) Richter 

Eleusine indica (L.) Gaertn. 

Leptochloa filiformis (Lam.) Beauv. 

Spartina alterniflora Loisel. 

Spartina alterniflora var. pilosa (Merr.) 
Fern. 

Spartina caespitosa A. A. Eaton 

Spartina cynosuroides (L.) Roth 

Spartina patens (Ait.) Muhl. 

Spartina patens var. juncea (Michx.) 
Hitche. 


od +4+4+4+4++ 


+ + ++ + 
+ + ++ + 

+ + ++ + + 

+ ++4++ + + + 


18 Rhodora [FEBRUARY 


Me. N.H. Vt. Mass. R.I. Conn. 
II. CHLORIDEAE— Cont. 
Spartina pectinata Link + + 
Spartina pectinata var. Suttiei (Farwell) 
Fern. + + 


E 
+ 


+ 


EE 


II. FEsrTUCEAE 


Briza media L. 

Briza minor L. 

Bromus arvensis L. 

Bromus brizaeformis Fisch. & May. 

Bromus ciliatus L. 

Bromus ciliatus var. intonsus Fern. 

Bromus commutatus Schrad. 

Bromus Dudleyi Fern. 

Bromus erectus Huds. 

Bromus inermis Leyss. 

Bromus inermis f. aristatus (Schur.) 
Fern. 

Bromus japonicus Thunb. 

Bromus Kalmii Gray 

Bromus latiglumis (Shear) Hitche. 

Bromus latiglumis f. incanus (Shear.) 
Fern. 

Bromus marginatus Nees 

Bromus marginatus var. seminudus Shear 

Bromus mollis L. 

Bromus mollis f. leiostachys (Hartm.) 
Fern. 

Bromus purgans L. — 

Bromus purgans f. glabriflorus Wieg. 

Bromus purgans f. laevivaginatus Wieg. T 

Bromus racemosus L. 

Bromus rigidus Roth var. Gussonii 
(Parl.) Coss. & Dur. 

Bromus rubens L. 

Bromus secalinus L. 

Bromus sterilis L. 

Bromus squarrosus L. 

Bromus tectorum L. 

„Cynosurus cristatus L, 

Dactylis glomerata L. 

Dactylis glomerata var. ciliata Peterm. 

Dactylis glomerata var. detonsa Fries 

Dactylis glomerata var. vivipara (Lange) 
Carpenter 

Diplachne maritima Bickn. 

Diplachne uninervia (Presl) Parodi 

Distichlis spicata (L.) Greene 

Eragrostis capillaris (L.) Nees 

Eragrostis Frankii C. A. Mey. 

Eragrostis hypnoides (Lam.) BSP. 

Eragrostis intermedia Hitche. 

Eragrostis megastachya (Koel.) Link 

Eragrostis multicaulis Steud. 

Eragrostis pectinacea (Michx.) Nees 

Eragrostis pilosa (L.) Beauv. 

Eragrostis poaeoides Beauv. 

Eragrostis spectabilis (Pursh) Steud. 


— 
+ + ++ 
+++ 4++14+4+4+ + 
444+ 


EE 
c 
+ +4+++ + +44+44+4+4+4+ + 


TER +4 + oda 
+ 
— 


+ 
+ 


I+1+ 
HEHEH cboc cB +4+4+4+4+4+4+4+4++ + 


> 
+ 


4 
4 

TRO + 

TEE GG BEA ++ + 


de 
44 

TER ++ 
TE 


+ 
+ 


+++ + ++ 
+ +4+++ 4A + 
+ +++ +++ 
EGRE 
^o ++ 
++4++++ T4. 


1946] Bean,—Report of Committee on Plant Distribution 19 


Me. N. H. Vt. Mass. R.I. Conn. 
II. Festucear—Cont. 


Eragrostis spectabilis var. sparsihirsuta 
Farwell 

Festuca capillata Lam. 

Festuca elatior I,. 

Festuca obtusa Spreng. 

Festuca ovina L. 

Festuca ovina var. duriuscula (L.) Koch 

Festuca ovina f. hispidula (Hack.) 
Holmb. 

Festuca prolifera (Piper) Fern. 

Festuca rubra L. 

Festuca rubra var. commutata Gaud. 

Festuca rubra f. megastachys (Gaud.) 
Holmb. 

Festuca rubra var. multiflora (Hoffm.) 
Asch. & Graeb. 

Festuca rubra f. squarrosa (Fries) 
Holmb. 

Festuca rubra var. trichophylla Gaud. 

Festuca saximontana Rydb. 

Glyceria acutiflora Torr. 

Glyceria borealis (Nash) Batchelder 

Glyceria canadensis (Michx.) Trin. 

Glyceria Fernaldii (Hitche.) St. John 

Glyceria fluitans (L.) R. Br. 

Glyceria grandis Wats. 

Glyceria grandis f. pallescens Fern. 

Glyceria laxa Scribn. 

Glyceria melicaria (Michx.) Hubb. 

Glyceria obtusa (Muhl.) Trin. 

Glyceria pallida (Torr.) Trin. 

Glyceria septentrionalis Hitche. 

Glyceria striata (Lam.) Hitche. 

Glyceria striata var. stricta (Scribn.) 
Fern. 

Molinia caerulea (L.) Moench. 

Pappophorum mucronulatum Nees 

Phragmites communis Trin. var. Ber- 
landieri (Fournier) Fern. . 

Poa alpigena (Fries) Lindm. f. 

Poa alsodes Gray 

Poa angustifolia L. 

Poa annua L. 

Poa Chapmaniana Scribn. 

Poa compressa L. 

Poa glauca Vahl . 

Poa languida Hitchc. 

Poa laxa Haenke 

Poa nemoralis L. 

Poa nemoralis var. glaucantha (Gaud.) 
Reichenb. 

Poa palustris L. 

Poa pratensis L. 

Poa saltuensis Fern. & Wieg. 

Poa saltuensis var. microlepis Fern. & 
Wieg. 


TEE 
T4 
I +++++ 
L4 


+++ 
a 


++ 
+ t+ + dd 


+ + +++ 
l 
+ + + ++ + +H++H++ 


an 


+ +4++4++ + 
E 


EE 


a 
+t+++4+4+4+ +444 


+ ++4+4+4+4+ GB +444 ++ 
++4+4++ 


+ HEHEHEH 4+4+4++4+4+4 


++ + 
+ 


++ t+ HHHH +44 4+4+44+44+44+ +444 
+- 


++ ++ 444+ 
++14++ +++ + 
+ + +++++ + 
+ + + 4+ 

TR + + + HHH 4+ 


+ +++ 
+ +++ 
+++ 
+++ 
++ 


20 Rhodora [FEBRUARY 


Me. N.H. Vt. Mass. R.I. Conn. 
II. Festucear—Cont. 


Poa sylvestris Gray — 
Poa trivialis L. + + 
Puccinellia distans (L.) Parl. 
Puccinellia distans var. angustifolia 
(Blytt) Holmb. 
Puccinellia fasciculata (Torr.) Bickn. 4 
Puccinellia maritima (Huds.) Parl. + + 
Puccinellia Nuttalliana (Schult.) Hitche. + EE 
Puccinellia paupercula (Holm) Fern. & 
Weath. var. alaskana (Scribn. & Merr.) 
Fern. & Weath. + 
Schizachne purpurascens (Torr.) Swall. + 
Schizachne purpurascens f. albicans Fern. + 
Scleropoa rigida (L.) Griseb. 
Triodia flava (L.) Smyth 
Triplasis purpurea (Walt.) Chapm. 
Vulpia megalura (Nutt.) Rydb. 
Vulpia myurus (L.) K. C. Gmel. 
Vulpia octoflora (Walt.) Rydb. var. 
tenella (Willd.) Fern. 


+ + 
+ 


+ | 


Lu 


TECH 
-— 


++ +++ 
++ +++ 


+ +++ 
+ 

+ + +++ ++ 

+ + +++ ++ 


E 
+ 
+ 


III. HORDEAE 


Agropyron pungens (Pers.) R. & S. 

Agropyron repens (L.) Beauv. 

Agropyron repens f. aristatum (Schum.) 
Holmb. 

Agropyron repens f. pilosum (Scribn.) 
Fern. 

Agropyron repens var. subulatum 
(Schreb.) Reichenb. 

Agropyron repens var. subulatum f. 
heberhachis Fern. 

Agropyron repens var. subulatum f. 
setiferum Fern. 

Agropyron repens var. subulatum f. Vail- 
lantianum (Wulf. & Schreb.) Fern. 

Agropyron repens f. trichorrhachis 
Rohlena 

Agropyron Smithii Rydb. 

Agropyron trachycaulum (Link) Malte 
var. glaucum (Pease & Moore) Malte 

Agropyron trachycaulum var. majus 
(Vasey) Fern. 

Agropyron trachycaulum var. novae- 
angliae (Scribn.) Fern. 

Elymus arenarius L. var. villosus E. Mey. 

Elymus canadensis L. 

Elymus canadensis f. glaucifolius (Muhl.) 
Fern. 

Elymus caput-medusae L. 

Elymus riparius Wieg. 

Elymus villosus Muhl. 

Elymus villosus f. arkansanus (Scribn. & 
Ball) Fern. 

Elymus virginicus L. + + 


+ +++ 
+ + 
+ + 
+ + 4+ 4 


+ 
E 


-L 
-= 


+++ 4 + + + 44 
+ 


-= 
++ + + + + + +4 
-+ 


-H 


+++ + + ++ 4+ 4 
+ 
E 


+++ + + 


=- 


4 
4 
++ ++ + + 4 4+ 4 
++ ++ + +44 
D 
++ +4+4++ + + 


1946] Bean,—Report of Committee on Plant Distribution 21 


Me. N.H. Vt. Mass. R.I. Conn. 
III. HonpEAE—Cont. 
Elymus virginicus f. hirsutiglumis 
(Seribn.) Fern. + + + 
Elymus virginicus var. glabriflorus 
(Vasey) Bush + + 
Elymus virginicus f. australis (Scribn. & 
Ball) Fern. E 
Elymus virginicus var. halophilus 
(Bickn.) Wieg. 
Elymus virginicus var. jejunus 
(Ramaley) Bush 
Elymus virginicus var. submuticus Hook. 
Elymus Wiegandii Fern. 
Elymus Wiegandii f. calvescens Fern. 
Hordeum aegiceras L. 
Hordeum jubatum L. 
Hordeum marinum Huds. 
Hordeum murinum L. 
Hordeum nodosum L. 
Hordeum pusillum Nutt. 
Hordeum vulgare L. 
Hordeum vulgare var. trifurcatum 
(Schlecht.) Alefeld 
Hystrix patula Moench 
Hystrix patula var. Bigeloviana (Fern.) 
Deam 
Lolium multiflorum Lam. 
Lolium multiflorum var. diminutum 
Mutel 
Lolium perenne L. 
Lolium temulentum L. 
Lolium temulentum var. leptochaetum 
Br: 
Nardus stricta L. 
Secale cereale L. + 
Triticum aestivum L. + 


+ + + 4 
+ + 
+ + + 4 


++ 
— 
+ 


d 
-+ 


+ +4+4++ +++ 
ot 


++ 


+++ 4+ ++ cd FAM + ++ + + 
+ 
NE + 


++ + +++ GM 
++ 
++ + +++ ++ ++ + 


+++ 
E 


Es 


For an explanation of names in the above list which are not to 
be found in Gray’s Manual, the following references may be 
consulted: Fernald, nHopoRnA XVIII. 177 and XXXV. 258 
(Spartina); Fernald, RHoponA XXXII. 63 and XXXV. 316, 
Hitchcock, nHoponA VIII. 211, and Wiegand, ruopora XXIV. 
89 (Bromus); Fernald, RHopoRA XXXV. 137 (Dactylis); Fernald, 
RHODORA XL. 108 (Eragrostis multicaulis); Hubbard, RHODORA 
XVIII. 235, Fernald, rnopora XXXIV. 209, XXXV. 132 and 
XXXVII. 250 (Festuca); Fernald, ndHoponA XL. 107 (Vulpia); 
St. John, rHoporA XIX. 75, Hubbard, rHopora XIV. 186, and 
Fernald, nHopomA XXXI. 211 (Glyceria); Fernald, RHODORA 
XXXIV. 211 (Phragmites); Hubbard, rHopora XVIII. 235 
(Poa palustris); Fernald & Wiegand, rHoporA XX. 122 (Poa 
saltuensis and variety); Fernald & Weatherby, RHopora XVIII. 


22 Rhodora [FEBRUARY 


1 (Puccinellia); Fernald, RHopoRA XXX. 161 (Agropyron); 
Wiegand, rHopoRA XX. 81, and Fernald, RHoporA XXXV. 187 
(Elymus); Fernald, RHoDORA XXIV. 229 (Hystrix patula var. 
Bigeloviana as Asperella Hystrix var. Bigeloviana). Additional 
names not given in Gray's Manual may be found in “Manual of 
the Grasses of the United States" by A. S. Hitchcock, Washing- 
ton, 1935. 

Geographically, the ranges of the groups here considered are 
well divided between northern and southern, as in our preceding 
report, (RHODORA XXXVIII. 263-271). The grasses in this list 
belong in geographie groups which have been used and defined 
in previous reports. As usual, varieties and forms which seem to 
have no geographic significance within our area are omitted. 
There are no strictly calcicolous representatives, nor are there 
grasses which have a range covering the region east of the Con- 
necticut River only. 

I. Generally distributed:—Agropyron repens, Elymus vir- 
ginicus, Glyceria canadensis, G. Fernaldii, G. grandis, G. striata, 
Poa angustifolia, P. palustris and P. pratensis. 

Agropyron repens, the familiar witchgrass of fields and gardens, 
seems not to have been extensively collected inland, perhaps 
because of its general abundance in settled areas; Glyceria 
Fernaldii has but one report from Cape Cod, none from Nan- 
tucket, Martha's Vineyard or Rhode Island, but has been col- 
lected at several stations in Connecticut, which is apparently 
near its southern limit of distribution. 

Poa alpigena and P. angustifolia are generally merged in this 
country with an all-inclusive P. pratensis L. Professor Fernald 
supplies the following memorandum. “In both P. pratensis and 
P. angustifolia all or nearly all of the culms bear erect or strongly 
ascending tufts of new green leaves from the basal sheaths, where- 
as P. alpigena has the culms chiefly arising from among old dried 
leaves at the tips of last year's stolons, the basal leaf-tufts all or 
nearly all on separate prolonged stolons or offsets. In P. pra- 
lensis the soft to firm culms are compressed at base and often 
geniculate, 2-3 mm. thick at base; the basal leaves flat or flattish, 
as broad as the thickness of the culm; glumes lanceolate to ovate, 
nearly straight. P. angustifolia has the firm and erect culms 
terete at base, there 1-2 mm. thick; basal shoots with some 


1946] Bean,—Report of Committee on Plant Distribution 23 


(often numerous) filiform to involute blades much more slender 
than the culm; glumes narrowly lanceolate, the second one arch- 
ing. In both these species the lemma is copiously webbed at 
base, the intermediate nerves glabrous, but in P. alpigena the 
nerves are pubescent. The latter boreal species extends south 
to Newfoundland, the Magdalen Islands, Prince Edward Island, 
northern Maine, and the alpine region of Mount Washington, 
New Hampshire." 

II. Rather general except in southeastern Massachusetts;— 
Agropyron trachycaulum var. glaucum, Bromus ciliatus var. in- 
tonsus, Glyceria borealis, G. melicaria, Poa saltuensis. 

Southward Glyceria borealis extends to northern Rhode Island 
and Connecticut, but apparently is absent from the central and 
southern sections of these states. Glyceria melicaria is missing 
from Rhode Island and eastern Massachusetts, save for one 
station on the Merrimack River. Poa saltuensis is not present 
in southeastern New Hampshire, eastern Massachusetts or Rhode 
Island. 

III. Northern A. (with numerous stations south of 43°): 
Agropyron trachycaulum var. novae-angliae, Bromus ciliatus, 
Glyceria striata var. stricta, Poa nemoralis. 

IV. Northern B. (with not many stations south of 43°): 
Agropyron trachycaulum var. majus, Poa glauca, P. saltuensis var. 
microlepis. 

Agropyron trachycaulum var. majus has been collected in Maine 
in the Katahdin and Kineo regions, in Washington County and 
sparingly westward along the coast; in New Hampshire at 
Northumberland, Woodstock and in the White Mountain Region; 
and in Vermont at Canaan. It is also reported in the Vermont 
Flora from Willoughby and scattered stations in the western 
part of the state. 

V. Alpine: Festuca prolifera, F. saximontana, Poa laxa, P. 
alpigena (except in northern Maine). 

Festuca prolifera has been collected on Mount Washington and 
Katahdin as has also Poa lara. The latter also is found on 
Mount Mansfield. Festuca saximontana has been collected at 
Smuggler’s Notch and Poa alpigena on Mount Washington. 

VI. Neither northern Maine nor southeastern Massachusetts: 
Elymus riparius, Festuca obtusa, Hystrix patula, H. patula var. 
Bigeloviana. 


24 Rhodora |. . [FEBRUARY 


Elymus riparius, as its name indicates, is a plant of river 
shores. Festuca obtusa, Hystrix patula and its variety are plants 
of rich mainly deciduous woods. None of these species extend 
east of the Kennebec valley. Festuca obtusa, in fact, has but one 
Maine station (Vassalboro). The absence of these species from 
northern and eastern Maine might give a false impression of the 
distribution of the species as a whole as Hystrix and Festuca 
obtusa are both found on North Mountain, Nova Scotia and in 
the St. John Valley, New Brunswick. 

VII. Chiefly the three southern states:—Bromus Kalmii, B. 
purgans, Elymus canadensis, E. villosus, Eragrostis capillaris, E. 
pectinacea, E. spectabilis, E. spectabilis var. sparsihirsuta, Glyceria 
acutiflora, G. pallida, G. septentrionalis, Poa languida, Triodia 
flava, Vulpia octoflora var. tenella. 

While these species are chiefly found in southern New Eng- 
` land, some of them do extend into Maine, Bromus Kalmii reach- 
ing Oxford County, Maine, and Elymus canadensis reaching 
Maine, as well as northern New Hampshire and northern Ver- 
mont (also New Brunswick). Many of these species have also 
been collected in western Vermont. Several of these species are 
absent from western Massachusetts and from southeastern Mas- 
sachusetts, Martha's Vineyard and Nantucket. These include 
Bromus Kalmii, B. purgans, Elymus canadensis, E. villosus, 
Eragrostis capillaris, Poa languida. 

The typical smooth form of Eragrostis spectabilis has been 
collected at three stations in southern New Hampshire, at two 
in the Boston area, on Martha’s Vineyard and Nantucket, and 
at Bridgeport, Connecticut. The variety sparsihirsuta, which is 
poorly named, since it is usually conspicuously hirsute, is ex- 
tremely abundant and extends considerably further northward. 

VIII. Western New England only:—Eragrostis Franki, E. 
hypnoides. Eragrostis hypnoides occurs along Lake Champlain 
and Otter Creek in Vermont, near the Housatonic River in 
Western Massachusetts and Connecticut, and on the Connecticut 
River as far north as the Hanover region. ‘There is a single 
station on the Stillwater River at Orono, Maine. Eragrostis 
Frankii is more restricted with stations at Manchester and 
Walpole, New Hampshire, Westminster, Vermont, and numerous 
stations along the lower Connecticut River and westward. 


1946] Bean,—Report of Committee on Plant Distribution 25 


IX. Maritime (in the vicinity of the coast with no inland 
stations) :—Dziplachne maritima, Distichlis spicata, Elymus arena- 
rius var. villosus, E. virginicus var. halophilus, Puccinellia fasicu- 
lata, P. maritima, P. paupercula var. alaskana, Spartina alterni- 
flora, S. alterniflora var. pilosa, S. caespitosa, S. cynosuroides, S. 
patens, S. patens var. juncea, Triplasis purpurea. 

Diplachne maritima is at Seabrook, New Hampshire, has one 
report from the Boston region, and is occasional from Falmouth 
and Nantucket, Massachusetts, westward along the coast. Dis- 
tichlis spicata is common as far north as Cumberland County, 
Maine with outlying stations at Rockland and South Thomaston. 
Elymus arenarius var. villosus is a boreal species abundant on the 
Maine coast westward to Cape Elizabeth, with isolated stations 
at Hampton and Isles of Shoals, New Hampshire, Cape Ann and 
Provincetown, Massachusetts. Puccinellia fasciculata is known 
from Great Duck Island, Maine, Plum Island, Massachusetts, 
and from Cape Cod westward. Puccinellia maritima is more lo- 
cal, ranging from Newport, Rhode Island, to Casco Bay, Maine, 
with eastern outposts at Ocean Point and Isle au Haut. Pucci- 
nellia paupercula var. alaskana, a boreal species, is abundant 
westward to New Hampshire, and has isolated stations at Well- 
fleet, Hyannis, Gay Head and Cuttyhunk, Massachusetts, 
Westerly, Rhode Island, and Old Lyme, Connecticut. Spartina 
caespitosa is rather local, occurring from the Thames River, 
Connectieut, to Seabrook, New Hampshire, with a single station 
at South Thomaston, Maine. Spartina cynosuroides has been 
collected in Massachusetts at Salisbury, Nantucket, and Sand- 
wich, Dennis and Brewster on Cape Cod; and at several stations 
from the mouth of the Connecticut River westward. Spartina 
patens var. juncea is common as far north as Plum Island, Massa- 
chusetts and Seabrook, New Hampshire. T'riplasis purpurea is 
frequent northward to York, Maine. 

X. Miscellaneous :—4A gropyron repens var. subulatum, Bromus 
Dudleyi, B. latiglumis, Festuca rubra, Glyceria laxa, G. obtusa, 
Phragmites communis var. Berlandieri, Spartina pectinata. 

Agropyron repens var. subulatum, while essentially a plant of 
seashore areas, also follows the rivers and sand plains inland. 
Bromus Dudleyi, an early blooming grass closely related to B. 
ciliatus, might. be considered as belonging in the Northern A 


26 Rhodora [FEBRUARY 


group were it not for the fact that it seems to be absent from 
northern Maine and northern Vermont. Although it has been 
collected only at scattered stations it is often abundant where it 
does occur. Bromus latiglumis does not occur in southeastern 
Maine (east of the Kennebec River), in southeastern Massachu- 
setts, or in Rhode Island. Glyceria laxa, a plant of moist soil, 
often flourishing in wet peat and wet glades of the Canadian type 
of forest, is frequent in Washington County, Maine, on Mount 
Desert, and in the Penobscot Bay region. It has also been col- 
lected at Milford, Monhegan and South Berwick, Maine, Wolfe- 
boro and Hooksett, New Hampshire, Everett, Massachusetts, 
and Colebrook, Connecticut. Glyceria obtusa has the general 
range of a typical coastal plain species, save that it is apparently 
absent from Nantucket. "There is no specimen available from 
that island, nor is it listed in Bicknell’s Flora. It does occur, 
however, on the other outlying islands, Martha's Vineyard, the 
Elizabeth Islands and Block Island. Phragmites communis var. 
Berlandieri is abundant locally along the coast of Massachusetts, 
. Rhode Island and Connecticut. There is no New Hampshire 
record, but there are scattered stations on the Maine coast as far 
east as Frankfort on the Penobscot River. Inland its distribu- 
tion is erratie with scattered stations in northern Maine, western 
Vermont, western Massachusetts and Connecticut. Spartina 
pectinata is very abundant along the entire coast and occurs inland 
on sandy soil especially along lakes and rivers as far north as the 
St. Johns River in Maine, and Lakes Memphramagog and 
Champlain in Vermont. Naturally it is absent from mountain- 
ous regions. 

There are à very large number of introduced grasses in the 
three tribes considered in this report. Many of these are waifs 
or easuals which have been found in vacant lots in cities and have 
appeared only once or twice. Wool waste has brought in many 
species. Others like the Festucas, Lolium and Cynosurus have 
come in with grass seed. Several of these grasses have become 
very persistent weeds, as in the case of Bromus tectorum, Hordeum 
jubatum, and in some cities, Eleusine indica. 

The Boston region has produced a large number of these 
grasses, partly because of the extensive areas of unoccupied land, 
and partly because so many metropolitan botanists collected 


1946] Fernald,—Studies on North American Plants 27 


extensively in these waste places, especially during the days when 
ballast was unloaded by incoming vessels. 

Among the wool-waste plants are Chloris elegans, C. cucullata, 
the Boutelouas, and Hordeum pusillum. 


R. C. BEAN 
C. H. KNOWLTON 
AR HILE 


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


TECHNICAL STUDIES ON NORTH AMERICAN PLANTS 


M. L. FERNALD 
(Continued from page 16) 


1. SALIX ERIOCEPHALA Michx. Fl. Bor.-Am. ii. 225 (1803) is 
represented by a good branch (except for broken leaf-tips) of the 
foliage ‘‘oblongo-ovalibus, basi subretusis, serrulatis", which my 
note of 1903 described “foliage of oblong-leaved cordata”, and a 
flowering branch which clearly gave the name to the species, ^S. 
diandra: ramulis minutim tomentosis: . . . amentis ovalibus, 
confertim villosissimis", “HAB. in regione Illinoensi", my note on 
it being “flowering branch near discolor". The type is material 
of the tomentulose-branched S. missouriensis Bebb in Garden 
and Forest, vii. 379 (1895). It has been erroneously placed 
with S. discolor as S. discolor Muhl., var. eriocephala (Michx.) 
Anderss. in DC. Prodr. xvi?. 225 (1868), the very large precocious 
aments and long (up to 1 cm.) capsules having deceived those 
who did not consider its other characters, into thinking it S. 
discolor. Michaux’s “foliis oblongo-ovalibus, basi subretusis" is 
not good for S. discolor which becomes relatively local in southern 
Illinois and adjacent eastern Missouri. Michaux collected his 
S. eriocephala ‘in regione Illinoensi". That meant southern- 
most Illinois, for Michaux went down the Ohio, camped at the 
mouth of the Wabash and then proceeded to the Mississippi near 
the mouth of the Ohio. Here S. missouriensis abounds (‘Plants 
of the Lower Wabash Valley", Robt. Ridgeway, no. 1580), Ball 
explicitly referring to it “in Illinois along the Ohio River near its 


28 Rhodora ` [FEBRUARY 


junction with the Mississippi” (Ball in Deam, Shrubs Ind. 52), 
just the region where Michaux got his S. eriocephala. The very 
weak photograph which I got of the foliage of Michaux’s type 
might almost as well have been taken from Ridgway, no. 1580 
from the Lower Wabash, from Glatfelter’s material from Crève 
Coeur, St. Louis Co., Missouri, or from Ball & Over, nos. 2233, 
2235 and 2246 from South Dakota—these all characteristic 
broad-leaved representatives of S. missouriensis. 

2. S. corpata Michx. l. ce. (1803). This, although omitted 
from Index Kewensis, has nothing to do with the later published 
and generally more southern S. cordata Muhl. It was from Lake 
St. John, the entire treatment being 


CORDATA. S. ramulis foliisque villosis: foliis cordato-ovalibus, 
acuminatis, argute serrulatis; stipulis foliaceis, maximis. 
H AB. in Canada, ad lacum S. Joannis. 


The shrub was very familiar to me when I studied Michaux’s 
type, for only three years earlier I had been collecting it along 
the Aroostook River in northern Maine, hence my memorandum: 
“The most extreme broad-leaved pubescent form of the Aroos- 
took R., once taken by me for S. adenophylla [with absolute 
correctness as it proves].” See later discussion. 

3. S. INCANA Michx. l. c. (1803), not Schrank (1789). My 
memorandum accords with the long-held identification: ‘‘The 
true candida with flocculent pubescence. The spec. labelled 
‘Lac Mistassins et Riv. des Goelands'." 

4. S. LONGIROSTRIS Michx. l. c. 226 (1803). My note says 
merely “One of the tristis forms". It is generally treated as 5. 
tristis Ait. (1789). 

5. S. CAROLINIANA Michx. l. c. (1803). Although S. carolini- 
ana is commonly placed in the unquestioned synonymy of S. 
nigra Marsh., with which, in 1903, I was very familiar, I did not 
recognize the Michaux material of his S. caroliniana. Michaux 
identified it with the “S. pentandra?” of Walt. Fl. Carol. 243 
(1788), which had ‘foliis glabris serratis nitidis lanceolatis" and 
which, if it at all resembled the Eurasian S. pentandra L., could 
not have looked very much like S. nigra Marsh., with, to quote 
Deam's Flora of Indiana, “leaves linear-lanceolate". | Michaux's 
S. caroliniana was described as follows: 


Rhodora Plate 996 


Photo, B. G. Schubert 


SALIX RIGIDA (S. cordata Muhl.): Fic. 1, leaf of type of S. cordata Muhl., X 1, after 
Muhlenberg; Fics. 2 and 3, staminate and pistillate flowering branches, X 1; FIG. 4, 
portion of fruiting ament, X 5. 


Rhodora Plate 997 


aS yor 


"2 
AE ud 


Kho oan i 
/4 6 (rm 


Photo, B. G. Schubert 


SALIX CORDATA Michaux (S. adenophylla Hook.): FIG. 1, type of S. adenophylla, X ca. 
Lo: FIG. 2, margin of leaf of type of S. adenophylla, X 10; Fras. 3 and 4, leafy tip and 
fruiting ament, X 45, from probable type-region of S. adenophylla. 


Rhodora Plate 998 


Photo. B. G. Schubert 


SALIX CORDATA Michaux. FIGs. 1, 2, 3 and 6 from Michaux’s TYPE-REGION: FIGs. 1 
and 2, leafy tips, X 45; FIGs. 3 and 4, stipules and leaf-bases, X 5; FIGs. 5 and 6, leaf- 
margins, X 10; ric. 7, staminate ament, X 45. 


Rhodora Plate 999 


SNR 


Photo. D. G. Schubert 


SALIX CORDATA Michaux: FIG. 1, staminate flowering branch, X 45; FIG. 2, pistillate 


flowering tip, X 45; FIG. 3, fruiting ament, 45; FIG. 4, portion of young, and FIG., 5, por- 
tion of flowering staminate ament, showing blackish bracts, X 10. 


Rhodor: Plate 1000 


Photo. B. G. Schubert 


SALIX CORDATA Michaux: Fic. 1, fruiting branch, X 1; FIG. 2, lower surface of mature 
leaf, showing delicate venation, X 10; FIG. 3, portion of fruiting ament, showing short 
pedicels, X 10. 


|Rhodor: Plate 1001 


Photo. B. G. Schubert 


SALIX SYRTICOLA : FIG. 1, pistillate, and FIG. 2, staminate branch, X 45, from TYPE. 


Rhodor: Plate 1002 


P > 


Photo. B. G. Schubert 


SALIX SYRTICOLA: FIG. 1, portion of stipule, petiole and leaf-base, X 5; FIG. 2, 
petiole and leaf-base, X 5; ria. 3, portion of stipule, X 5; FIG. 4, venation of lower 
leat-surtace, X 10; FIG. 5, portion of flowering pistillate ament, X 5; FIG. 6, portion 
of staminate ament, showing pale bracts, X 5, from TYPE. 


Rhodora Plate 1003 
i 


WB Photo. B. G. Schubert 


SALIX RETICULATA: FIG. 1, portion of staminate ament, X 10. 

S. RETICULATA, Var. SEMICALVA: FIG, 2, portion of TYPE, X 46; FIG. 3, staminate ament 
and lower surface of leaf, X 3, from TYPE; FIG. 4, portion of staminate ament, X 10, 
from TYPE, 


1946] Fernald,—Studies on North American Plants 29 


CAROLINIANA. S. foliis lanceolatis, subtiliter arguteque serrulatis, 
subsessilibus: staminibus 4-6: amenti foeminei squamis 
oblongis, minutissime partimque lanuginosis; ovariis 
oblongis, glabris. 

S. pentandra ? WALT. 
OBS. Affinis S. triandrae. 
H AB. in Carolina et Georgia. 


In studying the American willows Schneider had before him 
the vast accumulations in all the larger American herbaria and it 
is significant that, with all these collections before him, he was 
unable to find any typical S. nigra from much of North and South 
Carolina and Georgia: “A very well-known eastern species the 
range of which seems to extend along the Atlantie coast from 
southern New Brunswick to northern North Carolina, and west- 
ward through northwestern South Carolina and northern Georgia 
(from where I have not yet seen typical material) to central and 
eastern Alabama . . . , southern Arkansas", etc. (Schneid. i 
Journ. Arn. Arb. i. 6 (1919)). In S. nigra, according to Sargent, 
Silva, ix. 104, “The stamens vary from three to five in number"; 
similarly, Andersson, the most accurate student ever to work on 
Salix, said in DC. Prodr. xvi?. 200 (1868) “masc. 3-5-andris". 
Michaux’s '4-6" slightly exceeds this and his *amenti foeminei 
squamis . . . partimque lanuginosis" is not too well described 


by Sargent’s “scales . . . coated on the inner surface with pale 
hairs" nor by Andersson's ‘squamis in amentis . . . foemineis 


. . glabriusculis vel basi et- margine villosis’’. 

In view of the great rarity in or absence from much of the area 
of “Carolina et Georgia" of Salix nigra and, likewise in view of 
Michaux's “Opss. Affinis S. triandrae’’, it would seem the obvious 
procedure to look for some common species of Carolina and 
Georgia which looks like the Eurasian S. triandra L. and which 
has 4 to 6 stamens, and the scales of the pistillate aments woolly 
at base. This is obviously S. longipes Shuttlew. ex Anders. 
(1868) and especially S. Wardi Bebb (1895), which Schneider 
treats as S. longipes, var. Wardi (Bebb.) Schneid. Typical 5. 
longipes was recorded by Schneider in Journ. Arn. Arb. i. 25 et 
seq., ‘from Cuba to northern Florida . . . and from . . . ad- 
jacent southeastern Georgia . . . , South Carolina . . . , .. 
eastern North Carolina . . . and i in the southeastern corner of 
Virginia", the barely or hardly separable var. Ward? extending 


30 Rhodora [FEBRUARY 


to the District of Columbia “where it apparently reaches the 
most northern point of its distribution”. One can hardly look at 
characteristic specimens of S. triandra without seeing marked 
resemblance in outline and breadth of leaves to those of S. 
longipes (including S. Wardz), but he would scarcely think of the 
more linear- and narrower-leaved S. nigra. Although Schneider 
in various papers talked around the subject, I fail to find him 
getting down to a concrete statement of the characters of S. 
longipes (and Wardi). Sargent's full description in his Silva, 
l. c. 107, of S. Wardi emphasizes “leaves lanceolate to ovate- 


lanceolate . . . or. . . linear-lanceolate . . . stamens . . . three to 
SIX", while 5. longipes (as S. occidentalis Koch (1828), not Walt. 
(1788)) was defined with “leaves . . . lanceolate, . . . scales 

. oblong-obovate . . . and villous on the back . . . stamens 


five or six". Similarly, Ball, in his detailed description of S. 
longipes (including Ward?) in Deam's Shrubs of Indiana, 44, says 
"Jeaf-blades narrowly lanceolate to lanceolate, . . . scale ob- 
lanceolate or obovate, villous, . . . stamens 5-8". Assembling 
these modern statements and comparing with Michaux’s we get 
for S. longipes (including S. occidentalis sensu Sargent and S. 
Wardi): leaves lanceolate (narrowly or broadly); scales of ament 
oblong-obovate or oblanceolate, villous on the back; stamens 
3-8. Michaux said: leaves lanceolate; scales minutely and partly 
woolly; stamens 4-6; furthermore his S. caroliniana came from 
Carolina and Georgia, where S. longipes abounds and where S. 
nigra is rare or local. 

Two more points. Quite unfamiliar with Salix longipes 
(Wardi) in 1903, I entered only a query against Michaux's S. 
caroliniana. This was “Form of S. cordata?". That was only 
an off-hand suggestion, but Bebb, who set up the species, S. 
Wardi, in Gard. and For. viii. 363 (1895), had originally published 
it as S. nigra, var. Wardi in Ward, Guide to Flora of Washington, 
114 (1881). He then (1881) spoke of a form of the latter which 
"might be easily mistaken (in the absence of aments) for an 
extravagant growth of S. cordata". Again, in 1895, he wrote: 
“The statement made when this Willow was first described that 
in some of its forms the leaves alone, with their ample stipules, 
might easily be mistaken for 5. cordata, finds striking exemplifica- 
tion in Professor Short’s specimen in the Gray Herbarium, which 


1946] Fernald,—Studies on North American Plants 3l 


two no less competent salicologists than Mr. Carey! and Professor 
Andersson have mistaken for ‘S. cordata angustata’. Indeed it 
is apparent from the description that this identical specimen 
served as the type of S. cordata angustata, 1? forma discolor, 
Andersson (DC., Prod. xvi?. 252)." When the foliage of Mi- 
chaux's type of S. caroliniana reminded my then quite inexperi- 
enced eye of that of S. cordata (surely not of S. nigra) I was in 
distinguished company, for it is an honor to approach the class 
with the discerning John Carey, the highest of honors to get near 
the limited group of most cautious salicologists with Nils Johan 
Andersson! 

Schneider states with seeming finality regarding Salir caro- 
liniana that “unfortunately no type specimen seems to exist 
in Michaux's herbarium at Paris"—Journ. Arn. Arb. iii. 64 
(1921). But, from what I have already noted, it is evident that 
Schneider did not at all understand the types at Paris of Mi- 
chaux's 5. eriocephala and S. cordata and probably never studied 
them. These types and that of S. caroliniana were all there in 
1903, when I studied and photographed some of them; of course, 
since the invasion of Paris by Hitler’s ravaging hordes they 
may now be missing; but shortly before the “blitzkrieg” in which 
Paris was invaded at the opening of the recent war the TYPE was 
there, for Cintract took the photograph of it which is before me. 
This photograph shows the relatively broad young leaves paler 
beneath than above and the toothing of Salix longipes, not of S. 
nigra. lam satisfied that S. CAROLINIANA Michaux. (1803) is S. 
longipes Shuttlew. (1858), i. e. S. Wardi (Bebb) Bebb (1895). 

S. CORDATA Michx. Fl. Bor.-Am. ii. 225 (early 1803); Poiret in 
Lam. Encycl. Méth. vi. 661 (1804); not Muhl. in Ges. Naturf. 
Freunde Neue Schr. iv. 236, t. 6, fig. 3 (late 1803 or early 1804). 
S. adenophylla Hook. Fl. Bor.-Am. ii. 146 (1839); Schneid. in 


Journ. Arn. Arb. i. 158 (1920) in part (excl. S. syrticola) ; St. John, 
Viet. Mem. Mus. Mem. 125: 79 (1922); Raup in Sargentia, iv. 


1 In his Salices Amer. no. 22, Joseph Barratt had Salir longipes under a nomen 
nudum (in Index Kewensis and also cited by Schneider, so that I am not here pub- 
lishing a useless name), with the following explanation: “22. Salix Pitcheriana* 
Barratt, mss. Hab. Arkansas.—Dr. Pitcher. Sea Islands of Georgia. This unde- 
scribed species is allied to S. nigra. I possess specimens which have been obligingly 
communicated by John Carey, Esq. of New York”, etc. John Carey was not properly 
edified. On one of his labels of Georgia material he wrote: ‘‘I always supposed this 
to be S. nigra of Lin [who had no such species]: (no doubt it is of Ell.) but Dr. Barratt 
who calls himself the great authority for our willows names it a new species". ‘‘Au- 
thorities'" beware! 


32 Rhodora [FEBRUARY 


111 (1943).—Southeastern Labrador Peninsula to James Bay, 
Ontario, south to Newfoundland, Nova Scotia, northern Maine, 
eastern Cape Cod, northern New York, Simcoe and Bruce Cos., 
Ontario, and northern Michigan. PraArks 997-1000. 

Salix cordata Michx. has been wrongly guessed, ever since 
Willdenow, to be identical with the later S. cordata Muhl. (see 
pp. 14 and 28 and plates 995 and 996). In his Species Plantarum, 
iv?. 666 (1806), Willdenow took up the later S. cordata Muhl., 
expanding Muhlenberg’s original description to read “ramis 
glabris viridibus. Folia tripollicaria oblongo-lanceolata acuminata 
basi cordata, margine argute serrata, serraturis cartilagineis, 
utrinque glabra", ete.; and at the same time he maintained 5. 
rigida Muhl. as a distinct species, although others have not been 
able to do so. S. rigida was thought to be distinguished by 
“ramis viridibus superne purpurascentibus, junioribus pubescenti- 
bus. Folia tripollicaria rigida oblongo-lanceolata acuminata basi 
subcordata, margine argute serrata, serratura infime elongata apice 
glandulosa” ete., not very convincing differences, especially when 
Muhlenberg’s original figures (our PLATE 995, FIG. 1 and 996, 
FIG. 1) are compared. However, in S. rigida, with glabrous 
oblong-lanceolate subcordate leaves, Willdenow doubtfully 
included “S. (cordata) ramulis foliisque villosis, foliis cordato- 
ovalibus acuminatis argute serrulatis, stipulis foliaceis maximis. 
Mich. amer. 225?”. There he had a really different species, which 
has positively cordate and narrowly oval or ovate leaves densely 
villous when yoüng, and often to maturity, and coming originally 
from Lake St. John, which is more than 600 miles north of 
` Muhlenberg’s region (Lancaster County, Pennsylvania) and with 
a Hudsonian or Hudsonio-Canadian (instead of Alleghenian- 
Carolinian) flora. As stated on a previous page (28) the type 
of S. cordata Michx. from Lake St. John is surely of the northern 
species with densely pubescent branchlets and young foliage, 
which is common from southeastern Labrador Peninsula to James 
Bay, a species (PLATES 997-1000) of which many sheets from 
Lake St. John are before me. In its smallest-leaved develop- 
ments it is quite identical with the type of S. adenophylla Hook. 
(PLATE 997, rias. 1 and 2) from “Labrador. Dr. Morrison", the 
latter region. being presumably the Cóte Nord of the eastern part 
of Saguenay County, Quebec, which in Hooker's (and Morrison's) 
time was included in “Labrador”. At least the type of S. adeno- 


1946] Fernald,—Studies on North American Plants 33 


phylla could perfectly well have come from the Natashquan 
River, the old ‘‘American Harbour,” for Natashquan material 
(PLATE 997, FIGs. 3 and 4) is very like it. The photograph of 
Hooker's type of S. adenophylla was sent to the late Professor 
Sargent and is preserved, along with some leaves and portions of 
an ament at the Arnold Arboretum. In its details it is quite like 
specimens from Newfoundland, the Cóte Nord, Lake St. John, 
Aroostook River, Maine, James Bay and elsewhere in the range 
of Michaux's species; and the photograph and fragments exactly 
agree with Hooker’s detailed account of his S. adenophylla: 


8. S. adenophylla; ramis brevibus subrobustis lanatis, foliis ovatis basi 
cordatis acutis subcoriaceis fere omnino sessilibus reticulatim veriosis 
argute serratis serraturis elongatis glanduliferis lana sericea dense obsitis 
demum aetate nudiusculis, stipulis ovato-cordatis grosse glanduloso- 
serratis, amentis foemineis elongatis pedunculatis, capsulis ovatis acumi- 
natis glaberrimis, stylo elongato, stigmatis lobis fissis. 

HAB. Labrador. Dr. Morrison.—I know no species like this, well 
marked as it is by the copious long narrow serratures to the leaves tipped 
with a gland, so that the leaf looks as if it were fringed with pedicellated 
glands. These leaves are an inch or more long, clothed, even when fully 
grown, with long silky tomentum on both sides, but which is deciduous on 
the oldest leaves . . 


Although the teeth on young and just expanding leaves of 
Salix cordata Michx. (and S. adenophylla) may be prolonged and 
gland-tipped, they are often lower, blunter and less evidently 
ending in glands, in such specimens the toothing approaching that 
of S. rigida Muhl. (S. cordata Muhl.). Furthermore, the narrow- 
est-leaved S. cordata resembles the broadest-leaved S. rigida but 
the blades are more generally cordate in the former than in the 
latter. Measurements of all mature foliage in the Gray Her- 
barium gives the following results, to which I add the far more 
significant characters of the aments. 


S. CORDATA: leaves broadly oblong-lanceolate to ovate, mostly strongly 
cordate, the mature ones 3-13 (av. 6.5) em. long and 2-5.5 (av. 3.5) cm. 
broad; staminate aments terminating leafy axillary branchlets, the leaves 
well developed at anthesis, the bracts with whitish beard; pistillate aments 
dense, in anthesis with appressed-ascending ovaries, in fruit with the capsules 
crowded on very short pedicels shorter than to barely exceeding the bracts. 

S. RIGIDA: leaves oblong-lanceolate, subcordate, rounded or tapering or 
attenuate at base, the mature blades one eighth to one third as broad as long, 
4.5-15 cm. long, 0.9-4.5 em. broad; staminate aments subtended by short 
bracts barely expanded at anthesis; pistillate aments in anthesis with widely 
(often horizontally) divergent ovaries, in fruit with the widely divergent 
capsules on pedicels as long as to much longer than their subtending bracts. 


hi 


34 Rhodora [FEBRUARY 


Typical Salix cordata Michx. has heavily villous branchlets 
and young leaves, the pubescence inclined to persist on the 
mature foliage. In the eastern part of its range it is repre- 
sented in many areas (along certain rivers, etc.) by a glabrous or 
glabrescent extreme, which has the aments, stamens, short pedi- 
cels and cordate leaf of the typical extreme of the species, but 
with the teeth less often prolonged and gland-tipped than in 
that shrub. With leaf-outline and aments of S. cordata and 
occurring only in Newfoundland, eastern Canada and northern 
Maine, mostly north of S. rigida, it seems to be an extreme of 
the northern species. This is 


S. conDATA Michx., var. abrasa, var. nov., ramulis petiolisque 
glabris vel glabratis; foliis costa excepta glabris vel glabratis.— 
Newfoundland and Gaspé Peninsula, Quebec, to Nova Scotia 
and northern Maine. The following are characteristic. NEw- 
FOUNDLAND: Birchy Pond Stream, Fernald & Wiegand, no. 3149; 
Harry’s River (or Brook), Fernald & Wiegand, no. 3150; Force- 
le-Plain, Harry’s Brook, A. B. Kennedy, nos. 305 and 387; 
Riverview Camp, Grand Codroy River, Pease & Edgerton, no. 
27,120. Quesec: R. Ste. Anne des Monts, Gaspé Co., July 
14-17, 1906, Fernald & Collins, nos. 484 (ryrE in Herb. Gray.), 
485 and 486; R. Petite Cascapedia, Bonaventure Co., Victorin, 
Rolland & Jacques, no. 33,845; Bonaventure R., Bonaventure Co., 
Aug. 4-8, 1904, Collins, Fernald & Pease (Pease, nos. 5831 and 
5897); junction of Restigouche and Matapedia Rivers between 
Quebec and New Brunswick, Rousseau & Boivin, nos. 32,037 and 
32,082; Grande-Décharge, Lac St.-Jean, Victorin, Rolland & 
Meilleur, no. 45,872. New Brunswick: Tom's Island, Resti- 
gouche R., July 30, 1896, G. U. Hay; lower Tobique River, Oct., 
1945, G. D. Chamberlain. Nova Scoria: Salt Springs, Pictou 
Co., Perry, Wetmore, Hicks & Prince, no. 10,134; Truro, Col- 
chester Co., J. G. Jack, no. 3633; Wellington, Shubenacadie 
Grand Lake, Halifax Co., Fernald, Bartram & Long, no. 23,739 
(transitional); Landsdown, Digby Co., J. G. Jack, no. 3704. 
Maine: Fort Kent, Aroostook Co., Fernald, nos. 2473-2475; 
Pease, no. 2578. ' 


Schneider, who considered Salix cordata Muhl. and S. cordata 
Michx. identical and who put them both into § Cordatae, treated 
S. adenophylla Hook. (which is really very small-leaved 5. 
cordata Michx.) as the type of a separate § Adenophyllae. He 
was unable to distinguish from the latter northern species the 
very different shrub of sands about Lake Michigan, S. syrticola 
Fernald in Ruopora, ix. 225 (1907), (our PLATES 1001 and 1002), 


1946] Fernald,—Studies on North American Plants 35 


Schneider saying “Unfortunately Fernald did not see Hooker’s 
type of which I have before me an excellent photograph and some 
fragments from the Kew Herbarium . . . When he proposed his 
new species he did not know of the specimens collected by Ma- 
coun, Ross and Spreadborough in the James Bay region. They 
connect the original habitat of (probably southern) Labrador and 
that of S. syrticola. The only difference between the forms of 
James Bay and those of the Great Lakes, so far as I can judge, is 
in the length of the styles, which measure about 1.5 mm. in 
typical S. adenophylla, while they rarely are longer than 1 mm. 
in S. syrticola"—Schneid. in Journ. Arn. Arb. i. 158, 159 (1920). 
Now it so happens that, in 1903 (again in 1930) the author of 
Salix syrticola had closely studied Hooker's type of S. adeno- 
phylla and he has many times studied colonies of S. adenophylla 
in Newfoundland, Quebec and northern Maine. The type, from 
the northeastern and rather inhospitable limit of the specific 
range, was unusually small-leaved and heavily pubescent, but 
the characters of this species (as the earlier S. cordata Michx.) are 
shown in PLATES 997-1000. Whether S. syrticola is fully dis- 
tinct from S. cordata it is too soon to assert with finality. In 
general the two are very different in many characters, but in 
northern Michigan and about the Lake Huron shores of Ontario 
some shrubs indicate possible transitions which are no more 
tangible than the “mongrels’”’ into which other species of Salix 
regularly “spawn” (to quote Bailey's picturesque phrase) when 
they meet and cross. Until very recently we did not know good 
staminate aments and the plans to secure them last June from 
the type-region of the northern shrub, Lake St. John, or from 
northern Maine were thwarted by the erratic weather, when 
abnormally late frosts after an abnormally early opening of 
spring blighted the flowers or fruits as well as vegetative tips of 
many species occurring from eastern Canada to the Southeastern 
States. Most fortunately, however, in late June and early July, 
Fathers Dutilly and Lepage secured beautiful freshly flowering 
staminate, as well as pistillate, material near James Bay. This 
differs at once from S. syrticola in many characters. As to the 
staminate aments, they are borne at the tips of well developed 
leafy branchlets, the bracteal leaves very much more developed 
than are those of S. syrticola during anthesis, and the beard of the 
bracts is much whiter than the fuscous beard in 5. syrticola. 


36 Rhodora [FEBRUARY 


These characters of the staminate aments, accompanied by 
parallel ones in the pistillate, and by different outline, toothing 
and reticulation of leaves, different toothing of stipules, ete., lead 
to the following statement of contrasts. 


S. CORDATA: Leaves broadly lance-oblong to oblong-ovate, long-acuminate, 
gradually tapering from below or near the middle, the mature ones 3-13 cm. 
long; each margin with 25-90 (av. 55) forward-arching at first often gland- 
tipped but soon glandless mostly simple teeth; the mature petiole (2-) 5-35 
(av. 13) mm. long; the mature lower surface with secondary veins slender 
and relatively low: mature and larger stipules with 0-22 (av. 14) gland-tipped 
or mostly glandless teeth on the longer margin; staminate aments on leafy 
branchlets, the leaves well-grown at anthesis; the blackish to brown bracts 
with white beard: pistillate aments in maturity 2-6 cm. long; bracts narrowly 
obovate, fuscous or brown, their bright white beard only slightly longer; 
ovaries in anthesis appressed-ascending, in fruit more spreading, on pedicels 
shorter than bracts. PrATEs 997-1000. 

S. svRTICOLA: Leaves oblong-ovate, acute or abruptly short-acuminate 
from well above the middle, the mature ones 3.5-9.5 em. long; each margin 
with 81-137 (av. 105) horizontally or subhorizontally divergent and perma- 
nently gland-tipped prolonged often compound teeth; the mature petiole 
2-10 (-15) (av. 6.3) mm. long, thicker than in the preceding; mature lower 
surface with secondary veins coarse and rather prominent: mature and 
larger stipules with 24—40 (av. 32) mostly gland-tipped and straight teeth on 
the longer margin: staminate aments in full anthesis subtended by leaves 
only about one-fourth grown; the oblong pale brown bracts with ashy-white 
to fuscous beard: pistillate aments in maturity 6-8 cm. long; flowers soon 
divergent, the mature capsules on pedicels nearly as long as to longer than 
blade of the oblong pale brown bract, the beard of the latter very long, ashy; 
style 0.7-1.5 mm. long. Pxrares 1001 and 1002. 


With such an abundance of distinctive characters, I find myself 
incapable of following Schneider in treating S. cordata (S. 
adenophylla) and S. syrticola as absolutely the same. Schneider 
states that the “styles . . . measure about 1.5 mm. in typical S. 
adenophylla [i. e. cordata], while they rarely are longer than 1 mm. 
in S. syrticola" (Schneider, Journ. Arn. Arb. i. 159). Rather 
naturally, however, my measurements of the style of S. syrticola 
accurately coincide with those of Ball in Deam's Shrubs of 
Indiana, 0.7-1.5 mm. I am not, as already emphasized, con- 
vinced that S. cordata (S. adenophylla) and S. syrticola are con- 
specifie, although, as also noted above, at the northern border of 
the range of the latter some transitional specimens may possibly 
occur. So they do at the border-lines or coincident ranges of 
many willows. That is one of the reasons the willows are diffi- 
cult; they will cross. 

Incidentally, Salix syrticola has its greatest development on 
the dunes of Lake Michigan, especially toward the southern end 
of the Lake, a region famous for the isolation there of prevailingly 


1946] Fernald,—Studies on North American Plants 37 


southern, rather than northern species. Although the dunes sup- 
port Pinus Banksiana and a few other common Hudsonian and 
Canadian species, they are famous largely on account of the 
remarkable assemblage of far-isolated southern or southeastern 
species. These include Aristida tuberculosa, **Massachusetts to 
Georgia and Mississippi; around the southern end of Lake 
Michigan and in . . . Wisconsin, Indiana, Illinois", ete. (Hitch- 
cock); Panicum auburne, ‘‘Massachusetts to northern Florida and 
Louisiana; Arkansas; Indiana, near Lake Michigan" (Hitchcock); 
Eleocharis geniculata (E. caribaea), Florida to Texas and southern 
California, north near coast to North Carolina, sands of Great 
Lakes, southern Ontario to Michigan and northwestern Indiana; 
E. melanocarpa, localized in northern Florida and Georgia, south- 
eastern Virginia, southern New Jersey to southeastern Massa- 
chusetts, eastern Texas, and dunes of Lake Michigan; Psilocarya 
nilens, occurring from eastern Texas to Florida, thence north to 
southeastern North Carolina, otherwise highly localized in Sussex 
County, Virginia, Cape May region, New Jersey, Suffolk County, 
Long Island, Plymouth County, Massachusetts, and dunes of 
Lake Michigan; and so on to Stachys hyssopifolia, a very charac- 
teristic species occurring from southern New England and south- 
eastern New York to South Carolina and isolated in southern 
Michigan and northwestern Indiana; or Lycopus amplectens Raf. 
(L. sessilifolius Gray), one of the most definite of species, occur- 
ring from Mississippi to Florida, thence to North Carolina, from 
southern New Jersey to southeastern Massachusetts, and among 
the dunes of Lake Michigan. In view of this striking southern, 
rather than prevailingly Hudsonian, relationship of the flora of 
the dunes of Lake Michigan, it is wise to hesitate before too 
positively asserting that S. syrticola of the dunes of Lake Michi- 
gan is, in spite of its many distinctive characters, identical with 
the shrub which prevails in the Hudsonio-Canadian area from 
Hudson Bay to southern Labrador and Newfoundland.  Primar- 
ily upon S. syrticola which he misidentified with S. adenophylla, 
Schneider set up his § Adenophyllae, although, as quoted by me 
on p. 15, he admitted that he was perplexed *'to draw a sharp line 
between [§ Cordatae] . . . and the [§] Adenophyllae". If S. cor- 
data Michx. and S. cordata Muhl. are, as Schneider incorrectly 
says, identical, though clearly of the same section, and if, as it 


38 Rhodora [FEBRUARY 


seems to me, S. adenophylla is the same as S. cordata Michx. (not 
Muhl.), then § Adenophyllae Schneid. has a very slim basis for 
separation from $ Cordatae. 

The automatic abandonment of the later name, Salix cordata 
Muhl., not Michx., necessitates the following transfer: 

S. RIGIDA Muhl., forma mollis (Palmer & Steyerm.), comb. 
nov. S. cordata Muhl., forma mollis Palmer & Steyerm. in Ann. 
Mo. Bot. Gard. xxv. 770 (1938). 

S. RIGIDA Muhl., var. angustata (Pursh), comb. nov. S. an- 
gustata Pursh, Fl. Am. Sept. ii. 613 (1814). S. cordata Muhl., 
6. angustata (Pursh) Anderss. in DC. Prodr. xvi?. 252 (1868). 

Typical Salix rigida Muhl. has the leaves broadly rounded to 
subcordate or slightly cordate at base, one sixth to one third as 
broad as long, the mature blades 1.5-4.5 (av. 2.75) cm. broad. 
In var. angustata the blades taper to base or are very gradually 
rounded, one eighth to one fourth as broad as long, the mature 
ones 0.9-2.2 (av. 1.5) em. broad. 


2. ADDITIONAL NAMES AND TRANSFERS IN SALIX— 


X S. Jesupi. S. pameachiana sensu Anders. in K. Svensk. 
Vet.-Akad. Handl. vi. 48 (1867), not Barratt (1840). S. alba, 
var. pameachiana Anders. in DC. Prodr. xvi? 212 (1868), not S. 
pameachiana Barratt, basonym. S. alba X lucida Bebb in 
Gard. and For. viii. 423, 424, fig. 57 (1895). 

Named for its discoverer, HENRY Gmisworp JESUP (1826- 
1903). Salix pameachiana Barratt, Salices Amer. (1840) “‘grow- 
ing about the Pameacha stream in this town [Middletown, Con- 
necticut]” and, therefore, not a personal name as assumed by 
Andersson and by Schneider, who regularly used a capital initial, 
was said by Barratt to be “the intermediate of S. vitellina and 
the former [S. decipiens or fragilis]". Carey in Gray, Man. 428 
(1848) treated it as S. alba, var. vitellina X S. fragilis. It has 
nothing to do with the very striking X 5. Jesupi. 

S. INTERIOR Rowlee, var. exterior, var. nov., foliis plus minusve 
persistenter sericeis breviter subacutis oblongo-lanceolatis denti- 
bus suppressis, lamina 2-7 cm. longa ad 1.5 cm. lata.—Beaches 
of Aroostook River, Maine: Caribou, July 18, 1902, Walliams, 
Collins & Fernald (ryeE in Herb. Gray.; isorvyrE in Herb. N. E. 
Bot. Cl.); Fort Fairfield, June 28, 1931, Fernald & Weatherby, 
no. 2432. PENNSYLVANIA: banks of Susquehanna River, above 
MeCall's Ferry, York Co., Sept. 13, 1864, T. C. Porter (Herb. 
Phil. Acad.; fragments and tracing in Gray Herb.). 


1946] Fernald,—Studies on North American Plants 39 


In its very short and broad leaves with suppressed teeth quite 
distinct from narrower-leaved typical Salix interior, in which the 
usually divergent teeth are prominent. S. interior, named for 
its extensive inland development, or its forma Wheeler? (Rowlee) 
Rouleau, extends from Alaska to Oklahoma and Arkansas, and 
eastward in the North to the St. Lawrence River in Quebec and 
the Connecticut in western New England, with a slight but iso- 
lated occurrence along the St. John in New Brunswick and the 
lower Restigouche in Quebec. Var. exterior is close to the eastern- 
most edge of the specific range. Although the name S. longifolia 
Muhl. (1803) is still much used, it was clearly antedated by the 
wholly different S. longifolia Lam. (1778). 

S. RETICULATA L., var. semicalva, var. nov. (TAB. 1003, FIG. 
2-4 et TAB. 1004, FIG. 2-4), squamis masculis flavescentibus 
dorso glabris vel glabratis; antheris flavescentibus; squamis 
foemineis flavescentibus vel fulvis; capsulis sparse breviterque 
pilosis.—Limestone barrens and gravels, northern and north- 
western NEWFOUNDLAND: Quirpon Island, Wiegand, Gilbert & 
Hotchkiss, no. 27,939; Cook Point, Pistolet Bay, Fernald & Gil- 
bert, no. 27,936; Anse aux Sauvages, Pistolet Bay, Fernald, 
Wiegand & Long, no. 27,940; Sandy (or Poverty) Cove, Straits of 
Belle Isle, Fernald, Long & Dunbar, no. 26,586; between Name- 
less Cove and Mistaken Cove, straits of Belle Isle, Wiegand, 
Pease, Long & Hotchkiss, no. 27,935; Flower Cove, July 17, 1920, 
Mary E. Priest; Capstan Point, Flower Cove, Fernald, Long & 
Dunbar, no. 26,585; south of Flower Cove, Wiegand, Pease, Long 
& Hotchkiss, no. 27,934; Brig Bay, Fernald, Long & Dunbar, no. 
26,587; St. John Island, Fernald, Wiegand, Long, Gilbert & 
Hotchkiss, nos. 27,937 and 27,938; Ingornachoix Bay, Fernald & 
Wiegand, no. 301; Gargamelle Cove, Ingornachoix Bay, Fernald, 
Long & Fogg, nos. 1578 (rvPE in Herb. Gray., July 20, 1929)— 
1580; Pointe Riche, Fernald, Long & Fogg, no. 1581. 

The two collections (Woodworth, no. 143, and Abbe & Odell, no. 
210), from the Torngat region of northern Labrador, have the 
pale bracts and the minutely puberulent ovaries of var. semicalva. 
I have seen no staminate material from that area. 

Typical arctic-alpine Salix reticulata (PLATE 1003, Fic. 1 and 
1004, ric. 1) has the bracts of the pistillate aments dark (deep 
purplish to blackish), those of the staminate aments heavily 
villous at base on the back (as well as on the inner surface), the 
anthers dark (Schneider in Journ. Arn. Arb. iii. 92 (1921), in his 
key saying “Antherae violaceae" for S. reticulata as opposed to 


40 Rhodora [FEBRUARY 


* Antherae flavae" for S. vestita), the ovaries and capsules heavily 
white-tomentulose. The arctic shrub (especially in Eurasia) 
may often have silky pubescence on the young leaves, this per- 
sisting near the base of the leaf on the back; and in European 
(typical) specimens the peduncle is usually villous. The New- 
foundland representative of the species is glabrous from the first 
or with the young peduncles only sparsely pilose. Its aments 
have yellowish-brown to fulvous bracts, these in the staminate 
aments only weakly pilose or promptly glabrescent, the anthers 
pale or yellow, while the pubescence of the capsules is so short 
and fine that the purplish color of the capsule is scarcely ob- 


scured. 
(To be continued) 


LyYTHRUM ALATUM IN Martnge.—On 10 Aug., 1945, Mrs. Cleora 
D. Adams of Hartford sent me for identification a plant that she 
collected on 8 August in Belgrade, Kennebec County. I deter- 
mined it as Lythrum alatum Pursh, and Mr. C. A. Weatherby 
verified my finding. We thought that this might be its first 
known occurrence in Maine, but Prof. Fay Hyland writes me that 
he has it from Fryeburg, and Ralph Bean tells Mrs. Adams that 
he has collected it in Clinton. 

Mrs. Adams gives this note on its location. “Growing in a 
patch 10-12 ft. across, noticeable from. highway a few hundred 
feet away in damp meadow in bend where R. 135 leaves R. 11, 
before reaching cemetery which is on both sides of the road, and 
before R. 135 crosses R. 27. 

‘Three species of butterflies were working on the flowers: one 
blaek swallow-tail, several sulphurs, and several smaller ones 
which I think were pearl erescents".—Jonw C. Parry, Canton, 
Maine. 


Volume 48, no. 565, including pages 1-16 and plates 993 and 994, was issued 
ó January, 1946. 


Rhodora Plate 1004 


Photo. B. G. Schubert 


SALIX RETICULATA: FIG. 1, portion of fruiting ament, X 10. 


S. RETICULATA, Var. SEMICALVA: FIG. 2, fruiting plant, X 45; FIG. 3, fruiting ament, X 5; 
FIG. 4, portion of fruiting ament, X 10. 


MAR 


& 1946 


Rhodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL ! Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. March, 1946. No. 567. 


CONTENTS: 


Contributions from the Gray Herbarium of Harvard University— 
No. CLX. Technical Studies on North American Plants. 
M ue EAA (continued): — SLUT ASTU 41 


Observations on two Ecological Races of Allium tricoccum in 
Kalamazoo County, Michigan. Clarence R. Hanes and 
EN CORDE uL IAEA. ICE a 61 


Muhlenbergia setosa an Untenable Name. F.J. Hermann. ...... 63 


Does Habenaria cristata still grow in New England? 
MN Lille. es FONE a 64 


The New England Botanical Club, Ine. 


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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, 
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(if available) of not more than 24 pages and with 1 plate, 40 cents, numbers of 
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Address manuscripts and proofs to 
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Subscriptions (making all remittances payable to RHODORA) to 


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by Merritt LYNDON FERNALD and ALFRED CHARLES KINSEY 
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Section Axillares, by M. L. Fernald. 183 pp., 40 plates, 31 maps. 1932. 
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j- JOIO 


Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. March, 1946. No. 567. 


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


TECHNICAL STUDIES ON NORTH AMERICAN PLANTS 


M. L. FERNALD 


(Continued from page 40) 


In fact, Salix $ Reticulatae has strongly diverged in Newfound- 
land and the adjacent Labrador Peninsula from orthodox 
characters of the section, Schneider in Bot. Gaz. lxvii. 44 (1919) 
finding “the characters of the RETICULATAE . . . further 
changed by the inclusion of S. lezolepis [endemic in western New- 
foundland] with glabrous ovaries”. S. ledolepis (PLATE 1005), 
with habit of very coarse-stemmed S. reticulata but with strictly 
glabrous bracts and ovaries, is endemic, so far as known, on Table 
Mt., Port-au-Port Bay, 150 miles south of the southern known 
limit of S. reticulata var. semicalva. In the same general area 
with the embarrassing S. leiolepis the coarse and usually upright 
S. vestita 1s highly complex. Furthermore, this very definite 
coarse species of the Reticulatae, with very short petioles, the 
coriaceous leaf-blades usually heavily clothed beneath with dense 
and lustrous silky white hairs, and with staminate aments 1-1.5 
em. long, has diverged on Eskimo Island, west of the Straits of 
Belle Isle, and locally on walls of the Shickshock Mts. of the 
Gaspé Peninsula, as S. vestita, var. psilophylla Fernald & St. John 
in Vict. Mem. Mus. Mem. 126: 44 (1922), with the membrana- 
ceous leaves glabrescent beneath, the staminate aments 1.7-2.5 
em. long. Again, along the Straits of Belle Isle the Reticulatae 
have thrown off another endemic, S. jejuna Fernald in RHODORA, 


42 Rhodora [Marcu 


xxviii. 177 (1926), this tiny shrub (PLATE 1006) differing from 
S. reticulata in its very short (instead of long) petioles and short- 
(instead of long-) peduncled fruiting aments, and in the very long 
ascending villi of the papillate capsule, and from the Newfound- 
land S. reticulata, var. semicalva still further in its fuscous or 
dark purple very pubescent bracts. Furthermore, this unique 
little species of northernmost Newfoundland may flower twice in 
the same summer. In June or early July the fruits are ripe but 
one of the original collections, of July 16, had already fruited, 
while the new, vigorous shoots were already producing new 
flowering aments, without waiting for the next summer. Since 
neither of the endemic Newfoundland species of $ Reticulate have 
been illustrated, I am showing them in PLATES 1005 and 1006. 


S. vesTITA Pursh, forma mensalis, f. nov., trunco prostrato; 
foliis 1-1.6 em. longis; amentis fructiferis 5-6 mm. longis.— 
NEWFOUNDLAND: mossy knolls on the limestone tableland, alt. 
200-300 m., Table Mountain, Port-au-Port Bay, June 16 and 
17, 1914, Fernald & St. John, no. 10,824, specimens distributed 
under an identical but unpublished varietal name. 


Forming dense prostrate mats 1.5 dm. broad, with tiny leaves 
only 1-1.6 em. long, and fruiting aments 5-6 mm. long; in strong 
contrast with the erect or ascending typical Salix vestita (up to 1 
m. high) which has leaves 1.5-7 cm. long, the fruiting aments 
0.6-1.5 cm. long. 

The section Reticulatae Fries is unique among the diandrous 
Salices. In the tremendously extensive series of sections of 
diandrous willows with persistent bracts the aments or their 
supporting branchlets are axillary; terminal or subterminal, and 
the flowers are subtended by 1 or 2 slender to stout simple glands 
ornectaries. In the round-, roundish- or obovate- and reticulate- 
leaved § Reticulatae the peduncled aments are falsely terminal, 
borne just below the tip of the branchlet and on the side of the 
stem opposite the terminal leaf; and the glands of both staminate 
and pistillate flowers often form a false disk (as in Populus) with 
the margin lobed. "Thus making a transition in its gland and in 
several other characters to Populus, the section was set up by A. 
Kerner in 1860 as a genus, Chamitea. Of this differentiation the 
late Professor C. E. Moss wrote in his Cambridge British Flora, 
ii. 25 (1914). 


1946] Fernald,—Studies on North American Plants 43 


“S. reticulata possesses so many remarkable characters, showing it to 
be, in spite of the great difference in habit, intermediate in several respects 
between Populus and species of Salix in general, that there is little wonder 
that Kerner . . . suggested it should be placed in a new genus. How- 
ever, the remarkable characters possessed by 5. reticulata are so distributed 
among the other more primitive species of Salix that its generic separation 
from them cannot be maintained; and indeed Kerner himself at a later 
date accepted this view. The characters by which S. reticulata recalls 
Populus are the suckering habit, the long petioles, the broad laminae, 
and the perianthoid nature of the nectary. In its androecium, however, 
it has become a thorough Salix, more so even than S. pentandra, which 
has rather broad laminae, a double nectary, and, as a rule, 5 stamens at 
least. It seems to us that S. pentandra and S. reticulata diverged long 
ago from a primitive Salicalian stock, that each has retained a few of the 
Populus-like characters which this ancestral hypothetical group pos- 
sessed, and that each of these species or their ancient allies have given 
rise to the other species of Salix, some of which . . . exhibit interesting 
features of convergent development." 


The very primitive $ Reticulatae consists of only a few localized 
species: (1) S. reticulata of arctic-alpine range on calcareous soils, 
extending south to the higher mountains of Eurasia and in 
America in very local areas to northwestern Newfoundland 
(as var. semicalva), shores of Hudson Bay and southern Alaska 
and the Aleutian Islands; (2) S. vestita Pursh, with localized 
varieties, of the Labrador Peninsula, Newfoundland, Anticosti 
and Gaspé, west side of Hudson Bay, Cordilleran region of south- 
ern Alberta and British Columbia to northern Montana and 
eastern Oregon; (3) S. leiolepis and (4) S. jejuna, Newfoundland 
endemies; (5) S. nivalis Hook. of alpine regions of the Rocky 
Mts.—evidently a primitive section, consisting only of a few dis- 
joined relicts. 

'The pentandrous willows, on the other hand, such as those of 
$8 Nigrae, Pentandrae and Bonplandianae, are relatively southern 
(some even tropical) and their species, S. nigra Marsh., Hum- 
boldtiana Willd., lucida Muhl., Bonplandiana Kunth, etc., have 
broadly continuous ranges. Although presumably, as Moss 
pointed out, of as great antiquity as $ Aeticulatae, both series 
showing primitive characters, the pentandrous and chiefly more 
austral willows show no more evidence of relict-endemism than 
do the relatively modern diandrous species, such as S. rigida 
Muhl., S. humilis Marsh., S. discolor Muhl. or S. Bebbiana 
Sargent. § Reticulatae, however, has remained somewhat static 
and relict-endemism is one of its striking peculiarities. In view 


44 Rhodora {Marcu 


of its concentration in northern and western Newfoundland and 
its association there with hundreds of other relict-species of both 
plants and animals, those who believe the present distribution of 
nonaggressive plants and animals of as great or greater signifi- 
cance as the remote occurrence of chance fossils, find themselves 
unable to subscribe to the insistence of certain geologists and 
others, that life on Newfoundland and in adjacent areas was 
wholly obliterated by Wisconsin ice. It would be most difficult 
to demonstrate that in that area the relatively modern sections 
of diandrous Salix have in a few thousand years given rise to 
localized shrubs with more primitive floral characters, 


SALIX, § Uva-ursi, sect. nov., a § Herbaceae Borrer differt 
trunco valde ligneo vix subterraneo, ramulis valde foliosis; 
foliis firmis nec rotundatis nec valde reticulatis subtus albidis; 
amentis multifloris; bracteis valde sericeis; stamine plerumque 1. 
Tyre S. Uva-ursi Pursh. 

It is most difficult to see any close relationship of the eastern 
boreal American Salix Uva-ursi and the cireumpolar S. herbacea 
L. The latter has its trunks and main branches subterranean, 
stoloniferous and rooting at the nodes, only the short ascending 
filiform branchlets above ground, these bearing 2-4 reticulate 
rounded slender-petioled leaves which are green on both sides, 
and subterminal 2-8-flowered tiny aments, with nearly glabrous 
bracts, and the staminate flowers with 2 stamens. S. Uva-ursi 
is a strongly ligneous prostrate shrub, forming extensive super- 
ficial and very leafy carpets; the firm leaves not rounded, whit- 
ened beneath and not conspicuously reticulate, also short- - 
petioled; the many-flowered aments with long-silky bracts; the 
stamen solitary (rarely 2). 


S. ARCTICA Pallas, var. antiplasta (Schneider), comb. nov. 
S. anglorum Cham., var. antiplasta Schneider in Bot. Gaz. lxvi. 
134 (1918). 

SALIX, $ Argyrocarpae, sect. nov. Frutex 0.2-1.7 m. altus; 
foliis subtus micaceo-sericeis; stipulis minutis fugaceis; amentis 
fructiferis laxis; capsulis micaceo-sericeis; pedicellis elongatis 
glandulas duas 3-4-plo superantibus; staminibus 2. Typr S. 
argyrocarpa Anders. 


The boreal and alpine Salix argyrocarpa of northeastern 
America, like its associate, S. Uva-ursi, stands so far apart from 
other willows that it deserves a place in the system of Sections. 


Rhodor: Plate 1005 


Photo. B. G. Schubert 


SALIX LEIOLEPIS, all figs. from TYPE: FIG. 1, portion of shrub, X 1; FIG. 2, branchlet 
and portions of leaves, X 5; FIG. 8, portion of fruiting ament, showing glabrous bracts 
and capsules, X 10. 


Rhodora Plate 1006 


Photo. B. G. Schubert 


SALIX JEJUNA: FIG. 1, two portions of TYPE, X 1; FIG. 2, a small dense shrub, X 1; FIG. 
3, branch, X I, showing 2nd flowering in midsummer; FIG. 4, expanding bud and stipule 
(left), X 10; ria. 5, fruiting ament and leaf, X 5; FIG. 6, portion of fruiting ament, 
showing villous bracts and capsules, 10. 


1946] Fernald,—Studies on North American Plants 45 


Andersson, Bebb and Schneider have tried to place it in some 
section and the latter close student of the genus finally left it 
unanchored. 

S. GLAUCOPHYLLOIDES Fernald, forma lasioclada, f. nov., 
ramulis persistenter griseo-velutinis. Type from Robinson’s 
Brook, southwestern Newfoundland, August 10, 1930, Rachel B. 
Kennedy, no. 470, in Herb. Gray; flowering material from the 
same shrub, coll. June 7, 1930, no. 254. 

Differing from the glabrous- and lustrous-branched typical S. 
glaucophylloides in its densely gray-velvety branchlets. Found 
. through much of the range of typical S. glaucophylloides. 

S. GLAUCOPHYLLOIDES Fernald, var. albovestita (C. R. Ball), 
comb. nov. S. glaucophylla Bebb, var. albovestita C. R. Ball in 
Journ. Wash. Acad. Sci. xxix. 492 (1939). 

Unfortunately the name Salix glaucophylla Bebb (1881) is 
antedated by the same name for quite different species by Besser 
(1822) and by Andersson (1851). Typical S. glaucophylloides, 
occurring on gravelly shores in calcareous areas from Newfound- 
land to northern Ontario, south to the Gaspé Peninsula, northern 
New Brunswick and northern Maine, is a coarse shrub or small 
tree up to 5 m. high, its oblong to lanceolate or ovate leaves 
glaucous beneath and lustrous above, these about half-grown at 
anthesis, the aments subtended by 3-5 leaves. Its pistillate 
aments are dense, in maturity 2-6 cm. long, the capsules on 
pedicels only 1-1.5 mm. long. Var. glaucophylla (Bebb) Schnei- 
der, localized about the Great Lakes, is a low shrub (1-2.5 m. 
high) its aments expanding before the leaves are well grown, the 
pistillate aments lax and subremotely flowered, becoming 6-10 
cm. long, with fruiting pedicels 2-4 mm. long. Var. albovestita 
is similar to the latter and found on dunes of the Great Lakes 
from New York and southern Ontario to Michigan. Its branch- 
lets are densely pubescent and the young (sometimes the old) 
leaves are clothed with dense white pubescence. 

8. HUMILIS Marsh., var. hyporhysa, var. nov. Frutex 1-3 m. 
altus; ramulis fertilibus 2-5 mm. crassis; foliis glabratis vel 
subtus sparse puberulis, subtus valde rugoso-reticulatis, lamina 
matura 0.7-2 (-3) em. lata; amentis masculis 1-3 em. longis 
1-2.3 em. crassis; amentis fructiferis 2-8 cm. longis.—S. humilis, 
var. rigidiuscula sensu Rob. & Fern. in Gray, Man. ed. 7: 362 
(1908) not the basic S. humilis, var. longifolia, f. rigidiuscula 
Anders. (1897).— The commoner variety southward, from Florida 


46 Rhodora [Marcu 


to eastern Texas, north on or near the coastal plain to eastern 
Connecticut, Long Island, New Jersey and’ eastern Pennsyl- 
vania, and inland to West Virginia, Ohio, southern Michigan, 
southern Wisconsin, Iowa and Oklahoma. Tyre from NEw 
Jersey: open thickets bordering brackish marshes, Manahawkin, 
Ocean Co., July 23, 1923, Bayard Long, no. 28,011 (in Herb. 
Gray.). 

S. Humils, var. microphylla (Anders.), comb. nov. <S. tristis 
Ait. Hort. Kew. iii. 393 (1789). S. tristis microphylla Anders. in 
Öfv. Svensk. Vet.-Akad. Frósh. xv. 126 (1858). S. humilis, var. 
tristis (Ait.) Griggs in Proc. Ohio Acad. Sci. iv. 301, t. x (1905). 

S. HUMILIS, vàr. MICROPHYLLA, forma tortifolia, f. nov., foliis 
"valde spiraliter tortis.—MassAcnusETTS: Plymouth, Wm. Oakes, 
with manuscript label bearing an unpublished varietal name 
(rype in Herb. Gray.); dry hill, Plymouth, Sept. 22, 1853, Wm. ` 
Boott; another sheet, with copied (not original) label, “Tpswich, 
Masaschusetts ex herb. William Oakes", with typical flat-leaved 
and twisted-leaved branches mixed. 


Oakes’s material was evidently collected prior to 1848. The 
type-sheet has above his label the stamped memorandum 
“Manual, 1847”. Gray’s Manual 425 (1848) has under 5. tristis 
the note “A variety occurs with very small and rigid contorted 
leaves." 


S. nuMILIS, var. MICROPHYLLA, forma curtifolia (Fernald), 
comb. nov. S. tristis, forma curtifolia Fernald in RHODORA, 
xxxvi. 195 (1934). 

S. HUMILIS, var. MICROPHYLLA, forma festiva (Fernald), comb. 
nov. S. tristis, forma festiva Fernald in RHODORA, |. c. (1934). 

S. anACILIS Anders., var. textoris, var. nov., capsulis ad 9 mm. 
longis; foliis maturis glabratis 4-10 em. longis ad 2 cm. latis 
evidenter serrato-dentatis, dentibus apice glanduliferis. (S. 
petiolaris sensu Pursh and later American authors, not J. E. 
Smith)—Southern Quebee to Manitoba, south to New Bruns- 
wick, New England, northern New Jersey, northeastern and 
central Pennsylvania, Ohio, Indiana, Illinois, northern Iowa and 
Nebraska. Type: Massacuuserts: border of boggy meadow 
near Concord River, Bedford, May 11 and June 30, 1930, Fernald, 
Weatherby & Anderson in Pl. Exsicc. Gray. no. 447 (in Herb. 
Gray.; risorYPES in many herbaria). 

Typical Salix gracilis is smaller and generally more northern, 
occurring from Quebec to Alberta, south to northern Massachu- 
setts, western Connecticut (local), northern New York, northern 
Michigan, northern Wisconsin and Minnesota. It is S. petio- 
laris, vars. rosmarinoides (Anders.) Schneid., and angustifolia 


1946] — Fernald,—Studies on North American Plants 47 


Anders., characterized by capsules only 5-7 mm. long, and leaves 
entire or only obscurely denticulate, the mature ones 2.5-7 cm. 
long and 3-11 mm. broad. | 

Var. textoris (of the basket-maker) is so named because of the 
memoranda by Joseph Barratt (Salic. Amer.) and others, Barratt 
(1840) stating that “Mr. Hopkins, an experienced basket-maker 


. assures me that the green osier . . . furnishes the best 
twigs of any Willow he knows . . . The twigs are hard, tough, 
and elastic, and twist well for handles . . . It furnishes long, 


smooth twigs with small buds; the twigs are less tapering than is 
usual, which enhances their value to the basket-maker." 

It seems extraordinary that the identity of our low shrub, 
Salix gracilis, should, for more than a century and a quarter, 
have been confused by all students of Salix with the British tree, 
S. petiolaris J. E. Smith, Trans. Linn. Soc. vi. 122 (1802), Engl. 
Bot. xvi. t. 1147 (1803) and Fl. Brit. iii. 1048 (1804), ete. Our 
S. gracilis is a slender shrub with erect green to olive-brown 
tenuous and flexible branches 1-3 m. high; aments with narrow 
(linear-lanceolate to narrowly oblanceolate or spatulate) yellow- 
ish or pale brown bracts, leaves linear or narrowly lanceolate, 
entire to short-serrate-dentate, and 2.5-10 cm. (1-4 inches) 
long by 3-20 mm. (14-34 inch) wide, with stipules usually quite 
wanting, very rarely present on the sprouts but then minute and 
caducous. S. petiolaris, at first known from meagre material 
sent by Dickson from Scotland, was soon better known and the 
treatment by Forbes, Salict. Woburn. 45 t. 23 (1829), gave a 
good account of it: *A bushy tree, with slender, spreading [not 
strongly ascending] purplish, or dark-brown [not green or oliva- 
ceous] branches. Leaves about 4 inches long, and nearly 1 broad 
[in S. gracilis 1-4 inches long, L£-34 inch broad]. Stipules 
lanceolate, serrated [in S. gracilis wanting] . . . Scales rounded, 
notched [in S. gracilis elongate, entire, and yellowish; Smith's 
original diagnosis said “black, hairy, obovate, often notched"]. 

. Stalk of the germen as long as the adjoining scale", though 
in the plate shown as shorter [in S. gracilis much longer]. Forbes's 
beautiful plate would scarcely be taken as made from our slender 
and upright shrub; and European specimens, distributed as S. 
petiolaris, show a dense, curling, soft pubescence on the leaves 
and young branchlets, whereas the pubescence of young leaves 


48 Rhodora [Marcu 


(rarely if ever occurring on the branchlets) of S. gracilis is minute, 
silky and closely appressed. Although Schneider talked all 
around the subject in Journ. Arn. Arb. ii. 16-24 (1920), there is 
no indication in his discussion that he actually compared our 
species with true S. petiolaris. The latter name seems to have 
been wholly misapplied to our material. 


EXPLANATION OF PLaTEs 995-1006 


PraATEsS 995 and 996, SALIX RIGIDA Muhl. (S. cordata Muhl.). PrATE 995: 
FIG. 1, leaf of TYPE of S. rigida, X 1, after Muhlenberg; ria. 2, fruiting branch, 
X 1, from Hinsdale, New Hampshire, May 15, 1919, C. F. Batchelder; Fria. 3, 
portion of young pistillate ament, showing characteristically divergent flowers, 
X 5, from Lebanon, New Hampshire, May 4, 1884, G. Č. Kennedy; FIG. 4, 
fruiting ament, to show small basal bracts, X 44, from Buckland, Massachu- 
setts, May 21, 1906, F. F. Forbes; ric. 5, portion of latter ament, to show 
elongate pedicels, X 5. PLAare 996: rra. 1, leaf of S. cordata Muhl., X 1, 
after Muhlenberg; ric. 2, staminate flowering branch, X 1, from Shelburne, 
New Hampshire, May 21, 1920, Walter Deane; ric. 3, pistillate flowering 
branch, X 1, from Lebanon, New Hampshire, May 4, 1889, Kennedy; FIG. 4, 
portion of half-mature pistillate ament, showing long pedicels, X 5, from Hins- 
dale, New Hampshire, May 15, 1919, C. F. Batchelder. 

PLATES 997-1000, S. corpata Michaux (S. adenophylla Hook.). PLATE 
997: ria. 1, type of S. adenophylla, X ca. V$; ria. 2, margin of leaf of TYPE of 
S. adenophylla, showing porrect gland-tipped teeth, X 10; rra. 3, tip of leafy 
branch, X 41$, from Natashquan River, Saguenay County (“Labrador”), 
Quebec, July, Aug., 1912, C. W. Townsend; rig. 4, fruiting ament, with 
foliaceous bracts, X 44, from Townsend specimen. PLATE 998, rias. 1, 2, 3 
and 6, from specimens from Michaux's TYPE-REGION, Lake St. John, Quebec: 
FIG. 1, leafy tip, X 46, from Vauvert, Lake St. John, Victorin, no. 16,362, as 
S. adenophylla; ria. 2, tip of leafy shoot, X *$, from Roberval, Lake St. John, 
July 23, 1895, as S. adenophylla, J. G. Jack; ria. 3, stipules and leaf-bases, 
X 5, from no. 16,362; FIG. 4, stipule and leaf-base, X 5, from Rivière Petite 
Cascapedia, Gaspé Pen., Quebec, Victorin, Rolland & Jacques, no. 33,846; 
FIG. 5, leaf-margin, X 10, from Fort Fairfield, Maine, Sept. 19, 1900, Fernald, 
as S. adenophylla; r1G. 6, leaf-margin, X 10, from no. 16, 362; ric. 7, s'aminate 
ament, X 414, from Moosonee, mouth of Moose River, James Bay, Ontario, 
Dutilly & Lepage, no. 14,002. Pr ATE 999: rra. 1, staminate flowering branch, X 
4£, from Moosonee, mouth of Moose River, James Bay, Ontario, Dutilly & 
Lepage, no. 14,002; ria. 2, pistillate flowering tip, X *$, from Rupert House, 
Ungava, Dutilly & Lepage, no. 14,032; rra. 3, fruiting ament, X 4$, from no. 
14,032; ria. 4, portion of unexpanded staminate ament, showing blackish bracts, 
X 10, from no. 14,002; ria. 5, portion of expanded staminate ament, X 10, from 
no. 14,002. Prate 1000: rro. 1, fruiting branch, X 1, from Wellington, Ontario, 
June 3, 1902, James Fowler, as S. adenophylla; ria. 2, lower surface of mature 
glabrate leaf, showing delicate venation, X 10, from Michaux’s TYPE-REGION, 
Ile-aux-Couleuvres, Lake St. John, Quebec, Victorin, no. 16,371; ria. 3, portion 
of immature pistillate ament, to show short pedicels, X 10, from Rupert House, 
Dutilly & Lepage, no. 14,032. 

Ptares 1001 and 1002, S. syrticota Fernald. Puatre 1001, both figs., 
from TYPE, Lake Michigan, near Chicago, Bebb, Herb. Sal. no. 2, as S. adeno- 
phylla: ria. 1, pistillate, and rra. 2, staminate branch, X 44. Puare 1002: 
FIG. 1, portion of stipule, petiole and leaf-base, X 5, from Saugutuk, Michi- 
gan, August 15, 1896, C. F. Wheeler; ria. 2, petiole and leaf-base, X 5, from 
Dune Park, Indiana, Umbach, no. 95; rra. 3, portion of stipule, X 5, from no. 
95; FIG. 4, lower surface of mature leaf, showing venation, X 10, from New 


1946] Fernald,—Studies on North American Plants 49 


Buffalo, Michigan, Lansing, no. 3265; FIG. 5, portion of flowering pistillate 
ament, X 5, from TYPE; FIG. 6, portion of staminate ament, X 5, from TYPE. 


Puates 1003 and 1004, S. RETICULATA L. and var. sEMICALVA Fernald. 
PLATE 1003: FIG. 1, portion of staminate ament, showing villous-based and 
blackish bracts, X 10, of typical S. nEgTICULATA from Dovre, Norway, W. 
Boeik (?). Frias. 2-4, var. SEMICALVA: FIG. 2, portion of TYPE, X 4; FIG. 3, 
staminate ament and lower surface of leaf, X 3, from TYPE; FIG. 4, portion of 
staminate ament, showing pale and glabrous bracts, X 10, from TYPE. PLATE 
1004, ric. 1, portion, X 10, of fruiting ament of the typical S. RETICULATA 
from Torne Lappmark, July 19, 1927, Samuelsson & Zander. Fias. 2-4, var. 
SEMICALVA, all from Gargamelle Cove, Ingornachoix Bay, Newfoundland, 
Fernald, Long & Fogg, no. 1580: ria. 2, fruiting plant, X 44; FIG. 3, fruiting 
ament, X 5; FIG. 4, portion of latter, to show pale glabrous bracts : and sparsely 
pubescent capsules, AO: 

PraTE 1005, S. LEroLEPIs Fernald, all figs., from TYPE: FIG. 1, portion of 
plant, X 1; FIG. 2, branchlet and portions of leaves, X 5; ria. 3, portion of 
ament, to show glabrous bracts and capsules, X 10. 

PraATE 1006, S. zEJUNA Fernald: ric. 1, two portions of type, X 1; FIG. 2, 
a denser plant, X 1, from Four-Mile Cove, Straits of Belle Isle, Newfound- 
land, Fernald, Wiegand & Long, no. 27,949; FIG. 3, branch, X 1, showing 2nd 
flowering in midsummer, from east of Big Brook, Straits of Belle Isle, New- 
foundland, July 16, 1925, Fernald, Wiegand & Hotchkiss, no. 27,986; Fria. 4, 
expanding bud and stipule, X 10, from no. 27,986; ric. 5, fruiting ament and 
leaf, X 5, from no. 27,949; rra. 6, portion of same ament, showing villous 
bracts and capsules, X 10. 


III. NOMENCLATURAL TRANSFERS IN POLYGONUM 


POLYGONUM AMPHIBIUM L., var. STIPULACEUM Coleman, forma 
hirtuosum (Farwell), comb. nov. P. amphibium, var. margina- 
tum, forma hirtuosum Farwell in Papers Mich. Acad. Sci. i. 93 
(1923). 

P. AMPHIBIUM, var. STIPULACEUM, forma simile, f. nov., terres- 
tre vel subterrestre, ramis adscendentibus glabris vel minute 
pubescentibus; ochreis cylindricis; foliis lanceolatis, breviter 
petiolatis. "l'vPE from moist open meadow, Lisbon, New York, 
June 25, 1916, O. P. Phelps, no. 1551 (in Herb. Gray). 

P. AMPHIBIUM, var. STIPULACEUM, forma fluitans (Eaton), 
stat. nov. P. amphibium, var. a. natans Michx. Fl. Bor.-Am. i. 
240 (1803), not Moench, Enum. Pl. Hass. 189 (1777). P. 
natans (Michx.) Eaton, Man. ed. 3: 400 (1822). P. amphibium, 
var. aquaticum Torr. Fl. No. Mid. U.S. i. 404 (1824), not Leysser, 
Fl. Hals. ed. alt. 95 (1783). P. fluitans Eaton, Man. ed. 6: 274 
(1833). Persicaria fluitans (Eaton) Greene, Lfts. i. 26 (1904). 


In Ruopona, xxvii. 125-130, 146-152 and 156-166 (1925) 
Stanford discussed very clearly the heteromorphic series which 
at various times and by various authors has been treated as 
Polygonum amphibium L., his conclusions including, among 
other points, the segregation of the aquatic plant with thick and 
relatively short spikes and glabrous peduncles as a strictly North 


50 _Rhodora (Marcu 


American species, P. natans (Michx.) Eaton, with terrestrial 
phases. His main arguments for separating these from the 
Eurasian P. amphibium were the facts that in America the ter- 
restrial form often has the summit of the ochrea flaring into a 
horizontally divergent foliaceous and  bristly-ciliate flange, 
which does not occur in the Old World; that the terrestrial forms 
of the American plant have a less harsh leaf-margin; that the 
floating leaves of the American plant are more elliptic or elliptic- 
oval than in that of the Old World; that in true P. amphibium 
"the lateral veins of mature leaves [are] nearly straight and 
meeting the mid-vein nearly at right angles", whereas in the 
American P. natans they are more curved and meet “the midvein 
at an angle of about 60?" (Stanford, pp. 157, 158); and the och- 
reolae of P. natans are narrower and more tapering than in true 
P. amphibium. Other differences, measurements of achenes, 
calyx, etc., break down in the two series and the angle by which 
the veins join the midrib proves to vary too much in both series. 
So far as I can make out the really significant differences are the 
more lance-oblong or narrowly trowel-shaped leaf in P. amphib- 
ium, the harsher and shorter pubescence of the terrestrial foliage, ' 
the broader ochreolae and the more slender and often more 
elongated spike of the Eurasian series, with a tendency to longer 
peduncles. These are all relative characters, whereas the flowers 
and achenes are so similar that I am forced back to the long- 
established uniting of the two series as a single circumboreal 
species, P. amphibium L. 

The North American variety cannot take the first varietal 
name given it, P. amphibium, var. a. natans Michx. (1803), 
nomenclatural basis of P. natans (Michx.) Eaton (1822), for 
there was already a var. natans Moench (1777) for the floating- 
leaved European plant. The next varietal name for our plant, 
P. amphibium, var. aquaticum Torr. (1824) likewise duplicated 
an identical name given the aquatic European plant in 1783. 
Singularly enough, the first available varietal name for the Amer- 
ican series seems to be one applied to the most distinctively 
American phase of the species, var. stipulaceum Coleman, Cat. 
Fl. Pl. S. Pen. Mich. 32 (1874). Since Coleman’s work is a 
relatively scarce one I here give his account: Under P. am- 
phibium he had the conventional var. aquaticum, ascribed to 


1946] Fernald,—Studies on North American Plants 51 


Linnaeus, with P. fluitans Eaton as a synonym, and var. terrestre 
Willd. Then came 


Var. stipulaceum, with leaves and flowers like P. amphibium; the 
leaves, possibly a little narrower, and with salver form stipules, like P. 
orientale, found growing in sandy soil near Gd. Rapids. 


That was a clear description of the distinctively North Ameri- 
can terrestrial plant which had been described as P. Hartwrightii 
Gray in Proc. Am. Acad. viii. 294 (1870), the plant subsequently 
called P. amphibium, var. Hartwrightii (Gray) Bissell in Rnopo- 
RA, iv. 104 (1902) and P. natans (Michx.) Eaton, forma Hart- 
wrightii (Gray) Stanford in Rnopona, xxvii. 160 (1925). As the 
first valid varietal name, var. stipulaceum has to be taken up to 
include the many vegetative American forms. ‘This might seem 
to those who think of the American plant as the aquatic phase 
with oblong long-petioled floating leaves, which, becoming 
stranded, will change rapidly to the terrestrial phases, like put- 
ting the cart before the horse. The floating transmutation, 
however, is the exceptional one. Over much of the area, where 
lakes and ponds are scattered, the terrestrial var. stipulaceum 
occupies thousands and thousands of square miles of swamp, 
meadow and swale, where it makes vast carpets and never gets 
the opportunity to ‘stretch into permanently standing water. 
To those who know such extensive areas var. st?pulaceum seems 
the normal development of the species. 

As to var. stipulaceum, forma simile, that plant of swale, 
meadow and shore, although simulating the Old World P. am- 
phibium, forma terrestre (Leers) Blake in Ruopora, xv. 164 (1913) 
and Moss, Camb. Brit. Fl. ii. 115 (1914), which was based 
nomenclaturally on P. amphibium, var. terrestre Leers (1775), is 
really a parallel form of var. stipulaceum without the foliaceous 
flanges. 

It is in some ways fortunate that the name P. fluitans Eaton is 
available as a nomenclatural basis for the formal name of the 
common lacustrine extreme of the species in America, the P. 
natans (Michx.) Eaton, forma genuinum Stanford in RHODORA, 
xxvii. 158 (1925), for, since the varietal names given this extreme 
by earlier authors were later homonyms, we should be faced by a 
large handful of Greene's so-called species and might have to 
transfer to the formal category for the common aquatic plant 


52 Rhodora [Marcu 


growing from Labrador to Alaska and south into the Northeast- 
ern, Central and Western States one of several inappropriate 
binomials, such as Persicaria purpurata Greene, P. mesochora 
Greene or P. oregana Greene. 

Amos Eaton started off his Polygonum fluitans (1833) in a 
somewhat contradictory manner. In 1822 he had published P. 
"natans (floating knotweed)" from ‘‘Whiting’s pond, 5 miles 
south of New Lebanon springs", saying definitely “It is the P. 
amphibium. Var. natans of Mx". In 1833, however, he changed 
his mind, renaming the plant of Whiting's Pond P. “fluitans, 
Ea. . . swimming knotweed" and saying “Finding this to be a 
new one, not var. natans of Mx. I give it à new name." There 
is nothing about the lacustrine plant of western Massachusetts 
and adjacent eastern New York (rvPr-region) to separate it from 
Michaux's P. amphibium, var. natans from Lake St. John, 
Quebec, as shown by the photograph of it before me. And even 
Greene, who saw many species where others see only one, ad- 
mitted the probable identity, saying in his typically sophisticated 
and plausible style, under his Persicaria fluitans (Polygonum 
fluitans Eaton): 

“Amos Eaton as early as 1840 [i. e. 1833] gave the name P. 
fluitans to what, from the description as well as the locality, we 
must conclude to have been that here described anew. I do not 
know where that St. John's Lake is which Michaux cites as the 
habitat of his var. natans; but I suspect it to be some northern 
lake now known by another name, and lying within the habitat 
of P. fluitans, in which case that may be an older, though a 
merely varietal designation which would in my view be of no 
consequence." 

Since Michaux's Polygonum amphibium, var. natans was the 
nomenclatural basis of P. natans (Michaux) Eaton (1822), the 
plant later (1833) becoming P. fluitans Eaton, Greene's dismissal 
of the name natans as ‘‘merely a varietal designation" showed 
woeful lack of knowledge of the literature for a self-styled his- 
torian. So did his ignorance of *Where that St. John’s Lake is 
which Michaux cites", for the big Lake St. John, lying at the 
head of the Saguenay, in the Districts of Lake St. John and 
Chicoutimi, was on the old route of early explorers from the 
lower St. Lawrence to Hudson Bay, was much mentioned by 


Rhodora Plate 1007 


Photo. B. G. Schubert 


Xxnis Bavanpi, all figs. from TYPE: FIG. 1, four plants, X 1; FIG. 2, two spikes, 
x 5; FIG. 3, leaf-tip, X 10; FIGs. 4 and 5, lateral sepals, X 10; FIG. 6, four seeds, 
x 10. 


Rhodora Plate 1008 


Photo. B. G. Schubert 


XNYRIS BREVIFOLIA: FIG. 1, bases of plants, X 1, from TYPE-REGION; FIG. 2, basal 
leaf, X 10; FIG. 3, group of spikes, X 1, from Michaux’s TYPE, after photo. by 
Cintract; FIG. 4, spike, X 5. 

X. FLABELLIFORMIS, all figs. from tsorypr: FIG. 5, base of plant, X 1; FIG. 6, tip of 
leaf, X 10; FIG. 7, spike, X 5. 


1946] Fernald,—Studies on North American Plants 53 


Michaux in his journals of exploration and was repeatedly cited 
as the type-locality of plants described by him, and it is shown 
(and named), almost north from the city of Quebec, as a roundish 
blue spot on all maps of Canada, or even of North America, 
which I find in recent atlases. Nevertheless some botanists 
express admiration of Greene and his methods. 

P. AnrFOLIUM L., var. pubescens (Keller), comb. nov. P. 
sagittatum, var. pubescens Keller in Bull. Soc. Roy. Bot. Belg. 
xxx?. 45 (1891). P. arifolium, var. lentiforme Fernald & Griscom 
in Ruopona, xxxvii. 167 (1935). 

It is too bad to be forced to abandon a diagnostically descrip- 
tive name, var. lentiforme (from the lentieular achene), for a non- 
distinctive one, since both typical southeastern Polygonum ari- 
folium and the northern var. pubescens (or lentiforme) are pubes- 
cent, the former much more so than the latter. 

When Dr. Robert Keller described the northern variety as a 
pubescent variety of P. sagittatum, in a paper entitled Remarques 
sur quelques espéces du genre Polygonum de l'Herbier de Jardin 
Botanique de l'État à Bruxelles, he unwittingly indicated the 
weakness of that famous herbarium in representative North 
American material and, incidentally, unfamiliarity with the 
standard floras and manuals of eastern North America, in which 
the Linnean P. arifoliwm (1753) was described with the pubescent, 
large, long-acuminate leaves and the hispid-glandular axis of the 
inflorescence which set off Keller's P. sagittatum, var. pubescens 
(a single specimen from Troy, New York) as “une variété bien 
caractérisée” of his true P. sagittatum, defined "foliis sagittato- 
lanceolatis nudis vel margine setulis ciliolatis subtus nervo inermi 
vel plus minusve aculeolato" ; var. pubescens "foliis sagittatis late 
ellipticis, longe acuminatis, . . . subtusin nervo medio nervisque 
secundariis aculeolatis, foliis infra densius pubescentibus pilis 
stellatis supra parce pubescentibus pilis. simplicibus adpressis 
pilisque stellatis". 

As early as 1788 Walter described P. arifolium "foliis hastatis 
pilosis magnis"; Elliott (1817) knew its stem “towards the sum- 
mit with capitate hair and a stellated pubescence. Leaves on 
long petioles, hastate, with the auricles acute, pubescent"; 
Torrey (1824) knew its “Leaves on long aculeate petioles . . . 
acuminate, with short scattered hairs on the upper surface, 


54 Rhodora [Marcu 


minutely papillose beneath” and to these characters he added in 
Fl. N. Y. (1843) “peduncles glandularly hispid”. Even if 
Keller did not know these and the more detailed American 
treatments of succeeding years he might have found Meisner in 
his Monograph (1826) defining P. arifolium “foliis hastatis 
acuminatis, utrinque adpresse pilosis", these characters repeated 
in Meisner's treatment in DeCandolle's Prodromus (1856), with 
the other distinctive character of Keller's P. sagittatum, var. 
pubescens, “pedunculis subglanduloso-hispidulis". Had Keller 
looked further he would doubtless have found that the flowers 
of the latter have 6 stamens and bifid style (the achene, conse- 
quently, biconvex), whereas his true P. sagittatum would have 
shown him 8 stamens and a trifid style (consequently a trigonous 
achene). 

Since the type and only specimen cited by Keller came from 
Troy, New York, it is obviously of the less pubescent northern 
plant which Griscom and I described as P. arifoliwm, var. lenti- 
forme, we not imagining anyone in these days referring members 
of that long and generally understood species to the very different 
P. sagittatum. Even the most inexperienced of our “Jumpers”’ 
would hardly argue for the specific uniting of P. sagittatum and 
P. arifolium, provided, of course, that he knew the plants and 
their morphological characters. 

P. cruiNopE Michx., forma erectum (Peck), stat. nov. Var. 
erectum Peck, N. Y. State Mus. Rep. xlvi. 129—repr. 49 (1893). 
Var: breve Peck, Bull. N. Y. State Mus. vi. 120 (1899). 

The low and upright plants ending in panicles and without 
twining tips, comparable with bush-beans, as contrasted with 
twiners. 

IV. NOVELTIES IN OUR FLORA 


(Plates 1007-1020) 


CAREX cRINITA Lam., var. brevicrinis, var. nov., a var 
typica recedit spicis foemineis 4-10 (plerumque 6-7) cm. longis 
densifloris omnino foemineis vel ad apicem masculis; squamis 
imis perigyneis aequantibus ad duplo longioribus, squamis 
superioribus perigyneis vix aequantibus ad paulum longioribus; 
perigyneis valde inflatis 3-4 mm. longis 2-3 mm. latis.—North 
Carolina to eastern Texas, north to southern New England, 
Kentucky and Missouri. The following are characteristic. 
MASSACHUSETTS: edge of cedar swamp, southwest of Beachwood, 


1946] Fernald,—Studies on North American Plants 55 


Cohasset, July 22, 1938, Griscom & Svenson (as C. Mitchelliana) ; 
maple swamp, Spring Hill, Harwich, July 30, 1919, Fernald & 
Long, no. 18,127 (as C. Mitchelliana). RnopE IsLanp: Provi- 
dence, May, 1845, Thurber. Connecticut: Bridgeport, June 8, 
1910, H. S. Clark; New Fairfield, July 19 and 20, 1912, Blewitt. 
DisrRIcT oF CoLuMBIA: May 30, 1899, Steele; May 22, 1890, 
Steele (as var. gynandra). VirGINIA: sphagnum swamp, 114 
miles northwest of Williamsburg, June 12, 1921, Grimes, no. 
3693; Powhatan Swamp, 1% mile southwest of Five Forks, 
James City Co., June 20, 1922, L. F. & F. R. Randolph; sphagnous 
boggy margin of spring-fed pond, Century House, northeast of 
Burgess, Dinwiddie Co., July 23, 1938, Fernald & Long, no. 8614; 
wooded alluvial bottomland of Rowanta Creek, near Rowanta, 
Dinwiddie Co., June 8, 1938, Fernald & Long, no. 8413 (TYPE in 
Herb. Gray.; isorvPE in Herb. Phil. Acad.); inundated swamp 
along Quarrel Creek, **Chamblis bigwoods", Seward Forest, near 
Triplett, Brunswick Co., May 10, 1945, Fernald, no. 14,795. 
NORTH CAROLINA: swamp, edge of Newbridge Creek, 2 miles 
from Currituck Sound, Currituck Co., July 1, 1922, Randolph & 
Randolph; marsh, Lake Raleigh, Wake Co., May 11, 1937, 
Godfrey; low field, 7 miles northwest of Chapel Hill, June 28, 
1927, Wiegand & Manning, no. 414. Kentucky: without 
stated locality, Short (as C. Pseudo-Cyperus, this corrected by 
Dewey). TENNESSEE: low, wet grounds, Wolf Creek, August 15, 
1900, Ruth, no. 497. Missounr: rare in low ground, Courtney, 
May 25, 1902, Bush, no. 1714. LouisIANA: western section, 
Hale. Texas: without stated locality, Wright. 


Carex crinita, var. brevicrinis has puzzled its collectors. Some 
of the northern collections were placed with the then recently 
redefined C. Mitchelliana M. A. Curtis!, because they would not 
go satisfactorily into typical C. crinita, in which the pistillate 
spikes only rarely have staminate tips, the long awns of the lower 
scales are two to three or four times as long as and the upper awns 
definitely longer than the relatively small perigynia (2-3-rarely 
3.5 mm. long and 1-2 mm. broad). Var. brevicrinis most often 
has the pistillate spikes with a staminate tip (as frequently in var. 
gynandra (Schwein.) Schwein. & Torr.), its lower scales are rarely 
twice as long and the upper ones shorter than to barely longer 
than the large and strongly inflated, mostly obovate perigynia, 
these 3-4 mm. long and 2-3 mm. broad. 

Although superficially suggesting var. gynandra, the usually 
more southern or coastwise var. brevicrinis differs at once in its 


1 See Weatherby in RHODORA, xxv. 17-20 (1923). 


56 Rhodora [Marcu 


smooth lower leaf-sheaths (those of var. gynandra being harsh), 
in its relatively longer lower foliaceous bract, in its relatively 
shorter scales and in its more strongly inflated and crumpled 
perigynia. In the relatively northern and upland var. gynandra 
the lowest foliaceous bract ranges from 1.2-4 (av. 2.5) dm. long, 
this averaging twice the length of the axis of the inflorescence. 
In var. brevicrinis, on the other hand, the lowest bract is from 
(2.5-) 3-4 (av. 3.6) dm. long, this averaging three and a third 
times the length of the axis of the inflorescence. 

Although Carex crinita, var. brevicrinis has been mistaken for 
C. Mitchelliana, it is really very different. C. Mitchelliana has 
the leaves and lower foliaceous bract only 2.5-9 mm. broad (in 
var. brevicrinis thinner and darker, 6-12 mm. broad) the lower 
bract 0.8-3 (av. 1.8) dm. long; the tight or scarcely inflated and 
definitely nerved granular-papillate ovate to ovate-lanceolate 
perigynia 2.5-3.5 mm. long and only 1.4-2 mm. broad. 


Xynis (8 BnEviroLrAE) Bayardi, sp. nov. (ras. 1007). Planta 
annua vel biennis; foliis pallide viridibus lanceolato- vel lineari- 
ensiformibus 1.5-4 cm. longis ad 4 mm. latis, plus minusve 
curvatis, membranaceis, margine albido minute denticulato, 
apice subaeuto breviter curvato; scapis filiformibus 1-3 dm. 
altis glabris; vaginis basilaribus scapi perbrevibus brunneis; 
spicis paucifloris ellipsoideo-ovoideis acutis vel subacutis 5-6 mm. 
longis 3—4.2 mm. crassis; bracteis intermediis elliptico-ovalibus 
brunneis margine albido-hyalinis, area dorsali viridi distincta; 
sepalis lateralibus liberis curvatis oblique lineari-lanceolatis 
3-3.5 mm. longis 0.3-0.5 mm. latis, ala carinali angustissima in 
parte tertia superiore minute denticulatis; seminibus ellipsoideo- 
fusiformibus 0.45-0.5 mm. longis 0.2-0.24 mm. latis, pallide 
stramineis apicibus brunneis.—Sussex County, VIRGINIA: wet 
sandy and peaty shore of Airfield Millpond, southwest of Wake- 
field, September 11, 1945, Fernald & Long, no. 14,922 (TYPE in 
Herb. Gray; isorvrE in Herb. Phil. Acad.). 


Xyris Bayardi (for one of its discoverers, BAYARD Lona) is the 
northernmost member of Xyris $ Brevifoliae, only three other 
members of this chiefly tropical section, mostly annuals and 
biennials, being known in the United States. From all three X. 
Bayardi is abundantly distinct. The basic X. brevifolia Michaux, 
Fl. Bor.-Am. i. 23 (1803), shown in our pL. 1008, Fics. 1-4, 
described from the low country of Georgia, is rare except in 
Florida. Its firm leaves (rics. 1 and 2) are very narrowly linear 


1946] Fernald,—Studies on North American Plants 57 


(1-2 mm. wide), arching to long attenuate tips, firm and glabrous- 
margined; whereas the very thin and membranaceous leaves of X. 
Bayardi are broader (up to 4 mm. broad), with short and barely 
acutish tips, the white margin minutely denticulate. The 
sheaths at the base of the scape in X. Bayardi are so short as 
scarcely to be seen, while in X. brevifolia (Fia. 1) they are pro- 
longed and commonly much overtopping the basal foliage. The 
fruiting spike of X. Bayardi is ellipsoid and acute or acutish and 
only 3-4.2 mm. thick; whereas X. brevifolia (ria. 3, X 1, from 
the Michaux TYPE) has the mature spikes oblate or depressed- 
globose and broad-tipped, finally 6-8 mm. thick. The very 
narrow (0.3-0.5 mm.) lateral sepals of X. Bayardi are minutely 
denticulate only toward the summit of the keel; the broader 
(0.6 mm.) sepals of X. brevifolia, to quote Malme in the North 
American Flora, have the “keel ciliate-scabrid from near the base 
to the apex". At its type-station (the only one yet known) X. 
Bayardi was mature, but with lingering marcescent corollas, on 
September 11th. Whether it begins flowering before summer we 
do not yet know. It is significant, however, that most of the 
material of X. brevifolia, whether in flower or in fruit, from Florida 
was collected in March, April and May, only exceptionally up to 
midsummer. 

A second annual species, occurring from Florida to Mississippi, 
| ds Xyris flabelliformis Chapman, Fl. So. U. S. 499 (1860), our 
PL. 1008, rias. 5-7. X. Bayardi has flabelliform clusters of basal 
leaves as in Chapman's species, but they are thinner, paler, 
broader and blunter than in X. flabelliformis, and with minutely 
denticulate, instead of entire margins. Furthermore, the basal 
sheaths of X. flabelliformis run up the scape (rra. 5), while in X. 
Bayardi they are almost hidden. The spike of X. Bayardi might 
be taken for that of X. flabelliformis (ria. 7) but its bracts are 
broader, more appressed and more rounded at tip. X. Bayard 
can hardly be forced into X. flabelliformis. 

The only other species of § Brevifoliae described from north of 
the Tropics is X. Drummond?i Malme from Alabama, described 
as a perennial with smooth leaves, sheaths extending up the 
scape, and lateral sepals 0.8 (in X. Bayard? 0.3-0.5) mm. broad 
and ciliate-scabrous on the keel. I have seen no material but it 
can hardly be made to include X. Bayardi. 


58 Rhodora [Marcu 


It was only by the rarest of good luck that Xyris Bayardi was 
discovered. Mr. Long’s and my discovery that in Sussex and 
Southampton Counties, Virginia, there are at least two large 
ponds (Airfield in Sussex, Whitefield in Southampton) which we 
had not previously known and which, during a dry season, have 
broad sandy and peaty beaches, was recounted in RHODORA, xlv. 
372 and 373 (1943). We discovered these ponds with their 
broad beaches (then “perhaps 50 feet wide up to the bushes") 
in early July, 1942. Then gasoline-rationing cut us off from 
working them. Two futile attempts (after heavy rains) were 
made by me to explore them later (see Ruopora, |. c. 381 and 
xlvii. 103-105), so that the end of gasoline-rationing was the 
signal for Long and me to hurry to our headquarters at Waverly 
and to try, in September last, for the pond-shores. Again alas! 
An abnormally rainy summer had completely overflowed White- 
field Pond, Brittle’s showed no beach, but on September 11th 
and 12th a very narrow rim, still wet from drowning, was visible 
at Airfield, this very soon obliterated by more rain. Luckily 
we got there in time for preliminary exploration of the little 
emersed belt of shore. Here had been the only known station 
for Rhynchospora filifolia between southeastern North Carolina 
and Cape May, New Jersey; that and other species collected in 
early summer indicated an unusual pond-shore for eastern 
Virginia. The southern shore of Airfield is the best we saw, the 
northern being more uniformly pretty swaley, with Panicum 
verrucosum, Cephalanthus occidentalis, Xyris caroliniana, X. 
ambigua Beyrich (in all sizes) and other aggressive plants crowd- 
ing out all others. Immediately we got Sabatia difformis (L.) 
Druce, the first from between North Carolina and eastern Mary- 
land. Then Eriocaulon decangulare, very local in Virginia. 
Psilocarya nitens, previously unknown between southeastern 
North Carolina and Cape May was locally abundant; and we 
were (and still are) utterly baffled by the members of the alliance 
of Hypericum canadense. I had thought that I had this series 
under control, but its behavior on the peaty shore of Airfield Pond 
is seriously disturbing the latest treatments. I may stave off 
the evil day until another season of observation. And here, 
nestled among the coarser vegetation were the whitish-green 
fan-like tufts of a tiny annual Xyris: the newly described X. 
Bayardi. We saw it in only one small area, a few rods wide, but 


1946] Fernald,—Studies on North American Plants 59 


further exploration another season, when the water is much 
lower, will, we hope, reveal it in quantity. Airfield Pond still 
needs a season of intensive botanizing! 


PEDICULARIS CANADENSIS L., var. Dobbsii, var. nov. (TAB. 
1010 et 1009, FIG. 2 et 3), vix cespitosa, caudicibus lateralibus 
horizontaliter divergentibus elongatis plus minusve repentibus.— 
New York to Minnesota, south to northern Florida, Alabama, 
Louisiana and eastern Texas. The following are characteristic. 
New York: along Babcock Trail near Jim Pond, Black Rock 
Forest, Orange Co., Raup, no. 7215; Taughannock Ravine and 
vicinity, Ulysses, F. P. Metcalf, no. 7148. New JERSEY: 
Columbus, Burlington Co., May 11, 1921, H. B. Meredith. 
West VIRGINIA: along East Fork of Greenbrier River, Pocahon- 
tas Co., Greenman, no. 252.  NonTH Carouina: Highlands, May 
27, 1901, E. E. Magee. FLonipA: without statement of locality, 
Chapman. | ONTARIO: Tobermory, Bruce Co., Krotkov, no. 7771. 
MicHiGAN: Carp Creek, Cheboygan Co., Ehlers, no. 392.  Ixpi- 
ANA: Jackson Tp., Wells Co., May 11, 1908, Deam. ‘TENNESSEE: 
oak barrens, 7 miles east of Crossville, Cumberland Co., May 13, 
1933, C. A. & Una F. Weatherby, no. 6254 (TYPE, in Herb. Gray); 
Jackson, Bain, no. 442. ALABAMA: about 3 miles north-north- 
east of Marion, Perry Co., Harper, no. 3702. WISCONSIN: 
Preble, Brown Co., May 19, 1887, J. H. Schuette. ILLINOIS: 
Mt. Morris, May 20, 1909, Sherff; about L4 mile east of western 
boundary of Henry County and 14 mile south of U. S. Route 6, 
July 1, 1945, Raymond J. Dobbs. MiNNESOTA: Clearwater Co., 
M. L. Grant, no. 2702; Spring Grove, Rosendahl, no. 276. Towa: 
Homestead, Iowa Co., May 11, 1925, Shimek. ARKANSAS: 
Little Rock, Demaree, no. 18,795. Louisiana: Perkins, Calcasieu 
Parish, Pennell, no. 10,212. Kansas: Leavenworth Co., Hitch- 
cock, no. 788. OkrAHOMa: Arkansas National Forest, Sebastian 
Co., E. J. Palmer, no. 39,309. Texas: Camp Fannin, 8 miles 
northeast of Tyler, Smith Co., /7. E. Moore, Jr., no. 673. 


Var. Dobbsii, which was called to my attention by Mr. Ray- 
MOND JOSEPH Dosss of Geneseo, Illinois, keen student of the 
flora of Henry County, is a striking departure in habit from the 
densely cespitose typical Pedicularis canadensis (PLATE 1009, 
FIG. 1). At the northeastern corner of the range of P. canadensis, 
whence, undoubtedly, came Kalm’s material which was described 
by Linnaeus, the densely crowded crowns are erect or strongly 
ascending (PLATE 1009, FIG. 1) and the large series before me 
from southern Quebec and New England shows no appreciable 
tendency to the lax to stoloniferous habit of var. Dobbsii. The 
densely cespitose extreme also extends westward and southward 


60 Rhodora [Marcu 


essentially to the limits of the specific range, except for the more 
western var. fluviatilis (Heller) Macbride, and it is general in the 
large New England series before me; it is evidently the only 
variety occurring in this area. Whether var. Dobbsii has a 
different habitat and whether it more generally has fusiform 
roots (as in the type) are matters for close field-observation. 
Such fusiform roots are apparently rare in typical P. canadensis— 
perhaps only seemingly so through careless collecting. 

UrnicULARIA JUNCEA Vahl, forma virgatula (Barnhart), stat. 
nov. U. virgatula Barnhart in Bull. Torr. Bot. Cl. xxxiv. 580 
(1908). Stomoisia virgatula (Barnhart) Barnhart in Britton & 
Brown, Ill. Fl. ed. 2, iii. 232, fig. 3875 (1913). 

Ever since I first began collecting with him in Virginia Mr. 
Bayard Long has insisted that Utricularia juncea and U. virgatula 
are phases of one species, comparable with U. subulata L. and its 
forma cleistogama (Gray) Fernald in RHopora, xxiii. 291 (1922). 
Although we usually find U. virgatula growing with U. juncea I 
have been reluctant to merge them; but on the shore of Airfield 
Millpond in Sussex County, Virginia we found in September 
last such perfect intergradation in all the large areas of the plants 
between large-flowered U. juncea and small-flowered U. virgatula, 
that I give up. Only by arbitrary sorting and very arbitrary 
exclusion of many intermediates can two piles of similar speci- 
mens be made. By this doubtful process some of our earlier 
collections have been forced apart. It is significant, therefore, 
that many numbers in the Gray Herbarium show that their 
collectors did not mechanically sort them into two or three piles: 
J. A. Allen from Atsion, New Jersey, Aug. 16, 1879, with smallest 
and transitional corollas; Commons, Laurel, Delaware, Aug. 19, 
1880, both large and smallest corollas; R. R. Tatnall, no. 2759 
from Kent Co., Delaware, large and intermediate corollas, the 
latter U. juncea, forma minima Blake in Contrib. Gray Herb. no. 
lii. 89 (1917); Heller, no. 1222 from Princess Anne Co., Virginia, 
both typical U. juncea and forma virgatula; and so on to Missis- 
sippi. The experience at Airfield Millpond finally convinced me. 


CHRYSOPSIS MARIANA (L.) Ell., forma efulgens, f. nov. ligulis 
nullis.—Sussex County, VirRGINIA: dry pine woods, State Game 
Sanctuary, northwest of Newville, Sept. 13, 1945, Fernald & 
Long, no. 14,996 (rvrE in Herb. Gray., rsorvrE in Herb. Phil. 
Acad.); dry pineland 3-4 miles west of Waverly, Sept. 16, 1945, 
Fernald & Long, no. 14,997. 


(To be continued) 


Rhodora Plate 1009 


Photo. B. G. Schubert 


PEDICULARIS CANADENSIS: FIG, 1, plant, X 24. 
P. CANADENSIS, var. DoBBSII: FIG. 2, characteristic base, X 34; FIG. 3, flowering 
raceme, X 1. 


Rhodora Plate 1010 


Photo. B. G. Schubert 


PEDICULARIS CANADENSIS, var. DonnBsit, both figs. from rype: FIG. 1, base of plant, 
showing fusiform roots (center), X 23; FIG. 2, fruiting raceme, X 1. 


1946] Hanes and Ownbey,— Allium tricoccum 61 


SOME OBSERVATIONS ON TWO ECOLOGICAL RACES 
OF ALLIUM TRICOCCUM IN KALAMAZOO 
COUNTY, MICHIGAN 


CLARENCE R. HANES AND MARION OWNBEY 


REPEATED observations on Allium tricoccum Ait. both in the 
wild and in the garden have revealed the existence of two well- 
defined ecological races of this species in Kalamazoo County, 
Michigan. One race prefers low moist woodlands, where a single 
patch may often occupy several acres of low ground. The other 
is found, usually, in small scattered patches only, in upland beech 
and maple woods. The two races rarely, if ever, occur together. 
For convenience in the following discussion they are referred to 
as “Race A" and “Race B", respectively. 


Race A.—Plants of Race A are in every way conspicuously larger than 
are those of Race B, both in the wild and under cultivation. The leaf- 
blades vary from 22 to 26 cm. long, and from 4 to 7 em. broad, and are 
more nearly elliptic in outline than are those of Race B. The sheaths and 
well-developed petioles are reddish, and the scape is from 25 to 34 cm. 
tall. From herbarium specimens, and from descriptions in existing 
manuals, it appears that this is the common race eastward, and, therefore, 
likely the typical element of the species. 


In Kalamazoo County, Race A is very abundant in Section 20, 
Prairie Ronde Township, where it occurs with swamp birch, 
tamarack, white elm, red ash, ete. It also occurs in moist soil in 
sections 14 and 33, Climax Township, in Section 19, Brady Town- 
ship, and along the Kalamazoo River in sections 27 and 28, 
Comstock Township. At Cooper’s Glen, north of Kalamazoo 
(Section 27, Cooper Township), it grows on a wooded hillside 
which may be somewhat springy. 


Race B.—The leaf-blades of Race B vary from 20 to 22 em. long, and 
from 1.3 to 3 em. broad. They are more lanceolate than elliptic in outline. 
The sheaths and petioles are greenish, and the latter are often very short. 
The scape is less than 25 cm. tall. 


Race B occurs principally in upland woods of beech and maple. 
It has been found in sections 4, 5, 16, 19, 22, 24, 26, and 30, 
Prairie Ronde Township, and in Section 18, Schoolcraft Town- 
ship. Mostly, it occurs on the western side of the county. In 
Section 30, Prairie Ronde Township, only, has it been observed 


62 Rhodora [MancH | 


to occur in marshy ground. It grows also in Porter Townsbip, 
Van Buren County. . 

In localities where Race B is the prevalent form, occasional 
plants with the typical narrow leaves, but with sheaths of a slight 
pinkish cast, may be found. Also, among the broad-leaved 
plants of Race A, infrequent individuals may fail to develop the 
characteristic reddish color in the sheaths and petioles. This is 
the extent to which intermediacy or intergradation between the 
two races may be said to occur. 

An interesting physiological difference between the two races 
is their differential development when grown side-by-side as they 
have been in the first author’s garden for the last eight years. 
Race A appears from a week to ten days earlier in the spring than 
Race B, even though the weather may be continuously warm. 
In 1941, for instance, Race A was 2.5 cm. above the ground on 
April 6, whereas Race B had not yet appeared on April 13. On 
April 18, Race A was 21 em. tall, while Race B stood only 9 em. 

The determination of the proper taxonomic designations for 
these two entities is complicated by the description in 1808 of 
Allium triflorum Raf., and the present inaccessibility of authentic 
material of either this or of A. tricoccum, if it still exists. The 
original description of neither is conclusive. Aiton stresses ‘the 
nude, semiterete scape, lanceolate-oblong, plane, glabrous leaves, 
globose umbel, and solitary seeds, while Rafinesque describes the 
stem as cylindrical, 3-flowered, and shorter than the plane, cunei- 
form, lanceolated, acute, multinervous leaves. Rafinesque states 
that A. triflorum came from North Pennsylvania, whereas North 
America alone is indicated as the source of A. tricoccum. In 
1770, this would have meant only the Atlantic slope. 

No systematic attempt has been made to determine the distri- 
bution of the two races outside of Kalamazoo County, although 
Mr. C. A. Weatherby writes that narrow-leaved specimens with 
no red color in the petioles from as far east as South Berwick, 
Maine, are preserved in the Gray Herbarium, but that these are 
not so extreme in these respects as is the mid-western material. 
Also in the Gray Herbarium, according to Mr. Weatherby, are 
specimens collected in 1877 by Dr. J. H. Burdick at Milton, 
Wisconsin, which he sent to Dr. Gray with the following descrip- 
tions: 


1946] 


Purple var.—No. 1 


Leaves narrowly elliptic or 
elliptic-lanceolate tapering into 
a purple petiole (above ground) 
about one-third the length of the 
leaf, which is bright green and 
shining. 

Scape deep purple. 


Hermann,— Muhlenbergia setosa 


Green var.—No. 2 


Leaves lanceolate, pale green 
and glaucous.  Petiole hardly 
any (above ground), white. 
There is no purple about the 
plant. 


Scape pale green. 


63 


Dr. Gray and Dr. Watson were not disposed to recognize 
Burdick's varieties as anything more than response to environ- 
mental factors. Burdick wrote twice later, maintaining his 
opinion, and in one letter suggested a varietal name for the race 
with colored scapes. He does not seem to have published on the 
subject. His letters are preserved at the Gray Herbarium. 

Bulbs of both races were sent to the second author in 1940, and 
the differences pointed out remain constant in the experimental 
garden at Pullman, Washington. In spite of its late start, Race 
B flowers several weeks in advance of Race A in the garden under 
Washington conditions. In Michigan, such a lag has not been 
noted. The differences shown by the two races are such as are 
frequently shown by diploid and tetraploid races of the same 
species, but the chromosome number of both Race A and Race B 
as determined by Dr. Hannah C. Aase (unpublished) is diploid, 
n = 8. 

Schoolcraft, Michigan; State College of Washington. 


MUHLENBERGIA SETOSA AN UNTENABLE NAME.—In his realign- 
ment of the common rhizomatous species of Muhlenbergia of 
eastern North America (RHoporRA 45: 221-239. 1943) Professor 
Fernald has demonstrated that the eastern plant which had for 
so long been passing under the name Muhlenbergia racemosa 1s 
amply distinct from true M. racemosa (Michx.) BSP. of the 
interior. For the eastern segregate he took up the name Muhlen- 
bergia setosa (Spreng.) Trin. ex Hook. f. & Jackson, Ind. Kew 2: 
269 (1897), later (RHoponaA 47: 198. 1945) corrected to M. 
setosa (Biehler) Trin. ex Hook. f. & Jackson. There is, however, 
an earlier Muhlenbergia setosa which was overlooked by the com- 
pilers of the Index Kewensis, namely M. setosa (HBK.) Kunth, 
Rev. Gram. 1: 63 (1829), based upon Podosaemum setosum HBK. 
Although the latter name is currently treated as à synonym of 


* 


64 Rhodora [Marcu 


M. microsperma (DC.) Kunth, it meets all the requirements for 
valid publication. Muhlenbergia setosa (Biehler) Trin. ex Hook. 
f. & Jackson is therefore a later homonym and must be replaced 
by Muhlenbergia glomerata (Willd.) Trin., a name discussed in 
some detail by Professor Fernald (Ruopora 45: 232-235. 1943). 
This necessitates the transfer of the more northern M. sctosa 
var. cinnoides (Link) Fernald as follows: 

M. GLOMERATA (Willd.) Trin. var. cinnoides (Link), comb. 
nov. Dactylogramma cinnoides Link, Enum. Hort. Berol. 2: 248. 


1833 (RuoponaA 45: 238. 1943).—F. J. HERMANN, Bureau of 
Plant Industry Station, Beltsville, Md. 


Dors HABENARIA CRISTATA STILL GROW IN New ENGLAND?— 
The only New England specimens known to me of //abenaria 
cristata (Michx.) R. Br. are three sheets in the Herbarium of the 
New England Botanical Club, collected by the late E. Williams 
Hervey on Smith's Neck, Dartmouth, Bristol County, Massa- 
chusetts, in early August of 1905 and of 1908. "This is, appar- 
ently, the only station northeast of New Jersey. Is it still 
there?—M. L. FERNALD. 


Volume 48, no. 566, containing pages 17-40 and plates 995-1004, was issued 
6 February, 1946. 


Hovora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL . Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. April, 1946. No. 568. 


CONTENTS: 


Contributions from the Gray Herbarium of Harvard University— 
No. CLX. Technical Studies on North American Plants. 


M.L.Fernald (concluded): -n m r o 2.0022. a 65 
Previously unreported Plants from Minnesota. Olga Lakela. .... 81 
Bidens hyperborea var. typica. Norman C. Fassett. ........... 82 
Genus Palafoxia in Texas. V. L. Cory. .......... eese 84 
North American Representatives of Alisma Plantago-aquatica. 

ML RENAA a a a VG A 86 


Sporadic Appearance of Epipactis Helleborine. M. L. Fernald. 88 


The New England Botanical Club, Ine. 


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


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Address manuscripts and proofs to 
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Subscriptions (making all remittances payable to RHODORA) to 


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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 


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EIGHT WEST KING ST., LANCASTER, PA. 


EDIBLE WILD PLANTS OF EASTERN NORTH AMERICA 
by MERRITT LYNDON FERNALD and ALFRED CHARLES KINSEY 


Practical discussion of edibility and directions for recognition and prepara- 
tion of more than 1000 wild plants. 422 pp., introd. and detailed index, 124 
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MEMOIRS OF THE GRAY HERBARIUM. A series of illustrated quarto 


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No. I. A Monograph of the Genus Brickellia, by B. L. Robinson. 150 
pp., 96 fig. 1917. $3.00. 


No. Ill. The Linear-leaved North American Species of Potamogeton, 
Section Axillares, by M. L. Fernald. 183 pp., 40 plates, 31 maps. 1932. 
$3.00. 


No. IV. The Myrtaceous Genus Syzygium Gaertner in Borneo, by E. D. 
Merrill and L. M. Perry. 68 pp. 1939. $1.50. 


No. V. The Old World Species of the Celastraceous Genus Microtropis 
Wallich, by E. D. Merrill and F. L. Freeman. 40 pp. 1940. $1.00. 


Grav Herbarium of Harvard University, Cambridge 38, Mass. 


Rhodora Plate 1011 


Photo B, G, Schubert 


x SOLIDAGO HIRTIPES, all figs. from TYPE: FIG. 1, portion of plant, X 5; FIG. 2, pedi- 
celled heads, X 5; ric. 3, pedicels, X 10; ric. 4, upper, and FIG. 5, lower surface of leaf, 
x 10. 


Rbodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. April, 1946. No. 568. 


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


TECHNICAL STUDIES ON NORTH AMERICAN PLANTS 


M. L. FERNALD 
(Continued from page 60) 


» 

X SounipAGo (8 EvrHAMIA) hirtipes, hybr. nov. (? S. gramini- 
folia (L.) Salisb., var. Nuttallii (Greene) Fernald X S. microce- 
phala (Greene) Bush). Tas. 1011. Planta robusta ad 1.5 m. 
alta; foliis lineari-lanceolatis divergentibus 3—5-nerviis utrinque 
scabro-puberulis, in axillis plus minusve fasciculatis; foliis pri- 
mariis 4-6 mm. latis; corymbo ad 4.5 dm. lato ramis valde 
adscendentibus, ramulis ultimis densissime griseo-hirtellis; capi- 
tulis glomerulatis vel solitariis pedicellatisque; involucro glutinoso 
pallido cylindrico (sicco cylindrico-turbinato) 3-4 mm. longo; 
phyllariis pallide stramineis apice adpresso viride.—Sussex Co., 
VIRGINIA: roadside thicket about 1144 miles north of Waverly, 
Sept. 13, 1945, Fernald & Long, no. 15,015 (TYPE in Herb. Gray; 
ISOTYPE in Herb. Phil. Acad.). 


X Solidago hirtipes is a very puzzling plant, which may prove 
to be a fully established species. In its very broad and flat- 
topped corymb (with inclination to form "stories" at different 
heights) and in its very slender heads it at once suggests the 
common southern S. microcephala (PLATE 1012, Fics. 4-6); but . 
that characteristic species has the very narrow and often longi- 
tudinally folded primary leaves only 1-2 mm. broad, 1-nerved 
(only rarely with any trace of lateral nerves), and subtending 
abundant suppressed axillary branches. Its sparsely hirtellous 
pedicels are mostly 1-headed; while its very narrow leaves are as 
gray-puberulent as in X S. hirtipes. In its great stature, broad 


66 Rhodora ” TAPRID 


3-5-nerved leaves and strong tendency to glomerulate heads the 
new plant is as.near S. graminifolia, var. Nulttallit. In that 
plant (PLATE 1012, rias. 1-3), however, the suppressed axillary 
branches are few or wanting, the leaves less puberulent, the 
pubescence of the lower side usually confined to the ribs, the 
corymb with convex-topped secondary corymbs, the branchlets 
and pedicels as hirtellous as in X S. hirtipes (or even more so) 
but ending in densely crowded glomerules of broader and more 
subcampanulate heads. 

I am looking upon X S. hirtipes as probably daid from 5. 
microcephala and S. graminifolia, var. Nuttallii, though further 
. experience may show it to be wholly separable from them. The 
type-colony is likely to spread; as it is, we took only selected 
small plants of it. 


XANTHIUM Chasei, sp. nov. (TAB. 1013, rra. 1, et 1014). Plan- 
ta a X. strumario differt petiolis scabro-hispidis foliorum laminis 
crassis supra scabris; fructi corpore glabro vel minute puncticu- 
lato lucido olivaceo ellipsoideo-ovoideo vel subgloboso 1.8-1.6 
cm. longo 6-9 mm. crasso, rostris basin versus crassis glanduloso- 
puberulisque porrectis vel suberectis remotis rectis 4-5 mm. 
longis vel ad apicem exigue excurvatis vel incurvatis, exteriore 
facie 100-200 aculeis approximatis armato, aculeis anguste sub- 
ulatis rectis vel superne curvatis glabriusculis vel basin: versus 
glandulosis 2-3.5 mm. longis.—ILLINOIS: bottomlands of Illinois 
River near Peoria, Oct. 1, 1919, Virginius H. Chase, no. 3398, as 
X. globosum Shull (Herb. Chase., Herb. Gray.); Sept. 12, 1920, 
Chase, no. 3474, as X. globosum (Herb. Chase., Herb. Gray.); 
Sept. 15, 1945, Chase, no. 8205 (TYPE in Herb. Gray.; ISOTYPE in 
Herb. Chase.). 


Although originally identified as Xanthium globosum, a natural 
identification since there was then no good material readily 
available of Shull's species, X. Chasei, named for its discoverer, 
VinaiNIUS HEBER CnasEk, differs at once from that species (our 
PL. 1015, rras. 1 and 2) in several characters which appear in all 
the specimens before me. -In X. globosum the leaves are thinnish 
to membranaceous and the ripe bur is light brown, with the body 
only 0.9-1.1 em. long and 4.5-7 mm. thick, while the 50-80 
prickles visible on one face are remote and 4-6 mm. long. This 
specles is widespread from Illinois and Kentucky to Kansas and 
is represented in Mr. Chase's series by fine specimens from the 
bottomlands near East Peoria, his nos. 8206 and 8207. X. 


1946] Fernald,—Studies on North American Plants 67 


Chase differs in its thick and more scabrous foliage and especially 
in its burs. These are olivaceous or greenish brown, the body 
1.3-1.6 em. long and 6-9 mm. thick, with 100-200 prickles visible 
on one face, these rather crowded and only 2-3.5 mm. long. 
So far as the abundant material shows, the fruit of X. Chasei 
scarcely overlaps in any point the distinctive characters of X. 
globosum. 

In its very short prickles Xanthium Chasei might suggest the 
European X. strumarium L. (our Pr. 1013, FIGs. 2 and 3), a 
species with us only casually adventive! but apparently not 
naturalized near the port of Boston; but the two differ in many 
points and the natural habitat of X. Chasez is not ballast and 
rubbish about eastern seaports. In X. strwmarium the petioles 
are minutely soft-pilose and the leaf-blades submembranaceous 
and barely scabridulous above; whereas the petioles of X. Chasei 
(on bottomlands where many plants tend toward glabrescence 
and thin leaves) are harshly scabrous, as is the upper surface of the 
thick leaf-blade. In X. strumarium the mature burs are closely 
and finely pilose, with the body only 5-7 mm. thick, the beaks 
only 1-2 mm. long, while the 15-50 slender prickles visible on 
each face have broad interspaces separating their bases. In X. 
Chasei, on the other hand, the mature burs are 6-9 mm. thick, 
glabrous (except for glandular punctation), the beaks 4-5 mm. 
long, the 100-200- bulbous-based prickles visible on one face 
crowded. 

There is no chance that Xanthium Chasei has anything to do 
with the large-fruited series with strongly villous burs: X. italicum 
Moretti with fulvous burs villous-hirsute, the body 1.3-1.8 cm. 
long and 6-8 mm. thick, the subulate basally long-hirsute beaks 
5-7 mm. long, the divergently long-villous prickles mostly 4—7 
mm. long (see PL. 1015, Frias. 3 and 4); X. oviforme Wallr. with 


LIn the Gray Herbarium there are 3 sheets of Xanthium strumarium from the 
United States: (1) a fragment of doubtful origin in a series of plants said to have been 
collected by the late Stephen P. Sharples in Chester County, Pennsylvania, from 
1858-1864; (2) branches from a single tall plant, found by the writer on October 20, 
1912 (with fresh anthers), in rubbish back of Crescent Beach, Revere, Massachusetts; 
(3) portion of a single individual found on October 1, 1927, by S. F. Blake (Blake, no. 
10,549) on a sandy beach at Hull, Massachusetts. In early November, 1945, Dr. 
Bernice G. Schubert and I followed the beach from south of Crescent Beach to well 
north of that area but no Xanthium strumarium could be found. Similarly, the sandy 
and gravelly beaches in Hull, like those in Revere, yielded only X. echinatum, italicum 
and chinense. ; 


68 Rhodora - [APRIL 


the body of the fulvous bur 2-2.5 cm. long and 1.2-2 cm. thick, 
the stoutish beaks 7-10 mm. long, and the densely crowded 
prickles 7-10 mm. long; and other species. Nor can it be forced 
into the very definite X. inflecum Mackenzie & Bush (our PL. 
1015, rias. 5 and 6), which occurs on bottomlands and prairie of 
Illinois, Missouri, Kansas and Arkansas, for X. inflecum has the 
glabrous to hirtellous body of the bur 1.3-2 cm. long and 6-8 mm. 
thick, the strongly inflexed to overlapping or crossing beaks 5-7 
mm. long, and the prickles 4.5-10 mm. long. 

The glabrous or glabrescent burs of Xanthium Chasei might, to 
some minds, mean that it is X. chinense Mill. or X. curvescens 
Millspaugh & Sherff. The widely dispersed X. chinense (our PL. 
1015, rias. 7 and 8), however, has the fewer reddish-tinged 
prickles very remote at base and 4-7 mm. long, the more inflexed 
or hamate beaks up to 7 mm. long. As to X. curvescens (our PL. 
1016, rias. 1-4), described from the shores of Lake Champlain, 
that doubtful species, illustrated with the original description in 
Field, Mus. Pub., Bot. Ser. iv., no. 2, pl. xi, the burs shown in 
pl. viii, figs. 28-29 (1919), has the body of the fruit only 3.5-5 
mm. thick, with only 30-50 stout and upwardly curving prickles 
visible on each face. It thus has characters strongly suggestive 
of the Old World X. orientale L. (Pr. 1017). In fact, very typical 
X. orientale (our PL. 1017, rias. 5-8) is an abundant weed about 
Montreal, thence up the Richelieu Valley to the shores of Lake 
Champlain (perhaps better stated in reverse order), and the most 
. slender Old World fruits (our rra. 3) closely match those of X. 
curvescens (PL. 1016, rias. 1—4) and even of X. leptocarpum Mills- 
paugh & Sherff. (our Pr. 1016, rias. 5 and 6). In fact, when they 
published X. curvescens and X. leptocarpum the authors of those 
names saw their resemblance to X. orientale, saying 

Because of its strongly bent prickles and beaks, we were disposed at 
first to regard this spevies as a form of the European X. orientale L. But 
in the many fruiting specimens of X. orientale examined from Europe, we 
have found the fruiting involucres to be not only considerably larger, but 
brownish rather than reddish, also much more pubescent and the prickles 
nearly always more numerous. In its narrow, reddish, remotely aculeate 
fruits, this species suggests the next, X. leptocarpum, the type of which 
was collected likewise in western Vermont, about three years earlier. 
Indeed, as a species, it seems to lie just half-way between X. orientale 
and X. leptocarpum, and for a time we suspected it of being a hybrid 


between these two species. But the apparent absence of true X. orientale 
from all of North America would seem to make an assumption to this 


1946] Fernald,—Studies on North American Plants - 69 


effect purely gratuitous. Nor do we feel inclined to regard our plant as 
an anomalous race or variety of X. leptocarpum, since the arcuate char- 
acter of its prickles is a character that holds with a high degree of uni- 
formity throughout the specimens of the corresponding X. orientale of 
Europe. 


Had they realized the establishment of X. orientale from the 
shores of Lake Champlain to the streets, waste lots and wharves 
of Montreal, their first impression might have prevailed. At any 
rate, X. Chasei has nothing to do with X. orientale, and the 
doubtfully separable X. curvescens and X. leptocarpum. 

If only its short beaks and prickles were taken into account 
and all other characters ignored or overlooked (a tendency too 
. apparent in some recent so-called studies of Xanthium), Xanthium 
- Chasei might superficially be placed with X. echinatum Murr. and 
X. varians Greene, for in all three of these species the beaks are 
unusually short and stout and the prickles short. There, how- 
ever, the resemblance ends. Both X. echinatum (our Pr. 1018) 
and X. varians (PL. 1019) have the olive-shaped drab to pale 
brown mature bur densely long-hirsute, the stout beaks 3-6 
mm. long and with very stout hispid bases 2-3 mm. thick, the 
remote prickles mostly hamate, hirsute-villous below the middle 
and 3.5-5 mm. long. By recent authors the inland X. varians 
has not been distinguished from the coastal X. echinatum; but 
the two are well distinguished both by characters of leaf and bur 
and by their strikingly different habitats, X. echinatum confined 
to sea-beaches, dune-hollows and borders of saline marshes along 
the coast from southern Maine to Virginia; X. varians occurring 
inland, on shores and damp prairie, from western Quebec to 
northern Alberta, south to northern New York (St. Lawrence 
drainage), Iowa, South Dakota, Saskatchewan and Oregon. In 
the halophytie X. echinatum the young stem is whitened above 
with harsh and short hispidity, the later full-grown leaves are 
broadly cordate or subcordate-ovate, unlobed or very shallowly 
lobed, with undulations mostly longer than deep (Figs. 1 and 2), 
and the beaks of the fruit (Frias. 3-6) tend soon to inflex and 
finally to become approximate or with their tips crossing, like 
the bill of a crossbill. The inland X. varians (Pr. 1019), on the 
other hand, has the greener stem with sparse pubescence, the 
larger leaves more often rhombic or cuneate-based, their margins 
subacutely dentate with teeth nearly or quite as high as broad, 


70 Rhodora [APRIL 


and the erect or nearly erect beaks straight or nearly so (Fries. 2 
and 3). Only by the most superficial treatment could X. 
Chasei be crowded into either of these apparently different 
species or geographic varieties for its bur is essentially glabrous 
and lustrous, olivaceous, and with crowded nearly glabrous 
prickles only 2-3.5 mm. long.! 

I realize that in his somewhat novel (and we hope, immature) 
concept Cronquist, following the cue of Wiegand, who saw more 
hybrids than pure strains in several variable groups, has inti- 
mated in RHopona, xlvii. 403 (1945) that most if not all our 
species of Xanthium are variations of one species, the European 
X. strumarium, for he finds that “The determination of species 
of Xanthium has become a formidable task, undertaken by many 
botanists only when it becomes unavoidable, and then with seri- 
ous misgivings”, exactly the situation with Carex, Potamogeton, 
Festuca and many other groups with inconspicuous flowers, 
which are regularly dodged by those who want to work only on 
pretty flowers. Nevertheless, there are scores of very real species 
in these genera and they have been clearly recognized by some of 
the most thorough and wise students of plants, for we bow with 
profound respect to such sound students of Carex, for example, 
as Willdenow, Schkuhr, Schweinitz, Torrey, Dewey, Francis 
Boott, William Boott, Tuckerman and many others, although 
the casual botanizer lets the genus alone. Similarly, that the 
study of Xanthium bristles with difficulties none will gainsay; 
but, paradoxically, the harder the points in this genus the easier 

1It seems hardly necessary, in orienting Xanthium Chasei, to illustrate three species 
found in the eastern United States, with which it could not be confused. The bur of 
X. pensylvanicum Wallr. is very similar to that of X. chinense but, whereas the body 
of the latter is glabrous or merely glandular-punctate and lustrous, that of X. pensyl- 
vanicum is short-pilose with pale pubescence, and the 200 or more prickles visible on 
one face are mostly glandular-hispid below the middle, the bases of the prickles about 
as broad as the interspaces. In X. chinense the interspaces are broader than the 
nearly or quite smooth bases of the fewer prickles (100—150 visible on one face). 
X. pensylvanicum thus stands between X. chinense and X. italicum. X. speciosum 
Kearney has a bur (including beaks and prickles) 3—4 cm. long and 2-3 cm. thick, the 
almost filiform-setose prickles 7-9 mm. long and so crowded as to have their bases 
practically hidden, the beaks 6-11 mm, long. X. Wootoni Cockerell could be confused 
only with X. orientale, but it differs in its fewer heavily glandular and straightish 
prickles and in having the porrect (instead of strongly incurving) beaks more distant 


from the upper prickles. The burs of all four of the species are beautifully illustrated 
by Millspaugh & Sherff or by Widder. 


1946] Fernald,—Studies on North American Plants 71 


the identifications, for, whereas young and relatively tender 
flowering plants and even those with well-grown but immature 
fruit are most difficult to distinguish (being annuals of very simi- 
lar habit and foliage), the fully mature and very hard burs alone 
are sufficient for identification. The somewhat discouraged 
verdict of Wiegand, which Cronquist accepts in lieu of working 
out the characters, is not final: “It is probably wise to treat all 
North American Xanthiums as one species except X. spinosum 
L.and possibly X. strumarium L.and X.echinatum Murr.  X.stru- 
marium, however, is scarcely distinct, and with more study may 
also be included. X. echinatum may be a real species, as it has a 
distinct coastal range and seems to behave as though genetically 
distinct". Cronquist is “in complete agreement with Wiegand's 
observations, except that X. strumarium sens. strict., seems no 
more than varietally distinct from our plants, and that I am 
quite unable to see any sort of taxonomic unit in X. echinatum.” 

Inability to see what others clearly see is not a sin; neither 
does it necessarily clarify a question. To those who have some 
understanding of the characters in the genus, Wiegand's recogni- 
tion of X. echinatum will appeal. Several close students have 
devoted themselves to the difficult and often thankless task of 
working out the characters of Xanthium: (1) Wallroth, Mono- 
graphischer Versuch über die Gewdchs-Gattung Xanthium, Wallr. 
Beitr. i. 219-244 (1844), Wallroth recognizing 21 species, 7 of 
them North American; (2) Millspaugh & Sherff, Revision of the 
North American Species of Xanthium, Field Mus. Pub., Bot. Ser. 
iv. no. 2 (1918), with very fine illustrations of the burs, this very 
detailed study also recognizing 21 species but all of them found 
in North America, the authors having gone to great pains to 
identify the types of earlier-described species; (3) Widder, Die 
Arten der Gattung Xanthium in Fedde, Repert. Beih. xx. (1923), 
a very conservative and careful study with clear illustrations of 
the fruits, the original text completed before the publication of 
the monograph of Millspaugh & Sherff, but that study summar- 
ized in the Nachtrag. Widder maintained most of the species 


1That Wallroth was not a visionary, as one might infer from those who can see 
nothing in his species of Xanthium, is apparent from his great monograph of Agri- 
monia, published also in his Beitráge, for his four new North American species, A. 
gryposepala, rostellata, microcarpa and pubescens, are all maintained by those who 
have closely studied the genus. 


72 Rhodora Jäme 


recognized by Wallroth and by Millspaugh & Sherff. If, as we 
have recently been told, all of them, except X. spinosum, are 
modifications of a single species, which in its typical form has . 
scarcely gained a foothold (and that near a large Atlantic port) 
in America, it is amazing that not one of the really careful stu- 
dents of the genus should have suspected the species to be of 
such instability. 

Besides the three great monographs of Xanthium, there have . 
been many significant but briefer studies, these noted by Mill- 
spaugh & Sherff and by Widder. From our viewpoint as sig- 
nificant as any is that of the physiologist, Professor Charles A. 
Shull, Physiological Isolation of Types in the Genus Xanthium, in 
Bot. Gaz. lix.! 474—483, with 7 blocks of figs. (1915). Desiring 
material for physiological study, Shull went to an old field near 
Lawrence, Kansas, where plants of Xanthium with 3 quite dif- 
ferent typés of bur abounded. “It was thought that the various 
forms were possibly the result of promiscuous crossing of varieties 
or elementary species, and that a year or two of guarded pollina- 
tion would be necessary to purify the strains." Shull took 
essentially uniform burs from a single plant each of the three 
extremes, what he eventually described as a true species, his X. 
globosum, the wide-ranging X. pensylvanicum and X. canadense 
(now known by the earlier name X. chinense). Successive 
generations from these fruits in carefully checked plots proceeded | 
to come true and not to show any Mendelian segregation. 
“This result was wholly unexpected, as it was believed that 
hybridization could hardly have been avoided in nature". 
Shull recognized not only the differences in burs but those of 
cotyledons even, as well as of pigmentation of the stems and 
prickles and shape of the seeds and color of their coats; and, 
having started out a skeptie, with the popular preconception 
that the differences of burs and other characters he saw were the 
result of *promiscuous crossing", he frankly concluded his experi- 
mental study by defining à new species, and on his last page he 
unblushingly predicted that *"There are probably a number of 
new species of Xanthium still undescribed in America". X. 
Chasei is one of them. 


i'Through the perverse fatality which dogs editorial movements the signature of 
the Gazette on which Shull's paper begins bears the note ‘‘Botanical Gazette, vol. 69]"' 


Rhodora Plate 1012 


Photo B. G. Schubert 


SOLIDAGO GRAMINIFOLIA, Var. NUTTALLII: FIG, 1, glomerules, X 5; Fra. 2, branchlet 
and pedicel, X 10; ria. 3, lower surface of leaf, X 10. 
S. MICROCEPHALA from TYPE-REGION: FIG. 4, pedicelled heads, X 5; FIG. 5, branchlet 


and pedicels, X 10; FIG. 6, upper surface of leaf, X LO. 


Rhodora Plate 1013 


Photo B. G. Schubert 


NANTHIUM CHASEI! FIG. 1, TYPE, X ?/;. 
X. STRUMARIUM: FIG. 2, bur, X 2; FIG. 3, bur, X 5. 


1946] Fernald,—Studies on North American Plants 73 


On the Illinois bottomlands near Peoria Xanthium Chasei has 
remained constant, just as X. echinatum does along the coastal 
sands or as does the distinctive X. speciosum Kearney, brought 
unintentionally in western fleeces to New England woolen-mills, 
whence it has washed down-river and made constant and con- 
sistent colonies far from their native haunts. 

Cronquist’s last paragraph on Xanthium reads: 

Although both species [all-inclusive X. strumarium, involving all 
American members of DeCandolle’s § Euxanthium, and X. spinosum, the 
single species recognized by him in $ Acanthozanthium DC.] of Xanthium 
have now become cosmopolitan weeds, and X. strumarium was well 
established in Europe four hundred years ago . . . , it seems probable 


that they originated in the new world. Except for a few species of 
Ambrosia, the subtribe Ambrosinae is otherwise exclusively American. 


That statement implies that the several endemics in Xanthzum 
§ Euranthium which are known only in the Old World, such 
unique plants as X. inaequilaterum of DeCandolle (who was no 
*splitter"), found from Java and Borneo to Japan and the coast 
of southeastern China, or as X. indicum Konig (X. Roxburghii 
Wallr.) of tropical and subtropical India, Sumatra and Java,— 
the implication is that such endemic species of the Old World 
originated in America, but entirely quit the New World in favor 
of the Old. This interpretation is again illustrated by X. 
spinosum, often assumed to be indigenous (as perhaps it is) in 
South America, but certainly not in eastern North America, 
where it is a weed of relatively recent introduction. Linnaeus 
described it “Habitat in Lusitania" and Widder, whose citations 
of South American and North American specimens occupy less 
than 1 page, gives up 6 pages to citations of European and 
African specimens. Whether this semicosmopolitan weed is 
native in South America I cannot say, but the learned Charles 
Pickering wrote in his Chronological History of Plants, 976 . e 
; (1879): “Transported to North America as late perhaps as 1814 SEES z 

. (as may be. inferred from the silence of Walter, Michaux, and. VM e 


? 


2 ds Pursh) , Was found by Nuttall in 1818 near dwellings. from. BENE V ros 


ee ~ nah to Washington, and not foréseeing that it would become | a YA ELO 
SEC NER troublesome, was introduced by him as he informed me into the - dem d 3 
environs of Philadelphia; . . . In the Southern Hemisphere, by - 
European colonists also, was probably carried across the Andes 

into Chili (Beechey voy. 57, and A. Dec.)", Hooker & Arnott, 


74 Rhodora [APRIL 


in their Botany of Captain Beechy’s Voyage, recording it from 
Valparaiso, surely not an undisturbed locality. Finally, in re- 
gard to the nativity of X. spinosum, the article; Xanthium spino- 
sum in Neolithic deposits in Bulgaria, by W. B. Turrill in Kew 
Bull. for 1923: 190 (1923) is significant: 


Xanthium spinosum is one of the commonest ruderal plants of Central 
Europe and the Mediterranean region. It has generally been considered 
by recent writers to be a native of South America. The strong arguments 
in favour of this view will be found summarized by Thellung in his Flore 
adventice de Montpellier . . . . Another opinion, which was current 
among earlier botanists, that Xanthium spinosum was native in South 
Russia, was rejected. by Ascherson . . . . It may also be noted that 
according to L. Simonkai and Karl Flatt . . . both A. Florentin and also 
C. Spegazzini have collected fossil fruits (‘false-fruits’) of X. spinosum 
from the Pliocene beds in the Tertiary formation of the Pampas. 

"During a visit to Bulgaria last summer Dr. N. Stoianoff of Sofia 
University gave me some semi-fossilized fruits which careful comparison 
at Kew has shown to be those of Xanthium spinosum. These were ob- 
tained during excavations in prehistoric deposits . . . east of Sofia, . . . 
The deposits are of Neolithic age . . . 

"Whatever may be the real history of Xanthium spinosum in the Old 
World this discovery would seem to indicate that the species existed in 
South Europe long before the dates (1700-1750) accepted by Thellung for 
its first introduction and establishment. 


Surely Neolithie Bulgarians did not come to America to get it. 
Xanthium, like Ambrosia, obviously has species native to the 
Old World as well as to the New. Wallroth thought that he saw 
characters to separate American material from typical European 
X. spinosum. And, although Cronquist would reduce all Xan- 
thium to the' two species (i. e. the two sections of DeCandolle 
and others), the European X. strumarium L. and X. spinosum 
L., it is most difficult to believe that the very distinct and charac- 
teristic X. ambrosioides Hooker & Arnott, the Argentinian pro- 
cumbent plant with bipinnate blunt-lobed leaves, a species which 
has to be taken into account by those who watch the waste lots 
of New England, can possibly be forced into X. spinosum. 
Hooker & Arnott were ultraconservative, yet they saw what is a 
really good species in X. ambrosioides; so did Widder; so do others. 

HELIANTHUS ATRORUBENS L., var. alsodes, var. nov. (TAB. 
1020), foliis imis longe petiolatis anguste ovatis vel late lanceolatis 
acutis vel subacutis plerumque serratis vel serrato-dentatis; 
foliis supernis anguste ovatis vel lanceolatis.—Dry open woods, 


thickets and clearings, Virginia, North Carolina and upland of 
southeastern Kentucky and eastern Tennessee. VriRGINIA: dry 


1946] Fernald,—Studies on North American Plants 75 


land about 3 miles north of Williamsburg, James City Co., R. W. 
Menzel, no. 289; dry open thickets, Virginia Beach, Princess 
Anne Co., Sept. 10, 1935, Fernald, Long & Fogg, no. 5127 (TYPE 
in Herb. Gray.; isoryPr in Herb. Phil. Acad.); rich woods and 
bushy clearing north of Double Bridge, southwest of Jarratt, 
Sussex Co., Fernald & Long, no. 11,474; Bedford Co., Oct. 15, 
1871, '"leafy-stemmed form", A. H. Curtiss; *Mts., Carol. & 
Virginia", Asa Gray. NonrH CAROLINA: sand-ridge at Atlantic, 
Cartaret Co., Godfrey, no. 6447; pineland near Goldsboro, Wayne 
Co., Godfrey, no. 6576; sand-ridge near Roseboro, Sampson Co., 
Godfrey, no. 5699; sand-ridge at Carolina Beach, New Hanover 
Co., Godfrey, no. 6368; pine woodland, Raleigh, Godfrey, no. 6601; 
thicket, Raleigh, Godfrey, no. 6892; dry thicket, north of Spruce 
Pine, Mitchell Co., Wiegand & Manning, no. 3378; dry open 
woods, Biltmore, Buncombe Co., Bilt. Herb., no. 499*; rich 
wooded mountain side near Asheville, A. H. Curtiss, no. 6526; 
open woods, Henderson Co., Aug. 24, 1881, J. D. Smith (as 
Rudbeckia fulgida); open woodlands, Highlands, Bilt. Herb., 
no. 499°. KENTUCKY: roadsides and fields east of Cumberland 
Falls, Whitley Co., McFarland, no. 59. TENNESSEE: gravelly 
oak-pine woods, Jamestown, Fentress Co., Svenson, no. 4100. 


Helianthus atrorubens consists of the two rather marked varie- 
ties: var. normalis Kuntze, Rev. Gen. i. 343 (1891), and the 
narrower-leaved plant here described as var. alsodes. The 
species started with the detailed description and life-size plate of 
Corona solis minor, disco atro-rubente of Dillenius, Hort. Elth. 111, 

`t. xciv (1732) and the Helianthus foliis ovatis crenatis of Grono- 
vius, both cited without further description by Linnaeus, who 
obviously took his epithet from Dillenius (whose plant was from 
Carolina) but gave the “Habitat in Virginia". There was no 
specimen of the species in the Linnean Herbarium at the time of 
writing Species Plantarum; and the Gronovian description of the 
Clayton specimen, with "folia asperata paucis auritis, ex adverso 
binis auriculatis", suggests the Dillenian plate, in which there are 
tiny leaves (“auricles”) at the bases of some large primary leaves. 
. Such tiny undeveloped axillary branchlets occasionally occur in - 
the plant with ovate or oval bluntish leaves (var. normalis) but - 
I have not seen them in var. alsodes. The Dillenian plate — 
(which may stand as TYPE of the species) is of the characteristic — 
plant found from Florida to Louisiana, north to Virginia and 
Tennessee, with the basal leaves ovate or oval and crenate to 
dentate, these and the lower cauline with blades one half to four 


76 — S Rhodora [APRIL 


fifths as broad as long and blunt or merely subacute, their bases 
abruptly contracted to the winged petiole, the sessile or subsessile 
median cauline leaves ovate or oval. In this plant the middle 
phyllaries are broadly rounded at summit or abruptly short- 
acuminate.' Var. alsodes, although found in the northeastern 
third of the specific range, has the lower long-petioled leaves 
usually more gradually tapering from petiole to blade, and the 
most often serrate or serrate-dentate blade is acute or acutish, 
two to five times as long as broad; the lowest sessile cauline blades 
lanceolate or narrowly ovate. In var. alsodes the middle phyl- 
laries (FiG. 3) are usually less broadly rounded at summit. 

It is barely possible that Helianthus atrorubens, var. alsodes is 
the plant described as H. sparsifolius Ell. Sk. ii. 415 (1823) from 
the “western districts of Georgia", as contrasted with his H. 
atrorubens apparently common in his own region of southeastern 
South Carolina. Elliott’s “leaves ovate, acute, coarsely toothed” 
is good, but “abruptly contracted into a petiole” not so good; 
furthermore, since he defined H. atrorubens as having “leaves 
spathulate’’, it would be unwise, his specimens of H. sparsifolius 
not being preserved at Charleston, to take up the name for var. 
alsodes. From the citation by E. E. Watson in Papers Mich. 
Acad. ix. 343 (1920), in the synonymy of H. atrorubens, of H. 
gracilis Bertoloni one might infer that Bertoloni had our plant. 
It is noteworthy, however, that Watson omitted the citation of 
Bertoloni's plate, which he evidently had not seen. H. gracilis 
Bertoloni, Misc. Bot. vii. 41, t. vi, fig. 1 (1868), shown life-size, 
is a monocephalous plant with lower entire leaves barely petioled, 
extending gradually as linear-lanceolate blades to the single 
peduncular summit of the stem, the phyllaries linear or lanceolate 


1'The following are characteristic specimens of var. NORMALIS. VIRGINIA: west of 
Mt. Hope Church, Southampton Co., Fernald & Long, no. 11,472; northwest of 
Round Gut, southwest of Franklin, Southampton Co., F. & L., no. 11,473. NORTH 
CAROLINA: west of Jacksonville, Onslow Co., Godfrey, no. 6461; Winston-Salem, 
Forsyth Co., Godfrey, no. 6103; Biltmore, Buncombe Co., Bilt. Herb., no. 499*; South 
of Tuxedo, Henderson Co., Wiegand & Manning, no. 3377; Highlands, Aug. 2, 1902, 
Magee; Joy, Burke Co., Hunnewell, no. 12.981. Sours CanmorrNa: Eutawville, 
Eggleston, no. 4997; Marietta, Greenville Co., Wiegand «& Manning, no. 3370. 
Groraia: Middle Ga., 1846, Porter. TENNESSEE: Chilhowee Mt., A. H. Curtiss, no. 
1439. Lovi8IANA: without stated locality, Drummond. 

It is not improbable that var. normalis tends to grow in damper soil than var. 
alsodes. Of the 16 sheets before me of var. alsodes, of which habitat is given, all are 
from dry situations. Of the 13 of var. normalis with habitat noted 2 are from sa- 
vannas, 1 from pine-barren bog. 


Rhodor: Plate 1014 


Photo D. G. Schubert 


NANTHIUM CHASEL, FIGS. 1, 2 and 3 from rype: FIG, 1, portion of plant, X 1; Fic. 2, bur, 


X 2; FIG. 3, portion of bur to show beaks and bulbous-based prickles, X 5; FIG. 4, bur, 
X 2, and Fic. 5, its summit, X 5. 


Rhodora Plate 1015 


Photo B. G. Schubert 


NXANTHIUM, BURS, X 2, BEARS, X 5; FlGs, | and 2, X. GLoposum, from plant raised from 
seed of TYPE: FIGs. 3 and 4, N. ITALICUM; rics. 5 and 6, N. INFLEXUM; FIGs, 7 and 8, X. 
CHINENSE. 


Rhodora Plate 1016 


Photo DB. G. Schubert 


NTHIUM, entire burs X 2, enlargements X 5: FIGS. 1—4, X. CURVESCENS, FIGS. 1 


A 
2 from TYPE; FIGs. 5 and 6 from TYPE of X. LEPTOCARPUM. 


X 
d 


an 


Rhodora Plate 1017 


Photo B. G. Schubert 


NANTHIUM ORIENTALE: FIGS. 1, 3, 5 and 7 burs, X 2; Fics. 2, 4, 6 and 8 beaks, X 5; FiGs, 
1-4 from European plants, eras. 5-8 from plants adventive in America, 


1946] : Fernald,—Studies on North American Plants 77 


(“squamis linearibus", also *lanceolatis"). It surely has nothing 
to do with H. atrorubens. In fact Bertoloni knew that, specially 
saying “Ab Heliantho atrorubente L. diversus". His plate sug- 
gests H. heterophyllus Nutt. 

Another and really quite different species, Helianthus silphio- 
ides, is included by Watson under his aggregate H. atrorubens, a 
somewhat unexpected attitude, in view of the many minor vari- 
ants in other series recognized by him as “new species", minor 
variants which forced the obliging but always cautious Dr. C. C. 
Deam to write in his Flora of Indiana, “I at first attempted to 
construct a key to our species using Watson's determinations", 
and to state in conclusion: “I have excluded 15 species that have 
been reported for the state". Surely H. silphioides Nutt. in 
Trans. Am. Phil. Soc. vii. 366 (1841), found from “the plains of 
Arkansa" and southeastern Missouri up the Mississippi drainage 
to southwestern Indiana, thence south to Alabama and Louisiana, 
later described from along the Mississippi in Kentucky as H. 
kentuckiensis McFarland & Anderson in Am. Mid. Nat. ix. 139, 
t. 10, (1924),—surely this very distinct plant cannot be satis- 
factorily referred to the more eastern H. atrorubens, as is done by 
Watson. In fact, Nuttall specially noted after the detailed de- 
scription of H. silphioides, “Allied to H. atrorubens but” etc., just 
as Bertoloni had done after his description of the equally distinct 
H. gracilis. It almost appears as if both mentions of H. atroru- 
bens had been mistaken as identifications with it. 

To be sure, Dr. W. A. Anderson, coauthor of H. kentuckiensis, 
subsequently pointed out! that H. kentuckiensis is certainly iden- 
tical with H. silphioides Nutt., as well as with the type of H. 
atrorubens, var. pubescens Kuntze, l. c. At that time Anderson 
considered the tall and very leafy plant, with slenderly short- 
petioled cuneate- or truncate- to cordate-based leaves running 
high and nearly uniformly up the stem, the pubescence of stem 
and roundish leaves short and close, as a Mississippi Valley 
extreme of H. atrorubens of “the Atlantic coastal plain, and . . . 
the mountains of North Carolina, Tennessee and Georgia’’, the 
latter with the stem hispid to villous below (our rra. 2), the 
leaves ''ovate to oblong-lanceolate, . . . tapering or abruptly con- 
tracted into a winged petiole”, the “lower surfaces of the leaves 


1W. A. Anderson in RHODORA, xxxiv. 1—4 (1932). 


78 Rhodora [APRIL 


smoothish, with long hairs on veins and petiole’. Anderson 
further noted that “Before publishing Helianthus kentuckiensis 
as a new species, Dr. McFarland and the writer sent a specimen 
to Mr. E. E. Watson who was then preparing a monograph of 
the genus. It was at his suggestion that the plant was described 
as a new species. Later, when his monograph appeared, Watson 
disposed of H. kentuckiensis as ‘not a Helianthus’.” — 

The photograph of the type of Helianthus silphioides, taken by 
Dr. S. F. Blake at the British Museum, and the original plate of 
H. kentuckiensis might have come from the same material, and, 
after studying the type of Otto Kuntze’s H. atrorubens, var. 
pubescens (the “varietal name . . . not a happy one") from 
Cairo, Illinois, Anderson found it quite like the others. Dr. 
Anderson sent a root from the type-colony of H. kentuckiensis to 
the Harvard Botanic Garden, the crown then carrying a number 
of stems. That original plant, increasing by many deep-seated 
subterranean stoloniform roots and, as Mr. Francis Lazenby, the 
Superintendent, informs me, not here coming from the regularly 
scattered achenes, is now a clone or colonial plant of about 50 tall 
uniformly leafy fertile stems. Such a root-system, with the 
many elongate and stoloniform deep roots ending in vegetative 
buds and completely covering an herbarium-sheet is wonderfully 
displayed by a plant, mounted on three sheets, sent to the Gray 
Herbarium by Mr. Ralph M. Kriebel from a colony covering 
about 100 yards in Lawrence County, Indiana (Kriebel, no. 
3965), tentatively placed with H. scaberrimus. Although the 
Kriebel plant has the leaves less rounded at base than in most 
typical H. silphioides, it is well matched by the foliage of the 

upper half of the stem of Demaree, no. 16,543 from Arkansas, as 
well as that of Buckley's material from Alabama. Its involucres, 
flowers and achenes are those of H. silphioides.. How unlike the 
habit of H. atrorubens! The latter, with which I have become 
very familiar in the field, has a simple, short, horizontal or ascend- 
ing premorse rhizome which rarely attains a length of 4 cm. It 
does not form clones, but the plants regularly have a single 
flowering stem, arising from the axil just beyond the remnants 
of the stem of the preceding year. Most of its leaves are borne 
from near the base of the tall stem, these on very long upwardly 
winged petioles. Far separated from these subbasal leaves there 


1946] Fernald,—Studies on North American Plants 79 


is usually a pair of smaller but petioled leaves, borne one sixth to 
one fourth the way up the stem; then far above them a few much - 
smaller and eventually merely bracteiform leaves up the almost 
scapose upper two thirds of the stem. With such differences of 
subterranean habit, pubescence, shape and distribution of leaves 
and geographic isolation, H. silphioides and H. atrorubens seem 
to me as distinct as any two species of Helianthus. 


HELIANTHUS LAETIFLORUS Pers. var. rigidus (Cass.), stat. 
nov. Harpalium rigidum Cass. Dict. Sci. Nat. xx. 200 (1814). 
Helianthus scaberrimus Ell. Sk. n. 423 (1823). Helianthus 
rigidus (Cass.) Desf. Cat. Hort. Par. ed. 3: 184 (1829). Hel. 
rigidus, forma flavus Deam, Fl. Ind. 975 (1940). 

H. LAETIFLORUS, var. subrhomboideus (Rydb.), comb. nov. 
H. subrhomboideus Rydb. in Mem. N. Y. Bot. Gard. i. 419 (1900). 
H. scaberrimus, var. subrhomboideus (Rydb.) Farwell in Am. 
Midl. Nat. viii. 278 (1923). 


When Deam defined Helianthus rigidus, forma flavus he 
“started something". Up to that time the very obvious division 
of Helianthus into a series with lobes of the disk-corollas purple, 
as opposed to those with corolla-lobes yellow, had been almost 
unchallenged, although it was well known that the commonly 
cultivated H. annuus may have ''sports" with yellow, instead of 
the more frequent dark purple disk. As early as 1789 Lamarck, 
in his Dict. iii. 86, got confused and described as the quite 
different H. atrorubens L., a species then cultivated in the Jardin 
du Roi in Paris, a very scabrous plant with spatulate-oval 
harshly scabrous acuminate and triple-nerved leaves which 
tapered to short petioles; heads very showy, resembling those of 
Rudbeckia * mais qui sont plus grandes & ont plus d'éclat", 
these on long scabrous peduncles; the phyllaries erect and lanceo- 
late; the disk “d’un jaune foncé". So obviously not the Lin- 
nean H. atrorubens, the plant of Lamarck was redefined by Per- 
soon as H. laetiflorus Pers. Synop. ii. 476 (1807). H. laetiflorus, 
passed on to other large European gardens, was several times well 
illustrated and is well known as a native or as a garden-escape 
through much of eastern North America. When he discussed it 
in the Synoptical Flora Asa Gray, still clinging to the primary 
divisions, “Disk . . . dark purple or brownish” as opposed to 
“Disk yellow", placed H. laetiflorus immediately after H. rigidus 
but as the first species with “Disk yellow", and defined it ‘“‘ Re- 


80 Rhodora [APRIL 


sembling tall forms of the preceding, similarly scabrous or hispid”, 
etc., the plant recognized from ‘Indiana, Illinois, Wisconsin". 
When Deam said, under ''Excluded Species", of H. laetiflorus 
“I am excluding the species from our flora and referring speci- 
mens so named to the yellow flowered form of Helianthus rigidus 
(Cass.) Desf.", he was right as to the identity but, unfortunately, 
he overlooked the fact that the specific epithet laetzflorus started 
in 1807, while H. rigidus was based on a name which was first - 
published in 1814. 

I am treating Helianthus rigidus as a variety rather than a 
color-form. In general it has thicker leaves than typical H. 
laetiflorus and its natural range is more western. Similarly, 
although Deam and, before him, E. E. Watson, got nothing out of 
H. subrhomboideus, which admittedly passes into western phases 
of H. laetiflorus, var. rigidus, it stands apart on several characters. 

Var. rigidus has a very scabrous stem 0.7-2.5 dm. high; var. 
subrhomboideus a slender and less scabrous (especially above) 
stem 0.15-1.2 m. high. Var. rigidus has 7-15 leafy nodes, the 
lance-oblong to lance-ovate leaves acuminate to long tips, the 
longer blades up to 3 dm. long; but var. subrhomboideus has only 
6-9 nodes, the leaves subrhombic and barely acute to bluntish, 
the larger ones only 0.5-1.5 dm. long. In var. rigidus the phyl- 
laries are lanceolate to narrowly ovate; in var. subrhomboideus 
more oblong or oblong-oval. Although westward it merges into 
typical var. rigidus, it has come east to Quebec and New England, 
chiefly via transcontinental railways, from plains of the West. 
Here it is very pure and shows little or none of the transition to 
var. rigidus which occurs farther west. 


EXPLANATIONS OF PLATES 1007-1020 


Prare 1007, Xyris Bavanpi Fernald, all figs., from TYPE: Fie. 1, four 
plants, X 1; FIG. 2, two spikes, X 5; ria. 3, tip of basal leaf, X 10; FIGs. 4 and 
5, lateral sepals, X 10; Fic. 6, seeds, X 10. 

Puate 1008, X. BRevIroLIA Michaux and X. FLABELLIFORMIS Chapman. 
Fias. 1-4, X. BREVIFOLIA: FIG. 1, bases of plants, X 1, from the general type- 
region, between Guyton and Ogeechee Rivers, Effingham Co., Georgia, 
Harper, no. 920; FIG. 2, basal leaf, X 10, from no. 920; FIG. 3, group of spikes, - 
X 1, from the Tyre of Michaux, after photo. by Cintract; ria, 4, spike, X 5, 
from no. 920. Fies. 5-7, X. FLABELLIFORMIS, all figs. from ISOTYPE: FIG. 5, 
base of plant, X 1; rra. 6, tip of basal leaf, X 10; ria. 7, spike, X 5. 

Prate 1009, ric. 1, PEDICULARIS CANADENSIS L.: characteristic plant, X 24, 
from Brooklin, Maine, A. F. Hill, no. 1419. Fies. 2 and 3, var. DOBBSII 
Fernald: ria. 2, characteristic base of plant, X 34, from Spring Grove, Minne- 
sota, Rosendahl, no. 276; Fic. 3, flowering raceme, X 1, from no. 276. 

PLATE 1010, P. CANADENSIS, var. DonBssir, both from TYPz: FIG. 1, base of 
plant, X 24; FIG. 2, fruiting raceme, X 1. 


Rhodor: Plate 1018 


Photo B. G. Schubert 


NANTHIUM ECHINATUM: FIG, 1, fruiting branch, X 1; ria. 2, margin of large leaf, X 1; 
FIGS. 3 and 5, burs, X 2; rics. 4 and 6, beaks, X 


Rhodora Plate 1019 


Photo B. G. Schubert 


XANTHIUM VARIANS: FIG. l, portion of IsovyPE, X 1; FIG. 2, bur, X 2; FIG. 3, summit 
of bur to show beaks, X 5. 


Rhodor: Plate 1020 


Photo B. G. Schubert 


HELIANTHUS ATRORUBENS, var. ALsODEs, all figs. from type: FIG. 1, plant, X 34; 
FIG. 2, pubescence of base and petioles, X 2; ric. 3, involucre, X 2. 


1946] Lakela,—Plants from Minnesota 8I 


Puate 1011, X SoripAGOo HIRTIPES Fernald, all figs. from TYPE: FIG. 1, 
portion of plant, X l4; FIG. 2, pedicelled heads, X 5; FIG. 3, pedicels, X 10: 
FIG. 4, upper, and FIG. 5, lower surface of leaf, x 10. 

PLATE 1012, S. GRAMINIFOLIA (L.) Salisb., var. NuTTALLII (Greene) Fernald 
and S. MICROCEPHALA (Greene) Bush. Fics. 1-3, S. GRAMINIFOLIA, var. 
NvurrALLH, all from near Alexandria, Virginia, Blake, no. 8697: Fic. 1, glomer- 
ules, X 5; FIG. 2, branchlet and pedicel, X 10; rra. 3, lower surface of leaf, 
X 10. Fies. 4, 5 and 6, S. MICROCEPHALA, all ‘from the Pepe repo, Sumter 
Co., Georgia, Harper, no. 636: FIG. 4, pedicelled heads, X 5; rra. 5, branchlet 
and pedicels, X 10; rra. 6, upper surface of leaf, X 10. 

PraTE 1013, XaNTHIUM CHaskr Fernald and X. srrumarrum L. Fra. E; 
X. CHASEI, TYPE, X 3/; Fias. 2 and 3, X. srRUMARIUM: FIG. 2, bur, X 2, 
from Bavaria, Killermann, in Herb. Exsic. Bavar., no. 1226; ria. 3, same bur, 


5. 

PraTE 1014, X. Cnaskr Fernald: ria. 1, portion of TYPE, X 1; FIG. 2, bur, 
X 2, from TYPE; FIG. 3, portion of bur, to show beaks and bulbous-based 
prickles, X 5, from TYPE; FIG. 4, bur, X 2, from V. H. Chase, no. 3474; Fia. 5, 
summit of bur, X 5, from no. 3474. 

PrATE 1015, burs, X 2, and beaks, X 5, of X. GLoBosuM Shull, X. ITALICUM 
Moretti, X. INFLEXUM Mackenzie & Bush and X. CHINENSE Miller. Fics. 
l and 2, X. GLOBOSUM, from plant raised by Sherff from seed of TvPE. Fras, 
3 and 4, X. ITALICUM, from Venetia, Béguinot in Fl. Italica Exsice., no. 2774. 
Figs. 5 and 6, X. IiNFLEXUM, from Courtney, Missouri, Bush, no. 1806. Fics. 
7 and 8, X. CHINENSE, from Ogdensburg, New York, Phelps, no. 1215. 

PrATE 1016, burs of X. curvescens Millspaugh & Sherff and X. LEPTO- 
CARPUM Millspaugh & Sherff, smaller figs. X 2, larger, X 5. Fras. 1-4, X. 
CURVESCENS: FIGS. 1 and 2, from TYPE; FIGS. 3 and 4, from Burlington, Ver- 
mont, Sept. 8, 1918, N. F. Flynn (identification by Sherff). Frias. 5 and 6, 
X. LEPTOCARPUM, from TYPE. 

PraATE 1017, burs of X. ORIENTALE L., figs. 1, 3, 5 and 7, X 2, others X 5; 
FIGS. 1 and 2, from Austria, Fritsch in Fl. Exsic. Austro-Hung., no. 3068; 
FIGS. 3 and 4 (2 burs), from France (as X. macrocarpum DC.), ex Herb. Cosson- 
Germain; FIGs. 5 and 6, from Richelieu River, St. Hilaire, Quebec, Pease, no. 
12,955; rias. 7 and 8, from LaTortue, Quebec, Victorin, no. 21,254. 

PLATE 1018, X. kecuiNATUM Murray: FIG. 1, fruiting branch, X 1, from 
Revere, Massachusetts, Sept. 17, 1882, Herbert A. Young; ria. 2, margin of 
large leaf, X 1, from Marshfield, Massachusetts, August 28, 1898, C. H. 
Morss; FIG. 3, bur, with arching beaks, X 2, from Tisbury, Massachusetts, 
F. C. Seymour, no. 2029; ric. 4, summit of bur, X 5, from no. 2029; rra. 5, bur, 
with tightly crossing beaks, X 2, from Newcastle, New Hampshire, Sept. 19, 
1901, E. F. Williams; ria. 6, summit, X 5, of same bur as in fig. 5. 

PraTE 1019, X. vARIANS Greene: Fic. 1, portion of ISOTYPE, X 1; FIG. 2, 
bur, X 2, from The Dalles, Oregon, Suksdorf, no. 193; ria. 3, summit of bur, 
X 5, from no. 193. 

PLATE 1020, HELIANTHUS ATRORUBENS L., var. ALSODES Fernald, all figs. 
from TYPE: FIG. 1, plant, X 24; FIG. 2, pubescence c of base nie petioles, x 2; 
FIG. 3, involucre, x2. 


ex d d 


yi abon UNREPORTED PLANTS FROM BUS Wn" 
San TRIVIALIS L. Several years ago Dr. C. O. Rosendahl collected 
= . P. trivialis growing in association with P. sylvestris Gray and 


. Milium effusum L., in northeast Iowa, less than one mile from 
the Iowa-Minnesota state line. It appears to be the closest 
known station to that of Duluth. On July 19, 1945, the writer 
discovered the species on the bank of Tischer Creek at the foot of 


82 Rhodora [APRIL 


Hunters Hill. Collection no. 6007 was made from a colony 
growing in wet moss in deep shade under willows, not far from a 
colony of Listera auriculata Wieg., a rare orchid in Minnesota. 
P. trivialis is an interesting addition to the adventive flora 
established on the higher slopes of Hunters Hill, e. g. P. Chaixii, 
P. nemoralis, Milium effusum and Luzula nemorosa. 

HELENIUM NUDIFLORUM Nutt. is extending its range farther 
into the interior west from Michigan and north from Missouri. 
In Minnesota the writer discovered a colony of the species on 
Highway 23, about two miles south of the Holyoke Junction, in 
Carlton Co. Collection Lakela no. 5860 was made on Sept. 4, 
1944, from a colony of scattered individual plants growing in 
shallow moss-mats with species of Juncus and Carex, along a 
partly cleared border of the roadside thicket. 

. EQUISETUM ARVENSE L., f. CAMPESTRE (C. F. Schultz) Klinge 
.was discovered in water-filled depressions made by removal of 
sod, in a brookside-meadow northwest of Woodland Ave. in 
Duluth. Specimens no. 5985 collected June 23, 1945, have more 
chlorophyll-containing tissue in fertile stems than later formed 
plants. Strobili continued to develop through August. The 
writer is obliged to Dr. R. M. Tryon, Jr., University of Minne- 
sota, for determination of the form.—OrcA LAKELA, Duluth 
State Teachers College. 


BIDENS HYPERBOREA VAR. TYPICA 
NorMan C. FASSETT 


Bidens hyperborea was described in 1901! from material col- 
lected at Rupert House, James Bay, by J. M. Macoun in 1885, 
and its specific identity with plants on estuaries from the St. 
Lawrence to New England was later demonstrated.? 

“The treatment of this variable species by the present writer 
(1925c)? can be regarded only as tentative. Var. typica Fassett 
came from Rupert House, James Bay, Canada, and is described 
as being simple and monocephalous, with entire or few-toothed 
leaves. Plants matching this description were collected by 
Svenson and Fassett on the estuary of the Restigouche River, 


1 Greene, Pittonia iv. 257 (1901). 
2 Fernald, Ruopona xx. 146-150 (1918). 
3 Fassett, RHopona xxvii. 166-171 (1925). 


1946] Fassett,—Bidens hyperborea var. typica 83 


New Brunswick; this collection is cited (Fassett, 1925c) as No. 
889 under var. laurentiana. The unbranched plants with small 
achenes formed colonies distinct from those of the larger variety 
also present on this estuary, and were considered in 1925 to be 
mere dwarfed forms of it. This is probably the case, but it is 
possible that the type collection from James Bay, the only collec- 
tion known, may represent small forms of a larger plant present 
on the estuaries of rivers entering Hudson Bay. The final word 
may not be spoken until complete collections have been made 
in that region.’”4 

Copious collections of Bidens hyperborea have recently been 
made by Rev. Arthéme Dutilly, O. M.I. and Rev. Ernest Lepage, 
at Rupert House and East Main River; while it would be over- 
optimistic to try to speak “a final word", these collections show 
conclusively that the type material does represent only dwarfed 
forms such as may be found in almost any colony of any species 
of Bidens. "These new collections total 46 individuals, of which 
16 are simple and monocephalous like the type, 17 are slightly 
branched above, and 13 are copiously branched. The leaves, 
except on the smallest individuals, are conspicuously toothed, 
and the achenes are for the most part definitely larger than 
those of the type specimens, the outer being 4.5-7.5 mm. long, 
the inner 6.0-9.0 mm. long, and the awns 3.0-4.5 mm. long. In 
short, just as small individuals of var. laurentiana are inseparable 
` from the type material of B. hyperborea, so well-developed 
plants from the type locality are inseparable from similar indi- 
viduals of var. laurentiana. B. hyperborea var. laurentiana 
Fassett, RHopona xxvii. 169 (1925), therefore becomes a syno- 
nym of B. hyperborea var. typica Fassett, l. c., p. 167. 

The Dutily & Lepage collections, all of which have been 
loaned to me by Father Lepage, are as follows: Rupert House, 
51° 30’ N., August 31-September 2, 1944; nos. 13576, 13633 & 
13660; East Main River, 52? 15' N., August 28-29, 1944, nos. 
13485 and 13522. For the loan of the type of Bidens hyperborea 
from the National Herbarium of Canada I am indebted to Dr. 
A. E. Porsild. 


DEPARTMENT OF BOTANY, 
UNIVERSITY OF WISCONSIN 


4 Fassett, Proc. Boston Soc. Nat. Hist. xxxix. 103 (1928). 


84 Rhodora .. [Arni 


THE GENUS PALAFOXIA IN TEXAS 
V. L. Conx* 


. Mx No. 31195, which 1 had labeled as Palafoxia linearis Lag., 
was determined tentatively by Dr. Blake as being Othake macro- 
lepis Rydb. I know he was right in finding it not to be P. 
linearis, and my subsequent study of the material convinces me 
that he also was correct in using the interrogation mark after the 
O. macrolepis. Upon my request and in connection with a sum- 
mary of my study of this material, Dr. Blake has made a further 
examination of the material and has reported that it seems to 
represent a valid new species. 


PALAFOXIA riograndensis, new species. Plant annual; stems 
3-4.5 dm. high, up to 5 mm. broad at base, strigose and hispid 
throughout, branched at base and above, the branchlets hispid 
and densely glandular, especially above; leaves linear, up to 5 
em. long and 3 mm. broad, petiolate, strongly hispid; petioles 
slender, 1 cm. long or less; peduncles slender, densely glandular, 
up to 6 cm. long, the shorter ones usually more than 2 cm. long; 
involuere narrowly turbinate-campanulate, about 1 cm. long and 
0.5 em. broad; involucral bracts in a single series, 4-6, usually 5, 
each closely embracing an outer achene, linear, hispid and gland- 
ular, 9-10 mm. long, with rose-colored tips; ray-flowers wanting; 
disk-flowers twice as many as there are involucral bracts, 5.5-6.5 
mm. long, the corolla cleft nearly to the slender tube which is 
2-2.5 mm. long, the lobes linear; achenes tapering downwards, 
8-10 mm., usually about 9 mm. long, 0.7-0.9 mm. broad at apex, 
the inner and outer ones markedly different; outer achenes 
glabrate to sparsely strigose, epappose, or pappus minute, to 
2-3 mm. long, or frequently cup-like with 4 very short lobes, 
usually less than 0.5 mm. long, not at all scarious; inner achenes 
densely pubescent, with 4 squamellae about 6 mm. long with 
strongly excurrent midrib and scarious margins.! 


* Range Botanist, 'Texas Agr. Expt. Station, Sonora, Texas. 

! PALAFOXIA riograndensis sp. nov. Annua strigosa hispida 3—4.5 dm. alta caule basi 
5 mm. diam. ramosa, ramuli hispidi glandulosi. Folia linearia ad 5 cm. longa 3 mm. 
lata hispida petiolata, petiolis gracilibus 1 cm. longis vel brevioribus. Pedunculi 
graciles dense glandulosi ad 6 cm. longi (plerumque ultra 2 cm.). Involucra anguste 
turbinato-campanulata ca. 1 mm. longa 0.5 cm. lata. Phyllarii uniseriales 4—6 
(plerumque 5) achaenia exteriora amplectentes lineares hispidi glandulosi 9-10 mm, 
longi apice rosei. Corollae ligulatae desunt; disci corollae ca. 10, 5-6.5 mm, longae 
profunde lineari-lobatae fere ad tubum gracilem 2-2.5 mm. longum.  Achaenia basi 
angustata ca. 9 mm. longa apice 0.7-0.9 mm, diam. interiora ab exterioribus perspicue 


differentia, exteriora glabrata vel parce strigosa epapposa aut pappum minutum ad . 


2-3 mm. longum aut brevissime quadrilobatum ad 0.5 mm. longum nec scariosum 
gerentia; interiora dense pubera squamellas 4 ad 6 mm. longas costa excurrente mar- 
ginibus scariosis gerentia. 


Ne 


1946] ... Cory,—Genus Palafoxia in Texas 85 


TyPE specimen is designated as No. 31195, which is deposited 
at the Gray Herbarium, and isotype material is deposited at other 
herbaria. This collection was made on October 26, 1938, in a 
flat wash near the Rio Grande about three and one-half miles 
southeast of Presidio in Presidio County, Texas. We have seen 
this plant nowhere else, but it is assumed to grow elsewhere along 
the Rio Grande in the Big Bend Area of Texas, and in strong 
probability to occur more widely and in greater abundance across 
the river in Mexico. It seems fitting to associate the species in 
name with the river along which it grows. 

Othake macrolepis Rydb., a species occurring 550 or more air- 
. line miles north-northeast in Bent County, Colorado, has been 
reported from Texas: Demaree 7723, three miles north of Lub- 
bock, May 27, 1930. Lubbock is approximately 325 miles from 
Presidio and only about one hundred twenty-five miles from the 
Colorado locality. I have not seen Mr. Demaree’s specimen, 
but I presume it is well authenticated. In O. macrolepis the 
heads are said to be short-pedunculate, whereas in P. riogranden- 
sis they are relatively long-pedunculate. Among other differ- 
ences between these two species ours has much narrower leaves 
(ca. 3 mm. vs. ca. 8 mm.), a lesser number of involucral bracts 
(5 vs. 8-12), shorter corolla-tube (2-2.5 mm. vs. 5 mm.), longer 
achenes (9 mm. vs. 7 mm.) and fewer squamellae (4 vs. 6-8). 

The pappus of the new species is more nearly that of Palafoxia 
linearis than that of Othake macrolepis, and one well-known 
botanist did as I did by referring my material to Palafoxia 
linearis largely through this character. It would seem that this 
linking of two species, which some botanists consider as being of 
different genera, would be good support for placing these three 
species in a single genus, and this necessarily would be Palafozia. 
In treating the Rio Grande species in this manner, it seems well 
to propose new combinations for those species known from Texas 
hitherto as being of the genus Polypteris and Othake. 

Recently I have examined the publication, A MONOGRAPH- 
IC STUDY OF THE GENUS PALAFOXIA AND ITS IM- 
MEDIATE ALLIES by Elizabeth Ammerman Baltzer in which 
the treatment given is substantially that of Rydberg in the 
NORTH AMERICAN FLona. The author did not have Demaree’s 
plant from Lubbock, nor did she have my plant from Presidio. 


86 Rhodora [APRIL 


In my opinion she is right in considering Othake robustum Rydb. 
as being a variety of O. roseum Bush, but I do not agree with her 
in placing O. macrolepis Rydb. as a variety of O. texanum (DC.) 
Bush. From this viewpoint five new combinations for Texan 
species are required. 

ParaArFoxiA rosea (Bush), new comb. Othake roseum Bush in 
'Trans. Acad. Sci. St. Louis 14: 175, 1904. 

PALAFOXIA ROSEA, var. robusta (Rydb.), new comb. Othake 
robustum Rydb. in N. Am. Fl. 34, Pt. 1, 60, 1914. 

PALAFOXxIA macrolepis (Rydb.), new comb. Othake macro- 
lepis Rydb. in Bull. Torr. Bot. Club. 37: 332. 1910. 

PaLAFOXIA Reverchonii (Bush), new comb. Othake Rever- 
chonii Bush in Trans. Acad. Sci. St. Louis 14: 180. 1904. 

Pavaroxia sphacelata (Nutt. ex Torr.), new comb.. Stevia 
sphacelata Nutt. ex. Torr. in Ann. Lyc. N. Y. 2:214. 1828. 

I am indebted to Dr. I. M. Johnston and to Dr. S. F. Blake for 
critical study of the plant material, and to Dr. L. H. Shinners for 
assistance in preparation of the Latin description. 


Present address: 


INSTITUTE OF TECHNOLOGY AND PLANT INDUSTRY, 
SouTHERN METHODIST University, Dallas 5, Texas. 


Tue NORTH AMERICAN REPRESENTATIVES OF ALISMA PLAN- 
TAGO-AQUATICA.—To one who has long known the two broad- 
leaved representatives of Alisma Plantago-aquatica L. in North 
America it is a surprise to see growing in Europe the typical 
plant, for it commonly has lilac or roseate petals (our two plants 
with them white), while the stamens, ovaries and styles are 
markedly different from ours. To be sure, the late Professor 
Gunnar Samuelsson treated our larger-flowered northern plant 
as & North American subspecies, A. Plantago-aquatica, subsp. 
brevipes (Greene) Samuelsson in Arkiv fór Bot. xxiv^, no. 7: 19 
(1932), based upon A. brevipes Greene, Pittonia, iv. 158 (1900). 
When we compare the latter plant (which occurs across North 
America, from Quebec to British Columbia, south to Nova. 
Scotia, New England, Maryland, Michigan, Iowa, Nebraska, 
New Mexico, Arizona and northern Mexico) with true Old World 
A. Plantago-aquatica, surprisingly definite characters are found 
to separate them. These are concisely stated below: 


1946] Fernald,—Alisma Plantago-aquatica 87 


True ALISMA PrANTAGO-AQUATICA. Petals commonly lilac or roseate, 
sometimes white; anthers 1-1.25 mm. long; style 1-1.25 mm. long, erect, 
. Straight, median at summit of flowering carpel, the latter tapering to style. 

A. TRIVIALE Pursh, Fl. Am. Sept. i. 252, mostly (1814). A. brevipes Greene. 
Petals usually white; anthers 0.6-0.8 mm. long; style 0.5-0.7 mm. long, 
strongly curved, borne at ventral side of summit of margin of the flowering 
earpel, the dorsal keel of the latter broadly rounded. 


It would seem that our boreal Alisma triviale is well distin- 
guished from the Old World plant. 

The differentiation of our relatively northern Alisma triviale 
and the relatively southern A. subcordatum Raf. (Florida to 
Texas and Mexico, north to New England, New York, southern 
Ontario, Michigan, Wisconsin, Minnesota and Nebraska) was 
made by Pursh. Michaux, Fl. Bor.-Am. i. 218 (1803) had A. 
Plantago only from Canada, where A. triviale abounds. Pursh, 
Fl. Am. Sept. i. 252 (1814), took over, with unimportant change, 
Michaux’s diagnosis of the Canadian plant, citing Michaux as 
his basis, but stretching the range from. “Canada to Florida", 
with the new name A. triviale. Then from * New Jersey and 
Pennsylvania" his new A. parviflorum (p. 253) with “flowers 
small". It is quite evident that Pursh intended the larger- 
flowered and generally more northern plant as A. (riviale, the 
smaller-flowered and generally more southern plant as his A. 
parviflorum. Rafinesque, however, had already defined the latter 
in 1808 as A. subcordatum. 

The essential bibliography and the best specific characters of 
the two follow: 

A. TRIVIALE Pursh, Fl. Am. Sept. 1. 252 (1814) in large part at 
least. A. Plantago sensu Michx. Fl. Bor.-Am. i. 218 (1803), not 
L. Syst. ed. 10, 11. 993 (1759). A. Plantago, s. americanum 
Roem. & Schultes, Syst. vii. 1598 (1830). A. Plantago, var. 
triviale (Pursh) BSP. Prelim. Cat. N. Y. Pl. 58 (1888). A. 
Plantago-aquatica, var. triviale (Pursh) Farwell in Ann. Rep. 
Comm. Parks and Boulev. Detroit, xi. 44 (1900). <A. brevipes 
Greene, Pittonia, iv. 158 (1900). A. superbum Lunell, Bull. 
Leeds Herb. ii. 5 (1908). A. Plantago-aquatica, ssp. brevipes 
(Greene) Samuelsson in Arkiv fór Bot. xxiv^. no. 7: 19 (1932). 
A. Plantago-aquatica, var. brevipes (Greene) Victorin (wrongly 
ascribed to Samuelsson), Fl. Laurent. 615 (1935).—Flowers 9-13 
mm. broad; sepals broadly scarious-margined, in anthesis 3—4 
mm. long; petals 3.5-6 mm. long; stamens twice as long as 
ovaries; anthers ovoid, 0.6-0.8 mm. long; style about equaling 
carpel at anthesis; fruiting heads chiefly 4-7 mm. in diameter; 
achenes 2.2-3 mm. long. 


88 Rhodora [APRIL 


A. sUBCORDATUM Raf. in Med. Repos. ii. 5 (1808). A. parvi- 
florum [as parviflora] Pursh, l. c. 253 (1814). A. Plantago, Q. 
parviflorum (Pursh) Torr. Fl. N. Mid. U. S. 362 (1824). A. 
Plantago-aquatica, var. parviflorum (Pursh) Farwell, l. c. (1900).— 
Flowers 3-3.5 mm. broad; sepals very narrowly margined, in 
anthesis 2-2.5 mm. long; petals 1-2 mm. long; stamens only 
slightly exceeding ovaries; anthers subspherical, 0.3-0.5 mm. 
long; style about one fourth as long as ovary; fruiting heads 
mostly 3-4 mm. broad; achenes 1.5-2 mm. long.—M. L. FERNALD. 


THE SPORADIC APPEARANCE OF Eprpactis HELLEBORINE.—It 
has repeatedly been noted, since it first appeared as a naturalized 
plant in North America, that Epipactis Helleborine (L.) Crantz 
will suddenly appear in well known woodlands, wooded parks, 
ravines or thickets as a single individual, soon as several and in 
a few years as a relatively abundant species. This behavior is 
apparently not restricted to the American colonies, derived from 
European progenitors. Picking up, almost at random, Ben- 
tham's Handbook of the British Flora, ed. 4 (1878), I read 
(p. 457) under E. latifolia (L.) Sw., a variation of E. Helleborine: 
“Not unfrequent in Britain, but often appearing only in single 
specimens".—M. L. FERNALD. 


Volume 48, no. 567, containing pages 41—64 and plates 1005-1010, was issued 
4 March, 1946. 


0 1946 


odora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED eee 
ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS 


Associate Editors 


Vol. 48. May, 1946. No. 569. 

CONTENTS: 
Notes on certain Plants in the Gray’s Manual Range. 

Hart award Sherry rar cer IEEE oP ee? a 89 
Botanical Visits to Forts Clark, Mandan and Union in North 

Dakota FeO PMc lene ete. a ee 98 
Populus balsamifera of Linnaeus not a Nomen ambiguum. 

Ee ROU CGU SE Cu. cU EL ee 103 
Size, Shape and Number in Astragalus caryocarpus Fruits. 

D uw Hac oo os LIEN LL. PE un 111 
Spiraea latifolia var. septentrionalis in Virginia. 

bh T I ee ye EMI CUL 112 
Helianthus—a Correction. M. L. Fernald. ................... 112 


The New England Botanical Club, Ine. 


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


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at $4.00. Some single numbers from these volumes can be supplied only at ad- 
vanced prices (see 3rd cover-page). Somewhat reduced rates for complete sets can 
be obtained on application to Dr. Hill. Notes and short scientific papers, relating 
directly or indirectly to the plants of the northeastern states, will be considered for 
publication to the extent that the limited space of the journal permits. Forms may 
be closed five weeks in advance of publication. Authors (of more than two pages of 
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Address manuscripts and proofs to 
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No. HI. The Linear-leaved North American Species of Potamogeton. 
Section Axillares, by M. L. Fernald. 183 pp., 40 plates, 31 maps. 1932. 
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Grav Herbarium of Harvard University, Cambridge 38. Mass. 


Rhodora 


Plate 1021 


5). 


8, 194 


author, Julv 2 


wger colonies of ILIAMNA REMOTA var. TYPICA on Altorf Island (photograph by 


« 


One of the | 


Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. May, 1946. No. 569. 


NOTES ON CERTAIN PLANTS IN THE GRAY’S 
MANUAL RANGE 


EARL EDWARD SHERFF 


Plates 1021-1024 


ILIAMNA REMOTA Greene, Leaflets Bot. Crit. 1: 206. 1906.— 
It is now more than eighteen years since this rare species, there- 
tofore known only from an island in the Kankakee River (at 
Altorf, about nine miles northwest of Kankakee, Illinois), was 
discovered growing on Peters Mountain in Virginia. Straus- 
baugh & Core, in an early volume of RHopona (34: 142. 1232), 
have presented, under the synonym Phymosia remota (Greene) 
Britton, an interesting account of its discovery at the latter 
habitat. Their paper was followed closely by another, entitled 
“Phymosia remota in captivity” (S. C. Wadmond, op. cit. 207). 
Both of these papers were so stimulating to lovers of our rarer 
plant species that it has seemed worth while to supplement them 
with certain additional notes and remarks. 

Rev. E. J. Hill, who collected the first herbarium material 
June 29, 1872, was at that time a teacher in the Kankakee High 
School! Apparently he did not realize for some years the im- 
portance of his find. It became referred to Sphaeralcea aceri- 
folia Nutt. in Asa Gray’s Synoptical Flora (1: 317. 1897). On 
August 1, 1899, Dr. Edward L. Greene revisited the exact locality 
where Mr. Hill had collected in 1872 and obtained more speci- 
mens. He later assigned them to his new genus Iliamna, in 
which he erected for them the new species I. remota. His cited 


! For a good account of his long and fruitful life, see Mrs. Agnes Chase's article, 
with plate, RHODORA 19: 61—69. 1917. 


90 Rhodora [May 


herbarium specimens were all of his own collecting (none by Hill), 
so of course the type specimen would be a plant by Greene.’ 

In 1908, Fernald, Rnopona 10: 52, renamed Greene’s Kanka- 
kee plant Sphaeralcea remota. 

In 1912, Dr. N. L. Britton was concluding his revisional work 
upon Britton & Brown’s Illustrated Flora (edit. 2) and wished 
to secure an ampler assortment of specimens of the Kankakee 
plant. He wrote to the authorities of Field Museum of Natural 
History (now the Chicago Natural History Museum) for aid. 
Dr. Jesse M. Greenman, then of that institution, accordingly 
undertook to journey to the type locality. The elderly Mr. Hill, 
then in his seventy-ninth year, and myself were invited to ac- 
company him. Arriving in the forenoon of August 3rd at Altorf, 
just northeast of the island’s upper end, we forded the river at a 
point somewhat upstream, using a horse-drawn carriage. Se- 
curing a boat on the opposite shore, we rowed to the southwestern 
shore of the island and anchored. We were led presently by Mr. 
Hill with surprising directness and accuracy to the very spot 
where he remembered having collected some forty years before. 
The plants of /liamna remota—to use Greene’s original name 
unmodified—were numerous and both Dr. Greenman and myself 
secured a few herbarium specimens (Greenman 3530, Sherff 1600) 
and ripe seeds for Dr. Britton’s studies, also for distribution to 
various herbaria. It may be added here that Britton published 
the new combination Phymosia remota the following year (Britton 
& Brown loc. cit. 2: 522), and that, since then, several authors, 
among then Strausbaugh & Core (loc. cit.), have accepted 
Britton’s name. 

For some years before his death (in 1923), Dr. C. F. Millspaugh, 
the Curator of Botany in those days at Field Museum of Natural 
History, was keenly interested in the rare Kankakee plant (cf. 


2 Greene’s description of the island's position as ‘‘some twelve or fifteen miles 
above the city of Kankakee" was inexact. The island was indeed ''just opposite a 
small village [or what may once have been a village] called Altorf' as added by him, 
but the distance by road on either side of the Kankakee River measures roughly nine 
miles from the business center (particularly the courthouse) in Kankakee. 

3 Dr. Isaac Bayley Balfour, Regius Keeper of the Royal Botanic Garden at Edin- 
burgh, was among those sent some of the seeds. He wrote me perhaps three years 
later that the seeds had been planted on the grounds of his institution and had resulted 
in large, beautiful, flowering specimens that seemed well established. (Those who 
attempt to germinate seeds of this species might well scarify them first, between 
pieces of emory paper, or else soak them for 24 hours before planting, as mentioned 
in the above cited paper by Wadmond.) 


9) 


y author, July 28, 194 


INA REMOTA var. TYPICA on Altorf Island (photograph b 


One of the many smaller colonies of ILIAN 


1946] Sherff,—Plants in the Gray’s Manual Range 91 


Wadmond, loc. cit.). In 1916, on his suggestion, Mr. O. E. 
Lansing, Jr., of Field Museum, and myself were commissioned to 
make botanical trips to certain Illinois localities of special 
interest,‘ among them that on the Kankakee island. At this last 
locality we collected numerous specimens of /liamna remota 
(Lansing & Sherff 8, on rocky, grassy slope at Altorf Island, 
Aug. 23), supplementing these with a liberal quantity of mature 
seeds. Returning to Kankakee, we were graciously escorted by 
the late Judge Arthur W. De Selm to his residence and shown a 
number of fine flowering and fruiting specimens of I. remota 
growing in his garden. "These he had raised from seed personally 
obtained in the type locality several years before. 

Shortly afterward, Clute (Amer. Botanist 26: 127-129. 1920), 
writing under the caption, ‘‘The rarest American plant," told of 
finding, after long and what seemed destined to be an unsuccessful 
search in the type locality, “a single plant of the rare mallow.” 
He removed this plant to his private grounds, where it grew 
vigorously. Reading further: ‘‘We secured an abundance of 
seeds and hope by another year to have done something toward 
modifying its rarity." Strausbaugh & Core (op. cit. 146) cited 
Clute's paper and stated that, if his observation was correct, 
“it must then be apparent that the Virginia station is now the 
only known place in the world where Phymosia remota [t. e., 
Iliamna remota Greene] is growing as a wild plant, and since 
there are at the present time not more than 50 plants at this 
station the species must be regarded as an exceedingly rare one 
that may soon become extinct." 

'The experience recorded by Clute seemed to my mind some- 
what astonishing. In 1912 and again in 1916 I had observed 
that the more elevated, flat, level stretch of land on the Kankakee 
island (sometines called Altorf Island) was cultivated as a corn 
field. The Iliamna plants were found beginning at the edge of 
the corn field and extending a few meters down the more or less 
steep and wooded slope leading to the river. They were, we may 
say, in a narrow belt of open-woods habitat. Since this narrow 
belt was below where a plough would ever strike, it had seemed 
logical to assume that even with continued cultivation of the 


5 It is a curious coincidence that one of these trips was described by me in the very 
issue of Ruopora containing Mrs. Chase's biographic sketch of Mr. E. J. Hill (vol. 19, 
April, 1917; see pp. 74 & 75). 


92 Rhodora [May 


plateau-like field adjacent to this foothold they would be able to 
persist. 

Deciding at last to see for myself how Iliamna remota had 
fared on the Kankakee island, I made a return visit on July 23rd, 
1945, to the type locality. To my great delight, I found that 
farming had been abandoned and that there were several hun- 
dreds of Iliamna plants flourishing there. From a distance, 
they displayed in several places a massed effect and, in general 
habit, somewhat suggested hollyhocks. A good proportion 
were robust, several-stemmed, 1-1.7 meters tall, and abundantly 
flowered. The majority were on the open, level expanse of the 
island, where they may indeed have become re-established subse- 
quent to a one-time cultivation. A fair proportion, however, 
were on the upper part of the marginal rocky, grassy slope where 
I had seen them in earlier years.? At the time of day the plants 
were visited (one to two hours before true noon) the temperature 
was above 90° F. and the sunlight very bright. The flowers 
were fully expanded, although Wadmond reported (op. cit. 209) 
that on cloudy days the flowers never opened fully. Some five 
dozen herbarium specimens were obtained (my no. 5021) for dis- 
tribution to herbaria later on and one small living plant removed 
for my garden. The specimens were kept for more than four 
hours slightly moistened in a closed, standard-sized, metal 
household clothes-boiler, then put in a plant press. "Their 
flowers had remained fully expanded meanwhile. 

On the 28th of the same month, I returned to the type habitat, 
in company with Dr. G. S. Daston of the Chicago Natural 
History Museum. Photographs of the growing plants were 
taken (PLATES 1021 and 1022), two more plants were dug up for 
removal to my garden (where all three plants are now safely 
established), ripe seeds were obtained, and further observations 
were made. These may be given briefly: The soil was not 
gravelly as had been reported, but distinctly a brown, sandy clay. 


5 This time alone, Those who would visit this spot, if not good oarsmen, should 
have preferably a companion along. The river, while usually only 3 to 15 dm, deep in 
July and August, is very swift and has many treacherous currents, making progress 
uncertain and difficult. 

* It seems likely that Clute and his companions who visited the island in 1920 went 
to the wrong part of it. The older plants have remarkably long, thick, mostly hori- 
zontal roots and on the grassy, unmolested slope there seems little reason why the 
plants should have appeared successively in 1872, 1912, and 1916, then vanished by 
1920, and finally become established again by 1945. 


Rhodora Plate 1023 


ILIAMNA REMOTA var. TYPICA from Sherff 5021, type locality on Altorf Island, July 23, 
1945 (photograph by author from specimen in Gray  Herb.). 


1946] Sherff,—Plants in the Gray’s Manual Range 93 


In open, level situations colonies were up to 5 meters or more 
across, and essentially a pure stand (pl. 1021), the individual 
stems not tending to sprawl as they did where plants were grow- 
ing singly at the edge of the treeless plateau, on grassy slopes in 
partial shade. Commonly the plants were accompanied by the 
darkened and dead stems of the previous year (these show clearly 
in pl. 1022). Leaves nearly always had a broadly triangular 
terminal lobe, this widening regularly from apex to base and 
being subtended by wide, obtuse sinuses. A few plants had 
some leaves, especially on their branches, with terminal lobe ob- 
long or subcuneately narrowed below and subtended with sharp 
sinuses, as shown in Strausbaugh & Core’s plate for a Virginia 
plant. Some plants had numerous suberect, straightish, axillary 
branches + 2 dm. long, but for most plants these were unde- 
veloped or absent. The flowers had a very pleasing but delicate 
fragrance (indeed, the dried specimens after a lapse of three 
months are strikingly fragrant). The larger flowers measured 
easily 5 cm. in diameter. The petals were irregularly more or 
less emarginate and usually very inequilateral, with one terminal 
lobe much exceeding the other. 

In 1936, Wiggins published * A Resurrection and Revision of 
The Genus Iliamna Greene" (Contrib. Dudley Herb. Stanford 
Univ. 1: 211-230, pl. 20). Many data, references, and considera- 
tions advanced by him as being germane to a proper understand- 
ing of Iliamna remota and its congeners must be omitted here. 
I have been constrained, however, to follow Wiggins in taking up 
Greene's name Zliamna remota for the Kankakee plant, which, 
for more precise handling, may be known as var. typica (var. 
nov.)/. With the Kankakee plant, Wiggins merged the Virginia 
form, as indeed had been done by Strausbaugh & Core (vide 

7 Since preparing the text for this paper, I have learned of the recent remarkable 
discovery by Dr. S. W. Witmer of Goshen College, of the var. typica in Elkhart 
County, northern Indiana. Professor Witmer has very kindly sent me two specimens 
for examination and granted permission to announce his discovery. Under date of 
Nov. 23, 1945, he wrote: ‘‘One of these specimens I collected July 4, 1944, the day I 
discovered this species growing wild at the station to be mentioned below. The 
other specimen I collected from the same station on Aug. 28, 1945. 

“These plants were found 2 miles east of New Paris, Elkhart County, Indiana, at a 
point where the Wabash R. R. crosses the Elkhart River. I located four colonies of 
these plants over a stretch of ground extending about 430 yards from the river east- 
ward along the north side of the railroad right-of-way. The four colonies included 


about 73 more or less distinct clumps. On the latter date of collection many of the 
plants had grown between 5 and 6 feet tall.” 


94 Rhodora [May 


supra). However, the Virginia plant was known to be montane 
in its habitat, while the var. typica had been thought of as a 
prairie plant. Thus, for example, Gleason, writing some years 
ago on the vegetational history of the middle-western United 
States (Annals Assoc. Amer. Geographers 12: 39-85. 1922) men- 
tioned “Phymosia remota” in particular as one of certain plants 
participating in the eastward advance of our prairies and there- 
fore being “accompanied or followed by some specific evolution.” 
“These plants," he added, ‘are now confined to the eastern arm 
of the Prairie Province but in each case have their nearest re- 
lated species much farther west." 

Suspecting that the Virginia plant might be distinct, I visited 
the Peters Mountain locality at Narrows, Virginia, in August, 
1945, but was unable to locate a single specimen. Later, through 
the very kind aid of Mr. Henry H. L. Smith, Principal of the 
Narrows High School, the friendly and generous cooperation of 
Mr. James Hubert Browning, one of his Senior students, was 
enlisted. Mr. Browning instituted a fresh search upon Peters 
Mountain and within a week’s time succeeded in finding, at an 
altitude of ‘900 feet above New River", i. e., at about 2400 feet 
above sea-level, two plants. Six leaves and an abundance of 
ripe fruits were sent me for record purposes. The habitat was 
described as at Narrows. 

Meanwhile, through the good offices of my warm personal 
friend, Dr. P. D. Strausbaugh, of West Virginia University, the 
further assistance of Dr. Earl L. Core of the same institution and 
of Dr. E. Meade McNeill of Concord College, Athens, West 
Virginia, was obtained. Indeed, it happened that Dr. Core, who 
was the actual discoverer of the Peters Mountain Iliamna in 
1927, had ascended Peters Mountain as recently as the month 
before my own fruitless ascent, but during the time at his disposal 
was unable to find any specimens. That Jliamna might still be 
found, however, was thoroughly believed by Dr. McNeill who, on 
taking a class up Peters Mountain some nine years earlier, had 
found a colony of more than a hundred plants. Professor 
MeNeill volunteered to revisit the mountain in an attempt to 
rediscover specimens. This he did with two companions shortly 
afterwards. Without much difficulty they finally came upon an 
area a few rods long, containing numerous individuals and small 


1946] Sherff,—Plants in the Gray’s Manual Range 95 


clumps or even colonies. Early in October, 1945, spurred on by 
the enthusiasm and the zealous efforts put forth by Professor 
MeNeill in my behalf, I visited Narrows once again and on Oct. 
5th, in his company, ascended Peters Mountain to the newly 
discovered site. Specimens were obtained for our gardens, also 
for distribution to herbaria (McNeill & Sherff no. 1). Our 
ascent was made at a point 0.9 mile down (7. e., north and north- 
west along) the highway (as measured, in the automobile, from 
the northeast end of the New River bridge at Narrows) along 
New River. We went up eastwardly and somewhat northwardly. 
Our plants grew at the very crest of the rocky ridge or shoulder 
of the mountain, the altitude being about 2500 feet (no speci- 
mens were found at 2000 feet, the altitude reported originally by 
Strausbaugh & Core)*. The habitat was of the open-woods 
type, with numerous shrubs and small or stunted trees such as 
Crataegus, and naturally permitted wind and light an easy access. 
'The soil in the various patches of earth, often mere pockets in 
the rock, was a very black humus and when wet (as it was at the 
time of our ascent) appeared identical with the muck often en- 
countered in swamp or marsh habitats of our prairie states. 
'The plants all were in or just past their late fruiting state. In 
stature, they were much smaller than in var. typica of the Kanka- 
kee habitat. Their height was mostly 6-9 dm., not mostly 1-1.7 
m. as in var. typica®. The leaves had blades seldom 1 dm. 
(and very rarely 1.25 dm.) broad or long, while in var. typica 
leaf-blades 1.5 dm. or more in length and breadth are common. 
Some specimens could be found with terminal leaf-lobe as de- 
scribed above for most leaves of var. typica, but the majority of 
leaves were as shown in Strausbaugh & Core's illustration (loc. 
cit., fig. 1) and had the terminal lobe oblong or subcuneately 
narrowed below and subtended with sharp sinuses (see pl. 1024).!° 


8 On my abortive trip in August I made extensive searches at levels of from 1900 to 
2300 feet. Numerous outcropping rocks were observed, as described by Strausbaugh 
and Core for the habitat, but seemingly the habitat had been altered since their time 
because of heavy forest-cutting and subsequent reforestation. This change was re- 
marked upon likewise by Core when he ascended in July. 

* Strausbaugh and Core in their ‘‘ description of the Virginia plant” gave the height 
as ''0.6-2 m. tall, or taller," but neither Dr. McNeill nor I could find any plants 
appreciably over a meter tall. 

10 The terminal lobe is especially apt to be widened at or slightly below the middle, 
much as in some material of Iliammna latibracteata Wiggins of the western United 
States. 


96 Rhodora [May 


A subsequent examination of additional herbarium specimens 
from Peters Mountain, collected there by Fogg and by Sharp 
(and very kindly lent me by Dr. J. M. Greenman, Curator of 
the Herbarium of the Missouri Botanical Garden), has been 
made but no constant differences have been found in flowers or 
fruits. It appears that the Peters Mountain plants (vide pl. 
1024) are specifically identical with the Kankakee plants (vide 
pl. 1023) but varietally distinct. I have named them after Dr. 
Earl L. Core who, as a member of the West Virginia University 
Botanical Expedition in 1927, discovered the first specimens: 


ILIAMNA REMOTA var. Corei, var. nov.—A varietate typica 
plantis humilioribus plerumque sub 1 m. altis foliis minoribus 
plerumque sub 1 dm. latis et lamina sub 1 dm. longis lobo termin- 
ah saepius oblongo vel infra subcuneate angustato et sinibus plus | 
minusve acribus subtento differt. 

SPECIMENS EXAMINED: James Hubert Browning, at 900 feet 
above New River, end of Peters Mt., in Narrows, Virginia, 
September, 1945 (two plants found; leaves and fruits in Chi. Nat. 
Hist. Mus.); John M. Fogg, Jr., 15047, alt. 3000 feet, on dry, 
wooded, rocky shoulder of Peters Mt., along New River north of 
Narrows, Jul. 17, 1938 (Mo. Bot. Gard.); McNeill & Sherff 1, in 
black humus or muck, among rocks at top of ridge, alt. about 
2500 feet, Peters Mt., Narrows, Oct. 5, 1945 (TYPE, Gray Herb.: 
ISOTYPES, Carnegie Mus., 2 sheets; Chi. Nat. Hist. Mus.; Cornell 
Univ.; Delessert Herb.; Gray Herb.; Kew Bot. Gard.; McNeill 
Herb. in Concord Coll.; Univ. Minnesota ; Mo. Bot. Gard.; N. Y. 
Bot. Gard., 2 sheets; Mus. Nat. Hist. Paris; Phila. Acad. Nat. 
Sci., 2 sheets; Stanford Univ., 2 sheets; U.S. Nat. Mus., 2 sheets; 
West Virginia Univ.); Aaron J. Sharp, on rocky exposure of 
Peters Mt., Narrows of New River, June 29, 1940 (Mo. Bot. 
Gard., 2 sheets); West Virginia University Bot. Exped., Peters 
Mt., The Narrows (Carn. Mus.; N. Y. Bot. Gard., where dated 
Jul. 20, 1929). 


VALERIANELLA CHENOPODIFOLIA (Pursh) DC.—Deam (FI. 
Indiana 890. 1940) cites definitely or positively a single locality 
in Indiana for this species, namely Studebaker’s woods, St. 
Joseph County, where it was collected by Nieuwland in 1912 and 
again in 1919. For the past five years I have noted specimens 
growing at certain spots in bottom-land or flood-plain woods 
along Trail Creek, just south of U. S. Highway 20 and perhaps 
two miles southeast of Michigan City. One stand, covering a 
quarter of an acre or more near the bridge at à small cross-road, 


Rhodora Plate 1024 


ILIAMNA REMOTA var. Corer from McNeill & Sherff 1, alt. about 2500 feet, Peters 
Mt. at Narrows, October 5, 1945 (photograph by author from rvprkE in Gray Herb.). 


1946] Sherff,—Plants in the Gray’s Manual Range 97 


was so dense as to resemble somewhat a field of buckwheat 
(Fagopyrum esculentum Moench) in flower. Numerous speci- 
mens (my no. 5020, May 13, 1945) have been distributed to the 
larger herbaria. 

MONARDA FISTULOSA var. TYPICA f. ALBESCENS Farw.—This 
white-flowered forma I have already noted elsewhere (Torreya 
45: 68. 1945) as occurring on a shady bank along South Broadway 
Road, near Hastings, Michigan, in the summer of 1944. Several 
dozen flowering stems were counted in the same spot, Aug. 5, 
1945. Nowhere was there evident an intergradation with the 
typical purplish- or lavender-flowered form that abounded near 
by. Specimens were collected (my no. 5022) for distribution to 
the larger herbaria. 

VERBASCUM PHLOoMoIDES L.—Deam (Fl. Indiana 834. 1940) 
notes the recent discovery (about 1925) of V. Phlomoides in 
Indiana, at a place near Burnettsville. He cites several addi- 
tional localities where he himself found it. I have kept no 
records but have observed it at several points along highways, in 
old pastures and near fences, in central and northwestern Indiana, 
in August, 1945, and north of Chicago Heights, Illinois, in Sep- 
tember, 1945. Pennell (Scroph. E. Temp. N. Amer., Acad. Nat. 
Sc. Phila. Monogr. 1: 173. 1935) gives Minnesota as the north- 
westernmost state for its range in the United States. In July 
of 1945, I chanced upon a locality in Michigan where it was the 
dominant or at least most conspicuous species for many acres of 
ground. This was an old field (D. McCallum farm) just north of 
the Pine Lake in the northwest quarter of Hope Township, 
Barry County. Specimens (my no. 5023) were collected on 
Aug. 6th for distribution to herbaria. Descriptions in manuals 
were read and compared with the actual specimens in the field. 
It became at once obvious that some authors had relied for 
measurements and other details all too much upon herbarium 
specimens. For Verbascum Phlomoides these are apt, in many 
respects, to be misleading." The larger plants do not make 
herbarium specimens readily, because of their gigantic leaves 


u Cf. Britton & Brown, lllustr. Fl. edit. 2. 3: 174. 1913. Here the flowers are said 
to be “usually in a solitary elongated spike-like raceme.” Yet in practically all of the 
larger plants there are several racemes arising from below the principal or central one. 
Indeed, it is because of the characteristic candelabrum effect thus produced that V. Phlo- 
moides can so often be told at a considerable distance from the ubiquitous V. Thapsus L. 


98 Rhodora [May 


and their large multiple inflorescences. Consequently the col- 
lector turns to the smaller, more tractable plants for making 
specimens. u 

From the living plants the following notes were taken: Larger 
plants + 2 meters tall and having the principal raceme 7.5-9.3 
dm. long; often 12 or more long, erect branches from at to a few 
em. below base of principal raceme and having a raceme + 0.5 as 
long as principal one. Principal lower leaves (excluding basal 
ones) on these larger plants often 4.5-5 dm. long. The acumi- 
nate tip of the median and upper leaves is usually a conspicuous 
feature. 

CENTAUREA MACULOSA Lam.—In 1908, Gray’s New Manual 
(edit. 7) gave (p. 861) New England to New Jersey as the range 
for C. maculosa in the United States. Deam more recently 
(Fl. Indiana 1104. 1940) gave its range as extending west to 
Michigan, Wisconsin, and Minnesota. For Indiana he was 
reluctant to admit the species as being well enough established 
to deserve listing. In adjacent southern Michigan, however, I 
have seen several small colonies during the past four years and 
these all seemed thoroughly established, especially so in Barry 
County and at a country spot near Howell. My no. 5025 was 
collected in quantity near a fence in a hay field, one half mile east 
of Leach Lake, northeast of Hastings, Aug. 8, 1945. 


CHICAGO TEACHERS COLLEGE and CurcAGo NATURAL 
History MUSEUM. 


BOTANICAL VISITS TO FORTS CLARK, MANDAN 
AND UNION IN NORTH DAKOTA* 


O. A. STEVENS 
Plates 1025 and 1026 


Through the enthusiastic assistance and guidance of my friend, 
Roy N. Bach, I was able to visit the site of Fort Mandan where 
Bradbury and Nuttall collected in 1810 and that of Fort Union 
where Audubon collected and illustrated birds and mammals in 
1843. Maximilian was also at Fort Union in 1833. His artist, 


* Contribution from the Department of Botany, North Dakota Agricultural Ex- 
periment Station, published with the permission of the Director. 


1946] Stevens,—Botanical Visits to Forts in North Dakota 99 


Bodmer, gave us an excellent illustration of Fort Union and of 
the Indians, fortunately just before they were decimated by 
small pox. 

Maximilian (1) gave us the best general account and Bodmer’s 
classic illustrations are the best of the Indians of that time, but 
botanical contributions of this trip were few. Special interest 
attaches to Audubon’s visit from the fact that he had engaged 
Isaac Sprague as assistant artist. Asa Gray soon discovered 
Sprague and for many years he was Gray’s illustrator. The 
beautiful figures (unlabeled) of Penstemon albidus, Echinacea 
angustifolia and Psoralea esculenta in Audubon’s birds (2) are un- 
doubtedly Sprague’s work. In other figures, one is more in 
doubt how much Audubon modified the work of his assistant. 
In another paper (3), I have discussed this further. 

On June 21, 1945, we reached the site of Fort Union on the 
bank of the Missouri, about thirty rods east of the Montana 
State line. A tall flag pole was erected by the Great Northern 
Railway in 1935. The North Dakota Historical Society pur- 
chased the site in 1938 and has recently constructed a stone 
marker. A few posts stand to mark the walls but a gravel pit of 
recent date came near to destroying the site before the State 
intervened. Much of the material of the original fort was 
removed and used at Fort Buford a few miles farther east. Here 
the powder house still stands and the site is preserved. 

I wanted to collect something from the fort-site. What would 
it be? Certainly not the too prevalent Conringia or Descurainia 
Sophia. I noted only one plant of Penstemon albidus and de- 
cided on a bit of Sphaeralcea coccinea, Collomia linearis and the 
latest arrival, Camelina microcarpa. 

A mile or two farther east we found a road leading into the 
hills where Sprague and Audubon botanized. Penstemon albidus 
was still rare. Psoralea esculenta was fairly common and Mr. 
Bach found one plant with 13 racemes. Echinacea was just 
showing flower buds. Astragalus pectinatus was in full bloom 
and A. bisulcatus well begun. It seems strange that Audubon 
would not have mentioned these for they are striking. A few 
miles farther north we saw really breath-taking views of A. 
bisulcatus—great clumps dotting the small flats just below the 
foot of the buttes. 


100 Rhodora [May 


The afternoon was hot and we did not feel very ambitious. 
A creeping plant attracted my attention as we started down the 
hillside. It was Phlox alyssifolia (No. 818), which I had seen in 
the garden of J. Clayton Russell at Beach, North Dakota. He 
had found it about 30 miles over in Montana but we had not yet 
collected it in North Dakota. Here it seemed to have flowered 
only sparingly. On some hills farther north it was in better 
condition, with faded flowers and finally one fresh one, 25 mm. 
wide. Dr. E. T. Wherry writes me that there is a specimen of it 
from Fort Buford in the Britton Herbarium at the New York Bo- 
tanical Garden, also one from “ Phinney, N. D.,” collected by C. 
Lockwood in 1898, in the Field Museum Herbarium. We have 
been unable to trace this locality. 

Just before leaving the first hill, I was astonished to come upon 
Astragalus Drummondii (No. 817), new to North Dakota. The 
single plant had been half eaten off. A few miles farther north, 
we found one more plant and still farther, many plants, about 
seven miles south of U. S. Highway No. 2 and about 15 miles 
west of Williston. It looks so much like the prevalent A. 
pectinatus that one could hardly stop to examine each clump, 
and this may be why it had not been detected earlier. 

On June 22, Audubon (4) had written (2: 52): “found a num- 
ber of wild roses in bloom, quite sweet scented, though single and 
of a very pale rose color." It was June 23, when crossing the 
river at Elbowoods, we came suddenly on a bank of the first 
roses, R. Woodsii. My impression at first was that they were 
quite uniform in color and distinctly paler than our eastern R. 
blanda, but later we noticed various hues, more like the familiar 
“arkansana.” 

On August 7, 1938, we visited the site of the Indian village at 
Fort Clark. Here, also, the botanizing was disappointing. Be- 
hind the village-site was as pure a stand of Bouteloua gracilis as 
one could wish to see. The lodge-rings and most of the village- 
site were covered with Agropyron Smithii and a few weeds, 
especially Lactuca scariola and L. pulchella. Between the circles 
was a heavy growth of Iva xanthifolia and Chenopodium. Audu- 
bon (4) wrote of the camp being overrun with Chenopodium 
album, but it was doubtless C. Berlandieri. I finally collected a 
single plant of Euphorbia glyptosperma, large enough for seven 


Plate 1025 


a 


Rhodor 


“(CIS SutNoo[) 
JO] IOMOT 38 SOYSNE, “UBM JOG spooa JO SOUT, yep 


9 OYTO WILY A `N “UOJULAR IVAU IAY oftuyp JO YHOU jsn[ ADWITIA VSLVAIH DIQ JO ƏS 


TFGI “Yo ^N fiog Ag oyoyd ny — 7DojuoDap dipsoydoyy aav y3 aoddn pur 
YQIM ‘SBUL o3po[ JV sot 


Plate 1026 


Rhodora 


“ydaq ysy pur suwy ([^N As9j1n02)) 'IPGI ‘peysdneg LULIS Aq OJOYG — "urvos [BOD 9jtu3t[ UIG v st 
a33nq jo do} paxmoq out[ x18([ — ^sd322]nut unuoborsy pu Diza4421]nz) ‘wyofibuo) visimajip ‘snuumpyjosfisyy se sjuv[d qons sey 
393jnq jo osuq W PYS ~pipsaydaysy ATJSOUL 3t JO 3J9[ 29 sqnays 'snuirp44 A[quqoad st 3914 IBU =—“YYBLI 3v spunour Avy peus uo 
pue ssvis ul Dijund() “oljsLiaqoBIBYO ALAA JIB PUNOABIIO] 3j9[ ut 932nq JO JOO] WOI] YSMMMO PUB SUOTFBULIOY — "urme(p uosLrmr) 
pesodoud jo puo 459M Ao[9q jsní puv usr[ JoNUB]Y HMOJ JO IHS VAOGR ‘AAT Linosstyy JO yuRq uo 933nq Au[o pououjop v 


1946] Stevens,—Botanical Visits to Forts in North Dakota 101 


specimens (No. 384). The site of the large Indian village just 
north of the Knife River was chiefly in a farmyard pasture. 
Here also, Iva xanthifolia was one of the most conspicuous 
. plants. It is strange how the lodge-circles still show, especially 
as seen from the air (PLATE 1025). 

Much confusion has attached to the location of these historic 
places, especially that of “Fort Mandan,” so often mentioned by 
Nuttall and Bradbury. In discussing Arnica fulgens, Maguire (5) 
has enlarged upon Pennell’s error (6) in assuming that the Fort 
Mandan of Nuttall was the same as that of Lewis and Clark. 
The latter structure was already partly destroyed when the 
party returned from the west coast and the site later was washed 
away by the river. A marker has been established by the North 
Dakota Historical Society on higher ground. This is on the 
east ("north" of the journals), bank about three miles north of 
the present railway station of Fort Clark, which is on the west 
side or eight miles south of the mouth (where it enters the 
flood-plain) of the Knife Hiver. 

The “Fort Mandan” of Nuttall and Bradbury was about an 
equal distance north of the Knife River on the west (‘‘south’’) 
side of the Missouri. It is more correctly known as Fort Manuel 
Lisa, a trading and administrative post operated by that noted 
character. Truax (7) has given the best discussion of its loca- 
tion. Some maps show a town named Mannhaven, which was à 
river grain-elevator there. The bluffs between this place and the 
Knife River are high and steep. Northward, the river swings to 
the west, leaving a wide “bottom.” The “fort”? was located in a 
sort of amphitheater formed by the confluence of several short, 
deep ‘‘coulees” or ravines. The immediate bluffs are of the 
“Bad Land" butte type, presenting bare clay walls on the most 
exposed places, but the north and more gentle slopes are well 
covered with vegetation. The present plan seems to be to locate 
the west end of a huge dam across the river about three miles 
above the site. This may destroy the natural appearance of the 
place and cause great changes in flora. 

Eriogonum flavum is quite a common plant on the hills. Æ. 
multiceps, described from Maximilian's collections, which is more 
closely restricted to eroded clay slopes, grew on the buttes at the 
site. E. pauciflorum is also attributed to the region, but so far, I 


102 Rhodora [May 


have failed to find it. Astragalus tenellus, which seems to inhabit 
gravelly clay, was abundant on one ridge, and Thermopsis 
rhombifolia grew nearby on the clay slope. Hedysarum boreale, 
which had been overlooked until recent years, grew on the slope 
which was covered with Juniperus horizontalis and herbaceous 
plants. Erigeron glabellus grew on the protected side at the foot 
of the buttes and part way up the slope. This appears to be the 
characteristic habitat for it. We did not observe var. asper along 
the river. Both were reported by Nuttall as flowering in August, 
but this must have been an error, or possibly based on late indi- 
viduals, for June is the normal time. 

I had hoped to make extensive topotype collections of the 
Upper Missouri River plants, but completion of the project is 
very dubious and it seems desirable to record the preceding notes 
and a few on some of the plants. On June 29, 1941, we tried to 
find the small stream where Bradbury had described (8) the lilies 
on June 24, 1810, forming a “‘scarlet stripe as far as the eye could 
see." If one were to ascend the river to this point and then 
climb the bluffs, he would be amazed to see the rugged hills:soon 
give way to rolling, fertile farm-land. After considerable wan- 
dering by car, we found only one coulee which seemed to fit 
Bradbury’s description. There were only a few lilies present, 
though the date was right for them. 

I have been especially interested in Lactuca ludoviciana. All 
descriptions state that it has yellow flowers, but my conclusion is 
that it never did have such. In my first years of botanizing in 
eastern Kansas, I had called the flowers blue. We were a little 
too early at Fort Mandan for this plant. In the lily coulee were 
a few plants with the first flowers just open. Fortunately we 
were at the right time of day, for I have watched them at Fargo 
and found them open only from about 9 to 11 A. M. The rays 
were pale lilac as I have found them elsewhere. They may be 
nearly white or bluish. In dried specimens colors are notoriously 
misleading. I have noted L. scariola appearing bluish when dry. 

Nuttall’s lack of localities and his vague references to plants 
which ranged ‘‘to the northern Andes" have led to many specu- 
lations as to how far he might have gone. It seems more probable 
that he did not go far enough to see even the bluffs of the Little 
Missouri. It is not likely that he often crossed the Missouri 


1946] Rouleau,—Populus balsamifera of Linnaeus 103 


Maguire (5) (p. 425) suggested that Nuttall might have reached 

the White Earth vicinity for Arnica fulgens. This is neither 

necessary nor probable. In traveling along the Missouri in 1945, 

we did find it most abundant in the high, open coulees in Williams 

County, but it occurs southward through North Dakota and 

locally eastward to the central part of the State. In 1943, we 

were astonished to find it near McCanna (No. 683) in Grand 

Forks County. There is a specimen in the Brenckle Herbarium 

collected in western LaMoure County in 1903. I did not succeed 

in finding it there in 1944, but it easily may have been exter- 

minated by breaking or pasturing of the prairie. Dr. Brenckle 

writes me that it disappeared soon after he collected it. 

(1) Maximilian’s Travels in North America. In Thwaites, Early Western 
Travels. Vols. 22-25. 

(2) Audubon, J. J. Birds of America, oct. ed., Vol. 7, 1844. 

(3) Stevens, O. A. Audubon’s Journey up the Missouri River, 1843. N. D. 
Hist. Quart. 10: 62-82. 1943. 

(4) Audubon, Maria R. Audubon and his Journals, 2 vols. 1898. 

(5) Maguire, B. A Monograph of the Genus Arnica. Brittonia 4: 386-510. 
1943. 

(6) Pennell, Francis W. ‘Travels and Scientific Collections of Thomas Nut- 
tall. Bartonia No. 18: 1-51. 1936. 

(7) Truax, A. L. Manuel Lisa and his North Dakota Trading Post. N. D. 


Hist. Quart. 2: 240-246. 1928. 
(8) Bradbury's Travels. In Thwaites, Early Western Travels, Vol. 5. 


NonrH DAKOTA AGRICULTURAL COLLEGE, Fargo. 


POPULUS BALSAMIFERA OF LINNAEUS NOT A 
NOMEN AMBIGUUM 


ERNEST ROULEAU 


The binomial Populus Tacamahacca Miller has been generally 
accepted for the northern Balsam Poplar since Farwell published 
his note (8) on the nomenclature of Populus balsamifera Linnaeus. 
Sargent (36), Rehder (84), Redman (82, 33), Davy (6), Cansdale 
et al. (3) and Mansfeld (24) have also discussed the question and 
decided in favor of Populus Tacamahacca Mill. instead of P. 
balsamifera L., rejecting the latter as a nomen ambiguum. The 
major argument stressed by these authors for its rejection was the 
inferred basing of the Linnaean species upon a Catesby collec- 
tion of Populus deltoides Bartram ex Marshall from Carolina. 
This opinion was not shared by House (12) and Valckenier- 


104 Rhodora [May 


Suringar (40). Houtzagers (14) has more recently re-examined 
the question but does not commit himself, accepting the names 
in use in recent manuals: Populus Tacamahacca for the Balsam 
Poplar and P. deltoides for the Cottonwood. 

Moss (30) and Schinz & Thellung (37) have even gone so far as 
to reduce Populus candicans Ait. to the synonymy of P. Tacama- 
hacca Mill. Farwell (8), following this conception, adopted the 
name Populus Tacamahacca var. lanceolata (Marshall) Farwell for 
our indigenous tree, so that the native Balsam Poplar has become 
a variety of a tree known in cultivation only. These authors 
have wrongly ascribed Populus candicans to the synonymy of 
P. Tacamahacca. Miller's original description (26) and figure 
(28) describe and illustrate staminate flowers: “The katkins are 
like those of the black Poplar, but the number of stamina in the 
male flowers is uncertain, from eighteen to twenty-two. The 
female flowers I have not fully examined, but by the male kat- 
kins I have been induced to place it in this genus". It is a rec- 
ognized fact that Populus candicans Ait. is a cultivated tree, 
known only in the pistillate condition. 

While preparing monographic studies in the section Tacama- 
haca of the genus Populus for North America, the author has had 
occasion to study this question. 

Linnaeus' original description in Species Plantarum (16) was 
as follows: 
balsamifera 4. POPULUS foliis subcordatis oblongis crenatis. Hort. cliff. 

460. Roy. lugdb. 82. 
Populus foliis cordatis crenatis basi nudis, petiolis tere- 
tibus. Wach. ultr. 294. 
Populus nigra, folio maximo, gemmis balsamum odoratis- 
simum fundentibus. Catesb. car. I. p. 34. t. 34. 
Populus foliis ovatis acutis serratis. Gmel. sib. I. p. 152. 
Habitat in America septentrionali. h 

It is worthwhile examining in detail each of the references 

given by Linnaeus. 


1. Hort. cLIFF. 460 (15). 
The description is as follows: 


4. Populus foliis cordatis crenatis. 
Populus nigra, folio maximo, gemmis balsamum odoratissimum fun- 
dentibus. Catesb. ornith. 34, t. 34. 


1946] Rouleau,—Populus balsamifera of Linnaeus 105 


Crescit in Carolina Americes juxta aquas. Communicata ab IU. 
Boerhaavio. 

Cum etiamnum apud nos non floruerit, secum determinare nequeo. 

Tam similis est antecedenti ac umquam affinis, differt foliis magis 
cordatis, obtusis, foliisque balsamo obunctis; inter stipulas liqui- 
dissimum balsamum mazima in copia datur. 


Consequently, it is evident that Linnaeus' description was 
based upon a specimen communicated to him by Boerhaave. 
The only North American Poplar cultivated then in Holland was 
the Balsam Poplar, the ‘‘Populus similis: Arbor: resinosa: 
altera" of Boerhaave (2), based on Bauhin (1), the *"Tacamahaca 
foliis crenatis" of Hermann (10) and Plukenet (31). The 
descriptions of these pre-Linnaean authors refer to trees culti- 
vated in Holland, at Leyden mostly. The references given by 
these authors to Monardes (29) and Hernandez (11) are based on 
the analogy of the resinous gum of the buds of the Balsam Poplar 
with the gum of Bursera Simaruba (L.) Sarg. or some allied 
species, known in Mexico and Central America as “tacamahaca” 
or 'tacamaco". Many other plants received the name of 
“tacamahaca” for the same reason. 

It should be noted that Linnaeus in his Species Plantarum 
does not limit the distribution of Populus balsamifera to Carolina 
(apparently taken from Catesby) but extends it to “America 
septentrionali". 

Wein (43) gives an interesting discussion of the important 
question of the introduction of American trees into Europe. 

Whether there is a specimen of this species in the herbarium of 
Clifford is not known to the author, though there is a TYPE- 
SPECIMEN in the personal herbarium of Linnaeus, which seems to 
have been generally overlooked. It was there when he was 
writing his Species Plantarum and bears his annotation ‘‘4. 
balsamifera". It is of a short shoot consisting of six leaves with 
rounded bases, two of which have the base a little subcordate 
and asymmetrical. This type matches very easily numerous 
collections of Populus balsamifera made in Canada and the 
United States. One will find an illustration of the type and a 
letter of Jackson attesting the authenticity of the type-specimen 
in Houtzagers (14). The Catesby reference will be discussed 
later. 


106 Rhodora [May 


2. Roy. LUGDB. 82 (35). 


Royen only repeats the “specific name" found in Linnaeus’ 
Hortus Cliffortianus and in Catesby’s Natural History of Caro- 
lina. 


3. WacH. ULTR. 294 (41). 


4. Populus foliis cordatis, crenatis; basi nudis; petiolis teretibus. 
Populus foliis cordatis, crenatis. Linn. hort. 


This description can apply only to the Balsam Poplar, the 
“petiolis teretibus" and “basi nudis" leaving little doubt. 


4. CATESB. CAR. I, p. 34, t. 34 (4). 


Populus nigra, folio maximo, gemmis Balsamum odoratissimum fun- 
dentibus. 
The BraAck Porran of Carolina. 


Tuis Tree grows only near rivers, above the inhabited parts of Carolina. 
'They are large and very tall. In April, at which time only I saw them, 
they had dropt their seeds; which, by the remains, I could only perceive 
to hang in clusters, with a cotton-like consistence covering them. Upon 
the large swelling buds of this Tree sticks a very odoriferous balsam. 
The leaves are indented about the edges, and very broad, resembling in 
shape the black Poplar, described by Parkinson. 


Catesby's plate shows a vigorous shoot the leaves of which 
are truly cordate and toothed down to the junction of blade and 
petiole. The petioles, moreover, have at their junction with the 
blade a very strong rounded appearance. It may be safely 
assumed that it is a rather poor illustration of Populus hetero- 
phylla L. The unreliable characters of Catesby's plate enum- 
erated above were certainly sufficient to mislead Linnaeus as to 
its identity, and to make him think it represented a vigorous 
shoot of the Balsam Poplar. Specimens of Populus heterophylla 
in the Gray Herbarium match perfectly Catesby's plate (7. e. the 
long shoots). 

Catesby's specimen preserved in the British Museum has little 
to do with his illustration. It is a well developed short shoot of 
Populus deltoides (certainly not collected in April) with deltoid 
leaves and crenate teeth which begin at a comparatively long 
distance from the junction of blade and petiole (as is the case in 
the section Azgeiros). 

From Catesby’s description, supplemented by a more detailed 
one (5) wherein he states “Its leaves are large, smooth on one 


1946] Rouleau,—Populus balsamifera of Linnaeus 107 


side . . . The foot-stalks are long, remarkably flat, and of 
a reddish colour, as are the larger veins of the leaves . . . ”’, 
it is evident that the plate and the specimen in the British Mu- 
seum represent two different species. The plate represents 
Populus heterophylla and the description is a mixture of charac- 
ters of Populus heterophylla and P. deltoides. Linnaeus seems to 


have relied more on the plate than on the description. 


5. GMEL. SIB. I, P. 152, t. 33 (9). 


This reference to Gmelin is to a Poplar of section Tacamahaca 
which has been later segregated as Populus suaveolens Fischer. 


Through the ‘‘doctrine of residues", we must keep the name 
Populus balsamifera for the major element included by Linnaeus 
in his description, 7. e. the common American Balsam Poplar of 
which there is a type specimen in the Linnaean herbarium. This 
name (excl. syn. Catesby and Gmelin) should be reinstated for 
the tree which for the last quarter of a century has been passing 
as Populus Tacamahacca Miller. 

Supplementary references or notes given by Linnaeus in litera- 
ture subsequent to the Species Plantarum make even clearer 
what tree he was describing. 

In his Systema Naturae, eds. 10 and 11 (19, 20), the descrip- 
tion is as follows: 
balsamifera 4. P. fol. ovatis crenatis. Duham. arb. 2. t. 182. f. 6. Trew. 

ehret. t. 46. 

Linnaeus changes his ‘‘specific name". The adjective ovatis 
is more in harmony with the short-shoot leaves of Populus 
balsamifera. The reference to Duhamel’s figure (7) is to that of 
a leaf of the American Balsam Poplar. Similarly, the reference 
to Trew (39) leads to a handsome colored plate of the American 
Balsam Poplar. 

Again in his Systema Naturae, eds. 12 and 13 (21, 22), Linnaeus 
gives the following description: 


balsamifera 4. P. fol. ovatis serratis subtus albidis, stipulis resinosis. 
Populus foliis subcordatis inferne incanis, 
superne atroviridibus. Mill. dict. 7. Tacamahaca foliis 
crenatis. Pluk. alm. 360. t. 281. f. 2. Mill. dict. app. 
Trew Ehret. t. 46. Folia ovato-oblonga, subtus alba super- 
ficie vix conspicue tomentosa venis nudis reticulata. 


108 Rhodora [May 


It is very interesting to note that here Linnaeus himself 
cites Miller (25). In the eighth edition of his Gardener’s Dic- 
tionary (where binomial nomenclature was adopted, at least in 
part), Miller repeats the same diagnosis for Populus Tacama- 
hacca as the one given in the seventh edition (i. e. the reference 
given by Linnaeus). His more detailed English description 
differs only in phrasing from that given in the earlier edition. 
The other references have already been discussed. 

Finally, in Species Plantarum eds. 2 and 3 (17, 18), we read: 


balsamifera 4. POPULUS foliis subcordatis denticulatis. Hort. 

cliff. 460. Roy. lugdb. 82. 

Populus foliis cordatis crenatis basi nudis, petiolis 
teretibus. Wach. ultr. 294. 

Populus nigra, folio maximo gemmis balsamum odo- 
ratissimum fundentibus. Catesb. car. I. p. 34. t. 34. 
Duham. arb. 2. p. 178. t. 38. f. 6. 

Populus foliis ovatis acutis serratis. Gmel. sib. I. p. 
152. t. 33. 

Habitat in America septentrionali. h 


The “specific name" is again changed. The synonyms given 
are the same as those listed in Species Plantarum (ed. 1). The 
only addition is a reference to Duhamel which follows the one to 
Catesby. This indicates very well that Linnaeus believed 
Catesby’s plate to represent the Balsam Poplar. Duhamel’s de- 
scription (7) leaves absolutely no doubt: 

. mais il n'y en a point qui en répande autant, & d'une aussi agréable 
odeur, que celui de l'espece à feuilles ovales, no. 6, qu'on nomme pour 
cette raison Baumier. 

Je n'en ai jamais vu de grand, ses feuilles sont ovales, plus larges du 


cóté de la queue qu'à l'extrémité, terminées en pointe, dentelées finement 
par les bords, vertes en dessus, d'un blanc un peu jaunâtre par dessous. 


More evidence can be found in other authors. Thomas 
Martyn (27), who re-edited the Gardener's Dictionary of Miller, 
adds Populus Tacamahacca Mill. to the synonymy of Populus 
balsamifera L. for the following reason: “Mr. Miller's figure and 
description agrees with the American Tacamahaca as it appears 
in our gardens". Miller's plate (28), also reproduced in the ninth 
edition (27) of the Gardener's Dictionary, is a better-than-average 
illustration of the American Balsam Poplar. Willdenow (44) 
adds two post-Linnaean references: first to Wangenheim (42), 
whose illustration represents a short-shoot leaf of Populus bal- 


1946] Stevens,—Botanical Visits to Forts in North Dakota 109 


samifera; second, to Houttuyn (13), who describes the North 
American Balsam Poplar. Other authors of this period (such as 
Muenchhausen, DuRoi, Burgsdorff and others) only repeat the 
references given by Linnaeus (including Catesby and Gmelin) 
in his works published after Species Plantarum (ed. 1). 

From the above notes one can see that the citation by recent 
authors of Populus balsamifera Muenchhausen, DuRoi or Auct., 
not Linnaeus, as identical with Populus Tacamahacca Miller, is 
in reality Populus balsamifera Linnaeus sensu Linnaeus down to 
Farwell, Sargent, Rehder, etc., who rejected it as a nomen 
ambiguum. The assertion by only a few botanists that a Lin- 
naean name is an ambiguous name or that it has been variously 
applied is not a sufficient reason for rejecting it. 

~The name Populus balsamifera L. (excl. syn. Catesby and 
Gmelin) which was correctly accepted and understood by most 
dendrologists (Loudon, Spach, Wesmael, Koch, Schneider, Henry, 
etc.) must be kept for the American Balsam Poplar. 

Encouraging help and advice in the preparation of this article 
have been given by Messrs. Fernald and Weatherby of the Gray 
Herbarium, to whom the author expresses his heartiest thanks. 
A helpful background has also been acquired through the reading 
of Svenson's “On the descriptive method of Linnaeus" (38). 


BIBLIOGRAPHY 


(1) Bauhin, C., Pinax Theatri Botanici, etc., p. 430. 1623. 

(2) Boerhaave, H., Index alter Plantarum, quae in Horto academico Lug- 
duno-batavo aluntur. Vol. 2, p.211. 1727. 

(3) Cansdale, G. S., Beak, P. G., and Day, W. R., The Black Poplars and 
their hybrids cultivated in Britain. Imp. For. Inst. Univ. Oxford, 
pp. 17-18. 1938. 

(4) Catesby, M., The Natural History of Carolina, Florida and the Bahama 
Islands, etc., p. 34, t. 34. 1731. (second edition revised by Edwards 
in 1754, third edition revised by Edwards in 1771). 

(5) Catesby, M., Hortus britano-americanus, or a curious collection of trees 
and shrubs, etc., p. 32, t. 61. 1763. (reprinted in 1767 under a differ- 
ent title, but only slightly differing from the original edition). 

(6) Davy, J. B., Two suggested Nomina ambigua. Forestry, 10: 166-1068. 
1936. 

(7) Duhamel du Monceau, H. L., Traité des arbres et arbustes qui se cultivent 
en France en pleine terre. Vol. 2, pp. 178-182, pl. 38, fig. 6. 1755. 

(8) Farwell, O. A., Necessary changes in Botanical Nomenclature. RHo- 

DORA, 21: 101-103. 1919. 
(9) Gmelin, J. G., Flora sibirica, sive historia plantarum Sibiriae. Vol. 1, 

p. 152, t.33. 1747. 

(10) Hermann, P., Paradisi Batavi Prodromus sive Plantarum exoticarum in 
Batavorum Hortis observatarum index. p. 379. 1689. 

(11) Hernandez, F., Rerum medicarum Novae Hispaniae thesaurus seu 
plantarum, animalium et mineralium mexicanorum Historia, etc. p. 55. 
1651. 


E 


110 Rhodora [May 


(12) House, H. D., Annotated List of the Ferns and Flowering Plants of New 
York State. N. Y. State Mus. Bull. 254: 258-260. 1924. 

(13) Houttuyn, M. (Christmann, G. F. in), Des Ritters Carl von Linné voll- 
ständiges Pflanzensystem, etc., vol. 2, pp. 451—453. 1777. 

(14) Houtzagers, G., Het geslacht Populus in vervand met zijn beteekenis 
voor de Houtteelt. pp. 61—69, 226-227. 1937. 

(15) Linnaeus, C., Hortus Cliffortianus, etc. p. 460. 1737. 

(16) Linnaeus, C., Species Plantarum, etc. (ed. 1), p. 1034. 1753. 

(17) Linnaeus, C., Species Plantarum, etc. (ed. 2), p. 1464. 1763. 

(18) Linnaeus, C., Species Plantarum, etc. (ed. 3), p. 1464. 1763. 

(19) Linnaeus, C., Systema Naturae, etc. (ed. 10), vol. 2, p. 1294. 1759. 

(20) Linnaeus, C., Systema Naturae, etc. (ed. 11), vol. 2, p. 1294. 1760. 

(21) Linnaeus, C., Systema Naturae, etc. (ed. 12), vol. 2, p. 656. 1767. 

(22) Linnaeus, C., Systema Naturae, etc. (ed. 13), vol. 2, p. 656. 1770. 

(23) Linnaeus, C., Mantissa Plantarum Altera, Generum, etc., p. 499. 1771. 

(24) Mansfeld, R., Zur Nomenklatur der Farn- und Blütenpflanzen Deutsch- 
lands. III. Fedde Rep. Spec. Nov. 46: 59-64. 1939. 

(25) Miller, P., The Gardener's Dictionary, etc. (ed. 7), no. 7. 1759. 

(26) Miller, P., The Gardener's Dictionary, etc. (ed. 8), no. 6. 1768. 

(27) Miller, P., The Gardener's Dictionary, etc. (ed. 9), no. 5, t. 261. 1797. 
(edited by Thomas Martyn). 

(28) Miller, P., Figures of the most Beautiful, Useful and Uncommon Plants 
described in the Gardener's Dictionary, etc., pp. 174-175, t. 261. 1760. 

(29) Monardes, N., Simplicium medicamentorum ex Novo orbe delatorum, 
quorum in medicina usus est, historia, etc. pp. 5-6. 1579 (Latin 
translation of an earlier work in Spanish language published in 1569). 

(30) Moss, C. E., The Cambridge British Flora, vol. 2, p. 13. 1914. 

(31) Plukenet, L., Opera omnia botanica in sex tomos divisa. IV. Almages- 
tum botanicum, etc., p. 360. 1696. 

(32) Redman, K., Nomenclature confusion of Populus candicans Ait. Journ. 
Am. Pharm. Ass., 31: 140-141. 1942. 

(33) Redman, K., Nomenclature confusion in the case of the Balsam Poplar 
or Tacamahac. Journ. Am. Pharm. Ass., 31: 220-223. 1942. 

(34) Rehder, A., Proposed amendments to the International Rules of Botani- 
cal Nomenclature. Journ. Arn. Arb., 10: 55. 1929. 

(35) Royen, A. van, Florae Leydensis Prodromus, etc. p. 82. 1740. 

(36) Sargent, C. S., Notes on North American Trees. V. Journ. Arn. Arb., 
1:61-65. 1919. 

(37) Schinz, H. und Thellung, A., Weitere Beitrüge zur Nomenclatur der 
Schweizerflora (V). (Beitrüge zur Kenntniss der Schweizerflora (XV). 
Viert. Naturf. Gesell. Zürich, 60: 349. 1915. 

(38) Svenson, H. K., On the descriptive Method of Linnaeus. RHODORA, 47: 
273-302; 363-388, pl. 990-991. 1945. 

(39) Trew, D. C. J., Plantae selectae quarum imagines ad exemplaria naturalia 
Londini in Hortis Curiosorum nutrita manu artificiosa doctaque pinxit 
Georgeius Dionysius Ehret, etc. p. 15, t. 46. 1756. 

(40) Valekenier-Suringar, J., Die Anwendung der internationalen Nomenkla- 
tur-regeln. II.—Populus balsamifera, tacamahaca, candicans und 
deltoides. Eine kreuzweise Anderung von Namen. Mitt. Deut. 
Dendr. Ges., 41: 29-33. 1929. 

(41) Wachendorff, E. J. van, Horti Ultrajectini Index. p. 294. 1747. 

(42) Wangenheim, F. A. J. von, Beitrag zur deutschen holzgerechten Forst- 
wissenchaft die Anpflanzung Nordamericanischer Holzarten, etc., p. 
85, pl. 28, fig. 59, 1787. 

(43) Wein, K., Die erste Einführung nordamerikanischer Gehólze in Europa. 
II. Mitt. Deut. Dendr. Gesell., 43: 119-138. 1931. 

(44) Willdenow, C. L., Species Plantarum, etc., vol. 4, pp. 805-806. 1805. 


INSTITUT BOTANIQUE, UNIVERSITÉ DE MONTRÉAL. 


1946] Stevens,—Astragalus caryocarpus Fruits 111 


SizE, SHAPE AND NUMBER OF ASTRAGALUS CARYOCARPUS 
Fruits.—This species seemed to bloom in unusual profusion in 
the spring of 1945 and I noted much variation in color of flowers, 
from dark purple to pinkish and bluish hues. Fruits were pro- 
duced in abundance and their apparent variation in shape sug- 
gested a study of them. Collections were made at two places in 
northern Richland County, North Dakota, July 7, and in Clay 
County, Minnesota, July 13. From July 18 to 21, collections 
were made in Morton and Slope Counties, North Dakota. In 
each case, 10 or more fruits were taken from 15 to 20 individual 
plants and carefully measured while fresh. The following table 


summarizes the results. 
Average ratio, 


No. of Length length-width 
Place plants Aver. Max. Min. Aver. Max. Min. 
Leonard, N. D., 1 18 2.0 2.2 1.8 1.01 1.09 .91 
Leonard, N. D.,2 24 2.0 2.3 1.5 1.05 1.15 .97 
Muskoda, Minn. 15 2.0 2.2 1.9 1.01 1.09 . 96 
Mandan, N. D. 22 2.0 2.2 1.9 1.00 1.06 . 96 
Amidon, N. D. 15 1.8 2.0 E .89 .91 .85 


'The specimens from Slope County seem to run a trifle shorter 
than the others but the difference is scarcely significant. The 
general trend is clear and the variation less than I had expected. 
'The fruits are just about 2 em. long, varying only from slightly 
longer than wide to the reverse. Thickness was measured for 
only a few and it seemed rather uniform, about three-fourths of 
the width. 

Gray’s Manual describes the fruits as ‘‘ ovoid-globular, more or 
less pointed." This is certainly not true of our material. The 
usual outline is quadrate, occasionally slightly obovoid. Some- 
times there is a distinct mucronation from the style-base, but 
more often only a faint point. 

A large plant on a roadside cut in Emmons County, had an 
expanse of fully a meter and bore 163 fruits. Smaller plants 
found later were even more prolifie. We wondered whether size 
of fruit might be related to number, but counts indicated this 
was not true. The usual number per main stem was only | to 5, 
but occasionally reached 15 or 20 on an individual stem. ‘Three 
plants in Slope County bore 98, 110 and 124 fruits respectively, 
the last a plant with an expanse of about 3 dm.—O. A. STEVENS, 
North Dakota Agricultural College, Fargo, North Dakota. 


112 Rhodora [May 


SPIRAEA LATIFOLIA VAR. SEPTENTRIONALIS IN VIRGINIA.—On 
September 9, 1945, in company with Bernice M. Speese and 
Earlene Atchison, the writer collected Spiraea latifolia (Ait.) 
Borkh., var. septentrionalis Fernald on the very top of Hawksbill 
Mountain (elevation of 4049 feet), Page County, Virginia. 
There a small but dense patch grew in soil among the rocks. 
Some of the plants were in flower; others were in fruit. 

Specimens are in the Gray Herbarium (Baldwin 5464). They 
were identified by Prof. M. L. Fernald, who wrote in a letter of 
October 16, 1945: “This variety occurs on the Labrador Peninsula 
and in Newfoundland and south to the Magdalen Islands and 
subalpine and alpine regions of Mt. Katahdin, Maine, and the 
White Mountains, New Hampshire, also on Keweenaw Penin- 
sula, Michigan. It is distinguished from true S. latifolia by the 
dense cylindric to ovoid panicles, without elongate lower 
branches, and the relatively large flowers. It is a nice addition 
to the mountain-flora of northern Virginia."—J. T. BALDWIN, JR., 
The Blandy Experimental Farm, Boyce, Virginia. 


HELIANTHUS—A CorRECTION.—In a recent discussion of the 
confusion which has prevailed regarding certain species of 
Helianthus I became acutely infected by the germ of confusion 
and incorrectly cited, on page 79 of the April number of RHODORA, 
H. rigidus, forma flavus in the synonymy of H. laetiflorus, var. 
rigidus. H. rigidus, forma flavus = typical H. laetiflorus (with 
yellow disk).—M. L. FERNALD. 


Volume 48, no. 568, containing pages 65—88 and plates 1011—1020, was issued 
8 April, 1946. 


JUN 8 1940 


Rhodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. June, 1946. No. 570. 


CONTENTS: 


Notes on Aquatic and Prairie Vegetation in Southwestern 
Minnesota. R. M. Tryon, Jr. and J. W. Moore. .......... 113 


Notes on Compositae of the Northeastern United States, III. 
Inuleae and Senecioneae. Arthur Cronquist. ............. 116 


Amelanchier spicata not an American Species. M. L. Fernald. 125 


A Monograph of Amelanchier (Review). M.L.F............ 129 
Varieties of Lycopodium inundatum. M. L. Fernald. ......... 134 
New Combination in Chrysobalanus. Lyman B. Smith. ....... 136 


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TRbooora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. June, 1946. No. 570. 


NOTES ON AQUATIC AND PRAIRIE VEGETATION 
IN SOUTHWESTERN MINNESOTA! 


R. M. Tryon, Jr. AND J. W. Moore 


During the course of plant collecting in Minnesota in 1945?, 
it occurred to us that a trip to the southwestern part of the state 
might yield some interesting plant records. Early in the season 
some plants of western distribution were found in temporary 
rock pools on Sioux Quartzite in Cottonwood County. Knowing 
of the presence of similar conditions in Rock County we traveled 
down to Luverne with high expectations of making some worth- 
while discoveries. 

A few miles north of Luverne in Mound Township the Sioux 
Quartzite forms a series of gently rounded hills higher than the 
surrounding countryside. The southern extremity of the rock 
is marked by a sheer rock wall. On the top the glacial drift is 
very thin. The rock is close to the surface and exposed in many 
places. Numerous pools were present in the early summer 
between the low rock ridges and in shallow depressions. They 
occur dispersed in a typical rocky prairie habitat. Later in the 
summer these pools usually become dry. 

Probably the unusually wet spring and early summer were 
responsible: for the rich aquatic flora encountered. Dormant 
seeds had perhaps germinated for the first time in several years, 
since the area had been visited several times in the past and 

1 Contributions from the Herbarium of the University of Minnesota I. 


? The field work was supported in part by a grant in aid of research from the Gradu- 
ate School of the University of Minnesota. 


114 Rhodora [JUNE 


many of the plants present this year were not represented in 
the collections previously made. 

The distribution of species in Minnesota is based on specimens 
in the Herbarium of the University of Minnesota and published 
reports. Distribution outside of the state is based on current 
manuals and special treatments. All specimens cited are in the 
Herbarium of the University of Minnesota. Unless otherwise 
indicated collections were made by the authors on July 8, 1945, 
in Rock County, Minnesota. The following list includes five 
species and two forms reported for the state for the first time and 
seven species that are very rare in the state. 

Around a fairly large rock pool about three miles north of 
Luverne the following plants were found growing: 


ĪSOETES MELANOPODA Gay ex Dur., 17557, in mud; 17559, in 
2-4 inches of water at a similar pool about a quarter mile south- 
west of the large pool. These collections represent the first 
evidence of the occurrence of this species in Minnesota. 

SCHEDONNARDUS PANICULATUS (Nutt.) Trel., 17556. This 
grass grew in the prairie at the edge of the rock pool. A single 
collection of the species is represented in the University of Minne- 
sota Herbarium: Pipestone, Pipestone County, July, 1895, M. 
Menzel. Schedonnardus was reported for Minnesota by Upham! 
as Schedonnardus texanus Steud. (Lepturus paniculatus Nutt.) as 
occurring on “Rocky hills, Mound township, Rock county, 
Leiberg.” The nearest known occurrence for this species is 
in South Dakota. 

HETERANTHERA LIMOSA (Sw.) Willd., 17554. This blue- 
flowered species is another aquatic which has not previously been 
reported for Minnesota. The species has been reported from 
South Dakota. 

ELATINE TRIANDRA Schkuhr f. INTERMEDIA Seubert, 17551. 
This form, new to Minnesota, is the aquatic phase which grows 
in shallow water. It has been reported from Wisconsin and 
South Dakota. The species, Elatine triandra, was reported by 
Sheldon? as having been collected in pools near Cannon river, 
Burnside Township, Goodhue County, Minnesota, by A. P. 
Anderson, August, 1893. We have not been able to locate 
specimens of this, but is it likely that they represent f. intermedia 
as they grew in shallow pools. 

CALLITRICHE HETEROPHYLLA Pursh, 17552. Although re- 
ported for Minnesota by Muenscher?, this species was not pre- 

1 Upham, Warren, Catalogue of the Flora of Minnesota, p. 169, 1884. 


? Sheldon, E. P., Minnesota Botanical Studies 1: 16, 1894. 
3 Muenscher, W. C., Aquatic Plants of the United States, p. 262, 1944. 


1946] Tryon and Moore,—Aquatic and Prairie Vegetation 115 


viously represented from the state in the Herbarium. An addi- 
tional collection with mature fruit was made later in the summer 
on October 19, 1945, J. W. Moore & N. L. Huff, 18431, in an 
intermittent creek at Mound Spring State Park, a short distance 
northeast of the rock pool under discussion. 

LIMOSELLA AQUATICA L., 17548. One previous collection had 
been made in Minnesota: Pipestone, Pipestone County, June, 
1895, M. Menzel. 

PLAGIOBOTRYS SCOPULORUM (Greene) I. M. Johnston, 17553. 
The nearest known locality for this species is western South 
Dakota. Prior to this year it was not known to occur in Minne- 
sota. Earlier in the season very young plants were obtained 
from a rock pool on a Sioux Quartzite outcrop in Delton Town- 
ship, Cottonwood County, Minnesota, May 29, 1945, J. W. 
Moore & J. W. Posz, 16935. 

HORDEUM PUSILLUM Nutt., 17550. This species is of rare 
occurrence in Minnesota. Upham! first reported the species 
from Blue Earth County as collected by Leiberg. Other collec- 
tions from Minnesota are: Pipestone, Pipestone County, June, 
1895, Menzel and May, 1932, Fellows; Brown County, July, 1938, 
Rosendahl, 6926. 


The following species were associated with those listed above: 
Koeleria cristata (L.) Pers., 17561; Carex Eleocharis L. H. Bailey, 
17564; Juncus Dudleyi Wiegand, 17562; Allium canadense L., 
17566; Delphinium virescens Nutt., 17560; Potentilla pennsylvani- 
ca L. var. strigosa Pursh, 17565; Linum sulcatum Riddell, 17549; 
Gratiola neglecta Torr., 17555; Verbena simplex Lehm., 1755€. 

In a prairie pasture about four miles north-northwest of Lu- 
verne rock pool plants were again encountered; some were the 
same as those previously collected and some were additional 
ones. The water in some of these pools had evaporated leaving 
a moist layer of mud. Other pools still had an inch or two of 
water in them. 


ManSILEA vESTITA Hook. and Grev., 17578. This was grow- 
ing in some abundance at the margin of a pool. The only pre- 
vious collection from Minnesota is: Pipestone, Pipestone County, 
September 15, 1938, J. W. & M. F. Moore, 10550. 

IsoETES MELANOPODA Gay ex Dur., 17582. 

MYOSURUS MINIMUS L., 17586. Previous collections are: Big 
Stone County, June, 1901, Holzinger & Anderson; Chippewa 
County, May, 1909, Moyer; Pipestone County, May, !935, 
Rosendahl et al., 3052. 


1Upham, Warren. Catalogue of the Flora of Minnesota, p. 169, 1884. 


116 Rhodora [JUNE 


TILLAEA AQUATICA L., 17584. This interesting little plant 
has escaped detection in Minnesota until now. According to 
the reported range for North America, our collection is à new 
record for both Minnesota and the Great Plains Region of the 
United States and Canada. 

ELATINE TRIANDRA Schkuhr f. TERRESTRIS Seubert, 17585. 
'The terrestrial form in drying mud is new to Minnesota. 

HYDRANTHELIUM ROTUNDIFOLIUM (Michx.) Pennell, 17579. 
This species is also known from Lac qui Parle County where it 
was collected by Moyer and from Lyon County where it was 
collected by N. L. Huff. 

LIMOSELLA AQUATICA L., 17581. 

PLANTAGO ELONGATA Pursh, 17587. We are reporting this 
species from Minnesota for the first time, although there is one 
previous specimen preserved in the Herbarium: Pipestone, 
Pipestone County, June 15, 1931, Fellows. 


The following species were collected at the same locality in the 
adjacent prairie: Stipa spartea Trin., 17574; Hordeum jubatum L., 
17583; Anemone canadensis L., 17570; Lepidium densiflorum 
Schrad., 17571; Rosa arkansana Porter, 17572; Scutellaria parvula 
Michx., 17573; Acerates lanuginosa (Nutt.) Dene., 17568; Ver- 
bena simplex Lehm., 17580; Erigeron strigosus Muhl. ex Willd., 
17569; Achillea lanulosa Nutt., 17577. 


Department of Botany, 
UNIVERSITY oF Minnesota, Minneapolis. 


NOTES ON THE COMPOSITAE OF THE 
NORTHEASTERN UNITED STATES 


III. INULEAE AND SENECIONEAE 
ARTHUR CRONQUIST 


My condensation of the Antennarias of the northeastern 
United States (RHoporA 47: 182-184. 1945) has been criticized 
at some length by Fernald (RHoporA 47: 221-235; 239-247. 
1945). At the root of the trouble in Antennaria is the problem 
of apomixis. Some workers, especially those in Europe, have 
contended that each apomict should be treated as a distinct 
species, since it is self-perpetuating, and can, at least theoretically, 
be differentiated from all other apomicts by morphological 
minutiae. There are three noteworthy objections to such a pro- 


1946] Cronquist,—Notes on Compositae of the U. S., III 117 


cedure. First, it represents a radical departure from the tradi- 
tional concept of a species, ill-defined though that concept may 
be. Second, even in apomictic groups, apomixis is not always 
obligatory. Occasional normal sexual reproduction may occur, 
or, in some part of the range, be quite common. Fernald has 
admitted this to be true in Antennaria. These sexually repro- 
ducing plants may transcend the variation of several apomictic 
races, as has been demonstrated in Crepis acuminata by Babcock 
and Stebbins. One is then faced with the choice between dis- 
tinguishing as separate species the normally variable offspring of 
the sexual plants, or using apomixis itself as a specific criterion. 
Few botanists would defend either procedure, in theory. Third, 
the number of apomictic races may be so great that in practice it 
becomes impossible to distinguish them clearly. In a hypotheti- 
cal case of completely obligatory apomixis, there is no doubt that 
apomict a and apomict z are readily distinguishable, even though 
they may occur in the same area. Apomicts d, g, 0, and s, as 
they are successively discovered and studied, are likewise dis- 
tinguishable, although the differences between them are not so 
great as between a and z. But when apomicts a, b, c, d, e, f, g, 
hu pulmone Du I SU row x wand 2 Wave, been: 
discovered and named, a wholly confluent series is formed. 
Having once started splitting, the worker must either continue 
to describe new species ad infinitum, or undo his previous work 
on the group. The latter rarely happens. It is apt to take a 
relative new-comer, with an unbiased viewpoint and no axes to 
grind, to plunge into things and restore some degree of sanity to 
the situation. 

The alternative to treating each apomict as a separate species 
is to group them, insofar as possible, into units comparable in 
taxonomic significance, variability, and clarity of definition, with 
the ordinary sexual species. The principal difficulty here is that 
the normal specific lines may be obscured by the persistence, as 
apomietie races, of the occasional interspecifie hybrids. It may 
then become necessary to admit more than the usual amount of 
intergradation between species, in order to obtain units otherwise 
comparable to the strictly sexual species. A recent treatment 
embodying this alternative approach is that of Crepis by Babcock 
and Stebbins. After the most intensive and thorough study 


118 Rhodora [JUNE 


that has yet been given to a large American genus, these authors 
concluded that the only practical systematic approach was to 
admit the existence of numerous more or less distinguishable 
apomicts within a single species, while also admitting an unusual 
amount of intergradation between species. 

In considering the Antennarias of the manual range, I was 
unable to distinguish consistently and with reasonable certainty 
the great mass of recent segregates. The characters, as shown 
by our specimens, are just not as constant as has been claimed. 
I therefore found it necessary to prepare a new and more con- 
servative treatment. Although monographic study, making use 
of the material from all the major herbaria (which is beyond the 
scope of any floristic treatment) might conceivably warrant 
certain adjustments in my treatment of varieties, I am convinced 
that the entities considered do properly fall into only three well- 
founded and reasonably distinct species. The concluding para- 
graph of Stebbins’ reply (RHopora 38: 367-369. 1936) to Fer- 
nald’s reduction of A. virginica and its variety argillicola to 
varietal status under A. neodioica is wholly pertinent: “I recog- 
nize that there is overlapping between A. virginica and A. neo- 
dioica in some characteristics, but since I am aware of an equal 
amount of overlapping between A. fallax and A. Parlinii (par- 
ticularly in the southern and western portions of their range), 
A. neglecta and A. petaloidea (chiefly in Wisconsin), A. petaloidea 
and A. neodioica (throughout the north central states), and A. 
Parlinii and A. Brainerdii (in central New York) I feel that, in 

he interests of consistency, the reduction of A. virginica to a 
variety calls for a similar reduction of A. Parlinii, A. petaloidea, 
A. Brainerdii, and probably other species now recognized in the 
floras of eastern North America." 

It is my opinion that the first opinion expressed by Fernald 
regarding the validity of A. virginica is correct, and that Stebbins’ 
subsequent comments are wholly justified. Stebbins’ indication 
of the particular areas in which intergradation is most noticeable 
is of particular significance. It cannot be too strongly empha- 
sized that apparently constant differences in one locality may be 
broken down completely in some area, if apomixis is involved. 
The plants which break down the differences between two other- 
wise well-characterized apomicts may be still other apomicts 


1946] Cronquist,—Notes on Compositae of the U. S., II 119 


(since potentially there is an apomict for every genotype) or they 
may be sexually reproducing plants showing the normal varia- 
bility involved in cross-breeding. In either case the result is 
the same: differences which appear constant in one restricted 
area may vanish in some other area. With apomictic groups, 
even more than with sexual groups, therefore, the whole range of 
variability of the species, throughout its whole geographic dis- 
tribution, should be considered before any program of segregation 
is initiated. 

There are, however, some valid criticisms in Dr. Fernald’s 
paper. I overlooked the existence on the Gaspé peninsula of 
two boreal and cordilleran species, A. alpina (L.) Gaertn. and 
A. umbrinella Rydb. These are the species which he reports as 
A. vexillifera and A. subviscosa, respectively. A. vexillifera Fern. 
is merely one of a host of recent segregates from A. alpina (L.) 
Gaertn. a highly variable circumpolar species. Doubtless 
several varieties should be recognized, but their taxonomy and 
nomenclature are yet far from clear. For the present I think it 
best to associate A. vexillifera Fern. with A. alpina (L.) Gaertn. 
var. canescens Lange. 

Antennaria subviscosa Fern. appears to me to be identical with 
the northern and western A. umbrinella Rydb. This species is 
interesting in that it shows the most ideal transition, sometimes 
even on the same head, between the contrasting phyllary-types 
of the A. alpina group and the A. dioica group (including A. 
neglecta, A. plantaginifolia, A. parvifolia, A. microphylla, and 
others). In the former the cellular structure of the scarious part 
of the phyllary is more or less evident under 25 diameters magni- 
fication, and the pigment (varying from dirty green to brownish 
or nearly black) is in the cell walls. In the latter the cells are 
laterally compressed so that their raised walls are closely parallel 
and give the phyllary-tip a finely striate appearance, and the 
pigment, if any (mostly whitish or pink), is diffuse. Most of 
the western specimens of A. umbrinella have glabrous achenes, 
but Hitchcock and Martin 5600, from Nye County, Nevada, 
recently so determined by Dr. S. F. Blake, has them papillate, as 
in A. subviscosa. Since apparently no other characters are as- 
sociated with it, papillosity of the achenes would seem to fail in 
this case as a specific criterion. 


120 Rhodora [JUNE 


Two collections from the same station in western Minnesota 
form the basis for Dr. Fernald’s inclusion of A. aprica Greene in 
the manual range. I concur in the identification of the speci- 
mens, which mark the eastern limit of the known range of the 
species. Unfortunately the name A. parvifolia Nutt. antedates 
A. aprica Greene and applies to the same species. This was 
pointed out to me recently by Dr. Blake, and I confirmed it by 
examining Nuttall’s isotype at Philadelphia. Of more than 30 
specimens of A. parvifolia examined for achaenial papillae, about 
half were seen to be papillate at 25 diameters magnification, a 
few which seemed glabrous at 25 diameters were seen to be 
sparsely and minutely papillate when examined at 50 diameters, 
and the rest were apparently glabrous even at 50 diameters. 
In some cases specimens of the same collection, looking very 


1 Dr. Fernald informs me that Nuttall's material of Antennaria parvifolia at the 
British Museum contains plants of A. rosea Greene and A. microphylla Rydb., as well 
as A. aprica Greene, and suggests a need for further clarification. (It is my present 
opinion that A. microphylla, A. rosea, and a number of other western segregates prop- 
erly constitute a single polymorphic species, but these names are here used in the more 
restricted sense.) In accordance with Article 52 of the Rules, it becomes necessary to 
settle upon one of these as the basic element of A. parvifolia, to which the name should 
be restricted. Both A. microphylla and A. rosea may be excluded on the basis of 
Nuttall's description. The leaves of A. microphylla rarely exceed 1 cm. in length, and 
are certainly not ''half or three-quarters of an inch long . . . [and] somewhat rhom- 
boidally spathulate’’. A. rosea, on the other hand, occasionally has leaves as large as 
those described by Nuttall, but may be excluded by its pink phyllaries. In the diag- 
nosis of the species Nuttall says that the involucral bracts are yellow. In the more 
general discussion he says in one place that they are yellow, and in another that they 
are purple. Obviously here the two color-ty pes are being included in the same species; 
just as obviously, the pink (or purple) type must be considered a subordinate element, 
since it is mentioned only in the general discussion and is deflnitely excluded by the 
preliminary diagnosis. It should further be noted that occasional otherwise repre- 
sentative specimens of A. aprica have a distinct pink cast to the phyllaries, but it is 
to be assumed that the part of the type-collection of A. parvifolia which Dr. Fernald 
refers to A. rosea is not of this nature. One further difficulty remains: both A. 
microphylla and A. aprica have white rather than yellow phyllaries. Either of these, 
particularly A. aprica, may develop a yellowish tint in drying, however, and it must 
be remembered that the drying equipment available to Nuttall was not up to present- 
day standards. Both A. microphylla and A. rosea reach the edge of their range in 
the vicinity of the type-locality of A. parvifolia (‘‘On the Black Hills and plains of the 
upper part of the Platte”), and are there uncommon. A. aprica, however, abounds in 
that region. Except as noted above, Nuttall's description and comments apply very 
well to A. aprica Greene; one of the two exceptions may be explained by inadequate 
means of drying specimens, and the other by the inclusion of a definitely subordinate 
element (A. rosea) with the more characteristic specimens. Certainly Nuttall's 
description and comments are more applicable as a whole to A. aprica than to either 
A. rosea or A. microphylla. It therefore seems plain that the name Antennaria 
parvifolia Nutt. should properly be applied to that part of Nuttall's type which has 
pale phyllaries and comparatively large leaves (these leaves, incidentally, being much 
smaller than those of the common eastern species). This part is conspecific with A. 
aprica Greene, which is therefore reduced to synonymy. 


1946] Cronquist,—Notes on Compositae of the U. S., III 121 


much alike superficially, varied from apparently glabrous to 
evidently papillate. Although papillosity of the achenes is 
doubtless valuable in some cases as a taxonomic character, it 
seems that Mr. Porsild and Dr. Fernald have overestimated its 
importance. Linnaeus’ maxim, ‘‘Characterem non constituere 
Genus, sed Genus characterem", is equally true of species. 

In conclusion, the species of Antennaria known to occur in 
northeastern United States and the portion of adjacent Canada 
south of the St. Lawrence River, as I understand them, may be 
keyed as follows: 


1. Involucral bracts brown or dirty blackish green throughout: 
leaves about 1.5-5 mm. wide; Gaspé, Que., and northward... A. alpina. 
1. Involucral bracts (at least the inner) white or whitish toward 
the tip; leaves often much larger. 
2. Upper surface of the leaves nearly or quite as densely hairy 
as the lower, glabrate, if at all, only in extreme age; 
Gaspé, Que., and northward; w. Minn. and westward. 
3. Pistillate involucres mostly 5-7 mm. high; terminal 
scarious portion of the outer involucral bracts more or 
less discolored and brownish; Gaspé, Que., and north- 
SW ATC eect cree ud p ue RA ds ioia ciega dites e l d A. umbrinella. 
3. Pistillate involucres mostly 8-11 mm. high; terminal 
scarious portion of the outer bracts, as well as the 
inner, white (or occasionally becoming yellowish in 
drying) and shining; w. Minn. and westward........ A. parvifolia. 
2. Upper surface of the basal leaves distinctly less pubescent 
than the lower, sooner or later glabrate; throughout our 
range (A. solitaria toward the south only). 
3. Basal leaves and those at the ends of the stolons rela- 
tively small, less than 1.5 cm. wide, 1-nerved or ob- 
Bolireiv-dsnerved 4$. 50. Wee Se d EIS UE EE ela A. neglecta. 
3. Basal leaves and those at the ends of the stolons rela- 
tively large, prominently 3-5-nerved, the larger ones 
1.5 em. wide or more. 
4 Heads: several deese cse dao te IN S A. plantaginifolia. 
4 wHeadsisolitary $3 90 tienen Geet uM c M qp A. solitaria. 


GNAPHALIUM PURPUREUM L. var. purpureum, var. nov. Gnaph- 
alium purpureum L. Sp. Pl. 854. 1753, sens. strict. 

Gnaphalium saxicola Fassett is as yet known from only a very 
few collections in Wisconsin, but is approached by occasional 
specimens from elsewhere in the range of G. obtusifolium. At 
least until a larger series of specimens demonstrates its morpho- 
logic and genetic discontinuity, it seems better treated as a 
variety of G. obtusifolium. 


GNAPHALIUM OBTUSIFOLIUM L. var. saxicola (Fassett), comb. 
nov. Gnaphalium saxicola Fassett, RHopora 33: 75. 1931. 


122 Rhodora [JUNE 


Erechtites megalocarpa Fern. was described from saline coastal 
marshes in Massachusetts. It has subsequently been reported 
from Rhode Island and Long Island, and specimens in the local 
herbarium of the New York Botanical Garden extend the known 
range to Ocean County, New Jersey. It differs from E. heracii- 
folia (L.) Raf. in its fleshiness, broader heads, and larger achenes 
with more numerous nerves, as well as some minor and inconstant 
tendencies or trends. The succulence is not preserved in the 
herbarium, and dried specimens of the two are superficially very 
similar. The several technical differences would at first seem to 
be of specific importance, but a study of our material shows no 
real discontinuity. In this connection, it is interesting to note 
that Millspaugh and Chase (Field Mus. Pub. Bot. 3: 146. 1904) 
describe the Yucatan plants of E. hieraciifolia as having achenes 
up to 3.8 mm. long, although Fernald, in noting the large achenes 
(4-5.5 mm.) of E. megalocarpa indicates that those of E. hieracii- 
folia are only 2-3 mm. long. It seems reasonable, therefore, to 
treat E. megalocarpa as merely à well-marked ecotype of saline 
coastal marshes. 


ERECHTITES HIERACIIFOLIA (L.) Raf. var. megalocarpa (Fern.), 
comb. nov. E. megalocarpa Fern. Rnopona 19: 24. 1917. 

In trying to determine the differences between Petasites 
palmatus (Ait.) Gray and P. vitifolius Greene I was led into a 
general consideration of the P. frigidus group, to which they 
belong. The principal differences which have been used for 
taxonomic segregation within the group are in the size, shape, 
and lobing of the leaves, but it soon became apparent that these 
differences, while striking, are far from constant. The differ- 
ences adduced by Rydberg in the relative lengths of the throat 
and teeth of the corollas of the perfect flowers are unstable and 
apparently wholly without taxonomic value. 

It is my opinion that only two closely related species should be 
recognized in this group, namely P. frigidus Fries and P. sagitta- 
tus (Banks) Gray. Both species are variable, rendering a simple 
and concise statement of their differences most difficult. The 
leaves of P. sagittatus vary from merely a little wavy and callous- 
denticulate (especially in smaller specimens) to more commonly 
conspicuously dentate with 20-35 teeth on each side. Those of 
P. frigidus, on the other hand, vary from coarsely toothed, with 


1946] Cronquist,—Notes on Compositae of the U. S., III 123 


5-15 teeth on each side (in smaller forms), to more or less dis- 
tinctly lobed (in larger forms), and then often with more numer- 
ous teeth. The venation, toothing, and lobing of P. frigidus is 
pinnipalmate in small forms, becoming progressively more strong- 
ly palmate in larger ones; in P. sagittatus the venation is pinni- 
palmate throughout, the leaves often becoming conspicuously 
longer than broad. These differences, while seemingly weak, are 
in practice sufficiently constant to leave very few if any doubtful 
specimens, and the two species can generally be distinguished at 
a glance. 

The appearance of P. sagittatus is sufficiently characteristic so 
that only one segregate, P. dentatus Blankinship, has been pro- 
posed. This was reduced by Rydberg in the North American 
Flora. The extremes of variation in P. frigidus, on the other 
hand, seem so different that nearly a dozen segregates have been 
proposed. ‘These rest almost entirely on foliar characters, how- 
ever, and are seen to be confluent when a large series of speci- 
mens is examined. The facts were seen in their proper perspec- 
tive as long ago as 1865, when Herder recognized two American 
and one European variety of P. frigidus, in addition to the cir- 
cumpolar typical one. His summary of the situation is worthy 
of quotation: “Diese hoch-nordische Pflanze zeigt eine grosse 
Verschiedenheit in der Configuration und Zahnung ihrer Blatter, 
so dass, wenn man nur einzelne Exemplare aus einer Gegend hat, 
man leicht in den Fall kommt, auf ihre oft eigenthümliche 
Blattgestaltung hin, Arten zu gründen." It remains only to 
transfer his names from the segregate genus Nardosmia to 
Petasites and refer the more recent names to their proper places. 


PETASITES FRIGIDUS (L.) Fries var. genuinus (Herder), comb. 
nov. Nardosmia frigida var. genuina Herder, Bull. Soc. Nat. 
Mosc. 1865: 372. Petasites gracilis Britton, Bull. N. Y. Bot. 
Gard. 2: 186. 1901. P. alaskanus Rydb. N. Am. Fl. 34 (4): 314. 
1927. P. Warrenii St. John, Res. Stud. State Coll. Wash. 1: 
109. 1929. Leaves relatively small, pinnipalmately veined, 
coarsely and irregularly toothed, but only scarcely or obscurely 
lobed. 

PETAsITES FRIGIDUS (L.) Fries var. corymbosus (R. Br.), 
comb. nov. Tussilago corymbosa R. Br.! App. Parry’s First Voy. 

1 Dr. Fernald in personal correspondence has raised a question as to the identity of 


Tussilago corymbosa R. Br. The origina] description is clear enough, noting that the 
leaf-lobes are equal to 14 to 14 of the “radius” of the leaf, and are themselves toothed, 


124 Rhodora [JUNE 


279. 1824. Nardosmia corymbosa Hook. Fl. Bor.-Am. 1: 307. 
1833. Nardosmia frigida var. corymbosa Herder, Bull. Soc. Nat. 
Mose. 1865: 372. Petasites trigonophyllus Greene, Leafl. 1: 180. 
1906. P. vitifolius Greene, loc. cit. P. nivalis Greene, Pitt. 2: 
18. 1889. P. corymbosus Rydb. Bull. Torrey Club 37: 460. 1910. 
P. hyperboreus Rydb. N. Am. Fl. 34 (4): 312. 1927. P. frigidus 
var. hyperboreoides Hulten, Fl. Aleutian Isl. 328. 1937. Leaves 
mostly pinnipalmate, distinctly lobed, the lobes seldom extending 
more than half way to the base. 

PETASITES FRIGIDUS (L.) Fries var. palmatus (Ait.), comb. nov. 
Tussilago palmata Ait. Hort. Kew. 3: 188. 1789. Nardosmia 
palmata Hook. Fl. Bor.-Am. 1: 308. 1833. Nardosmia Hookeri- 
ana Nutt. Trans. Am. Phil. Soc. II. 7: 288. 1841. Nardosmia 
frigida var. palmata Herder, Bull. Soc. Nat. Mose. 1865: 372. 
Nardosmia speciosa Nutt. Trans. Am. Phil. Soc. II. 7: 288. 1841. 
Petasites palmatus var. frigidus Macoun, Cat. Can. Pl. 1: 553. 
1886.  Petasites speciosus Piper, Mazama 2: 97. 1901.  Peta- 
sites speciosus var. frigidus Henry, Fl. S. Br. Columb. 311. 1915. 
P. Hookerianus Rydb. N. Am. Fl. 34 (4): 314. 1927. Leaves 
distinetly palmate, the lobes generally extending more than half 
way to the base. 


the teeth mucronulate. The leaf size given, 1 14-2 14 inches broad, is within the range 
of normal variation of the entity to which Hooker, Herder and I have successively 
attached the name, although many of the more southern specimens, in particular, are 
often much larger. A photograph at the Gray Herbarium of some fragments in the 
British Museum which are supposed to represent type material collected by Captain 
Sabine on Melville Island shows a plant similar in size, habit, and general characteris- 
tics to what has been assumed to be an isotype in the Gray Herbarium, the two being 
apparently of the same collection. Both of these differ from Brown's description, 
however, not only in the smaller leaves (barely 3 cm. wide in the Gray Herbarium 
specimen, apparently even narrower in the other), but also in having the leaves merely 
toothed, and seem to represent the typical phase of P. frigidus as understood by me. 
The flowering stalks do have the corymbiform inflorescence described by Brown (a 
character now generally conceded to be of no taxonomic value here), but in neither 
specimen is the accompanying basal leaf organically attached to the flowering stem. 
In the Flora Boreali-Americana Hooker distinguishes Nardosmia corymbosa (based on 
Tussilago corymbosa R. Br.) from N. frigida solely on its leaf-outline, and cites speci- 
mens by Parry, Sabine, and Richardson. Since Brown was describing plants collected 
on the Parry expedition as well as the Sabine expedition, the name may well have 
been founded partly or even wholly on the Parry collection. On the basis of Brown's 
clear and explicit description, in which he specifically contrasted the leaves of T. 
corymbosa with those of T. frigida, I find it difficult to believe that the specimen in the 
Gray Herbarium, or the specimen which Dr. Fernald has photographed in the British 
Museum is authentic material of T. corymbosa. There may have been an error in the 
labeling or the mounting, or the Sabine collection may have been mixed, since 
Petasites frigidus is certainly to be expected on Melville Island. In any case, the 
basal leaves accompanying the specimens mentioned at the Gray Herbarium and the 
British Museum can scarcely be considered to form part of the true type of Tussilago 
corymbosa R. Br. Under the circumstances, and especially since he evidently had 
authentic material at his disposal, I think it proper to continue Hooker's interpreta- 
tion of the name. 


1946] Fernald,—Amelanchier spicata 125 


As might be expected, the species increases progressively in 
size with amelioration of the habitat, so that var. genuinus is the 
most reduced, and the var. palmatus averages the largest. P. 
speciosus, of the Pacific coastal states, has the leaves a little less 
deeply lobed, on the average, than do the ordinary forms of the 
transcontinental P. palmatus, but the differences are slight and 
inconstant, and many specimens are quite indistinguishable. 

More collections are needed before the precise limits of range 
of each variety may be defined. In general, the var. genuinus is 
the most boreal of the three, and the only circumpolar one. It 
is known to extend south in the high mountains to Washington. 
The var. corymbosus is found through much of the American 
range of var. genuinus, but extends farther south, reaching 
Gaspé, Que., Mich., and Minn. The var. palmatus is the most 
southern of the three, extending as far as Mass., on the east 
coast, and Calif., on the west. 


New York BOTANICAL GARDEN 
New York, N. Y. 


AMELANCHIER SPICATA NOT AN AMERICAN SPECIES 
M. L. FERNALD 
(Plates 1027-1030) 


The name Amelanchier spicata (Lam.) K. Koch has dodged in 
and out of American treatments of the genus but was supposed 
to have been finally dismissed by Wiegand in his critical studies 
of the genus, when he wrote in Rnopona, xiv. 123 (1912): 


Those who have seen these specimens and the type specimen of 
Crataegus spicata [of Lamarck, basis of A. spicata], as well as specimens 
of our Eastern American stoloniferous fine-toothed Amelanchier agree 
that they all appear to be one and the same thing. Flowers and leaves 
in the exsiccatae match those of this stoloniferous species as well as one 
could wish, as do also the flowers and leaves in the original descriptions. 
The original description of C. spicata, however, gives the height of the 
plant as from two to three times that of the native European species of 
Amelanchier, which, figured out, would mean about 2-5 m. [6:/,-16?/,, 
feet]. Willdenow gives the height as 2-2.5 m. In addition, Mr. 
Alfred Rehder has stated to the writer that the plant in European 
gardens commonly passing under the name A. ovalis [A. ovalis sensu 
Borkh., equaling A. spicata, not A. ovalis Medicus, the native European 
species] is not low and stoloniferous but tall and fastigiate. 


126 Rhodora [JUNE 


Wiegand considered that Amelanchier spicata was “of hybrid 
origin” and, believing the European tradition that it came from 
Canada, he guessed that it might have been a hybrid of the tall 
and fastigiate or arborescent A. canadensis (L.) Medic. (A. ob- 
longifolia (T. & G.) Roemer) and the freely stoloniferous and 
colonial dwarf A. stolonifera Wiegand. The latter conjecture 
seems not well supported by the detailed characters of A. spicata, 
as shown by the photograph of the type taken by Professor 
Alfred Rehder, reproductions, X 1, of which are shown in our 
PLATE 1027, and the tall fastigiate habit sufficiently demonstrates 
that A. spicata has nothing, except generically, to do with the 
low and freely stoloniferous and loosely colonial shrubs of eastern 
North America. 

In spite of this conclusive evidence from those who have long 
known the importance of such characters in the genus, the name 
Amelanchier spicata (Lam.) K. Koch suddenly pops up again! as 
indubitably belonging to three low (“0.3-2 m. tall") native 
North American ‘‘surculose colonial shrubs", A. stolonifera and 
humilis Wiegand and A. austromontana Ashe. Since this inter- 
pretation by Professor George Neville Jones is so at variance 
with those of Wiegand and of Rehder, while the photograph of 
the type of Crataegus spicata and Lamarck's original account of 
it show complete departure in foliage, flowers, fruit and habit 
from the characters of these three American species, it becomes 
necessary to check the type and the description of C. spicata 
(our PLATE 1027) with some care. 

The aggregate Amelanchier spicata sensu G. N. Jones, not of 
anyone else, is defined, in part, as Low surculose colonial shrubs 
0.3-2 m. tall; . . . lateral veins 7-9 pairs, . . . distantly ar- 
ranged . .. ; ... sepals... usually recurved from the 
middle after anthesis”. Now, if we examine Professor Rehder’s 
photograph of the type of Crataegus spicata (portions shown in 
our PLATE 1027) it will at once be evident that the lateral veins 
do not stop at “9 pairs . . . distant", as defined by Jones, but 
exceed that maximum and, as compared with the really distant 
lateral veins of A. stolonifera (our PLATE 1029, rras. 3 and 4, and 
PLATE 1030, rra. 2, both from the TYPE) are relatively approxi- 


1 In G. N. Jones, Am. Species of Amelanchier, Ill. Biol. Mon. xx. no. 2: 51 (1946). 
See review in this journal, pp. 129-134. 


1946] Fernald,—Amelanchier spicata 127 


mate. Similarly, the TYPE of A. humilis (see Jones, plate xv) 
and what Jones takes as an isotype of A. austromontana (Jones, 
plate xvi) have the lateral veins fewer and more distant than in 
the type of A. spicata, besides more elongate blades with more 
uniformly subacuminate tips. In foliage, then, A. stolonifera, 
humilis and austromontana are very inadequate matches for the 
leaves of A. spicata. In the type of A. spicata (PLATE 1027, 
FIG. 2) the sepals and petals are erect, not a very convincing 
match for the “sepals usually recurved from the middle after 
anthesis" of the three American shrubs (see our PLATE 1029 
FIGS. 1 and 2, and 1030, ria. 1), while in ours the petals diverge. 
It is to me quite evident from the photograph of the type- 
specimen of A. spicata that in flowers, as well as leaves, it is not 
any of the American shrubs which Jones identifies with it, he 
saying (p. 57) “it becomes clearly evident that Lamarck’s Cra- 
taegus spicata is the common and widespread small serviceberry 
of eastern North America". Wiegand, Rehder and others had 
already clearly shown that it is not, for the Lamarck shrub is 
fastigiate and up to 16 or more feet (““—5 m.") high, not a very 
convincing similarity to “Low surculose colonial shrubs 0.3-2 m. 
tall." i 
Lamarck’s own account should be convincing: 


8. ALISIER à épis, Crataegus spicata. Amelanchier du Canada à 
petites fleurs. 

Cet arbrisseau a beaucoup de rapport avec le précédent; mais il 
s'éléve deux ou trois fois davantage. Ses feuilles sont pétiolees, ar- 
rondies, dentées, vertes en dessus, páles en dessous, & glabres des deux 
côtés dans leur parfait dévéloppement. Elles sont alors à-peu-prés 
aussi larges que longues, & ont un pouce & demi de diametre ou environ. 
Les fleurs sont blanches, petites, & naissent sur des grappes un peu 
étroites, qui ressemblent à des épis. Ces grappes sont munies de 
petites bractées linéaires, colorées, caduques, & qui sont plus longues 
que les pédoncules propres de chaque fleut; ce qui n'a point lieu dans 
l'espece précédente. Les pétales sont une fois plus courts que ceux de 
l'Amélanchier, & ses baies sont une fois plus grosses que les siennes, 
c'est-à-dire, ont au moins la grosseur des fruits du Prunelier ou Prunier 
épineux. On prétend que cet arbrisseau se trouve au Canada. On le 
cultive au Jardin du Roi & dans les jardins des Curieux, ainsi que le 
suivant. (v. v.)—Lam. Encycl. Meth. i. 84 (1783). 


Immediately following this account Lamarck described as 
Crataegus racemosa the American fastigiate shrub or tree which 
was common in European gardens and which Linnaeus had de- 


128 Rhodora [JUNE 


scribed as Mespilus canadensis, the species correctly called 
Amelanchier canadensis (L.) Medic., but known until recently as 
A. oblongifolia (T. & G.) Roemer!, an American species to which 
various American students (Robinson, Rehder and others) have 
sometimes referred shrubs which Jones now places in his hetero- 
morphic “A. spicata’. If Crataegus spicata were, as we are now 
positively assured, ‘‘the common and widespread small service- 
berry of eastern North America", in which Jones includes ele- 
ments which others have considered variations of A. canadensis 
(Crataegus racemosa Lam.), it is astounding that Lamarck did 
not suggest the resemblance but distinctly said that his new C. 
spicata “has much the aspect of the preceding, but rising two or 
three times as high". As to "the preceding", it was the one 
native European species (our PLATE 1028) of Amelanchier, the 
type of the genus, Mespilus Amelanchier L., called by Lamarck 
Crataegus rotundifolia Lam. l. c., the species correctly called A. 
ovalis Medicus but often known as A. vulgaris Moench or A. ro- 
tundifolia (Lam.) Dum.-Cours. Wiegand's figures for the height, 
2-5 m. (or up to nearly 17 feet) and Rehder's statement that A. 
spicata is a tall and fastigiate shrub, already quoted, do not seem 
like Jones's *Low, surculose colonial shrubs 0.3-2 m. tall". 
Altogether the original account by Lamarck gives as little support 
to the recent misidentification of A. stolonifera et al. as A. spicata 
as does the photograph of the type. 

Lamarck compared his Crataegus spicata only with the one 
European species, his Crataegus rotundifolia, the Amelanchier 
ovalis Medicus, but it was much taller and with larger leaves and 
fruit, the leaves of C. rotundifolia “petites en comparaison de 
celles des autres espéces". When we examine characteristic 
material of A. ovalis (our PLATE 1028), we note the leaves with 
outline much as in the type of A. spicata, with their veins simi- 
larly subapproximate; the sepals and petals erect; and the pedicels 
and hypanthium densely pilose (ria. 2). It is difficult to escape 
the conclusion that A. sp?cata is closely related to the European 
A. ovalis. If it is desired, on account of its much greater height 
and fastigiate development, to consider it of hybrid origin, why 
not invoke A. canadensis (oblongifolia), as is done by Wiegand 


1 For detailed study of this identification and for photograph of the type of Mespilus 
canadensis see Ruopona, xliii, 560—563, plate 672, fig. 1 (1941). 


Rhodora Plate 1027 


cai 


Photo B. G. Schubert, after Alfred Rehder 


Types, X 1, of AMELANCHIER SPICATA (Lam.) K. Koch (Crataegus spicata Lam.). 


Rhodora Plate 1028 


Photo B. G. Schubert 


The European AMELANCHIER OVALIS: FIG. 1, flowering and fruiting branches, X 1; 
FIG. 2, portion of flowering raceme, X 5. 


Rhodora Plate 1029 


Photo B. G. Schubert 


AMELANCHIER STOLONIFERA from TYPE-SERTES! FIG. 1, flowering branch, X 1; FIG. 2, 
portion, X 5, of raceme just past anthesis; rics. 3 and 4, lower and upper faces of mature 
leaves, X 1. 


Rhodor: Plate 1030 


Photo B. G. Schubert 


AMELANCHIER STOLONIFERA from TYPE-SERIES: FIG. 1, branch with half-grown fruit, X 1; 
FIG. 2, leaves of vegetative sprout, X 1. 


1946] Fernald,—A Monograph of Amelanchier 129 


and by Rehder (Man.), but without dragging in the very different 
A. stolonifera. In A. canadensis, which had long been cultivated 
in European gardens and which, like A. spicata, “on cultive au 
Jardin du Roi", the leaves are broadly rounded at summit, the 
young hypanthium densely pilose and the sepals soon erect or 
strongly ascending. 

The insistence that Amelanchier spicata originated in America 
is based on rather unsatisfactory evidence. Lamarck did not 
assert that it came from Canada, but that “They pretend or 
claim that it is found in Canada” (“On prétend que cet arbrisseau 
se trouve au Canada”). That was a qualified statement, not a 
clear statement that it is Canadian. It seems more probable 
that it originated in the Jardin du Roi, possibly as a cross of A. 
canadensis and A. ovalis. It is time to exclude without quibble 
the name A. spicata (Lam.) K. Koch from North American 
treatments of Amelanchier. 


EXPLANATION OF PLATES 1027-1030 


PLATE 1027, AMELANCHIER spicata (Lam.) K. Koch, both figs. enlarged 
to X 1, from photograph of Type of the basic Crataegus spicata Lam. (from 
negative taken by Professor Alfred Rehder and preserved in the Herbarium of 
the Arnold Arboretum): Fics. showing vegetative leafy shoot with subap- 
proximate lateral nerves, the blades broadly rounded to subtruncate, the 
calyx-teeth and petals strongly ascending. 

PLATE 1028, AMELANCHIER OVALIS Medic.: Fic. 1, flowering and fruiting 
branches, X 1, from Alsace, Spach (note broadly rounded to subtruncate 
leaves with subapproximate ribs; also erect calyx-lobes and petals); FIG. 2, 
portion of flowering raceme, X 5, showing densely pilose pedicel and calyx; 
also erect calyx-teeth and petals. 

PLATE 1029, AMELANCHIER STOLONIFERA Wiegand, all figs. from TYPE: FIG. 1, 
flowering stem, X 1, showing long smoothish pedicels and spreading calyx- 
teeth and petals; FIG. 2, summit of raceme just past anthesis, X 5, showing 
glabrous pedicels and calyx and spreading calyx-teeth; rics. 3 and 4, lower 
and upper faces of mature (autumnal) leaves, showing remote ribs, X 1. 

PLATE 1030, AMELANCHIER STOLONIFERA Wiegand, both figs. from TYPE: 
FIG. l, stem with half-mature fruit, showing short-acuminate leaves, long 
glabrous pedicels and spreading calyx-teeth; ria. 2, leaves of vegetative sprout, 
showing distant ribs. 


A MONOGRAPH OF AMELANCHIER.—Perhaps no genus of the Rosaceae in 
North America, except of course Rubus and Crataegus, has offered so much 
of perplexity and has had such contradictory treatment as Amelanchier. 
Wiegand brought much light into the problem and established by his 
detailed studies, published from 1912 to 1921, the more important char- 
acters of growth-habit, flowers, fruit and foliage of the several eastern 
North American species. Wiegand, however, too much accepted tradi- 
tional misidentifications and apparently did not always trace the older 
names to their actual types; his strength lay in his tireless field-observa- 


130 Rhodora . [JUNE 


tions and clarification of morphological characters. Most of the types 
have subsequently been checked and the names of several species, of which 
the morphological characters were accurately and painstakingly worked 
out by Wiegand, have, consequently, been readjusted. Now comes a 
beautifully printed and well illustrated monograph of 100 pages and 23 
plates by Professor George Neville Jonest. Professor Jones dismisses at 
once the decision of Wiegand and very many others with extensive field- 
experience that hybridization occurs to any extent in Amelanchier, al- 
though he does actually admit the single hybrid, x A. grandiflora Rehder 
and in a note concedes a cross of A. Bartramiana and A. canadensis. 
His conclusion is based on cytological studies by others in the subfamily 
Pomoideae, Sax having reported, as quoted by Jones, "that the pure 
species of Amelanchier that he studied are diploids, but two natural inter- 
mee hybrids are tetraploids" and Moffett having stated that ''the 
chromosome number of the Pomoideae is a ‘secondary basic number (un- 
balanced relative to the primary basie number) and the derived series of 
polyploids . . . are secondary polyploids'." Jones goes on: “It seems 
possible, therefore, that polyploidy may have played a more important 
part in the differentiation of genera and species in this subfamily than 
has heretofore been realized, since a change in chromosome balance is 
usually accompanied by a change in the morphological characters of the 
plant”. Isn't it common knowledge that the vast number of strikingly 
different and fully appreciated apples and pears (such strikingly dissimilar 
apples as Baldwin, Gilliflower, Lady and Red Siberian, or such contrasted 
pears as Bartlett, Seckel and Beurre Bosc) are the result of hybridization? 
Ability to hybridize is the basis of pomology. If the Pomoideae did not 
freely lend themselves to crossing, apples and pears would be pretty poor 
food. Groups in which hybridization (therefore allopolyploidy) is rare 
or difficult to achieve are not horticultural successes! Professor Jones 
dismisses generally recognized hybridization in the Pomoideae because 
“polyploidy may have played a more important part"; but he does not 
seem to recognize the elementary fact that allopolyploidy is an accompani- 
ment of hybridization! Witness the distinction between the two types of 
polyploidy as clearly indicated in Julian Huxley’s Evolution: the Modern 
Synthesis" (p. 334): 


Next we have the various phenomena of polyploidy in which a multiplica- 
tion of whole genomes or chrothceome-sete occur. As already mentioned, 
polyploidy is of two fundamentally distinct types: autopolyploidy in which 
the chromosome sets are all of the same kind, derived from the same s cies, 
and initial allopolyploidy, in which they are of different kind, derived from 
two distinct species. The actual doubling is in both cases due to the suppres- 
sion of division of a cell after division of the chromosomes has taken place, but 
whereas this is the primary event in autopolyploidy, in allopolyploidy it is sub- 
sequent to hybridization. 


The generally recognized hybrid of A. Bartramiana and A. laevis, which 
Dr. Jones evidently, from the citation of specimens, has known only as 
dried material collected by others, is treated as a full species, A. neglecta 
Eggleston, although the specimens loaned to Dr. Jones for study often 
have his validation as “X A. neglecta"! Most other hybrids, of which 


1 Georcr NEvILLE Jones. American Species of Amelanchier. Illinois Biological 
Monographs, xx. no. 2. 100 pp. 23 plates. Univ. Ill. Press, Urbana, Ill. 1946. $1.50, 
paper-cover; $2.00, cloth-cover. 


1946] Fernald,—A Monograph of Amelanchier 131 


Wiegand had very many, are placed without reservation in single recog- 
mre species because, by the author’s unique reasoning, they are poly- 
ploids. 

A total of 18 species is admitted for the whole continent, an ultra- 
conservative number which many, who have had more extended field- 
acquaintance with the shrubby members, will doubtless be inclined to 
augment. Pubescence (usually a pretty variable and too often a passing 
character) is largely relied upon, the first division of the eastern species 
depending on “Top of ovary glabrous" as opposed to “Top of ovary 
tomentose, usually densely so, rarely with only a few trichomes”, while 
under the latter the eastern species are again divided on more or less per- 
sistence of pubescence. This may be the best that can be done, but it is 
unfortunate that so fluctuating a character as persistence of pubescnce 
has to be made fundamental. The characters of hypanthia, sepals, petals, 
styles, etc., used to separate species, seem more stable; and growth-habit, 
which is well understood by those who, like Wiegand, spent many seasons 
in studying the genus in the field over an extensive area, is really important. 

Dr. Jones reproduces, most happily, photographs, so far as they were 
accessible to him, of types or isotypes, a large proportion of the originals 
borrowed from the Gray Herbarium. In two cases only tracings made by 
others and borrowed by him are given, as in plate XI, fig. 1, a tracing 
made by the late Benjamin L. Robinson and bearing in his hand the 
explanation, or in plate XVII, fig. 2, another tracing made by Dr. Robin- 
son (with his characteristic handwriting nearly covering the sheet). 
These illustrations are invaluable. 

In some cases, at least, the photographs of the types lead one to wonder 
at the identifications. For instance, plate XI, fig. 2, shows a small and 
not very clear photograph of Crataegus spicata Lam., basonym of A. 
spicata (Lam.) K. Koch, which Dr. Jones feels to be “clearly . . . the 
common and widespread small serviceberry of eastern North America": 
A. humilis Wiegand, A. stolonifera Wiegand and the more localized and 
southern A. austromontana Ashe. Whether those who have for many 
years known the stoloniferous and loosely colonial low A. stolonifera and 
A. humilis will be satisfied that the calcicolous inland A. humilis and the 
oxylophytie A. stolonifera, chiefly of the Atlantic slope, are identical is 
very doubtful; and it is extremely doubtful if anyone but Dr. Jones will 
be satisfied that either of these dwarf and colonial American shrubs is 
identical with the non-colonial, fastigiate and very tall Crataegus spicata 
of Lamarck. The latter, a shrub of unknown nativity, supposed or 
claimed to have come from Canada (“On prétend que cet arbrisseau se 
trouve au Canada"), was cultivated in Paris in 1783. Lamarck stated 
that his C. spicata strongly resembled the widespread European species, 
Mespilus Amelanchier L., which under Amelanchter is known as A. ovalis 
Medicus, A. vulgaris Moench or A. rotundifolia (Lam.) Dum.-Cours., 
the species which had been called by Lamarck Crataegus rotundifolia. 
Lamarck said that his C. spicata was two or three times taller (“Cet 
arbrisseau [Crataegus spicata] a beaucoup de rapport avec le précédent 
[C. rotundifolia], mais il s'éléve deux ou trois fois davantage"). Since 
Lamarck's C. rotundifolia was 3-5 feet high (“la hauteur de trois à cinq 
pieds"), that would make C. spicata measure "about 2-5 m." (as pointed 
out by Wiegand who is quoted by me in the illustrated study of Lamarck's 
type on p. 125, with plate 1027 in this number of RnHopon4) in 


132 Rhodora [JUNE 


height. The North American A. humilis is consistently lower, 0.3-1.5 m. 
high (“3-12 dm.", Wiegand, and also Rosendahl & Butters, Trees and 
Shrubs Minn.; “4-15 dm.”, Deam, Shrubs Ind.), while A. stolonifera 
(details shown in plates 1029 and 1030 in this number) is quite as low 
(“3-12 dm.", Wiegand). Dr. Jones, discussing the habit of species, says 
(p. 10): “Certain other species, e. g., A. spicata, are usually dwarf, but 
under exceptionally favorable conditions may reach a height of two me- 
ters". That still falls far short of the maximum of 5 meters implied by 
Lamarck for the cultivated type of A. spicata; and if, as Dr. Jones believes, 
the southern A. austromontana belongs to all-inclusive A. spicata, it is 
too bad that he restricted the height to “0.3-2 m.", for A. austromontana 
was originally described by Ashe as “not exceeding 4 m. in height" (a 
back-handed way of implying at least more than 2 m.). The photographs 
of types of A. spicata, A. stolonifera, A. humilis and A. austromontana are, 
fortunately, all shown by Dr. Jones and outlines of characteristic leaves 
are displayed separately. While the leaves of the type of A. spicata (see 
plate 1027 in the article preceding this review) are quite broadly rounded 
to subtruncate at summit, those of all three American types are of nar- 
rower and more elongate outline and more gently rounded or tapering to 
the short-tipped apex and the veins of A. spicata are relatively more 
approximate. The three Americans are closely akin; the type of A. 
spicata evidently an alien, with leaves closely matched in outline, toothing 
and venation by very many specimens of the European A. ovalis Medic. 
(Mespilus Amelanchier L.) (See plate 1028 in preceding article). 

If, as would seem from the photograph of the type, Crataegus spicata 
Lam. is a variation or perhaps a hybrid of the European Amelanchier 
ovalis, then it would be distinguished from our shrubs by several char- 
acters, including, according to Rehder's Manual, “Styles free” in A. ovalis 
as opposed to “Styles connate at least at base" in the remainder of the 
species, this character sometimes considered to have such magic that it 
is relied upon as the final argument in keeping apart as genera Pyrus 
(“Styles free") and Malus (‘Styles connate at base")'. If it be main- 
tained that Amelanchier spicata is not a phase of or derivative from A. 
ovalis, it can, at least, not be satisfactorily identified with three or more 
lower and habitally very different American species. 

Further differences from the interpretations of Wiegand and others, 
like the reduction outright to A. sanguinea (Pursh) DC. of A. huronensis 
and A. amabilis Wiegand, need not here be discussed. They at least 
show very strong departures in interpretation of characters. 

Outline maps with dots indicate the ranges. These, it must be assumed, 
give a general picture of the range, and when such strongly contrasted 
areas as those of the western A. pallida (map 13) and of the eastern A. 
Bartramiana (map 1) are shown the contrast is striking. When, however, 
A. stolonifera, humilis and austromontana are merged (as A. ‘‘spicata’’) 
one does not know how, from the map, to sort out the different elements 
which others recognize as true species. In some cases more attention to 
exact geography would have been desirable. For instance, map 2 is said 
to show the range of A. neglecta. It does in a general way, but since this 
shrub is not cited (or known) from the Magdalen Islands, it is at least 
disconcerting to see the outlines of that archipelago blackened out by a 


1 For an analysis of the reputed but sadly inconstant morphological characters 
separating Malus from Pyrus see RHopona, xlv. 450 (1943). 


1946] Fernald,—A Monograph of Amelanchier 133 


round dot. From Nova Scotia a single collection is cited, from Meteghan, 
at the western end of the Province, but unsupported dots are shown all 
the way to northern Cape Breton, more than 500 miles northeast of the 
one recorded station at Meteghan. Again, the great Province of Quebec 
seems to have been as much of a puzzle to the map-maker as it was when, 
in his Monograph of Symphoricarpos in Journ. Arn. Arb. xxi. 216 (1940), 
he cited the type of S. albus as coming from “Quebec, Ontario, Canada". 
On the map of A. neglecta a single row of six equally spaced dots runs from 
the tip of the Gaspé Peninsula to the Richelieu River south of Montreal 
and no dot is shown north of the River St. Lawrence. Five stations are 
cited (instead of six): Gaspé Bay; Montmagny (more than 300 miles to 
the southwest); Montmorency Falls (a famous tourist-resort about 40 
miles west of Montmagny and nortH of the St. Lawrence); Lac Long 
(about 65 miles northwest of Montmorency and also NortuH of the St. 
Lawrence); and “Ottawa River" (hundreds of miles long and joining the 
St. Lawrence FROM THE NORTH). A mere outline of the general range 
might have suggested the main point; the failure carefully to consult an 
atlas would be less evident. 

All in all the new monograph of Amelanchier is a very neat piece of 
presswork. Its illustrations of types are well reproduced and most help- 
ful, the general ranges approximately shown (though slipping on unfa- 
miliar territory), its descriptions detailed, the recent identifications by 
others of types of American species accepted (but the specific segregations 
and the hybrids of Wiegand not admitted) and the citations of specimens 
very full (though said to be incomplete). In the latter particular, with 
what Wiegand and many others have always considered quite separate 
elements here merged, it would have helped others if specimens of the 
different included elements could have been indicated by some sign. 
Where individual judgment so radically differs as in this “Monograph”, 
especially through obviously unequal degrees of field-work and consequent 
understanding of the different characters and, likewise, through failure 
to recognize the well known fact that allopolyploids are hybrids, it would 
have helped, so long as the largest portion of the work is given over to 
very full listing of specimens, if the original identifications by Wiegand of 
his own species, for instance, had been noted. That would partly sepa- 
rate the different elements. Then the shrub which Wiegand and scores 
of other critical students (Butters, Deam, Fassett, Rehder, Rosendahl and 
many more) have considered A. humilis would not (when merged with the 
habitally similar A. stolonifera under the impossibly American and habit- 
ally very dissimilar A. *'sp?cata") appear to occur from Newfoundland to 
Georgia, where it is not found. 

In the enumeration of numbered specimens (admittedly only a part of 
those seen) the author of the latest treatment of Amelanchier has the very 
modest score of 32, Wiegand of 85, a score which could and certainly 
should have been most profitably increased if Wiegand’s authoritative 
series of 696 sheets left at Cornell University and his earlier authentic 
series of 63 sheets at Wellesley College—a total, with the 85 cited assumed 
to be duplicates, of more than 750 sheets, which largely formed the bases 
of Wiegand’s detailed publications over ten years—had been consulted. 
Most surprisingly, however, there is no indication in the acknowledgments 
that the new author on Amelanchier was interested to see this great funda- 
mental mass of authentic material. One other close field-student of the 


134 Rhodora [JUNE 


genus over many years, one who understands the southeastern species, is 
credited with having participated in the collection of 183 cited numbers; 
another, who has closely watched the group in the field for half a century 
and who largely accompanied the painstaking Wiegand, with about 225. 
These are mere trivialities compared with Wiegand’s record of more than 
750 but impressive when compared with a total score of only 32. If these 
clear evidences of field-experience and intensive work on and under- 
standing of the genus had been reversed, perhaps the results would be 
very different.—M. L. F. 


THe VARIETIES OF LYCOPODIUM INUNDATUM.— The circum- 
boreal Lycopodium inundatum L., if it had stayed in the cool- 
temperate areas of Eurasia, would present no troublesome 
problem. In North America, however, it is one of a series of 
species and varieties which extend, as a somewhat perplexing 
group, all the way from Newfoundland to the Tropics. By 
many authors it has been considered to be confluent with the 
coarse L. alopecuroides L. of the southeastern United States and 
the West Indies, a species characterized by its low-arching 
or recurved-procumbent stem rooting at the tip, its leaves 
linear- or narrowly lance-attenuate and with the similar sporo- 
phylls usually bristly-ciliate, the upright densely leafy fertile 
branches 0.5-1.5 em. thick, the strobile 1-2.5 cm. thick, the 
spores with coarsely reticulate base and with the apical half 
covered with coarse papillae crowded into rows. Thus Hooker 
in his British Ferns (1861) wrote, under L. inundatum, pl. 51: 
“L. Carolinianum and L. alopecuroides, and not a few others, 
supposed species, must be added to this list, if we were to make 
the exotic synonymy complete" ; then, to add to the bibliographic 
confusion, he cited as a synonym ‘‘Lycopopium Bigelovii. 
Oakes and Tuckerman in Sillim. Journ.". The latter unverified 
reference Hooker obviously borrowed from Spring, Mon. Fam. 
Lycopod. pt. ii. 33 (1849), who had cited as a synonym of L. 
inundatum L.: ** L. Bigelovii Oakes et Tuckerm.! prius (ex Tuckerm. 
in Sill. Journ. of Nat. Hist.)." Oakes and Tuckerman made no 
such binomial! Instead, Tuckerman in Am. Journ. Sci. and Arts 
(conducted by Professor Silliman and Benjamin Silliman, Jr.), 
xlv. 47 (1843) had under L. inundatum two varieties: “8. Bigelovii, 
(mihi): majus, ramis subramosis elongatis, folis acuminatis 
sparsim denticulatis s. integris. L. Carolinianum, Bigel. Fl. 
Bost. p. 384.—y. alopecuroides, (mihi): . . . L. alopecuroides L.", 


1946] Fernald,—Varieties of Lycopodium inundatum 135 


etc. The significant point is that var. @. Bigelovi?, based in part 
on L. carolinianum Bigel. Fl. Bost. ed. 2: 384 (1824), not L., 
Bigelow’s plant with ‘‘peduncle . . . long, slender", coming 
from Cape Cod (Sandwich), was based by Tuckerman primarily 
on a collection from “Plymouth, Oakes and Tuckerman”. Spring 
confused the collectors with the author and wrongly made them 
the authors of a species they did not describe; Hooker followed 
suit and in copying Tuckerman’s synonymy, neglected to em- 
phasize that he meant L. caroliniamum sensu Bigelow, not L. 
As pointed out by me in RHopora, xxiii. 100 (1921), the type of 
Lycopodium inundatum, var. Bigelovii, the Oakes and Tucker- 
man plant from Plymouth, is inseparable from L. adpressum 
(Chapm.) Lloyd & Underwood or L. Chapmani Underwood, 
this well illustrated in Britton & Brown, Ill. Fl. ed. 2, i. fig. 104 
(1913), their fig. 105, purporting to show L. alopecuroides, being 
made from a specimen of L. inundatum, as shown by the short 
creeping sterile branches, the slender fertile branch, and the 
ovate-based sparsely toothed sporophyll. In some later writings, 
as in RHopona, xxxviii. 382 (1936), I lost the thread and con- 
fused two varieties, taking up again L. inundatum, var. adpressum 
Chapm. for what had earlier been described as var. Bigelovii 
and treating, incorrectly, as var. Bigelovii a coarser variety 
which had been described from southern New England and New 
Jersey as L. inundatum, var. robustum R. J. Eaton in RHonopa, 
xxxiii. 202 (1931). As I now understand the three varieties of 
L. inundatum they may be defined as follows. 
Leading creeping terminal shoot elongating from 1.5-10 
(when submerged -15) cm. beyond the last of the 1 or 2 
(very rarely 3) fertile branches; fertile branches 0.5-7 
(rarely -10) cm. high; strobile 0.8—4 cm. long, 6-10 mm. thick, 
its ovate- or lanceolate-based bracts with loosely ascending 
to spreading tips; transcontinental northern plant. . L. inundatum (typical). 
Leading creeping terminal shoot elongating from 0.7-4 dm. 
beyond the last of the mostly 2-10 fertile branches; the 
latter 0.5-4 dm. high; strobile (1.5-) 2-15 cm. long, its 
bracts with lanceolate bases; southeastern and eastern 
coastwise plants. 
Fertile branches stiffly erect (flexuous only when drowned), 
2-5 mm. thick, densely covered with loosely ascending to 
spreading leaves mostly 5-8 mm. long; strobiles 7-14 
mm. thick at base, including tips of loosely ascending to 
spreading elongate (5-8 mm.) bracts; sporangia mostly 
wadon undor the bracts c ee ee Var. robustum. 


Fertile branches erect or flexuous, 1.5-3 mm. thick, more 
openly covered with appressed or ascending leaves 3—5 


136 Rhodora [JUNE 


mm. long; strobile 3-7 mm. thick, with closely appressed 
or ascending shorter (3-6 mm.) bracts; outlines of spor- 
angia (at least in dry specimens) often evident through 
the thin bracts. ..,.. 000.000.0000 ccc sess. Var. Bigelovii. 


L. iNuNDATUM L. Sp. Pl. 1102 (1753).—Damp peaty or sandy 
shores, swamps or bogs, Newfoundland to Alaska, south to Nova 
Scotia, New England, New Jersey, Pennsylvania, mountain- 
region to western Virginia and West Virginia, northern Ohio, 
northern Indiana, northern Illinois, Minnesota, Idaho and 
Oregon; Eurasia. 

Var. noBusTUM R. J. Eaton in RnHopona, xxxiii. 202 (1931). 
Var. Bigelovit sensu Fernald in Rnopona, xxxviii. 382 (1936), 
not Tuckerman.—Eastern Massachusetts, south on or near 
coastal plain to Florida and Louisiana. 

Var. ROBUSTUM, forma furcatum (Fernald), comb. nov. Var. 
Bigelovii, forma furcatum Fernald in Rnopona, xliv. 377 (1942). 

Var. BiaELovir Tuckerm. in Am. Journ. Sci. xlv. 47 (1843). 
L. carolinianum sensu Bigelow, Fl. Bost. ed. 2: 384 (1824), not 
L. (1753). Var. appressum Chapm. in Bot. Gaz. iii. 20 (1878), 
altered to Var. adpressum Chapm. Fl. So. U. S. ed. 2: 671 (1883). 
L. alopecuroides, var. adpressum (Chapm.) Chapm., Fl. So. U. S. 
ed. 3: 638 (1897). L. adpressum (Chapm.) Lloyd & Underwood 
in Bull. Torr. Bot. Cl. xxvii. 153 (1900). L. Chapmani Underwood 
in Maxon in Proc. U. S. Nat. Mus. xxiii. 646 (1901). L. alopecu- 
roides, ssp. adpressum (Chapm.) Clute, Fern Allies, 118, pl. 5 
(1905). L. alopecuroides, forma adpressum (Chapm.) Clute in 
Fern. Bull. xvii. 48 (1909).—Florida to eastern Texas, north on 
or near coastal plain to southeastern New Hampshire and Nova 
Scotia, where freely passing into typical L. inundatum. 

Var. BrGELovrr, forma polyclavatum (McDonald), comb. nov. 
L. adpressum, f. polyclavatum McDonald in Fern Bull. ix. 9 
(1901). L. alopecuroides, var. adpressum, f. polyclavatum 
(McDonald) Clute in Fern Bull. xvii. 45 (1909). L. inundatum, 
var. adpressum, f. polyclavatum (McDonald) Fernald in Rmo- 
DORA, xli. 405 (1940).—M. L. FERNALD. 


New COMBINATION IN CHRYSOBALANUS.—As a preliminary to 
issuing the next century of the Plantae Exsiccatae Grayanae it 
becomes necessary to make the following new combination: 

CHRYSOBALANUS pallidus (Small), comb. nov.  Geobalanus 


pallidus Small, Fl. Miami, 81, 200 (1913)—Lyman B. SMITH, 
Gray Herbarium. 


Volume 48, no. 569, including pages 89-112 and plates 1021-1026, was issued 
6 May, 1946. 


a 


rt 
m UA 


Rhodora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. July, 1946. No. 571. 


CONTENTS: 


Contributions from the Gray Herbarium of Harvard University— 
No. CLXII. Identifications and Reidentifications of North 
American Plants. M. L. Fernald. ........cccccacccsvccccn 137 


Relict Boreal Plants in Southeastern Minnesota. J. B. Moyle. .. 163 


Another later Homonym. Charles L. Gilly. ............ suu. 163 


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JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. July, 1946. No. 571. 


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


IDENTIFICATIONS AND REIDENTIFICATIONS OF 
NORTH AMERICAN PLANTS 


M. L. FERNALD 
(Plates 1031-1050)! 


During a somewhat searching study of different groups in the 
flora of eastern North America it is found imperative, if we are 
properly to identify our plants, to study the original descriptions 
and their citations and, when possible, to see the types or good 
photographs of the types. Otherwise, as in too many cases, we 
make wrong guesses and encumber taxonomy with erroneous 
and quite misleading names. In some of the present series of 
studies attempts have been made to place the names of the 
plants studied on a surer basis than heretofore. 


THE IDENTITY OF QUERCUS LAURIFOLIA (PLATES 1031—1036). 
Quercus laurifolia Michx. Hist. Chénes Am. Sept. tt. xvii. and 
xviii (1801), has so long been misinterpreted, at least since 
Elliott in 1824, that it becomes necessary to study in some detail 
the original material of Michaux and the two species which have 
figured in the misinterpretation of his clearly defined and beauti- 
fully illustrated species. 

1 The cost of preparing and engraving the plates met in part through grants from 


the AMERICAN PHILOSOPHICAL Society and from the DEPARTMENT or BIOLOGY OF 
HARVARD UNIVERSITY. 


138 Rhodora [JULY 


The two principal elements involved in the problem are (1) 
Q. hemisphaerica Bartram as validated by Willd. Sp. Pl. iv!. 443 
(1805), the southern tree (our PLATES 1035 and 1036) with full 
hemispherieal or subspherical head; the coriaceous, dense and 
opaque, evergreen or only tardily deciduous (at least overwin- 
tering) leaves, with foliage of the fruiting branches oblong- 
lanceolate or -oblanceolate to narrowly elliptic or oblong, quite 
glabrous on both lustrous surfaces, cuneate at base, either acute 
or obtuse, 3-10 em. long and 1-3 em. broad, very short-petioled; 
leaves of vigorous vegetative sprouts often sinuate-dentate or 
-serrate; cup shallowly saucer-shaped, 1-1.5 cm. broad and 3-5 
mm. high, with strongly flattened-rounded base; the nut 0.8-1.5 
em. long; the tree occurring on the outer coastal plain from 
Florida to eastern Texas, north to southeastern Virginia; and (2) 
the tree with longer, broader and thinner leaves membranaceous 
and showing, by transmitted light, a clear and intricate reticula- 
tion, the glabrate lower surface often with tufts of pubescence in 
the axils of the lateral veins; the cup deeply saucer-shaped, 
shallowly cup-shaped or turbinate, 1.5-2 cm. broad and 7-10 
mm. high; this being the species described in detail by Sargent as 
his Q. rhombica Sargent in Bot. Gaz. Ixv. 430 (May 15, 1918), our 
PLATES 1031 to 1033, with “Leaves rhombic, rarely oblong- 
obovate to lanceolate, acute or rounded . . . at apex, cuneate 
at base, . . . at maturity thin, . . . 9-12 cm. long and 3.5-5 
cm. wide, with . . . slender primary forked veins; . . . falling 
gradually in early winter; . . . cup saucer-shaped to cup-shaped 
... From Q. laurifolia it differs in the shape of its thinner 
leaves . . . and in its larger fruit with much deeper cups". As 
a variety Sargent described material from a single tree, his var. 
obovatifolia (our PLATE 1034) which differed “in the obovate 
leaves at the ends of the branches, rounded or slightly 3-lobed or 
undulate at the broad apex”. 

On a succeeding page (432) Sargent discussed what he took to 
be true Quercus laurifolia Michx., with “leaves, which are thicker 
than those of Q. Phellos L., are dark green, very lustrous, and 
glabrous . . . usually not more than 7-8 em. in length;" and 
(p. 433) saying “The laurel oak is not evergreen. Late in the 
winter the leaves begin gradually to turn yellow and then brown, 
and when the buds begin to swell at the appearance of spring 


Rhodora Plate 1031 


Photo D. G. Schubert 


QUERCUS LAURIFOLIA Michx.: FIGs. 1 and 2, foliage and fruit, X 1, after Michauz; 
FIG. 3, fruit of Q. rhombica Sargent, after Sargent; ria. 4, leaf, X 1, of Q. laurifolia, 
rar. hybrida Michx. or var. obtusa Willd., after Michauc. 


Rhodora Plate 1032 


Photo B. G, Schubert 


QUERCUS LAURIFOLIA Michx.: eras. | and 2, fruiting branch and terminal leaves, X 1, 
from Virginia; FIG. 3, leaves, X 1, of PARATYPE of Q. rhombica Sargent. 


1946] Fernald,—Identifications of North American Plants 139 


drop almost simultaneously”. Later, in his Man: Trees N. Am. 
ed. 2: 261 (1922) the “much deeper cup” *saucer-shaped or cup- 
shaped" of Q. rhombica became merely “a saucer-shaped cup”, 
although the artist saw it and represented it (our PLATE 1081, 
FIG. 3) as definitely turbinate and half as high as broad. In this 
edition of Sargent's Manual the earlier and inclusive description 
of Q. laurifolia was not sufficiently altered to bring out all the 
contrasts between the two species as conceived by Sargent, for 
the “thicker” leaves were here described as “at maturity thin", 
although the cup of the species with narrow coriaceous and over- 
wintering leaves was correctly illustrated and described as ‘‘a 
thin saucer-shaped cup". 

In his original discussion of Quercus laurifolia in his sense 
(1. e. Q. hemisphaerica), with leaves ‘‘usually not more than 7-8 
cm. long" and, as stated in his Manual, ‘34’ [7. e. 1.8 em.] wide”, 
Sargent commented on the plates which Michaux published with 
the original description of Q. laurifolia: “On the branches figured 
by MicHavx the leaves are generally elliptic but sometimes 
slightly broadest above the middle, acuminate at the ends [in 
Michaux's var. hybrida (our PLATE 1031, rra. 4) ‘obovate and 
rounded at apex’’], and 6-12.5 cm. in length". Furthermore, 
Michaux's plate (our PLATE 1031, FIGs. 1 and 2) showed, in case 
of typical Q. laurifolia, a fruiting branch with leaves mostly 2—4 
cm. broad, and a photograph in the Gray Herbarium of his type 
of the typical form shows very thin and translucent foliage 
(PLATE 1033, FIG. 1), the intricate reticulate venation too strongly 
suggesting that shown by transmitted light in leaves of the type 
of Q. rhombica (PLATE 1033, ria. 3). Instead of being unusual the 
leaves shown by Michaux of his original Q. laurifolia and its var. 
hybrida seem to me inseparable from those of thoroughly char- 
acteristic Q. rhombica, originally described with leaves “9-12 cm. 
long and 3.5-5 em. wide, with stout conspicuously yellow mid- 
ribs and slender primary forked veins" ; and the type of the acute- 
leaved form (var. acuta Willd.) preserved at Paris could easily 
have come from material of the tree illustrated by Michaux. 
Still further, Michaux described the ‘‘cupule un peu turbinée" 
and illustrated it (our PLATE 1031, ria. 2) as pretty definitely so, 
just as in his Manual (our rra. 3) Sargent had so illustrated the 
cup of Q. rhombica. 


140 Rhodora [JULY 


It is most difficult for me to believe that, when Michaux 
described and illustrated as his new Quercus laurifolia, with 
evidently thinnish leaves (the type showing intricate venation 
even in the photograph) on a fruiting branch, the blades up to 
12.5 cm. long and 2-4 em. broad, many of them in outline (as 
well as size) quite like those of Q. rhombica, the cup definitely 
turbinate and shown as half as high as broad, the tree with these 
characters having the thin leaves often with tufts of hair persist- 
ing beneath in the axils of the veins,—it is most difficult to follow 
Sargent in believing that Michaux was wholly mistaken and 
really intended to describe a tree with much shorter and narrower, 
dense, coriaceous, overwintering, glabrous leaves and ‘‘thin 
saucer-shaped" cups. I see no way but to treat Q. rhombica as 
typical Q. laurifolia. 

One week before the publication of Quercus rhombica, Ashe 
picked up the Q. laurifolia @. obtusa of Willdenow, Sp. iv!. 428 
(1805), a substitute preferred by Willdenow for Q. laurifolia 
hybrida Michx., and made the specific combination Q. obtusa 
(Willd.) Ashe in Torreya, xviii. 72 (May 8, 1918), thus antedating 
Q. rhombica of May 15, 1918. Ashe, like Sargent and others, 
took as Q. laurifolia the species which Willdenow had properly 
defined in 1805 as Q. hemisphaerica, for Ashe said of his Q. obtusa: 
“This tree is undoubtedly closely related to Q. laurifolia as 
generally understood, but it can be readily separated from it by 
the leaves of vigorous shoots, which in Q. laurifolia are irregularly 
toothed, while in Q. obtusa the margins are entire. The cup also 
has a very pointed base, while the base of the cup in Q. laurifolia 
is flat.” Had Ashe looked at Michaux’s plate of the original Q. 
laurifolia (our PLATE 1031, FIGs. 1 and 2) he would have seen 
quite entire leaves and a cup with “a very pointed base", not at 
all “flat”. He could also have seen that Michaux's diagnosis 
clearly said ‘‘Cupula subturbinata’’. 

Somewhat later, Ashe, realizing that Quercus rhombica of 
May 15 was Q. obtusa of May 8, published a note in RHODORA, 
xxiv. 78 (1922), making the reduction, still apparently not 
knowing that Q. obtusa and Q. rhombica were phases of true Q. 
laurifolia. He then picked up Sargent’s very unsatisfactory Q. 
rhombica, var. obovatifolia and made the combination Q. obtusa 
obovatifolia (Sarg.) Ashe, this variety, according to him, differing 


1946] Fernald,—Identifications of North American Plants 141 


“only in its spatulate leaves with rounded apices.” Sargent 
had said ‘‘obovate leaves at the ends of the branches, rounded or 
slightly 3-lobed or undulate at the broad apex’’, his type (our 
PLATE 1034) and only cited material being from “A single tree" 
from Beaumont, Texas, E. J. Palmer, no. 12747; Sargent acci- 
dentally omitting the last figure and adding: ‘The terminal 
leaves of this variety, . . . show a relationship with Q. nigra 
L.". They certainly do. 

The more significant bibliography of Q. laurifolia is as follows: 

Q. LAURIFOLIA Michx. Hist. Chênes Am. Sept. t. xvii (1801) 
and Fl. Bor.-Am. ii. 197 (1803). Q. laurifolia hybrida Michx. 
Hist. Chênes Am. Sept. t. xviii (1801). Q. laurifolia a. acuta and 
6. obtusa Willd. Sp. Pl. iv!. 428 (1805). Q. laurina Raf. Alsog. 
Am. 27 (1838), not Humb. & Bonpl. (1809). Q. laurifolia Q. 
obtusa Pursh, Fl. Am. Sept. ii. 627 (1814). Q. Phellos, var. 
laurifolia (Michx.) Chapm. Fl. So. U. S. 420 (1860). Q. aquatica 
6. laurifolia (Michx.) A. DC. in DC. Prodr. xvi?. 68 (1864), in 
part. Q. hybrida (Michx.) Ashe in Proc. Soc. Am. For. xl. 88 
(1916), not (Chapm.) Small (1903). Q. obtusa (Willd.) Ashe in 
Torreya, xviii. 72 (May 8, 1918). Q. rhombica Sargent in Bot. 
Gaz. lxv. 430 (May 15, 1918).—Florida to eastern Texas, north 
on coastal plain to Cape May, New Jersey. 

My reason for including as separate names in the above syn- 
onymy both Quercus laurifolia Q. obtusa Willd. (1805) and Q. 
laurifolia B. obtusa Pursh (1814) is, that Pursh did not cite 
Willdenow as the author of his variety. From the start this 
name has been the source of bibliographie or nomenclatural 
errors which make one blush for the performances of taxonomists 
of renown. In his Report on Forests of North America, 152 
(1884) Sargent gave in his synonymy of Q. laurifolia “Q. obtusa 
Pursh, Fl. Am. Sept. ii. 627". This unintentional new binomial, 
which Pursh had not made, promptly appeared, correctly in this 
instance, in Index Kewensis under Quercus as “obtusa, Sargent 

. insyn’’.; and “Quercus obtusa Pursh ” appears, as if made by 
Pursh, in Sudworth’s Nomenclature of the Arborescent Flora of 
the United States (1897), in Trelease’s writings and elsewhere, 
where some sort of verification might have been expected. Since 
the original Q. laurifolia var. hybrida Michx. (1801) was the sole 
basis of Q. laurifolia 6. obtusa Willdenow (1805), the intricate and 
not very intelligent confusion of the two names must, unhappily, 
be investigated. 


142 Rhodora [JULY 


In Small, Fl. S. U. Se. 350 (1903), something quite different and 
based on Quercus aquatica Walt., var. hybrida Chapman, Fl. So. 
U.S. 421 (1860), this having nothing to do with Q. laurifolia, var. 
hybrida of Michaux, appeared as Q. hybrida (Chapm.) Small, 
although Small, with his customary “lack of time", left others to 
hunt up the place of publication of the basonym. Now Q. 
aquatica of Chapman's time was what has proved to be Q. nigra 
L. (1753) and his var. hybrida was evidently a form of it, with 
"leaves oblong or wedge-oblong, entire, emarginate, or 3-lobed 
at the summit", etc., surely having nothing to do with Q. lauri- 
folia hybrida Michx. (our PLATE 1031, rra. 4). Nevertheless, 
having subsequently decided that his Q. hybrida (Chapm.) 
Small was not worth maintaining, Small dropped it; but, under 
the already much discussed and nonexistent Q. “obtusa (Willd.) 
Pursh”, he gave a single synonym, “Q. hybrida (Michx.) Small’, 
Man. Se. U. S. 428 (1933), thus making still another useless 
binomial, this one invalid because published in synonymy! 
But when he published his Q. hybrida (Chapm.) Small in 1903, 
the author was not retarded because of the identical binomial 
(Q. hybrida) already published four times before, by Brotero in 
1804, by Bechstein in 1843, by Hampton in 1886 and by Houba 
in 1887. As described by Chapman the basic name seems to 
have belonged to a tree very unlike the latter; but, as illustrated 
by Britton & Shafer, N. Am. Trees, fig. 254 (1908), its leaves 
and their venation are those of Q. laurifolia, the cup that of Q. 
hemisphaerica. Probably Q. hybrida (Chapm.) Small was an 
appropriate name. 

In his monumental Histoire des Chénes de l'Amérique Septen- 
trionale (1801) Michaux showed as Quercus aquatica the species 
now known to be Q. nigra L. in plates xix-xxi. Plate xx showed 
several different leaves, not all of them belonging to Q. nigra 
(aquatica) as now understood. Fig. 2 (our PLATE 1035, rra. 1), 
especially, was a vigorous young shoot of an unnamed variety 
with the narrow leaves eut as in those of sprouts of Q. hemi- 
sphaerica. In a special paragraph at the end of his discussion of 
Q. aquatica Michaux commented on such foliage, saying: 

On trouve sur les dunes sablonneuses en Géorgie et en Floride, une 
variété de cet arbre à feuilles étroites, dentées irréguliérement, et qui 


conserve ses feuilles pendant tout l'hiver. C'est probablement cette 
variété que BARTRAM a nommée QuERCUS dentata (narrow leaved winter 


Rhodora Plate 1033 


Photo B. G. Schubert 


QUERCUS LAURIFOLIA Michx.: FIG. 1, venation of lower leaf-surface, X 3, from 
photograph of the TvPE by Cintract; ric. 2, foliage, X 1, of type of Q. rhombica Sar- 
gent; FIG. 3, venation of lower surface of leaf of the latter, X 3. 


Rhodora Plate 1034 


Photo B. G. Schubert 


QUERCUS LAURIFOLIA: portions, X 1, of type of Q. rhombica, var. obovatifolia 
"argent. 


1946] Fernald,—Identifications of North American Plants 143 


green oak), et dans un autre endroit, Quercus dentata s. hemisphoerica. 

On doit la rapporter à notre Chêne aquatique. Voyez BARTRAM 

Travels, p. 14, 28 et 320. 

Referring to Bartram, p. 14, one finds an account of “Our 
turkey of America" and nothing about any oak; but on p. 24 
(probably intended by Michaux) there is a list of the trees and 
shrubs “growing on the banks of this sequestered river" which 
includes the undefined name Q. dentata. Page 28 yields, as 
growing near Savannah, “5. Q. dentata", which in a footnote 
is explained as “5. Narrow-leaved Wintergreen Oak”; and on 
p. 320 a long list includes “Quer. dentata, s. hemispherica". 
The last entry, Quercus dentata or hemispherica, makes the names 
alternative and therefore illegitimate, even if they had been 
accompanied by diagnoses, which they lacked. Index Kewensis 
solved the identity of these two alternative names by stating that 
one “= aquatica", the other “= nigra". This is one way out of 
the problem. The usually very cautious Stephen Elliott in- 
cluded in his Sk. Bot. S. C. and Ga. ii. 596 (1824), under Q. 
laurifolia the tree which Willdenow had properly validated as 
Q. hemisphaerica, with “Leaves nearly perennial, sessile, oblong- 
lanceolate, nearly acute, tapering at base, entire, glabrous on both 
surfaces" ; and he cited in the synonymy Q. hemispherica Bartram, 
and in addition to Michaux's two plates of true Q. laurifolia 
“perhaps also t. 20. f. 2". His more detailed account of the tree 
with “a large handsome hemispherical head" with leaves “very 
glabrous on both surfaces . . . , those of the young plant toothed 
and irregularly sinuate. . . . Cup shallow, nearly sessile", was 
of Q. hemisphaerica, rather than of Michaux's Q. laurifolia. It is 
probable that Elliott’s misidentification in 1824 set the standard 
by which later authors have misidentified Q. laurifolia of 
Michaux. Elliott’s further remarks are illuminating: 

The figure in Mich. Querc. t. 20. f. 2. exactly resembles the young 
plants of this species. And as this oak, though growing in dry soils, is 
more known by the name of “Water Oak," than by any other appella- 
tion, it is not impossible that Michaux may have been misled by its 
popular denomination to insert a figure of it among the real Water Oaks. 

I have always considered this as the real Q. Hemisphaerica of Bar- 
tram. It certainly is the species to which his [i. e. Willdenow’s] de- 
scription most appropriately applies. 

Grows in rich sandy soils along the margin of swamps, appearing to 


take the place of the live oak as you leave the margin of the ocean, but 
growing also with the live oak on the sea-islands. 


144 Rhodora [JULY 


I realize that by Index Kewensis and by Sargent, Silva, vill. 165 
(1895) Q. hemisphaerica is reduced, without a word of explanation, 
to Q. nigra L. To be sure, Trelease in his American Oaks, Mem. 
Nat. Acad. Sci. xx. 160 (1924), speaking of Willdenow’s her- 
barium, said “forms occur as representatives of Q. hemisphaerica 
as he meant to use that name", but that these are the hetero- 
phyllous sprouts of several species. Willdenow may have had 
mixed material (and probably did, just as did Linnaeus for many 
species) but his diagnosis, “Q. foliis perennantibus oblongo- 
lanceolatis indivisis trilobis sinuatisque lobis mucronatis utrinque 
glabris. W.", and his basing of the species on Michaux's interpre- 
tation above quoted and Michaux's t. 20, fig. 2, leave no doubt as 
to what he primarily meant. Trelease’s suggestion that Q. hemi- 
sphaerica might have been sprouts of Q. nigra or of Q. Phellos 
does not dispose of Willdenow’s definite “foliis perennantibus ” 
nor of the Michaux figure, for both Q. nigra and Q. Phellos have 
promptly deciduous leaves. 

Sargent and, after him, Trelease, also cite in the synonymy of 
Quercus nigra “Quercus hemisphaerica, var. nana, Nuttall, Gen. 
ii. 214 (1818)", although Nuttall had no such variety under Q. 
hemisphaerica. Instead, this is what he said: “11. hemispherica, 
Bartram. Willd. also Q. aquatica, Willd. Gà. nana, Q. nana, Willd." 
Willdenow having published as a species Q. nana, Nuttall's Q. 
aqualica, var. nana was a new combination (but not under Q. 
hemisphaerica). If we do not take up Q. hemisphaerica Bartram 
ex Willd. for the evergreen or partly evergreen species of the 
outer Coastal Plain of the South, we must coin à new name. 
Whoever does that will have some difficulty in demonstrating 
that André Michaux in 1801, Willdenow in 1805 and Elliott in 
1824 were all wrong. "There has been trouble enough; no good 
can come from making more. One whole precious week has been 
lost in working out these unedifying details. That is enough. 

Professor W. C. Coker in Journ. Elisha Mitchell Sci. Soc. 
xxxii. plates 2-4 (1916) shows some beautiful illustrations of 
Quercus hemisphaerica (as the traditional “laurifolia’”’). The 
name hemisphaerica, when applied to these photographs, seems 
not to need the restrictive first two syllables. In the trade this 
tree has become known as ‘Darlington Oak", because of its 
early introduction as an evergreen ornamental to the streets of 


1946] Fernald,—Identifications of North American Plants 145 


Darlington, South Carolina. Professor Coker’s account of the 
persistence of the foliage of ‘‘one of the handsomest and most 
ornamental oaks to be found in any country” should be repeated: 


One of the most remarkable qualities of the Laurel Oak is its habit 
of holding its leaves through the entire year throughout most of its 
range. Towards its northern limit and when planted outside of the 
coastal plain this habit is modified to a varying degree, depending on 
the climate. In the low country of South Carolina and along the gulf 
to New Orleans the tree is nearly or completely evergreen, but at 
Darlington and Hartsville, S. C., the leaves fall slowly through the 
winter, usually beginning at the tips of the branches and proceeding 
inwards, so that by February or March only the center remains decid- 
edly green, with scattered green leaves in the peripheral parts.—Coker, 
l. c. 40. 


Back TO CAREX ROSTRATA.—In RuHopora, xliv. 324-331, 
plates 715 and 716 (1942) I felt it necessary, in following strict 
priority, to abandon the long-familiar name Carex rostrata 
Stokes (1787) for the earlier C. inflata Hudson (1762 in part, 
emend. 1778). This most unfortunate change of name for one 
of the most common and generally known Carices of cool- 
temperate regions around the northern hemisphere was induced 
by the decision of the late A. B. Rendle and James Britten in 
1907, that the change was unavoidable. The decision of these 
usually quite accurate gentlemen, neither of whom understood 
Carex, was further argued and accepted by those most careful 
students of nomenclature, Schinz & Thellung (1908) and Mans- 
feld (1938), as well as the less cautious Druce and many others 
in Europe. There seemed no escape from C. inflata. 

Now, however, the most accurate of recent British students of 
Carex, Mr. E. Nelmes of Kew, has given a very detailed analysis 
of Hudson's C. inflata in Journ. Bot. lxxx. 109-112 (1942), 
Hudson's original C. inflata of 1762 having been a sad mixture, 
clarified by his redefinition of it as a single specific element in 
1778. Approaching the problem with full understanding of 
British Carices, Nelmes considers every possibility and, most 
happily, concludes: * From the available evidence it seems to me 
that Carex inflata Huds., Fl. Angl., ed. 1, cannot be identified 
with any one species: it may be C. vesicaria L., or, less likely, C. 
rostrata Stokes, or, least likely, C. laevigata Sm.; but it is most 
likely to represent a mixture. Carex inflata Huds., Fl. Angl., ed. 


146 Rhodora [JULY 


2, is almost certainly C. laevigata 8m.". Everyone who knows 
Carex will give a sigh of relief. The four varieties recognized by 
me and illustrated in 1942 should now be called 


CAREX ROSTRATA Stokes in Withering, Brit. Pl. ed. 2, ii. 1059 
(1787). C. ampullacea Gooden. in Trans. Linn. Soc. ii. 207 
(1794). C. inflata sensu Rendle & Britten in Journ. Bot. xlv. 
444 (1907), not Hudson, Fl. Angl. 354 (1762), emend. ed. 2: 412 
(1778). 

Var. anticostensis (Fernald), comb. nov. C. inflata, var. 
anticostensis Fernald in Rnopona, xliv. 329, pl. 715, figs. 5 and 6 
(1942). 

Var. UTRICULATA (Boott) Bailey in Proc. Am. Acad. xxii. 67 
(1886). C. utriculata Boott in Hook. Fl. Bor.-Am. ii. 221 (1839). 
C. utriculata, var. minor Boott, l. c. (1839); and many other 
synonyms cited by me in RmHopona, xlvi. 330 (1942), these 
including C. inflata, var. utriculata (Boott) Druce in Bot. Soc. & 
Exchange Club Brit. Isl. Rep. ix. 141 (1930). 

Var. AMBIGENS Fernald in Ruopora, iii. 51 (1901). C. inflata, 
var. ambigens (Fernald) Fernald l. c. xliv. 330, pl. 715, figs. 7 
and 8 (1942). 


LEMNA VALDIVIANA, Not L. cvcLosTAsA.— Although in his 
Revision of the North American Lemnaceae, printed in advance, 
Nov. 1, 1897, from the Ninth Ann. Rep. Mo. Bot. Gard. (1898), 
Dr. C. H. Thompson took up the name Lemna cyclostasa for one 
of the most distinct members of the genus, the authors of Gray, 
Man. ed. 7 (1908), rejected that name in favor of the validly pub- 
lished L. valdiviana Philippi (1864). Had Thompson refrained 
from precipitate haste to get out his Revision “in Advance" and 
verified his bibliography it would have been well; for, even now, 
some authors of volumes on aquatics, in spite of the correction 
made in ed. 7 of the Manual, still use the name L. cyclostasa. 
Thompson’s bibliography was as follows: 

LEMNA cvcLosTASA (Ell) Chev. Fl. Par. 2: 256. 1827. Schleid. 

Linnaea. 13: 390. 1839. Lemna minor var.? Cyclostasa Elliott, Bot. 8. 

Carol. and Ga. 2: 518. 1824. L. Valdiviana Ph. Linnaea. 33: 239. 


1864. L. Torreyi Aust. in Gray, Man. Bot. 479. 1867. [5th ed.]. L. 
abbreviata Hglm. Engler's bot. Jahrb. 213: 298. Jan. 1895. 


Had Thompson been less content with the lax and unverified 
type of bibliography which characterizes so much of the work of 


his sponsor, Trelease, he would have searched in vain in Cheva- 
lier's Flore générale des Environs de Paris for any mention what- 


Rhodora Plate 1035 


Photo B. G. Schubert 


QUERCUS HEMISPHAERICA Bartram ex Willd., both figs. X 1: FIG. 1, the original 
figure of Michaux, cited by Willdenow; Fic. 2, leaves of young tree of Q. laurifolia 
sensu most authors, not Michaux, from Florida 


Rhodor: Plate 1036 


Photo B. G. Schubert 


QUERCUS HEMISPHAERICA, both figs. X 1: ric. 1, fruiting branch from Georgia of Q. 
laurifolia sensu most authors, not Michx.; FIG. 2, cups and acorn from Florida. 


1946] Fernald,—Identifications of North American Plants 147 


ever of L. cyclostasa, a species unknown in Europe. Chevalier 
had the regular European L. trisulca, polyrrhiza, arhiza, minor 
and gibba but not L. cyclostasa of tropical and warm-temperate 
America. Had he looked at Schleiden’s Monograph of Lemna- 
ceae in Linnaea, xiii. 390 (1839) he would have found under L. 
minor the synonym “L. cyclostasa Elliot". Elliott had no such 
species but he did describe L. minor, var. ? cyclostasa; Schleiden 
incorrectly attributed the binomial to him but, since the binomial 
was published in synonymy, it is invalid. Presumably Thompson 
copied his references directly from Index Kewensis, which gives 
“eyclostasa, Ell. ex Chev. Fl. Par. ii. 256; Schleid, in Linnaea, 
viii. (1839) 390 — minor". But everyone with much experience 
knows that the earlier volumes of Index Kewensis must be taken 
as very untrustworthy guides. If anyone can find L. cyclostasa 
in Chevalier, he will make a clarifying discovery; but the reduc- 
tion by Index Kewensis of the species to L. minor shows equal 
lack of understanding. As to Thompson’s reference to L. 
abbreviata Hegelm., consultation of his Systematische Übersicht 
der Lemnaceen in Engler's Jahrb. (1895) immediately shows that 
Hegelmaier was not treating it as a species. Unfortunately, 
Hegelmaier adopted a misleading method of designating his 
varieties, by enclosing the specific name and “var.” in parenthe- 
ses, just as in this country there are always some nonconformists, 
who refuse to follow conventional practices. Hegelmaier gave 
numbers to all recognized species, “8. Lemna angolensis Welw.’’, 
"9. Lemna valdiviana Philippi", “10. Lemna minima Phil.", 
but under, 9. L. valdiviana, he had, without numbers “Lemna 
(valdiviana var.) abbreviata" and ‘Lemna (valdiviana var.?) 
platyclados". L. valdiviana, var. abbreviata was not intended as 
à species. 

It has seemed important to dissect this case, in view of the 
faith trustingly reposed in “the books" by those who follow the 
uncritical American principle of accepting anything they see in 
print—a method not creditable when extended, as it so regularly 
is, into scientific bibliography. LEMNA VALDIVIANA Philippi has 
right-of-way until it is demonstrated that it is antedated by a 
validly published binomial. 


148 Rhodora [JULY 


STENANTHIUM IN THE EASTERN UNITED STATES (PLATES 
1037-1041) .— 

STENANTHIUM GRAMINEUM (Ker) Morong, var. robustum 
(S. Watson), stat. nov. S. robustum 8. Watson in Proc. Am. Acad. 
xiv. 278 (1879). PrATES 1039 and 1040. 

S. GRAMINEUM, var. micranthum, var. nov. (TAB. 1041), 
caule 0.25-1 m. alto, basin versus 1.5-5 mm. diametro; foliis 
subrigidis corrugatis 0.4—1 cm. latis; perianthiis 3-4.5 (-5) mm. 
longis.— Upland woods, western Virginia and eastern Tennessee 
to northwestern Florida and Alabama. Tyre from open woods, 
Caesar’s Head, South Carolina, July 28, 1881, John Donnell 
Smith (Herb. Gray.). 

The so-called specific distinctions between Stenanthiwm 
gramineum and S. robustum have proved to be very evasive, 
Britton in Britton & Brown, Ill. Fl. ed. 2: i. 489 (1913) giving 
seemingly good differences, his key reading: 


“Leaves 2'"-3' wide; capsule reflexed............-..--. 1. S. gramineum. 
Leaves 3’’-10” wide; capsule erect............0-00-55- 2. S. robustum." 


In the illustrations, however, these characters are not very 
definitely shown, the figures showing flowering pedicels spreading 
in both and no contrast in flowers or fruit, while the comment 
under S. robustum implies lack of full conviction: * Apparently 
distinet from the preceding species, though closely related". 
Small, Man. 277 (1933), gives a key which looks good: 


“Perianth white: capsule deflexed, about 8 mm. long..... 1. S. gramineum. 
Perianth green: capsule erect, fully 10 mm. long....... 2. S. robustum." 


In both cases the ranges are given as very extensive, S. robustum 
said by Small to grow in “Acid swamps, various provinces, 
S[outhern] S[tates] to Ark., Ohio, and Pa.". 

When, however, the material of the Stenanthium of the eastern 
United States is even casually studied the keys are found not to 
unlock the door. The type of S. gramineum is Helonias graminea 
Ker in Curtis's Bot. Mag. xxxix. t. 1599 (Nov. 1, 1813), Ker 
there describing the “flowers small, . . . white, suffused with 
purple on the outside", and very many specimens, otherwise 
quite like Ker's plate, have the perianth green or bronze to 
purplish, as well as whitish. Furthermore, Watson's original 
description of S. robustum clearly said ‘‘perianth-segments 3 or 
4 lines long, white or green". As to the “erect” capsules of the 


1946] Fernald,—Identifications of North American Plants 149 


latter, as opposed to “capsule deflexed” in the former, it is not 
reassuring to one’s confidence in the character, to examine a 
series of S. robustum and then one of the usually smaller-flowered 
S. gramineum. Watson seems to have picked out to constitute 
his S. robustum stout and broad-leaved specimens with flowers 
the largest in the series, these largest plants having ‘been orig- 
inally included in the last", the “last” being the more slender and 
narrower-leaved plants with perianth “2 or 3 lines long" and 
called by Watson “S. ANGUSTIFOLIUM, Gray", based on Veratrum 
angustifolium Pursh. Pursh had the plant which, shortly before, 
had been described by Ker as Helonias graminea, Pursh not 
realizing that this plant was later to be distinguished by Small 
on account of its “Perianth white", for Pursh brazenly wrote 
“Flowers greenish yellow". Pursh’s species came from “high 
mountains of Virginia and Carolina" and a specimen (with an 
unpublished name) collected by him and bearing his label 
“Veratrum [unpublished trivial name somewhat equivalent to 
angustifolium], P. Salt pond mountain", is in his herbarium at the 
Academy of Natural Sciences of Philadelphia. This, with 
perianths obviously herbaceous (not white and petaloid), and 
about 6 mm. long, which may pass as presumably the type, has 
the capsules nearly erect, not at all ''deflexed". Similarly, 
plants which have the medium-sized perianth and firm narrow 
leaves of true S. gramineum may have the capsules as erect as in 
theoretically ideal S. robustum (PLATE 1039, ria. 3). Such speci- 
mens of S. gramineum are shown in PLATE 1037, FIG. 3, from 
Pisgah Forest, North Carolina, House, no. 4040. Others, with 
the pedicels no more erect than in some Pennsylvanian specimens 
of S. robustum (PLATE 1040, ric. 3, from New Texas, Lancaster 
County). Such fruiting racemes are shown in PLATE 1038, this 
specimen from Springfield, Ohio. 

Most unfortunately Watson designated no type for his Sten- 
anthium robustum and, still more unfortunately, he left others to 
find out just what he intended by his species of ‘‘ Pennsylvania to 
Ohio, Tennessee and South Carolina." A single sheet only 
bears his inscription “S. robustum, n. sp." in a delicate pencil- 
notation easily overlooked. This, PLATE 1039, Fics. 1 and 2, 
was from Sligo Furnace, Clarion Co., Pennsylvania, Aug. 1859, 
J. R. Lowrie (distributed by Porter) and must stand as the 


150 Rhodora [JULY 


TYPE. As will be seen from the photograph, the capsules on the 
lateral racemes are on almost as divergent pedicels as those of 
S. gramineum (PLATE 1038, FIG. 3). I get little satisfaction out 
of “Perianth green: capsule erect" as opposed to “Perianth 
white: capsule deflexed”. 

Taking the one sheet which Watson marked S. robustum as the 
type, the stout and very leafy stem and the broad leaves, accom- 
panied by perianths 8 or 9 mm. long, set the standard; and it at 
once becomes apparent that such plants, with bases of stem up 
to 1.5 em. thick (as opposed to a maximum of 1 em.), with larger 
leaves 1-3 em. broad (as opposed to 4-15 mm.) have the very 
smooth and translucent leaves thin or membranaceous and with 
the very many (up to 61) nerves immersed in the tissue (PLATE 
1039, rias. 2 and 4, PLATE 1040, rra. 2). In the usually more 
southern S. gramineum and var. micranthum, on the other hand, 
the firm or coriaceous and mostly narrow leaves have prominent 
elevated ribs, these giving the upper leaf-surface a closely fur- 
rowed or corrugated appearance when dry (PLATE 1037, ria. 2, 
PLATE 1038, FIGS. 2 and 6, and PLATE 1041, FIG. 2)!. Occasionally 
plants with all other characters of typical S. gramineum (PLATE 
1038, Fic. 3) will diverge in foliage, showing thin but narrow 
leaves (FIG. 4) too strongly approaching those of S. robustum. 
The failure of this last character, so hopefully worked out, leaves 
nothing absolute to separate the larger-flowered S. gramineum 
and the smaller-flowered S. robustum as species. 

Turning in the other direction, we find along the mountains 
from western Virginia and eastern Tennessee southward a 
series of slender-stemmed and narrow-leaved plants with flowers 
strikingly smaller than in typical S. gramineum. The perianths 
are either herbaceous or petaloid in texture, sometimes greenish 
or purplish, sometimes white, and the sepals and petals vary 
from lance-attenuate to linear-oblong and bluntish. The inflor- 
escence is strongly suggestive of smaller panicles of typical S. 
gramineum. This plant evidently passes through its larger- 
flowered individuals into the smaller-flowered extreme of 5. 
gramineum. Iam calling it var. micranthum (PLATE 1041). 

1 These differences stand out sharply, without aid of a hand-lens, through the 
reading-glasses prescribed after the removal of a cataract. 'Thelate K. K. Mackenzie 


boasted of his ''microscopic vision". It is hoped the the present writer will not see 
what others can not visualize! 


Rhodora Plate 1037 


Photo B. G. Schubert 


STENANTHIUM GRAMINEUM, var. TYPICUM: FIG. 1, portion of flowering plant, X 1; 
FIG. 2, portion of leaf, X 2; rra. 3, portion of fruiting plant, X 1. 


Rhodora Plate 1038 


Photo B. G. Schubert 


STENANTHIUM GRAMINEUM, var. TYPICUM: FIG. 1, portion of inflorescence, X 1; FIG. 
2, portion of leaf, X 2; FIG. 3, portion of fruiting panicle, X 1; FIG. 4, portion of leaf, 
X 2; ria. 5, terminal fruiting raceme, X 1; FIG. 6, portion of leaf, X 2. 


Rhodora Plate 1039 


Photo B. G. Schubert 


STENANTHIUM GRAMINEUM, var. ROBUSTUM: FIG. 1, portion, X 1, of TYPE of S. robus- 
, I , 

tum S. Watson; FIG. 2, portion of leaf, X 2, from TYPE; FIG. 3, portion of terminal 

fruiting raceme, X 1; FIG. 4, portion of leaf, X 2. 


Rhodora Plate 1040 


Photo B. G. Schubert 


STENANTHIUM GRAMINEUM, var. ROBUSTUM: FIG. l, portion of inflorescence, X 1; 
FIG. 2, portion of leaf, X 2; ria. 3, summit of inflorescence, X 1; FIG. 4, terminal 
flowering raceme, X 1. 


1946] Fernald,—Identifications of North American Plants 151 


Thus, from the coarse and relatively northern S. gramineum, 
var. robustum through the commoner and much wider-ranging 
true S. gramineum, thence on to the tiniest of the series, the 
southern var. micranthum, we have a confluent series, the two 
most extreme variations with distinctive areas of development. 
I see no way but to treat them as three geographic varieties. 
The material in the Gray Herbarium alone suggested three 
probable species. The large representation in the United States 
National Herbarium, the herbarium of the Academy of Natural 
Sciences of Philadelphia, and that of the New York Botanical 
Garden, for the use of which I am greatly indebted to Drs. 
Maxon, Pennell and Seaver, respectively, have destroyed all my 
illusions! As the best I can now do I propose the following defi- 
nition of the three varieties. 

a. Stem (dry) 4-10 mm. thick at lowest exposed internode; 

leaves rather crowded below, rapidly diminishing be- 
low panicle, firm to coriaceous, mostly opaque, the 
larger ones 4-15 mm. wide, their prominently raised ribs 
producing a corrugated surface; panicle lax, the branches 
distant, the flowers mostly subremote along the often 
flexuous branches; perianth 3-8 (-10) mm. long; cap- 
sules ovoid-urceolate, 6-9 mm. long, on spreading to 


reflexed pedicels; seeds 5-5.5 mm. long. 
Stem 0.5-1.9 cm. high, 4-10 mm. thick at base; perianth 


S 10 mm. longi — 4. pt sae a S. gramineum var. typicum. 
Stem 0.25-1 m. high, 1.5-5 mm. thick at base; perianth 
B. 5.05) mum dang... ev esI I es Var. micranthum. 


a. Stem (dry) 7-15 mm. thick at lowest exposed internode, 
up to 1.8 m. high; leaves crowded and numerous nearly 
up to panicle, thin and membranaceous, translucent, 
the larger ones 1-3 cm. wide, their ribs mostly immersed 
in the tissue; panicle usually dense, with flowers crowded 
along the stiffly ascending branches; perianth 5-10 mm. 
long; capsules oblong-subcylindrie, 9-10 mm. long, 
crowded and ascending to horizontally spreading; seeds 
TO mmdong 5 cts ay ec NEL E.A Var. robustum. 


S. GRAMINEUM (Ker) Morong, var. typicum. S. gramineum 
(Ker) Morong in Mem. Torr. Bot. Cl. v. 110 (1894). Helonias 
graminea Ker (Gawler) in Curtis’s Bot. Mag. xxxix. t. 1599 
(Nov. 1, 1813). Veratrum angustifolium Pursh, Fl. Am. Sept. i. 
242 (1814). Xerophyllum gramineum (Ker) Nutt. Gen. i. 236 
(1818). Veratrum (subgen. Stenanthium) angustifolium (Pursh) 
Gray in Ann. Lyc. N. Y. iv. 120 (1837). S. angustifolium 
(Pursh) Kunth, Enum. iv. 190 (1843). S. angustifolium, *S. 
gramineum (Ker) Kunth, l. c. (1843).—Rich woods, thickets and 
borders of bottoms, northwestern Pennsylvania to Illinois and 
Missouri, south to southeastern Virginia, upland to northwestern 


152 Rhodora [JULY 


Florida and Alabama, and to Louisiana and eastern Texas. 
PraATES 1037 and 1038. 

Var. MICRANTHUM Fernald, supra.— Upland woods, western 
Virginia and eastern Tennessee to northwestern Florida and 
Alabama. PLATE 1041. 

Var. RoBUsTUM (S. Watson) Fernald, supra. S. robustum 8. 
Watson in Proc. Am. Acad. iv. 278 (1879).—Southeastern Penn- 
sylvania, Maryland and District of Columbia to Indiana. 
PrATES 1039 and 1040. 


Tue IDENTITY OF SISYRINCHIUM ANGUSTIFOLIUM (PLATES 
1042-1044).—In 1865, in the 4th edition of his Manual, Asa 
Gray recognized one species of Sisyrinchium in eastern North 
America, the all-inclusive S. Bermudiana L., with two varieties: 
var. anceps (S. anceps Cav.) with “broadly winged scape” and 
var. mucronatum (S. mucronatum Michx.) “with slender and 
narrowly winged scape”. By the 6th edition of the Manual 
Watson (and Coulter), conscious that the continental North 
American plants are not the restricted S. Bermudiana of Bermuda, 
recognized two species: S. angustifolium Mill., with “Scape . . . 
simple, the spathe solitary and terminal", and S. anceps Cav. 
with “Scape .. . usually branching and bearing 2 or more 
peduncled spathes". Then came Bicknell’s amazing revelation 
that in temperate North America there are very many morpho- 
logically and geographically segregated species, although Bicknell 
went much farther in his splitting than others have been able to 
follow. Nevertheless, in Atlantic North America Bicknell 
clearly differentiated at least a score of species which all careful 
students must recognize. So absorbed, however, was Bicknell in 
his newly discovered small world that he did not carefully trace 
to their sources some of the earlier-named plants. 

One of his earliest papers, his Studies in Sisyrinchium—IIT: 
S. angustifolium and some related Species new and old, in Bull. 
Torr. Bot. Cl. xxvi. 335-349 (1899), started with the basic as- 
sumption that the identity of the first segregate from the Linnean 
all-inclusive S. Bermudiana was settled. His S. “angustifolium 
Mill.", with three synonyms (besides S. Bermudiana L. in part), 
S. gramineum Lam. (1783), S. anceps Cav. (1788) and S. mon- 
tanum Greene (1899), was described as "stiff and erect . . . 
Leaves . . . 1.5-2.5 mm. wide (1-3.5 mm.) . . . : stems simple 
and leafless, or occasionally bearing a single leaf subtending 1 or 2 


1946] Fernald,—Identifications of North American Plants 153 


branches . . . , 1-2 mm. or even 3 mm. wide, wing-margined 

spathes erect, green or sometimes purplish-tinged; outer 
bract 2-6 em. long, surpassing the inner one 1.5—4 cm., rarely less 
than twice its length, . . . clasping for 2-6 mm. at base . . . ; 
interior scales silvery white, narrow, usually about half the length 
of the inner bract: flowers 1-8, violet-blue; perianth 10-12 mm. 
long; . . . pedicels erect or nearly so, 17-25 mm. long, shorter 
or slightly longer than the inner bract: capsules 4-6 mm. high, 

. . pale, but often clouded with brownish-purple". 

Then Bicknell continued: ‘This species is far more widely 
distributed than any other one of its genus, ranging from New- 
foundland and New Jersey to Saskatchewan and Montana and 
southward along the eastern mountains to Virginia [only Moun- 
tain Lake, 4000 ft. alt. cited] and in the west to southern Colora- 
do". In his introductory paragraph Bicknell had said: ‘‘The 
common Blue-eyed Grass of the eastern states, Sisyrinchium 
angustifolium Miller, may be taken as representative of a section 
of the genus Sisyrinchium, embracing those species having 
simple leafless stems with terminal spathes . . . the simple- 
stemmed species are, as a group, of more northern and alpine 
distribution than those which develop pedunculate spathes from 
one or more leaf-bearing nodes, while, on the other hand, the 
species having a definitely compound system of branching are 
all distinctive southern". 

Thus was set up for all of us, who have found it easier to follow 
the course of least resistance, a standard misconception of what 
Miller was supposed to have meant by Sisyrinchium angusti- 
folium, what Lamarck had as S. gramineum and Cavanilles as S. 
anceps, although Miller and Cavanilles had described or shown 
the spathes as peduncled and all had cited or illustrated plants 
with peduncles, and with pedicels far from “erect or nearly so, 

. Shorter or slightly longer than the inner bract" of Bicknell’s 
S. “angustifolium”. 

Starting with Sisyrinchium angustifolium Miller, Gard. Dict. 
ed. 8, sp. no. 2 (1768), we get a brief account of a plant cultivated 
in England, with peduncles and small pale blue flowers: 

2. SisyniNCHIUM (Angustifolia) foliis lineari-gladiolatis pedunculis 
longioribus [than in the preceding S. Bermudiana]. Sisyrinchium with 


linear sword-shaped leaves, and longer foot-stalks to the flower [longer than 
in true S. Bermudiana]. Bermudiana graminea, flore minore caeruleo. 


154 Rhodora [JULY 


Hort. Elth. 49. Grass-leaved Bermudiana with a smaller blue flower. 


The second sort grows naturally in Virginia; this hath a perennial 
fibrous root, from which arise many very narrow spear-shaped leaves 
about three inches long, and searce an eighth part of an inch broad, of a 
light green colour, and entire. The stalks rise about three inches high; 
they are very slender, compressed and bordered like those of the first 
[S. Bermudiana], and have short, narrow, sword-shaped leaves, whose 
base embrace them; they are terminated by two small pale blue flowers, 
inclosed in a two-leaved sheath, standing upon longer foot-stalks than 
those of the other. 


The identification by Miller of his Sisyrinchium angustifolium 
with long peduncles and small pale blue flowers on “longer” 
foot-stalks, with the plant described and illustrated by Dillenius 
as cultivated in the English gardens in 1732, has real significance, 
although Bicknell did not note it, just as he apparently did not 
even read Miller's description in Gard. Dict. ed. 8 (1768), the 
first place of publication of the binomial S. angustifolium. 
Instead, Bicknell started S. angustifolium from the polynomial 
ed. 7 (1759), where neither the name nor the adjective angusti- 
folium is found. Here is what Miller gave in 1759: 


2. SiSYRINCHIUM foliis lineari-gladiolatis, pedunculis longioribus. 
Sisyrinchium with linear Sword-shaped Leaves, and longer Foot Stalks 
to the Flower. This is the Bermudiana graminea, flore minore caeruleo. 
Hort. Elth. 49. Grass-leaved Bermudiana with a smaller blue Flower. 


This nonbinomial account, which alone was cited by Bicknell, 
was followed by the same statement as in ed. 8, that the plant 
“grows naturally in Virginia’, etc. 

Now, if we turn to the basic treatment by Dillenius, in his 
Hortus Elthamensis, 49, t. XLI, fig. 49 (1732) we find his Ber- 
mudiana graminea, flore minore caeruleo described, not merely as 
“three inches high", as evidently misquoted by Miller, but up to 
a foot high: “PLANTA est palmaris, dodrantalis & pedalis"; the 
leaves full green (“folia . . . saturanter viridia"); the stem 2- 
forked (“Caulis . . . in summitate bifariam dividatur"). The 
plate shows as fig. 48 a plant of true Sisyrinchium Bermudiana, 
with erect and but slightly exserted pedicels, as described by 
Bicknell for S. angustifolium sensu Bickn., not Mill.; and as fig. 
49 (our PLATE 1042, ric. 1) Bermudiana graminea, flore minore 
caeruleo, two plants, the taller 1.6 dm. high and with forked scape, 
the other with simple scape. Both show narrow sepals and petals 


Rhodora Plate 1041 


Photo B. G. Schubert 


STENANTHIUM GRAMINEUM, var. MICRANTHUM: FIG. 1, inflorescence, X 1, of TYPE; 
FIG. 2, portion of leaf, X 2; ric. 3, portion of fruiting plant, X 1. 


Rhodora Plate 1042 


SISYRINCHIUM ANGUSTIFOLIUM Mill.: FIG. 1, TYPE, 1. e. Bermudiana gramineo, flore minore 
caeruleo Dillenius, t. xli. fig. 49, X 1, after Dillenius; ria, 2, S. anceps Cav., X 1, after 
Cavanilles. 

S. MUCRONATUM Michx.: FIG. 3, Sisyrinchium caeruleum parvum, gladiato caule, Virgini- 
anum, after Plukenet, cited by Lamarck under S. gramineum Lam., in part, not Mill. 


1946] Fernald,—Identifications of North American Plants — 155 


5-10 mm. long and, most important of all, slender fruiting pedicels 
reflexed or recurving from the spathes, quite as in S. graminoides 
Bicknell, Bull. Torr. Bot. Cl. xxiii. 133, pl. 263 (1896), for Bick- 
nell's plate (our PLATE 1043, FIGs. 1 and 2) shows bifurcate and 
simple stems from the same base, quite as did that of Dillenius, 
and 2-4 arched to recurving elongate pedicels, while his descrip- 
tion of his S. graminoides contains the following items: plant 6 
inches to 2 feet high (up to a foot acc. to Dillenius), green or 
somewhat glaucous (full green acc. to Dillenius); leaves and stem 
“mostly 11$"-2" wide" (Dillenius showed them approximately 
the same, 3-4 mm.); “Stem dividing . . . into two... 
branches. Not infrequently simple and leafless scapes rise 
among the normally branched ones, simulating the stem of S. 
angustifolium” (Dillenius said “Caulis . . . in summitate bifari- 
am dividatur" but, like Bicknell, showed also a simple stem); 
“Capsules . . . disposed to be spreading or even recurved on 
slender pedicels" (Dillenius showed them definitely so), not 
erect or nearly so as shown in S. angustifolium sensu Bicknell 
(our PLATE 1043, Fra. 3), not Miller. 

It is fortunate that Dillenius gave a good illustration of 
Sisyrinchium graminoides Bicknell (as Bermudiana, flore minore 
caeruleo), for in his discussion Dillenius refers to it, as a “figura 

. vitiosa exhibita", the Sisyrinchium | caeruleum parvum 
gladiato caule Virginianum of Plukenet, Phyt. t. lxi. fig. 1 (our 
PLATE 1042, ric. 3), for the latter figure, although *'vitiosa" for 
the Dillenian species, was a good one of the common Virginian S. 
mucronatum Michx.!, the type of which I studied in 1903 and 
which is definitely the plant we now know under that name. 

Next among the older species included by Bicknell in his 
usually simple- and winged-stemmed and relatively northern 
Sisyrinchium “angustifolium’’, with erect or nearly erect short 
pedicels is S. gramineum Lam. Encycl. i. 408 (1783). Lamarck 
gave a clear description of a Virginian plant which can hardly be 
waved aside, as one might be tempted to do, and he also gave as 
synonyms the conventional S. Bermudiana var. a L., which 
included at least the Dillenian plant, already discussed, and the 
Plukenet illustration (our PLATE 1042, rra. 3) of a Virginian plant, 
which is evidently S. mucronatum Michx. (1803). That La- 
marck in 1783 had before him a really new species is possible, 


156 Rhodora | [JULY 


from his description. That it was the plant of Plukenet’s t. 61, 
fig. 1 is seemingly apparent and that it was the characteristic 
plant, common at least in eastern Virginia, which in 1803 Michaux 
described from Pennsylvania as S. mucronatum seems possible 
from Lamarck’s very clear description: 


1. BERMUDIENNE graminée, Sisyrinchium gramineum. Sisyrinchium 
caule simplici alato, spathis inaequalissimis flores superantibus. N. 
Sisyrinchium angustifolium. Mill. Dict. n°. 2. Sisyrinchium caeru- 
leum parvum, gladiato caule, Virginianum. Pluk. Alm. 348. t. 61. 
f. I. Bermudiana graminea, flore minore caeruleo. Dill. Elth. 49. t. 
41. f. 49. Sisyrinchium Bermudiana. Lin. var. a. 

Les feuilles de cette plante sont très-étroites, linéaires, graminées, 
lisses ou sans nervures bien remarquables, & s’engainent à leur base par 
le côté, comme celles des Iris: les tiges sont presque filiformes, simples 
pour l'ordinaire, comprimées, bordées dans leur longueur de deux 
petites alles ou membranes courantes, & hautes de six ou sept pouces. 
Chaque tige est terminée par deux écailles spathacées, fort inégales, 
l'extérieure étant une fois plus longue que l'autre, & dépassant toujours 
les fleurs, qui sont petites, bleuátres, & communément au nombre de 
deux. Cette plante croît naturellement dans la Virginie; on la cultive 
au Jardin du Roi. J}. (v. v.) 


In Virginia the only species of Sisyrinchium known with con- 
sistently unforking stems are S. “angustifolium” sensu Bicknell, 
already somewhat discussed, and in Virginia definitely montane; 
S. mucronatum Michx., frequent; and the southern and capillary- 
leaved S. capillare Bicknell, a very rare and very distinct species 
with paired sessile spathes, of flat pineland or bog of the Coastal 
Plain, only twice found in the southeasternmost counties. That 
S. gramineum of Lamarck could have been S. mucronatum is 
evident from the slender (“almost filiform") stems bordered by 
2 little wings; whereas, if it were S. “angustifolium” sensu Bick- 
nell, which is less likely to have reached the Jardin du Roi from 
Virginia, we should expect less emphasis on the slenderness. 
Whether Lamarck had either of these becomes, perhaps fortu- 
nately, unimportant, since he cited as an unquestioned synonym 
S. angustifolium Mill. Dict. no. 2. By the International Rules 
Lamarck's S. gramineum is, therefore, illegitimate because 
(Art. 60!) “there was a valid name for the group to which it was 
applied, with its particular circumscription, position and rank". 
Thus the identity of the plant actually described by Lamarck 
becomes an academic question only. S. MucronatuM Michx., 
published without entangling bibliography, stands. 


1946] Fernald,—Identifications of North American Plants 157 


Returning to Bicknell's enumeration of supposed synonyms of 
his S. “angustifolium” we next come to S. anceps Cav. Diss. vi. 
345, t. exc, fig. 2 (1788), and we almost repeat the story. This 
plant (our PLATE 1042, ria. 2), “Habitat in Virginia’, described 
“Caule simplici ancipite saepius aphyllo", was shown as either 
simple or with a fork at the axil of the cauline leaf, just as in the 
Dillenian plate of S. angustifolium and in Bicknell’s plate (our 
PLATE 1043, rias 1 and 2) of his S. graminoides, while the long- 
exserted pedicels are too suggestive of Bicknell's illustration of his 
S. graminoides. Here, then, is the same old plant of European 
gardens, S. angustifolium Mill. (not sensu Bicknell) and, to cap 
the climax, Cavanilles calmly gave as the same as his reputedly 
new S. anceps the following synonyms: S. Bermudiana var. « L.; 
S. gramineum Lam.; S. angustifolium Mill., the pre-Linnean S. 
caeruleum parvum, gladiato caule, virginianum of Plukenet, Alm. 
348, t. 61, fig. 1, and the much discussed Bermudiana graminea, 
flore minore coeruleo of Dillenius. Then, for full measure, 
Cavanilles added a new synonym: *S. Minus. R. H. P. [Roy. 
Hort. Paris, 7. e. Jardin du Roi, whence Lamarck got his material 
of S. gramineum]|". Thus, S. anceps started off with all the pre- 
viously published names for the North American plants encum- 
bering it; and by the same Rule which invalidates the name S. 
gramineum Lam. the name S. anceps is also illegitimate. 

The last name cited by Bicknell as synonymous with his 
Sisyrinchium *'angustifolium" is S. montanum Greene, Pittonia, 
iv. 33 (1899), described from the mountains of Colorado. Here, 
at last, is a resting-place for the commonest and most widespread 
species of the genus in Canada and the Northern States, but, 
even so, the common plant of the East and Northeast cannot 
really rest. Most of the Northeastern plants have the leaves and 
stem full green, the green or purple-suffused capsules becoming 
dull brown or greenish-purple, when ripe changing to almost 
blackish (PLATE 1044, ria. 3). Greene’s S. montanum, an ISO- 
TYPE shown in our PLATE 1044, FIGs. 1 and 2, is pale or whitish- 
green, with the capsules whitish-brown or pale-stramineous. 
In the East it occurs along the limy valleys of the Gaspé Penin- 
sula and Anticosti; it is also in western New York, along the 
Ottawa River, on the limestone of Bruce Peninsula and in 
northern Michigan and Minnesota. So far as I can find there is 


158 Rhodora |JuLy 


no published name for the common deeper green and darker- 
fruited and not prevailingly calcicolous eastern plant. 

To summarize, the species of SisyRINCHIUM here specially dis- 
cussed are 


SISYRINCHIUM ANGUSTIFOLIUM Mill. Dict. ed. 8, 2 (1768), 
based largely, if not wholly on Bermudiana graminea, flore minore 
caeruleo Dill. Hort. Elth. 49, t. XLI, fig. 49 (1732). S. Bermudi- 
ana L. Sp. Pl. 954 (1753) at least as to citation of Dillenius. 
S. foliis lineari-gladiolatis, pedunculis longioribus Mill. Gard. 
Dict. ed. 7, no. 2 (1759). S. gramineum Lam. Encycl. i. 408 
(1783) in part. S. anceps Cav. Diss. vi. 345, t. exc, fig. 2 (1788). 
Ferraria pulchella Salisb. Prodr. 42 (1796), in part, as based on 5. 
anceps Cav. (illegitimate substitute). S. graminoides Bicknell in 
Bull. Torr. Bot. Cl. xxii. 133, plate 263 (1896). Our PLATES 
1042, rias. 1 and 2, and 1043, Fras. 1 and 2. 

S. MONTANUM Greene, Pittonia, iv. 33 (1899). PLATE 1044, 
FIGS. 1 and 2. S. angustifolium sensu Bicknell in Bull. Torr. 
Bot. Cl. xxvi. 336 (1899), in part only. S. septentrionale Bicknell 
in Bull. Torr. Bot. Cl. xxvi. 452 (1899), the slenderest extreme. 
Plant pale green, not much if at all darkened in drying; capsules 
whitish-brown or pale-stramineous, not becoming blackish in 
age.—Caleareous and other mountains and river-gravels and 
openings, western Newfoundland to Mackenzie and northern 
British Columbia, south to Anticosti Island and Gaspé Peninsula, 
Quebec, southern Ontario, western New York, Michigan, north- 
ern Indiana, northern Illinois, Iowa, Nebraska, Colorado, Utah 
and southern British Columbia. All material I have seen from 
Michigan and Manitoba northwestward and westward belongs 
here. Characteristic eastern specimens (selected from a larger 
series) are: NEWFOUNDLAND: dry limy barrens, upper slopes and 
tablelands, alt. 200-300 m., Table Mountain, Port-au-Port Bay, 
Fernald & Wiegand, no. 4252, Fernald & St. John, no. 10,814 (both 
as S. septentrionale). QueBEC: Anticosti I., graviers calcaires, 
many colls., from Victorin & Rolland-Germain (some of them with 
Louis-Marie): no. 20,333 (Rivière Vaureal), 20,334 (R. au Sau- 
mon), 20,335 (R. à la Patate), 24,233 (R. Jupiter), 24,234 and 
27,108 (Pointe Sud-Ouest), 27,109 (R. Chicotte), 27,110 (R. 
Dauphine), 27,111 (R. Natiskotek), 27,112 (R. McKane), 27,113 
(R. du Brick); alluvium, banks of Grand River, Gaspé Co., 
June 30-July 3, 1904, Fernald; alluvial thicket between the Forks 
and Brulé Brook, Little Cascapedia River, Bonaventure Co., 
July 29 and 30, 1904, Collins, Fernald & Pease (Pease, no. 4971); 
alluvium of Little Cascapedia R., Collins & Fernald, no. 56; 
Rivière Petite Cascapedia, Victorin, Rolland & Jacques, no. 
33,781; alluvium, Grand Cascapedia River, July 12-15, 1905, 


Rhodora Plate 1043 


Photo B. G. Schubert 


SISYRINCHIUM ANGUSTIFOLIUM Mill.: Fics. 1 and 2, portions of Bicknell’s illustra- 
tions of S. graminoides Bicknell. 

S. MONTANUM, Var. CREBRUM: FIG, 3, reduced from Bicknell’s illustration of S. an- 
gustifolium sensu Bickneil, not Mill. 


Rhodora Plate 1044 


Photo B. G. Schubert 


SISYRINCHIUM MONTANUM Greene: FIG. 1, portion, X 1, of ISOTYPE. 
S. MONTANUM, Var. CREBRUM Fernald: FIG. 2, fruiting portions, X 1, of TYPE. 


1946] Fernald,—Identifications of North American Plants 159 


Williams, Collins & Fernald; alluvium of Bonaventure River, 
Bonaventure Co., Aug. 5, 6 and 8, 1904, Collins, Fernald & Pease 
(Pease, no. 5780); gravelly beach, Carleton Point, Carleton, 
July 21, 1904, Collins & Fernald; dry calcareous sand back of 
beach, Tracadigash Point, Carleton, Fernald & Weatherby, no. 
2426; sur le barachois, Carleton, Victorin, Rolland & Jacques, 
no. 33,576; Chicoutimi, Saguenay River, Aug. 3, 1892, Kennedy. 
New York: low field 24 miles northwest of Waterloo, Seneca 
Co., Wiegand, Eames & Randolph, no. 11,801; low meadow, 
Victor, Ontario Co., House, no. 17,489. ONTARIO: east of Davis 
Swamp, Ottawa, J. Macoun, no. 86,675; Casselman, J. Macoun, 
no. 4387, crevices and talus of hornblendic cliffs and ledges, 
Cloche Peninsula, Manitoulin Distr., Fernald & Pease, no. 3244; 
dry limestone flats, Cloche Peninsula, Fassett, no. 14,570; Hay 
Bay, Tobemory, Bruce Peninsula, Krotkov, no. 7281; open glade, 
Jack Fish, Thunder Bay Distr., Pease & Bean, no. 23,547; 
Sleeping Giant, Thunder Bay Distr., Taylor, Losee & Bannan, 
no. 915. 

S. MONTANUM, var. crebrum, var. nov. (TAB. 1044, FIG. 3) 
foliis saturante viridibus; capsulis viridibus vel purpurascentibus 
deinde plus minusve atrobrunneis.—Foliage and stem deep 
green, drying dark or blackish; capsules green or purple-tinged, 
in age fuscous-brown or blackish.—Southwestern Greenland; 
Newfoundland to Ontario, south to Nova Scotia, New England 
and Pennsylvania, and mts. to western Virginia (according to 
Bicknell). Type from Clark's Point, Southwest Harbor, Maine, 
August 17, 1890, Edward L. Rand (in Herb. Gray.). 

In addition to the type the following, from twenty times as 
many sheets, are cited as characteristic. GREENLAND: Niina- 
tàrssüak, 65° 25’ N., 49° 50’ W., August, 1932, J. Iversen. NEw- 
FOUNDLAND: by rill on steep siliceous slope of Joan Plains Hill, 
Bay Bulls, Fernald, Long & Dunbar, no. 26,528; dry soil in fields, 
near Topsail, Conception Bay, Howe & Lang, no. 1219; Glen- 
wood, Fernald & Wiegand, nos. 5196 and 5198; dry burned 
crests north and east of Tilt Cove, F. & W., no. 5199; Grand 
Falls, F. & W., no. 5195; Port Saunders, F. & W., no. 3094; Cow 
Head, F. & W., no. 3091; river-gravel, mouth of McKenzie 
River, Bonne Bay, Fernald, Long & Fogg, no. 1531; Lark Harbor, 
F. L. € F., no. 188. QuEBEc: Natashquan, St. John, no. 90,327, 
Victorin & Rolland, no. 28,543; Cap de l'Est, Anticosti, Victorin 
& Rolland, no. 27,105; graviers de platiéres, Riviére la Loutre, 
Victorin & Rolland, no. 24,235; La Madeleine, Gaspé Co., 
Rousseau, no. 31,099; Mont-Saint-Pierre, Gaspé Co., Victorin, 
Rolland & Jacques, no. 33,229; peaty headland, Cap au Renard, 
Gaspé Co., Fernald & Pease, no. 24,968; Isle-aux-Courdes, 
Victorin, no. 4192; Lac St.-Jean, Victorin, no. 15,957; Ile-aux- 
Basques, Co. de Temiscouata, Victorin, Rolland & Jacques, no. 


160 Rhodora [JULY 


33,042; sable, nord de Mont-Laurier, Senneterre, Victorin, 
Rolland & Dominique, no. 204. MaAGpALEN IsLANDs: dry sandy 
field, Wolf Island (Pointe du Loup), Fernald, Bartram & Long, no. 
7214. Prince Epwarp IsLanp: dry clearings, Alberton, 
Fernald & St. John, no. 7213. New Brunswick: dry gravel- 
pavement, Belledune Point, Fernald & Pease, no. 24,969; sandy 
shore, Bathurst, S. F. Blake, no. 5387; damp place in field, Deep 
Cove, Grand Manan, C. A. & Una F. Weatherby, no. 5530. 
Nova Scotia: Barrasois River, Cape Breton, G. E. Nichols, no. 
611; gravelly beach, Guysborough, Rousseau, no. 35,352; gravelly 
beach of Third Lake, Windsor Junction, Fernald & Long, no. 
20,806; seepy banks and moist fields, Yarmouth, Bissell, Pease, 
Long & Linder, no. 20,793. Marne: dry fields, Blaine, Fernald, 
no. 2493; Peaked Mt., Clifton, Aug. 22, 1897, Fernald; roadside, 
Newry, Pease, no. 28,376; dry pasture, Roque Bluffs, Aug. 1, 
1911, C. H. Knowlton; boggy meadow, Harriman Point, Brook- 
lin, A. F. Hill, no. 1347; old orchard, Lincolnville, G. B. Rossbach, 
no. 148; Blackstrap Hill, Falmouth, Bissell & Chamberlain, no. 
373; York Harbor, Aug. 25, 1896, Bicknell. New HAMPSHIRE: 
dry pasture, Martin Location, Coés Co., Pease, no. 14,251; Dur- 
ham, Hodgdon, no. 4000; Hookset, June 7, 1921, C. F. Batchelder. 
VERMONT: field, Concord, Essex Co., Pease, no. 28,576; Wells 
River, Pease, no. 29,462; sand plain, Swanton, S. F. Blake, no. 
3153; Manchester, M. A. Day, no. 171. MASSACHUSETTS: Essex, 
June 13, 1896, E. F. Williams; Groton, Harris & Smith, no. 2678; 
Boxborough, Hubbard & Torrey, no. T521; wet field, Stoughton, 
June 2, 1909, Kennedy; Plymouth, Fernald & Hunnewell, no. 
15,083; dry sandy oak barrens, Harwich, Fernald, no. 16,601; 
South Ashburnham, F. F. Forbes, no. 1085; dry gravelly bank, 
Ashfield, June 19, 1921, Churchill et al.; dry open soil, Middle- 
field, Fernald & Long, no. 9262. Ruopr IsrANp: Cumberland, 
May 30, 1911, R. A. Ware; Middletown, May 30, 1908, E. F. 
Williams; Block Island, June 16, 1917, R. P. Marshall. CoN- 
NECTICUT: dry field, Franklin, May 31, 1906, Woodward; moist 
field, South Britain, Blewitt, no. 624; wet meadow, Waterbury, 
Blewitt, no. 567; dry hill, Southington, L. Andrews, no. 489; 
Oxford, June 8, 1902, Harger; meadows, Stratford, June 5, 1898, 
Eames. New York: moist open field, Madrid, Phelps, no. 340; 
Canton, Phelps, no. 108; Fort Ann, July 15, 1917, Burnham; 
Oneida Castle, Maxon, no. 152; Caroline, Hames & MacDaniels, 
no. 3801; Dryden, F. P. Metcalf, no. 2025. PENNSYLVANIA: 
wooded roadside, Muncy Valley, Sullivan Co., Fosberg, no. 
15,036; grass-flat in open woods, Leroy, Bradford Co., June 21, 
1941, W. F. Westerfeld. Ontario: dry clearing, Temagami 
Forest, W. R. Watson, no. 6701; open field, Central Island, 
Toronto, June 18, 1936, R. J. Eaton; crevices of sandstone, 
Corbeil Point, Algoma Distr., T. M. C. Taylor et al., no. 705. 


1946] Fernald,—Identifications of North American Plants 161 


SoME ORCHIDS OF THE MaNuAL Rance (Prares 1045-1048). — 


HABENARIA CLAVELLATA (Michx.) Spreng., var. ophioglos- 
soides, var. nov. (TAB. 1046), a var. typica recedit folio infimo 
sessile, vix basi prolongato, ovale vel oblongo vel late oblanceo- 
lato 1-4 cm. lato, 3-17 cm. longo, longitudine bis vel etiam 
sexies majore quam latitudine.—Newfoundland and Cóte Nord, 
Que., to western Ontario, south to Nova Scotia, New England, 
New Jersey, Michigan and northern Wisconsin. Type from 
Nova Scotia: boggy clearing near Porcupine Lake, Arcadia, 
Yarmouth Co., Sept. 2, 1920, Fernald & Long, no. 20,839 (in 
Herb. Gray.; isotype in Herb. Phil. Acad.). 


Typical Habenaria clavellata was based on Orchis clavellata 
Michx. Fl. Bor.-Am. ii. 155 (1803) from Carolina. As shown by 
the Michaux TYPE (our PLATE 1045, rra. 1), which, unfortunately, 
has the tip of the backward-folded leaf covered by the label, 
he had the characteristic Carolinian and broadly southern plant 
with the relatively narrow leaf tapering to a subpetiolar base, a 
full-length leaf from North Carolina shown as Fig. 2. Typical 
H. clavellata has the single large leaf narrowly oblong, oblanceo- 
late or spatulate, gradually tapering to a rather definite petiolar 
base, the leaf varying from 5.5 to 23 cm. long and 0.7-2.7 em. 
broad (one twelfth to one fifth as broad as long). All material 
in the Gray Herbarium from Florida westward to Louisiana and 
northward to Virginia and Tennessee is typical H. clavellata, 
while this extreme of the species pushes locally northward into 
the southern area of var. ophioglossoides. 

The latter, var. ophioglossoides, has oval, oblong or broadly 
oblanceolate leaves rounded to tapering to an essentially sessile 
(not subpetiolar) base, the blade 3-17 cm. long and 1-4 cm. 
broad (one sixth to one half as broad as long). It is, as a series, 
more northern than typical H. clavellata, its dominance in the 
North indicated in the following check of specimens in the Gray 
Herbarium and that of the New England Botanical Club: from 
NEWFOUNDLAND, typical H. clavellata 0, var. ophioglossoides, 21; 
QUEBEC, typical clav. 0, var. oph. 8; PRINCE EDWARD ISLAND, 
typical clav. 0, var. oph. 4; New Brunswick, typical clav. 0, 
transition 1, var. oph. 7; Nova Scoria, typical claw. 0, transition 
1, var. oph. 23; MAINE, typical clav. 0, transition 6, var. oph. 75; 
New HAMPSHIRE, typical clav. and transition 8, var. oph. 42; 
VERMONT, typical clav. and transition 1, var. oph. 12; Massa- 


162 Rhodora [JULY 


CHUSETTS, typical clav. and transition 41, var. oph. 17; RHODE 
ISLAND, typical clav. 7, var. oph. 1; CoNNEcTICUT, typical clav. 6, 
var. oph. 0; New York, typical clav. 8, var. oph. 5; ONTARIO, 
typical clav. 0, var. oph. 6; MicHIGAN, typical clav. 0, var. oph. 6. 
Too little material is at hand from New Jersey, Pennsylvania 
and Ohio to Minnesota and Iowa to indicate the relative fre- 
quency of the two in those states; but the strong development 
southward of typical H. clavellata and northeastward of var. 
ophioglossoides is clear. 

From the citation in Ames, Enum. Orchids U. S. and Canada, 
44 (1924) one would infer that, in publishing the basic Orchis 
clavellata, Michaux assigned it earlier synonyms which might 
render his later name (of 1803) untenable. The first citation 
in the Enumeration, under synonymy of H. clavellata reads: 
“Orchis clavellata Michaux, Flora Boreali-americana, vol. 2, 
p. 155 (1803), exclude synonymy in part." Reference to 
Michaux's treatment shows, however, merely the description of 
the new species, with “Has. in Carolina"! As originally pub- 
lished Orchis clavellata had no synonymy, so that the validity of 
the name is unblemished. 

Orchis tridentata Muhl. ex Willd. Sp. Pl. iv. 41 (1805) from 
Pennsylvania, with ‘Folia caulina tria, infimum lanceolato- 
ensiforme dodrantale" must be the typical relatively southern 
Habenaria clavellata, for the very narrow leaf “nine inches long" 
can apply only to that, and the material from Lancaster County 
before me is of typical 77. clavellata. 


(T'o be continued) 


! Michaux's Flora Boreali-Americana was based on his own collections made during 
many years of pioneer botanical exploration in eastern North America, just as the 
Flora Virginica of Gronovius was based on collections made in Virginia by John Clay- 
ton, the latter book prepared with the collaboration of the young Linnaeus who, later, 
based many binomials on the Clayton plants described by Gronovius. 'This is common 
knowledge, clearly stated in the two pioneer books. It is, therefore, a bit startling to 
read in the North American Flora, xxviii B?, in the treatment of the Umbelliferae, items 
like the following: Ptilimnium capillaceum (Michx.) Raf., resting on Ammi capillaceum 
Michx. Fl. Bor.-Am., with ‘TYPE rLocarrrv: ‘In campestribus Carolinae’, collector 
unknown”; or Thaspium trifoliatum (L.) Gray, resting on Thaspia trifoliata L., from 
Clayton, with Linnaeus citing '' Gron. virg. 31", said to have its type from '' ' Virginia’, 
collector unknown''. Surely, the editors could have found out. Such entries are 
similar to the enumerations of states from which species are known, with the state 
from which the type came omitted. 


Rhodora Plate 1045 


IERR MIS- PARIS 


Herbier de V Ame 


Fue Sentonty 


Photo B. G. Schubert 


HABENARIA CLAVELLATA: FIGS. 1 and 2, Type of Orchis clavellata Michx., X 1, after 
Cintiact; FIGs. 3-6, leaves X 1, from various localities. 


Rhodora Plate 1046 


Photo B. G. Schubert 


HABENARIA CLAVELLATA, var. OPHIOGLOSSOIDES: FIGS. 1 and 2, portions of TYPE, 
X 1; FIGs. 3-6, leaves, X 1, from various localities. 


1946] Gilly,—Another later Homonym 163 


Retict BOREAL PLANTS IN SOUTHEASTERN MINNESOTA.— 
It has long been known that several typically boreal plants occur 
as relicts in Winneshiek and Allamakee counties of northeastern 
Iowa. These include Abies balsamea (L.) Mill., Mertensia 
paniculata (Ait.) G. Don, Rubus pubescens Raf. and Rhamnus 
alnifolia L'Hér. Recently, in June 1941, this relict area was 
found to extend into Fillmore County of southeastern Minnesota 
and to be represented by a small balsam-fir stand near the town 
of Wycoff. The firs form a fairly dense stand along the north 
slope of a steep and narrow valley and seem to have perpetuated 
themselves by layering. In addition to many species quite 
typical of this region, the following typically boreal species were 
found associated with the firs; Rubus pubescens Raf., Rhamnus 
alnifolia L'Hér., Cornus canadensis L., Pyrola secunda L., Pyrola 
chlorantha Sw. and Mertensia paniculata (Ait. G. Don. With 
the exception of this locality, the last species is known in Minne- 
sota only from the north shore of Lake Superior and the remainder 
are largely limited to the coniferous forest region of northern 
Minnesota.—JonN B. Movrz, Minnesota Department of Con- 
servation, St. Paul. 


ANOTHER LATER Homonym.!'—While going through the cards 
of the most recent issue (No. 189) of the Gray Herbarium Index, 
my attention was again drawn to a later homonym which has 
been in use for twenty years in the Sapotaceae. Because of the 
prior publication of Manilkara emarginata H. J. Lam (Bull. 
Jard. Bot. Buitenz. III 7: 241. 1925) for an endemic species of 
the Hawaiian Islands, Manilkara emarginata (L.) Britton & 
Wilson (Bot. Porto Rico 6: 366. 1926), based on Sloanea emar- 
ginata L. (Sp. Pl. 512. 1753), was illegitimate when published 
as a name for the “wild dilly” of southern Florida, the Florida 
Keys and the Bahama Islands. If one associates this northern- 
most western-hemisphere member of the genus with several 
Antillean entities, as Cronquist recently has done (Bull. Torrey 
Club 72: 550-562. 1945), the proper name for the species- 
aggregation would be Manilkara Jaimiqui (Wright) Dubard 


1 Thanks are due Mrs. Lazella Schwarten, librarian of the Arnold Arboretum, for 
her kindness in checking several bibliographic references. 


164 Rhodora [JULY 


(Ann. Col. Mus. Marseille III 3: 16. 1915; based on Mimusops 
Jaimiqui Wright in Griseb. Cat. Pl. Cub. 164. 1866). 

However, if one considers the “wild dilly” to be a distinct 
species—and I am at present inclined to take this stand—its 
proper appellation would seem to be Manilkara bahamensis 
(Baker) Gilly, comb. nov. (based on Achras bahamensis Baker in 
Hook. Icon. 18: ¢. 1795. 1888). Synonymy of this species 
would also include! Mimusops floridana Engl. (Engl. Bot. Jahrb. 
12: 524. 1890), Mimusops bahamensis (Baker) Pierre (Not. 
Sapot. 37. 1891) and Mimusops emarginata (L.) Britton 
(Torreya 11: 129. 1911).—CnanrzEs L. Gritty, Department of 
Botany, Iowa State College, Ames, lowa. 


1The name ‘‘Mimusops parvifolia Radlk.” (not Mimusops parvifolia R. Br. Prodr. 
531. 1810; nor Mimusops parvifolia Kurz, For. Fl. Br. Burma 2: 124, 1877; nor 
Manilkara parvifolia (Kurz) H. J. Lam, Bull. Jard. Bot. Buitenz. III 7: 269. 1925) 
is not included in the legitimate synonymy of this species for the reasons stated in 
the following sentences. Although Pierre (Not. Sapot. 37. 1891) and most sub- 
sequent authors have cited this name as ‘‘Mimusops parvifolia Radlk.", Radlkofer 
published it as Mimusops parviflora (Sitzb. Akad. Wiss. München 12: 344. 1882), 
basing his new combination upon ‘‘Achras zapotilla var. parviflora Nuttall”, which he 
cites on the authority of A. Gray. Nuttall, however, published this variety as Achras 
zapotilla B parvifolia (N. Am. Sylv. 3: 28, t. 90. 1849). Unfortunately Asa Gray in- 
cluded Nuttall's varietal name in synonymy under Mimusops Sieberi A. DC., which 
he believed to be the correct name for the ‘‘wild dilly”, as Achras zapotilla var. parvi- 
flora (Syn. Fl. N. Am, 2: 69. 1878). This erroneous citation by Gray seems best 
handled as a superfluous renaming of Nuttall's variety; Achras zapotilla var. parviflora 
Nutt. in A, Gray, therefore, is an illegitimate name under Article 60 of the Internation- 
al Rules of Botanical Nomenclature. Subsequent usage of this varietal epithet parvi- 
flora, including its elevation to specific rank by Radlkofer, is equally illegitimate; this 
particular epithet parviflora is, therefore, unavailable for transfer to the genus Manil- 
kara. 


Volume 48, no. 570, including pages 113-136 and plates 1027—1030, was 
issued 4 June, 1946. 


op c 


Ww 


Dodota 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Clahy by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED beside seni f 
ALBERT FREDERICK HILL - Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. August, 1946. No. 572. 
CONTENTS: 
‘Lhe Genus Liatris. L. Q- Gaer in «225 a eee s 165 


Betula glandulosa at a low Altitude in New Hampshire. 
Charlotte. Endicott Wilde. ...... 00.000. cech RR uo eV Cus 183 


Contributions from the Gray Herbarium of Harvard University— 
No. CLXII. Identifications and Reidentifications of North 
American Plants. M. L. Fernald (continued). ............ 184 


Application of the Name Euphorbia maculata L. F. R. Fosberg. 197 


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Vol. 48. August, 1946. No. 572. 


THE GENUS LIATRIS 
L. O. GAISER 


Tue North American genus Liatris has been considered one of 
unusual difficulty. Variability in and intergradations between 
the species are undoubtedly responsible for the bewildering prob- 
lems in specific determination. When comparisons of various 
collections, used for the growing of cytological material, were 
made with herbarium specimens difficulties arose in the determi- 
nation of species, and the project grew further into an examina- 
tion of types and a taxonomic revision. It has seemed that the 
present summation of results might meet some others’ needs as 
well as my own. 

Preliminary work was begun in 1928 by the study of living 
cultures grown in a garden at Crediton, Ontario, from plants 
collected by the generous help of many botanists of the United 
States. Recognition of the great kindness of all those who con- 
tributed to that phase of the work will be made elsewhere. 
Many species have also been collected in the field on a trip in 
the south-central States, westward through Arkansas and south- 
ward through Texas to the Gulf of Mexico. The present treat- 
ment is, however, largely the result of an examination of the 
mass of material at the Gray Herbarium, where work was begun 
in the summer of 1938, at the New York Botanical Garden, where 
many types of Small and Rydberg are available, and the National 
Herbarium, Washington, which possesses a great number of 
specimens collected by E. S. Steele. Other herbaria have also 
been visited and particular collections, such as the Greene types 


166 Rhodora [AvaUsT 


at Notre Dame University, the Nuttall and Pursh specimens at 
the Philadelphia Academy of Sciences, and the Gates specimens 
at the Brooklyn Botanical Garden, have been studied. As well, 
loans from other herbaria, as of Lunell's varieties of L. scariosa 
from the University of Minnesota and the University of Indiana 
have been of invaluable help. Representative collections re- 
ceived, as from the University of Oklahoma, the A. & M. College 
of Texas, the University of North Carolina, the National Mu- 
seum of Canada, and the Fowler Collection from Queen's Uni- 
versity, Kingston, have also been greatly appreciated. To all 
the curators of these and to the owners of private herbaria who 
have been such a help in making available to me so many speci- 
mens for study and comparison, I am greatly indebted and wish 
to express sincere gratitude. To Prof. I. M. Johnston, I grate- 
fully acknowledge the stimulation and encouragement which 
involved me in undertaking this taxonomie study. I was im- 
measurably helped by having Prof. M. L. Fernald show me the 
photographs of type specimens in the herbarium of the Linnaean 
Society, London. To Mr. C. A. Weatherby, I am especially 
indebted for having obtained for me during his visit abroad to 
the herbaria and museums in London, Paris, and Geneva during 
the summer of 1939, photographs of many other type specimens, 
without which this study could not have progressed. From all 
the staff at the Gray Herbarium has come much appreciated 
help in oecasional conferences which I am happy to acknowled ge, 
as I am also appreciative of the ever generous help of the li- 
brarian, Miss R. D. Sanderson. To students who have done 
what must have been exacting typing for them, goes my most 
sincere gratitude for their help in the preparation of this manu- 
script. 

In citing specimens from various herbaria the following ab- 
breviations will be used: 


G—Gray Herbarium, Harvard University; NY—New York Botanical 
Garden; US—National ‘Herbarium, Washington; P—Academy of Natural 
Sciences, Philadelphia ; B—Brooklyn Botanical Garden; ND—Greene 
Herbarium, Notre Dame University; M—University of Minnesota, 
Minneapolis; I—Indiana University, Indianapolis; T—College of A. & 
M., College Station, Texas; O—University of Oklahoma, Norman; NC— 
University of North Carolina, Chapel Hill; Ot—National Museum, 
Ottawa; Q—Queens University, Kingston, Ontario; To—University of 
Toronto, Ontario; W—University of Wisconsin, Madison; N—University 


1946] Gaiser,—The Genus Liatris 167 


of Nebraska, Lincoln; OA—Herbarium of Prof. O. Ames, Ormond, 
Florida; HB—Herbarium of Mr. Hubert B. Brown, Toronto; WH—Her- 
barium of Mr. W. H. Herriot, owned by Mr. Monroe Landon, Simcoe, 
Ontario: F—University of Florida, Gainesville. 


JENERAL DISCUSSION 
BiBLIOGRAPHY OF THE GENUS AND ITS CHIEF SUBDIVISIONS 


LiaTmRIs Schreb. Gen. Plant. ii. no. 1263, 542 (1791) (nomen 
conservandum); Endl. Gen. Plant. no. 2270, 368 (1836-1840); 
Benth & Hook. Gen. Pl. ii. no. 73, 248 (1873) ; Engl. & Prantl, Nat. 
Pflanzenfam. iv. Abth. 5, 142 (1890). Lacinaria Hill., Veg. Syst. 
iv. t. 49 (1762); Porter & Britt. Mem. Torr. Bot. Club, v. 313 
(1894). Laciniaria Hill, Hort. Kew. 70 (1769) ; O. Ktze. Rev. Gen. 
i. 349 (1891).  Psilosanthus Neck. Elem. i. 69 (1790). 

Section I. EurrATRIS DC. Prod. v. 128 (1836).  Liatris as 
limited by Cassini, Dict. Sci. Nat. xxvi. 235 (1823). 

Section II. Supraco (Gaertn.) DC. Prod. v. 129 (1836). 
Suprago as limited by Cass. Dict. Sci. Nat. li. 385 (1827). 


(GENERIC CHARACTERS 


Lrarris Schreb. Perennial herbs, generally from ovoid or 
globular corms, but sometimes with much flattened or quite 
elongate rootstocks; leaves elongate, linear to ovate-lanceolate, 
sessile or petiolate, more or less conspicuously punctate with 
impressed and resinous dots, the radical leaves usually much 
longer than the cauline; stems with numerous spirally arranged 
leaves, these diminishing upwards as bracts subtending flower- 
heads: heads of flowers in cymosely disposed spikes, racemes or 
panicles, rarely a loose open cyme; the heads of 3 to numerous 
(up to 70) similar, tubular flowers on a naked receptacle sur- 
rounded by an involucre of imbricated phyllaries in several 
series: phyllaries lanceolate, ovate, oblong or orbicular; mu- 
cronate-, acute-, or obtuse-tipped, herbaceous, narrowly or 
broadly petaloid with ciliate or deeply erose margins: corolla 
phlox-purple, or rarely white, regular, usually glabrous without, 
commonly dotted with scattered resinous droplets; tube cylin- 
drical, usually exceeding the pappus, or twice as long as the 
pappus; throat hardly at all or slightly perceptible, glabrous 
within or with little or much pilosity; lobes 5, equal, ovate, acute, 
erect, or more or less spreading, glabrous or pilose within: 
stamens 5, included, filaments filiform, equally inserted below 
the middle of the corolla-tube, glabrous or with tiny outgrowths; 
anthers short, oblong, about as long as the filaments: style stiff, 
bifid and exserted after anthesis: achene somewhat cylindrical 
but pointed at base, about ten-ribbed, pubescent on ribs and 
more finely between ribs: pappus of 12-40 bristles, sessile in one 
or more series, plumose or barbellate. 


168 Rhodora [AUGUST 


North American herbs: United States, although not west of 
the Rocky Mts., southern Canada, and most northerly Mexico. 


THE SPECIES 

Thirty-two species with their varieties and ten hybrids, have 
been here recognized, though the number of the latter is not 
definitive. In some cases the wide variety in individuals of an 
interspecific hybrid has made description impossible. As there 
is much evidence of intermediate material it is a genus in which 
it is difficult to draw sharp lines of distinction between the species. 

The greatest number of species occur in Florida, at least 
twelve being known from that state. At the western limits of 
extension, as in the states of Colorado and Montana or in the 
province of Alberta, there are probably two species, L. punctata 
Hook. and L. ligulistylis (Nels.) K. Sch. In the New England 
States there is probably but one species, L. borealis Nutt. ex 
MacNab and some intergradations with the species of adjacent 
districts. In southern Ontario, two species and one stable 
hybrid occur in the sandy stretches along Lake Erie and southern 
Lake Huron, L. cylindracea Michx., L. aspera Michx., X L. 
sphaeroidea Michx. and a third species, L. spicata (L.) Willd., 
only in a limited region along Lake St. Clair. A few species, 
such as those of the series Spicatae (except L. microcephala 
(Small) K. Sch.) and occasional ones, such as L. ligulistylis, 
grow in low grounds around ponds or even in wet lands. Most 
of the species, however, grow in warm sandy places, as in remain- 
ing oak-hickory forests or, as across Texas, along dry railroad 
tracks. Other species, e. g. L. turgida Gaiser, have been more 
limited to the mountains of the Carolina region, where the 
Appalachian ranges have harbored a rich assortment of species 
and extensive colonial growth. 


Gross MORPHOLOGY 


UNDERGROUND STEM. Being perennial plants, all species of 
Liatris have a thickened underground stem from which fibrous 
roots spread out to anchor the plant. During the first summer 
of the seedling's growth there develop a few radical leaves above 
what appears as a slightly thickened tap-root, but at the end of 
the season an apical bud is developed from a small erown and this, 
in the second year, produces the first flowering stalk. During 


1946] Gaiser,— l'he Genus Liatris 169 


successive summers the stem thickens, becoming globular or 
remaining ovoid in most species of all series, except the Spicatae, 
Pycnostachyae and some of the Punctatae. Very singular is the 
clustered, slender, tuberous formation of L. Garberi A. Gray. 
Except for the roots that come off basally and the buds that 
develop annually from the central summit into aerial stalks, this 
stem lacks any investment, such as a membranous scaly coat, 
and could be spoken of as a naked corm, sensu Gray (Gray’s Bot. 
Textb. i. 61-2 (1879)). In accordance with this interpretation 
we have chosen to use the term corm. In one species there 
occurs a variation of the generally rounded depressed shape, 
which appears definitely related to soil conditions. In L. elegans 
(Walt.) Michx., as it is found from Florida westward through the 
Southern States to Texas and Arkansas, the underground part is 
commonly small and globular or subovoid. However, specimens 
from the Carizzo region of Texas, with no marked distinguishing 
difference of above-ground parts, show an elongated tap-root- 
like growth up to ten times the usual length of the small ovoid 
form. Mr. Parks of College Station, in describing this region, 
writes as follows!: 


“The Carizzo sand is an outcrop of the Carizzo formation of the Eocene 
Age. In Medina, Bexar, and Wilson counties it is a very conspicuous 
part of the landscape. There is a wooded ridge reaching a height of 
nearly two hundred feet above the surrounding country. It is a sand 
dune covered by a climax oak-hickory association. The soft white sand 
in which the trees grow bears very little vegetation, however in the spring 
and late fall when the rains occur it is very prolific of annual plants 
which are of a very short existence. It is needless to say that most of 
these plants have deep tuberous roots”. 


Thus it seems clear that, in common with other plants of the 
region, the effects of varied soil conditions are shown in the 
modification of shape in the subterranean growth of L. elegans 
and such plants are here recognized as a variety (var. carizzana). 

In some of the species of the Spicatae and the Pycnostachyae 
series, the corm, while growing slightly deeper, broadens com- 
paratively more during the successive years, sometimes to a width 
of 10 centimeters or more, thus providing a widened erown from 
which many flower-stalks arise. Such plants have been found 
to live for more than fifteen years, probably representing the 


! By private communication. 


170 Rhodora [AUGUST 


hardiest of the shallow-rooted species, which advantage, along 
with the provision of numerous flowering spikes, makes them the 
favored species in perennial flower-borders. Some buds of broad, 
old stems grow into new corms and upon separation or death of 
the old become new individuals'. 

Some species, yet not all, of the Punctatae series show à more 
pronounced elongation during the thickening of the young 
seedling’s subterranean growth into a tough rootstock that grows 
into the soil, often to the depth of 50 em. or more. Rarely there 
has been found a modification of it into a somewhat more pros- 
trate rhizome which sends off aerial shoots at points along its 
surface, as in L. densispicata Bush, in the Anoka sand-dunes of 
Minnesota. Lack of any such elongation helps to mark off, we 
believe, such species as L. mucronata DC. and L. angustifolia 
Bush from L. punctata with which they have been confused. 

In the course of a study of internal secretions of some of the 
Compositae Mayberry (Sc. Bull. Univ. Kans. xxiv. 8 (1936)) 
described resinous secretory canals in the cortex of the under- 
ground stem, as large pockets, 0.9 mm. in diameter, in three 
species of Liatris (L. pycnostachya, L. punctata and L. aspera). 
In the roots, aerial stems and leaves less conspicuous canals, 
from .03 to .01 mm. in diameter, were also reported. 

Lear. The leaves of most of the species are linear or lanceo- 
late, though in some they are obovate or ovate-lanceolate. The 
early radical leaves, coming from the crown, are usually much 
longer than the cauline ones. "The latter arise spirally around 
the flowering stalks and generally show a reduction in length 
from the base of the stem upwards as they become the subtend- 
ing bracts of the axillary flower-heads. In some species, however, 
the leaves are abruptly shortened and there results a long strict 
spike with setaceous bracts which hardly project beyond the 
flower-heads. Commonly the leaves are sessile; but sometimes 
the broader basal leaves are narrowed into a petiole. The leaves 
are generally firm; in some linear forms they are even coriaceous, 
as in L. laevigata Nutt., though in broader forms they are less so. 
All variations from glabrous to scabrous blades are found and in 
many cases within a single species. For example, there can be 


! In the series Spicatae a number of horticultural names have arisen, which have 
not been exhaustively treated here, 


1946] Gaiser,—The Genus Liatris 171 
associated with the more distinctive characteristics by which we 
recognize the species L. ligulistylis, such as the few large heads 
with erect, loose phyllaries, any condition from the complete lack 
of to a complete covering of hairs on upper and lower surface of 
the leaves. Thus pubescence has been found to be of little use 
as a character for specific differentiation. In all species, not only 
in those of the Punctatae series, the leaves have small sunken 
resinous glands but they may be less conspicuous in some than in 
others. Pubescence of the leaves may or may not be accom- 
panied with cilia along the margin of the leaf. 

INFLORESCENCE. After a period of producing radical leaves, 
the flowering stalks appear singly on young, or severally (in 
groups of 2-20) in larger, older plants where a broader crown has 
developed. These stems are often stiff, tall and erect (as in L. 
spicata and L. pycnostachya), and the floriferous portions, being 
covered with numerous closely-developed flower-heads, have 
been commonly spoken of as spikes, though the development is 
determinate rather than indeterminate, and they are therefore 
false spikes. In all species the apical flower-head is the first to 
open. All variations from strictly sessile, through short-pedi- 
cellate to long-branched-peduneulate flower-heads occur. In 
many (as especially in the Scariosae) there is found variation 
from a straight spike-like to a pyramidal and paniculate inflores- 
cence, the result of a gradation from the apically sessile flower- 
heads successively through longer-pedicellate ones to those low- 
ermost which have become very much elongated- and branched- 
pedunculate. In some species the stems are more slender and 
tufted with flower-heads more scattered in a raceme-like arrange- 
ment which, being again determinate, is correctly a racemiform 
cyme. In one species (L. cymosa (H. Ness) K. Sch.) there is 
considerable branching and the large flower-heads are borne 
more distantly in a loose dichotomous cyme-arrangement, 
giving quite a different appearance from that generally found in 
the genus. However, that there are intermediate conditions 
between this and the false spikes, when the less proximate heads 
occur in a rhipidial cyme-arrangement, is seen in L. cylindracea 
(to which series we have attached L. cymosa), and L. squarrosa 
(L.) Willd. of the closely related series. Thus, though for strict 
correctness the inflorescence should be spoken of as reversely or 


172 Rhodora [AUGUST 


falsely spicate and a reversed or determinate raceme, it is ex- 
pected that the explanation here will suffice to permit an under- 
standing of the unqualified terms used throughout the discussion 
of the genus. The rachis of the inflorescence is sometimes 
glabrous and may be striate, as exemplified in L. microcephala. 
More frequently it is pubescent above and glabrous below, 
though occasionally the entire stalk is puberulent or covered 
with pubescence of short cinereous or longer hairs. 

FLowrn-HEkAp. The heads in Liatris contain from 3-4 to 
50-70 flowers. Exceptional are the terminal heads!, which 
regularly have a greater number of flowers in almost all species. 
In some species there is a marked increase in the apical head, as 
in L. ligulistylis. In certain species which normally have few 
heads, such as L. cylindracea and L. squarrosa (L). Willd., 
reduction to one-headed specimens often occurs, perhaps under 
unfavorable conditions. This has given rise to naming one- 
flowered varieties, which seems inadvisable to the author. Also 
in other species having a generally spicate inflorescence of nu- 
merous close heads occasional specimens show terminal ones very 
much larger than the rest, as, for instance, in L. punctata var. 
turgida Lunell (Amer. Mid. Nat. v. 241 (1918)). Since specimens 
with such exceptional terminal fusion of heads have been found 
to occur in different species, again varietal ranking has not here 
been recognized. Heads having few flowers may be short, 
cylindrical, ca. 6-8 mm. long in L. microcephala, or quite long and 
cylindrical as in L. punctata—ca. 1.5-2 cm. long. With more 
flowers per head, as in the Graminifoliae series, where approxi- 
mately one dozen is an average number, the stoutish cylindrical 
buds give, when the flowers open, heads that are somewhat 
turbinate in shape. Many-flowered heads are hemispherical, 
ovoid or almost globular, their outward appearance varying, 
largely due to the nature of the phyllaries. Measurements given 
were taken, whenever possible, from heads with open flowers, 
the width at the tip of the corolla and the length from there to 
the base of the head. 

In species of Liatris there is a wider variation in the form of the 
phyllaries than of the leaves: In those of few-flowered heads 


! In stating the number of flowers per head in any description of species, heads 
other than the terminal one, being more uniform, are referred to. 


1946] Gaiser,— The Genus Liatris 173 


there may be small outer foliaceous, linear or ovate, closely 
appressed bracts that become gradually more membranous and 
generally more elongate inwardly. They may, however, vary 
generally from acute or mucronate to obovate, and may be 
glabrous, punctate or pubescent. They may also become re- 
curved and slightly colored, though rarely are they petaloid. 
In the exceptional L. elegans (Walt.) Michx., the phyllaries have 
become much elongated and colorful, either pink or white and, 
being loosely arranged, give an altogether distinctive appearance. 
In species with many-flowered heads there occur the same varia- 
tions in shape, margin and texture of the many phyllaries as in 
those with few-flowered heads. But there have developed as 
well many petaloid forms, many of which, without further 
classification, have sometimes been ascribed to L. scariosa, 
which had thus become a fine “melting pot". In L. ligulistylis 
these petaloid phyllaries with deeply erose margins stand loosely 
erect. In L. aspera Michx. a combination of petaloid and crisped 
condition is found. In other species, as L. borealis, there is only 
a narrow scarious margin which may become finely ciliolate, and 
the phyllaries stand loosely erect, while in L. scariosa, phyllaries 
of similar margin may be recurved. The endless intermediates 
and variations between these prime types has given rise to much 
confusion. In L. squarrosa (L.) Willd. where the recurved con- 
dition of the phyllaries has not been accompanied with petaloid 
development, intermediates between the appressed and the 
squarrose types have likewise been numerous. In L. pycno- 
stachya Michx. the degree to which the phyllaries are reflexed 
also varies. In the Spicatae and Pycnostachyae series there is 
considerable variation from colorful to strictly green herbaceous 
involucre but care must be taken to compare specimens of the 
same age since, after flowering time when the seeds mature, 
phyllaries that had color usually lose it and become green. 
FLowER. Corolla. The corolla is tubular, slightly dilated 
upwards, with an almost imperceptible narrowing at the throat, 
and with ovate to lanceolate, acute lobes from 14-14 the length 
of the tube. The length! of the corolla varies from ca. 5 mm. in 
smaller species to ca. 20 mm. in members of the Squarrosae 
series. The wide-spreading lobes of two sections represented by 


! Measurements were made after boiling the corolla. 


174 Rhodora [AUGUST 


L. squarrosa and L. cylindracea, having the largest corollas, have, 
since the time of Nuttall’s observation ‘‘internally villous” 
(Nutt. Gen. ii. 132 (1818)), been characterized as “hairy within" 
(Gray, Man. Bot. ed. 7). Colorless cellular outgrowths, ca. 2 
mm. long in L. cylindracea, can be seen projecting along the mar- 
gin of the lobes in fresh flowers without the aid of a hand-lens. 
The throat of the corolla-tube of many species appears almost 
translucently clear both in pressed and fresh specimens, but in 
others does not. Within the tube of all species, five stamens, 
having filaments about equal in length to the laterally united 
anthers, are inserted about the middle of the throat. By 
splitting the tube with a fine needle and examining it with a dis- 
secting lens, small colorless cellular hair-like outgrowths ca. 
50-100 u long (which is only a fraction of the length of those on 
the lobes of Squarrosae species) are sometimes found to be abun- 
dant about and below the region of stamen-attachment in the 
throat. In one series, Tenuifoliae, smaller outgrowths come from 
the filaments. Quite unexpectedly, the presence of this pilosity 
within the throat of the corolla proved to coincide with other 
diagnostic characters in some pairs of species that were other- 
wise difficult to separate, and specific differentiation was thereby 
strengthened: as, for example, L. spicata without and L. gramini- 
folia (Walt.) Willd. with hairs; or L. ligulistylis without and L. 
aspera with hairs. Care must be exercised since on hasty exam- 
ination fungal filaments, sometimes found in herbarium specimens 
might be mistaken for such pilosity. 

Commonly on microscopic examination of herbarium speci- 
mens, tiny resinous droplets are seen distributed over the corolla, 
but, as the occurrence of such probably transitory secretions was 
not consistent for all specimens of a species, no proof of their lack 
in any species was found. 

Cotor. Liatris flowers are all of approximately one color, 
somewhere near phlox-purple by Ridgway's color chart, though 
they vary in intensity of the shade, except for the occasional 
white forms. The latter differ in no way from the species’ 
descriptions except for the corolla-color. The one general! 
exception is in L. elegans, where the phyllaries, which are so much 


! A singular example of a yellow corolla is found in L. elegans f. Fisheri Standley 
(Field Mus. Pub. xi. 275 (1936)), a plant with ‘‘both the flowers and the long petal- 
like tips of the bracts lemon-yellow in color”. 


1946] Gaiser,—The Genus Liatris 175 


expanded and which add much beauty, as the name of the species 
suggests, to the heads are white and green accompanying white 
or mauve-tinted flowers, or there may be pinkish-green. bracts 
accompanying the white ‘flowers. Through southern Arkansas 
and Texas, where I have observed the species, the white- and 
purplish-flowered plants seemed equally abundant and successful 
and yet but one species could be recognized. The author’s 
experience in transplanting white-flowered forms of at least four 
other species along with those of normal color (L. aspera’, L. 
punctataà?, L. spicataà?, and L. cylindricea?), has been that the 
white forms never survive longer than about one year after 
transplantation, though the companion plants of normal color 
have lived on for à number of years. This experience was borne 
out by that of Mr. A. C. Edinborough*of Baljennie, Saskatchewan, 
who transplanted the white-flowered form of L. punctata to soil 
similar to that of its native habitat in the Eagle Hills. Appar- 
ently, in species of Liatris the white-flowered forms merely 
represent weaker mutants; they are therefore not being given 
varietal rank. 

Apart from the reference to one sweet-scented variety, La- 
cinaria scariosa var. trilisioides Farwell (Rep. Mich. Acad: Sci. 
xvii. 170-171 (1916)), which has not been encountered, only one 
species, L. tenuifolia Nutt., has been reported* as noticeably 
fragrant. Where hundreds and thousands of plants‘grew the air 
was said to be “delicately scented, much like that of the Buddleia 
slightly modified, and sufficient to arrest one's attention". » The 
observer noticed that many butterflies and bees had been drawn 
to the spot and also that the three albino plants found in the 
region were likewise fragrant. 

Fruit. The fruit of Liatris is a ribbed, somewhat cylindrical 
achene, finely pointed at the base, varying in length from ca. 3 
mm. in some species to ca. 10 mm. in the largest. In color, as 
in size, considerable difference can be noted between unfilled 
fruits and completely matured ones, and therefore care must be 
taken when giving color from examination of herbarium speci- 

! Found near Pt. Edward, Lambton Co., Ontario, Aug. 26, 1938, no. 168. 

? Received from Mr. A, C. Edinborough, Baljennie, Sask. Aug. 24, 1939, no. 202, pl. 1. 

* Found at Walpole Is., Lambton Co., Ontario, Aug. 26, 1938, no. 166. 

4 Found at Turkey Point, Norfolk Co., Ontario, Aug. 27, 1940, no. 213. 


* By private communication. i 
* By private communication from Mrs. H. T. Butts, Ormond Beach, Fla, Oct. 1944. 


176 Rhodora [AUGUST 


mens. Mostly the mature achenes are of a brown color, but in 
some species they approach black. "The number of ribs was not 
uniform; in most specimens ca. 10 was the average. In all 
species the achenes are pubescent along the ribs and more finely 
and inconspicuously so between the ribs. Though in some 
cases achenes showed angles, appearing as if developed by close 
approximation during growth, no specific coordination could 
be found. 

Paprus. One must be careful in stating the color of the 
pappus, since if not mature, it is colorless, as in L. ligulistylis, 
but purple when mature. 

'There is considerable difference in the appearance of the pappus 
of Liatris species. In L. punctata for instance, the tuft of bristles 
appears feathery or plumose to the naked eye. Upon closer 
examination there are found 20-30 setae, from 8-10 mm. in 
length, with lateral cilia considerably longer than the diameter 
4 of the seta, a condition called setose-plumose by J. Small, New. 

& Phytol. xvi.—xviii. (1917-1919). The bristles are arranged in- 
definitely in one to two series. In other species, for example L. 
gracilis Pursh, where there are 30-40 bristles 4-5 mm. long per 
achene, there is no downy appearance, as the setae have pro- 
jecting outgrowths about as long as, or only slightly longer than, 
the diameter of the seta—this called setose-barbellate (Small, 1. ¢.). 
This character was used by Cassini in the division of the genus 
into three subgenera, two of which, Suprago and Euliatris, were 
made sections by DeCandolle; the third is now a separate genus, 
Trilisa DC. These sectional divisions of the genus are here 
retained. 

Though there is never uniformity in the length of all the 
bristles of an achene, the over-all length of the pappus, obtained 
by measuring the longest bristles, generally exceeds the length of 
the achene, and is shorter than, or about equalling, the corolla. 
In a few cases as in L. Helleri Porter, it is only about half the 
length of the corolla-tube. 


KEY TO THE SECTIONS AND SERIES 
a. Pappus barbellate, the lateral cilia only 3-6 times the diam- 
eter of the seta, so that they are har diy visible without the 
use of a lens. Section I SUPRAGO (Cass.) DC....b. 
b. Heads 3-20-flowered. oblong, with phyllaries mostly erect... .c. 
c. Inflorescence spicate, the heads mostly sessile; leaves 
numerous, gradually reduced to bracts subtending the 
heads; no pilosity within the corolla-tube. . . .d. 


1946] Gaiser,—The Genus Liatris 177 


d. Phyllaries obtuse, appressed and never recurved....... SPICATAE. 
d. Phyllaries acuminate and recurved at the tips... . PYCNOSTACHYAE. 
c. Inflorescence loosely spicate or racemose, with sessile or 
variously peduncled heads; corolla pilose within the 
tube....e. 
e. Phyllaries obtuse, ciliolate; inflorescence frequéntly 
panicled; heads 5-15-flowered................. GRAMINIFOLIAE. 
e. Phyllaries lanceolate-acuminate, appressed to the tips, 
not ciliolate; heads 3-5-flowered................. PAUCIFLORAE. 
c. Inflorescence spicate, with the basal rosette of leaves 
changing abruptly to setaceous bracts; corolla-tube 
not pilose, but with short hairs on the filaments of the 
stamens; heads with 3-6 flowers and few loosely erect 
phylane n DIOE Lr ee IS res TENUIFOLIAE. 
b. Heads 15-70-flowered, hemispheric; the numerous broad 
phyllaries loosely erect, bullate or partly recurved; 
corolla pilose within the tube (except in L. ligulistylis) . .. .SCARIOSAE. 
a. Pappus plumose, the lateral cilia 15 or more times the diam- 
eter of the seta, so that these appear plumose to the naked 


eye. Section II Eunrarris (Cass) DC..... fe 
f. Heads 4-8-flowered, dence yn inflorescence spi- 
cate....g. 
g. Phyllaries with prolonged petaloid tips: corolla not at all 
puosezwithinjo 5 0 eis EN en EU hrs ae ELEGANTES. 
g. Phyllaries herbaceous and appressed, or with tips only 
free; corolla quite pilose within the tube............. PUNCTATAE. 


f. Heads 15-60-flowered, of more nearly isodiametric-cylin- 
drical proportions; inflorescence loosely cymose to cy- 


mose-racemose....h. 
h::Phyllifiés appressed ariede aar as). Oats LBS CYLINDRACEAE. 
h. Phyllaries recurved or loosely spreading.............. SQUARROSAE. 


DESCRIPTION AND CLASSIFICATION OF THE SPECIES 


SERIES I. SPercATAE. Mostly glabrous plants with numerous 
spikes from globular to large-crowned, perennial stocks; leaves 
linear, gradually diminishing upwards from the long basal ones; 
heads numerous, 4-18-flowered, 1-1.5 cm. long with phyllaries 
erect, oblong, mostly obtuse, appressed, never recurved; corolla- 
tube non-pilose within; achene 4-7 mm. long.—From New York 
and southern Ontario to South Dakota and Colorado, south to 
Florida, the Gulf States and New Mexico, with only one species 
not of moist habitats. 


a. Flowering stems stout, 6-15 dm. high; spikes dense; heads 
4—18-flowered; phyllaries mostly obtuse. ...b. 
b. Plants with narrowly linear leaves; on moist meadows or in 
damp woodlands east of the Mississippi Vue M 1. L. spicata. 
b. Plants with broadly linear leaves; on the bottomlands in 
the Western Plains States and the mountains of New 
Mexico as s c eee NM E Nr 2. L. lancifolia. 
a. Flowering stems more flexible, 3-8 dm. high; head 3-6- 
flowered; phyllaries obtuse, 'obtuse-mueronulate or acu- 
minate....c. 
c. Glabrous plants; corms rounded. . . .d. 


178 Rhodora [AUGURT 


'd. Phyllaries narrowly oblong, obtuse; leaves «narrowly 
linear, diminishing gradually upwards; more tufted 
plants with several stems oi fewer heads in racemes 
» 8. L. microcephala. 
d. Phyllaries ovate to lanceolate, acuminate; basal leaves 
long, narrowly linear, the upper cauline ones abruptly 
changing to short subulate bladés; plants of 1—4 spicate 
Blems:.« Sees scele desee es be 4. L. acidota. 
c. Hirsute plants; roots elongated and ‘tuberous; spicate; 
phyllaries oblong and obtuse-mucronulate; from South — . 
Florida Only ......llsseeeeseeeeee e 5. L. Garbert. 


1. Liarris sPICATA (L.) Willd. Rootstock globose in young 
plants, enlarged and shallow in old plants by separation of parts 
permitting considerable vegetative propagation: stems stiff and 
tall, 6-15 dm. high, glabrous, only rarely hirsute: leaves numer- 
ous, linear or linear-lanceolate, glabrous or sparingly hirsute 
along the veins; lower ones 1—4 dm. long, 5-20 mm. wide, gradu- 
ally shorter toward the summit of the stem: inflorescence a 
dense spike 3-7 dm. long: heads subcylindrical, 4—18-flowered, 
usually sessile along the spike, although basal heads may become 
peduncled, 8-15 mm. long and 5-10 mm. thick at time of flower- 
ing; phyllaries appressed, sometimes glutinous, elliptical-oblong, 
mostly obtuse but sometimes slightly acuminate, herbaceous, 
glabrous and having a narrow scarious margin frequently pur- 
plish at flowering time; corolla phlox-purple, lacking any pilosity 
within the tube, 6.5-9 mm. long; achene 4—6 mm. long; pappus 
5-7 mm. long, barbellate-setose (not plumose to the naked eye).— 
Sp. Pl. iii. 1636 (1803). Serratula spicata L. Sp. ii. 819 (1753) 
(excl. synonomy). 

Var. typica. A thick-spiked, marsh-loving plant mostly from 
the more northern latitudes of the range of the species and around : 
mountain lakes and bogs in the southern Appalachian region: 
stems frequently 5 mm. in diameter at the base and 10-15 dm. 
tall, glabrous or basally with few scattered hairs: leaves glabrous; 
the wider basal ones linear-lanceolate, 1 dm. or more long, 5-20 
mm. wide: inflorescence usually a dense spike 3-7 dm. long; the 
heads of 10-18 flowers 1-1.5 em. long and ca. 1 cm. thick, cylin- 
drieal, with an appressed involucre that is sometimes adherent 
by its glutinous nature; phyllaries glabrous, mostly green or 
somewhat purplish at time of flowering, with a narrow scarious 
margin: corolla usually phlox-purple, occasionally white.— 
Serratula spicata L. Sp. ii. 819 (1753), excl. synonyms., Liatris 
spicata Willd. Sp. Pl. ii?. 1636 (1803); Andr. Bot. Repos. t. 401 
(1804); Curtis's Bot. Mag. t. 1411 (1811); Sweet, Brit. Fl. Gard. 
ser. 2, t. 49 (1823); Torr. & Gray, Fl. N. Am. ii. 73 (1841); Torr. 
Fl. N. Y. 325, t. 47 (1842); Gray, Synop. Fl. i. 111 (1884). 
L. macrostachya Michx. Fl. Bor.-Am. ii. 91 (1803). L. spicata 
$. macrostachya DC. Prodr. v. 130 (1836). L. magnifica Hort. 


1946] Gaiser, — The Genus Liatris 179 


Hand-List of Herbaceous Plants Cultivated in the Royal Gar- 
dens, Kew 263 (1895), nomen. L. spicata f. albiflora Britton, Bull. 
Torr. Bot. Club, xvii. 124 (1890). Lacinaria spicata var. albi- 
flora Britton, Mem. Torr. Bot. Club, v. 314 (1894) Laciniaria 
spicata f. albiflora House, Bull. N. Y. State Mus. 243-4, 69 (1923). 
Lacinaria spicata var. foliacea Farwell, Amer. Mid. Nat. ix. 260 
(1925). 

Moist or marshy land from Long Island, New York, to Florida, 
and westward to the Mississippi, from the Lake St. Clair region 
of Ontario and Michigan to the Gulf of Mexico.— M ASSACHU- 
SETTS (possible garden escapes). Essex Co.: Lawrence, with- 
out collector's name, 1877 (G). WORCESTER Co.: roadside near 
Quinsigamond, Sept. 21, 1932, N. P. Woodward (G). CON- 
NECTICUT (possible garden escape). Without stated locality, 
Hitchcock (NY). NEW YORK. ArBaNY Co. (possible escapes): 
sandy plains, near Londonville, Aug. 14, 1937, H. D. House, 
24958 (G, NY); near Londonville, Aug. 13, 1934, H. D. House, 
121980 (G, NY). Surrork Co.: White Mills, Long Isl., Sept. 19, 
1887, J. F. Poggenburg (G). QurENS Co.: swamps, Woodhaven, 
Long Isl. Sept. 14, 1892, A. Brown (NY); (albino), Forbell's 
Landing, Long Isl., Aug. 29, 1890, G. D. Hulst (NY). PENN- 
SYLVANIA. Without stated locality: Aug. 1832, C. J. Moser 
(NY). BUTLER Co.: low field, Aug. 8, 1923, S. S. Dickey, 76 
(G). Bucks Co.: near Argus, Ridge Valley, Aug. 14, 1923, C. D. 
Fretz (G). Monraomery Co.: thicket in old brick-yard near 
West Telford, Aug. 23, 1909, W. M. Benner (G). DELAWARE 
Co.: open woods, Aston Mills, Oct. 16, 1926, F. W. Pennell, 
13118 (NY). Berks Co.: rather moist soil in a low meadow, 
2.4 mis. n. e. of Geigertown, Aug. 1, 1942, W. C. Brumbach, 3388 
(G); CHESTER Co.: Oxbow of Octoraro Creek, above Octoraro, 
Sept. 23, 1928, F. W. Pennell, 14607 (NY); LANCASTER Co.: 
without stated locality, Aug. 1858-64, S. P. Sharples (G), Sept. 
18, 1868, T. C. Porter (US). York Co.: McCall's Ferry, Sept. 9, 
1893, A. A. Heller & E. G. Halbach, 1273 (NY, US); dry woods, 
MeCall's Ferry, Sept. 2, 1898, A. MacElwee (US). NEW 
JERSEY. Without stated locality, A. Gray (G), P. D. Knies- 
kern (G).. Sussex Co.: Franklin Furnace, Aug. 1, 1884, O. E. 
Pearce (US); Aug. 15, 1895, W. N. Van Sickle (US). Passaic Co.: 
Clifton (white form), Aug. 18, 1891, G. V. Nash (NY); moist 
ground along rwy., Newfoundland, Aug. 3, 1893, W. M. Van- 
Sickle (US). SoMERsET Co.: in rich woods, Watchung, Aug. 8, 
1930, H. Moldenke, 1358 (NY); in field at edge of woods, on 
First Mt., Watchung, July 31, 1937, H. Moldenke, 9996 (NY); 
along roadside, in woods, Watchung, Aug. 1, 1924, H. Moldenke, 
2111 (NY); near Bernardsville, (forma albiflora), Aug. 12, 1890, 


! A note on this sheet states: ''spreading from cultivation’’, so that it seems probable 
that when introduced in a favorable location, this species may become naturalized. 


180 Rhodora [A UGUST 


Miss A. M. Vail (NY). MippLEsEX Co.: Perth Amboy, Aug. 
25, 1893, L. H. Lighthipe (US). Ocean Co.: Bayhead, Barnegat 
Bay, Aug. 24, 1892, J. R. Churchill (G); Bayhead (forma albi- 
flora), Aug. 29, 1890, L. H. Lighthipe (NY); border brackish 
marsh, 2 mis. e. of Manahawkin, Aug. 13, 1936, J. M. Fogg, 
11172 (G). DELAWARE. NrwcAsTLE Co.: near Delany’s 
Chapel, Sept. 1899, W. M. Canby (G). Kent Co.: Brandywine, 
1843, E. Tattnall (G). MARYLAND. BarvriMonE Co.: Ca- 
tonsville, Aug. 10, 1873, Morong (NY), E. Foreman (NY). 
VIRGINIA. Without stated locality: Dr. Bauer (NY). 
Farrrax Co.: (possible escape), near mouth of Difficult Run, 
Sept. 25, 1909, F. W. Pennell (US). Monrcomery Co.: Blacks- 
burg, Aug. 3, 1895, W. A. Murrill (NY). Grves Co.: Brush Mt., 
2 mi. e. of Newport, Aug. 30, 1933, E. J. Alexander, J. H. Everett, 
& S. D. Pearson (NY). WEST VIRGINIA. Hampsuire Co.: 
Millbrook, Aug. 11, 1940, W. M. Frye, 154, 157 (NY). FATETTE 
Co.: Nuttall, Aug. l1, 1891, C. F. Millspaugh, 1115 (NY); New 
River, Cotton Hill, July 6, 1929, W. V. U. Biol. Exped. (G). 
NORTH CAROLINA. BuNCOMBE Co.: slopes of Cedar Cliff 
Mt., Aug. 24, 1897, Biltmore Herb., 579 d (NY, US); Biltmore, 
Aug. 1894, Biltmore Herb. 579 (US). Hrnprrson Co.: Hender- 
sonville, Aug. 22, 1891, J. D. Smith (G, US). TRANSYLVANIA 
Co.: Green Knob in Pisgah Forest, Aug. 1908, H. D. House, 
3682 (G). SOUTH CAROLINA. KrxnsHAw Co.: Camden, 
July 27-28, 1906, H. D. House, 2658 (NY, US). Braurort Co.: 
Beaufort district, 1882, J. H. Mellichamp, 399 (US); Bluffton, 
1886, J. H. Mellichamp, 9, 10 (US); GEORGIA. CATOOSE 
Co.: Chickamauga Creek, near Springgold, Aug. 6-12, 1895, 
J. K. Small (NY, US). Dapk Co.: along C. &. D. Rwy., Look- 
out Mt., July, 1898, A. Ruth, 675 (NY); Lookout Mt., July, 1898, 
A. Ruth, 658 (US). RicuMonp Co.: Augusta, 1832, Drummond 
(G). SuMTER Co.: sandy bog, Aug. 26, 1896, R. M. Harper 
(NY). CorqvurrT Co.: moist pine barrens, between Moultrie & 
Kingwood, Sept. 22, 1902, R. M. Harper, 1652 (NY). FLORIDA. 
Without stated locality, ex. Herb. Chapman, Columbia College 
(NY). FRANKLIN Co.: Apalachicola, Aug. 20, 1872, Biltmore 
Herb., 579 b (G). Bav Co.: open sandy moist ground, Lynn 
Haven, Oct. 12, 1911, C. Billington, 168 jus). WALTON Co.: 
boggy places, in pine barrens, Argyle, Oct. 2, 1901, A. H. Curtiss, 
6923 (G, NY); moist soil, in pine barrens, ‘Crestview, April 21, 
1899, Biltmore Herb., 579 a (US). ORANGE Co.: ‘flatwoods, 
Vineland, Oct. 17, 1929, F. Vasku & E. West (NY). ONTARIO. 
LAMBTON Co.: meadows, Squirrel Isl., O. A. Farwell, 7103 (G, 
isotype of Lacinaria spicata var. foliacea) ; Sarnia, Aug. 11, 1894, 
C. K. Dodge (G, Ot); borders of marshes, Pt. Edward, R. St. 
Clair, Aug. 11, 1884, J. Macoun (G, Ot); marshes, Pt. Edward, 
Aug. 12, 1901, J. Macoun, 26616 (NY, Ot); meadows on Squirrel 


1946] Gaiser,—The Genus Liatris 181 


Isl., Sept. 3, 1924, O. A. Farwell, 7102 (G). Essex Co.: Sandwich, 
July 27, 1901, J. Macoun (G, NY); damp thickets, Sandwich, Aug. 
4, 1892, J. Macoun, 22751 (G, NY, Ot); near Mineral Springs, 
Windsor, July 31, 1894, A. W. Cody (Q). MICHIGAN. TuscoLa 
Co.: near shore of Lake Huron, 9 mis. e. of Bay City, Aug. 3, 1940, 
H. A. Gleason, 9913 (NY). Kent Co.: Grand Rapids, Aug. 3, 
1886, A. A. Crozier (US, ND). Sr. CrArR Co.: banks of Gov- 
ernmental Canal, Lake St. Clair, July, 1893, T. Morong (NY); 
Port Huron, C. K. Dodge, Aug. 6, 1896 (G); Aug. 11, 1894 (NY); 
Fort Gratiot (within Port Huron), 1829, Dr. Pitcher (NY). 
OAKLAND Co.: near Lake Orion, Aug. 3, 1913, B. F. Chandler 
(US). WasHTENAW Co.: low marshy meadow, 1.3 mis. e. of 
Dexter, Aug. 20, 1937, F. J. Hermann, 9141 (G, NY, US). Jack- 
soN Co.: Watkins Station, Aug. 4, 1892, C. F. Wheeler (US); 
low ground, 10 mis. s. of Jackson, Aug. 24, 1906, S. H. Camp & 
D. R. Camp (US); without stated locality, Aug. 12, 1896, S. H. 
Camp & D. R. Camp (US). KarAMAzOO Co.: swampy soil, n. 
of Pawpaw Lake, Texas twsp., July 23, 1930, C. R. Hanes (NY); 
n. of Pawpaw Lake, the “Mud Hole" 4% mi. s. e. of Vicksburg, 
Aug. 8, 1940, F. W. Rapp, 3625 (NY). OHIO. Erie Co.: 
Castalia prairies, Aug. 7, 1895, E. L. Moseley (US). Lucas Co.: 
Dorr St., Toledo, Aug. 4, 1920, E. L. Moseley (G). Lorain Co.: 
Lorain to Huron, Aug. 24, 1924, R. J. Webb, 5476 (G), Aug. 24, 
1924, R. J. Webb & G. A. Cook, 1612 (G). TARK Co.: south 
swamp, Canton, Aug. 1912, June 25, 1912 (No. 13) Mrs. F. E. 
Case (US). CHAMPAIGN Co.: without stated locality, Aug. 9, 
1893, W. C. Werner (NY). Pickaway Co.: Kibler's bog, 4% mi. s. 
of Circleville, July 28, 1936, Bartley & Pontius, 39 (NY). Mownr- 
GOMERY Co.: Dayton, Aug. 15, 1881, A. Foerste (US). Ross 
Co.: Frankfort (some white), Aug. 9, 1935, D. Demaree, 11496 
(G, US). INDIANA. Co. UNDETERMINED: low ground, e. of 
Chicago, Aug. 30, 1891, W. S. Moffatt, 1627 (US). Laxe Co.: 
moist prairie l4 mi. n. of Griffith, Aug. 29, 1916, C. C. Deam, 
21324 (US); moist prairie ditch, !4 mi. n. of Griffith, Aug. 29, 
1916, C. C. Deam, 21330 (US); Hammond, old beaches ‘‘Lake 
Chicago", Sept. 14, 1909, E. S. Steele, 147 (G, US); Buffington 
to Pine, old beaches “Lake Chicago", Sept. 20, 1909, E. S. 
Steele, 181 (G); open place in oak woods, 4% mi. n. of Griffith, 
Aug. 29, 1916, C. C. Deam, 21342 (US); Whiting, Aug. 29, 1893, 
N. L. Britton (NY); dry sands, Pine, Aug. 31, 1895, L. M. Um- 
bach (US); low sands, Pine, Aug. 27, 1897, L. M. Umbach (US); 
swales, Pine, Aug. 19, 1898, L. M. Umbach (US). Nose Co.: 
low sandy and marl border, Eagle Lake, Sept. 14, 1916, C. C. 
Deam, 21889 (US). MansHaALL Co.: Lake Maxinkuckee, July 
12, 1899 (No. 731), July 28, 1899 (No. 848), B. W. Evermann, 
(US); near Lake Maxinkuckee, 1900, J. T. Scovell & H. W. 
Clark, 1430 (NY, US), Aug. 26, 1900, J. T. Scovell & H. W. 


182 Rhodora [AUGUST 


Clark, 848 (NY); Plymouth, Sept. 21, 1909, H. W. Clark (US). 
STARKE Co.: low sandy border, s. e. side of Bass Lake, Aug. 22, 
1916, C. C. Deam, 21084 (US); open places in sandy woods, s. e. 
side of Bass Lake, Aug. 22, 1916, C. C. Deam, 21038 (US); sandy 
soil, Sept. 28, 1940, C. M. Elk (NY). Newton Co.: fallow 
sandy field, 5 mis. n. & 1 mi. w. of Enos, July 27,1940, R. C. 
Friesner, 14697 (NY). Tipron Co.: w. of Goldsmith, Aug. 2, 
1913, Mrs. C. C. Deam, 13921 (G, US); along the right of way of 
Erie Rwy., 4% mi. w. of Goldsmith, Sept. 3, 1914, C; C. Deam, 
15362 (NY). ILLINOIS. McHenry Co.: (albino), McHenry, 
Aug. 23, 1935, H. C. Benke, 5760 (G). Cook Co.: Chicago, H. 
H. Babcock, 1860, Dr. Scammon (NY, US); South Chicago, 
76th & Stoney Isl., Aug. 4, 1913, H. H. Smith, 5737. (G); waste 
places, Chicago, Aug. 16,.1897, L. F. Ward (US); in vacant lot, 
Chicago, Sept. 2, 1893, L. F. Ward (US); near Pullman, Aug. 27, 
1893, G.B. Sudworth (US); W. Pullman, near 119th St., Aug. 8, 
1907, J. M. Greenman, 1972 (G); along rwy., Lake Villa, Aug. 
10, 1906, H: A. Gleason & F. D. Shobe, 243 (G); wet intervals, old 
beaches, Indiana Harbor, Sept. 15, 1909, E. S. Steele 148 (US); 
Cicero, July 20, 1896, Mrs. A. Chase (NY); Riverdale, Aug. 
1909, J. M. Greenman, 2866 (G, US); vacant lot, Evanston, July 
4, 1911, E. E. Sherff (US); Morgan Park Ridge, Sept. 8, 1907, 
R. A. Dixon & C. A. Gage, 789 (US, Q). KENTUCKY. CALLO- 
WAY Co.: between Murray & Pine Cliff Ferry, July 23, 1937, 
L. B. Smith & A. R. Hodgdon, 4202 (G, NY, US). TENNES- 
SEE. Roserrson Co.: rocky hills, Aug. 19, 1897, H. Eggert 
(NY). Dickson Co.: dry oak barrens, Burns, Aug. 17, 1939; 
H. K. Svenson, 10485 (B). Cocke Co.:3 mis. w. of Wolf Creek 
Sta., Aug. 30, 1897, T. H. Kearney, 746 (NY). CUMBERLAND 
Co.: Mayland, Aug. 16, 1934, Porter & Harbison, 3059 (NY). 
VAN Buren Co.: grassy swamps, between Spencer & Cagle, Aug. 
1938, H. K. Svenson, 9678 (B). Correte Co.: oak barrens n. of 
Manchester, Aug. 6, 1938, H. K. Svenson, 8959 (G, B); swamp, 2 
mis. s. of Manchester, Aug. 18, 1940, H. K. Svenson, 10606 (B); 
low fields, near Tullahoma, Aug. 10, 1899, Biltmore Herb., 579e 
(NY, US); dry oak barrens, Tullahoma, Aug. 24, 1930, H. K. 
Svenson, 4231 (G, P, B). CuksTER Co.: borders of thickets; 
1892, S. Bain (US); fields, Henderson, S. M. Bain, June, 1892 
(No. 81) (NY), Aug. 1892 (No. 81) (G). FRANKLIN Co.: wet 
places in oak barrens, between Tullahoma & Estill Springs, Aug. 
13, 1939, H. K. Svenson, 10490 (B). ALABAMA. Without 
stated locality: M. Lea (G). Hare Co.: dry chalky prairies, 
1 mi. n. w. Rosemary, Aug. 23, 1934, R. M. Harper, 3254 (G. 
NY, US). Ler Co.: Auburn, Aug. 11, 1897, F. S. Earle & C.F. 
Baker, 1160 (NY). BvurLER Co.: grassy pine barrens, near 
Bolling, Aug. 28, 1885, J. D. Smith, 427 (US). Henry Co.: 
dryish swamp border, 8 mis. n. of Headland, Aug. 10, 1927, K. 


1946] Wilde, 


Betula glandulosa in New Hampshire 183 


M. Wiegand & W. E. Manning, 3174 (G). MISSISSIPPI. 
WAYNE Co.: Waynesboro, Aug. 8-9, 1896, C. L. Pollard 1248 
(G, NY, US, ND). JACKSON Co.: Ocean Springs, Aug. 14, 
1889, Herb. F. S. Earle (ND). Harrison Co.: Biloxi, Aug. 23, 
1898, S. M. Tracy 4886 (NY). WISCONSIN. Kenosua Co.: 
Pleasant Prairie, 3 mi. d of Kenosha, Aug. 7, 1941, E. P. 
Kruschke, K-41- 175 (G), K-41-179 (G) (albino). MISSOURI 
(possible garden escape). ST. Louris Co.: St. Louis, July 7, 
1910, E. E. Sherff, 288 (G). LOUISIANA (possible garden 
escapes). HRaPripEs Co.: Alexandria, J. Hale (6). ORLEANS 
Co.: New Orleans, 1835, Dr. Ingalls (NY). 


(To be continued) 


BETULA GLANDULOSA AT A LOW ALTITUDE IN New HAMPSHIRE. 
—While vacationing in Jackson, N. H., in late September, a few 
years ago, we drove up the Black Mountain Road, turning right, 
near Whitney’s, into the Dundee Road. After a short way we 
parked our car in front of a house, on the right side of the road— 
a house which we used to call “the house with the chimney 
outside." We walked through the yard, past the house and 
barn, and went a short distance into an open upland meadow- 
like pasture, to enjoy the view. This location was on the lower 
north-east slope of Tin Mountain, possibly 400: feet below the 
summit, which is only 2025 feet. 

As we sat there, I examined the low growth around me, and 
saw, to my great amazement, what looked exactly like: Betula 
glandulosa, which I had never collected before except in the 
alpine region of the Great Range. However, Professor Fernald 
has identified the. specimens I sent him and this establishes, 
evidently, an exceptionally low altitude: for B. glandulosa.— 
CHARLOTTE ENpicotr WILDE, Canton, Massachusetts. 


184 Rhodora [AUGUST 


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


IDENTIFICATIONS AND REIDENTIFICATIONS OF 
NORTH AMERICAN PLANTS 


M. L. FERNALD 
(Continued from page 162) 


HABENARIA PsYcODES (L.) Spreng., forma varians (Bryan), 
stat. nov. Var. varians Bryan in Ann. Mo. Bot. Gard. iv. 37, pl. 
5, fig. B (1917). 

H. riMBRIATA (Ait.) R. Br., forma mentotonsa, f. nov., labelli 
lobo terminali euneato integro vel apice breviter eroso-dentato; 
petalis integris.—MaiNEÉ: meadow, Hamilton Cove, Lubec, 
Washington County, August 2, 1909, Fernald, nos. 1662d (TYPE 
in Herb. Gray.), 1662e and 1662g. 

Quite like typical Habenaria fimbriata but with entire petals 
and narrowly cuneate entire or obscurely short-dentate or erose 
terminal division of the lip. Entire petals are frequent in both 
H. fimbriata and the smaller-flowered H. psycodes, and the lip of 
H. psycodes, forma ecalcarata (Bryan) Dole is entire. At Hamil- 
ton Cove H. fimbriata, forma mentotonsa (with shaved chin) is 
relatively common, mixed with typical H. fimbriata, but I cannot 
follow Correll who, in Bot. Mus. Lfls. Harv. Univ. vii. 65 (1938), 
calls this plant the characteristic slender-racemed one with 
greenish-white or rose-tinted flowers, the hybrid of H. lacera 
(Michx.) Lodd. and H. psycodes, the always scanty and relatively 
insignificant plant known as X H. Andrewsit Marcus White ex 
Niles, Bog-Trotting for Orchids, 258 with plate (1904). 

Correll’s vast aggregation of relatively typical Habenaria 
psycodes, H. fimbriata and H. lacera, var. terrae-novae Fern. in 
Rnopona, xxviii. 21 (1926) under the unsatisfactorily blanketing 
name X H. Andrewsii can appeal to no field-botanist who for 
decades has known the various elements involved. Much New- 
foundland M. lacera, var. terrae-novae is included under his re- 
modeled X H. Andrewsii, although no true M. lacera is found in 
Newfoundland, where its smaller-flowered var. terrae-novae occurs 
by thousands on boggy barrens, tundra and treeless alpine areas, 
almost always apart from H. psycodes of richer, often alluvial, 
thickets and meadows. On Sable Island, 100 miles out-to-sea off 


1946] Fernald,—lIdentifications of North American Plants — 185 


Canso, Nova Scotia, the only Fringed-orchid is H. lacera, var. 
lerrae-novae. 

Very similarly, although Habenaria fimbriata, forma mentotonsa 
occurs in eastern Washington County, Maine, it is significant 
that in the many pigeonholes of Fringed-orchids in the Her- 
barium of the New England Botanical Club I can find neither 77. 
lacera nor H. psycodes (parents of true X H. Andrewsii) from that 
county. Both seem to stop their eastern extension in coastwise 
Maine in Hancock County, 70-90 miles to the southwest of 
Cutler. 

Similarly, nine tenths of the specimens in the Gray Herbarium 
and that of the New England Botanical Club which have been 
annotated (some of them cited) as X Habenaria Andrewsii are 
characteristic H. fimbriata (including the TYPE of H. fimbriata, 
forma albiflora Rand & Redfield) or H. psycodes. X H. Andrew- 
sii, as well as Fleur-de-lis, Blackberry blossoms, Yellow Clin- 
tonia, Indian Pipes, “white, innocent twigs of apple" and other 
non-orchidaceous plants, was illustrated in Bog-Trotting for 
Orchids. The life-size photograph shows racemes 2-2.5 cm. 
thick; and the description calls for *Labellum about 14-14 inch 
(8-12.5 mm.] broad". In his very detailed account of H. psy- 
codes X lacera, Andrews, in Ruopora, ni. 246 (1901), said: 


“Lower leaves as in H. lacera . . . , width to 3 cm. . . . Average 
width of lip about 12 mm. . . . cleft asin H. lacera . . . Glands 
of pollen-masses . . . elliptical or slightly kidney-shaped"', and, 


on p. 247, “All in all the characteristics of the hybrid seem to 
show a stronger influence of H. lacera". "The distinctive charac- 
ters of H. lacera and of H. fimbriata, besides color and dissection 
of lip, include the following. H. LACERA: largest lower leaves 
1-3.5 cm. broad; raceme 2-6 cm. thick; perianth 5-6 mm. long; 
lip 1-1.5 em. long and broad, its terminal division cuneate into a 
very slender claw; glands of anther oblong-linear. H. FIM- 
BRIATA: largest lower leaves 2.5-9 cm. broad; raceme 5-9 em. in 
diameter; perianth 9-12 mm. long; lip 1.5-2 cm. long, 1.8-3 cm. 
broad, its dilated terminal division short-stalked or subsessile; 
glands suborbicular. In all except the narrow and fringeless 
terminal division of its lip H. fimbriata, forma mentotonsa is very 
characteristic H. fimbriata, growing, as said, far from H. lacera or 
H. psycodes. In view of these many considerations it is toler- 


186 Hhodora [AUGUST 


ably certain that the great group of amateur and professional 
botanists who have assembled the large representation of H. 
lacera, psycodes and fimbriata in the herbarium of the New 
England Botanical Club and in the Gray Herbarium, for the most 
part with correct identifications, have not all been wrong. 


CLEISTES DIVARICATA (L.) Ames, var. bifaria, var. nov. (TAB. 
1048), var. typica recedit planta plerumque 1.5-5 dm. alta 
pedunculo 0.3-1.6 dm. longo; sepalis longioribus 3-4.5 cm. 
longis; petalis 2-3 em. longis, 5-10 mm. latis.—Upland woods, 
mountain-crests and slopes, Cumberland Plateau and Mountains 
of Kentucky and Tennessee and Blue Ridge of western North 
and South Carolina, coming out to peats and pine barrens of the 
Coastal Plain from eastern North Carolina to Florida, thence to 
Louisiana. Map 2! Type from summit of Table-rock Moun- 
tain, Burke Co., North Carolina, July 2, 1891, Small & Heller, 
no. 285 (Gray Herb., isorvPEs in several other herbaria). 

Arethusa divaricata L. Sp. Pl. 951 (1753), typonym of Cleistes 
divaricata (L.) Ames, Orchidaceae, vii. 21, pl. 108 (1922), was 
based on Serapias radicibus palmato-fibrosis, caule unifloro of 
Gronovius from Virginia (photograph before me) and upon 
Catesby's plate 58 of his Helleborine Lilii folio caulem, ambiente, 
etc., represented as having an extraordinarily large flower (with 
sepals 6.4-7.3 cm. long, petals 6-7 cm. long and lip 7 em. long). 
The Clayton material, shrunken by drying, is more modest; its 
dried and distorted sepals up to 4.2 cm. long, petals to 3.6 em. 
and lip slightly over 4 cm. long. The Clayton material represents 
a small-flowered extreme of the plant which locally follows the 
Coastal Plain from southern New Jersey to northern Florida 
(MAP 1). The Catesby drawing is presumably exaggerated in 
size. I have had through the courtesy of the Curators the 
advantage of studying, besides that in the Gray Herbarium and 
the Ames Herbarium, all the material at the United States 
National Herbarium, the New York Botanical Garden, the 
Academy of Natural Sciences of Philadelphia and the Brooklyn 
Botanic Garden. These collections show that there are two 

! Since the map was engraved, specimens from additional sii oni in the Cumber- 
land Mountains of Tennessee have been sent me for study by Professor Jessé Mj 
Shaver of George Peabody College for Teachers, at Nashville. They add three dots 
for Tennessee. i , 

? One Florida specimen of 1888, bearing the intriguing data, ‘‘wedding trip’’, has 


not been entered on the map; neither have I selected it as the type of var. bifaria 
(in two parts or on two sides), 


Rhodora Plate 1047 


Photo B. G. Schubert 


CLEISTES DIVARICATA, all figs. X 1: FIGs. 1 and 2, median leaf and flower from 
eastern Virginia (type-region); FIG. 3, flower from southern New Jersey. 


Rhodora Plate 1048 


LES ^ 


Photo B. G. Schubert 


CLEISTES DIVARICATA, var. BIFARIA, all figs. X 1: FIG. 1, upper half of plant from 
TYPE-series; FIGs. 2-6, flowers from various localities. 


1946] Fernald,—Identifications of North American Plants 187 


Map 1, Range of typical CLEISTES DIVARICATA; MAP 2, of var. BIFARIA. 


rather strongly defined varieties passing as Cleistes divaricata. 
The plant which is confined to the Coastal Plain, northward to 
southern New Jersey but in the South not found west of northern 
Florida, seems to be true C. divaricata. This plant (our PLATE 
1047) in its best development is usually 4 or 5 dm. high, the whole 
series ranging from 2.2-7.2 (av. 4.5) dm. high, while the peduncle 
(between the base of the median leaf and the floral bract is 0.9-2 
(av. 1.5) dm. long. The median leaf ranges from 6.5-15 (av. 10) 
em. long; the lateral sepals 4-7 em. long; petals 3.5-5 (by Catesby 
shown up to 7) em. long and 8-14 mm. broad; the ovary and 
stipe during anthesis 2.5-4.5 (av. 3.25) em. long. 

Throughout much of the Southeast, from Florida to eastern 
North Carolina, west to Louisiana, chiefly on the Coastal Plain, 
and inland on the Blue Ridge (up to open summits) to North 
Carolina and on the Cumberlànd Plateau and Mountains of 
Tennessee and Kentucky, the plant is generally smaller in most 
parts, var. bifaria (from its two areas of development). In var. 
bifaria (PLATE 1048) the-stem is rarely 6:5 dm. high, usually 
ranging from 1.5-5 dm., with the peduncle 0.3-1.6 (av. 1) dm. 
long and the median leaf 3.5-13 (av. 7.6) cm. long. Its flower is 
conspicuously smaller, though sometimes approaching that of 
var. typica, with longer sepals 3-4.5 em. long, petals only 2-3 


188 Rhodora [AUGUST 


em. long and 5-10 mm. wide, and ovary and stipe during anthesis 
1.2-3.5 (av. 2.6) em. long. 

Occurring on the ancient Cumberland Mountains and Plateau 
and along the ancient Blue Ridge, var. bifaria seems to be the 
biological type of the species, which, on withdrawal of the 
Cretaceous and then the Tertiary seas from the country to the 
east and south, largely moved out to the Coastal Plain. "There, 
in new environment, it has given rise to the larger-flowered 
extreme (nomenclatural type of the species) which has followed 
locally northward to southern New Jersey. 

As indicating the confusion heretofore of typical Cleistes 
divaricata of the Atlantic Coastal Plain and var. bifaria of the 
southern Atlantic and the Gulf Coastal Plain, as well as the 
mountains, there is a sheet in the Britton Herbarium, originally 
in the herbarium of the late Professor Lewis R. Gibbes of the 
College of Charleston, South Carolina, labeled in the hand of Dr. 
John K. Small as from “Flat Rock, S. C.". This original label, 
which, like all original labels, would never be altered or written 
upon by those who fully respect original documents, was unfor- 
tunately altered by a later botanist, who knew the Carolina 
Mountains, to “N. C." instead of the original S. C., and the 
original label further desecrated by the misinformative addition 
“Henderson Co., N. C.". The specimen is of typical Atlantic 
Coastal Plain C. divaricata, which is apparently unknown in 
Henderson County or elsewhere on the Blue Ridge or on the 
Cumberland Plateau or Mountains. In view of the fact that 
Gibbes lived at Charleston and that there 1s another sheet of 
material with his original handwriting on the labels (one **Sum- 
merville, 20 May, 1859", the other “Flat Rock, 12 June, 1858. 
L. R. G.") it would seem that the specimen with unjustifiably 
altered label came from Flat Rock on Flat Rock Creek, which 
drains into Wateree River, a tributary of Santee River, in Ker- 
shaw County, on the Coastal Plain of South Carolina, north of 
Camden and northwest of Sumter and Charleston. 

CALOPOGON PULCHELLUS (Salisb.) R. Br., var. latifolius (St. 
John), stat. nov. Forma latifolius St. John in Proc. Bost. Soc. 
Nat. Hist. xxxvi. 69, pl. 1, fig. 4 (1921). Limodorum tuberosum, 
f. latifolium (St. John) House in Bull. N. Y. State Mus. no. 243- 


244: 51 (1923), as to name only. Cathea pulchella, f. latifolia (St. 
John) House, 1. c. no. 254: 244 (1924), as to name only. 


1946] Fernald,—Identifieations of North American Plants 189 


The original material is much more than a broad-leaved ex- 
treme of Calopogon pulchellus, a species which, even in the same 
area, may have the leaf varying from narrowly linear and only 
2-4 mm. wide up to lanceolate or lance-oblong and up to 2 cm. 
wide, while very extreme and gigantic plants (up to 9.75 dm. 
high), may have the leaf up to 3-5 cm. broad. In this typical C. 
pulchellus, either very narrow- or very broad-leaved, the leaf is 
usually solitary and much shorter than the elongate scape. The 
type of var. latifolius has the leaves often paired and broadly 
lance-oblong to narrowly oblong-ovate, only twice to six times as 
long as broad and greatly overtopping the very short scape, 
while its heavily dark-coated tuber is much larger (2 em. thick) 
than in any typical C. pulchellus I have ever seen. The type is 
past flowering, but other material, also from Sable Island, is 
flowering. This is narrower-leaved and has either paired or 
single leaves, although its scape is much shorter than to barely 
overtopping the leaf or leaves; furthermore, some material from 
the Magdalen Islands is stongly transitional to var. latifolius. 
This plant of Sable Island and, less typically, of the Magdalen 
Islands is not the Newfoundland Limodorum tuberosum, var. 
nanum Nieuwland in Am. Midl. Nat. iii. 130 (1913). The latter 
is merely typical Calopogon pulchellus at its bleak northern limit, 
0.7-2 dm. high, with scape much overtopping the leaf, the raceme 
reduced to 1—4 flowers, merely the smallest extreme of the species, 
just as plants of southeastern Virginia 6-9.75 dm. (pretty close 
to 1 m.) high, with the leaf 3-5 ecm. broad and the 10-20 flowers 
4—4.5 em. broad, are the largest. The paired and short leaves 
and the large tuber of the type of var. latifolius give the plant 
(past flowering), as shown in St. John's figure, the aspect of 
Liparis! 

SPIRANTHES TUBEROSA Raf., var. Grayi (Ames), comb. nov. 
S. Grayi Ames in RHopona, vi. 44 (1904). S. simplex Gray, 
Man. ed. 5: 506 (1867), not Griseb. Fl. Brit. W. Ind. 641 (1864). 

As noted by me in RHopora, xlviii. 6 and 10 (1946), the name 
Spiranthes tuberosa Raf., Herb. Raf. 45 (1833) antedates by seven 
years the name S. Beckii Lindley (1840), the latter name cur- 
rently used for the very slender and tiny-flowered plant which 
Ames correctly, except for the overlooked S. tuberosa, named S. 
Gray? in 1904. It is fortunate, at least, to be able to dismiss the 


190 ; Rhodora [Aveusp 


name S. Beckii, for Lindley made a sad mess of his original publi- 
cation of it in his Genera and Species of Orchidaceous Plants, 472 
(Sept. 1840). There, in the fashion of many British botanists of 
his day (and too often of the present day), he chose the British 
use of the name S. gracilis, rather than the earliest use of it. 
Consequently, he took up S. gracilis, as of Hook. (we now would 
say sensu Hook.), Fl. Bor.-Am. ii. 202, t. 203 (1839), with the 
synonymy copied directly from Hooker.. Hooker mis-cited the 
combination as starting in Bigelow, Fl. Bost. ed. 2: 322 (1824), 
Bigelow having called it Neottia gracilis: Hooker cited his. 5S. 
gracilis (Bigel.) Hook. as having the “Has. Canada; and Lake 
Huron (Dr. Todd) to Fort Franklin, on the Mackenzie River. 
Dr. Richardson. Drummond” and. his plate beautifully showed 
the Canadian S. lacera (Raf.) Raf., l. c. 44 (1833), discussed and 
illustrated by me in Rnopona, |. c. 5-9, pl. 993 (1946). Lindley, 
maintaining S. gracilis sensu Hooker: (1839), assumed that 
Drummond, who actually explored northward to northern Cana- 
da, had collected it much farther south, consequently he inter- 
preted the Drummond. citation given by Hooker as meaning 
"Louisiana", then for good measure he added *'etiam in. Baha- 
mis"! S. gracilis (Bigelow) Beck, Bot. 333 (1833) and 5$. 
gracilis (Bigelow) [sensu] Hooker (1839), although two different 
species so far as the plants are concerned, both go back nomen- 
claturally tothe same type. 

Having thus temporarily saved the:name Spiranthes gracilis 
sensu Hooker (1839), Lindley's next problem was to dispose of 
the earlier S. gracilis (Bigelow) Beck (1833). That was quickly 
accomplished by renaming the latter S. Becki? Lindl..l. c. (1840), 
with the additional synonyms Neottia tortilis [sensu] Elliott 
(1822) [not Swartz (1800)], and Limodorum praecox Walt. (1788) 
basis of S. praecox (Walt.) S. Watson (1890). Nomenclaturally 
alone the name S. Becki? Lindl. is doubly illegitimate. If it was, 
as he said, the same as the earlier S. gracilis (Bigelow) Beck he 
should have used the latter name for it; if, however, it was also 
the same as Limodorum.. praecox Walt, (1788) Lindley should have 
retained this specific name. Taxonomically, furthermore, S. 
Beckii Lindl. was as hopeless a muddle as could be imagined, for 
it was concocted from elements of several different species. 
Limodorum praecox Walt., originally deseribed with fibrous roots 


1946] Fernald,—Identifications of North American Plants — 191 


and ensiform leaves (“radicibus fibrosis, foliis ensiformibus’’) 
etc., is a plant with long and mostly linear firm leaves extending 
up the stem, the relatively coarse spike with heavily pubescent 
rachis, bracts and ovaries, the perianth 4-6 mm. long, etc., the 
perianth of S. tuberosa being only 2-3 mm. long. Nevertheless, 
Lindley described his S. Becki? as “perfectly glabrous. The 
flowers are very minute . . . S. glaberrima, foliis omnibus radi- 
calibus anguste ovalibus" etc. If, furthermore, it were N. 
tortilis sensu Elliott, it would be very difficult to reconcile 
Lindley's description with Elliott’s “foliis radicalibus linearibus 
. . . Stem pubescent towards the summit. Leaves . . . of the 
root linear lanceolate, nine to ten inches long . . . Bracteal 
leaves pubescent” ete. In view of the vertical, finger-like, usual- 
ly solitary tuber of S. tuberosa (“S. Beckii" of most recent 
authors) it is illuminating that Lindley knew nothing of this 
character nor did those authors with whose descriptions he as- 
sociated his name. Furthermore, since his S. Beckii was '' per- 
fectly glabrous" as is S. tuberosa, it is significant that Lindley 
said in his Latin diagnosis ‘‘ovario puberulo", a character be- 
longing to S. praecor. The ‘lip [with] . . . a remarkably lax 
cellular texture" applies to S. tuberosa, but the description and 
cited synonyms otherwise are so confused that it is certainly 
fortunate that the name given by Lindley is illegitimate.! 

Spiranthes tuberosa consists of two strongly marked geo- 

! Hooker, under his Spiranthes gracilis (i. e. S. lacera), a plant with glabrous inflores- 
cence, and which Hooker correctly described ''foliis radicalibus ovatis petiolatis”, 
cited as synonyms the much earlier Ophrys aestivalis Michx. Fl. Bor.-Am. ii. 157 (1803) 
and Neottia tortilis Pursh, Fl. Am. Sept. ii. 589 (1814), “(non Sw.)". Lindley, under 
Spiranthes gracilis, cited the same synonyms.  Evidently neither he nor Hooker 
studied very closely the Michaux description [and specimens] nor the description by 
Pursh; otherwise they would not have cited them under the wholly glabrous S. gracilis, 
with leaves all basal and ovate, for Michaux definitely described his Ophrys aestivalis: 
“O. scapo folioso: foliis glabris, lanceolatis, acutissimis; spica pubescente, spirali” etc. 
and he suspected that it might be the Limodorum praecor of Walter, O. aestivalis 
occurring ‘ʻa Pensylvania ad Carolinam’’. The type of Ophrys aestivalis. a species 
which I do not find accounted for in recent American literature, as shown in one of 
Cintract’s photographs before me, consists of two full plants, with linear-lanceolate 
leaves extending up the stem, the longer blades about 2 dm. long. the slightly spiraling 
to secund spike with perianths 6 mm. long. Mounted with these two plants is a 
broken-off spike of Spiranthes cernua, which obviously was an inadvertent addition 
made by the mounter. Ophrys aestivalis Michx. (1803) is Spiranthes vernalis Engelm. 
& Gray (1845). Most fortunately, we do not have to displace the latter name, for 
there is an Old World Spiranthes aestivalis Richard (1818). 

As to Pursh's misidentification of Neottia tortilis, we need not here go into details, 


except to note that Pursh included under it Ophrys aestivaiis Michx., gave the same 
range as the latter, and described the leaves as linear. Enough said! 


192 | Rhodora [AUGUST 


graphic variations. Essentially all the material in the Gray 
Herbarium and that of the New England Botanical Club from 
New England, forty-five collections, has a relatively close spike 
with closely spiralling and often crowded and overlapping flowers, 
as in the type of S. simplex Gray, not Grisebach. This plant 
varies from 0.7-3 dm. (farther south to 4.5 dm.) in height, and 
its vertical tuber is thick and finger-like, usually solitary. This, 
as said, is the plant described by Gray as S. simplex and correctly 
renamed by Ames S. Grayi. All the material in the Gray Her- 
barium from the southernmost states, from Florida to eastern 
Texas, north to South Carolina, has the spike strongly secund, 
without or with few spiral twists in the rachis and the relatively 
few flowers distant and not overlapping. From North Carolina 
to New Jersey both variations, with some transitions, occur, the 
plant often reaching a height of 5.25 dm., while its roots are 
usually more slender and not infrequently 2 or even 3. This is 
true S. tuberosa Raf. which was described with “spic. gracilis vix 
spiralis secunda . . . pedal." 

Dr. Schubert has made dissections of flowers from several speci- 
mens of each extreme and, while each series shows some variation 
in the degree of toothing and shape of the lip, there appears to 
be nothing constant except the relatively dense and strongly 
spiralling spike and usually thicker tuber to separate var. Gray: 
from the usually more southern typical S. tuberosa. 

Gray's Spiranthes simplex, type of S. Grayi and of S. tuberosa, 
var. Grayi, had “scape... bearing a small narrow (rarely 
1-sided) spike of very short flowers (perianth 1"—1Y$" long)". It 
came from *E. Mass. (Nantucket, Dr. Robbins), New Jersey 
(C. F. Austin, &c.), and Delaware, Wm. M. Canby." Gray’s 
original sheet contains the Nantucket material from Robbins, 
which is the dense-spiked S. tuberosa, var. Gray?; a series of six 
quite similar plants collected by himself (''&c.") in the pine 
barrens of New Jersey (the Austin material evidently not re- 
tained by him), and three characteristic plants (one of them 
misplaced by the mounter) with the “rarely 1-sided" spike from 
Canby, but marked as from “Salisbury, Maryland" (not ‘Dela- 
ware"), this Canby material being of typical S. tuberosa. 


1946] Fernald,—Identifications of North American Plants — 193 


CORALLORHIZA, NOT CORALLORRHIZA.—From the first edition 
of Gray’s Manual (1848) through the 6th edition (1890) the 
saprophytic woodland Coral-roots were rightly called Corallo- 
rhiza, although the genus was ascribed to Haller, whose definition 
of it was prior to 1753, in his Enum. Meth. Stirp. Helvet. i. 278 
(1742), Haller, who went back to Ruppius, then spelling the 
generic name Corallorhiza. In the 7th edition of Gray’s Manual 
(the Orchidaceae revised by Professor Oakes Ames) Haller was 
bracketed as the author prior to 1753, the post-Linnean author 
given as Robert Brown; and the spelling was changed to Corallor- 
rhiza. Although Robert Brown was there and in the later com- 
pendium of Ames, his Enum. Orch. U. S. and Can. 21 (1924), 
made the first post-Linnean author of the genus, Brown himself 
had cited the genus as starting after 1753 in Haller's Hist. Stirp. 
Helvet. ii. 159 (1768). That was correct, so far as it went, and 
Haller in 1768 had adopted the better Greek spelling, Corallor-. 
rhiza. Brown gave the common circumboreal species the specific 
name C. innata R. Br. in Ait. Hort. Kew. ed. 2, v. 209 (1813). 

In Gray’s Manual, ed. 7, and in his Enumeration of 1924 Ames 
took up for the original species of the genus, the latter said by him 
to date from 1813, a binomial dating from 53 years prior to 
Brown's publication, a case of putting prophecy before history 
which has puzzled many students, for the genus Corallorhiza 
and its species C. trifida were both clearly and very adequately 
published in Chatelain's Specimen inaugurale de Corallorhiza 
in 1760, the genus clearly diagnosed on p. 6, the species on p. 8. 
Here, so far as I can find, is the initial date (after 1753) for both 
CORALLORHIZA and its original species, C. trifida, which was 
based on Ophrys Corallorhiza L. (1753). We thus get rid of the 
situation wherein a binomial seems to have been published 53 
years earlier than the genus under which it was placed; but, at the 
same time, we can return to the long-familiar spelling of the 
generic name, since, by the International Rules of Nomenclature, 
the original spelling (in this case of Haller in 1742 as well as of 
Chatelain in 1760) must stand'. The correction of the first post- 


! Since the above was written the similar decision of Rendle and Britten in Journ. 
Bot. xlv. 442 (1907) has come to my attention: '''This genus was established by J. J. 
Chatelain ‘Specimen inaugurale de Corallorbiza' 1760. He names the species C. 
TRIFIDA, Which must stand, as the Linnean trivial Corallorhiza (under Ophrys) is 
inadmissable''. 


194 Rhodora [AUGUST 


Linnean author of the genus (but incorrectly as Corallorrhiza) 
was made in Britton & Brown, Ill. Fl. ed. 2, i. 574 (1913) but, 
singularly enough, in a work seeming to be authoritative, Schlech- 
ter’s Monographie der Gattungen und Arten in Keller & Schlech- 
ter, Monographie und Iconographie der Orchideen Europas und 
des Mittelmeergebietes, Fedde, Rep. Spec. Nov. Sonderbeiheft 
A, Lief. 9-10, 302, 303 (1928), the anomaly again appears: the 
genus Corallorhiza here started from Robert Brown in 1813, but 
its single European species given as “1. C. trifida Chatel., Spec. 
inaug. Corall. (1760), p. 8”! 

In current works on the flora of the northeastern United States 
the lip of Corallorhiza trifida is described as “white, not spotted” 
(Gray’s Man. ed. 7), “lip unspotted” (Wiegand & Eames, FI. 
Cayuga Lake Basin), “lip usually pure white" (A. M. Fuller, 
Studies on the Fl. Wisc. Part I: The Orchids), etc.; though rarely 
in America it is described, as by Morris & Eames (Our Wild 
Orchids), as ‘almost as often spotted as unspotted". Their 
discussion, however, shows that no distinction was being made 
between plants of North America and those of northern Eurasia 
and that they included Canada to the Arctic. In view of the 
usual lack of red or purple mottling of the lip in the United 
States and southernmost Canada it is worth noting that Chate- 
lain, in his original account of European C. trifida, said ‘‘labellum 

. album, punctis coccineis notatum", while Schlechter, 1. e., 
describing the European plant, says “die Petalen zuweilen 
rotpunktiert, Lippe weiss rotpunktiert”’. 

In 1916, Cockerell in Torreya, xvi. 231, getting in Colorado the 
common plant of the United States, with “lip whitish”, described 
it as Corallorhiza Corallorhiza coloradensis n. subsp., he then 
separating it because the true European plant, as shown by the 
enlarged figures of flowers published by H. Müller, has the throat 
“dotted with dark pigment". Almost a century earlier, however, 
Thomas Nuttall clearly understood the situation when he mono- 
graphed our species in his Remarks on the Species of Corallorhiza, 
indigenous in the United States in Journ. Acad. Nat. Sci. Phila. 
iii. 135-139, with plate (1823). Nuttall there defined his new 
C. "verna . . . petalis omnibus lineari-lanceolatis patentibus, 
labello oblongo immaculato basi bidentato apice recurvo ovato 
calcare obsoleto innata . . . whole plant except the lip, of a 


1946] Fernald,—Identifications of North American Plants — 195 


yellowish-green colour . . . Lip nearly white, without spots", 
etc. This species, C. innata in the sense of Muhlenberg, Amos 
Eaton and Nuttall's Genera, was based primarily on material 
from New England and in his *OBsERvATION" Nuttall wrote: 
“Mr. Eaton justly remarks the discrepancy of this plant with the 
species which I had erroneously considered the Corallorhiza 
innata of Europe"; but, pursued by the fatality which so often 
confuses those who attempt clarification, Nuttall proceeded in 
his discussion to ascribe to his new eastern American C. verna 
(which he had just correctly defined as “whole plant except the 


lip, of a yellowish-green colour . . . three outer petals lanceolate- 
linear spreading; the two inner . . . [of] nearly the same figure 
and colour. Lip nearly white, without spots, . . . the point 


ovate”) the distinctive characters of European C. innata! 
These were given (with obvious lapse or omission of a phrase) in 
his observation where he said of his new species: “It differs also 
from the European . . . principally in the oblong ovate form 
and whiteness of the inner lateral petals [characters of the 
European], also by the lip which is obtuse and spotted [the spots 
belonging to the European], and in the connivence of the two 
upper and outer petals with the inner [as shown in detailed 
figures of the ÉEuropean]". 

Only by those who see no difference between the Eurasian and 
the more boreal North American plant, with connivent sepals 
forming à hood, blunt oblong white petals and round-tipped 
spotted lip, and the temperate American plant with lanceolate 
sepals, linear-lanceolate yellow-green petals, lip abruptly tipped 
and unspotted, Nuttall's confusion of the two in his **OBsERVA- 
TION" will be applauded. By those who have carefully compared 
the two series 1t will be recognized that in the main the temperate 
North American plant with “Lip white, unspotted", is well 
separated from the Eurasian and Hudsonian North American 
C. trifida. 

In 1926, reporting on explorations in northern Newfoundland 
of a party of New England and more southern botanists who were 
all familiar with the narrow-petalled plant with unspotted lip, I 
recorded from near the Straits of Belle Isle (a Hudsonian to 
Subarctic area) a plant which differed from what we had been 
considering to be true Corallorhiza trifida. ‘The plant which was 


196 Rhodora [AUGUST 


troubling us had more purple; . . . the sepals purple or brown; 
the lip larger and with long rows of purple dots below the marginal 
notches”, ete.—Fernald in RHopona, xxviii. 93 (1926). "This 
plant was then identified with the original description and 
beautiful colored plate of the northern European C. ericetorum 
Drejer, a plant which is now generally considered an unimportant 
(small) phase of true C. trifida. Now, with the generous aid of 
Dr. Schubert in making dissections, it appears that the more 
northern plants in North America are true C. trifida, occurring 
from southern Greenland to Alaska (thence to northern Eurasia), 
south in tundra, peat, moss and peaty thickets or woods to 
northern Newfoundland, the Cóte Nord of Quebec, shores of 
Hudson Bay, northern Ontario, and along the mountains to 
Wyoming and Oregon. 

Meeting the southern limit of true Corallorhiza trifida in North 
America and sometimes overlapping it is the northernmost series 
of C. verna Nutt. Perfectly definite south ward, the latter seems 
northward to show transitions, whether through crossing or 
otherwise, and its treatment as a relatively southern variety of 
the more boreal and typical C. trifida seems justified. It thus 
becomes 

CORALLORHIZA TRIFIDA Chatelain, var. verna (Nutt.), comb. 
nov. C.verna Nutt. in Journ. Acad. Nat. Sci. Phila. iii. 134 (1823). 
C. odontorhiza (Willd.) Nutt., 8. verna (Nutt.) Wood, Class- Bk. 
ed. 2:531 (1847). C.innata R. Br., var. virescens Farr in Contrib. 
Bot. Lab. Univ. Pa. ii. 425 (1904). C. Corallorhiza, ssp. colora- 
densis Cockerell in Torreya, xvi. 231 (1916). C. trifida, var. 
virescens (Farr) Farwell in Papers Mich. Acad. Sci. xvi. pt. 1: 9 
(1941).—Details of flower well shown by A. M. Fuller, Bull. 
Pub. Mus. Milwaukee, xiv. no. 1, pl. 51 (as C. trifida) (1933).— 
Dry to moist woods, sometimes in bogs, Newfoundland to British 
Columbia, south to Nova Scotia, New England, New Jersey, 
Pennsylvania, mountains to Georgia and Tennessee, Ohio, 
northern Indiana, Wisconsin, Missouri, South Dakota, Colorado 
and Oregon. 

Details of flowers of typical Corallorhiza trifida are shown in 
many European works. A few specially accurate illustrations 
are the following: Reichenb. f., Ie. Fl. Germ. xiii-xiv. pl. 490 
(1851); H. Müller, Alpenbl. 77, fig. 21 (1881); Knuth, Handb. 
Blütenbiol. ii. pt. 2: 456 (1899); Correvon, Album Orchid. pl. 9 
(1899); Lindman, Svensk. Fanerogamfl. fig. 125, no. 6 (1918). 


1946] Fosberg,—Euphorbia maculata L. 197 


The largest-flowered species of Corallorhiza in temperate North 
America, C. striata Lindl., has a remarkably disrupted range: the 
Gaspé Peninsula; southwestern Quebec to western Ontario, south 
to northwestern New York and southern Ontario, Michigan, 
northern Wisconsin and northeastern Minnesota; southern 
Alberta and southern British Columbia, with tongues down the 
mountains to northwestern Wyoming, eastern Idaho and Cali- 
fornia; in the East preferring calcareous woodlands and growing 
chiefly at the bases of Thuja occidentalis. Throughout this 
broad range the plant (scape, sheaths and perianths) is of a warm 
madder-purple, with the sepals and 2 upper petals conspicuously 
3 (or 2)-striate with deep purple. At the easternmost limit of 
the range the stem, sheaths and perianth are yellow- or orange- 
brown, comparable with color-forms in C. maculata Raf.! and C. 
odontorhiza (Willd.) Nutt. This plant may be called 


C. STRIATA Lindl., forma fulva, forma nov., scapo vaginis 
perianthiisque fulvis. Type: arbor-vitae woods, cold walls of 
Percé Mt., Percé, Gaspé Co., Quebec, July 25, 1905, Williams, 
Collins & Fernald (Herb. Gray.). 


(To be continued) 


APPLICATION OF THE NAME EUPHORBIA 
MACULATA L. 


F. R. FOSBERG 


SVENSON, in his valuable discussion of the Descriptive Method 
of Linnaeus (RHoporA 47: 273-302, 363-388, 1945), disagrees 
with Wheeler’s interpretation of the Linnaean Euphorbia macu- 
lata. The latter (Contr. Gray Herb. n. s. 127: 76, 1939) applied 
this name, on the basis of the Linnaean specimen, to the upright 
species long known as E. nutans (or E. Preslii), regarding the 
Plukenet figure cited by Linnaeus as of secondary importance. 
Svenson, arguing that the latter figure is of equal significance 

! When, in RHopona, xxiv. 145—148 (1922), Bartlett defined (as varieties) the color- 
forms of Corallorhiza maculata Raf. in Am. Mo. Mag. ii. 119 (1817), he considered the 
yellow plants as relatively rare, while his purplish var. punicea is relatively common. 
He then concluded: ‘‘The deeply purple-stemmed var. punicea might with some 
reason be viewed as the biological type of the species, and therefore chosen, in the 
absence of a type specimen, as the nomenclatorial type as well". Bartlett and those 


who have followed him, but treating the color-forms as formae, apparently overlooked 
Raflnesque's statement that in the original C. maculata ‘‘the whole plant is vellowish''. 


198 Rhodora [AUGUST 


NN f J, & E N ) Wi 
ARRA 2 LN cy 2 inia 
ey Y 4 \ AN 
“aN 


EUPHORBIA MACULATA: FIG. 1, Plukenet’s figure (in part), Alm. t. 65, f. 8; 
FIG. 2, E. supina Raf. from Winchester, Massachusetts, Smith & Zimmerman; 
FIG. 3, Washington, D. C., Freeman. 


1946] Fosberg,—Euphorbia maculata L. 199 


Fic. 4, EUPHORBIA MACULATA: the speci- 
men in Herb. Linnaeus. 


with the specimen in 
illustrating Linnaeus' 
concept, used the case 
of E. maculata as an 
example, attempting 
to show that if the 
Plukenet figure is re- 
garded as the type, 
long-established us- 
age will be preserved 
and the name will con- 
tinue to be applied to 
the prostrate plant re- 
ferred by Wheeler to 
Euphorbia supina Raf. 

To test Svenson's 
conclusions I examin- 
ed the specimens of 
the two species in the 
herbarium of the U.S. 
National Arboretum, 
many of them anno- 
tated by Wheeler, and 
compared them with 
Svenson's reproduc- 
tion of Plukenet's fig- 
ure (RHODORA 47: pl. 
990, 1945). The re- 
sults were as follows: 

(1. The upright 
plant (our FIG. 3) fre- 
quently has the re- 
duced axillary shoots, 
referred to by Sven- 
son, as well developed 


200 Rhodora [AUGUST 


as in Plukenet's figure (our FIG. 1). (2.) The leaves of the upright 
plant practically always have petioles as long as or longer than 
those of E. supina (our ria. 2). (3.) Most important of all, the 
length of the leaves in the Plukenet figure is over half that of the 
internodes, corresponding to the upright plant (ric. 3) rather 
than the prostrate one (FIG. 2), which has them less than half, 
This gives a valuable clue to the scale of the drawing. 

'The Plukenet figure looks as though it represented an upright 
plant rather than a prostrate one, though this is inconclusive. 
Furthermore, in Linnaeus’ treatment nothing is said about a 
prostrate habit, though this character is striking enough so that 
it probably would have been mentioned. 

Loreover, the statement in the Mantissa (2: 392, 1771) 
“Euphorbia maculata similis E. hypericifoliae" lends weight to 
Wheeler’s conclusion, as E. hypericifoliae L. is superficially almost 
indistinguishable from Æ. maculata (sensu. Wheeler), while it 
bears no resemblance to the prostrate E. supina. Most impor- 
tant of all, the specimen preserved by Linnaeus and in his Her- 
barium when he prepared Species Plantarum (1753) is, as shown 
by a photograph (our ric. 4) provided by Professor Fernald, the 
upright plant which Plukenet had shown, not the prostrate one 
selected by Svenson. 

The addition of the photo. of the modern upright specimen to 
Svenson's plate removes all force from the similarity of his two 
illustrations. 

My conclusion, therefore, is that both the Plukenet plate and 
the entire Linnaean treatment apply to the upright plant, which 
should be called Euphorbia maculata L. The confusion surround- 
ing this species did not exist in Linnaeus’ concept, but was intro- 
duced by later misinterpretations. This example, consequently, 
does not seem to have much bearing on the problem of the ap- 
plication of Linnaean names to present-day concepts. 


Falls Church, Virginia. 


Volume 48, no. 571, including pages 137-164 and plates 1031-1046, was 
issued 16 July, 1946. 


SEP 17 1946 


Douora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chicf 


CHARLES ALFRED ERAI EMI 
ALBERT FREDERICK HILL 
STUART KIMBALL HARRIS 


Associate Editors 


Vol. 48. September, 1946. No. 573. 
CONTENTS: 

The Amos Eaton Herbarium. E. D. Merrill. .................. 201 

Notes on New Hampshire Plants. Alb/on R. Hodgdon. ....... 205 


Contributions from the Gray Herbarium of Harvard University— 
No. CLXII. Identifications and Reidentifications of North 


American Plants. M. L. Fernald (concluded). ............ 207 
The Genus Liatris. L. O. Gaiser (continued). ................ 216 
Edgar Burton Harger. Arthur E. Blewitt. ................2... 263 


The New England Botanical Club, Ine. 


8 and 10 West King St., Laneaster, 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 
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Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


September, 1946. No. 573. 


Vol. 48. 


THE AMOS EATON HERBARIUM 
E. D. MERRILL 


In 1937 Mr. Gustafson! published an account of what he took 
to represent an Amos Eaton herbarium. It is reported that this 
small collection, now preserved at the Rensselaer Polytechnic 
Institute, Troy, New York, contains 111 specimens labeled in 
Eaton's handwriting. That this book of specimens now at Troy, 
where Eaton taught, was thought to have been picked up in a 
southern farmhouse by a Union soldier in the Civil War strength- 
ens the idea as to the possibility or even the probability that this 
collection, which I have not seen, actually represents a student's 
herbarium corresponding to that of Mr. David L. Coe. Dr. 
Barnhart? has diseussed the Coe herbarium which was taken by 
Mr. Coe to Ohio soon after 1818, and over a century later was 
discarded by one of his descendants in Ashland, Oregon. Mr. 
Coe was one of Eaton's students at Williams College, in the class 
that actually financed the publication of the first edition of the 
latter’s Manual in 1817. 

In searching for the numerous unlisted technical names for 
plants, some deliberately, some inadvertently originated by 
Eaton in the eight editions of his Manual between 1817 and 1840, 
certain problems as to the identity of individual species arose. 
While most of the approximately forty new species actually pub- 
lished by Eaton can be placed from his descriptions, which are 
mostly short or very short, in some cases it seemed to be desirable 


! Gustafton, A. H. A Note on Amos Eaton's Herbarium. RHopona 39: 153-155. 


1937. 
? Barnhart, J. H. A  Century-Old American Herbarium. Jour. N. Y. Bot. Gard 
35: 241-245. 1934. 


202 Rhodora [SEPTEMBER 


to examine specimens that Eaton might have had before him, 
could they be located. Accordingly a serious attempt was made 
to locate Amos Eaton's herbarium in the hope that it might con- 
tain his actual types. Nothing further developed at Williams- 
town, Troy, or Albany, and then, because of Eaton's connections 
with Yale University, it occurred to me that something might be 
available there. This proved to be the case. The Amos Eaton 
herbarium, now the property of one of his great-grandsons, Mr. 
George Eaton, is deposited in the Osborn Botanical Laboratory, 
Yale University, and through the courtesy of Dr. E. W. Sinnott 
it was made available to me for examination. 

The first Eaton herbarium consists of four books nine by eleven 
inches and about two inches thick, each with about ninety sheets. 
On the back of each is the lettering “A. Eaton's Herbarium" and 
on the fly leaf of one volume is the signature of Sara C. Eaton, 
followed by the statement “from Rens. Institute, Troy, N. Y." 
This herbarium originally contained 1237 small specimens, many 
of them mere scraps, but a few of the specimens have been de- 
stroyed or lost. In the herbarium are representatives of five 
species that Eaton described as new, either on his own account 
or for other botanists, these being Xylosteum solonis Eaton, Sar- 
racenia heterophylla Eaton, Polygonum natans Eaton (P. fluitans 
Eaton), Lonicera hirsuta Eaton and Hydnum chrysorhizum Torr. 
ex Eaton. ‘These are the only “types” that were located repre- 
senting the forty species that he described as new either on his 
own account or for other botanists. "Therefore the fact that the 
Eaton herbarium is finally located is merely of slight historical 
importance, largely because he apparently did not consider it 
necessary or even desirable to retain specimens representing 
genera or species that were described as new in the various edi- 
tions of his Manual. Curiously there appear in the herbarium 
two specimens from Rafinesque which from the accompanying 
holographic descriptions of Rafinesque were collected at Fishkill, 
New York, both of which were considered by Rafinesque to be 
new, but which Eaton did not accept. Neither of the species 
was “new” at the time, but examination of these specimens 
enabled me to place two Rafinesque binomials that had appeared 
as nomina nuda in various papers published by that erratic 
botanist. 


1946] Merrill,—-The Amos Eaton Herbarium 203 

In Eaton's Manual, ed. 2, p. 5, 1818, he states that he then 
had representatives of about 1600 species in his herbarium. It 
seems to be clear that the specimens were at that time unmounted, 
and that the seleeted material was not mounted until after the 
third edition of the Manual appeared in 1822. While a few of 
the species that are described in the first or second edition of the 
Manual appear in the collection, such as Xylosteum solonis Eaton, 
Lonicera hirsuta Eaton, and Hydnum chrysorhizum Torr. ex 
Eaton, it is also to be noted that others that were not described 
until edition three appeared in 1822 also occur, namely, Sarra- 
cenia heterophylla Eaton and Polygonum natans Eaton. 

The material that Eaton elected to preserve consists for the 
most part of mere snips, mostly several to many species mounted 
on a sheet, the arrangement being in accordance with the Lin- 
naean classes. Apparently when the selected specimens were 
arranged for mounting, the names, in greatly abbreviated form, 
were penciled on the sheets, for in some cases the specimens in 
part cover these entries; later Eaton wrote the name in ink near 
each specimen. There are in general no notes, few localities are 
indicated, and no dates of collection other than an occasional 
indication of the month. The few localities that are given appear 
chiefly as N. H. = New Haven, A = Albany, W = Williamstown, 
etc. In most cases the species appear to be correctly named in 
accordance with the nomenclature of the time. There are usually 
three or four, sometimes six or more species mounted on the same 
small sheet, or in the case of the cellular eryptogams as many as 
twenty or more species on a single sheet. On the whole, ap- 
parently, this Eaton herbarium is exactly the type that he re- 
quired his students to prepare, if we may judge from the extant 
herbarium of David L. Coe, mentioned above; except that in the 
Coe collection the specimens are very much better than are those 
in Eaton's own herbarium. 

There is a second Amos Eaton herbarium at Yale University 
of which only one of the two books still exists. This covers classes 
I to X of the Linnaean system. The specimens are somewhat 
better than are those in the first and larger herbarium. There 
are 225 species in book one of this herbarium, many of them 
representing the same species as those in the first collection. In 
a few cases dates of collection are given, the latest being 1834. 


204 Rhodora [SEPTEMBER 


All the specimens are labeled, the label taking the form of a slip 
of paper pasted across the lower end of the specimen, bearing the 
name, sometimes a few other data, and always figures referring 
to the generic names used in the last (1840) edition of the Manual. 
The handwriting on the labels is not that of Amos Eaton but may 
represent that of John Wright, the co-author of this edition. In 
the front of the book, however, is a written list of the contents 
of the two volumes, and this is in Eaton's handwriting. From 
this index it is evident that there were in book two (which ap- 
parently no longer exists) about 278 additional specimens in 
classes XI to XX. No Eaton types were located in this second 
herbarium. 

It was at first thought that the Amos Eaton herbarium dis- 
cussed by Gustafson, mentioned above, might represent the miss- 
ing part of this second Eaton herbarium now at Yale University. 
This, however, cannot possibly be the case, judging from the 
data given by Mr. Gustafson. He inferred that perhaps Eaton 
has assembled large collections of botanical material in his 
travels, but I find no evidence in support of this idea. "That 
Eaton did receive rather extensive collections from his numerous 
correspondents is evident, and he certainly had the opportunity, 
as did John Torrey and the somewhat later Asa Gray, of building 
up an important historical collection of botanical specimens, an 
opportunity that he did not grasp. Whether the material that 
he did receive was discarded by Eaton himself as valueless and 
not worthy of preservation, or whether it was discarded by some- 
one else after Eaton's death, there seems to be no way of de- 
termining a century after the event. 

It had been hoped that there might be located in the Eaton 
herbarium not only specimens of the twenty new species that 
Eaton himself named and described, but that also there might 
be present specimens representing approximately the same num- 
ber of species named and described by Torrey, Hall, Aikin, Beck, 
'Tracy, and Le Conte, but actually published by Eaton for these 
authors. Only one of the latter was located, and of the former 
the holotypes of only four of the twenty species involved. 

Eaton never claimed to be a professional botanist. He was 
interested in botanical instruction and in popularizing the study 
of botany. His attitude was apparently always that of the 


1946] Hodgdon,—Notes on New Hampshire Plants 205 


amateur rather than that of the professional botanist. Becuase 
of the very simplicity of his text, and because of the obscurity of 
publication of approximately 200 new names proposed by him, 
a remarkably high percentage of these new names have remained 
for a century or longer unlisted in our standard indices, and hence, 
for the most part entirely overlooked by modern botanists. 
Only about fifty of the two hundred new Eaton names have 
previously been listed, and in these entries there are approxi- 
mately thirty-five corrections to be made, as Eaton actually 
published the names earlier than the current entries indicate. 
A special paper on these overlooked names will appear in a future 
issue of Bartonia. 


ARNOLD ARBORETUM 


Notes oN New HAMPSHIRE PrANTS.— In their recent report, 
completing their consideration of the distribution of the Gra- 
mineae of New England, Bean et al. specifically state that 
Phragmites communis Trin. var. Berlandieri (Fournier) Fern. has 
not been reported from New Hampshire. In the past two years 
I have twice collected this grass in Strafford County. However, 
not sufficient time has elapsed for these collections to have be- 
come available in herbaria. One of the new stations is an inland 
swamp in the township of Lee. There it grows amongst Alders 
and scattered specimens of Rhus Vernix L. and near a colony of 
Rhamnus alnifolia L'Hér. The second locality is in Durham, at 
the edge of a salt-marsh near the upper tidal limits of the Oyster 
River. The plants were observed there in August, 1945. 

Lemna valdiviana Philippi, according to herbarium records, has 
been found previously in New Hampshire only in Rindge in 
Cheshire County. Apparently it has not been reported from 
Maine. A number of years ago, in rapidly flowing water of the 
Bellamy River in the township of Madbury, I collected an inter- 
twining mass of vegetable matter which was finally traced to this 
species. The somewhat falcate fronds, each with one evident 
vein, characterize the species. 

Gerardia virginica (L.) BSP., the Downy False Foxglove, gets 
into New Hampshire in the Connecticut Valley in Hinsdale and 


1 Preliminary Lists of New England Plants, X XIV. RHoporA XLVIII, 17-27 (1946 


206 Rhodora [SEPTEMBER 


Alstead and far to the east in Durham!. I have collected the 
species in three widely separated areas in Durham-township. 
Two of the stations are in oak woods immediately adjacent to 
brackish water, the third is near Packers’ Falls in an oak woods 
on a dry slope. 

In the autumn of 1945, I took a small group of students to the 
Packers’ Falls area, particularly to point out an interesting type 
of oak-hickory forest. The trip was rewarded by our finding 
some vigorous specimens of Lechea villosa Ell. ‘This represents 
an extension of range northward and eastward in the state from 
Pelham and Milford. A further canvas of that part of Maine 
bordering on the Piscataqua River might yield this handsome 
member of the genus, although up to this time no reports from 
that state have been forthcoming. 

Similarly absent from Maine and not known heretofore from 
eastern New Hampshire, according to records, is that very unique 
hickory, Carya cordiformis (Wang.) K. Koch. In late August of 
1945 I found it as scattered specimens on warm slopes in perfectly 
natural situations adjacent to Great Bay in Durham, near the 
Newmarket line. There it was associated with Carya ovata 
(Mill.) K. Koch and at least one other member of the group, 
presumably Carya ovalis (Wang.) Sargent. It is interesting that, 
according to Hyland and Steinmetz in their recent careful and 
authoritative “Woody Plants of Maine", only one Hickory is 
certainly reported there as native—Carya ovata (Mill.) K. Koch. 
Again, perhaps in much botanized York County, Maine, in 
proper situations other additions to the Maine flora await 
discovery. 

Specimens of the above species from the stations mentioned are 
distributed among the following herbaria: the Gray Herbarium, 
the Herbarium of the New England Botanical Club, and that of 
the University of New Hampshire.—ALpBion R. Hopapon, Uni- 
versity of New Hampshire, Durham. 


1F, W., Pennell, “The Scrophulariaceae of Eastern Temperate North America’’, 
Academy of Natural Sciences of Philadelphia, Monograph No. 1, 1935, Map 94 
(above), shows this species as present in New Hampshire only in the Connecticut 
Valley and entirely absent from Maine. In his discussion, loc. cit. p. 385, Pennell 
comments on its host-relationship as follows: ‘‘It has been shown to be parasitic on 
the roots of Quercus prinus L. and doubtless of other species of white oaks”. Certainly 
in Durham where Quercus Prinus L. is not associated with the Gerardia and, in fact, 
has never been collected, the Downy False Foxglove is parasitic on the prevalent 
Quercus alba L. 


1946] Fernald,—Identifications of North American Plants 207 


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


IDENTIFICATIONS AND REIDENTIFICATIONS OF 
NORTH AMERICAN PLANTS 


M. L. FERNALD 
(Continued from page 197) 


Corypauis Halei (Small), Fernald & Schubert, comb. nov. 
Capnoides Halei Small in Bull. Torr. Bot. Cl. xxv. 137 (1898). 
Corydalis aurea, var. australis Chapm. Fl. So. U. S. ed. 2: 604 
(1883). Corydalis micrantha, var. diffusa Fedde, Repert. Nov. 
Spec. x. 380 (1912). 

Corydalis Halei seems to be a very distinct species of the 
southeastern Atlantic Coastal Plain and the Gulf Coastal Plain, 
extending inland, in sandy soil, to southeastern Missouri. Chap- 
man well described it as C. aurea Willd., var. australis but the 
crested keel of the outer petals, as well as its tendency to produce 
cleistogamous inflorescences, place the species nearer C. micrantha 
(Engelm.) Gray. In fact, Dr. Gray united it with C. micrantha 
in the Synoptical Flora, i!. 98 (1895). | C. micrantha, however, is 
an inland plant, found from Illinois to Nebraska, south to 
‘Tennessee, Arkansas, Oklahoma and Texas, with abundant and 
well developed foliage on the loosely ascending branches, the 
leafy summits mostly overtopping the longer terminal racemes. 
The latter are rather closely 3-12-flowered, and when fully ex- 
panded (in fruit) only 1.5-6 cm. long, with the lower and longer 
internodes 2-10 (-12) mm. long; its erect or strongly ascending 
fruiting pedicels are 0.5-3 mm. long, the thick-cylindric capsule 
scarcely torulose and 6-15 mm. long. C. Halei, on the other 
hand, is stiffly ascending, with leaves rapidly reduced upward, so 
that its terminal racemes overtop the foliage. The racemes 
quickly elongate to 0.5-2 dm., the 4—20 flowers and capsules all 
becoming remote, with the lower internodes of the rachis 1.5-2.5 
cm. long. Its capsules, on pedicels 2-5 mm. long, are slender, 
torulose and 1.5-2.5 cm. long, often outwardly arching. An 
isotype of C. micrantha, var. pachysiliquosa Fedde, 1. ¢., is a 
close match for an isotype of typical C. micrantha. 


208 Rhodora [SEPTEMBER 


ARABIS PERSTELLATA E. L. Braun, var. Shortii, nom. nov. 
Sisymbrium dentatum Torr. in Short, 3rd Suppl. Cat. Pl. Ken- 
tucky, 338 (1833), not Allioni (1785). A. dentata (Torr.) Torr. 
& Gray, Fl. N. Am. i. 80 (1838), not de Clairville (1811). Shortia 
dentata Raf. Aut. Bot. 17 (1840), at least as to ‘‘Sisymbr. do Tor. 
et Arabis"' Jodanthus dentatus (Torr.) Greene, Pittonia, iii. 
254 (1897). 

A. PERSTELLATA, var. phalacrocarpa (M. Hopkins), comb. nov. 
A. dentata, var. phalacrocarpa M. Hopkins in RHODORA, xxxix. 
169 (1937). 


Several acute botanists have called my attention to the fact 
that the name Arabis dentata (Torr.) Torr. & Gray (1838) is : 
later homonym, invalidated by de Clairville's A. dentata of 1811. 
I, naturally, passed the matter on to Dr. Milton Hopkins, before 
he became discouraged with the financial outlook for the uni- 
versity botanist and took to another and, we hope, more remun- 
erative profession. Dr. Hopkins agreed with me that the very 
distinct and localized A. perstellata E. L. Braun in RHODORA, 
xlii. 47 (1940), while differing from typical and wide-ranging A. 
dentata, must be considered an extreme variety of the same 
serles, many of the characters showing transitions in a large 
series of specimens. While it is anomalous that the plant of 
broad range should be treated as a geographic variety of a very 
local one, there seems to be no other specific name available. 
The usual source of many names which might be pieked up, 
Rafinesque, seems to have been satisfied to retain the trivial name 
for his Shortia dentata. It would now be helpful if he had em- 
ployed another. 


ASCLEPIAS VIRIDIFLORA Raf., var. linearis (Gray), comb. nov. 
Acerates viridiflora (Raf.) Eaton, var. linearis Gray, Syn. Fl. ii!. 
99 (1878). 


1 JOSEPH PHILIPPE DE CLAIRVILLE (1742-1830) was one of the most modest and self- 
effacing taxonomists I have ever encountered. His Manuel d' Herborisation en Suisse 
et en Valais (1811) was “Par l'Auteur de l'Entomologie helvétique”, without indica- 
tion of his name; while the 26-page Preface was unsigned— merely ‘‘21. Fevrier 1811”. 
How unlike the modern counterparts of the visionary Bombastus Paracelsus who 
imagine that they are helping by publishing such combinations as Phyton euphyton 
Bombastus Superbus, ssp. euphyton (Bombastus Superbus) Bombastus Superbus, 
comb. nov., var. euphyton (Bombastus Superbus) Bombastus Superbus, comb. nov.,— 
based on Phyton euphyton Bombastus Superbus in Journ, Erudit. clv. 00.000 (194?). 
So far as I can find no genus was ever dedicated to Bombastus Paracelsus. CLAIR- 
VILLEA Was defined by DeCandolle with the dignified dedication: ‘‘Dixi in memoriam 
cl. de Clairville Galli botanici et entomologici de historiá naturali Helvetica bend 
meriti", DC. Prodr. v. 636 (1836). 


1946] Fernald,—Identifications of North American Plants 209 


Since the characters supposed to separate Acerates from 
Asclepias so definitely break down the preceding combination is 
needed. Whereas typical Asclepias viridiflora and its var. 
lanceolata (Ives) Torr. extend eastward to the Atlantic States, var. 
linearis is an inland extreme, from western Ontario and Mani- 
toba south to Louisiana, Oklahoma and New Mexico. 


TRIODANIS VERSUS SPECULARIA (PLATES 1049 and 1050).—Un- 
der the title The Genus Triodanis Rafinesque, and its Relation- 
ships to Specularia and Campanula Dr. Rogers MeVaugh! has 
discussed in detail what he considers the strongest characters 
separating Specularia Heist. and Triodanis Raf. from Heterocodon 
Nutt. and Campanula L. Since the three species already recog- 
nized in the Manual range" as belonging in Specularia (and some 
new ones which occur in Missouri) are all treated as belonging to 
Triodanis, I have found it necessary to study with some care the 
characters relied upon as separating the American and Eurasian 
Triodanis from the chiefly Eurasian and North African Specu- 
laria. MeVaugh's statement of the strongest characters (as 
contrasted with those of the other two genera, which need not 
here be considered) are as follows: 


Specularia Heist. ex Fabr. Plants annual, branching above the 
middle; the flowers nearly sessile, clustered near the tips of the branches 
or corymbosely aggregated at the summit of the plant; flowers all open, 
or some imperfectly developed but open and not vestigial; corollas 
divided well below the middle, more or less rotate [this in contrast with 
Heterocodon]; filaments gradually widened to base, glabrous; capsules 
much elongate, linear, contracted at apex beneath the calyx-lobes, 
dehiscent at apex. Two species, western and southern Europe. 

Triodanis Raf. Plants annual, the branches, if any, from base or 
middle of the plant; flowers axillary, sessile or essentially so, the in- 
florescence spiciform; flowers from the lower nodes normally cleisto- 
gamous, with the corolla and androecium vestigial; some of the upper 
flowers or at least the terminal one usually open (all corollas sometimes 
open in T. coloradoensis), with expanded corollas divided below the 
middle [this, likewise, in contrast with the American Heterocodon]; 
filaments abruptly dilated and ciliate at base; capsule ovoid or clavate 
to linear or subulate, usually not abruptly contracted at apex, opening 
at the apex or (in T. perfoliata) at the middle or a little above it. Eight 
species, one chiefly Mediterranean, the others American. 


If these are, indeed, distinet genera, separated on constant 
morphological characters, in the sense of genera of Eichler, 


! McVaugh in Wrightia, i. 13—53 (1945) 


210 Rhodora [SEPTEMBER 


DeCandolle, Bentham & Hooker and Engler & Prantl, we should 
hardly expect so many qualifying expressions or exceptions in the 
statement of differential generic characters, especially when, in 
the detailed accounts of species so many additional exceptions 
to the generic definition are stated. "Taking up the reputed 
GENERIC differences in order, we, without monographie knowledge 
of details, note the following items. 

1. “Plants annual, branching above the middle" in Specularia, 
as opposed to "branches, if any, from base or middle of the plant" 
in the reputed genus Triodanis. If Rafinesque had made such a 
distinetion we should not be surprised; but, noting in passing, 
that small plants of European, southwestern Asiatic and North 
African! (not merely of “western and southern Europe") Specu- 
laria Speculum-Veneris (L.) Tanfan? (S. speculum A. DC.), 
the type of the genus Specularia, may be quite simple, we come 
to the very evident fact that of 59 individuals in the Gray Her- 
barium, these mostly validated by MeVaugh, 29 branch from the 
very base (PLATE 1049, FIG. 1) or just above it. Similarly, the 
second species allowed by MeVaugh to stay in Specularia, which 
he distinguished generically by “branching above the middle", 
the southern European (also North African—‘‘a Barbariá 
(Desf.)", A. DC. 1. e. 349) S. hybrida (L.) A. DC., shows in the 
Gray Herbarium 40 individuals with strictly or primarily basal 
branching (PLATE 1049, ria. 3) and 25 quite simple, but only 3 
branching definitely near (not * above") the middle. When, 
furthermore, we note that in the first species taken up by Mc 
Vaugh (l. c. 25) as generically distinct from Old World Specularia 
(because, as to branching at least, Specularia has the “branching 
above the middle", whereas Triodanis has “the branches, if any, 
from base or middle" (not “above the middle")—when we note 
MeVaugh's detailed description of his Triodanis coloradoensis 
(Buckl. MeVaugh (our PLATE 1050, ric. 1) as “with branches 
. . . from the nodes just above the middle of the plant (usually 

i“Syrid . , . et Barbaria’’—A. DC. Mon. Camp. 347 (1830); ‘‘Palestinae (Boiss!) 
Aegypti . . . (Bal!) ... Berythum Syriae . . . Africa  borealis"—Boissier, Fl. 
Orient. iii. 959 (1875). 

? Although the combination Specularia Speculum- Veneris appears in Index Kewen- 
sis, Suppl. 1 (1902) as made by Caruel in Parlatore, Fl. Ital. viii. 139 (1888), a check 
on the cover-page of vol. viii reveals the pertinent statement: *'CAMPANULACEE. 


JASMINACEE. OLEACEE. PER ENRICO TANFANI,” this author also cited 
at the beginning of the Campaniflore, p. 15. 


1946] T°ernald,—Identifications of North American Plants 211 


not with basal branches)”, it almost seems as if we were reading 
the contradictory writings of the author of Triodanis. The 
branching, as most taxonomists would suspect in annual weedy 
plants, is not a constant or even nearly constant GENERIC char- 
acter. 

2. Flowers “nearly sessile, clustered near the tips of the 
branches or corymbosely aggregated" in Specularia; “axillary, 
sessile or essentially so, the inflorescence spiciform" in Triodanis. 
The first point, whether the flowers are sessile or nearly so is not 
a generic difference and was put in, apparently, to contrast with 
the peduncles of Heterocodon; but when one views the inflores- 
cence of the basic Specularia Speculum-Veneris (our PLATE 1049, 
FIG. 2) and one of a species of the supposedly different genus 
Triodanis, T. coloradoensis (PLATE 1050, ria. 2), both figures from 
specimens validated by MeVaugh, and when he notes that 
Boissier, one of the truly great taxonomists, who clearly under- 
stood the Old World species, described S. Speculum ò. racemosa 
with “Inflorescentia . . . ut in Sp. falcatá racemosa” (Boiss. |. c. 
959), he stops to take a long breath. If Specularia falcata 
(Ten.) A. DC. is not a Specularia because its flowers are not 
“corymbosely arranged" as they are said to be in real Specu- 
laria, what about S. Speculum, var. racemosa which has the 
flowers racemose? And what about Triodanis coloradoensis, in 
which the flowers may be ‘‘corymbosely aggregated"? As a 
strong GENERIC difference this does not seem quite clear. 

3. In Specularia “flowers all open, or some imperfectly de- 
veloped but open and not vestigial’; in Triodanis “flowers from 
the lower nodes normally cleistogamous, with the corolla and 
androecium vestigial”. This distinctive character of Triodanis, 
however, is at once weakened farther on in the same paragraph 
by the statement that ‘‘all corollas [are] sometimes open in T. 
coloradoensis," this fact again stated in the full description of the 
quite unconventional Texan T. coloradoensis: "flowers . . . all 

prevailingly fertile and open". Cleistogamy is a common 
trait of some species or sections of genera or in some strains 
within species. In Utricularia it sometimes occurs in some 
plants and with every transition from truly cleistogamous flowers 
through intermediates to those with large and showy expanding 
corollas. In one large and perplexing American subgenus of 


212 Rhodora [SEPTEMBER 


Panicum the later and reduced panicles of apparently cleisto- 
gamous spikelets are diagnostic, yet no sound student of the 
grasses has suggested setting up Panicum, subg. Dicanthelium as 
a true genus. In Viola certain sections exhibit abundant 
cleistogamy, certain others not; yet, even to Rafinesque, Greene, 
Rydberg and Small they were all Viola. Late in the season 
Danthonia produces within and at the bases of the old sheaths 
specifically distinctive cleistogamous flowers, but if one species 
somewhere should not do so, we should hardly treat it, on that 
character alone, as another genus. Innumerable showy-flowered 
herbs, annual, biennial or perennial, will, under certain condi- 
tions, produce insignificant cleistogamous flowers. Yet, so far as 
I have seen, most of these cleistogamous individuals (except in 
case of Utricularia) have not been sorted out as separate genera. 
If Triodanis coloradoensis would obligingly stop its misbehavior, 
the case for a genus separate from Specularia would be less weak ; 
and the case would gain a little strength if the Old World S. 
hybrida (left by MeVaugh in Specularia) would stop producing 
"flowers . . . considerably reduced in size and apparently . . . 
a step in the direction of truly cleistogamous flowers" (MeVaugh, 
p.19). A little more and the step may prove fatal. 

4. The characters, corolla divided “well below the middle" 
and “‘below the middle", were obviously put in as a contrast with 
the really different corolla of Heterocodon. This section needs no 
further discussion. 

5. In Specularia “filaments gradually widened to base, glab- 
rous"; in Triodanis "filaments abruptly dilated and ciliate at 
base". However, in three of the eight species of Triodanis Mc- 
Vaugh allows that the filaments may gradually widen to base: T. 
biflora (R. & S.) Greene conceded to have “Filaments . . . , the 
proximal half gradually or abruptly expanded; and T. texana 
MeVaugh and T. Holzingeri MeVaugh with filaments similarly 
described. "That leaves (unchecked by me) filaments “glabrous” 
as opposed to "'ciliate at base". In Campanula the filaments are 
either abruptly dilated or gradually dilated at the ciliate or 
glabrous base. 

6. In Specularia “capsule much elongate, linear, contracted 
at apex beneath the calyx-lobes, dehiscent at apex"; in Triodanis 
“ovoid or clavate to linear or subulate, usually not contracted at 


Rhodora Plate 1049 


Photo, B. G. Schubert 


SPECULARIA SPECULUM-VENERIS, both figs. X 1: FIG. 1, lower half of plant, 
showing basal branching, and Fra. 2, inflorescence, both from France. 
S. HYBRIDA: FIG. 3, base, to show basal branching, from Italy. 


Rhodora Plate 1050 


h 


Photo B. G. Schubert 


SPECULARIA COLORADOENSIS (= Triodanis coloradoensis), both figs. X 1: FIG. 1, 
median third of plant, showing branching “above the middle" and fruits “corymbi- 
form aggregated"; ric. 2, flowering inflorescence, 


1946] Fernald,—Identifications of North American Plants 213 


apex, opening at the apex or (in T. perfoliata) at the middle or a 
little above it”. The apical opening is at once dismissed as 
occurring in both Specularia and Triodanis, though another 
character was overlooked in the latter, for in his key to the 
species of Triodanis emphasis is placed on “Capsule opening 
from base toward apex” in T. falcata (Ten.) MeVaugh. Further- 
more, since Alphonse DeCandolle in his classic Monographie des 
Campanulées (1830) described (p. 348) the capsule of Specularia 
Speculum-Veneris as either ‘cylindrica, medio inflata, basi et 
apice angustata" or “non proprié cylindrica, sed subfusiformi, 
utrinqué angustata’’, while Boissier (l. c. 959) defined S. Specu- 
lum, $. libanensis A. DC. with “capsulae abbreviatae interdum 
oblongae", the definition of the genus Specularia as having the 
capsule “linear” (a measurement of flat surfaces, not solids) is 
not convincing, especially since Triodanis is allowed “linear or 
subulate" capsules. Nor is the tip of the capsule a strong 
generic character: “contracted at apex" in Specularia, “usually 
not contracted at apex" in Triodanis. For, as already quoted 
from Alphonse DeCandolle, the type-species of Specularia may 
have “capsulé . . . utrinqué angustatá", although his S. hy- 
brida (which MeVaugh leaves in Specularia) has *'Capsula 
prismatica, basi apiceque abrupté constricta". Similar contrasts 
occur in American species of so-called Triodanis: for instance 
MeVaugh's description of T. coloradoensis (embarrassing species, 
always getting in the way!) contains "Capsule . . . oblong- 
linear or clavate [as also that of T. falcata (Ten.) MeVaugh] 
usually abruptly narrowed distally"; and they are “truncate” 
in T. Holzingeri McVaugh. 

From this recapitulation of the stated characters supposed to 
separate a genus, Triodanis Raf., from Specularia it must be 
apparent that the so-called generic characters set up in support 
of such a segregation fail at altogether too many points. I find 
myself following Endlicher, Alphonse DeCandolle, Boissier, 
Bentham & Hooker and Schónland (in Engler & Prantl) in keep- 
ing Triodanis in Specularia. I cannot follow Rafinesque, Greene 
and MeVaugh in considering it a clearly distinct genus. As my 
friend Pease remarks, '"Too much present-day writing is quanti- 
tative rather than qualitative." 

Although as a genus Triodanis seems to me very weak, the 


214 Rhodora [SEPTEMBER 


species defined under it seem to me very strong. It becomes 
necessary, therefore, to transfer three of them to Specularia. 
I dislike so to do, for it would have been much more satisfactory 
if the author who clearly worked them out had himself placed 
them in Specularia. 

SrEcULARIA lamprosperma (McVaugh), comb. nov. Trio- 
danis lamprosperma MeVaugh in Wrightia, i. 42 (1945). 

S. texana (McVaugh), comb. nov. Triodanis texana Me 
Vaugh, l. c. 43 (1945). 

S. Holzingeri (MeVaugh), comb. nov. Triodanis Holzingeri 
MeVaugh, l. c. 45 (1945). 


EXPLANATION OF PLATES 1031-1050 


PLATE 1031, QUERCUS LAURIFOLIA Michx.: FIGs. 1 and 2, foliage and fruit, 
x 1, from Michaux’s original plate; ric. 3, fruit of Q. rhombica Sargent, from 
Sarg. Man. ed. 2, fig. 239; ria. 4, leaf, X 1, from Michaux's original plate of 
Q. LAURIFOLIA, var. HYBRIDA, basonym of Q. obtusa (Willd.) Ashe. 

PLATE 1032, QUERCUS LAURIFOLIA Michx.: Frias. 1 and 2, fruiting branch 
and terminal leaves from Windman’s Mill, south of Sunbeam, Southampton 
Co., Virginia, Fernald & Long, no. 11,323; ria. 3, leaves of fruiting branch, 
originally and correctly identified as Q. laurifolia, later cited as a PARATYPE of 
Q. rhombica Sargent, and later identified as Q. obtusa (Willd.) Ashe, from 
Monroe, Ouachita Parish, Louisiana, E. J. Palmer, no. 8934. 

PLATE 1033, QUERCUS LAURIFOLIA Michx.: Fic. 1, venation of lower leaf- 
surface, X 3, of Michaux’s type, from photograph by Cintract; ria. 2, foliage 
of type of Q. rhombica Sargent; FIG. 3, venation, X 3 (by transmitted light) of 
lower leaf-surface of type of Q. rhombica. 

PLATE 1034, QUERCUS LAURIFOLIA Michx.?: portions of TYPE, X 1, of Q. 
rhombica, var. obovatifolia Sargent or Q. obtusa (Willd.) Ashe, var. obovati- 
folia (Sarg.) Ashe. 

Puare 1035, QUERCUS HEMISPHAERICA Bartram ex Willd., both figs. X 1: 
riG. 1, Q. aquatica, var. from Michx. Hist. Chénes Am. Sept. t. 20, fig. 2, 
cited by Willdenow; Fic. 2, toothed leaves of young tree, from Gainesville, 
Florida, Harbison, no. 35 in Herb. Arn. Arb. as Q. laurifolia. 

PLATE 1036, QUERCUS HkMISPHAERICA Bartram ex Willd., both figs. X 1: 
FIG. 1, fruiting branch, from Mayfield, Georgia, 1916, in Herb. Arn. Arb. as 
Q. laurifolia; ria. 2, cups and acorn from vicinity of Eustis, Lake County, 
Florida, Nash, no. 1663, as Q. Phellos, in Herb. Arn. Arb. as Q. laurifolia. 

PLATE 1037, STENANTHIUM GRAMINEUM (Ker) Morong: FIG. 1, portion of 
flowering plant, X 1, from Hendersonville, Henderson County, North Carolina, 
Biltmore Herb., no. 5616*; rra. 2, portion of leaf, X 2, from no. 5616°; FIG. 3, 
portion of fruiting plant, X 1, from “Pink Beds", 3500 feet alt., Pisgah Forest, 
North Carolina, House, no. 4040. 

PLATE 1038, SreNANTHIUM GRAMINEUM (Ker) Morong: Fic. 1, portion of 
inflorescence from near Merrifield, Virginia, Allard, no. 3234; FIG. 2, portion of 
leaf, x 2, from no 3234; ria. 3, portion of fruiting inflorescence, X 1, from 
Pisgah Mountain, Buncombe County, North Carolina, Biltmore Herb., no. 
3501^; ria. 4, portion of leaf, X 2, from no. 35015; ric. 5, portion of terminal 
fruiting raceme, X 1, from Springfield, Ohio, no. 11839, collector not stated; 
FIG. 6, portion of leaf, X 2, from same plant as FIG. 5. 

PLATE 1039, STENANTHIUM GRAMINEUM, Var. ROBUSTUM (S. Watson) Fer- 
nald: rra. 1, portion of TYPE of S. robustum S. Watson, X 1, from Sligo Furnace, 


1946] Fernald,—Identifieations of North American Plants 215 


Clarion County, Pennsylvania, August, 1859, J. R. Lowrie; FIG. 2, portion of 
leaf, X 2, from TYPE; FIG. 3, portion of terminal fruiting raceme, X 1, from 
Pleasant Grove, Pennsylvania, August 22, 1889, Small; FIG. 4, portion of leaf, 
x 2, from Pleasant Grove. 

PraATE 1040, STENANTHIUM GRAMINEUM, var. ROBUSTUM (S. Watson) Fer- 
nald: FIG. 1, portion of inflorescence, X 1, from Beaver, Pennsylvania, J. F. 
Mansfield; ria. 2, portion of leaf, X 2, from the Mansfield specimen; FIG. 3, 
summit of inflorescence, X 1, from New Texas, Lancaster County, Pennsyl- 
vania, Aug. 12, 1863, Porter; ric. 4, terminal flowering raceme, X 1, from 
Armstrong County, Pennsylvania, 1868, S. W. Knipe. 

PLATE 1041, STENANTHIUM GRAMINEUM, Var. MICRANTHUM Fernald: Fra. 1, 
inflorescence, X 1, of TYPE, from Caesar's Head, South Carolina, July 28, 
1881, John Donnell Smith; FIG. 2, portion of leaf, X 2, from TYPE; FIG. 3, 
portion of fruiting plant, X 1, from “Mts. Carol.", 1845, Asa Gray. 

PLATE 1042, SrsyRINCHIUM ANGUSTIFOLIUM Mill. and S. MUCRONATUM 
Michx. Fies. 1 and 2, S. ANGUSTIFOLIUM: FIG. 1, Bermudiana graminea, 
flore minore caeruleo Dillenius, Hort. Elth. t. XLI, fig. 49, X 1, basis of S. an- 
gustifolium; FIG. 2, S. anceps Cavanilles, X 1. Fia. 3, S. mucronatum Michx.: 
figure of Sisyrinchium caeruleum parvum, gladiato caule, Virginianum of 
Plukenet, Alm. 348, t. 41, fig. I, X 1, cited by Lamarck under his S. gramineum. 

PLATE 1043, SISYRINCHIUM ANGUSTIFOLIUM Mill. and S. Mo0NTANUM Greene, 
var. CREBRUM Fernald. Fics. 1 and 2, 8. ANGUSTIFOLIUM: portions of Bick- 
nell’s illustration of S. graminoides Bicknell, slightly reduced from Bull. Torr. 
Bot. Cl. xxiii, plate 263 (1896). Fic. 8, S. MONTANUM, var. CREBRUM: reduced 
from Bicknell's plate of S. angustifolium sensu Bicknell, not Miller, in Bull. 
Torr. Bot. Cl. xxiii. pl. 265 (1896). 

PraTE 1044, SISYRINCHIUM MONTANUM Greene and var. CREBRUM Fernald, 
both X 1, Frias. 1 and 2: portions of ISOTYPE of S. MoNTANUM, from Mancos, 
Colorado, Baker, Earle & Tracy, no. 113. Fra. 3, var. CREBRUM: portions of 
TYPE. 

Prate 1045, HABENARIA CLAVELLATA (Michx.) Spreng.: Frias. 1 and 2, 
TYPE of Orchis clavellata, X 1, after Cintract (note leaf—partly covered by 
label—with tip folded back); Fria. 3, leaf, X 1, from Blowing Rock, North 
Carolina, B. L. Robinson, no. 143; ria. 4, leaf, X 1, from Dedham, Massachu- 
setts, July, 1875, E. H. Hitchings; ria. 5, leaf, X 1, from Southington, Con- 
necticut, L. Andrews, no. 325; ric. 6, leaf, X 1, from Southampton, Long 
Island, St. John, no. 2655. 

PLATE 1046, H. CLAVELLATA, var. OPHIOGLOSSOIDES Fernald: rias. 1 and 2, 
portions of TYPE, X 1; ria. 3, leaf, X 1, from Tignish, Prince Edward Island, 
Fernald, Long & St. John, no. 7228; Fria. 4, leaf, X 1, from Glenwood, New- 
foundland, Fernald & Wiegand, no. 5208; FIG. 5, leaf, X 1, from Coffin Island, 
Magdalen Islands, Fernald, Long & St. John, no. 7229; FIG. 6, leaf, X 1, from 
'Trepassey, Newfoundland, Fernald, Long & Dunbar, no. 26,534. 

PLATE 1047, CLEISTES DIVARICATA (L.) Ames, all figs. X 1: FIGs. 1 and 2, 
median leaf and flower from southeast of Petersburg, Virginia (general type- 
region), Fernald, Long & Smart, no. 5742; FIG. 3, flower from Bennett, Cape 
May Co., New Jersey, June 20, 1909, Witmer Stone, “in presence of S. S. Van 
Pelt, B. Long, O. H. Brown & Dr. J. W. Eckfeldt". 

PLATE 1048, CLEISTES DIVARICATA, var. BIFARIA Fernald, all figs. X 1: 
FIG. 1, upper half of plant, from the rYvPE-sERIES, summit of Table-rock 
Mountain, Burke Co., North Carolina, Small & Heller, no. 285; ria. 2, flower 
from Jacksonville, Onslow Co., North Carolina, Moldenke, no. 1246; rra. 3, 
flower from back of Gatlinburg, Sevier Co., Tennessee, A. N. Leeds, no. 1139; 
FIG. 4, flower from Apalachicola, Florida, B. F. Saurman; ria. 5, flower from 
Spring Hill, Alabama, Æ. W. Graves, no. 853; FIG. 6, flower from near Jackson- 
ville, Florida, A. H. Curtiss, no. 4729. 

Prate 1049. Fics. 1 and 2, SPEcULARIA SPECULUM-VENERIS (L.) Tanfani: 
FIG. 1, lower half of plant, X 1, with characteristic basal branching (not merely 


216 Rhodora [SEPTEMBER 


"branching above the middle"), from France, June, 1890, S. E. Lassinome 
(identification validated by McVaugh); FIG. 2, flowering summit, X 1, of same 
specimen. Fic, 3, SPECULARIA HYBRIDA (L.) A. DC.: portion of plant, X 1, 
to show characteristic basal branching (not merely “branching above the 
middle"), from Etruria, Fiori in Fiori & Béguinot, Fl. Ital. Exsiec., no. 1962^i* 
(identification validated by McVaugh). 

PLATE 1050. SPECULARIA COLORADOENSIS Buckl. = Triodanis coloradoen- 
sis (Buckl.) MeVaugh, from specimens validated by MeVaugh: rrG. 1, median 
third of plant, X 1, showing branching ''above the middle" (not “branches, if 
any, from base or middle") and fruits “corymbiform aggregated" (not with 
“inflorescence spiciform") from New Mexico, Chas. Wright, no. 1432; FIG. 2, 
flowering inflorescence, X 1 (cf. PLATE 1049, rra. 2) from plant raised by Asa 
Gray from Wright's seed. 


THE GENUS LIATRIS 


L. O. GaisER 


(Continued from page 183) 


Var. TYPICA, forma montana (Gray), comb. nov., spikes short, 
ca. 3 dm. long; basal leaves 2-4 dm. long, 1.5-2 em. wide. Mostly 
from the mountains of North Carolina. L. spicata var. montana 
Gray, Synop. Fl. 1°. 111 (1884), in part. VIRGINIA. BATH 
Co.: vieinity of Millboro (alt. 480 m.), Aug. 24, 1907, E. S. 
Steele (US); vicinity of Millboro (alt. 510 m.), Sept. 5, 1900, 
E. S. Steele (US). NORTH CAROLINA. Mrrcueti Co.: 
Yellow Mt., Curtis (NY); Roan Mt., 1893, H. A. Edson (US); 
Little Roan Mt., (alt. 1800’), Sept. 9, 1892, C. H. Merriam 
(G, NY, US); Little Roan, July 19, 1880, J. D. Smith (US). 
BuNcoMBE Co.: Mt. Negro, July, 1841, A. Gray & J. Carey (G, 
type of L. spicata Willd. var. montana Gray); moist ground, 
slopes of Craggy Mt., July 13, 1897, Biltmore Herb., 579e (G, 
NY, US); wet hillside, vicinity of Montreat, Sept. 9, 1913, P. C. 
Standley & H. C. Bollman 10520 (US). Haywoop Co.: Frying 
Pan Gap & Big Baldy, Pisgah Forest (alt. 5000'-5300^), July 
25, 1925, P. A. Rydberg, 9517 (NY). Jackson Co.: damp rocks, 
summit of Devil's Court, House Peak (alt. 4800^), Sept. 1, 1882, 
J. D. Smith, 212 (US). SOUTH CAROLINA. Pickens Co.: 
cliffs, Table Mt. (alt. 2600’), Aug. 1896, J. K. Small (NY). 
GEORGIA. Dape Co.: woods, Lookout Mt., July, 1893, A. 
Ruth, 658 (US). 

Var. resinosa (Nutt.), comb. nov. A more slender, glabrous 
phase (hirsute only very exceptionally) from the drier barrens 
southward to the Gulf of Mexico: stems more slender than in var. 
typica, though frequently as tall, the upper two-thirds strict: 
leaves narrowly linear, the basal 2-4 dm. long and rarely exceed- 
ing 5 mm. in width, reduced upward quite abruptly: inflorescence 
spicate, 1.5-7 dm. long, when more elongated the heads some- 
times scattered and the inflorescence loosely spicate: heads 


1946] Gaiser,—The Genus Liatris 217 


cylindrical, usually of 4—7 flowers, sometimes up to 12, 8-12 mm. 
long and ca. 5 mm. thick at time of flowering; phyllaries usually 
correspondingly shorter than in var. typica but similarly ap- 
pressed, sometimes glutinous and adherent, obtuse, with narrow 
scarious margin, frequently colored at time of flowering.—Liatris 
resinosa Nutt. Gen. ii. 131 (1818), not of DC.; Chapman, FI. So. 
U. S. 192 (1860) and ed. 3, 211 (1897). Liatris sessiliflora Bertol. 
Mise. Bot. v. t. 2 (1846). Laciniaria vittata Greene, Pittonia, 
v. 57 (1902).  Liatris spicata var. B Torr. & Gray, Fl. N. Am. ii. 
73 (1841). 

Pennsylvania to Tennessee, south to Florida and Mississippi.— 
PENNSYLVANIA. Bucks Co.: Without stated locality, 1882, 
C. D. Fretz (Q). DkrLAwaRE Co.: Aston Mills, 1904, F. W. 
Pennell (US); serpentine, Williamson, Aug. 3, 1909, F. W. Pen- 
nell, 1413 (US). Lancaster Co.: vicinity of mouth of the 
Tuequan, Aug. 7, 1890, J. K. Small (US). MARYLAND. 
BALTIMORE Co.: near Baltimore, 1873, Dr. E. Foreman (US, NY); 
Catonsville, near Baltimore, Aug. 10, 1873, Morong (NY). 
DELAWARE. NEkwcasrLE Co.: gravelly roadside bank, w. 
of Chestnut Hill, 6 mis. s. of Newark, Elkton Rd., Sept. 9, 1937, 
R. R. Tatnall, 3532 (G). VIRGINIA. Sussex Co.: dry pine- 
lands, about 4 mis. n. w. of Waverly, July 26, 1936, M. L. Fernald 
& B. Long, 6417 (G); damp woods, bordering Assamoosick 
swamp, about 2 mis. n. e. of Homeville, Aug. 24, 1938, M. L. 
Fernald & B. Long, 9171 (G). DiNwipprE Co.: open argillaceous 
low woods, just e. of MeKenney, Oct. 13, 1941, M. L. Fernald 
& B. Long, 14035 (G). NORTH CAROLINA. Co. unde- 
termined: eastern. North Carolina, 1901, F. M. Jones (NY). 
Rowan Co.: Faith Post Office, Aug. 14, 1891, J. K. Small & A. 
A. Heller, 355 (G, NY, US, Q). Burke CO.: without stated 
locality, 1835, Mr. Curtis (NY). BuNcOoMBE Co.: rocky soil, 
on Cedar Cliff Mt., Aug. 17, 1904, Biltmore Herb., 579k (US). 
HENDERSON Co.: swamps of Muddy Creek, Aug. 19, 1881, J. D. 
Smath (US); open woods, Aug. 21, 1881, J. D. Smith, 184 (US); 
summit, Devil's Peak, Apr. 1882, J. D. Smith (US). Haywoop 
Co.: Looking Glass Rock, Pisgah Forest, Aug. 9, 1909, H. D. 
House, 4412 (US). Sampson Co.: pineland, near Roseboro, 
Aug. 5, 1938, R. K. Godfrey, 5716 (G). Onstow Co.: savannah, 
12 mis. w. of Jacksonville, Aug. 6, 1938, R. K. Godfrey, 5739 
(NY), savannah 8 mis. s. w. of Jacksonville, Sept. 1, 1938, R. K. 
Godfrey, 6465 (G). PENDER Co.: Big Savannah, Sept. 4, 1924, 
B. W. Wells (NY). Brapen Co.: near Bladensboro, Sept. 3, 
1908, W. W. Eggleston, 4028 (G, NY, US). New Hanover Co.: 
wet bog, near Wilmington, Sept. 8, 1927, W. H. Munter (G). 
BnuNswICK Co.: pineland, near Wilmington, Aug. 28, 1938, R. 
K. Godfrey, 6225 (C) F COLUMBUS Co.: pineland, near Halls: 
boro, Aug. 29, 1938, R. K. Godfrey, 6253 (G). SOUTH CARO- 


218 Rhodora [SEPTEMBER 


LINA. GREENVILLE Co.: rocky hillside, Caesar’s Head, J. D. 
Smith, Aug. 1881 (No. 185) (US), Aug. 5, 1881 (G); Caesar’s 
Head, Aug. 10, 1881, J. D. Smith (US); Greenville, Aug. 2, 1881, 
J. D. Smith (US). CHeEsTeRFIELD Co.: near MacBee, Aug. 
1936, W. Rhoades (G). DanrrNGTON Co.: sandhills, dry soil, 
Hartsville, July 27, 1920, J. B. Norton (US). FronENCE Co.: 
in a dry sandy place, about 1 mi. w. of New Hope, Sept. 8, 1938, 
R. Clausen & H. Trapido, 3550 (NY). Horry Co.: without 
stated locality, Oct. 8, 1934, G. F. Tarbox (NY); savannah, near 
Conway, Sept. 1, 1940, P. O. Schallert (G). GEonaETOWN Co.: 
grass-sedge upland bog or savannah, 5 mis. s. of Georgetown, 
Sept. 9, 1939, R. K. Godfrey, 8100 (G, NY). BERKELEY Co.: 
grass-sedge upland bog or savannah, 2 mis. w. of Jamestown, 
Sept. 11, 1939, R. K. Godfrey, 8182 (G, NY). GEORGIA. 
Without stated locality: Dr. Boykin (NY). Hancock Co.: 
Sparta, July 27, 1901, Biltmore Herb., 579h (US). Dopak Co.: 
damp pine barrens, Eastman, Oct. 20, 1893, C. Mohr (US). 
SumTer Co.: moist pine barrens, in s. e. part, Sept. 10, 1900, 
R. M. Harper, 620 (G, NY, US, B). Ware Co.: grassy pine 
barrens, Waycross, Aug. 23, 1900, Biltmore Herb., 579e. (US). 
CorqQuiTT Co.: between Moultrie and Kingwood, Sept. 22, 1902, 
R. M. Harper, 1652 (G, US). THomas Co.: Thomasville, Oct. 
16, 1903, Mrs. Taylor (G). FLORIDA. Without stated local- 
ity: ex Herb. J. Torrey (NY); Leavenworth (G); Chapman (G, 
NY). Co. UNDETERMINED: middle Fla., Chapman, 338 (NY). 
Duvar Co.: swamp, pine barrens, Oct. 30, 1893, A. Fredholm, 
394 (US); Jacksonville, Aug. 1896, L. H. Lighthipe (B); moist 
pine barrens, near Jacksonville, Oct. 15, 1894, A. H. Curtiss, 5311 
(NY, US); Jacksonville, A. H. Curtiss (US). BakkR Co.: Glen 
Saint Mary, Oct. 1927, H. H. Hume (G). CornuMBrA Co.: 
flatwoods n. of Lake City, Oct. 15, 1893, P. H. Rolfs (F). LEoN 
Co.: old field, Tallahassee, Oct. 20, 1942, Henry & Beggs (F). 
GADSDEN Co.: open pinelands, w. part of county, Aug. 30, 1931, 
H. Foster 114 (F). Frankin Co.: swamps, near Apalachicola, 
Aug. 20, 1872, Biltmore Herb., 579b (NY, US). Bax Co.: open 
moist ground, Lynn Haven, Oct. 12, 1921, C. Billington (US); 
Lynn Haven, Oct. 24, 1939, R. A. Knight (F). WASHINGTON 
Co.: Chipley, Aug. 31, 1942, C. M. Senner (F). WALTON Co.: 
boggy places, near Argyle, Oct. 2, 1901, A. H. Curtiss, 6923 
(US, Q); moist soil, de Funiak Springs, Sept. 26, 1900, Biltmore 
Herb., 579j (US). Crav Co.: 5 mis. w. of Penney Farms, Sept. 
22, 1939, W. A. Murrill (F). Putnam Co.: low pineland, along 
U. S. Hwy. 17, 10 mis. n. of Palatka, Sept. 24, 1940, Ruth & 
West (F). Union Co.: wet shaded edge of flatwoods, 2 mis. n. e. 
of Lake Butler, Oct. 19, 1941, W. A. Murrill (F). ArAcHUA Co.: 
flatwoods 3 mis. s. of La Crosse, Nov. 3, 1941, W. A. Murrill (F); 
low pineland, Fairbanks, Oct. 18, 1927, G. F. Weber & E. West 


1946] Gaiser,—The Genus Liatris 219 


(F); along road to Fairbanks, Oct. 28, 1936, H. K. Wallace & 
Miss L. Arnold (F); swamp s. w. of Worthington Springs, Oct. 
1945, H. H. Hume (F). '"TAvron Co.: open flatwoods, 14 mis. 
s. of Perry, Oct. 8, 1940, W. A. Murrill (F). Diıxıe Co.: flat- 
wood, 3 mis. n. e. of Shamrock, Aug. 28, 1937, Pasture Survey 
(F); 5 mis. s. of Cross City, Oct. 8, 1940, W. A. Murrill (F). 
FLAGLER Co.: “Korona meadow”, near route 1 n. of Ormond, 
Aug. 2, 1943, Mrs. H. T. Butts (G); low ground, US no. 1, Aug. 
23, 1944, Mrs. H. T. Butts (OA); near Rwy., s. of Dupont, 
June 27, 1944, Mrs. H. T. Butts (OA); Rwy. ditch, s. of Bunnell, 
spring of 1944, O. Ames (OA); flatwoods, 10 mis. e. of co. line, 
Hwy. 28 near St. Johns Park, Oct. 10, 1940, E. West and Miss L. 
Arnold (F). VOLUSIA Co.: flatwoods s. of Pierson, Sept. 17, 
1943, E. West & Miss L. Arnold (F). ORANGE Co.: sand scrub, 
Windermere, Oct. 2, 1929, F. Vasku (F); flatwoods, Orlando, Oct. 
9, 1929, F. Vasku (F); flatwoods, Vineland, Oct. 17, 1929, F, 
Vasku (F); low grassy pineland, Ft. Christmas, Sept. 24, 1927, 
O. F. Burger & E. West (F). HknNANDO Co.: near Bayport, 
July 15, 1919, R. N. Jones (US). BnEvARD Co.: in wet soil of 
prairie-like area in flatwoods, w. of Cocoa, Sept. 2, 1937, A. S. 
Rhoads (F). MaNarEE Co.: Bradentown, July 5, 1900, S. M. 
Tracy, 7078 (T); in moist ditch, Palma Sola along roadside, Aug. 
19, 1945, L. O. Gaiser, Mrs. H. T. Butts, Miss L. Arnold (F). 
OscEoLA Co.: swamp, Oct. 12, 1903, A. Fredholm, 6086 (G). 
OKEECHOBEE Co.: Okeechobee, Sept. 10, 1903, A. Fredholm, 
5999 (G); in low pineland, 4 mis. e. of Okeechobee, Sept. 11, 1929, 
H. O'Neill (US). CoLLIER Co.: prairie s. of Deep Lake, Dec. 7, 
1925, J. K. Small & W. M. Buswell (NY). DADE Co.: in 
Everglades Key, Long Pine Key, Aug. 25, 1937, Miss E. Scull 
(F). TENNESSEE. Davipson Co.: Nashville, Dr. Gattinger 
(US). Correr Co.: Tullahoma, Biltmore Herb., Aug. 7, 1899 
(No. 673), Aug. 20, 1897 (No. 579) (US); low woodland near 
Tullahoma, Oct. 2, 1902, Biltmore Herb., 579n. (US). ALA- 
BAMA. Without stated locality: Dr. Buckley (G, NY), 1831, 
Dr. Gates (G, NY, B, isotypes of L. sessiliflora Bert.), 1878, 
G. R. Vasey (US). Co. UNDETERMINED: near the Wash, Retaei, 
common, low pine barrens, Oct. 1877, C. Mohr (N Y). MARSHALL 
Co.: swamps near Albertville, July 10, 1899, Biltmore Herb., 
579m (NY). CuLLMAN Co.: near Garden City, July 28, 1938, 
H. K. Svenson 9418 (B). Ler Co.: Auburn, Aug. 11, 1897, 
F. S. Earle & C. F. Baker, 1160 (NY). AvtTauGa Co.: wet pine 
barrens, w. of Autaugaville, Aug. 2, 1938, H. K. Svenson & R. M. 
Harper, 9505 (G); flat pine woods, 1 mi. n. e. of Autaugaville, 
Aug. 2, 1938, R M. Harper, 3689 (G, NY, US). BurLrockx Co.: 
Culvers, Aug. 6, 1896, C. Mohr (US). MowTrcoMEny Co.: 
Pentulleh Creek, July 1884, C. Mohr, 16 (NY); dry open banks, 
Pentulleh, July 10, 1884, C. Mohr (US). Darras Co.: open 


220 Rhodora [SEPTEMBER 


swamp, along Mud Creek, w. of Selma, Aug. 1, 1938, H. K. 
Svenson & R. M. Harper 9790 (G). Batpwin Co.: Gateswood, 
S. M. Tracy, Oct. 30, 1903 (No. 8567) (G, NY, US, T), Oet. 30, 
1903 (No. 8568) (ND), Oct. 31, 1903 (No. 8571) (ND). Mo- 
BILE Co.: low flat pine barrens, Mobile, Oct. 10, 1871, C. Mohr 
(US); damp pine barrens, Aug. Geol. Surv., C. Mohr (US); low 
pine barrens, Mobile, Sept., 1878, C. Mohr (US); damp pine 
barrens, Mobile, Oct. 1878, C. Mohr (US); pine woods, Mobile, 
Oct. C. Mohr (US); Mobile, C. Mohr (US); Chickahasa River, 
Mobile, Oct. 7, 1894, C. S. Mohr & J. Sargent (US); near 
swamp, w. of Spring Hill, Sept. 1918, E. W. Graves, 595 (US); 
pine barrens, Spring Hill, Aug. 1919, E. W. Graves, 731 (US). - 
MISSISSIPPI. Jackson Co.: Ocean Springs, Aug. 31, 1891, 
A. B. Seymour & F. S. Earle, 132 (G), Aug. 31, 1895, J. Skehan, 
2795 (G), Sept. 6, 1889, F. S. Earle (ND); swampy open pine 
barrens, Moss Point, Aug. 8, 1933, O. Degener, 4973 (NY). 
Harrison Co.: grassy pine barrens, near Mississippi City, 
Sept, 14, 1885, J. D. Smith, 426 (US); Biloxi, Sept. 3, 1900, F. 
E. Lloyd & S. M. Tracy, 488 (NY), Sept. 19, 1898, S. M. Tracy, 
6350 (NY (ND, type of Laciniaria vittata Greene)), Aug. 23, 
1898, S. M. Tracy, 4886 (US, NY), S. M. Tracy (US), Aug. 20, 
1898, S. M. Tracy, 6353 (ND); in low pine clearing, Pass Chris- 
tian, Sept. 1879, A. B. Langlois (ND, type of Laciniaria elongata 
Greene); Cuevas (albino), Sept. 8, 1900, S. M. Tracy & F. E. 
Lloyd, 579 (US). 

Linnaeus, Sp. Pl. ii. 819 (1753), described Serratula spicata 
briefly as a plant with linear leaves, flowers sessile along a spike 
and a single stem. He based his diagnosis on a number of 
references, including that of Gronovius to Clayton's no. 237, 
from Virginia (Gron. Virg. 92 (1739)) and a Plukenet figure 
(Pluk. Alm. 190, t. 424, f. 6 (1696)), which is indeterminable. Of 
the other citations, the brief statement in Banister's catalogue 
(Banist. Virg. 1927 (1693)), though referring to something in the 
spicata group, is insufficient to determine the plant. Morison 
(Moris. Hist. iii. 137, t. 27 (1715)) quotes Banister but describes 
L. squarrosa and is cited by Linnaeus under both L. spicata and 
L. squarrosa, but in his second edition (1763) under L. squarrosa 
only. Dillenius’ figure (Dill. Elth. 85, t. 72 (1732)) was drawn 
from a cultivated plant and shows a stout branching garden 
specimen probably similar to the plant of Gronovius. The 
leaves are distinctly ciliate at the base and the phyllaries are not 
obtuse. In the second edition, Linnaeus (Sp. Pl. ii. 1147 (1763)) 
added to his description of the leaves: ‘basi ciliatis" and likewise 


1946] Gaiser,— The Genus Liatris 221 


referred to the second edition of Gronovius (Gron. Virg. 116 
(1762)). Liatris spicata, as based on Serratula spicata, was 
described by Willdenow (Sp. Pl. iii. 1636 (1803)) with very little 
further elaboration on Linnaeus’ description, except the state- 
ment that the bracts were linear and obtuse; and he again in- 
cluded all the same references. 

There is in the Linnaean herbarium a specimen labelled 
spicata, a photograph of which Dr. Fernald kindly showed me, 
which Linnaeus had in 1753. It shows a plant mounted in two 
sections, each about 30 em. long, with basal leaves about 9 mm. 
wide and an inflorescence 22 em. long with heads almost 1 em. 
long and not too crowded. Thus it agrees well with what is 
commonly called Liatris spicata as it grows in marshy places or 
as it is seen cultivated in flower-gardens. 

Examination of a photograph of the Clayton specimen in the 
herbarium of the British Museum, also seen through the kindness 
of Dr. Fernald, shows a plant having turbinate heads with loose 
bracts, rather than cylindrical heads with appressed bracts. 
The inflorescence is of 9 heads along a short raceme or spike, 
about 2 em. thick. The phyllaries appear oblong-oval. Only 
the uppermost leaves are present and these lengthen downward 
from short bracts ca. 12 mm. long subtending the basal heads to 
leaves about 3 em. long. Fortunately there is also in the Gray 
Herbarium a small packet containing a head, labelled in Dr. 
Gray’s hand: *L. spicata L. Clayt." Examination of one floret 
of this showed the corolla-tube to be pilose within and the little 
bract subtending the head distinctly ciliate on the margin. In 
all these characters the Clayton specimen 1s quite different from 
that of Linnaeus and from what is now generally considered L. 
spicata, fitting rather what has been called L. graminifolia. The 
name L. spicata, then, is correctly based on the Linnaean speci- 
men, exclusive of the Clayton plant and omitting that part of 
the description of the leaves by Linnaeus, “basi ciliatis", which 
probably was taken from the Clayton specimen. 

Michaux (Fl. Bor. Amer. ii. 91 (1803)), citing Serratula 
spicata L. as a synonym, described Liatris macrostachya, as 
having a very long spike of sessile flowers with appressed involu- 
eral bracts. A photograph of his type seen at the Gray Her- 
barium agrees well with the Linnaean specimen. Pursh (Fl. ii. 


222 Rhodora [SEPTEMBER 


507 (1814)) took up Michaux's L. macrostachya and brought 
Willdenow’s L. spicata and Walter's Anonymos graminifolius into 
synonymy, thus confusing the identity of this long-spiked plant 
(see no. 7). Elliott (Sk. ii. 273 (1824)) chose Dillenius' figure to 
represent a variety of L. spicata with longer, narrower leaves and 
applied to it the varietal epithet macrostachya, taken from 
Michaux's species. DeCandolle (Prod. v. 130 (1836)) chose the 
same epithet for his var. @. Otherwise, L. macrostachya has been 
commonly regarded as synonymous with L. spicata, and so it 
seems from photographs of the type specimens it should be. 

Torrey & Gray (Fl. N. Am. ii. 73 (1841)) and especially Gray 
(Syn. Fl. i?. 111 (1884)) excluded Clayton's plant and interpreted 
L. spicata as the thick-spiked plant represented in Curtis's Bot. 
Mag. t. 1411 (1811), Brit. Fl. Gard. t. 49 (1823) and in the figure 
of Dillenius, Hort. Elth. t. 72, f. 83 (1732). As mentioned above, 
Dillenius’ figure shows a large, branching plant of less spicate 
form with short leaves which are markedly ciliate at the base and 
probably represents a cultivated specimen of a plant similar to 
that of Gronovius. 

Two varieties of L. spicata have been recognized that need to 
be considered here: 


1. “var. 8, heads about 5-flowered; plant smaller—L. resinosa 
Nutt., . . . not of DC." T. & G. L e. ii. 73 (1841). 

2. “var. montana Gray, low and stout . . . rocky mountain- 
tops in Virginia and N. Carolina, where it abounds.” Gray, 
Syn. Fl. P. 111 (1884). 

Referring first to the latter, there are specimens in various 
herbaria, of a wide-leaved, stout-stemmed but short-spiked 
mountain form of the Alleghanies. But from boggy places or by 
lakes in the mountains have come plants with very tall spikes 
and broad leaves, so that it has seemed faulty to include in à 
mountain variety only those individuals that had probably 
grown in more exposed and less favorable places, resulting in 
shorter spikes. Examination of collections from the North 
Carolina mountains (well represented at the National Herbarium), 
with study of their habitats, seems to confirm the water-loving 
nature of this species. Probably in dry habitats the spike often 
becomes reduced. So in Biltmore Herb., no. 579d, from the 
slopes of Cedar Cliff Mt., Buncombe Co., North Carolina, one 


1946] Gaiser,—The Genus Liatris 223 


specimen at the National and one at Queen’s University Her- 
barium were found to be the low forms, while another specimen 
at the former herbarium and one at the New York Botanical 
Garden are hardly to be recognized as mountain specimens, 
resembling rather the usual var. typica. Those of reduced 
stature, if compared with those of normal var. typica from the 
same region, might represent an ecological variant or forma, but 
hardly merit varietal ranking. They have therefore been listed 
under var. typica as f. montana. 

Nuttall, Gen. ii. 131 (1818), described a plant collected in the 
pine forests of North and South Carolina as L. resinosa, for the 
resiniferous glands that are sometimes conspicuous on the bracts 
(Type in the Banks Herbarium, a photograph of which was 
obtained by Mr. Weatherby). A glutinous condition of the buds 
of plants of var. typica from much further north is sometimes 
found, so that this characterization for a new species is not dis- 
tinctive. However, the photograph of the type specimen bear- 
ing “Georgia” on the label, shows a very slender tall plant (more 
than three times the length of the herbarium sheet) with narrow 
long leaves and rather narrower lanceolate phyllaries. Further- 
more, others have described what appear to be variants in the 
South of the tall-spiked plant of open swamps.  Bertoloni, 
Misc. Bot. v. 10, t. 2 (1846), saw something new in slender plants 
collected in Alabama by Gates (isotypes in G, NY, and B) and 
named them L. sessiliflora. Greene, Pittonia iv. 315 (1901), 
also described L. vittata from a specimen from Mississippi, S. M. 
Tracy, no. 6350, near Biloxi, Harrison Co. Sept. 19, 1898 (ND). 
After examination of many specimens we have united these three 
as var. resinosa of Liatris spicata. The slenderer form, with 
long narrow leaves and fewer-flowered heads, has a more southern 
range, occurring southward chiefly from Virginia and Tennessee 
to Georgia, Florida and Alabama, in regions that may during the 
summer season become dry barrens. In Virginia and the Caro- 
linas it is unusual to find L. spicata var. typica except near the 
mountain lakes, while the slender extreme is abundant, and is 
still more so in Georgia, Alabama and Florida, though both 
varieties occur. "There are, of course, small differences in height, 
length of leaves and density of spike, in specimens from different 
localities throughout the entire range of the species, but generally 


224 Rhodora [SEPTEMBER 


the southern extreme with narrow leaves and slender spike of 
4-7-flowered heads having corolla-tubes without any pilosity, 
makes a true variety of L. spicata. In the northern latitudes of 
its range, when a specimen has grown in a location unfavorable 
for this marsh-loving species, the size of the heads and phyllaries 
and the thickness of the spike are often reduced so that one is 
reminded of specimens from Carolina, though the same long, 
narrow basal leaves may not accompany such depauperate forms. 
In most of the states where this slenderer variety and the thick- 
stemmed var. typica both occur there are present some confusing 
intermediates, frequently with very narrow leaves and yet larger 
heads of 12 and more flowers. Of these I have listed some below. 
Sometimes collections of the same collector's number and date 
comprise both varieties; as A. H. Curtiss, no. 6923, from boggy 
places near Argyle, Walton Co., Florida, Oct. 2, 1901, which at 
the National and Queen's Herbaria favor var. resinosa and at 
Gray var. typica; or R. M. Harper, no. 1652, from moist pine 
barrens between Moultrie and Kingwood, Colquitt Co., Georgia, 
Sept. 27, 1902, which at the New York Botanical Garden favors 
var. typica and at Gray and the National Herbaria var. resinosa. 
Intermediates seem frequently to occur in Florida. Likewise, 
some southern specimens suggest a variation in their long narrow 
phyllaries and very tall spikes, as Biltmore Herb., no. 579m, from 
a swamp near Albertville, Marshall Co., Alabama, July 10, 1899 
(NY). This kind of specimen was described by Greene (Pittonia, 
v. 57 (1902)) as Laciniaria elongata, type, A. B. Langlois, low 
pine-barrens at Pass Christian, Harrison Co., Mississippi, Sept., 
1876 (ND). We have included all such under the one variety 
resinosa. 

Intermediates between L. spicata var. typica and var. resinosa. 
—VIRGINIA. Sussex Co.: moist pinelands just southeast of 
Waverly, Sept. 10, 1937, Fernald & Long, 7660 (G). NORTH 
CAROLINA. Duptin Co.: savannah, 8 mi. west of Richland, 
Aug. 6, 1938, R. K. Godfrey, 5861 (G). CARTERET Co.: pineland 
near sea-level, Sept. 1, 1938. Godfrey, 6405 (G). PENDER Co.: 
savannah near Burgaw, Aug. 7, 1938, Godfrey, 5922 (G). 
GEORGIA. Warren Co.: Warrentown, July 24, 1910, Bilt- 
more Herb. 579g (NY, US). FLORIDA. Duvar Co.: moist 
pine-barrens near Jacksonville, Nov., A. H. Curtiss 1178 (G, 


NY). WakULLA Co.: St. Marks, Sept. 3, 1895, G. V. Nash 
2542 (G, NY, US, ND). VorusrA Co.: Deland Road, near 


1946] CGaiser,—The Genus Liatris 225 


Daytona Beach, Aug. 16, 1943, Mrs. H. T. Butts (G OA). 
ALABAMA. Curman Co.: Cullman, July 24, 1900, Biltmore 
Herb. 579f (US). 

The few native specimens of this complex from Louisiana 
which have been seen resemble var. resinosa rather than var. 
typica but they are quite pubescent. Of two specimens, Aug. 11, 
1919, and Sept. 1, 1922, Bro. G. Arsène, no. 11207 and no. 12517 
respectively, from Sulphur Springs, vicinity of Covington, St. 
Tammany Parish (US), the latter is similar to var. resznosa with a 
moderately hirsute stem, while the former is more densely so. A 
very hirsute condition does not usually go with L. spicata. Deam 
(Flora of Indiana, p. 912) states: “The rachis of all my plants is 
quite glabrous. Kriebel’s no. 3958, from Greene Co., has the 
rachis closely puberulent". In the examination of many speci- 
mens pubescence has but rarely been encountered elsewhere. 
Of the three Gates specimens from Alabama, which are the 
isotypes of Bertoloni’s L. sessiliflora, the one at the Gray Her- 
barium is the only one to have hirsute foliage; it was mentioned 
by Gray (Synop. Fl. P. 111 (1884)). Also on two sheets of 
collections from the Carolinas, both hirsute as well as glabrous 
plants are found; Aug. 29, 1938, R. K. Godfrey, no. 6253 from pine- 
land near Hallsboro, Columbus Co., N. C. (G), and Sept. 11, 
1939, R. K. Godfrey, no. 8182, 2 mis. w. of Jamestown, Berkeley 
Co., S. C. (G). In the recently seen herbarium of the University 
of Florida, in which are represented collections from eighteen 
counties of that state, variations from a few scattered cilia at the 
base of the leaves to densely pilose leaves and stems were found 
in four specimens from Alachua county and one each from Dixie, 
Union, Manatee and Leon counties. Since such varying degrees 
of pubescence were witnessed in plants from one locality it seemed 
that exceptional hirsute individuals of the generally glabrous L. 
spicata must be included in var. resinosa. L. Garberi, which is 
the truly hirsute species of the series Spicatae, occurs in the 
southern half of the peninsula, beginning to appear just about 
where L. spicata leaves off. 

Greene (Pittonia, iv. 315-16 (1901)) described Laciniaria sero- 
fina from a specimen of A. B. Langlois, Nov. 8, 1885, from Cov- 
ington, St. Tammany, Louisiana (ND), as a plant with a slender 
stem covered with hairs and having ovate outer phyllaries with 


226 Rhodora [SEPTEMBER 


ciliate margins and inner oblong ones with acute tips. It is of 
course possible that a few such mutations might occasionally 
arise. However, examination of a specimen of Aug. 20, 1903, S. 
M. Tracy, no. 8533, from Mendenhall, Mississippi (G, T) which 
resembles L. pycnostachya but shows only slightly recurved bracts 
while in other respects it is almost a match for the Louisiana 
resinosa-like plants, suggests that these may represent inter- 
grades with another species, as L. pycnostachya. They seem to 
make a series of intermediates between typical southern L. 
spicata, as can be seen in the Gates specimen (G, NY, B) or that 
of Oct. 30, 1903, of S. M. Tracy, no. 8567, from Gateswood, 
Baldwin Co., Alabama (G, NY, US, T), and L. pycnostachya 
Michx. (see no. 6). The suggestion of some hybridity seems more 
plausible since in Florida some plants approaching these have 
slightly ciliolate bracts and also pilosity in the corolla-tube, 
which is lacking in L. spicata but present in members of the 
Graminifoliae series. Such is a specimen of Oct. 8, 1930, F. S. 
Blanton & H. O'Neill, no. 6820, from low pineland, at edge of 
swamp, 10 mis. n. w. of Antioch, Paseo Co., Florida (US). In 
another collection of Sept. 3, 1895, of G. V. Nash no. 2542, from 
St. Marks, Wakulla Co., Florida, the specimen at the New York 
Botanical Garden has a hairy corolla and the one at Notre Dame 
is smooth. Until more specimens have been collected that 
would confirm recognizing the pubescent plants as a distinct 
species or variety we have considered them here as exceptional 
or as intermediates; and this includes Greene’s Laciniaria serotina 
which is left under the doubtful species. 

There is also some evidence in the southeastern coastal plain, 
as from south Maryland to Georgia where representatives of the 
Graminifoliae abound, that intergrades between them and L. 
spicata may occur. The collection of Oct. 4, 1929, J. H. Pyron, 
from Princeton, Clarke Co., Georgia (P) which has a stalk 93 em. 
tall, with long slender leaves and a spike 2.5 em. in diameter, has 
heads a little turbinate and corolla-tube slightly hairy, similar to 
L. graminifolia. A similar specimen of Sept. 15, 1916, Bro. F. 
Hyacinth, no. 950, Amendale, Prince George Co., Maryland (P), 
is another that looks tall and stiff like L. spicata and yet has a 
hairy corolla-tube. A specimen of Oct. 11, 1912, F. W. Pennell, 
no. 4852, from sandy pineland, Yemassee, Hampton Co., South 


1946] Gaiser,—The Genus Liatris 227 


Carolina (NY, P), while resembling the southern L. spicata var. 
resinosa in general, shows a hairy corolla and acute bracts that 
suggest L. regimontis. 

A modified specimen, such as that described by Farwell as 
Lacinaria spicata var. foliacea (Amer. Mid. Nat. ix. 260 (1925)), 
was found by the writer when collectiong on Squirrel Is., in the 
St. Clair River, Lambton Co., Ontario, and is considered to be 
the result of a plant of var. typica, found there at the northern 
limit of its range, having had the spike broken when young, per- 
haps by grazing cattle, thus stimulating greater growth of the 
remaining basal heads, their pedicels and the subtending bracts. 

In this species also, the white-flowered form seems to have 
been noticed fairly frequently and was first recognized as Liatris 
spicata forma albiflora by Britton (Bull. Torr. Bot. Club, xvii. 
124 (1890)), but as the albino has occurred in many of the other 
species, no special recognition of it is here made. 

Hereafter is described a hybrid from the northern limit of the 
range of L. spicata var. typica. 

X Liarris Steelei, hybr. nov. (L. spicata X sphaeroidea), 
'aulibus erectis glabris ca. 8.5 dm. altis e cormo compresso 3 em. 
lato 2 em. alto; foliis radicalibus argute lanceolatis 36 em. longis 
2 cm. latis, petiolo anguste alato longitudine tertiam partem 
laminae aequante, foliis caulinis basem versus radicalibus simili- 
bus ca. 20 em. longis, superne ad folia lineari-lanceolata vel 
linearia 10-1 em. longa 3-1 mm. lata abrupte reductis; inflores- 
centia e spica laxa 20-40 cm. longa; capitulis 20-25-floris sessili- 
bus vel breviter pedicellatis 1-1.5 em. longis anthesi 1-1.2 cm. 
latis tune subturbinatis; phyllariis glabris plerumque herbaceis, 
exterioribus ovatis vel oblongis marginibus scariosis angustis, 
medianis interioribusque oblongo-linearibus apice rotundato 
conspieue scarioso coloratoque; corolla purpurea, tubo 7 mm. 
longo intus piloso; achaeniis 5 mm. longis, pappo barbellato 6 
mm. longo.—IN DIANA. Porter Co.: on and among dunes, 
Dune Park (alt. 175-200 m.), Sept. 17, 1909, E. S. Steele 169 
(US 609031, TYPE, 609030, isotype). 

These two plants were observed to resemble L. spicata var. 
(ypica but to vary in their larger heads of more flowers and 
broader phyllaries with scarious, rather crisped margins. The 
leaves too are sharply lanceolate rather than linear-lanceolate as 
found in that species. 

These variations in characters suggested relationship to mem- 


228 Rhodora [SEPTEMBER 


bers of the Scariosae series and a search was made through the 
Steele collection. A specimen of X L. sphaeroidea, collected at 
the same time and place was discovered—no. 162b (US609017). 
As other specimens of this latter hybrid have come from this 
region of Indiana it seems quite likely that they represent one 
parent of the new hybrid. As discussed (see no. 18) X L. 
sphaeroidea has come to be quite a stable unit and has probably 
been a parent of other hybrids as well. "Though no specimen of 
L. spicata from the very same location was found, in the collection 
there is, surprisingly enough, one of Sept. 20, 1909, of E. S. Steele, 
no. 181, from the old beaches, Lake Chicago, Buffington to Pine, 
Lake Co., Indiana (US 609098), with the typical, long, linear 
leaves and usual heads of about 10 flowers of L. spicata. Thus 
presumably the second parent-species grew at least in the region 
and perhaps even in the near-by vicinity at the time of collection 
of the hybrid plants. l 


2. LIATRIS LANCIFOLIA (Greene) Kittell. A thick-stemmed 
glabrous plant 6 dm. high resembling L. spicata var. typica: 
leaves numerous; basal ones 2-3 dm. long and 1-1.5 em. wide in 
the center from which they taper equally to the base and obtuse 
tip; upper leaves shorter and bluntly lanceolate: inflorescence a 
dense or loose spike 1.5-3 dm. in length and 2-3 em. in diam.; 
heads ca. 12-flowered; phyllaries erect, glabrous, the outer 
triangular-ovate, the inner oblong and acute, mostly herbaceous, 
with narrow purplish ciliolate margins; corolla purple, 6 mm. 
long, smooth within; achene (immature) at least 3 mm. long; 
pappus 5 mm. long.—Tidestr. & Kittell, Fl. Arizona and New 
Mexico, 370 (1941). Laciniaria lancifolia Greene, Bull. Torr. 
Bot. Club xxv. 118 (1898). Lacinaria kansana Britt., Man. 
927 (1901). Liatris kansana Rydb., Fl. Prairies & Plains Cen- 
tral N. Amer. 781 (1932). 

Along river-bottoms from S. Dakota to Colorado and in the 
mountains of New Mexico.—STATE UNDETERMINED. J. 
D. Cooper (NY). SOUTH DAKOTA. Unton Co.: Wallace, 
Aug. 16, 1892, S. Dak. Agr. Exp. Sta. (US). NEBRASKA. 
Co. UNDETERMINED: Aug. 9, 1899, J. H. Holms (US). Kryapau 
Co.: Carns, Aug. 18, 1893, F. Clements, 2892 (G, US). Scorrs- 
BLUFF Co.: Platte Bottom, w. of Scottsbluff, Aug. 3, 1891, P. A. 
Rydberg, 139 (NY, US). Kearney Co.: Platte Valley at Fort 
Kearney, July 1891, H. Hapeman (US). KANSAS. Rivey Co.: 
prairie, Aug. 7, 1895, J. B. Norton, 215 (G, NY, US). Forp Co.: 
bottoms, south of river, Dodge City, Aug. 19, 1890, B. B. Smyth, 
163 (NY, type of Lacinaria kansana Britt., US). WYOMING. 


1946] CGaiser,—The Genus Liatris 229 


GosHEN Co.: moist meadows, Pratt, Aug. 1912, A. Nelson, 9667 
(G). COLORADO. Co. UNDETERMINED: Platte, Fremont 
(NY). Yuma Co.: Wray, Aug. 16, 1907, H. L. Shantz (NY). 
NEW MEXICO. CuHaves Co.: Roswell, South Spring R. 
(alt. 3800’), Aug. 25, 1900, F. S. Earle & E. S. Earle, 258 (NY, 
US); Roswell, South Spring R., Aug. 25, 1900, F. S. Earle (ND). 
LINcoLN Co.: in marshy land, south fork of Tularosa Creek, 
White Mts. (alt. 6800), July 31, 1897, E. O. Wooton, 254 (G, 
NY (ND type)). Orero Co.: along Tularosa Creek, Aug. 18, 
1899, Aug. 6, 1901, E. O. Wooton (US); Tularosa Creek, 3 mis. s. 
of Mescalera agency, July 19, 1928, C. B. Wolf, 2795 (G); rather 
swampy meadows, Mescalera Indian Reserv., Aug. 2, 1931, 
W. Huber (P). 

Essentially the same basic characteristics are common to L. 
spicata var. typica found in the states east of the Mississippi and 
to L. lancifolia which was described from marshy land in the 
White Mountains of New Mexico. The width of the leaves of 
the latter, described by Greene, l. c. as *broad in proportion", is 
somewhat unusual for a member of the Spzcatae series, though 
similar to L. spicata var. typica f. montana. Lacinaria kansana 
Britt. was described from a plant from river-bottoms in Kansas 
as having also blunter leaves. Rydberg l. e. treated Liatris 
kansana as a species occurring rather infrequently from S. Dakota 
to Colorado, i. e. in the western plains states. | 

When the New Mexico specimens are compared with those of 
the neighboring plains and allowance is made for some modifica- 
tion due to mountain habitats, the forms seem hardly distin- 
guishable. In both, the corolla-tube is smooth within so that 
one might consider these geographically separated segregates as 
varieties of L. spicata but for their separation by the expanse of 
the central plains states from the rest of that species. We are 
here considering the two as one species, L. lancifolia (Greene) 
Kittell. 

The species is readily separated from L. pycnostachya Michx., 
a common species in the western plains, by the appressed rather 
than recurved phyllaries. Occasional material is suggestive of 
a condition intermediate between the two, as the collection of 
1861, 7. J. Hale, Trempealeau, Trempealeau Co., Wisc. (G), 
with acute and ciliate-margined though erect phyllaries or those 

! In the Report of Fremont's Expedition, Washington, 1845, the reference on page 


90 of the Catalogue of Plants to “L. spicata (Willd.) north fork of the Platte Sept. 4” 
is probably to this specimen. 


230 Rhodora [SEPTEMBER 


of Aug., 1892, ex Herb. J. J. Thornber, Brookings, Brookings Co., 
S. Dak. (G), and of Sept. 3, 1891, ex Herb. J. M. Bates, Valentine, 
Cherry Co., Neb. (G), both having phyllaries that, though not 
recurved, are loose rather than appressed and have petaloid 
margins. As there was some pilosity within the corolla-tubes, 
confirmation of intergradation is given and may even suggest 
that it be between L. lancifolia and some species of the plains 
other than L. pycnostachya since that usually shows none or very 
few hairs in the tube. 


3. LIATRIS MICROCEPHALA (Small) K. Sch. Stems commonly 
tufted, slender, striate, glabrous, 3-7 dm. tall, from a small ovoid 
or subglobose corm, usually not more than 3 em. in diameter; 
leaves numerous, linear, glabrous with very few hairs along mid- 
vein or margin, the basal 5-10 cm. long, 2-3 mm. wide, the upper 
gradually reduced to the base of the inflorescence: heads in lax 
slender racemes, quite numerous, 4-6-flowered, cylindrical, 
somewhat narrowed, but not acutely, at the tips, ca. 8 mm. long, 
slender, on short, slender, erect peduncles about equalling them 
in length; phyllaries glabrous, appressed, obtuse, the outer ovate, 
the inner oblong, with only minutely ciliolate or merely slightly 
scarious margins, 5-6 mm. long: corolla-tube 5-6 mm. long, 
purple, lacking any pilosity within; achenes 3-4 mm. long; 
pappus short, 3.5-4 mm. long, about half as long as the corolla- 
tube, not plumose to the naked eye (i. e. with barbellate bristles). 
—Just, Bot. Jahresb. xxvi. pt. 1, 378 (1900). Lacinaria micro- 
cephala Small, Bull. Torr. Bot. Club xxv. 473 (1898); Man. S. E. 
Fl. 1334 (1933). Laciniaria polyphylla Small, Fl. Southeastern 
U. S. 1173, 1338 (1903). 

On sandstone outerops and dry barrens in North Carolina, 
Tennessee, Kentucky, Georgia and Alabama.—NORTH CARO- 
LINA. Pork Co.: Tryon, near Columbus, Aug. 4, 1897, E. C. 
Townsend (US); Tryon, Aug. 11, 1921, D. C. Peaítie, 1154 (NC). 
Henverson Co.: Flat Rock, Aug. 24, 1881, J. Donnell Smith 
(NY). GEORGIA. Co. UNDETERMINED: in shade of moun- 
tains, 1836, Drummond, 173 (NY). Rasun Co.: canyon, 
Tallulah Falls (alt. 1600’), Aug. 16, 1893, J. K. Small (G, 
NY, US). pe Kats Co.: on slopes and summit of Stone Mt. 
(alt. 1000'-1686^), Sept. 6-12, 1894, J. K. Small (G (NY, type 
of Laciniaria polyphylla Small), US); Stone Mt., Biltmore 
Herb. Sept. 8, 1897 (No. 4117a), Sept. 1, 1899 (No. 149362) 
(NY); Stone Mt.; L. R. Gibbes (NY), Sept. 1876, W. M. Canby 
(G); turf, rocks, n. w. base of Stone Mt., Aug. 17, 1927, K. M. 
Wiegand & W. E. Manning, 3175 (G); rocky soil, Aug. 2, 1912, 
F. W. Pennell, 4064 (US, P). HockpaALE Co.: 6 mis. s. w. of 
Logansville, Big Haynes Cr., Oct. 18, 1936, J. H. Pyron & R. 


1946] Gaiser,— The Genus Liatris 231 


McVaugh, 1126 (US). KENTUCKY. Co. UNDETERMINED: 
Mts. of Kentucky, Lexington, 1837, C. W. Short (NY). Mc 
CREARY Co.: on sandstone and sand, along South Fork, Cumber- 
land R., Sept. 16, 1934, E. Lucy Braun (G); along sandy shores of 
Upper Cumberland R., Sept. 13, 1940, F. T. McFarland & H. J. 
Rogers, S8 (G). TENNESSEE. Co. UNDETERMINED: On rock, 
e. Tenn. MacFarlands, A. Ruth, 1896 (No. 3762), 1897 (No. 
5804) (NY). DavipsoN Co.: Nashville, 1878, Dr. A. Gattin- 
ger (P). Moraan Co.: sandy margin of stream, Rugby, Aug. 
19, 1930, H. K. Svenson, 4094 (G, P, D); moist sandy glade, n. 
of Hoffman, Sept. 11, 1927, E. T. Wherry & F. W. Pennell, 
13942 (P). Cockr Co.: 3 mis. s. of Wolf Creek Sta., Aug. 30, 
1897, T. H. Kearney, 746 (US). CuMBERLAND Co.: gravelly 
oak woods, 6 mis. e. of Crossville, Aug. 15, 1930, H. K. Svenson, 
4173 (G, B); Ozone, July 14, 1929, W. A. Anderson, 1402 (G), 
H. M. Jennison & W. A. Anderson (G, O without No.); sandy 
soil, mountain-top, Aug. 24, 1890, D. C. Coffman (US). VAN 
Buren Co.: dry sandy ledge, bluff overlooking Can Creek Falls, 
July 19, 1935, J. K. U. Sharp & A. J. Sharp, 2955 (P). COFFEE 
Co.: dry oak-barrens, n. of Manchester, H. K. Svenson, Aug. 6, 
1938 (No. 8987) (G, B), Aug. 8, 1940 (No. 10607) (B); near 
Tullahoma, Sept. 24, 1933, E. J. Alexander, T. H. Everett & S. D. 
Pearson (NY). FRANKLIN Co.: Sewanee, 1897, K. Selden, 311 
(NY, F); sandstone rocks, St. Andrews, July 15, 1938, H. K. 
Svenson, 9583 (G, B). ALABAMA. Co. UNDETERMINED: 
in tufts on rocks, on the summit of mts., July 29, 1896, C. Mohr 
(US). pe Kares Co.: falls, above Valley Head, Sept. 20, 1899, 
Biltmore Herb., 14936b (NY). MamnsHALL Co.: rocky glades, 
along Short Creek, 5 mis. n. e. of Boaz, Aug. 29, 1933, R. M. 
Harper, 3108 (G, NY, P, US). Erowan Co.: on sandstone 
rocks, above Noccalula Falls, Lookout Mt., Aug. 31, 1911, R. 
M. Harper, 149 (G, NY, US). CurLMaN Co.: Cullman, Sept. 9, 
1897, H. Eggert (NY, ND, US); rocky glade, St. Bernard, Sept. 
15, 1937, Rev. Gattman (NY, P). Carnouwv Co.: on granitic 
rocks, dry hills, near Anniston & Chandler Springs, Sept. 20, 
1892, C. Mohr (US). '"lTAnnLEbEGA Co.: Blue Mts., Sept. 24, 
1893, C. Mohr (NY). Avrauaa Co.: flat pine-woods, n. e. of 
Autaugaville, Aug. 31, 1930, R. M. Harper (G, P); dry gravelly 
banks in open exposed places, banks of Autauga Creek, near 
Prattville, July 18, 1880, C. Mohr (US). Mowmok Co.: in 
mts., near Franklin, 1843, Rugel (NY). 

Although fire in the Herbarium of the University of Tennessee 
destroyed the type from which Small described Lacinaria micro- 
cephala (Dr. A. Gattinger 1888, in the Cumberland Mts. Tennes- 
see), observation of many specimens from this region in various 


herbaria confirms its identity as a distinctive species. 


232 Rhodora [SEPTEMBER 


Liatris microcephala has been frequently confused with L. 
graminifolia, probably because of a general similarity in size of 
plant and the somewhat similar distribution of heads. However, 
the prominent cilia found along the margins of the petiole in the 
latter species are lacking in the almost glabrous leaves of L. 
microcephala. The number of flowers in a head are also much 
fewer in this species and make a slender cylindrical head, whereas 
in L. graminifolia the head is thickened in bud and becomes more 
turbinate upon opening. Upon examination of the corolla-tube 
with a lens it is found to be clear in L. microcephala, as in L. 
spicata, whereas in L. graminifolia considerable pilosity is seen. 
It is by these characters that L. microcephala has been placed 
with the Spicatae rather than with the Graminifoliae. 


4. Liarris acipora Engelm. & Gray. Corm globose or 
slightly elongated, usually not more than 3 cm. in diameter, 
bearing a tuft of fibres, the remnants of previous basal leaves, as 
seen also in L. spicata and L. pycnostachya: stems slender but 
stiffly erect, 5-8 dm. tall, glabrous or puberulent, arising singly 
in young, but to the number of three or four in older plants: 
basal leaves very long, glabrous, linear-lanceolate, 2-4 dm. long, 
3-5 mm. wide, those of the slender spikes narrowly linear becom- 
ing abruptly shortened, erect and bract-like: heads numerous, 
loosely covering 10-20 cm. of the erect spikes, 3—5-flowered, 
cylindrical but narrowed acutely at the tip when in bud, about 
1 em. long; phyllaries few, glabrous, appressed, the outer ovate, 
the inner oblong-lanceolate, sometimes becoming purplish; 
corolla about 8-10 mm. long, phlox purple, lacking any pilosity 
within the tube; achene 4-5 mm. long; pappus ca. 7 mm. long, 
not plumose to the naked eye (i. e. with barbellate bristles).— 
Pl. Lindh. i. 10 (1845); Boston Jour. Nat. Hist. v. 218 (1845); 
Gray, Pl. Wright. i. 83 (1852), in part. L. mucronata sensu 
Torr. & Gray Fl. N. Am. ii. 70 (1841) not DC.  Lacinaria 
acidota O. Ktze., Rev. Gen. i. 349 (1891); Bush, Amer. Mid. Nat. 
xii. 318 (1931).  Lacinaria Halei Small, Bull. Torr. Bot. Club, 
xxvii. 281 (1900). Liatris acidota var. vernalis Engelm. & Gray, 
Pl. Lindh. i. 10 (1845). Probably Lacinaria brachyphylla Bush, 
Amer. Mid. Nat. xii. 317 (1931). 

Coastal plain region of Texas and Louisiana.—TEXAS. 
Without stated locality: 1843, Lindhetmer, 73 (G); Apr. 1842, 
Lindheimer (P); Sept. 1842, Lindheimer (G); 1836, Drummond, 
171 (G, paratype), Drummond, 146 (G, paratype); C. Wright 
(G, P), 155 (US); Sept. 1850, G. Thurber (G, NY, P). Co. unde- 
termined: wet prairies, Houston to the Brazos, 1843, Lindheimer, 
72 (G, type) (P). Trintrry Co.: 4 mis. e. of Riverside, Sept. 


1946] Galser,— The Genus Liatris Es 


29, 1934, V. L. Cory, 10476 (G). Nrwron Co.: 16 mis. s. of 
Newton, Oct. 4, 1934, V. L. Cory, 10966 (G, US); 21 mis. n. of 
Dewbeyville, Oct. 4, 1934, V. L. Cory, 10966 (G). HanniN Co.: 
2.6 mis. e. of Camp Jackson, Sept. 13, 1936, V. L. Cory, 19799 
(G, T); 7 mis. n. of Silsbee, V. L. Cory, 20036 (G); Kountze, Aug. 
5, 1939, H. R. Reed, 30 (G). Tyurr Co.: without stated locality, 
July 22, 1939, B. C. Tharp (G). Liserty Co.: Stilson, Sept. 3, 
1923, B. C. Tharp, 2633 (US); Cleveland to Dayton, Sept. 9, 
1937, B. C. Tharp (G). Harris Co.: pine woods, Houston, 
Aug. 9, 1921, R. Ferris & C. D. Duncan, 3266 (NY, P); Houston, 
July 23, 1915, G. L. Fisher, 1720 (US); Houston (specimen to 
right), 1915, E. J. Palmer, 8577 (P): Bammel Ranch (very 
abundant), July 18, 1925, Oct. 1, 1925, H. Ness (T); prairies, 
Harrisburg, 1874, J. F. Joor (US); Harrisburg, Sept. 2, 1875, 
Oct. 1894, J. F. Joor (US); Seabrook, Aug. 11, 1913, G. L. 
Fisher, 663 (US). WarLER Co.: without stated locality, Aug. 
14, 1898, F. W. Thurow, 12 (US). AvsrIN Co.: without stated 
locality, Oct. 15, 1939, B. C. Tharp (G). JEFFERSON Co.: Vidor, 
Sept. 1, 1923, B. C. Tharp, 2641 (US). CHAMBERS Co.: coastal 
prairies, Sept. 3, 1924, B. C. Tharp, 3179, 3180 (US). GaLvEs- 
TON Co.: 13 mis. n. w. of Galveston, Aug. 4, 1939, H. R. Reed, 
28 (G). Brazoria Co.: 12 mis. s. of Alvin, Dec. 6, 1918, H. C. 
Hanson (NY, P). Maracorpa Co.: moist sandy soil, along 
Rwy. s. of Van Vleck, Sept. 20, 1913, F. W. Pennell, 5512 (NY, 
P); 6.8 mis. w. of Palacios, Oct. 18, 1936, V. L. Cory, 20279 (G). 
JACKSON Co.: Edna, Dec. 9, 1930, J. A. Drushel, 6847 (US); El 
Toro, Nov. 11, 1931, J. A. Drushel, 8837 (US). CaLHouNw Co.: 
Port Lavaca, July 2, 1939, B. C. Tharp (G). LOUISIANA. 
Without stated locality: Hale, 334 ((G, unnumbered), (NY, 
type of Lacinaria Halet Small) P). CarcasrEgU Co.: low moist 
grassy field, along Rwy., 1 mi. e. of Lake Charles, July 20, 1938, 
D. S. Correll & H. B. Correll, 9638 (G); Lake Charles, Aug. 25- 
Sept. 10, 1938, K. K. Mackenzie, 475 (NY, P); Lake Charles, 
Aug. 7, 1897, S. M. Tracy, 3448 (NY, P); Lake Charles, Sept. 
1906, R. S. Cocks, 2922 (ND). 

Liatris acidota was described by Engelmann and Gray in 1845 
(Pl. Lindh. i. 10), when they distinguished it from De Candolle’s 
earlier species L. mucronata (Prodr. v. 129 (1836)). However, 
the two species are still confused and sometimes as well with 
L. punctata. 

The type of L. mucronata is a plant collected Nov. to Dec., 
1828, by Berlandier, no. 1926, in Texas (Herb. Geneva; isotypes 
in G, NY). Torrey & Gray (Fl. N. Amer. ii. 70 (1841)) main- 
tained L. mucronata and referred to 1t other specimens collected 
by Drummond, no. 171, and no. 146, from Texas (G), and by 


234 Rhodora [SEPTEMBER 


Hale from Louisiana (G, NY). When Engelmann and Gray de- 
scribed L. acidota from a plant of Lindheimer's collections, no. 
72, from wet prairies, Houston to the Brazos (G, NY), they 
selected as similar to it, the Drummond and Hale specimens. 
A spring-blooming plant collected by Lindheimer, no. 73, near 
Houston (G), was made L. acidota var. vernalis. However, 
they recognized their species as distinet from De Candolle's and 
wrote (l. c. page 10) as follows: 

"In the Flora of North America, this species, which is ap- 
parently common in western Louisiana and Texas, was hesitat- 
ingly referred to L. mucronata DC., from the character of which 
it differs in some respects, principally in the form of the involucral 
scales. But among Lindheimer’s plants, some specimens of what 
is no doubt the true L. mucronata DC. occur (which have been 
distributed in some sets probably mixed with L. acidola) and 
which render it clear that the present is a different, although very 
nearly allied species". 

Then in Gray, Synop. Fl. i?. 110 (1884), L. acidota was retained, 
not in the sense of Engelmann & Gray, but of L. mucronata of 
Torrey & Gray. "True L. mucronata was reduced to a variety, to 
which other Texas plants were referred. Specimens collected in 
1849 by Lindheimer, no. 940, in the neighborhood of Comanche 
Spring and in 1850, no. 941 at New Braunfels, were so named in 
the 18th report of the Missouri Botanical Garden (1907). Thus 
there was continued an association of L. mucronata and L. acidota 
that is hardly justifiable when the type specimens are examined. 
De Candolle in describing L. mucronata referred only to Ber- 
landier's specimen which, by following the route travelled by 
Berlandier (Geiser, Naturalists of the Frontier, Chap. III 
(1937)), was probably collected between Boerne and Comfort, 
in what is now Kerr and Kendall counties, of the blackland 
prairie region of Texas. Engelmann and Gray stated clearly, 
when describing L. acidota, that it came from the wet prairies 
"from Houston to the Brazos” (in the case of Lindheimer’s 
plant), “western Louisiana" (in the case of Dr. Hale’s plant); 
and, as Drummond’s letters to Hooker (Hook, Journ. Bot. i. 
(1835)) indicate that Drummond spent most of his time collecting 
in a region inland from Galveston, that plant too would have 
come from the wet coastal plain region. All the specimens I 


1946] Gaiser,— The Genus Liatris 235 


have seen that conform to the type of L. acidota have come from 
the coastal plain area. 

The authors’ original descriptions further differentiate the two 
species: 


L. acidota L. mucronata 
spike: elongate, strict strict 
heads: 3-flowered 5-flowered 
bracts: oblong-lanceolate, outer ciliate, obtuse, abruptly 
ovate, gradually becoming mucronate 


acuminate, cuspidate 


Further examination of the specimens emphasize such differ- 
ences as these: 


height: 4-6 dm. 3-4 dm. 

rachis: glabrous, almost naked leafy 

basal leaves: very elongate, diminishing not noticeably longer and 
abruptly gradually diminishing 

upper leaves: erect, short, almost progressively shorter leaves 
subulate bracts 

corolla tube: smooth inside pilose within 

pappus: not plumose to the naked eye plumose to the naked eye 
(i. e. with short lateral 
cilia) 

achene: 3 mm. long 5 mm. long 


Thus it seems that by the nature of the pappus alone the two 
species fall each into a different section of the genus, L. acidota 
in Suprago, and L. mucronata in Euliatris. Not only to this 
latter section but also to the same Punctatae series, belongs L. 
angustifolia Bush, and in the discussion of that species (see no. 
27) will be found further reference to the confusion of L. acidota- 
like plants with L. mucronata and L. angustifolia. 

In 1900 Small described Lacinaria Hale?, from Dr. Hale’s no. 
334, from Louisiana (NY), stating: “This species has heretofore 
been included in Lacinaria acidota, with which it has little or 
nothing in common and it may be separated by its fewer leaves 
and much smaller heads which are disposed in elongated inter- 
rupted spikes". It would seem that by this time the erroneous 
synonymy of L. mucronata and L. acidota had led to such con- 
fusion that plants of the blacklands and prairies were referred to 
as L. acidota and Small made a second attempt to give the slender 
coastal plain plant its own rating as Lacinaria Halei. However 
the name Liatris acidota Engelm. & Gray can be taken up for it. 

Bush (Amer. Mid. Nat. xii. 318 (1931)), in attempting to 
clarify L. acidota, made a comparison of a great many sheets of 


236 Rhodora [SEPTEMBER 


so-called L. acidota and L. punctata and included in L. acidota 
only specimens from the coastal plain region of Texas and 
Western Louisiana. Among them he included one of Aug. 25- 
Sept. 10, 1898, collected by K. K. Mackenzie, no. 475, in low 
prairies at Lake Charles, Louisiana. I have seen duplicates of 
this specimen at the New York Botanical Garden and the Phila- 
delphia Academy of Sciences and, finding them similar to the 
type plant, am agreeing with his interpretation. 

L. acidota, as here interpreted, shows in common with the 
other species of the Spicatae series (except L. microcephala) the 
tall spicate form and non-pilose corolla, as well as a preference 
for moist habitats. 


5. LIATRIS GARBERI A. Gray. From clustered, elongate, 
thickened, tuberous roots that spread out in a somewhat fingeroid 
manner from the crown of the plant; stems one to several in older 
plants, hirsute and 3-5 dm. high: basal leaves linear, 1-2 dm. 
long, ca. 5 mm. wide, almost glabrous but with cilia along the 
margin of the lower half that narrows into a clasping petiole; 
upper ones reduced from the base of the spike as they are in L. 
spicata: heads 6-7-flowered, 8-15 mm. long, cylindrical and some- 
what narrowed at the tip; phyllaries appressed, viscid-hirsute, 
ovate to lanceolate and acuminate- to mucronate-tipped, though 
not sharply, mostly entire but thin on the margins, sometimes 
ciliate; corolla tube 7-9 mm. long, smooth inside (i. e. lacking 
any pilosity), and phlox-purple; achene ca. 3 mm. long; pappus 
6-7 mm. long, barbellate.—Proc. Amer. Acad. xv. 48 (1880). 
Laciniaria Garberi O. Ktze. Rev. Gen. i. 349 (1891); Small, Man. 
S. E. Fl. 1333 (1933). Laciniaria Nashii Small, Fl. S. E. U.S. 
1175, 1338 (1903). Laciniaria chlorolepis Small, Man. S. E. Fl. 
1333 (1933). 

Low pinelands and damp flatwoods in the southern half of 
peninsular Florida.—FLORIDA. Without stated locality: 
1842-49, F. Rugel (US). Co. undetermined: s. Florida, A. P. 
Garber (G, type). OnaxaE Co.: St. John's River flats, Aug. 3, 
1935, L. H. Bailey & H. H. Hume (F). HrirrsBonovaH Co.: 
Tampa, Sept. 1877, A. P. Garber ((NY, type of Laciniaria chloro- 
lepis Small.), P (specimen to right), US); near Tampa, Nov. 19, 
1928, Mrs. C. A. Miles (NY); Tampa, Aug. 25, 1903, N. L. Britton 
& P. Wilson, 35 (NY); low pinelands, s. of Riverview, Aug. 22, 
1922, J. K. Small, J. W. Small & J. B. DeWinkeler, 10605 (NY). 
Brevard Co.: low open meadow, Melbourne, Sept. 24, 1927, 
O. F. Burger & E. West (F). INptaN River Co.: damp grassy 
prairie, halfway between Fellsmere and Sebastian, Aug. 13, 1925, 
R. M. Harper, 53 (G, NY). MawNarEE Co.: Palmetto, Aug. 21- 


1946] CGaiser,— The Genus Liatris 237 


23, 1895, G. V. Nash, 2480 (G, NY, P, ND, F); Bradentown, 
July 5, 1900, S. M. Tracy, 7078 (G, NY, US); pinelands, near 
Salt Springs, Myakka Pen., Aug. 27, 1922, J. K. Small, J. W. 
Small & J. B. DeWinkeler, 10612 (NY); flatwoods, Bradentown, 
Sept. 17, 1916, A. Cuthbert (F); flatwoods, Bradentown, Sept. 16, 
1916, A. Cuthbert (F). Sarasora Co.: flatwoods, 10 miles south 
of Venice, Aug. 18, 1945, L. O. Gaiser, Mrs. H. T. Butts & Miss L. 
Arnold (F). CHARLOTTE Co.: flatwoods ditch, s. of Punta 
Gorda, Aug. 18, 1945, L. O. Gaiser, Mrs. H. T. Butts & Miss L. 
Arnold (F). Parm Breaca Co.: low flatwoods, 5 mis. w. of Lake 
Worth, Aug. 27, 1943, E. West (F). Hrnpry Co.: in low pine- 
land, L6 mi. s. of La Belle, Aug. 14, 1929, H. O'Neill (US, F). 
Ler Co.: in pinelands, Fort Myers, Aug. 9, 1916, J. Standley, 313 
(G (NY, type of Laciniaria Nashit Small), US); sand, Ft. Myers, 
Sept. 5, 1928, G. F. Weber (F); around ponds, Fort Myers, July- 
Aug. 1900, A. S. Hitchcock, 155 (G, NY, US). CoLLIER Co.: 
Cat Tail Island, Big Cypress, Feb. 26, 1919, P. P. Sheenan 
(NY); pineland near Naples, Aug. 4, 1937, Miss E. Scull (F); 
flatwoods ditch, 9 mis. n. of Naples, Aug. 18, 1945, L. O. Gaiser, 
Mrs. H. T. Butts, Miss L. Arnold (F). 

This is a singularly interesting species since it is the only one 
occurring anywhere east of the Mississippi that has elongate, 
thickened roots. While west of the Mississippi there are L. 
punctata, with a long ramifying rootstock, and L. densispicata 
Bush, which sometimes sends up aerial shoots as from the nodes 
of a rhizome, L. Garberi gives the only example of a clustered root 
that is rather fleshy and fingeroid. In the general appearance of 
its spike, in specific characters of pappus and corolla, as well as 
its habitat in damp soil, it is related to the Spicatae rather than 
the Punctatae series. It is however the only really hirsute mem- 
ber of the series. From members of the Graminifoliae series it is 
separated by the non-pilose corolla-tube. On examination of 
three sheets of Sept. 1877, A. P. Garber, Tampa, Florida (NY, 
US, P), named Laciniaria chlorolepis Small l. c., no sharp dis- 
tinctions could be found to separate them from L. Garberi, 
while the fleshy root suggested close relationship to it. 

Series I]. PveNosrACHYAE. Glabrous to hirsute plants of 
numerous stiff spikes from large crowned perennial stocks; 
leaves linear, diminishing gradually upwards; paralleling west 
of the Mississippi the 5picatae series to the east; heads of approx- 
imately similar size and shape but with phyllaries acute to acum- 
inate and recurved to merely loosely appressed; eorolla-tube non- 
pilose within. 


238 Rhodora [SEPTEMBER 


6. Liarris PYCNOSTACHYA Michx. From an enlarged (though 
globose in young plants) woody rootstock often attaining 1 dm. 
in width: stems one to many, 6-15 dm. tall, stiff, striate, generally 
hirsute, sometimes glabrous: leaves numerous, linear, punctate, 
lower ones 1 dm. long, 4-5 mm. wide, hirsute or glabrous, grad- 
ually decreasing in length upwards and passing into bracts sub- 
tending the heads, which are 5-12-flowered, cylindrical, ca. 1 em. 
long, sessile, crowded in very dense spikes 1.5-3 dm. long, 2-3 em. 
in diameter, with a generally hirsute rachis; phyllaries herbaceous 
or purplish, lanceolate-acuminate or oblong with more or less 
acute tips, that are markedly squarrose, scarcely reflexed or 
merely lax and spreading; margin mostly ciliate when herbaceous 
but frequently merely crisped and sometimes petaloid; corolla 
phlox-purple, occasionally white, 7-9 mm. long, tube nonpilose 
or with very few hairs within; achenes 4-7 mm. long; pappus 6-7 
mm. long, barbellate.—Michx. Fl. Bor. Amer. ii. 91 (1803); 
Torr. & Gray, Fl. N. Am. ii. 74 (1841), x and $; Gray, Synop. FI. 
P. 110 (1884). L. brachystachya Nutt. Journ. Acad. Phil. vii. 72 
(1834) and Trans. Amer. Phil. Soc. vii. 284 (1841). L. Bebbiana 
Rydb., Brittonia, i. 99 (1931). Laciniaria Langloisii Greene 
Pittonia, v. 58 (1902). Liatris Langloisii (Greene) Cory, 
RHODORA, xxxviii. 407 (1936). Laciniaria macilenta Small, Man. 
S. E. Fl. 1333 (1933). Liatris pycnostachya f. Hubrichti E. 
Anderson, Bull. Mo. Bot. Gard. xxv. 122 (1937) (albino form). 

From the prairies of Indiana to South Dakota southward to 
Louisiana, Texas, and Oklahoma.—INDIANA. Jasper Co.: 
Without stated locality, Aug. 7, 1878, T. M. Coulter (G). New- 
TON Co.: in prairie, along Penn Rwy., just w. of Goodland, Sept. 
5, 1988, C. C. Deam, 59098 (O). Vrao Co.: in prairie habitat, 
along Vandalia Rwy., 4 mis. s. e. of Atherton, Sept. 23, 1917, 
C. C. Deam, 23994 (G). KENTUCKY. Co. undetermined: 
Barrens of Kentucky, 1835, C. W. Short (NY). WISCONSIN. 
Co. undetermined: Kalbs Farm, near East River, Aug. 5, 1878, 
J. H. Schuette (G, NY). Juneau Co.: Camp Douglas, July 16, 
1890, E. A. Mearns (NY). Corumsia Co.: Dells of Wisconsin, 
July 20, 1886, C. H. Sylvester (NY). SAvK Co.: Kilbourn, Aug. 
26, 1909, FE. S. Steele, 33 (G, NY). MirwaukEk Co.: Bay 
View, July 31, 1880, Dr. Hasse (NY). Dane Co.: Lake Geneva, 
Mrs. E. Bayer (US); Madison, Aug. 30, 1893, J. R. Churchill (G). 
Rock Co.: Clinton, along the Chicago N. W. Rwy., Sept. 1, 
1909, E. S. Steele, 109 (G). MINNESOTA. PENNINGTON 

30.: 24.5 mis. s. of Thief River Falls, July 18, 1934, J. B. Moyle, 
1326 (NY). Hvnnanp Co.: Park Rapids, July 15, 1940, H. 
A. Gleason, 9443 (NY). Pore Co.: Glenwood, Aug. 1891, 
B. C. Taylor, 3641 (G, NY); Montevideo, Aug. 6, 1899, L. 
Moyer (NY); Montevideo, Aug. 9, 1906, L. Moyer (NY). Hex- 
NEPIN Co.: St. Anthony Park, Minneapolis, July 18, 1888, 


1946] (aiser,-—The Genus Liatris 239 


J. H. Schuelte (G, NY); Fort Snelling, Aug. 19, 1889, £. A. Mearns 
(G); wet prairie, Fort Snelling, July 31, 1888, E. A. Mearns (G); 
Fort Snelling, Aug. 28, 1881, E. A. Mearns (NY); Fort Snelling 
Reservation, Aug. 7, 1909, C. O. Rosendahl, 2344 (G). GOODHUE 
Co.: Zumbrota, Aug. 1892, C. A. Ballard (NY). Murray Co.: 
Lake Shetek, July 1922, F. P. Metcalf, 1897 (G, NY); without 
stated locality, July 1900, J. M. Holzinger (NY). ILLINOIS. 
Without stated locality: S. B. Mead (G, NY), M. S. Bebb (US). 
Co. undetermined: prairies, 1837, C. W. Short (G); prairies, 1849, 
G. Vasey (G). McHenry Co.: Ringwood, G. Vasey, 2 (G). 
WINNEBAGO Co.: Fountaindale, M. S. Bebb (G (NY type of L. 
Bebbiana) US). Cook Co.: moist prairie, e. border of River- 
side, Aug. 24, 1909, E. S. Steele, 142 (G). Kane Co.: Aurora, 
Sept. 1884, T. E. Boyce, 1180 (G). Rock IsrAND Co.: moist 
prairies, near Barstow, Aug. 1889, F. E. McDonald (G). Knox 
Co.: damp prairie, Yates City, Aug. 1912, F. E. McDonald (G). 
McLean Co.: prairie, Bloomington, Aug. 1886, B. L. Robinson 
(G). Hancock Co.: Augusta, 1842, S. B. Mead (G); S. B. Mead, 
Herb. H. A. Gleason, 1208 (G). CHAMPAIGN Co.: Champaign, 
July 23, 1898, H. A. Gleason, 292 (G); Big Four Rwy. toward 
Maryville, vicinity of Urbana, Aug. 11, 1910, E. S. Steele (US). 
Cass Co.: Beardstown, Aug. 1842, C. A. Geyer (G). ApAMs Co.: 
prairie, w. of Camp Point, July 18, 1941, R. A. Evers, G. N. Jones 
& F.F. Jones, 569 (G). Canuouw Co.: prairie, Sept. 13, 1914, R. 
Ridgway 89 (G). Mapison Co.: wet prairies, Aug. 5, 1878, H. 
Eggert (G); Carbon, Aug. 5, 1878, H. Eggert (NY, US). Ricu- 
LAND Co.: near Olney, Sept. 6, 1914, R. Ridgway (G). WASHING- 
TON Co.: Irvington, Aug. 13, 1873, G. H. French (NY, US). 
IOWA. Without stated locality: Chapman (NY). Emmer Co.: 
Armstrong, Aug. 15, 1897, R. I. Cratty (G). Dickinson Co.: low 
prairie s. w. of Miller's Bay, w. of Okoboyi Lake, Aug. 11, 1916, 5. 
Shimek (G, US). FAvETTE Co.: Fayette, Aug. 1894, B. Fink (G). 
HarDIN Co.: vicinity of Iowa Falls, Aug. 1928, M. E. Peck (G). 
HAMILTON Co.: 2 mis. n. w. of Webster, Aug. 29, 1933, Miss A. 
Hayden, 412 (NY). MansHaALL Co.: La Moille to Marshall- 
town, Aug. 21, 1927, C. C. Lounsbury (O). Sronv Co.: Ames, 
Aug. 8, 1896, L. H. Pammel & C. R. Ball, 40 (G, NY, US); Ames, 
Aug. 10, 1872, J. C. Arthur (NY); Ames, Aug. 4, 1907, R. S. 
Jeffs (O). JonNsow Co.: Iowa City, Aug. 25, 1882, B. Shimek 
(US); Lake Macbride, July 1938, J. L. Lofek (NY). PowksHIEK 
Co.: Grinnell, M. E. Jones (NY). Darras Co.: without stated 
locality, Aug. 8, 1867, J. A. Allen (G). Harrison Co.: Wood- 
bine, 1875, L. H. Horpradt, 60 (US). Decarur Co.: prairies, 
Aug. 24, 1897, T. J. Fitzpatrick & M. F. L. Fitzpatrick, 13548 
(G). MISSOURI. Co. undetermined: Wolf Creek to Inde- 
pendence City, July 20, 1849, A. Fendler (G); Ozark region, St. 
Louis & San Francisco Rwy., July 25, 1896, N. M. Glatfelter 


240 Rhodora [SEPTEMBER 


(US). Crank Co.: Medill, Aug. 24, 1910, B. F. Bush, 9169 (NY). 
ADAIR Co.: Kirksville, Aug. 6, 1887, C. S. Sheldon (NY). Rous 
Co.: Hannibal, Sept. 8, 1916, J. Davis, 251 (G). Jackson Co.: 
prairies, Martin City, July 25, 1902, K. K. Mackenzie, 46 (NY); 
Buckner, Aug. 15, 1886, B. F. Bush (US). Sr. Lours Co.: St. 
Louis, Drummond (G); St. Louis, Aug. 1845, G. Engelmann (G); 
St. Louis, July 22, 1910, E. E. Sherf (G); wet prairie hillsides, 
Aug. 5, 1878, H. Eggert (US). Cass Co.: w. Belton, Aug. 4, 1902, 
K. K. Mackenzie, 96 (NY). Henry Co.: Clinton, July 31, 1911, 
W. L. McAtee, 3035 (G, US). Vernon Co.: near Horton, Aug. 1, 
1919, W. L. McAtee, 3043 (US). Pork Co.: 5 mis. n. of Bolivar, 
Aug. 1, 1937, J. A. Steyermark, 24046 (NY). WEBSTER Co.: 
without stated locality, Aug. 1883, Miss Hosmer (G). JASPER 
Co.: dry prairies, Webb City, July 26, 1908, E. J. Palmer, 1348 
(US); high prairies, Sarcoxie, July 15, 1914, E. J. Palmer, 6262 
(US); Sarcoxie, Aug. 14, 1897, C. H. Demetrio, 45 (NY); Car- 
thage, Sept. 4, 1913, F. W. Pennell, 5369 (NY). Onkcow Co.: 
3 mis. n. of Kosh Konong, Aug. 9, 1934, J. A. Steyermark, 14383 
(US). ARKANSAS. Without stated locality: Leavenworth 
(G), 1882, F. L. Harvey (US). Co. undetermined: northwest 
Ark., Aug. F. L. Harvey 12 (G). Benton Co.: 1899, E. N. 
Plank (NY). WasnuixGTON Co.: without stated locality, 1895, 
J. W. Blankinship, 17 (G). FRANKLIN Co.: moist meadow, 
Charleston, July 17, 1935, F. J. Scully, 372 (G). PRAIRIE Co.: 
Hazen, July 25, 1937, D. Demaree, 15460 (O); waste areas in 
Grand Prairie, Hazen, June 29, 1941, D. Demaree, 22293 (G). 
Lonoke Co.: Carlisle, July 31, 1938, D. Demaree, 18010 (QO). 
GARLAND Co.: open dry woods, Glenwood Rd., Hot Springs, 
July 1, 1938, F. J. Scully, 1072 (US). AnKANSAS Co.: Stuttgart, 
July 18, 1937, D. Demaree, 15424 (O). Crarx Co.: low pine 
woods, Gurdon, Oct. 23, 1893, C. Mohr (NY, US). Howarp 
Co.: Baker Springs, Oct. 2, 1909, J. H. Kellogg (US). LOUISI- 
ANA. Without stated locality: Leavenworth (NY), Hale (G, 
NY). Narcurrocues Co.: dry sandy woods, Oct. 3, 1915, E. J. 
Palmer, 8813 (US). Rares Co.: Alexandria, Hale (NY, US). 
SAINT TAMMANY Co.: Covington, Sept. 30, 1911, N. F. Peterson 
(NY); Covington, April 5, 1912, Bell & Peterson (NY); 1 mi. n. 
of Abita Spring, Aug. 13, 1912, F. W. Pennell, 4140 (NY); 1 mi. 
n. of Abita Spring, Aug. 17, 1912, F. W. Pennell, 4252 (NY). 
ACADIA Co.: wet prairies, Oct. 1, 1895, A. B. Langlois (ND, type 
of L. Langloisii Cory). Jerrerson Davis Co.: low prairies, 
Welsh, Sept. 10, 1915, E. J. Palmer, 8483 (US).  Carcasikv Co.: 
Lake Charles, Aug. 7, 1897, S. M. Tracy, 3447 (NY); moist 
pineland n. w. of West Lake, Sept. 23, 1913, F. W. Pennell, 5615 
(NY). SOUTH DAKOTA. Roserts Co.: White Rock, 1903, 
Mrs. H. D. Powell (G). NEBRASKA. Cass Co.: low meadows 
& prairies, Weeping Water, T. A. Williams (US); Wabash, Aug. 


1946] Gaiser, —The Genus Liatris 241 


1, 1889, T. A. Williams, 196 (US); Nehawka, G. D. Swezey, 189 
(NY). BurraLo Co.: prairies, Aug. 1922, W. E. B., 13071 (O). 
KANSAS. Hirky Co.: Manhattan, Aug. 9, 1892, J. B. Norton 
(US, ND). Miami Cb.: without stated locality, July 23, 1884, 
J. H. Oyster (NY). CHEROKEE Co.: without stated locality 
1898, L. V. Harvey (US). MowTGoMERv Co.: Caney, Aug. 31, 
1895, J. W. Blankinship (G). OKLAHOMA. Co. undeter- 
mined: Choctaw Agency!, July 19, 1853, Whipple’s Exped., J. 
M. Bigelow (NY, US); Red River, Dr. Pitcher (NY). OTTAWA 
Co.: in dry pasture, Ottawa, Aug. 29, 1913, G. W. Stevens, 2502 
(G, NY). Osace Co.: in low places in prairies, Copan, Aug. 18, 
1913, G. W. Stevens, 2143 (G, US, T, O). Tutsa Co.: near Daw- 
son, July 8, 1928, J. J. Meyer, 112 (O); e. of Dawson, July 8, 1928, 
Mr. Meyer (O); s. of Tulsa, July 22, 1937, P. V. Beck, 169 (O); 
Tulsa, July 17, 1928, Miss E. R. Force (O). Noste Co.: near 
Perry, Bayliff (O). CHEROKEE Co.: wet prairies, Aug. 19, 1895, 
J. W. Blankinship (G, US). Creek Co.: Sapulpa, July 27, 1894, 
B. F. Bush 218 (G, US). Muskogee Co.: moist prairies, 3 mis. e. 
of Muskogee, July 18, 1926, E. L. Little, 190 (O); Lot 1, July 23, 
1927, E. L. Little, 1992 (O); Lot 3, Aug. 28, 1927, E. L. Little, 
2453 (O). LrFLore Co.: Page, Sept. 8, 1913, G. W. Stevens, 
269114 (G); Page, July 11, 1914, O. W. Blakeley, 1497 (G); 
Talihina, Aug. 4, 1933, U. T. Waterfall (NY). Latimer Co.: 
Laura, June 16, 1930, O. M. Clark (O). PrrrsBURGH Co.: 
McAlester, July 8, 1894, C. S. Newhall, 2 (G). PowToroc Co.: 
near Stonewall, July 10, 1891, C. S. Sheldon, 126 (US). McCur- 
TAIN Co.: open sandy ground, Idabel, July 22, 1915, E. J. 
Palmer, 8372 (US). STEPHENS Co.: 8 mis. s. of Dixie, July 26, 
1905, A. H. Van Fleet (O). Cuocraw Co.: dry open ground, Ft. 
Towson, July 16, 1915, E. J. Palmer, 8304 (US); ?Ft. Towson, 
Leavenworth (G, NY). Bryan Co.: vie. of Durant, 1931, W. L. 
Blain, 222 (US). TEXAS. Without stated locality: 1843, 
Lindheimer, 74 (G); C. Wright (G). Morris Co.: Naples, June 
27, 1935, H. B. Parks & V. L. Cory, 14426 (T). SMITH Co.: 
Swan, June 10, 1902, J. Reverchon, 3305 (US). HovsroN Co.: 
sandy open bogs, Grapeland, Sept. 16, 1915, E. J. Palmer, 14406 
(US); open sandy bog, Grapeland, Sept. 22, 1917, E. J. Palmer 
12843 (US). Trinrry Co.: 5 mis. s. w. of Trinity, Sept. 29, 
1934, V. L. Cory, 10477 (G, T). Newron Co.: without stated 
locality, July 23, 1939, B. C. Tharp (G); 21 mis. n. of Dewey- 
ville, Oct. 4, 1934, V. L. Cory, 10963 (G). TYLER Co.: without 


1 In J. M. Bigelow— Report on Lieut. Whipple's Expedition, 1857, p. 96, Sec. 4, 
no mention is made of this species being collected. The maps and notes of country 
covered refer to Oklahoma. 

2 According to Barnhart, Journ. N. Y. Bot. Gard. xxii. 131 (July, 1921), Dr. 
Leavenworth, then a surgeon in the U. S, Army, was first stationed at Fort Towson 
(which is in Choctaw Co.), and it is probable that this collection was made in that 
vicinity. 


242 Rhodora [SEPTEMBER 


stated locality, July 22, 1939, B. C. Tharp (G). WALKER Co.: 
on blackland prairie, Huntsville, July 9-12, 1909, R. Dixon, 402 
(G, NY); without stated locality, 1920, S. R. Warren, 80 (US). 
HarDIN Co.: 2.6 mis. e. of Camp Jackson, Sept. 13, 1936, V. L. 
Cory, 19797 (G); (an albino) 2.6 mis. e. of Camp Jackson, Sept. 
13, 1936, V. L. Cory, 19798 (G); without stated locality, B. C. 
Tharp (NY); Silsbee, Sept. 1926, F. W. Pennell, 5596 (NY). 
Harris Co.: Cypress, Aug. 1877, S. Ball, ex Herb. Reverchon, 
752 (G); La Porte, Aug. 6, 1939, H. R. Reed, 314 (G) ; Bammel's 
Ranch, July 18, 1925, H. Ness (T); Pierce, July 19, 1901, S. M. 
Tracy, 7331 (NY). AvsriN Co.: San Felipe, Aug. 1832, T. 
Drummond, 142 (G); ca. 7 mis. n. e. of Bellville, July 22, 1939, 
B. C. Tharp (G); Austin, Aug. 17, 1922, B. C. Tharp, 1393 (US). 
JEFFERSON Co.: Fannett, Aug. 28, 1932, B. C. Tharp (G). 
GALVESTON Co.: 13 mis. n. w. of Galveston, Aug. 4, 1939, H. R. 
Reed, 29 (G). Fort Benn Co.: Rosenburg to Wallis, July 11, 
1929, B. C. Tharp, 7497 (G); Sugarland, Aug. 5, 1933, O. Degener, 
5174 (NY). Brazoria Co.: Angleton, Sept. 14, 1933, B. C. 
Tharp (G). Wuarton Co.: between Ganado & Wharton, Sept. 
19, 1921, R. S. Ferris & C. D. Duncan, 3253 (NY). MaATAGORDA 
Co.: Palacios, July 2, 1939, B. C. Tharp (NY); without stated 
locality, July 2, 1939, B. C. Tharp (G). Jackson Co.: near 
Edna, July 18, 1930, S. E. Wolff, 2421 (US); eastern part of 
county, July 22, 1939, B. C. Tharp (G). 


André Michaux (Fl. Bor.-Amer. ii. 91 (1803)) deseribed L. 
pycnostachya from *'the Illinois meadows” together with L. macro- 
stachya from Virginia and Carolina. "Though there is no refer- 
ence made to the collection of Liatris or Serratula in his journal 
(Proc. Amer. Phil. Soc. xxiv. no. 129, pp. 1-145 (1889)), he 
botanized along the Mississippi to the vicinity of Kaskaskia, a 
point 94 miles south of St. Louis. In the map of F. A. Michaux, 
(Travels to the westward of the Alleghany mountains, J. Maw- 
man, 350 pp. (1805)), that region east of the Mississippi was 
called Illinois. The descriptions of the two species are very 
similar, the most distinctive difference being the nature of the 
phyllaries, squarrose at the tips in the former and appressed in 
the latter. A second difference is in pubescence, stem hirsute 
and leaves pubescent in pycnostachya and cilia basally on the 
shining leaves of macrostachya. Much of the puzzle in limiting 
this species has been bound up with the varying expressions of 
these two characters. Michaux’s L. macrostachya seems none 
other than Linnaeus' L. spicata. 

Nuttall (Jour. Acad. Phil. vii. 72 (1834)) described a new 


1946] Gaiser,—The Genus Liatris 243 


species, L. brachystachya, for a very glabrous plant collected by 
Dr. Pitcher in Arkansas. He, however, made it synonymous 
with L. pycnostachya Michx. in a later publication (Trans. Amer. 
Phil. Soc. n. s. vii. 284 (1841)) and this procedure was also 
followed by Gray l. c. for the glabrous variety 6, that had been 
adopted by Torr. & Gray |. c. from the description of Nuttall’s 
species. Delimitation of hirsute and glabrous individuals as 
varieties seems useless because they occur without any special 
geographical range and various intermediates are so abundant. 

Examination of a photograph of Michaux’s type specimen 
(Paris), shows the phyllaries of the expanded heads as not mark- 
edly squarrose and neither sharply acute nor long-acuminate but 
rather oblong or lanceolate-acuminate and slightly reflexed 
(“apice reflexis" as given in Michaux’s handwriting on the label 
of the specimen). The rachis and leaves clearly show a hairy 
condition. 

Comparison of many specimens from the Mississippi basin 
shows a variation in phyllaries from long-acuminate, quite 
strongly recurved, to oblong with acute tips and less recurved, 
and to merely spreading. Also, although the stem is generally 
hirsute there is a range from plants with glabrous stems and 
almost glabrous recurved phyllaries without ciliate margins, 
through plants with hirsute stems and only scattered hairs along 
the midveins beneath and scant cilia on the margins of the outer 
phyllaries while the remainder have a crisped margin, to those 
with more generally hirsute leaves and phyllaries with ciliate 
margin to completely hirsute phyllaries. 

Rydberg (Brittonia i. 99 (1931)) described L. Bebbiana from 
Illinois, stating it was “related to L. pycnostachya but the outer 
bracts are ovate and merely acute, not lanceolate and long 
acuminate”. A detailed study of the type specimen (M. S. 
Bebb, Fountaindale, Winnebago Co., Ill., (NY)), has failed to 
disclose how this or any other fundamental character can sepa- 
rate it from L. pycnostachya Michx. 

The most extreme cases of only slightly recurving non-acumi- 
nate phyllaries, combined with a glabrous condition, approach L. 
spicata (as F. L. Harvey, no. 12, from N. W. Arkansas (G), or of 
1849, G. Vasey no. 2, Ringwood, McHenry Co., Ill. (G)). Or 
again, as referred to under L. lancifolia (see no. 2), in the more 


244 Rhodora [SEPTEMBER 


western states, intergrades between the two of that region are 
suggested by ovate, ciliate-margined or but slightly crisped 
phyllaries. An occasional specimen with very broad leaves, as 
in L. lancifolia, but with the more pycnostachya-like inflorescence, 
as of July, 1891, Hapeman, from Platte Valley, at Fort Kearney, 
Neb. (US), gives further evidence of this mixture. 

Greene (Pittonia v. 58 (1902)) described L. Langloisii from 
Louisiana, as allied to L. pycnostachya but well marked by its 
pale herbage and colorless involucral bracts, “wanting the purple 
tips of the kindred species". Examination of the type specimen 
(Oct. 1, 1895, A. B. Lamglois, from wet prairies, Acadia Co., 
Louisiana (ND)), showed that at the time of collection the spike 
had passed the blooming stage and was in seed, at which time 
(as noted in the introduction) the phyllaries of Spicatae and 
Pycnostachyae, if they have been colorful, usually become green. 
The strongly hirsute condition is marked, giving a glaucescent 
green color. But specimens from the more southern end of the 
range are frequently very hirsute without any correlation with 
lack of color of the phyllaries or smaller size of the achene. 
Thus the various combinations of herbaceous or purple-petaloid, 
glabrous or hirsute phyllaries, make it as difficult to draw sharp 
distinctions between L. pycnostachya and L. Langloisii, as 
acuminate or ovate, recurving or spreading phyllaries make it 
impossible to decide between L. pycnostachya and L. Bebbiana. 

Specimens from Louisiana, as one showing markedly recurved 
phyllaries (Sept. 10, 1915, E. J. Palmer, no. 8483, from low 
prairies, Welsh, Jefferson Davis Co., (US)) and another showing 
them less so (Oct. 3, 1915, E. J. Palmer, no. 8813, dry sandy 
wood, Natchitoches (US)) give further support to the explana- 
tion offered for puzzling hirsute forms allied to L. spicata (see 
no. 1), as possible intermediates between these two species at the 
southern end of their ranges. "The specimen of Aug. 20, 1903, 
S. M. Tracy, no. 8533, from Mendenhall, Simpson Co., Miss. 
(G, T) is still the only evidence I have seen of pycnostachya-like 
plants from Mississippi and since specimens from Louisiana, as 
that of A. B. Langlois and those of Bro. Arséne, show such vary- 
ing degrees of hirsuteness combined with spreading of phyllaries, 
I prefer to treat them together as intermediates between spicata 
and pycnostachya., 


1946] Gaiser,— The Genus Liatris 245 


As Gray (Synop. Fl. P. 111 (1884)) stated of L. pycnostachya: 
"Apparently this hybridizes with L. spicata: at least specimens 
oceur which are intermediate between the two species." This 
is probably as true for the conditions on the frontiers of their 
-occurrence (i. e. in Louisiana and Nebraska) as in the center of 
their overlapping ranges (specimen of Aug. 24, 1897, T. J. & M. 
F. L. Fitzpatrick, no. 13548, from prairies of Decatur Co., Iowa 
(G)). The number of intermediates is great and their variety 
so wide as to make impossible a type description of such hybrid 
populations. Since these two species are the most favored of 
the genus in perennial borders, many garden specimens showing 
different kinds of intermediacy are met with under a variety of 
horticultural names, not included here. 

Records of single collections of each of two other interspecific 
hybrids follow. 


X Liarris lüpcGwavi Standl. (L. pycnostachya X squarrosa). 
Upper portion of plant only seen: stem slightly pilose; leaves 
linear, pubescent on under surface, gradually decreasing in length 
upwards; spike 19 em. long and 3 em. wide; heads ca. 18-flowered, 
sessile, somewhat crowded, subtended by foliar bracts, longer 
than or about equalling the length of the heads; outer phyllaries 
lanceolate-oblong, acuminate, almost entirely green or purple- 
tipped, pilose, with ciliate margins and somewhat squarrose 
apices; inner phyllaries oblong, appressed, acute or acuminate, 
almost glabrous, purple-tipped and recurved; corolla purple, 10 
mm. long, inner surface of lobes hairy; achenes 3-5 mm. long; 
pappus 9 mm. long and short-plumose.— RHopona, xxxi. 37 
(1929).—ILLINOIS. RICHLAND Co.: east of Bethel Church, 
Aug. 26, 1928, R. Ridgway, 3265 (Field Museum no. 579880, 
TYPE, not seen; isotypes, G, US). 

This rare plant was found growing with L. pycnostachya and L. 
squarrosa and, though it resembles the former more closely in 
general appearance, it is intermediate in other characters, as in 
number, distribution and size of heads and in the pappus. In 
the outer green phyllaries, the long foliose braets subtending the 
heads and the hairy corolla lobes, it resembles L. squarrosa. 

X Liarris Frostii, hybr. nov. (L. aspera X pycnostachya). 
Caulis unus 10.3 dm. altus superne pilis albis crispatis dense 
hirsutus denseque foliosus solum visus; foliis basalibus lanceolatis 


1.5 em. longis, 6 mm. latis sessilibus paginis ambobus hirsutis, 
medianis gradatim. minoribus minusque hirsutis, superioribus 


246 Rhodora [SEPTEMBER 


longitudine capitula aequantibus haud hirsutis; capitulis multis 
inter se distantibus 13-floris sessilibus leviter campanulatis ca. 
13 mm. longis; phyllariis obtusis petaloideis marginibus pur- 
purascentibus, exterioribus brevibus oblongis marginibus cris- 
patis, interioribus apice leviter dilatatis plus petaloideis ca. 9 
mm. longis; corollis 9 mm. longis intus pilosis; achaeniis 4 mm. 
longis, pappo 7 mm. longo vix plumoso nisi manifeste barbellato. 
—MINNESOTA: Kawnrvour Co.: Spicer, Aug., 1892, W. D. 
Frost (G: TYPE). 


This single specimen, like which no others have been seen and 
about the collection of which there is no exact information, 
strongly suggests another case of hybridization, and between 
two species for which there had previously been no such report. 
It resembles L. pycnostachya, à species occurring throughout 
that part of Minnesota, specimens of which from the adjoining 
county of Polk have been seen, in the more general appearance 
of the stem with many narrow leaves, and a fairly dense spike of 
numerous elongate heads. However, in the size of the heads 
and the obtuse tips and petaloid, erisped margins of all the 
phyllaries, it resembles L. aspera. From the township of White- 
field, lying about 8 miles south of Willmar and about 20 miles 
south and west of Spicer, two specimens have been seen of the 
same collector, W. D. Frost, no. 4893, collected in the same 
month, one of which, though not typical, is near L. aspera 
(US884659) and the other, L. lgulistylis (US201918), two species 
abounding and also hybridizing freely in that district (see no. 20). 
On examination of the corolla-tube of this particular specimen, 
X L. Frostii, it was found to be pilose within as in L. aspera thus 
pointing more toward that parentage than to L. ligulistylis. 


Series III. QGmaAMriNIFOLIAE. Plants with generally more 
open inflorescences than Spicatae and Pycnostachyae, varying 
from stiff-spicate to branched-paniculate and slender-racemose; 
leaves mostly ciliate along the petioles or with scattered hairs on 
mid-veins, if not entirely hirsute; heads 3-20-flowered, .7-1.5 em. 
long, turbinate when flowers are open; phyllaries obtuse, with 
fine ciliolate margins mostly appressed when in bud but loosely 
spreading in mature heads; corolla-tube pilose within; achene 
3-5 mm. long. 

From the coastal plain of New Jersey to Florida and westward 
through and along the Appalachian ridge to the Gulf of Mexico. 


1946] Gaiser,— The Genus Liatris 247 


a. All leaves linear to linear-lanceolate; phyllaries spreading in 
mature turbinate heads. . . . b. 
b. Heads 5-15-flowered. . . .c. 
c. Phyllaries without prominent mid-vein and with ciliolate 
margin; inflorescence varying from slender few-headed 
racemes to loose spikes or panicles... .d. 
d. Pappus about as long as the corolla-tube....... 7. L. graminifolia. 
d. Pappus only half as long as the corolla; found only in 
the mountains of several adjacent counties i in western 
Nese aii ama aa E T LORI. 20 as 8. L. Helleri. 
c. Phyllaries with prominent midvein or keel and with en- 
tire margin; inflorescence generally loose, spicate to 
pamculates Mog M. c DERE po ll. L. regimontis. 
b. Heads 3-5-flowered; phyllaries without keel and quite 
ciliate; pedicels slender and often at right angles to the 
stem; heads short-turbinate when flowers are open....10O. L. gracilis. 
a. Lower leaves more bro: idly lanceolate, upper ones linear- 
lanceolate; heads 10-20-flowered; inflorescence more stiffly 
spicate; phyllaries without keel, obtuse, oblong to orbicular 
and appressed when in bud, with narrowly scarious or 
ciholate margins e o runc diee MN inae 9. L. turgida. 


7. LIATRIS GRAMINIFOLIA (Walt.) Willd. Corm globose, up to 
3 em. in diameter; stems single or few, sometimes striate, 3—7 dm. 
high, glabrous or with long scattered hairs or hirsute; basal 
leaves 10-15 cm. long, narrowly linear (2 mm. wide) or linear- 
lanceolate (5-10 mm. wide), with ciliate margins or with long 
hairs along the winged petioles and sometimes scattered over the 
lower surface; upper leaves reduced in length and width: inflores- 
cence either a long slender spike, or a fairly dense raceme that 
frequently becomes paniculate, or an open short raceme of a few 
scattered heads: heads sessile or pedicellate, of 5-15 flowers, 
10-15 mm. high, 6-12 mm. wide and somewhat turbinate in 
shape at the time of flowering; phyllaries thin, glabrous or hirsute, 
loose, not overlapping much laterally, narrowly linear to ovate- 
lanceolate, 1-4 mm. wide, obtuse, scarious and very finely 
ciliolate on the margin; corolla 6-8 mm. long, purple; pappus 5-6 
mm. long, barbellate; achene 3-4 mm. long.—Sp. Pl. iii. 1636 
(1803). Anonymos graminifolia Walt. Fl. Car. 197 (1788). 

From New Jersey and southern Pennsylvania southward to 
Florida and Alabama. 


KEY TO VARIETIES 


a. Inflorescence somewhat spicate, though often paniculate, and 
of quite numerous heads; phyllaries narrowly linear, 1-3 
mm. wide. E 

b. Leaves narrowly linear; stem and leaves with scattered 
hairs; heads small, with narrowly linear phyllaries ca. 1 
mm. wide; from the coastal plain region of Virginia and 

Lhe: Carolinas a4 era ee eh DD UR ee T var. typica. 
b. Basal leaves broader, linear-lanceolate; stem and leaves 
mostly glabrous, but with long hairs along the winged 
petioles; heads larger; of more general distribution in 

the northeastern range of the species.................... var. dubia. 


248 Rhodora [SEPTEMBER 


b. Stem and pedicels densely hirsute; leaves linear-lanceolate; 
of northern range... 2.6.00 es var. lasia. 
a. Inflorescence slender, somewhat virgate, with few scattered 
heads; phyllaries a little broader, 3-4 mm. wide and fre- 
quently rounded at the tip....c. 
e. Shorter plants of few leaves, the basal linear-lanceolate, 
much longer than the others; from mountains in Vir- 


ginia, North and South Carolina.................0.... var. Small. 
c. Taller plants with more numerous linear leaves; from 
Georgia, Alabama and Florida...................... var. elegantula. 


Var. typica. Stems tall, slender, with long scattered hairs: 
leaves numerous, generally with abundant long hairs; basal ones 
narrowly linear, 10-12 em. long and 2-4 mm. wide, with ciliate 
margins; upper ones reduced in lengtb, and in width to 1 mm.: 
inflorescence usually a long, slender, dense spike of 2 em. width 
when flowers are open, sometimes becoming paniculate: heads 
5-9-flowered, mostly sessile and erect, ca. 1 em. high, 5-6 mm. 
wide when flowers are open; phyllaries narrowly linear, ca. 1 mm. 
wide; corolla-tube 6-7 mm. long; pappus 4-5 mm. long; achene 
ca. 3 mm. long.—Anonymos graminifolia Walt. Fl. Car. 197 
(1788). LIATRIS GRAMINIFOLIA Willd. Spec. Pl. iii. 1636 (1803); 
Ell. Sk. ii. 274 (1824). 

New Jersey to Alabama, chiefly on the coastal plain.—NEW 
JERSEY.  OckEAN Co.: in sandy field, Forked River, Aug. 31, 
1937, H. N. Moldenke, 10172 (NY). BunuiNGTON Co.: Atsion, 
Aug. 1877, T. C. Martindale (US). CAMDEN Co.: Atco, Sept. 15, 
1904, T. W. Edmondson, 1201 (G). ArraANTIC Co.: Weymouth, 
Sept. 12, 1923, E. P. Killip, 12395 (US). Carre May Co.: on 
sand dunes, Five Mile Beach, Oct. 2, 1889, A. MacElwee 1413 
(NY); in sandy Rwy. bed, Wildwood, Aug. 6, 1928, A. Moldenke, 
4084 (NY); without stated locality, Sept. 20, 1894, C. Mohr 
(US). MARYLAND. Harrorp Co.: without stated locality, 
Aug. 10, 1873, J. W. Eckfeldt (P); along Bush R., n. of Bush R. 
Sta., Sept. 10, 1902, G. H. Shull (G). Baurimore Co.: near 
Baltimore, 1873, D. Foreman (US); Oakwood, near Baltimore, 
Sept. 6, 1896, A. Fredholm, 2224 (NY, US). ANNE ARUNDEL 
Co.: dry soil, right of road, from Muirkirk to Contee, Sept. 5, 
1910, A. H. Moore, 4832 (G). Prince GEonaGES Co.: low pasture 
land, vicinity of Lanham, June 28, 1910, W. R. Mazon, 4605 
(US); dry field, Carter’s Lane, Sept. 29, 1914, E. S. Steele (US). 
DELAWARE. Sussex Co.:s. of Robbins, Sept. 5, 1925, F. W. 
Pennell, 12865 (P); moist swale, 14 mi. e. of Ellendale, Sept. 23, 
1938, R. R. Tatnall, 4053 (G). DISTRICT OF COLUMBIA 
AND VICINITY. "Terra Cotta Swamp, Aug. 1895, D. L. 
Topping (US); vicinity of Washington, Oct. 1, 1876, L. F. Ward 
(US). VIRGINIA.  AnriNGTON Co.: 1 mi. e. of Rwy., Bareroft 
Station, Oct. 3, 1915, E. S. Steele (US); edge of sandy oak woods, 
on Washington Blvd., at Glebe Road, Sept. 28, 1940, F. J. 


1946] (raiser,— The Genus Liatris 249 


Hermann (NY); Rwy., Cowdons, Sept. 22, 1912, E. S. Steele, 
643156 (US). JAMES Ciry Co.: on dry sandy red soil by road- 
side, 2 mis. w. of Williamsburg, Sept. 27, 1921, E. J. Grimes, 
4625 (NY); dry barren soil, 2 mis. s. of W illiamsburg, Sept. 23, 
1921, E. J. Grimes, 4427 (NY); along C. & O. Rwy. 1 mi. w. of 
Lightfoot, Sept. 26, 1921, E. J. Grimes, 4496 (NY); dry sandy 
soll, along C. & O. Rwy. e. of Lightfoot, July 23, 1921, E. J. 
Grimes 4096 (NY). Prince GrorGE Co.: 1-3 mis. w. of New 
Bohemia, Sept. 24, 1927, E. T. Wherry & F. W. Pennell, 14415 
(P). CHESTERFIELD Co.: dryish swale, n. w. of Colonial Heights, 
Sept. 16, 1938, M. L. Fernald & B. Long, 9452 (G). Sussex Co.: 
molst pinelands, just s. e. of Waverly, Sept. 10, 1937, M. L. 
Fernald & B. Long, 7661 (( » US); damp sandy pine & oak woods, 
s. of Stony Creek, Sept. 1939, M. L. Fernald & B. Long 
11455 (G). DIN WIDDIE Co. ' ca. I mi. n. e. of Burgess, Sept. 13, 
1937, M. L. Fernald & 5b. Long 7662 (G); dry pine woods, n. W. 
of Carson, Sept. 14, 1937, M. L. Fernald & B. Long 7663 (G). 
PRINCESS ANNE Co.: along trailsides in pine-oak-hickory forest, 
Cape Henry, Sept. 3, 1940, F. E. Egler, 40-348 (NY); dry oak 
woods, The Desert, Cape Henry, Sept. 23, 1933, M. L. Fernald & 
L. Griscom, 2907 (G).  NANSEMOND Co.: dry sandy woods & 
adjacent clearings, Kilby, Sept. 11, 1935, M. L. Fernald & B. 
Long 5078 (G); Suffolk, Sept. 17, 1907, E. B. Bartram & B. Long, 
(US); Kilby, Sept. 11, 1985, M. L. Fernald, B. Long & J. M. 
Fogg, 5145 (G). SOUTHAMPTON Co.: clearing in pine & oak 
woods, w. of Branchville, Sept. 20, 1938, M. L. Fernald & B. 
Long, 9454 (G). GREENSVILLE Co.: mossy pineland e. of Slagle’s 
Pond, n. of Emporia, Sept. 20, 1938, M. L. Fernald & B. Long, 
9457 (G). NORTH CAROLINA. Without stated locality: 
M. A. Curtis (G). Co. undetermined: eastern North Carolina, 
Sept. 6, 1908, W. W. Eggleston, 4052 (G, NY). Nasu Co.: pine- 
land, at Middlesex, Oct. 9, 1938, R. K. Godfrey & T. Kerr, 6627 
(G); sandy soil, Rocky Mt., Oct. 17, 1912, F. W. Pennell (P). 
Durnam Co.: Duke Forest, ‘Sept. 16, 1932, H. L. Blomquist, 28 
(US);low open dry ground, Old Oxford Rd. , Sept. 24, 1938, H. L. 
Blomquist, 10519 (F). FonsyrH Co : Bronton suburbs, Winston 
Salem, Oct. 30, 1921, P. O. Schallert (Q). Martin Co.: swale at 
Williamston, Oct. 13, 1938, R. K. Godfrey, 7024 (G). Wake Co.: 
4 mis. n. of Raleigh, Oct. 10, 1938, R. K. Godfrey, 6688 (G). 
CHATHAM Co.: pine woodland, at Siler City, Oct. 12, 1938, R. K. 
Godfrey 6977 (G); along Pittsboro Road, near town, Sept. 29, 
1909, W. C. Coker (NY). HanNETT Co.: pineland near Lilington, 
Aug. 5, 1938, R. K. Godfrey 5673 (G). Ler Co.: open pine wood- 
land, at Sanford, Oct. 12, 1938, R. K. Godfrey, 6929 (G). Pam- 
L1CO Co.: pinelands at Grantsboro, Oct. 11, 1938, R. K. Godfrey & 
R. N. White, 6825 (G). Craven Co.: Newbern, Oct. 10, 1898, 
T. H. Kearney, 2231 (US). ONSLOW CO in dry sandy land, 10 


250 Rhodora [SEPTEMBER 


mis. s. of Jacksonville, Nov. 4, 1929, H. N. Moldenke, 116a (NY). 
New Hanover Co.: sandy pineland, Wilmington, Oct. 15, 1912, 
F. W. Pennell, 4907 (US, P); Wilmington, Sept. 20, 1888, G. 
McCarthy (US); vicinity of Wilmington, Nov. 1922, Mrs. M. H. 
Cummings (US). SOUTH CAROLINA. Co. undetermined: 
dry pineland, Santee Canal, Oct., /7. W. Ravenel (G). JASPER 
Co.: Ridgeland, Nov. 13, 1893, C. Mohr (US). Braurort Co.: 
without stated locality, 1882, Dr. Mellichamp (US). GEORGIA. 
Co. undetermined: dry rocky hillside, Oct. 8, 1893, C. Mohr 
(US). ALABAMA. MonmnirE Co.: Spring Hill, Sept. 28, 1878, 
C. Mohr (US). 

Var. puBIA (Barton) Gray. Generally the most robust- 
looking of all the varieties: stem striate and glabrous with scat- 
tered hairs: basal leaves linear-lanceolate, 6-10 cm. long, 5-7 mm. 
wide, with hairs along the winged petioles and sometimes scat- 
tered over the lower surface; upper ones reduced in length: heads 
erect, usually sessile or on short pedicels in a fairly dense raceme, 
but frequently becoming paniculate, 10—15-flowered and thus 
larger than in other varieties, 12-15 mm. high and 10-12 mm. 
thick at the time of flowering; phyllaries linear, ca. 2 mm. wide, 
obtuse and ciliolate on the margin; corolla purple (rarely white), 
6-8 mm. long; pappus ca. 5 mm.; achene 4 mm. long.— Liatris 
graminifolia var. dubia Gray, Man. ed. 1.191 (1848), ed. 2, 185 
(1856). Liatris dubia Barton, Mat. Med. ii. 223, t. 49 (1818); 
Gray, Synop. Fl. i?. 111 (1884), in part. Serratula spicata L. 
Sp. Pl. ii. 819 (1753), as to citations Gron. Virg. 92 (1739), Dill. 
Elth. 85 t. 72 (1732). Serratula pilosa Ait. Hort. Kew ed. 1, 
iii. 138 (1785). Liatris pilosa Willd. Spec. Pl. iii. 1636 (1803); 
Pursh, Fl. Am. Sept. ii. 508 (1814); Ell. Sketch n. 277 (1824); 
Ker-Gawl. Edwards Bot. Reg. t. 595 (1821). Liatris spicata Y 
racemosa DC. Prodr. v. 131 (1836). Liatris propinqua Hook. 
Bot. Mag. lxvii. t. 3829 (1841). Liatris virgata Nutt. Jour. 
Acad. Phil. vii. 72 (1834), and Trans. Amer. Phil. Soc. n. s. vii. 
284 (1841). 

New Jersey and Pennsylvania southward through North 
Carolina.—NEW JERSEY. Without stated locality: A. Gray 
(G); P. D. Knieskern (G). Co. undetermined: pine barrens, 
Sept. 1872, Ex. Herb. Miss M. Treat (G); pine barrens, Sept. 
1869, T. C. Porter (US). Ocean Co.: Tom's River, W . W. 
Denslow (NY); Quaker Bridge, Tom's River, 1860, D. C. Eaton 
(G, NY); pine barrens, 1 mi. w. of Tom's River, Sept. 12, 1930, 
J. A. Drushel, 8515 (US); sandy open depression, East Plains, 
Sept. 7, 1924, F. W. Pennell, 12918 (NY); dry pine barrens, 
Barnegat Pier, Sept. 1907, K. K. Mackenzie, 2919 (US); low pine 
barrens, Forked River, Sept. 18, 1893, L. H. Lighthipe (NY, US); 
Manchester, Aug. 28, 1879, N. L. Britton, 16 (G); Manchester, 
Aug. 31, 1878, G. Guttenberg (US); Manchester, Sept. 15, 1879, 


1946] Gailser,— The Genus Liatris 251 


A. Brown (NY); Manchester, Sept. 17, 1870, Mus. Nat. Hist. 
(NY). BumuiNGTON Co.: without stated locality, C. F. Parker 
(G); Chatsworth, Sept. 5, 1897, E. H. Eames (G); Woodmansie, 
Sept. 25, 1879, C. L. Pollard (US); in dry sandy soil, Sept. 3, 1867, 
C. F. Parker (G); wet sand, Atsion, Sept. 5, 1917, A. Gershoy, 
685 (G); Atsion, Aug. 22, 1879, O. D. Allen (G); Brown’s Mills, 
Sept. 11, 1864, E. Dillenbaugh (G); ArLantic Co.: Egg Harbor, 
Sept. 6, 1884, E. G. Knight (NY); Egg Harbor, Sept. 6, 1884, 
W. H. Manning (G); Egg Harbor, Sept. 6, 1884, L. F. Ward 
(US); sandy soil, Egg Harbor, Sept. 4, 1938, R. L. Schaeffer, 827 
(G); Hammonton, Sept. 6, 1913, W. M. Benner (G); in sand, 
Hammonton, Sept. 4, 1917, A. Gershoy, 684 (G); dry pine bar- 
rens, Hammonton, Aug. 24, 1907, Shreve & Miller (US). Cum- 
BERLAND Co.: Vineland (albiflora), 1872, Miss M. Treat (G); 
sandy pineland 3 mis. s. c. of Millville, Sept. 11, 1924, J. R. 
Pennell (NY). Care May Co.: dry sandy meadow, Briar 
Island, Sept. 25, 1915, B. Long, 13592 (G); Anglesea, Sept. 20, 
1891, C. D. Lippincott (G). PENNSYLVANIA. Bucks Co.: 
Quaker Bridge, 1882, C. D. Fretz (Q). DELAWARE. Sussex 
Co.: sandy woods, Rehoboth, Sept. 12, 1908, J. R. Churchill (G); 
edge of salt marsh, Fenwick Is., Sept. 16, 1934, R. R. Tatnall, 
2385 (G); Indian River Bay, Sept. 12, 1934, A. 5. Goodale, 77591 
(G). MARYLAND. Harrorp Co.: on dry bank along Bush R., 
n. of Bush R. Station, Sept. 10, 1902, G. H. Shull, 348 (NY, US); 
Savage Station, Sept. 4, 1905, H. D. House, 1527 (US). BALTI- 
MORE Co.: Baltimore, 1866, P. V. LeRoy (NY); without stated 
locality: Sept. 1888, G. L. S. Herb., 1179 (G); Springfield Road, 
Baltimore, 1888, J. H. Holmes (US); sandy clearing, shore of 
Bird R., 2 mis. e. of Whitemarsh, Sept. 16, 1938, F. J. Hermann 
(NY). Monrcomery Co.: Woodside, Sept. 20, 1896, H. Wolds 
(US). CarvERT Co.: en Miocene escarpment, at Little Cove 
Point, Oct. 15, 1937, H. A. Allard, 3827 (G, US). PRINCE 
GrorGEs Co.: n. of Riverdale, Sept. 23, 1916, E. S. Steele (US); 
dry woods, near Hyattsville, Oct. 13, 1919, F. W. Hunnewell 
6498 (G); sandy soil, Ardwick, Sept. 5, 1910, F. W. Pennell, 2647 
(NY); dry gravelly field, near Bladensburg, Oct. 3, 1926, S. F. 
Blake, 9734 (G); Bladensburg, 1883, McCarthy (US); dryish open 
knoll, vie. of Lanham, Sept. 24, 1910, W. R. Maxon & P. C. 
Standley, 6 (US); Silver Hill, Sept. 6, 1915, W. L. McAtee, 2346 
(US); in sphagnum bog, between Magruder & Benning, Sept. 6, 
1899, E. L. Morris, 309 (US). DISTRICT OF COLUMBIA 
AND VICINITY. Vicinity of Washington, L. F. Ward (NY, 
US); vieinity of Washington, Sept. 17, 1880, L. F. Ward (G); 
Sluice Run, vicinity of Washington, Sept. 14, 1879, L. F. Ward 
(U8); Washington, 1876, Vasey (US); Bates Road, vicinity of 
Washington, Sept. 28, 1888, E. S. Burgess (US); Terra Cotta, 
vicinity of Washington, Sept. 30, 1896, E. S. Steele (US); vicinity 


252 Rhodora [SEPTEMBER 


of Washington, Sept. 12, 1922, E. S. Steele (US); District of 
Columbia, Sept. 13, 1874, L. F. Ward, 736 (US); dry open woods, 
Brookland, Oct. 2, 1927, H. O'Neill (NY); Takoma Park, Oct. 6, 
1897, T. A. Williams (G); sandy woods, Takoma Park, Sept. 8, 
1904, H. D. House (US); Takoma Park, Sept. 6, 1905, H. D. 
House, 1533 (US); sandy woods, near Takoma Park, Aug. 25, 
1904, H. D. House, 335 (NY); sandy woods, Takoma Park, Sept. 
8, 1904, J. H. Painter, 1149 (US); Chesapeake Beach Junct., 
Oct. 5, 1911, A. Ruth, 45 (G). VIRGINIA. ARLINGTON Co.: 
Fort Barnard, near Cowdons, Oct. 1, 1916, E. S. Steele (G, NY, 
US); Cowdons, near Southern Rwy., Sept. 22, 1912, E. S. Steele 
(US). Farrrax Co.: Fort Myer, Oct. 24, 1895, E. A. Mearns 
(NY); Falls Church, Sept. 26, 1874, J. J. Carter (P); Mount 
Vernon, Sept. 27, 1902, F. L. Fisher (P). NORTHAMPTON Co.: 
Cape Charles City, Sept. 25, 1894, W. M. Canby, 823 (US); 
NORTHUMBERLAND Co.: dry woods, Coan, Oct. 17, 1916, J. 
Tidestrom, 8185 (G). RocksripGE Co.: Balcony Falls, Sept. 5, 
1883, Dr. & Mrs. N. L. Britton (NY). James Crry Co.: field, 
ca. 5 mis. w. of Toano, Oct. 1, 1939, R. W. Menzel (G); opening 
in flat pine woods, Williamsburg, Sept. 25, 1920, E. J. Grimes, 
3107 (G). CHESTERFIELD Co.: field, May, 1935, E. Veazey (G); 
dry field, near 8t. Elmo, Sept. 17, 1910, P. C. Standley, 5903 (US). 
CAMPBELL Co.: dry sunny hillside, at mouth of Otter R., near 
Altavista, Oct. 6, 1913, J. Fauntleroy, 610 (US); Lynchburg, 
Sept. 17, 1927, W. A. Murrill (F). Wyrne Co.: Walker Mt., 
Sept. 1, 1981, E. L. Core, 3372 (NY). SwvrH Co.: road between 
Marion & White Top Mt., Aug. 22, 1908, P. A. Rydberg (NY). 
Sussex Co.: open pine & oak woods, near Greensville line, s. of 
Jarratt, Sept. 18, 1938, M. L. Fernald & B. Long, 9453 (G). 
Princess ANNE Co.: dry sandy barrens, Cape Henry, Sept. 23, 
1933, M. L. Fernald & L. Griscom, 2908 (G); Cape Henry, Sept. 
23, 1933, M. L. Fernald & B. Long, 2909 (G). HarurrFax Co.: 
2 mis. n. e. of Clover, Sept. 23, 1927, E. T. Wherry & F. W. 
Pennell, 14391 (P). GREENSVILLE Co.: open thickets, clearings 
& woods, s. of Emporia, Sept. 20, 1938, M. L. Fernald & B. Long, 
9455 (G). NORTH CAROLINA. Avery Co.: slopes of 
Grandfather Mt., Sept. 25, 1898, W. M. Canby & C. S. Sargent 
& J. Muir, 71 (US). Burke Co.: without stated locality, 
1836, Curtis (NY). McDowkrr Co.: vic. of Graphiteville, 
Aug. 29, 1913, P. C. Standley & H. C. Bollman, 10086 (US). 
BuNcOMBE Co.: open hillside, vic. of Montreat, Aug. 29, 1913, 
P. C. Standley & H. C. Bollman, 10079 (US); bushy hillside, vic. 
of Montreat, Sept. 9, 1913, P. C. Standley & H. C. Bollman 
10497 (US); open woods, vic. of Montreat, Sept. 9, 1913, P. C. 
Standley & H. C. Bollman, 10527 (US). Pork Co.: Columbus, 
Aug. 10, 1897, E. C. Townsend (US). Craven Co.: Newbern, 
Oct. 10, 1898, T. H. Kearney, 22207 (US). SOUTH CAROLINA. 


1946] Gaiser,— The Genus Liatris 203 
) 


GEORGETOWN Co.: sandy pine woods, Oct. 12, 1934, F. G. Tarbox 
175-1 (NY). GEORGIA. Without stated locality: Nuttall 
(P, isotype of Liatris virgata Nutt.). 

Var. LASIA Fern. & Griscom. <A hirsute variety resembling var. 
dubia generally in arrangement and size of leaves, heads and 
flowers but with pubescent stem and leaves; the phyllaries too 
are similar, though the margins are distinctly ciliate.—RuHopora, 
xxxvii. 183 (1935). Liatris pilosa Lodd. Bot. Cab. t. 356 (1819). 

New Jersey, Delaware and Alabama.—NEW JERSEY. Co. 
undetermined: pine barrens, Sept. 1870, A. H. Smith (G). 
CAMDEN Co.: Lindenwold, Sept. 29, 1903, J. M. Fogg, 622 (G, 
type); south of R. P. R. Station, Lindenwold, Sept. 18, 1920, 
H. B. Meredith (G). CUMBERLAND Co.: 14 mi. s. of Seeley’s 
Mill, Bridgeton, Sept. 1, 1924, Beals & Bassett (G); 3 mis. s. e. of 
Millville (1 plant), Sept. 11, 1924, J. R. Pennell (NY). DELA- 
WARE. Kenr Co.:dry soil, Felton, Sept. 28, 1863, A. Commons 
(P). Sussex Co.: Rehoboth, Sept. 1908, C. S. Williamson (P). 
ALABAMA. MonmiLkE Co.: pine barrens, Spring Hill, Aug. 1919, 
E. W. Graves, 731 (US). 

Var. SMALLII (Britt.) Fern. & Griscom. Generally a shorter, 
more slender variety often with reddish non-striate stems of 3—5 
dm. and with fewer leaves: basal leaves quite long, linear- 
lanceolate (10-15 cm.) and 3-10 mm. wide, reduced upwards to 
braets usually shorter than the heads: inflorescence an open, 
short raceme of few scattered heads, averaging approximately 
one head per centimeter; rachis slender; heads generally sessile, 
8-12-flowered, 8-12 mm. high.—Ruopora xxxvii. 182 (1935). 
Laciniaria Smallii Britton, Man. 927 (1901). 

Southwestern Virginia to Georgia, chiefly in the mountains.— 
VIRGINIA. AUGUSTA Co.: Mt. Rogers, Elliot's Knob, Aug. 9, 
1893, A. A. Heller & E.G. Halbach 1179 (G, NY, P). Barn Co.: 
on more fertile slope, exposed shale barrens, along railroad eut 
just w. of tunnel, Millboro, Sept. 11, 1935, J. W. Adams & E. T. 
Wherry 2409 (G). Brprorp Co.: without stated locality, Sept. 
1, 1871, A. H. Curtiss (G (right plant), NY); Aug., A. H. Curtiss, 
(G); Aug. 30, 1872, A. H. Curtiss (ND). Roanoke Co.: dry 
woods on Chestnut Ridge, just s. of South Roanoke, Aug. 25, 
1942, C. E. Wood, 5298 (G). MowTrGoMEnv Co.: Blacksburg, 
July 30, 1895, W. H. Murrill (NY). GiuEs Co.: border of dry 
woods, Salt Pond Mt. (alt. 3800’), Aug. 4, 1937, J. M. Fogg, 
12942 (G); Salt Pond Mt., Aug. 1890, W. M. Canby (G). WYTHE 
Co.: Walker Mt., Sept. 5, 1899, ex Biltmore Herb., 4117h (NY); 
Wytheville, Lick Mt., Oct. 6, 1867, H. Shriver (G). Smytu Co.: 
Iron Mt., along Dickey Creek, Aug. 8, 1892, J. K. Small (G 
(NY, TYPE)); slopes & summits of Iron Mt., at Skull Gap, Aug. 
11-12, 1892, J. K. Small (US). WEST VIRGINIA. GREEN- 
BRIER Co.: Kate's Mt., Aug. 9, 1941, F. W. Hunnewell, 17466 


254 Rhodora [SEPTEMBER 


(G). Monroe Co.: Cove Creek, Sweet Springs, July 22, 1929, 
W. V. U. Biol. Exped. (G). NORTH CAROLINA. Yancey 
Co.: woods, Mt. Mitchell, Sept. 14, 1926, F. W. Hunnewell, 
16028 (G). HkNpERsoN Co.: Hendersonville, Sept. 18, 1898, 
Biltmore Herb., 4117d (NY); Flat Rock, Aug. 24, 1881, J. D. 
Smith (G). SOUTH CAROLINA. CursrER Co.: Chester 
State Park, Aug. 12, 1935, R. B. Mackintosh (G). GEORGIA. 
WILKEs Co.: without stated locality, Herb. J. A. Lowell (G). 
Var. ELEGANTULA (Greene) K. Sch. Stems slender, almost 
glabrous, 5-7 dm. high: leaves mostly linear; basal ones 10-25 
em. long, 3-4 mm. wide, glabrous, with broad ciliolate petioles, 
sometimes with woolly tufts on the petioles or scattered hairs on 
the under side: inflorescence a lax racemiform spike of sessile or 
slenderly pedicellate heads; rachis slender and somewhat virgate 
to stiffer and straight: heads more horizontal and campanulate in 
shape, of 8-12 flowers; phyllaries ovate to oblong with rounded 
less scarious tips; corolla-tube 6-7 mm. long; pappus 5-6 mm. 
long; mature achene 3 mm. long.—Just, Bot. Jahresb. xxix’. 569 
(1903). Laciniaria elegantula Greene, Pittonia iv. 316 (1901).— 
Georgia and Florida to Mississippi. —GEORGIA. Without 
stated locality: Boykin (G, NY, P). Lrserry Co.: near Sunbury, 
J. LeConte (NY). HasEnsHAM Co.: Tallulah Falls, 1846, T. C. 
Porter (P); dry hillsides, Tallulah Falls, Aug. 1899, A. Cuthbert 
(F); between Toccoa Falls & Tallulah Falls (alt. 1000'—1700^), 
Sept. 3, 1894, J. K. Small (NY). FrLovp Co.: Rome, Herb. Chap- 
man (US). Crankk Co.: dry oak woods, Athens, July 3, 1900, 
R. M. Harper 132 (G). GwiuxETT Co.: on Yellow River, near 
MeGuire's Mill (alt. 750’), Sept. 9, 1894, J. K. Small (NY). 
Ricumonp Co.: damp pineland, Augusta, Sept. 16, 1903, A. 
Cuthbert, 1004, 1004a (F); flat pineland, Augusta, Sept. 16, 1904, 
A. Cuthbert, 1004 (US); dry flat barrens, Augusta, Sept. 20, 1903, 
A. Cuthbert, 1004 (F). Warr Co.: Waycross, Sept. 17, 1909, 
W. W. Eggleston, 5099 (NY). MiırcaeLL Co.: Camilla, Sept. 20, 
1909, W. W. Eggleston, 5129 (G, NY, P, US). Lownprs Co.: 
dry woods, ca. 24% mis. e. of Valdosta, Sept. 8, 1902, R. M. 
Harper, 1614 (G, US). FLORIDA. Without stated locality: 
Chapman (G, NY, US), Leavenworth (G, NY). L&oN Co.: near 
Tallahassee, N. K. Berg (NY); Tallahassee, Oct. 10, 1914, R. M. 
Harper, 925 (G, NY, US). Liperty Co.: Aspalaga (Rwy. Sta. of 
Rock Bluff), Oct. 1892, Biltmore Herb., 4117b (G, US). ALA- 
BAMA. TaLLADpEGA Co.: open rocky woods, Chandler Springs, 
Talladega Mts., Sept. 20, 1892, C. Mohr (US); dry rocky hillsides, 
Oct. 1893, C. Mohr (US); rocky dry ridges, Chandler Springs, 
Sept. 1892, C. Mohr, la (NY); rocky dry ridges, Mts., Oct. 14, 
1893, C. Mohr, 1b (NY); on Talladega Mts., Sept. 1843, Rugel 
(NY). Jerrerson Co.: East Lake, Birmingham, Oct. 9, 1896, C. 
Schuchert (NY, US). TvscArLoosA Co.: e. of Tuscaloosa, Sept. 


1946] Galser,— The Genus Liatris 255 
23, 1933, R. M. Harper, 3121 (G, US); mixed woods, edge of pine 
barrens, 10 mis. n. e. of Tuscaloosa, Oct. 4, 1912, H. H. Bartlett, 
3337 (US). Ler Co.: Auburn, Oct. 18, 1896, F. S. Earle (ND, 
TYPE Of Laciniaria elegantula Greene); in dry open woods. sand 
or clay (common), Auburn, Sept. 28, 1899, F. S. Earle & E. S. 
Earle, 95 (G, NY, ND, US); Auburn, Oct. 9, 1898, F. S. Earle & 
C. F. Baker (NY, US), Sept. 18, 1897, F. S. Earle & C. F. Baker 
1343 (N Y), Sept. 20, 1896, C. F. Baker, 51 (NY), Oct. 10, 1896, 
C.F. Baker, 50 (NY), Sept. 1900, F. E. Lloyd & F. S. Earle (NY). 
MoNTGOMERY Co.: without stated locality, Sept. 21-23, 1909, 
W.W. Eggleston, 5141 (NY). Batpwin Co.: dry sandy pineland, 
Bay Minette, Sept. 6, 1912, F. W. Pennell, 4549 (P, US). MIS- 
SISSIPPI. Jackson Co.: Biloxi, Sept. 6, 1900, Tracy & Lloyd, 
560 (NY). 

Among the Walter specimens in the British Museum there is 
none labelled Anonymos graminifolia, but one labelled Chryso- 
coma affinis F. 309 (supposedly referring to Fraser) and with 
Nuttall's annotation Liatris in pencil, has been taken as un- 
doubtedly the plant described under the former name. Dr. H. 
K. Svenson kindly gave me notes and sketches made of the 
specimen when he was in London in 1938. These show the 
whole plant, including the basal corm, to be tall. His measure- 
ments of the inflorescence are “50 x 1.5 em." and his comment 
"narrow". The lower leaves are described as: “1 dm. x 2 mm. 
glabrous, except within the base which was not closely examined. 
The involucral bracts were rugose at the apex but not pubescent, 
shghtly ciliate at the margins". 

These careful observations, I believe, help us finally to recog- 
nize for Walter’s type the slender-stemmed, many- and very 
narrow-leaved plant found especially along the coastal plain of 
Virginia and the Carolinas that has passed under the general 
term Liatris graminifolia and which I have described as var. 
typica. Elliott (Sk. ii. 274 (1824)) seems to have had this variety 
in mind when he wrote his description of L. graminifolia. The 
species, as here defined, centers on Walter's specimen and de- 
scription. 

Barton (Mat. Med. ii. 223, t. 49 (1818)) states that after a 
thorough examination of specimens in the Muhlenberg Her- 
barium he could not satisfactorily refer a Liatris that he had, to 
any of the species described by Pursh, Michaux, or Willdenow 
and so named it L. dubia, clearly illustrating and carefully 


256 Rhodora [SEPTEMBER 


describing it. Although the specimen can unfortunately not be 
located at present in the herbarium of the Philadelphia Academy 
of Sciences, Barton’s plate and description including *'the 
striated stem covered with a sparse and hispid pubescence, 

the lower leaves longer and much wider than the upper . . . the 
upper leaves much smaller and linear, ciliated for the most part 
at the base", seems clear and directly applicable to many speci- 
mens in herbaria collected from back of the coastal plain, from 
New Jersey to Georgia and distributed under the name Liatris 
graminifolia. Gray (Man. ed. 1, 191 (1848)) first made the 
combination L. graminifolia var. dubia and this has been adopted 
here. 

As explained in the discussion of L. spicata (see no. 1) Clayton's 
plant from Virginia and Gronovius! description of it, which was 
cited by Linnaeus in the description of Serratula spicata and in 
turn by Willdenow in the description of Liatris spicata is a plant 
more commonly recognized now as L. graminifolia. "Torr. & 
Gray (Fl. N. Am. ii. 73 (1841)) and Gray (Synop. Fl. P. 111 
(1884)) had rightly excluded the Gronovian citation by Linnaeus 
from L. spicata. Gray included it in synonymy of L. gramini- 
folia, attributing that name to Pursh as the first to define the 
species correctly. Upon comparing the Barton plate with the 
Gronovius’ and Dillenius’ figures, it seems they are very similar, 
especially when allowing for the effect of cultivation, shown in 
the Dillenius’ figure, and considering Barton’s own comment: 
"this plant 1s one of a genus nearly all the species of which vary 
considerably, particularly in those marks usually supposed to 
be characteristic, as the sessile and pedicellated flowers". Fur- 
ther, as stated above, examination of a floret of the Clayton 
plant showed the corolla-tube to be pilose within and so are 
found to be the florets of the so-called graminifolia specimens. 
Clayton's plant may be cited, then, as by Gray, under L. gramini- 
folia and more correctly under var. dubia which 1s a large robust 
plant occurring abundantly in Virginia. While the two varieties, 
typica and dubia, overlap in the more eastern part of their range 
(as for example, two specimens of R. R. Tatnall from Delaware 
show, that of Sept. 23, 1934, no. 4053, from moist swale, L4 mi. 
e. of Ellendale, Sussex Co. (G) resembling typica and that of 
Sept. 16, 1934, no. 2385, from the edge of salt marsh, Fenwick Is., 


1946] Gaiser,— The Genus Liatris 257 


Sussex Co. (G) resembling dubia) no specimens of the former have 
come from the mountainous counties to the west. 

Still further confusion has been added by Aiton’s brief descrip- 
tion (Hort. Kew. ed. I. iii. 138 (1789)) of Serratula pilosa as “S. 
foliis linearibus pilosis, floribus axillaribus longe pedunculatis”’. 
Willdenow l. c. adopted the same description when transferring 
the species to Liatris. Although no Aiton specimen was found 
at the British Museum or Kew in the summer of 1939, there is 
fortunately a head of Aiton’s type “from the Banks Herbarium” 
at the Gray. From this specimen it can be learned that the head 
contains at least 10 flowers having a corolla ca. 8 mm. long and 
pappus 6 mm. long. The almost linear phyllaries are ea. 11 mm. 
long and 1-1.5 mm. wide and slightly narrowed and ciliate at the 
tip. The corolla-tube is also slightly pilose within. Also in the 
British Museum, Mr. Weatherby found a specimen from the 
herbarium of Goodenough, bearing, in the latter's handwriting, 
the label, “Serratula pilosa sp. nov. 1785". The photograph of 
this sheet shows a loose inflorescence, that by caleulations would 
measure about 3 dm. in height, bearing heads about 1 em. in 
length on leafy peduncles four to six times as long, giving the 
panicled appearance common in Liatris when grown in cultiva- 
tion. Two slenderer stems on the same sheet, each with one 
terminal head, may have been longer peduncles eut off lower 
down on the main stem. However, the size of the heads and 
character of the phyllaries are in agreement with those of Aiton's 
specimen. Unfortunately only the very narrow upper leaves are 
present on any of the stems and so the extent of their pilosity 
cannot be determined. Elliott (Sk. ii. 277 (1822)) added the note 
to his description of L. pilosa based on Willdenow, Pursh and 
Nuttall, “This variety is certainly not sufficiently hairy to 
have merited the trivial name which belongs to this species", 
and he included questioningly var. dubia Barton in synonymy. 
Support is given this opinion by the note of E. G. Baker found in 
the National Herbarium in connection with comparisons made 
of specimens sent him. He states of the type “Serratula pilosa 
Aiton, Hort. Kew. ed. 1. iii. p. 138. (1) The heads are axillary 
and rather long-peduncled. (2) The heads are rather larger. 
(3) The plant is pilose. (4) The involucral bracts are narrow 
and subacute. (5) Lodd. Cab. t. 356 is a figure of the true plant. 


258 Rhodora [SEPTEMBER 


The plant is not so thickly pilose as the figure. (6) The true 
plant seems inseparable from the plant figured in Barton's Med. 
Bot. i. t. 49 as L. dubia," Until the Aiton specimen may be 
again examined it seems to this author also that 1t would be best 
to include it in such synonymy. 

Torr. & Gray (Fl. N. Am. ii. 74 (1841)) described Liatris 
pilosa from the Aiton specimen and one of Mr. Read's collection 
from Seven Mile Mountain, Va., now in the Philadelphia Her- 
barium. However, later in Synop. Fl. ?. 111 (1884) it was made 
synonymous with Liatris graminifolia var. dubia and referred to 
as a variety with many large heads with the range “sandy pine 
barrens from New Jersey to Florida and Alabama". Examina- 
tion of the Read specimen (P), showed a plant with quite broadly 
lanceolate basal leaves, a spike 18.5 em. long with 9 distant 
larger heads ca. 1.5 em. long and 1.5 em. wide, and phyllaries 
with a narrow scarious margin. Of the specimen Torr. & Gray 
l. c. had written: ‘Plant nearly as stout as L. scariosa", and to 
the writer too it seems distinctive but more closely related to 
plants found more abundantly in the mountains, that are here 
described as L. turgida. (see no. 9). 

The really pubescent or hirsute variety of this species has been 
described by Fernald and Griscom from the northern part of its 
range as Liatris graminifolia Willd. var. lasia. The plate of L. 
pilosa (Loddiges Cab. t. 356 (1819)) shows a very pubescent 
plant, which could have belonged to this variety. 

With this species has been included a slenderer-stemmed plant, 
with long basal leaves, typified by a specimen of J. K. Small, 
Aug. 8, 1892, from Iron Mountain, Smyth Co., Va. (NY) and 
described as Laciniaria Smallii Britton (Man. 927 (1901)). 
Fernald and Griscom, |. c., finding a match for this in their 
specimen of Sept. 25, 1933, no. 2907 from the desert at Cape 
Henry, Princess Anne Co., Va. (G) reduced it to a variety of 
Liatris graminifolia, since it could not be separated from that 
species by any satisfactory characters. Comparison of speci- 
mens show that much of this species coming from the mountain 
regions of south-west Virginia and of adjoining states bear a 
resemblance to Small's plant from Smyth Co., in the loose 
infloresencec of sessile, 8-12-flowered heads beyond which the 
bract-like leaves do not extend, and the generally fewer leaves 


1946] Gaiser,—The Genus Liatris 259 


(basal ones being longer and wider) with scattered hairs on the 
under surface. Some specimens are, however, less slender and 
have a more spike-like inflorescence with heads of appressed 
almost orbicular phyllaries which on herbarium sheets have been 
questioningly referred to L. pilosa Ait. Until Aiton’s specimen 
may have been found, the evidence given from the single head at 
Gray and a photograph from the British Museum of Goode- 
nough’s specimen match var. dubia better than this variety from 
the mountains. Since the specimen of Read, from Seven Mile 
Mountain, Va. (P) referred to by Torr. and Gray, Fl. N. Am. ii. 
74 (1841) in their discussion of L. pilosa, seems to bear closer re- 
semblance to specimens as of Aug. 30, 1912, E. S. Steele, no. 24, 
from Afton, in the Blue Ridge Mountains, Nelson Co., Va. (US), 
it has been included under the new species L. turgida. Though 
L. Helleri Porter is similarly found limited to a small section of 
the Appalachian Mountains it seems to represent a different 
segregate and is distinguished from this species by the short 
pappus, the usually quite glabrous leaves and the few closely 
arranged heads. 

Greene (Pittonia, iv. 316 (1901)) described a species similar to 
Small’s but taller and of more southern range, from a specimen 
of Oct. 18, 1896, F. S. Earle, from Auburn, Lee Co., Ala. (ND) 
which he called Laciniaria elegantula. By the turbinate heads 
and narrowly lanceolate leaves with ciliate petioles it seems to 
belong to the graminifolia alliance, as here interpreted, rather 
than to the Scariosae as suggested in his description. After 
careful comparison of this plant with that of Small from Smyth 
Co., Va. and examination of a number of specimens in the 
herbaria it seems that one form passes into the other. Yet in 
the southern part of its range the taller and somewhat stiffer 
elegantula seems to become even stiffer so that it is still more 
difficult to make sharp delimitation of the most southern Florida 
material from the Greene type from Alabama. Thus, rather 
than broaden the conception of L. Smallii it has seemed more 
helpful to split off the more northern, short material as var. 
Smallii from the southern, taller plants, var. elegantula, and 
include therein the stiffer Florida ones. However, that these 
Florida specimens may be wider variants is again suggested by 
possible intermediates between this variety and L. gracilis, as 


260 Rhodora [SEPTEMBER 


in the specimen of Oct. 10, 1914, R. M. Harper, no. 225, 2 miles 
e. of Tallahassee, Leon Co., Fla. (G, NY). 

L. graminifolia is by far the largest species of the Graminifoliae 
series and out of it have probably arisen the more localized segre- 
gates that follow. 


8. Liarris HELLERI Porter. Rootstock in older plants large 
and shallow, usually 5-6 em. across and 2—4 cm. deep, giving rise 
to many radical glabrous linear-lanceolate leaves 2-3 dm. long and 
6 mm.-1 em. wide at the middle, diminishing to long, winged, 
non-ciliate petioles; cauline leaves diminishing uniformly upward 
in short-pyramidal form to bracts less than the length of the 
flower-heads; only a few scattered cilia at base of leaves or along 
veins: inflorescence-bearing stems 1 or 2, rarely more than 2 dm. 
high, with heads occurring closely and covering one third the 
length; heads turbinate at the time of flowering, 12-15 mm. high, 
7—10-flowered ; phyllaries oblong-ovate, with narrow scarious rim 
and beyond it a finely ciliolate margin; corolla purple, 5-7 mm. 
in length, slightly pilose within the tube; pappus scanty, barbel- 
late, only 2.5-4 mm. long, i. e. half the length of the corolla and 
hardly showing beyond the involuere at the time of flowering: 
achene 2.5-3.5mm. long.— Bull. Torr. Bot. Club, xviii. 147 
(1891). Lacinaria Helleri Porter, Muhlenbergia i. 6 (1900). 

Has been found only on the mountains of North Carolina.— 
NORTH CAROLINA. WaraAvGA Co.: Blowing Rock Mt., 
Aug. 18, 1890, A. A. Heller, 81 (NY, type (ND, US)), Aug. 18, 
1890, A. A. Heller, 82 (NY); Blowing Rock, Aug. 17, 1891, A. B. 
Seymour, 22 (G), Aug. 1928, W. C. Ferguson, 18 (NY); dry soil, 
exposed rocky ledge, Blowing Rock, Aug. 1, 1922, L. F. Randolph 
& F. R. Randolph, 1161 (G). CALDWELL Co.: ledges of Blowing 
Rock (elev. 4200’), Aug. 6, 1891, J. K. Small & A. A. Heller, 344 
(G, NY, P, Q, O, US, F (without no.)); e. of Blowing Rock, 
(3500-4000), Aug. 24, 1893, A. A. Heller, 1236 (NY, US); 
summit of Grandfather Mt., Sept. 25, 1898, W. M. Canby, C. S. 
Sargent & J. Muir, 70 (US); Grandfather Mt., Aug. 9, 1890, G. B. 
Sudworth, 95 (US); peaks of Grandfather Mt., Linville Station, 
Aug. 11, 1890, G. B. Sudworth, 115 (G, US). Avery Co.: high 
rocks, s. end Beech & Grandfather Mts. (alt. 1650 m.), Sept. 1-2, 
1915, E. S. Steele, 89 (G, NY). Burge Co.: Table Rock Mt., 
Aug. 2, 1890, A. A. Heller, 81 (NY), 82 (P). MīircaeLL Co.: 
mountain meadows, Roan Mt., July 10, 1894, C. Mohr (US). 


This species, of a very limited mountain range, seems distinc- 
tive in its very short pappus since others having both plains- and 
mountain-habitats have not shown this variation. Like the type, 
from Blowing Rock Mountain, all other specimens seen are short, 
indicative of growth only on exposed barren ledges. 


1946] Gaiser,—The Genus Liatris 261 


It is to be distinguished from varieties typica and dubia of L. 
graminifolia by these characters and from the shorter var. 
Smallit, which occurs in neighboring mountains of Virginia, by 
the slender stems bearing few distant heads. L. turgida of the 
same series, has a more robust, elongate inflorescence. 

9. Liarris turgida, sp. nov., cormo globoso (juniore) vel 
(vetustiore) irregular! compresso 4-5 cm. lato ad 4-5 caules 
emittente; caulibus saepe singulis suberassis saepe rubescentibus 
nec striatis 6-9 dm. altis glabris vel sparse vel dense pilosis pilis 
longis albis curvatis; foliis vix numerosis lineari-lanceolatis 
pagina inferiore et ad basem sparse vel subcopiose pilosis, 
basalibus plerumque quam apud L. spicatam vel L. gramini- 
foliam longioribus latioribusque saepe 10-15 em. longis 1 em. 
latis ad basem et ad apicem aequaliter angustatis petiolo alato, 
superioribus epetiolatis angustioribus fere linearibus ad bracteas 
quam capitula breviores reductis; spica simplice racemiformi; 
capitulis inter se 1-2 cm. distantibus sessilibus vel breviter 
pedicellatis 9-20-floris anthesi subturbinatis 1-1.5 cm. altis et 
latis; phyllariis plerumque purpureis adpressis glabris marginibus 
anguste scariosis vel ciliolatis, exterioribus ovatis aliquando fere 
orbiculatis, interioribus oblongis vel linearibus obtusis ea. 12 mm. 
longis; corollis purpureis 8-9 mm. longis, tubo intus piloso; 
achaeniis 4-5 mm. longis, pappo 6-7 mm. longo.— Mountain 
ridges of southwestern Virginia and adjoining West Virginia and 
North Carolina. 

Rocky woods, road to Royal Orchard, vicinity of Afton in the 
Blue Ridge Mts., alt. 600 m., Nelson Co., Virginia, Aug. 31, 1912, 
E. S. Steele 24 (US 643319, TYPE). 

VIRGINIA. Pace Co.: vicinity of Stony Man Mt. (alt. 
3600^), Blue Ridge near Luray, Sept. 3, 1901, E. S. Steele & Mrs. 
Steele, 241 (G, NY); vicinity of Stony Man Mt. (alt. 3500’), 
Blue Ridge near Luray, Aug. 28, 1901, E. S. Steele & Mrs. Steele, 
241 (NY, US); Honzman, Aug. 1891, Miss L. Smith (US); 
Hawksbill Mt., Aug. 30, 1891, W. Palmer, W. H. King (US). 
RocKINGHAM Co.: vicinity of Elkton, foot and slopes of Blue 
Ridge Mts., 1918, E. S. Steele (G).  AvGvsTA Co.: slopes near 
Jennings Gap, vicinity of Stribling Springs (alt. 570 m.), Aug. 
24, 1917, E. S. Steele, 23 (G); Little North Mt., vic. of Augusta 
Springs, Aug. 29, 1908, E. S. Steele (US 785731, -3, -4, -5); 
Elliott's Knob (1200 m.), Aug. 27, 1913, FE. S. Steele, 56, 72 (US); 
spur of Elliott’s Knob, vicinity of Augusta Springs, Aug. 29, 
1908, E. S. Steele (US 785727), Sept. 8, 1908 (US 785728, -9); 
vic. of Fordwick and Craigsville, 1913, E. S. Steele (G); Elliott's 
Knob, summit (alt. 1320 m.), vic. of Augusta Springs, Sept. 2, 
1908, E. S. Steele (US 785730); Elliott’s Knob (alt. 1200 m.), vic. 
of Augusta Springs, Sept. 2, 1908, E. S. Steele (US 785732). BATH 


262 Rhodora [SEPTEMBER 


Co.: vic. of Millboro (alt. 480 m.), Aug. 26, 1907, E. S. Steele (P); 
vic. of Millboro, Panther Gap, Sept. 6, 1907, E. S. Steele (US 
643315); railroad, vic. of Millboro, Aug. 21, 1907, E. S. Steele 
(US 643317); vic. of Millboro (alt. 510 m.) Aug. 29, 1906, E. S. 
Steele (US 590180). NeLrson Co.: rocky woods, road to Royal 
Orchard, vic. of Afton, in the Blue Ridge Mts. (alt. 600 m.), Aug. 
31, 1912, E. S. Steele, 24 (G, US 643318, -20, -22, -23) ; Afton, road 
to Royal Orchard, Sept. 19, 1909, W. S. McAtee, 1243, 1245 (US); 
high cliff, toward Humpback Mt., vic. of Afton, Sept. 13, 1912, 
E. S. Steele, 121 (US). RocknnipaE Co.: North Mt., near Lex- 
ington, Aug. 26, 1924, J. R. Churchill (G); vicinity of Goshen, alt. 
450 m., Sept. 4, 1904, E. S. Steele (US 494378, -80); mts., e. of 
Natural Bridge, Sept. 13, 1907, E. B. Bartram & B. Long (US). 
Bxpronp Co.: without stated locality, Sept. 1, 1871, A. H. Curtiss 
(G, left plant). Crara Co.: (and MoNnokE and ALLEGHANY, 
W. Va.) Co.: alt. 600 m., Aug. 22, 1903, E. S. Steele & Mrs. 
Steele, 86 (G, NY, US), Aug. 30, 1903, E. S. Steele & Mrs. Steele, 
166 (G, NY, US); Johns Creek Mt., Aug. 26, 1903, E. S. Steele 
(US); Potts Mt., Aug. 28, 1903, E. S. Steele (US). MONTGOMERY 
Co.: Blacksburg, July 20, 1895, W. H. Murrill (NY). Gres Co.: 
Brush Mt., 2 mis. e. of Newport, Aug. 30, 1933, E. J. Alexander, 
T. H. Everett & S. D. Pearson (NY); Salt Pond Mt., Bald Knob, 
Aug. 25, 1899, C. L. Pollard & W. R. Mazon, 69 (NY, US). 
WEST VIRGINIA. GREENBRIER Co.: dry upland, 1 mi. s. of 
White Sulphur Springs, Aug. 10, 1922, L. F. Randolph and F. R. 
Randolph, 1281 (G); White Sulphur Springs, July 16, 1892, A. 
Brown (NY); White Sulphur Springs, July 31, 1877, G. Gutten- 
berg (NY); dry woods, White Sulphur Springs, Aug. 27, 1903, K. 
K. Mackenzie, 359 (NY, US). Monror Co.: Peters Mt., vic. 
of Old Sweet Springs, Sept. 9, 1905, E. S. Steele (US). NORTH 
CAROLINA. Burxe Co.: dry woods, Jones Ridge, July 22, 
1933, F. W. Hunnewell, 12982 (G). BuNncomBE Co.: rocky 
roadside banks, near Black Mt., Aug. 23, 1927,!K. M. Wiegand & 
W. E. Manning, 3176 (G). 

It is a pleasure to adopt the specific epithet found on many of 
the National Herbarium sheets of Mr. E. S. Steele’s specimens 
under another generic name, since his rich collection from the 
mountains of Virginia has, in our mind, established the identity 
of the species. 

While the species here described may resemble L. graminifolia 
var. dubia in shape and distribution of the heads and L. spicata 
in the appressed ovate phyllaries, it cannot be merely referred to 
as an intermediate between the two. It is to be distinguished 
from the former by its rather stout stem, the loose raceme-like 
arrangement of heads showing somewhat appressed ovate, orbieu- 


1946] Blewitt,—Edgar Burton Harger 263 


lar or oblong and obtuse-tipped phyllaries, rather than loose, 
narrowly lanceolate, ciliolate-margined ones, and by lanceolate 
leaves having scattered hairs over one or both surfaces but lacking 
the marked basal cilia along a winged petiole. It resembles the 
variety Smalli of L. graminifolia most in the nature of the spike, 
of rather distant sessile heads beyond which the upper bract-like 
leaves do not extend, but it differs in having a stouter and stiffer 
rachis, usually with more heads and more orbicular appressed 
bracts, and in having leaves frequently covered with hairs, a 
character in which it also differs markedly from L. spicata. 

There are three specimens (US 643318, -19, -20) of collection 
no. 24 of E. S. Steele, Aug. 31, 1912, from the vicinity of Afton, 
Nelson County, and these show variation from an almost glabrous 
condition to an abundance of scattered hairs; of these, the inter- 
mediate condition was chosen as the type, though the description 
allows for the inclusion of all. Collected only from the mountain 
ridges of Virginia and the neighboring ones of West Virginia 
and North Carolina it has frequently been referred to as L. pilosa 
Ait. on herbarium sheets but until Aiton's specimen may have 
been found (see no. 7) the evidence gathered makes that name 
seem inapplicable to this plant. However, the one specimen of 
Read, from Seven Mile Mountain, Va. (P) referred to by Torr. & 
Gray Fl. N. Am. ii. 74 (1841) in their discussion of L. pilosa 
seems to have a closer resemblance and it is here included. 
Though L. Helleri Porter is similarly found limited to a small 
section of the mountains, it seems to represent a different seg- 
regate, distinguished from this species by the short pappus, the 
few, though closely placed, heads, and usually quite glabrous 
leaves. 


(To be continued) 


EpGar Burton HAarGer.—In the passing of Mr. Harger, the 
Connecticut Botanical Society has lost one of its most active and 
gifted members. He had a large part in the organization of the 
Society in 1903, became a charter member, and at the first meet- 
ing was elected corresponding secretary, an office in which he 
served with zeal and distinction for twenty-three years. On the 
death of Charles H. Bissell, he was elected president and served 


264 Rhodora [SEPTEMBER 


for nineteen years. In 1945, because of ill health, he was made 
honorary president. 

Harger was born in Oxford, Connecticut, February 5, 1867, 
lived for most of his life in the Quaker Farms section of that town, 
and died there February 22, 1946. He was a graduate of the 
Sheffield Scientific School of Yale University in the class of 1887. 
He took up the profession of surveyor and served his native town 
and its vicinity in that capacity for many years. He had been 
Judge of the Probate Court, Town Clerk and representative in 
the state legislature; he had also served on the Board of Education 
in Oxford and as treasurer and clerk of the Oxford Congregational 
Church, of which he was a life-long member, In 1903, he married 
Olive Platt, who died in 1926. Four children survive them. 

Harger was the author of a number of articles in Rhodora. As 
a particularly keen and active collector and amateur taxonomist, 
a member of the committee who compiled the Catalogue of 
Flowering Plants and Ferns of Connecticut (1910) and the Addi- 
tions to it (1931), he made a substantial contribution to the 
advancement of botanical knowledge in the northeastern United 
States. He was ever ready to share with all comers his extensive 
knowledge of the local flora and of natural history in general.— 
ARTHUR E. BrEwiTT, Waterbury, Connecticut. 


Volume 48, no. 572, including pages 165—200 and plates 1047 and 1048, was 
issued 5 August, 1946. 


OCT 25 1946 


Hovora 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL | Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. October, 1946. No. 574. 
CONTENTS: 
Frère Marie-Victorin. James Kucymiak. ............0002. sess 265 
Dicentra eximia in Vermont. Richard J. Eaton. .............. 272 
The Genus Liatris. L. O. Gaiser (continued). ................ 273 
Second Station for Corydalis flavula in Connecticut. 
Frederick W. Kilbourne. ........... eene 326 
Does Bartonia verna grow in Virginia? M. L. Fernald. ..... 327 


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Rbodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. October, 1946. No. 574. 


FRERE MARIE-VICTORIN* 
(1885-1944) 
JAMES KUCYNIAK 


In the passing of Frére Marie-Victorin, the inhabitants of 
French Canada mourned the loss of one of their noblest sons and 
foremost contributors to the development of Science among them. 
Much has appeared in local bulletins and newspapers, notably in 
those of the French language. A few papers only, perhaps be- 
cause little was known outside of Quebec of the biographical de- 
tails of his life, have appeared in the far more numerous scientific 
periodicals in the United States. This paper is presented in the 
hope of filling to a certain extent this want. 

Frére Marie-Victorin was born in Kingsey Falls, April 3, 1885. 
Christened Joseph-Louis-Conrad, he was one of the family of 11 
of Conrad Kirouac and Philomène Luneau. He was the only 
one of six sons to reach manhood. 

Soon after his birth, the family left Kingsey Falls and moved 
to Quebec City where his father opened up a profitable grain and 
flour business. It was here that Joseph-Louis-Conrad spent his 
early boyhood. He entered the grammar school of St. Sauveur 
Parish. Heading his class, he was awarded a three year scholar- 
ship which enabled him to continue his schooling under the re- 
ligious order of the Brothers of the Christian Schools. 

He soon sensed that his vocation in life was teaching and, to 
realize it more fully, decided to become à member of the Order 
which taught him. 


* Paper read as “The life and work of Frère Marie-Victorin'' at the 50th annual 
meeting of the Michigan Academy of Science, Arts and Letters, in Ann Arbor, on 
April 12, 1946. 


266 Rhodora [OcroBER 


At the age of sixteen, upon completing his studies at the 
Académie Commerciale, he entered Mont-de-la-Salle, the Order’s 
motherhouse, in Montreal, which was then located where the 
Montreal Botanical Garden at present stands. 

It was upon being accepted into the community, that he chose 
the name Frére Marie-Victorin. 

In 1903, he began his career as a teacher in the Collége de St- 
Jéróme, in a city some thirty miles from Montreal, in the foothills 
of the Laurentians. This was momentarily interrupted by the 
disclosure of pulmonary tuberculosis. Under doctor’s orders, he 
had to give up teaching and spend as much time as possible out- 
doors. So it was at twenty that France Bastien found him one 
day, pensively seated on a moss-covered boulder, with a flora in 
one hand and trying to tag a name on a broad-leaved Liliacea. 
He gave him a clue by telling the future botanist that it was the 
*ail-douce", the name under which the French-Canadian habi- 
tant knows the dog’s-tooth-violet. This bit of information was 
to start him definitely on his botanical career. 

A year later, his condition haviig improved sufficiently to en- 
able him to return to classes, he was posted to St. Leo's High 
School, in Westmount, a wealthy suburb of Montreal. That 
same year, he made the acquaintance of Frére Rolland-Germain 
who only a little later was to become the faithful companion on 
the many field trips which were to be undertaken in the years 
that lay ahead. 

His stay at St. Leo's lasted four years after which he was trans- 
ferred to the Collége de Longueuil, across the St. Lawrence from 
Montreal. Here he taught algebra, geometry and French com- 
position, but devoted most of his spare time to botany and the 
organizing of botanical outings with students. 

In spite of a heavy working-schedule, he continued making 
worthy contributions to more advanced botany. He had pub- 
lished some 39 papers before being signalled out to create a 
department of botany in the now independent Université de 
Montréal, which had just severed from being a Montreal affiliate 
of Quebec City's Université de Laval. He was reluctant to leave 
the Collége de Longueuil and, up until 1928, remained a part-time 
professor there. However, as the youthful and vigorous Labora- 
toire de Botanique began to make increasing demands on his 


1946] Kucyniak,—Frére Marie-Victorin 267 


time, he was obliged to give up his teaching assignment at Lon- 
gueuil and devote full time to the Université. 

Taken up with the preparation of lectures and mapping out of 
"lab" sessions, the selecting and forming of assistants who were 
to remain with him and carry on after he was gone, left little time 
for publishing. It was fully two years before any paper of tech- 
nical importance appeared: the first number of the ‘‘Contribu- 
tions du Laboratoire de botanique", now known as the ‘‘Contri- 
butions de l'Institut botanique de l'Université de Montréal" and 
which consist of original memoirs or reprints from various peri- 
odicals, principally the Mémoires de la Société Royale du Canada 
and Le Naturaliste Canadien. For his work on the *'Filicinées 
du Québec", the Université de Montréal awarded him a Docteur 
és Sciences. This was the second in the series of Contributions, 
and the first in a contemplated set of fascicles of a vast under- 
taking which would eventually form a complete and critical study 
of the flora of Quebec. Two years later, a similar paper appeared 
on clubmosses and in 1927 an issue each was devoted respectively 
to horsetails and gymnosperms. After another lapse of two 
years, a study on the Liliiflorae was published and this was fol- 
lowed in as many years by his treatment of the Araceae. Fully 
cognizant of the fact that, some seventeen years earlier, he had 
himself made a vibrant appeal for the urgent publication of a new 
illustrated flora of the Province of Quebec, he realized that at 
the rate the fascicles appeared it would be a good many years 
before an indispensable flora of the kind was completed. He 
therefore interrupted the series and started work upon what at 
first was eyed as a less ambitious project. This was to be a 
manual which, from the outset, was affectionately termed the 
“petite flore", in which illustrations, a description of and inter- 
esting notes on the species found within the inhabited regions of 
Quebec were to be treated. The result was La Flore laurentienne, 
a quarto of some 927 pages in which 1917 species of flowering 
plants and vascular cryptogams were described and illustrated, 
preceded by a phytogeographical study of the region in which a 
geobotanical foundation of Quebec was originally outlined. An- 
other innovation was the introduction of chromosome numbers 
for the first time in a flora. Taking time out to write the Flore 


268 Rhodora [OcroBER 


did not put an end to the Contributions which during his lifetime 
reached 51 in number, while others have appeared subsequently. 

On June 10, 1923, together with several other enthusiastic field- 
naturalists, he founded the Société Canadienne d'Histoire Na- 
turelle which, in its unassuming way, has contributed much to 
the diffusion of scientific knowledge through French-Canada. 
For years, he was its president. The society’s annual activities 
were brought to a close in a meeting devoted to what was termed 
the presidential address. This served as an excellent platform 
for the setting in motion of certain reforms or giving birth to 
many long-needed institutions. One can't help but admire the 
courage that it took to make as frank and unflattering an inven- 
tory of the shortcomings and of the attitude of French-Canadians 
towards pure science as is found in “La Science et nous" which 
Frére Marie-Victorin delivered in 1926. Not content to relate 
this sad plight of affairs, a remedy to the situation was proposed 
in advice given to educationalists as well as in the selection of 
studies which should constitute the curriculum in a faculty of 
science. These are the basic ideas of the 1930 address entitled 
"Les Sciences naturelles dans l'Enseignement supérieur". Five 
years later, with “La tâche des Naturalistes canadiens-frangais’’, 
an attempt was made to plot a future course offering new vistas 
in which research could be undertaken after having taken into 
full account the intellectual capacities and available resources. 
“La Science et notre Vie nationale" in 1938 ended this series with 
an examination of what had been done to date and reiterating a 
vigorous appeal in favor of what was deemed necessary for the 
future. 

In its turn, the Société fostered the idea of young naturalists' 
clubs to encourage all schoolchildren to take a closer interest in 
the study of nature. Frére Marie-Victorin followed its activities 
with the deepest interest. The countless articles he has written 
for their column, which appears weekly in the local French- 
Canadian newspaper *Le Devoir", or the leaflets of popular in- 
formation known as the “Tracts des C. J. N.”, issued at irregular 
intervals by the Société, will testify to that effect. The Cercles 
des Jeunes Naturalistes, as these clubs are known, have under 
his watchful eye grown increasingly and to-day reach almost a 
thousand in number, listing some 30,000 members in all, to whom 


1946] Kucyniak,—Frére Marie-Victorin 269 


half a million of the leaflets mentioned earlier are distributed 
annually. To further the satisfactory work carried out already, 
a more widespread development was inaugurated in the produc- 
tion of a program of radio broadcasts, known as “La Cité des 
Plantes", broadcast over a network of 13 Quebec radio stations. 
This consists of 26 weekly, fifteen-minute chats devoted to intro- 
ducing students the province over to the striking aspects of the 
plant world. 

Time unfortunately permits but mention here of the “Ecole de 
l'Eveil", where some 40 youngsters ranging in age from three to 
five, meet for an hour each week in a kindergarten of natural 
history. 

Hardly one to encourage the French-speaking workers in 
science to carry on in the isolated spheres of their respective 
fields, the secretary of the newly-formed Société, together with 
Léo Pariseau, Edouard Montpetit and Louis Dalbis founded 
the ACFAS, the French-Canadian equivalent of the A.A.A.8. 
Breaking away from an attempt to make it an affiliate of the 
Association Frangaise pour l'Avancement des Sciences, it has 
survived well on its own. "Though connected with the group in 
an official capacity over two relatively short periods, being its 
secretary for the 1924-1925 term and its president in 1937-38, he 
had nonetheless, in an unofficial capacity, contributed consider- 
ably to its progress. 

In 1929, having just returned from a lengthy trip through 
Africa, the Middle East and Europe, and highly impressed with 
the work being carried out in the Cape of Good Hope, Cairo, the 
Canary Islands and Cologne, Frére Marie-Victorin was firmly 
convinced of the necessity of a botanical garden for a city as large 
as Montreal. On the 14th of December 1929, “Le jardin botan- 
ique de Montréal" was a plea made before the members of the 
Société which, within six years, was well on its way to full realiza- 
tion. Under the pressure of a vigorous campaign backing the 
idea, the municipality's Director of Departments called upon 
Frére Marie-Victorin, fifteen years ago last April 8, to draft the 
initial project for a botanical garden. It took a little over a 
month to turn in the requested report. Within a year, an origi- 
nal sum was voted and preliminary work, such as draining and 
soil-leveling, was earried out on what was then known as Maison- 


270 Rhodora [OcroBER 


neuve Park. A greenhouse was erected as well as a small pa- 
vilion for administrational purposes. Definite realization of the 
project was begun in earnest with the creation on April 24, 1936, 
of the Municipal Commission of the Montreal Botanical Garden 
with Frére Marie-Victorin as its director. Intelligent planning 
and many arduous hours were put into this undertaking which, 
with the generous collaboration of the federal, provincial and 
municipal authorities, accomplished much within a short period 
of three years: the administration building with its offices, lecture 
rooms, auditorium seating some 500 people, "labs," library, her- 
barium and museum, the 22 service or propagation greenhouses, 
the gardens of ornamental plants and of economie plants, to 
mention but a few. 

The outbreak of war in 1939, together with a change in the 
provincial government, brought the intense activity centred on 
the building of the Garden to practically a standstill. Frère 
Marie-Victorin lived, however, to see almost three-quarters of 
the project realized and was ever confident that its completion 
was only a matter of time. 

One would think that, with the heavy duties imposed upon 
him by the founding of a department of botany, a task which in 
itself obliges one to keep a closer eye upon the rapid develop- 
ments occurring therein, would have constantly kept Frére 
Marie-Victorin the year round within easy reach of the prospering 
department of botany. If, however, in Eastern Canada, the 
Institut is to-day considered first and foremost in the field of 
taxonomy and phytogeography, it is thanks to the enthusiastic 
initiative which its founder put into field trips, a practice which, 
no matter how numerous the obligations nor how urgent the 
affairs of office, was never relayed to second place when the season 
for botanizing was in full swing. These surveys were carried out 
on quite a heavy scale for almost a quarter of a century. Though 
the Université was in no position financially to back all of them, 
the expenses were at times defrayed by those taking part them- 
selves and, for good measure, by grants from the National Re- 
search Council, the Federal Ministry of Mines, the Provincial 
Ministry of Game and Fisheries and the Quebec Office des 
Recherches. The first region to be explored botanically was the 
district of Lake St. John to which the summers of 1920 and 1921 


1946] Kucyniak,—Frére Marie-Victorin 271 


were devoted, the region itself being subsequently re-visited in 
1932, 1936 and 1937. Two summers later, a trip was made into 
the heart of the Gaspé peninsula where climbing one of the steep 
slopes in the wilderness proved too strenuous a physical effort for 
Frère Marie-Victorin. A severe heart-attack did not put an end 
to his taking part in further botanical excursions but limited 
them to those places which required but little bodily exertion to 
cover. The next five seasons were devoted to intense field work 
in the Mingan Archipelago as well as Anticosti Island. Several 
species new to science, Cirsium minganense, Botrychium minga- 
nense, Scirpus Rollandii, Solidago Victorinii, Aster anticostensis, 
were among the rewarding yields of this geologically as well as 
biologically interesting area. From 1930 on, the field trips were 
no longer of lengthy duration, seldom lasting more than a fort- 
night. Two successive seasons were devoted to combing the 
Baie des Chaleurs region and adjoining New Brunwsick. Initial 
work in Ontario was begun in 1932 and was centred mainly about 
the Great Lakes and the Georgian Bay which were re-visited in 
the summers of 1936 and 1937. In 1933, the upper Ottawa and 
the Abitibi districts were covered. During the course of a rather 
long season in 1936, several collecting trips were made to the 
little-known though highly-promising terrain of the Eastern 
Townships. A four-months sojourn in Cuba, in the winter of 
1938-1939, aided and abetted by Hermano Léon, resulted in the 
publication of an illustrated 400-page memoir, devoted entirely 
to the field trips carried out there. A second volume appeared 
covering subsequent visits to this island, while a third was in 
preparation before his untimely end. Reams of unpublished 
notes taken in minute detail on each herborization are housed in 
the archives of the Institut Botanique. 

When not undertaking major field trips, Frére Marie-Victorin, 
in company with the faithful co-worker on so many of them, to- 
gether with a lab assistant or two, would spend an afternoon, 
sometimes a day or even a long week-end botanizing within easy 
reach of Montreal, visiting such localities as the shores of the St. 
Lawrence near Longueuil; St. Jéróme, noted for its boggy lakes; 
Oka with its sand dunes and stands of pine and sweet-fern; St. 
Janvier, Farnham and Lanoraie with their interesting stretches 
of peat-bog; Cap-Rouge with its fresh-water estuary and charac- 


272 Rhodora [OCTOBER 


teristic flora of unique items, such as Gentiana Victorinii and 
Cicuta Victorinii; Ancienne-Lorette where the pleasure of botaniz- 
ing was coupled with the nostalgia of many agreeable boyhood 
memories and countless other places perhaps just as interesting 
botanically but far too numerous to enumerate here. 

For some time, Frère Marie-Victorin had been hoping to pay 
a visit to Black Lake where E. T. Wherry, after the geologist 
Harvie, had reported the rare serpentinicolous Oregon cliff-brake 
(Cheilanthes siliquosa). This station is one of the only three 
known in Eastern America, the others being Mt. Albert in Gaspé 
and Owen Sound, Ontario. So, early on the Saturday morning 
of July 15th, with Frère Rolland-Germain and three others, 
Frère Marie-Victorin set out for the station in the asbestos 
centre of the Province, a little over a hundred miles from Mon- 
treal. Stopping to botanize at several interesting localities on 
the way, it was well after lunch when their destination was 
reached and time to think of the return journey now that several 
stands of this exceedingly exacting fern had been found. Home- 
ward-bound, a brief stop was made at St. Norbert where Frère 
Marie-Victorin dropped in to chat with an old couple of relatives 
and call on a childhood friend in the village where he had spent 
so many summers as a youngster. When more than halfway on 
the way home, at Ste.Rosalie, some two or at most three miles 
east of St. Hyacinthe, a car coming out of the latter city crashed 
head-on into the party of botanists. The shock of the collision 
was too violent for Frère Marie-Victorin’s heart. After an un- 
successful attempt to administer a dose of the ever present 
"eoramine" which he always carried with him, he passed away 
by the roadside before efficient medical assistance could be 
administered. 


Montreal Botanical Garden 


DicENTRA EXIMIA IN VERMONT.—A single plant of Dicentra 
eximia (Ker.) Torr. was found on the western outskirts of 
Brattleboro, Vermont, by Dr. Somers H. Sturgis of Cambridge 
on May 10, 1946. It was growing in partial shade between a 
country road and a mountain brook on a high bank composed of 
mixed soils which appeared to have been deposited by freshets. 
Unfortunately, Dr. Sturgis dug the plant for his wild garden 


1946] Gaiser,—The Genus Liatris 273 


without realizing its interest as a rare adventitious species in 
New England. I revisited the station later but was unable to 
find any other specimens. Diagnostic fragments of the plant 
have been deposited in the Herbarium of the New England Bo- 
tanical Club for the record. Only one other New England speci- 
men has been brought to my attention, viz., from Rutland, 
Massachusetts, coll. Mrs. Rufus B. Dodge.—Ricnarp J. EATON, 
Lincoln, Massachusetts 


THE GENUS LIATRIS 


L. O. GAISER 


(Continued from page 263) 


10. LIATRIS GRACILIS Pursh. Corm small, globose, up to 3 
em. in diameter, in tall robust plants giving rise to a dense spread- 
ing fibrous root-system; stems slender to stiff and thick (up to 
0.7 em. in diameter at the base), often reddish, glabrous or more 
generally softly cinereous-pubescent, 2-10 dm. tall: leaves 
glabrous, ciliate or with few scattered hairs, punctate, short and 
numerous, lanceolate to linear; longest basal ones about 2 dm. 
long, 0.5-1 cm. wide, bluntly lanceolate, narrowed to a winged 
petiole with scattered cilia along the margin; other leaves reduced 
upwards to bracts, 1-2 cm. long and 1 mm. wide; inflorescence a 
simple raceme often ca. 3 dm. long, sometimes becoming panicu- 
late and twice as long; heads on short filiform, villous pedicels, 
at least as long as and often longer than the heads, divaricate 
and frequently at right angles to the pubescent rachis, often, in 
paniculate inflorescences, becoming slender bracteolate peduncles 
15 cm. long: heads the smallest of any species of the genus, 7-9 
mm. long and about as wide when the flowers are open, of 4—6 
flowers; phyllaries rather thin, oblong, obtuse, rarely tending to 
become acute, sometimes pubescent, ciliate on the margin, 
loosely erect: outer ones short, ovate, 2 mm. long and ca. 1 mm. 
wide; inner ones 5.5 mm. long and ca. 1.5 mm. wide; corolla from 
deep lavender to violet, 5-7 mm. long, densely pilose within the 
upper part of the tube and lower part of the throat; pappus 
hardly as long as the corolla, ca. 5 mm., short-barbellate; achene 
ca. 3 mm. long—Fl. Amer. Sept. 508 (1814); Gray, Synop. FI. 7’. 
111 (1884). Laciniaria gracilis O. Ktze., Rev. Gen. 1349 (1891), 
not, sensu Small, Man. S. E. Fl. 1834 (1933). Liatris pauciflos- 
culosa Nutt. Jour. Acad. Phil. vii. 71 (1834). L. lanceolata 
Bertol. Mise. Bot. v. 11, t. 3 (1846). Lacinaria laxa Small, 
Bull. Torr. Bot. Club, xxv. 472 (1898) and Man. S. E. Fl. 1334 
(1933). Liatris laxa. K. Sch. in Just, Jahresb. xxvi!. 378 (1900). 


274 Rhodora [OCTOBER 


South Carolina, Georgia, Alabama and especially Florida.— 
SOUTH CAROLINA. Burauvrort Co.: dry woods, St. Helena 
Isl., Sept. 1882, A. Cuthbert (F); St. Helena Isl., Sept. 1892, A. 
Cuthbert (NY); dry barrens, St. Helena Isl., Sept. 1894, A. Cuth- 
bert (F); flat pine woods, St. Helena Isl., Oct. 1902, A. Cuthbert 
(QN Y 874) F); Oct. 1903, A. Cuthbert (F). GEORGIA. Without 
stated locality: Nuttall (P), L. Conte (P), 1844, Dr. Harden (P); 
1831, Gates (NY). Co. undetermined: Atlantic & Gulf Rwy., 
Chapman (NY). GyxNN Co.: sandy pine land, Brunswick, Oct. 
10, 1912, F. W. Pennell, 4837 (P). Berrien Co.: rather dry 
flat pine barrens, ca. 114 mis. s. of Tifton, Sept. 26, 1902, R. M. 
Harper, 1683 (G, NY, US). THomas Co.: Thomasville, Oct. 16, 
1903, Taylor (G). Decatur Co.: Bainbridge, 1875, A. H. Cur- 
tiss (US). FLORIDA. Without stated locality: Chapman (G, 
NY, P, US), Chapman, 52 (US), A. W. Chapman, ex Herb. G. 
Thurber (G), A. P. Garber (US), Leavenworth (G, NY), Rugel 
(US). Co. UNDETERMINED: southern Florida, Chapman (NY); 
pine barrens, Chapman, ex Herb. J. A. Lowell (G); Fort Dallas, J. 
G. C. (NY). Duvar Co.: dry pine barrens, Jacksonville, Oct. 25, 
1894, A. H. Curtiss, 5312 (G, NY, US); dry pine barrens, near 
Jacksonville, Oct., A. H. Curtiss, 1181 (G, NY, P, Q, US); Jack- 
sonville, Oct. 25, 1893, A. H. Curtiss, 4446 (NY, US); Jackson- 
ville, Nov. 1891, Herb. W. G. Farlow (G); Jacksonville, Oct. 18, 
1898, A. H. Curtiss (G); dry pine barrens, near Jacksonville, Oct. 
18, 1898, A. H. Curtiss, 6290 (NC, F); Jacksonville, Nov. 14, 
1901, J. K. Small & G. V. Nash, 369 (NY). CoruwarA Co.: dry 
open woods, Camp Oleno, Oct. 1, 1939, Watson & Murrill (F). 
HawiLTON Co.: dry woods on Suwannee R., White Springs, Sept. 
30, 1941, E. West & Miss L. Arnold (F). Jerrerson Co.: high 
pine-oak woods, n. of Monticello, Oct. 7, 1940, W. A. Murrill (F). 
GADSDEN Co.: without locality, Oct. 24, 1880, C. Mohr (US); 
open pinelands, w. part of county, Aug. 30, 1936, H. Foster 115 
(F). Jackson Co.: Cypress, Oct. 21, 1941, R. A. Knight (F). CAL- 
HOUN Co.: without locality, Oct. 25, 1944, R. A. Knight (F). 
LrBERTY Co.: sandy pine barrens, Aspalaga, Oct. 1897, Biltmore 
Herb., 577a (G, NY, ND). FRANKLIN Co.: sandy pine barrens, 
Apalachicola, Oct. 3, 1882, Biltmore Herb., 577b (G, NY, NC, 
Q, US); dry pine barrens; Apalachicola, B. F. Saurman (P); 
Apalachicola, 1867, B. F. Saurman (P). Wasuineton Co.: 
Chipley, Oct. 13, 1943, R. A. Knight (F). Bay Co.: dry sandy 
ground, Lynn Haven, Oct. 14, 1921, C. Billington (US). Oxa- 
LOOSA Co.: Crestview, Oct. 23, 1939, R. A. Knight (F); Crest- 
view, Oct. 9, 1939, Mrs. G. Barrow (F). Santa Rosa Co.: 
sandy pine land, Milligan, Sept. 10, 1912, F. W. Pennell (NY, 
US). SaiNT Jonns Co.: St. Augustine, M. C. Reynolds (NY); 
pine barrens, M. C. Reynolds (US). PurNAM Co.: flatwoods, 
S. E. Adm. Bldg., Welaka, Oct. 26, 1944, A. M. Laessle (F). 


1946] Gaiser,—The Genus Liatris 275 


BnaprFonD Co.: Starke, Nov. 1893, Miss G. Gilbert (G). UNION 
Co.: on dry ground near highway n. of Worthington Springs, 
Oct. 20, 1945, H. H. Hume (F). ArAcHUA Co.: high open 
land, Warren’s Cave, Gainesville, Oct. 5, 1927, G. F. Weber 
& E. West (F); roadside, Archer Rd., Gainesville, Oct. 31, 1931, 
Miss L. Arnold (F); upper slopes, Devil's Millhopper, Gaines- 
ville, Oct. 23, 1932, Miss L. Arnold (F). TayLor Co.: 9 mis. s. 
of Perry, Oct. 8, 1940, W. A. Murrill (F). FLAGLER Co.: road- 
side just e. of Bunnell, Oct. 10, 1940, E. West and Miss L. Arnold 
(F). Marron Co.: dry woods, Belleview, Sept. 15, 1927, O. F. 
Burger & E. West (F). VorusriA Co.: low sandy ground, De 
Land road, Nov. 6, 1944, Mrs. H. T. Butts (OA). SEMINOLE Co.: 
high pineland, Altamonte Springs, Oct. 4, 1928, E. West (F). 
ORANGE Co.: Wekiwa Springs, Dec. 5, 1929, H. N. Moldenke, 
1920 (NY); Wekiwa Springs, Sept. 25, 1929, H. O'Neill (US, F); 
high pineland, Orlando, Sept. 24, 1927, O. F. Burger & E. West 
(F); high pineland, Zellwood, Sept. 24, 1927, O. F. Burger & E. 
West (F). LAKE Co.:4 mis. s. w. of Astor Park, Oct. 13, 1940, W. 
A. Murrill (F). SuwTER Co.: 3 mis. s. of St. Catherines, Oct. 14, 
1943, W. B. Tisdale (F). Crrrus Co.: Inverness, M. A. Noble 
(P); sandy dry oak-pine woods, 1 mi. s. of Floral City, Oct. 16, 
1945, H. H. Hume (F). Hernanpo Co.: vic. of Brooksville, 
Sept. 21, Oct. 19, 22, 1919, R. N. Jones (US). Brevarp Co.: 
low pine barrens, Okeechobee, Sept. 26, 1903, A. Fredholm, 6027 
(G); serub oakland, Okeechobee region, Oct. 20, 1903, A. Fred- 
holm, 6148 (G). HrirnLsBonouaH Co.: Tampa, Sept. 1877, A. P. 
Garber (P); dry sand, Oct. 13, 1904, A. Fredholm, 6424 (G); in a 
high hammock, Riverview, Oct. 8, 1930, F. S. Blanton, 6794 
(US); flatwoods n. of Riverview, Oct. 19, 1945, L. O. Gaiser, Mrs. 
H. T. Butts, Miss L. Arnold, Aug. 19, 1945 (F). PriNELLAS Co.: 
Tarpon Springs, Aug. 1894, C. S. Williamson (P); sandy open or 
light wooded space, St. Petersburg, Sept. 17, 1932, G. M. Hocking 
(F). Inpran River Co.: between Fellsmere and Sebastian, 
Aug. 13, 1925, R. M. Harper, 53 (US). OKEECHOBEE Co.: pine- 
land, Okeechobee, Oct. 12, 1941, J. H. Davis (F). MANATEE 
Co.: Tampa Bay, Herb. G. Thurber (G); Bradentown, Sept. 29, 
1900, S. M. Tracy 7105 (G, NY, US); Manatee, 1889, J. H. 
Simpson (US); flatwoods, at Palma Sola, Bradentown, Oct. 10, 
1920, A. Cuthbert (F); flatwoods, n. of Parrish, Aug. 19, 1945, 
L. O. Gaiser, Mrs. H. T. Butts, Miss L. Arnold (F). SARASOTA 
Co.: air field in flatwoods, Venice, Aug. 19, 1945, L. O. Gaiser, 
Mrs. H. T. Butts, Miss L. Arnold (F). Henpry Co.: wet pine- 
lands, Indian Reservation, Jan. 15, 1942, J. H. Davis (F). LEE 
Co.: in pineland, vicinity of Fort Myers, Oct. 29, 1916, Miss J. 
P. Standley, 415 (G, P, US); pine flatwoods, 5 mis. s. of Ft. 
Myers, Oct. 20, 1942, J. H. Davis (F). Browarp Co.: pinelands, 
Fort Lauderdale, Nov. 19-25, 1903, J. K. Small & J. J. Carter, 


276 Rhodora [OcroBER 


1470 (NY); Fort Lauderdale, Nov. 1903, J. K. Small & J. J. 
Carter, 1069 (P). Dape Co.: Cape Florida, Mar. 14, 1892, Mrs. 
J. H. Simpson, 537 (G (NY, TYPE of Lacinaria laxa Small)); 
pinelands, w. of Silver Palm, Nov. 21, 1916, J. K. Small, 7982 
(NY); pinelands, w. of Silver Palm, Nov. 21, 1916, J. K. Small, 
7974 (G, NY, F); pinelands w. of Rockdale, Nov. 18, 1916, J. K. 
Small, 7899 (NY); pineland prairie, Tamiami Trail, w. of Miami, 
Dec. 19, 1919, J. K. Small, N. L. Britton, & M. De Winkeler, 
9372 (NY); pinelands, near Camp Jackson ca. 35 mis. s. w. of 
Miami, Jan. 7-10, 1909, J. K. Small & J. J. Carter, 3043 (NY) 
pinelands, w. of Miami, Apr. 7, 1904, N. L. Britton, 475 (NY); 
pinelands, between Cocoanut Grove and Cutler, Oct. 31-Nov. 4, 
1903, J. K. Small & J. J. Carter, 719 (NY, P); pinelands, between 
Cocoanut Grove and Cutler, Oct. 31-Nov. 4, 1903, J. K. Small & 
J. J. Carter, 753 (NY); Buena Vista, Jan. 21, 1930, H. N. Mol- 
denke, 4850 (NY); s. of Miami, Mar. 27, 1930, H. N. Moldenke, 
5858 (NY); e. of Naranja, Jan. 14, 1909, J. K. Small & J. J. 
Carter (P). ALABAMA. Without stated locality: Gates (G, 
NY, P, B, isotypes of L. lanceolata Bert.), C. W. Short (P), Herb. 
G. Thurber (G), Herb. Torrey (NY). Co. undetermined: pine 
woods, Buckley (NY). Covineton Co.: Blue Springs, near 
McRae, Sept. 13, 1912, F. W. Pennell, 4615 (P). BarpwiN Co.: 
Gateswood, Oct. 31, 1903, S. M. Tracy, 8559 (G, NY, ND, T, 
US). Mose Co.: sandy dry pine woods, Mobile, Oct. 1878, 
C. Mohr (US); dry sandy pine barrens, Mobile, Oct. 1880, C. 
Mohr (US); Mobile, 1878, C. Mohr (US); mixed wood, s. of 
Mobile, Sept. 24, 1912, H. H. Bartlett, 3226 (G, US); 3217, in 
part (US); Whistler, Oct. 10, 1884, C. Mohr (US); Spring Hill, 
Aug. 3, 1897, B. F. Bush, 150 (NY); Spring Hill, Sept. 28, 1878, 
C. Mohr (NY, US), Oct. 1890 (NY), Oct. 20 (US); hillside, Spring 
Hill, Sept. 1919, E. W. Graves, 1324 (US); pine woods, w. of 
Spring Hill, Aug. 1918, E. W. Graves, 594 (US); pine ridges, 
Sept. 28, 1879, C. Mohr (NY). 


It was possible to compare a photograph of the type specimen 
of L. gracilis Pursh, obtained by Mr. Weatherby at Kew, and 
two specimens at the Philadelphia Academy of Science, of Gates 
from Alabama, one of which is probably the co-type, if not the 
type, of L. pauciflosculosa Nutt. While the phyllaries are slightly 
acute in one Gates specimen they are obtuse in the other and 
with their filiform, divaricate pedicels, closely resemble the Pursh 
specimen, except for the fact that Pursh described his plant as 
glabrous while Nuttall described the stem as puberulent. Since 
such a surface character alone has not been of diacritical signif- 
icance in this genus, and there is a wide variation in the degree of 


) 


1946] Gaiser,—The Genus Liatris 277 


pubescence in this generally ciliate species the two names seem 
synonymous, as does also L. lanceolata Bertol. of which isotypes 
were seen in four herbaria (G, NY, P, B). 

Careful examination of the plant of J. H. Simpson, no. 537, 
from Cape Florida (NY) which was described by Small (Bull. 
Torr. Bot. Club, xxv. 472 (1898)) as Lacinaria laxa showed no 
decisive characters by which it could be separated from L. gracilis. 

Abounding chiefly in Florida, though occurring also in South- 
ern Georgia and Alabama, L. gracilis is distinguishable from L. 
graminifolia var. elegantula which overlaps its northern range, by 
its smaller heads, flowers and achenes. 


11. LIATRIS REGIMONTIS (Small) K. Sch. Corm globose, 1.5- 
2 cm. in diameter; stem slender, glabrous, striate, 4-7 dm. high: 
leaves few, linear, mostly glabrous with marginal cilia, but some- 
times with hairs on upper surface as well as beneath; basal ones 
6-10 cm. long and ca. 5 mm. wide having a few cilia at base: 
inflorescence long, covering one half to three quarters of the 
length of the stem, with numerous sessile or peduncled heads 
forming a spike or slightly branched panicle: heads 6-12-flowered, 
ca. 10 mm. long, cylindrical but with the phyllaries slightly 
spreading rather than appressed, the mid-vein showing rather 
distinctly and forming a rather rigid keel; outer phyllaries ovate, 
mucronate-tipped, and with entire margins, the inner ovate- 
lanceolate with somewhat membranous margins; corolla 7 mm. 
long with appressed hairs at the base of the throat and upper 
part of the tube, sometimes appearing up to the base of the lobes; 
pappus barbellate, 3.5-4 mm. long: achene 3-3.5 mm. long.—Just, 
Bot. Jahresb. xxvi. pt. 1, 378 (1900). Lacinaria regimontis 
Small, Bull. Torr. Bot. Club, xxv. 473 (1898). Liatris carinata 
Coker, Pl. Life of Hartsville, 108 (1912). Lacinzaria carinata 
Small. Fl. S. E. U. S. 1174 (1903). 

Eastward from the mountains in Virginia, North and South 
Carolina.—VIRGINIA. WvrHE Co.: Wytheville, H. Shriver, 
9874 (P. NORTH CAROLINA. FonsvrH Co.: Winston 
Salem, Sept. 22, 1927, E. T. Wherry & F. W. Pennell, 14348 (P). 
Wake Co.: Raleigh, Sept. 6, 1908, W. W. Eggleston, 4052 (G, 
US); Burxe Co.: Bridgewater, Sept. 20, 1927, E. T. Wherry & 
F. W. Pennell, 14297 (P); WAxNE Co.: sand ridge, near Golds- 
boro, Sept. 3, 1938, R. K. Godfrey, 6562 (G, P). Moore Co.: 
Pinehurst, Aug.-Sept. 1897, O. Katzenstein (G); sandy places, 
Sept. 1, 1940, P. O. Schallert (G). CLEVELAND Co.: King's Mt. 
and vicinity (alt. 600'-1300^, Aug. 27-30, 1894, J. K. Small 
(NY, type); King's Mt. (alt. 2600'-3000^, Sept. 10, 1908, W. W. 
Eggleston, 4073 (NY, US); woodland, near King's Mt., Oct. 7, 


278 Rhodora [OCTOBER 


1902, Biltmore Herb., 15006 (NY). ScorrANDp Co.: 10 mis. s. 
of Aberdeen, Oct. 12, 1938, R. K. Godfrey, 6941 (G); sandhill, 
10 mis. n. of Laurenburg, July 14, 1938, R. K. Godfrey, 5045 (G). 
SOUTH CAROLINA. Co. undetermined: dry soil, Santee 
Canal, Oct., H. W. Ravenel (G). CHESTERFIELD Co.: 1 mi. w. of 
McBee, Sept. 7, 1939, R. K. Godfrey, 8055 (G, NY). ANDERSON 
Co.: dry slopes, Anderson, Aug. 10, 1912, J. Davis (NY); dry 
pine woods, Anderson, Aug. 15, 1919, J. Davis, 7794 (US). Oco- 
NEE Co.: without stated locality, A. P. Anderson, Sept. 9, 1898, 
1529, Sept. 11, 1898, 1546 (US); Dartrneron Co.: sandy pine 
woods, Society Hill (common), M. A. Curtis (G); low pine 
lands, Society Hill (1 stem to left), Sept. 8, 1904, Biltmore Herb., 
14935a (NY). Kersuaw Co.: 3 mis. w. of Bethune, Sept. 7, 
1939, R. K. Godfrey, 8014 (G, NY). Suwrzn Co.: local, Sumter, 
Oct. 21, 1913, Herb. E. B. Bartram, 3248 (P). LrxiNGToN Co.: 
6 mis. s. of Columbia, Aug. 7, 1939, R. K. Godfrey & R. M. Tryon, 
1247 (G, NY). EnpaErrELD Co.: sandy ridges, Sept. 22, 1883, 
J. D. Smith, 34 (US). GronaETOWwN Co.: pine barren clearing, 
5 mis. s. of Andrews, Sept. 13, 1939, R. K. Godfrey, 8190 (G, NY); 
sandy pine woods, Oct. 12, 1934, F. G. Tarbox Jr., 175-2, 175-3 
(NY). Arken Co.: Aiken, Sept. 12-15, 1909, W. W. Eggleston, 
5047 (G, NY). BERKELEY Co.: sandy soil, Monk's Corner, Oct. 
14, 1912, F. W. Pennell, 4888 (NY); near Monk's Corner, Sept. 
29, 1856, L. R. Gibbs (NY). Oraneesure Co.: Eutauville, Sept. 
6-11, 1909, W. W. Eggleston, 4952, 4989 (NY), 4966 (NY, P), 
5008 (G, NY). Braurort Co.: dry sandy bluff, Sept. 5, 1904, 
Biltmore Herb., 14935b (NY). GEORGIA. Without stated 
locality: Croom (G). 

This species is found in North and South Carolina, east of the 
mountains, where also the varieties typica and dubia of L. 
graminifolia occur. It is best distinguished from those varieties, 
having leaves with prominent hairs along the petioles or on the 
surfaces and turbinate heads with thin, ciliolate-margined phyl- 
laries, by almost glabrous leaves and almost entire-margined, 
somewhat keeled phyllaries, the outer appearing revolute. 

Laciniaria carinata described by Small, l. c., from the same 
general region, was included in Small, Man. S. E. Fl. 1333 (1933) 
under Lacinaria secunda (Ell.) Small. Examination of the speci- 
mens bearing this name in the herbarium (NY) shows a closer 
relationship to L. regimontis in the size of the heads and corolla- 
tubes (ca. 7 mm. long in regimontis and 10-12 mm. long in se- 
cunda) and in the lack of the puberulent stem of L. secunda. The 
phyllaries are perhaps slightly more membranous but if not 
“thick lax-tipped", as regimontis was described by Small, are 


1946] Gaiser,—The Genus Liatris 279 


nevertheless spreading and show the midvein projecting at the 
tip. It is here included in the synonymy of L. regimontis. 

The corolla-tube in all these specimens is pilose at the base of 
the throat, above the attachment of the filaments, and so shows 
relationship to the species of the Graminzfoliae series rather than 
to L. microcephala, which has a smooth corolla-tube and small 
cylindrical heads of appressed bracts. When found in a young 
inflorescence stage it casually resembles and could be confused 
with this species. 


Series IV. PaucrFLoRAE. Herbs of one or two stems, 
glabrous or short-pubescent, with numerous short, linear to 
linear-lanceolate leaves; inflorescence spicate, racemose or simply 
paniculate, usually erect (often reclining in L. secunda), of slender 
long cylindrical compact heads 1-2 cm. long, usually 3-6-flowered, 
with appressed glabrous, thin linear-acuminate to -acute phyl- 
laries; corolla-tube 10-12 mm. long, with only very few hairs 
within and few scattered ones at the base without; achene ca. 4 
mm. long.—From sandy pinelands of the coastal plain from 
North Carolina to Florida, Alabama and Louisiana. 


a. Slender sessile crowded heads densely erect, giving a narrow 

stiff spicate inflorescence, glabrous or short-pubescent; 

corolla-tube with slight pilosity within the base of the 
Dos ant pein.) > Pee es Wo ee ES «...12. L. Chapmanii. 

a. Slender short-pedicellate heads less crowded in loose racemes 

or simple panicles; corolla-tube with only a few scattered 

hairs within and outside at the base of the tube... . b. 

b. Stem and leaves glabrous; inflorescence erect, racemose or 
panioulatec s.. Lan tenaa PLATO Lud 13. L. pauciflora. 

b. Stem short-pubescent; leaves pubescent, or ciliate toward 

the base; inflorescence racemose, with heads frequently 
RT res n ee N awa E SR. re 14. L. secunda. 


12. LIATRIS CHAPMANI Torr. & Gray. Corm rounded or 
ovoid up to 2 cm. in diameter: stem usually single, stiff, minutely 
cinereous-pubescent or glabrous, very leafy, 3-7.5 dm. tall; 
leaves very numerous, linear-lanceolate, inconspicuously punc- 
tate, finely pubescent or glabrous, lower ones ca. 1.5 dm. long, 
and 5-10 mm. wide, forming a rosette at the base of the stem; 
upper leaves narrowed below the middle into a slender margined 
petiole, gradually reduced, 1.5 cm. long at the base of the spike; in- 
florescence a very dense, narrow spike, 1-3.5 dm. long, of slender, 
3—-5-flowered, cylindrical, nearly sessile heads, 1-1.8 cm. long, 
erect and generally closely appressed to the rachis; phyllaries 
oblong-lanceolate, acute or mucronulate, glabrous and with 
narrow scarious margins, sometimes purple; corolla phlox-purple, 
rarely white, ca. 12 mm. long, with very slight pilosity, if any, 
within the tube at the base of the throat; achene 4-5 mm. long; 


280 Rhodora [OCTOBER 


pappus 9-11 mm. long and barbellulate, with barbellae about 
equal to or twice the diameter of the seta.—Fl. N. Am. ii. 502 
(1843); Chapman, Fl. So. U. S. 191 (1860); Gray, Synopt. FI. i?. 
112 (1884). Laciniaria Chapmanii O. Ktze. Rev. Gen. i. 349 
(1891). Laciniaria Deamiae Lunell, Amer. Midl. Nat. ii. 163 
(1912). Lacinaria Chapmanii var. longifolia Nash, Bull. Torr. 
Bot. Club, xxiii. 106 (1896). 

Georgia and Florida.—GEORGIA. Decatur Co.: without 
locality, Sept. 15, 1880, Dr. E. A. Smith (NY). FLORIDA. 
Without stated locality: Chapman, ex Herb. C. Mohr (US); 
Curtiss, ex Herb. U. S. Dept. Agr. (US); ex Herb. Park Davis Co., 
3496 (US). Co. UNDETERMINED: sandhills, A. W. Chapman (G, 
(NY type)); on arid sandhills, Chapman (G); southern Florida, 
A. W. Chapman (NY, US). Leron Co.: Bellair, Sept. 3, 1895, 
G. V. Nash, 2547 (G, NY, US, ND). GapspEN Co.: dry sandy 
pine ridges, near Bristol & Quincy, Oct. 24, 1895, C. Mohr (US); 
on arid sandhills, Sept., Chapman (G); open pinelands, western 
part of county, Aug. 30, 1936, H. Foster, 113 (F). FRANKLIN Co.: 
dry sandy ridges, near Apalachicola, July 12, 1895, ex Biltmore 
Herb., 4114a (G, NY, US, ND); dry pine barrens, near Apalachi- 
cola, Oct. A. H. Curtiss, 5441 (NY); Apalachicola, Oct. Curtiss, 
(NY); dry pine barrens, near Apalachicola, Oct. 1888, A. H. 
Curtiss, 1182 ((G, US, without date) NY, ND)); St. Vincent 
Island, Oct. 30, 1910, W. L. McAtee, 1730, (US); St. Vincent 
Island, Oct. 30, 1910, W. L. McAtee, 1743 (US). Okaroosa Co.: 
East Pass, S. M. Tracy, 6362 (G, US, NC). Escamsīia Co.: 
barren scrubs near Pensacola, Sept. 26, 1901, A. H. Curtiss, 6918 
(G, NY, US, Q); high dry pine barrens, w. side of Escambia Bay, 
Sept. 20, 1910, R. M. Harper, 88 (G, NY, US). PurNaAM Co.: 
with low turkey-oak, along n. boundary, Welaka, April 28, 1940, 
A. M. Laessle (F). VorusrA Co.: dry scrub, Tomoka Ave., 
near Ormond, Sept. 24, 1944, Mrs. H. T. Butts (OA); flatwoods, 
s. of Seville, Sept. 17, 1943, G. West & Miss L. Arnold (F). 
ORANGE Co.: flatwoods, Orlando, Oct. 18, 1929, E. West & De 
Vane (F); sandhills, Windermere, Sept. 19, 1929, F. Vasku (F). 
HERNANDO Co.: Choocochattee Hammock, near Brooksville, 
Aug. 26, 1922, J. K. Small, J. W. Small & J. B. DeWinkeler, 
10604 (NY, F). Brevarp Co.: Okeechobee region, Aug. 13, 
1903, A. Fredholm, 5962 (G). Hr~usBoroucH Co.: Tampa, Aug. 
24, 1895, G. V. Nash, 2473 (G (NY, type of Lacinaria Chap- 
mannii var. longifolia Nash) US, ND, F). PriNELLAS Co.: sandy 
waste, near Veteran City, 8 mis. w. of St. Petersburg, Sept. 29, 
1907, Mrs. C. C. Deam, 2804 (G, NY, US, I, cotypes of Laciniaria 
Deamiae Lunnell). HiaHLANDS Co.: sandhills, near De Soto city, 
Aug. 31, 1922, J. K. Small, J. W. Small & J. B. DeWinkeler, 
10683 (G, NY); Avon Park, Sept. 5, 1934, J. K. Small, E. West, 
J.B. McFarlin (F). MawNaTEE Co.: (albino) high bank of Mana- 


1946] Gaiser,—The Genus Liatris 281 


tee River, Sept. 17, 1916, A. Cuthbert (F); high ridges, Braden- 
town, Sept. 16, 1916, A. Cuthbert (F); high bank of Manatee 
River, near Palma Sola, Sept. 16, 1916, A. Cuthbert (F); sand- 
hills along Manatee River, Bradentown, Sept. 17, 1916, A. Cuth- 
bert (F). SarNT Lucie Co.: low dune, 8 mis. s. of Fort Pierce, 
Oct. 4, 1941, E. Kurz (F). Ler Co.: pinelands, Deering Reserva- 
tion, Cutler, July 20, 1924, J. K. Small, J. W. Small & J. B. 
DeWinkeler, 11538 (G, NY). Correr Co.: Cypress Head, s. of 
Naples, Aug. 29, 1922, J. K. Small, J. W. Small & J. B. De 
Winkeler, 10619 (NY); in Ceratiola scrub, Naples, Oct. 17, 1941, 
J. H. Davis (F). Dave Co.: in sandy pineland, Buena Vista, 
Jan. 21, 1930, H. N. Moldenke, 483a (N Y); in dry sandy pineland, 
Buena Vista, Miami, Mar. 4, 1930, H. N. Moldenke, 6540 (NY); 
in pinelands, between Cocoanut Grove & Cutler, Oct. 31-Nov. 4, 
1903, J. K. Small & J. J. Carter, 710 (NY); Miami, Sept. 1, 
1939, E. West (F); in sand, Homestead, Sept. 7, 1928, G. F. 
Weber (F). 


L. Chapmanii is to be distinguished from L. pauciflora Pursh, 
also occurring in Florida, by its stiffer, narrow spike of crowded, 
erect, almost sessile heads rather than the usually pedicellate ones 
in an erect, simple panicle or raceme characteristic of that species. 
No distinctive characters could be found in the type of Laciniaria 
Deamiae Lunell to differentiate it from this species, nor did the 
length of basal leaves seem significant enough for varietal recog- 
nition of the Nash specimen from Tampa. 


13. LrATRIS PAUCIFLORA Pursh. Corm small, globose, 1-2 cm. 
in diameter: stem generally single, erect but slender, glabrous, 
striate, leafy, 2-5 dm. tall: leaves narrow, inconspicuously 
punctate; basal ones not abundant, glabrous with a few cilia on 
the petiole, 7-10 cm. long, reduced upwards to 1 cm. long and 1 
mm. wide at the base of the inflorescence: this 10-25 cm. long, a 
loose raceme or simple panicle: heads on very slender pedicels 
3-5 cm. long, 3-6-flowered; corollas projecting well beyond the 
tips of the involucre; phyllaries glabrous, ovate-lanceolate with 
acute to acuminate tips; corolla phlox-purple, 11-12 mm. long, 
with a few hairs scattered over the entire inner surface and outer 
base of the tube; achene 4-5 mm. long and sharply pointed; 
pappus 9-11 mm. long, with short barbellae about equal to or 
twice the diameter of the seta.—Fl. Am. Sept. ii. 510 (1814); 
DC. Prod. v. 131 (1838); Gray, Synopt. Fl. i?. 112 (1884), in part. 
Laciniaria pauciflora O. Ktze. Rev. Gen. i. 349 (1891). 

Mostly northern Florida.—FLORIDA. Co. UNDETERMINED: 
east Florida, Lt. Alden (NY); dry sand, high pineland, Dr. 
Leavenworth (NY). HawrirTroN Co.: dry woods, on Suwannee R. 
White Springs, Sept. 30, 1941, E. West & Miss L. Arnold (F). 


282 Rhodora [OCTOBER 


SUWANNEE Co.: dry pine barrens, near Live Oak, Sept. 12, 1901, 
A. H. Curtiss, 6896 (G, NY, US, Q). Watron Co.: dry pine 
barrens, between Freeport & Portland, Sept. 23, 1910, R. M. 
Harper, 90 (US right plant). Purnam Co.: just n. of Orange 
Springs, Sept. 23, 1939, Watson & Murrill (F). ArAcnuvA Co.: 
blackjack ridge, w. of Archer, Aug. 25, 1922, J. K. Small, J. W. 
Small & J. B. DeWinkeler, 10585 (G, NY); sandy open soil, 
Gainesville, Aug. 31, 1927, O. F. Burger (F); dry oak woods, 
Rochelle, Sept. 5, 1927, E. West (F). Drxr& Co.: pineland, 10 
mis. w. of Shamrock, Aug. 22, 1937, Pasture Survey (F). Marion 
Co.: high pine woods, Belleview, Sept. 15, 1927, O. F. Burger & E. 
West (F). Levy Co.: Bronson, Sept. 9, 1936, Pasture Survey 
(F). OnaNGE Co.: Clarcona, Jan. 20, 1900, Miss M. Meislahn, 
148a (US); Clarcona, Sept. 19, 1889, Miss M. Meislahn, 39 (US); 
Clarcona, Dec. 20, 1899, Miss M. Meislahn, 138a (US). Her- 
NANDO Co.: dry pineland, near Weekiwachee Spring, Sept. 4, 
1937, Mrs. M. F. Baker (F). Laxes Co.: vicinity of Eustis, Aug. 
16-25, 1894, G. V. Nash, 1711 (G, NY, US, ND). HirLsBo- 
ROUGH Co.: low pineland, 12 mis. n. of Tampa, Sept. 15, 1930, 
F. S. Blanton & H. O'Neill, 6639 (US); Tampa Bay, ex Herb. G. 
Thurber (G); Tampa Bay, Oct. 1877, A. P. Garber (US (P, left 
plant)); sandhill, Hillsborough (plant to right), Sept. 17, 1904, 
A. Fredholm, 6392 (G). 


For discussion see the next species. 


14. LIATRIS SECUNDA Ell. Corm small and globose, 1-2 cm. 
in diameter; stem usually one, slender, not stiff, puberulent, fre- 
quently reclining and 3-5 dm. in length; leaves not abundant, 
lanceolate, inconspicuously punctate, ciliate along the margin 
and towards the base; basal ones lanceolate, 7-10 cm. long, 5-8 
mm. wide; upper ones reducing in length to 1 em. and in width 
to 2 mm. wide: inflorescence a loose terminal raceme 1.5-2 dm. 
long with 3-6-flowered heads ca. 1.5 cm. long, borne single on 
peduncles 2-3 cm. long, frequently secund if plant reclining; 
phyllaries oblong-lanceolate, acute, sometimes acuminate, gla- 
brous but finely ciliolate along the margin; corolla phlox-purple, 
very slender, tubular, 11-12 mm. long, with a few hairs inside the 
full length of the tube and sometimes with few scattered hairs on 
outside at base of the tube; achene ca. 4 mm. long; pappus 8-9 
mm. long, barbellulate, barbellae about three times the diameter 
of the seta.—Sk. ii. 278 (1822(?)); Torr. & Gray, Fl. N. Am. ii. 
71 (1841); Chapman, Fl. So. U. S. 191 (1860). Lacinaria secunda 
(Ell.) Small Man. S. E. Fl. 1331 (1933). 

From the southern coastal plain region of North Carolina 
through South Carolina, Georgia, and central and western 
Florida, to Alabama and Louisiana.—NORTH CAROLINA. 
Without stated locality: ex Herb. Chapman (NY). New Haw- 


1946] Gaiser,—The Genus Liatris 283 


OVER Co.: dry sand barrens, Wilmington, Oct. 2, 1908, E. B. 
Bartram (NY); dry sand, Wilmington, Oct. 4, 1908, Oct. 3, 1909, 
E. B. Bartram (P); Wilmington, Aug. 1842, C. S. Williamson (P); 
near Wilmington, Oct. 1867, W. M. Canby (NY, P, US); 3 mis. s. 
of Wilmington, July 25, 1922, L. F. Randolph & F. Randolph, 
1023 (G); Wilmington, 1885, G. McCarthy (US); Wilmington, 
1880, Sept. 22, 1882, T. F. Wood (US); Greenfield Lake, at 
Wilmington, Aug. 7, 1938, R. K. Godfrey & B. W. Wells, 5914, 
(G). Brunswick Co.: in coarse sand of long-leaf pine barren, 
10 mis. n. of Southport, Sept. 12, 1941, R. K. Godfrey, 1189 (G, 
NY). SOUTH CAROLINA. Without stated locality: Gibbes, 
(G). CHESTERFIELD Co.: among scrub-oak & long-leaf pine, 1 
mi. w. of McBee, Sept. 7, 1939, R. K. Godfrey, 8084 (G, NY, P). 
DaRLINGTON Co.: Society Hill, M. A. Curtiss (G); sand hills, 
across Blast Creek, Hartsville (one plant), Aug. 22, 1908, W. C. 
Coker (NY). LEkxiNGTON Co.: sandy burned clearing, 14 mis. s. 
of Columbia, Aug. 7, 1939, R. K. Godfrey & R. M. Tyron, 1309 
(G, NY); sandy pine woods, near Columbia, Sept. 1889, Miss 
Crawford (US); 2 mis. n. e. of Columbia, Sept. 25, 1883, J. D. 
Smith, 32 (US). EpcaErrELD Co.: sandy blackjack-pine ridges, 
Sept. 22, 1883, J. D. Smith, 31 (G without no., US). GEORGE- 
TOWN Co.: in swamp, near Georgetown, Aug. 1935, W. Rhoades 
(G); Georgetown, Sept. 9, 1933, F. G. Tarbox, 710 (NY); pine 
barren clearing, 5 mis. s. of Georgetown, Sept. 9, 1939, R. K. 
Godfrey, 8123 (G, NY, P). Arken Co.: (albino) Aiken, Sept. 
12-15, 1909, W. W. Eggleston, 5044 (G, NY, US, P); Aiken, Sept. 
1869, H. W. Ravenel (US). GEORGIA. Without stated local- 
ity: ex Herb. Boston Soc. Nat. Hist. (G). RrcnwoNp Co.: barrens, 
Augusta, Sept. 1897, A. Cuthbert (NY, F); sand hills, Augusta, 
A. Cuthbert, Sept. 29, 1898, 310 (NY), Sept. 10, 1903 (NY), 
Sept. 8, 1902 (US); sand hills, sterile ridges, Augusta, Sept. 4, 
1902, A. Cuthbert, 785 (F); sand hills, no. of Augusta, Oct. 16, 
1937, J. H. Pyron & R. McVaugh, 1866 (US); Augusta, G. 
McCarthy, Aug. 1888 (NY, ND), Sept. 20, 1888 (US); Augusta, 
1849, S. T. Olney (G); Augusta, S. T. Olney (G); about Augusta, 
S. T. Olney & J. Metcalf (NY); Burge Co.: without stated local- 
ity, Sept. 10, 1897, H. Hopkins, 39 (NY). Wayne Co.: dry 
sandy ridge between Doctortown and Jesup, Sept. 14, 1903, 
R. M. Harper, 1997 (G, NY, US). FLORIDA. Watton Co.: 
between Freeport & Portland, Sept. 23, 1910, R. M. Harper, 90 
((G, NY (US, two plants to the left)). OxaroosA Co.: East 
Pass, Aug. 31, 1899, S. M. Tracy, 6362 (G, NY, ND, NC). 
Santa Rosa Co.: dry sandy pineland, Milton, Sept. 9, 1912, 
F. W. Pennell, 4571 (NY). OnawacE Co.: vacant lot, Orlando, 
Aug. 16, 1929, C. J. Williams (F); high pineland, Gotha, Sept. 2, 
1929, F. Vasku (F); pineland, Windermere, Aug. 27, 1929, F. 
Vasku (F); sandhill, Windermere, Sept. 10, 1929, F. Vasku (F); 


284 Rhodora [OcroBER 


high pineland, Windermere, Sept. 1, 1929, E. West (F); sandy 
woods, Windermere, Sept. 30, 1929, E. West (F); HILLSBOROUGH 
Co.: sandhills, Tampa, Sept. 17, 1904, A. Fredholm, 6392 ((G 
plant to left); Tampa, Oct. 1877, A. P. Garber (G, P (plant to 
right). ALABAMA. Covineron Co.: 1-2 mis. e. of Florala, 
Sept. 13, 1912, F. W. Pennell, 4631 (NY). LOUISIANA. Co. 
UNDETERMINED: 8. w. Louisiana, ez Herb. C. Mohr (US). 

Liatris pauciflora was described by Pursh as a glabrous plant, 
from a collection by Bartram in Georgia. Elliott, later described 
L. secunda as a reclining species, with short-pubescent stem, 
growing on the summits of dry sandhills and common near 
Columbia, South Carolina. This has generally passed into syn- 
onymy under L. pauciflora. Since, in his flora, Pursh made two 
divisions of the genus, tuberous and non-tuberous, and placed L. 
pauciflora in the latter group (DeCandolle doing likewise), it was 
judged by Nash (Bull. Torr. Bot. Club. xxii. 152 (1895)) that the 
description of this Pursh species was not that of a Liatris at all 
and that Liatris secunda Ell. was the legitimate name for L. 
pauciflora of Gray (Synop. FI. 1. c.). 

There is in the Banks Herbarium at the British Museum a 
specimen! of L. pauciflora Pursh, a photograph of which was ob- 
tained by Mr. Weatherby, labelled in pencil in the same hand- 
writing as is the type of L. gracilis Pursh, which would therefore 
seem to be the type of L. pauciflora Pursh. Examination of the 
photograph confirms the glabrous nature of the rachis of the 
inflorescence and a part of the stem, though much of the stem, 
the basal leaves and the underground stem are lacking (the last 
omission perhaps explaining the error Pursh made in classifying 
the plant). The heads are noticeably borne severally along 
slender, erect, peduncles 3-5 cm. long, thus making the inflores- 
cence a simple panicle. The heads and phyllaries are as described 
by Pursh. Specimens paralleling this type have been seen in her- 
baria mostly from the northern half of peninsular Florida. 

1 Though the type specimen bears no collector’s name, date or place of collection, 
in the recently published diary of John Bartram (Diary of a Journey through the 
Carolinas, Georgia, and Florida, July 1, 1765 to Apr. 10, 1766, John Bartram, anno- 
tated by F. Harper, Trans. Amer. Phil. Soc. n. s. xxxiii. pt. 1, 1-120 (1942)), mention 
is made of Serratula at Turtle Creek in Georgia, which would be Glynn Co., and very 
near the Florida border. Also in the report of William Bartram (Travels in Georgia 
and Florida, 1773-74, a report to Dr. John Fothergill, William Bartram, annotated 
by F. Harper, Trans. Amer. Phil. Soc. n. s. xxxiii. pt. 2, 121-242 (1943) reference to 
Serratula in Georgia is found twice: (1) Brier Creek, in Screven or Burke Co. and (2) 


the Ridge, referred to as south of the forking of the Tulagoo from the Savannah 1 
probably in Madison or Oglethorpe county, from the route shown on the map. 


1946] Gaiser,— Tbe Genus Liatris 285 


Though the type of L. secunda Ell. is unfortunately no longer 
available, there is at the Gray Herbarium a small envelope la- 
belled “L. secunda Ell. ex. Herb. Ell.", from which it is possible 
to see the nature of the puberulent stem and to make flower- 
measurements. The cylindrical head is 1.5 em. long and com- 
pact as that of L. punctata but with thin glabrous phyllaries, an 
achene only 4 mm. long, and the pappus not conspicuously 
plumose. The corolla when boiled is 12 mm. long, and has only 
a few scattered hairs within and without the tube. Specimens 
having similar characters are often reclining and the heads of the 
loose raceme often become secund. Herbaria show numerous 
specimens from the New Hanover and Brunswick coastal region 
of North Carolina, from South Carolina and Georgia but fewer 
from Florida and they are mostly from the western counties. 
The characters of the flower-parts of glaborus specimens growing 
in sandhills through the northern half of peninsular Florida that 
match the type of L. pauciflora are very similar to those of L. 
secunda but the plants are usually erect, the leaves are narrowly 
linear and the heads are frequently borne in a loose erect panicle. 
The only puberulent specimens seen from peninsular Florida 
came from Windermere, Gotha and Orlando, three very adjacent 
stations in Orange County, and one plant each of collections by 
Garber and Fredholm labelled as from Tampa and Hillsborough 
respectively. These two latter specimens might possibly have 
been obtained by the collectors elsewhere. Since the more 
northern specimens are persistently puberulent we are led to re- 
tain L. secunda Ell. as a separate species, as did Small, though 
not in synonymy with L. carinata Small (see no. 11). 

Confusion has resulted from failure to recognize the almost 
glabrous specimens from North and South Carolina of so-called 
L. carinata as resembling species of the Graminifoliae rather than 
the Pauciflorae series and in its given synonymy with L. secunda. 
As pointed out (see no. 11) L. regimontis, here including L. 
carinata, has a corolla only 7-8 mm. long and is quite pilose 
within at the base of the tube whereas the corolla of L. secunda 
is 10-12 mm. long and has only a few scattered hairs within and 
without. The length of the heads too is greater, 1.5-2 cm. long 
in L. secunda and L. pauciflora and only about 1 cm. in L. regi- 
montis. When these confusing glabrous specimens of the Caro- 


286 Rhodora [OCTOBER 


linas are thus classified as L. regimontis of the Graminifoliae, the 
Pauciflorae specimens of the same region all prove to be puberu- 
lent and a match for L. secunda Ell., just as the wholly glabrous 
specimens in Florida give confirmation to the species L. pauci- 
flora. No glabrous specimens from Georgia have been seen ex- 
cept the type plant. Though no locality was given by Pursh for 
Bartram’s plant from Georgia it could possibly have been col- 
lected very near the Florida border (see last footnote). 


That a glabrous species, L. pauciflora, should be generally 
found in a more southerly region than a related puberulent species 
of the same series, L. secunda, is in strong contrast to finding the 
rare hirsute form in the widespread species L. spicata, in some 
few plants of its variety resinosa from Florida and Louisiana. 
It is however in agreement with the condition obtaining in the 
series Tenuifoliae, where likewise the glabrous L. laevigata is 
limited to peninsular Florida, while the related L. tenuifolia, 
with a tendency to ciliate leaves, extends northward to South 
Carolina. 


Serres V. "TENUIFOLIAE. Herbs showing the tallest and 
most slender spikes of the genus, attaining a height of 12 dm., 
with basal rosettes of long filiform to linear coriaceous leaves, 
diminishing abruptly to short setaceous bracts; heads 4-6- 
flowered, ca. 1 cm. long, but not compact in appearance when 
mature due to the few narrow linear thin phyllaries that are not 
appressed when the flowers are open; corolla-tube non-pilose 
within but with short hairs on the filaments of the stamens; 
achenes 3-4 mm. long. 

From dry and moist lands from South Carolina to Big Pine 
Key, Florida. 


a. Plants with scattered hairs along stem; leaves dull, sparsely 
ciliate, very narrow, filiform; of more northern distribution 
15. L. tenuifolia. 
a. Plants completely glabrous; leaves glossy, coriaceous, non- 
ciliate and linear; peninsular Florida G L OTR 16. L. laevigata. 


15. LIATRIS TENUIFOLIA Nutt. Corm small, rounded, up to 2 
cm. in diameter: stems slender and spicate, 6-12 dm. high: leaves 
glabrous and punctate; lower ones long, linear, filiform, 10-25 cm. 
long, not more than 2 mm. wide, broadening at the point of 
attachment and often with scattered cilia at their bases, crowded 
into a rosette from which cauline leaves abruptly diminish to 
erect setaceous bracts 1 cm. or more long; heads 4—5-flowered, in 
a long, lax spike, long raceme or narrow panicle, sessile or on 


1946] Gaiser,—The Genus Liatris 287 


short, very slender pedicels 6-9 mm. long; phyllaries glabrous, 
the outer acute and only about half as long as the inner oblong- 
elliptic ones, 6-10 mm. long, with petaloid margin, frequently 
purple and with midvein often terminating in a short cusp; 
corolla phlox-purple, rarely white, 6-8 mm. long, smooth within, 
but filaments of the stamens with short hairs; achene ca. 3 mm. 
long, pappus 5-7 mm. long, barbellate, and not plumose to the 
naked eye; flowers said to be fragrant.'—Nutt. Gen. ii. 131 (1818); 
Ell. Sk. ii. 275 (18227). Laciniaria tenuifolia O. Ktze. Rev. Gen. 
Pl. i. 349 (1891). 

South Carolina to Florida and Alabama.—STATE UNDE- 
TERMINED.  Torrey (NY), Chapman, 7 (NY), Leavenworth 
(NY). SOUTH CAROLINA. Without stated locality: Hb. 
Gibbes (G, NY). CHESTERFIELD Co.: among scrub-oak and long- 
leaf pine, 1 mi. w. of McBee, Sept. 7, 1939, R. K. Godfrey, 8074 
(G, NY). DanriNGTON Co.: Society Hill, M. A. Curtis (G). 
RICHLAND Co.: sandy woods, near Columbia, Sept. 25, 1883, J. D. 
Smith, 2032 (US).  ArkEN Co.: Aiken, Sept. 12-15, 1909, W. W. 
Eggleston, 5045 (G, NY, US); dry scrub, Aiken, Sept. 1866, H. 
W. Ravenel (G); Aiken, Sept. 1869, H. W. R. (US). GEORGIA. 
Without stated locality: Boykin (G, NY). Ricumonp Co.: pine 
& blackjack hills, Summerville near Augusta, Sept. 21, 1883, J. D. 
Smith, 2033 (US); dry pine barrens, Augusta, A. Cuthbert (F); 
sandhills, high ridges, Augusta, Sept. 1899, A. Cuthbert, 312 (NY); 
sandhills, Augusta, Sept. 1899, A. Cuthbert, 1132 (F); sandhills, 
1 mi. n. of Mt. Lebanon Church, Oct. 17, 1937, J. H. Pyron & R. 
McVaugh, 1895 (US). Burge Co.: without stated locality, 
Sept. 15, 1897, M. H. Hopkins, 38 (NY). Screven Co.: oak 
ridge, in s. e. part of county, Sept. 29, 1940, D. Eyles, 7554 (G). 
Crisp Co.: sandy soil, Cordele, Sept. 18, 1901, Biltmore Herb., 
4116e (US). SuwrER Co.: high sandy banks of Flint R., Sept. 
10, 1900, R. M. Harper, 634 (G, NY, US). FLORIDA. With- 
out stated locality: Chapman (US, 4557 & 31492); Nuttall (P, 
isotype); Chapman, ex Herb. C. Mohr (US), ex Herb. Sartwell 
(US); Chapman (NY); Dr. Burroughs (NY); Curtiss, 188 (NY). 
Duvar Co.: dry pine barrens, Jacksonville, Oct. 24, 1894, A. H. 
Curtiss, 5310 (G, US); near Jacksonville, Oct. 13, 1893, A. H. 
Curtiss, 4447 (US); s. of Jacksonville, Aug. 1896, L. H. Lighthipe, 
339 (NY). CoruwsniA Co.: north of Camp Oleno, Oct. 6, 1940, 
W. A. Murrill (F). Hamittron Co.: dry woods on Suwannee R., 
White Springs, Sept. 30, 1941, E. West & Miss L. Arnold (F). 
Leon Co.: Tallahassee, N. K. Berg (NY); Tallahassee, Oct. 7, 
1895, Biltmore Herb., 576 (US). GapspEN Co.: dry soil, River 
Junction, Sept. 22, 1900, Biltmore Herb., 4116d (US); high pine- 
oak-woods near Havana, Oct. 6, 1940, W. A. Murrill (F). JACK- 
son Co.: without locality, Aug. 23, 1880, E. A. Smith (US). 


1 See introduction. 


288 Rhodora [OCTOBER 


FRANKLIN Co.: dry pine barrens, Apalachicola, 1867, B. F. 
Saurman (Q). Bay Co.: sandy moist open ground, Lynn Haven, 
Oct. 12, 1921, C. Billington (US). Watron Co.: dry sandy soil, 
Argyle, Aug. 25, 1899, Biltmore Herb., 4116b (US). SANTA 
Rosa Co.: dry sandy pineland, Sept. 9, 1912, F. W. Pennell, 
4575 (NY). Aracnua Co.: dry fields, Gainesville, Oct. 7, 1928, 
G. F. Weber (F); flatwoods, Gainesville, Oct. 14, 1927, Miss L. 
Arnold. (F); open fields, Archer Road, Gainesville, Oct. 12, 1931, 
Miss L. Arnold (F). GincHmisr Co.: dry woods, 6 mis. e. of 
Trenton, Oct. 5, 1940, E. West & Miss L. Arnold (F). TAYLOR 
Co.: 9 mis. s. of Perry, Oct. 8, 1940, W. A. Murrill (F). Marion 
Co.: 4 mis. s. of Belleview, Oct. 13, 1940, W. A. Murrill (F). 
Levy Co.: flatwoods, 5 mis. e. of Otter Creek, Nov. 11, 1939, 
Watson & W. A. Murrill (F). VorvusiA Co.: (typical and albino) 
dry soil, pine woods, s. of New Smyrna, Oct. 14, 1944, Mrs. H. T. 
Butts (OA). OnaNaE Co.: Lake Mary, Nov. 17, 1902, S. M. 
Tracy (US); pinelands, Gotha, Aug. 22, 1929, E. West (F); flat- 
woods, Orlando, Oct. 9, 1929, F. Vasku (F); (albino) flatwoods, 
Orlando, Oct. 22, 1929, De Vane & West (F). Laxe Co.: 5 mis. 
s. w. of Astor Park, Oct. 13, 1940, W. A. Murrill (F). CITRUS 
Co.: sandy dry oak-pine-woods, on U. S. 41, 1 mi. n. of Inverness, 
Oct. 16, 1945, H. H. Hume (F). Pork Co.: Haines City, Nov. 
1917, R. H. Young (US); Fort Meade, April 4, 1880, J. D. Smith 
(US); sandy place, July 27, 1940, P. O. Schallert (G). Hirrs- 
BOROUGH Co.: Tampa, Oct. 1877, A. P. Garber (G, US); dry 
sand, Oct. 18, 1904, A. Fredholm, 6422 (G). MaNarEE Co.: 
Bradentown, Aug. 10, 1900, S. M. Tracy, 7100 (NY, T); flatwoods, 
Bradentown, Oct. 30, 1916, Oct. 27, 1920, A. Cuthbert (F). 
Dave Co.: in tropical pineland, Miami, Nov. 28, 1933, F. 
Duckett, 242 (US); Buena Vista, Jan. 21, 1930, H. N. Moldenke, 
483a (NY); w. of Rockdale, Nov. 18, 1916, J. K. Small, 7907 
(NY). ALABAMA. Without stated locality: ex. Herb. G. 
Thurber (G). CHEROKEE Co.: dry soil, dry ridges, near Bristol, 
Oct. 25, 1895, C. Mohr (US). Covinaton Co.: Blue Springs, 
near McRae, Sept. 13, 1912, F. W. Pennell, 4616 (NY). 


See discussion following the next species, L. laevigata Nutt. 


16. LIATRIS LAEVIGATA Nutt. Corm stout, globular or some- 
what broadened, larger than that of L. tenuifolia, up to 4 cm. in 
diameter; stems shorter, frequently not attaining the extreme 
height of that species: lower leaves entirely glabrous, punctate, 
involute, long, linear, but not filiform, 2-3 dm. long, 2-8 mm. 
wide, broadening and sheath-like at the point of attachment, 
shining, coriaceous, in a rosette at the base, diminishing abruptly 
to short erect setaceous glabrous bracts: spike sometimes be- 
coming panicled, 3-6 dm. long; heads 4-6-flowered, usually 
sessile or on slender pedicels 6-9 mm. long; outer phyllaries short, 


1946] Gaiser,— The Genus Liatris 289 


narrowly acute: the inner oblong, elliptic, 6-10 mm. long, often 
with obtuse petaloid tips and frequently purple; corolla phlox- 
purple, 7-8 mm. long, smooth internally but filaments of sta- 
mens with short hairs; achenes ca. 3 mm. long; pappus 5-7 mm. 
long, barbellate, and not plumose to the naked eye.—Nutt. 
Trans. Amer. Phil. Soc. n. s. vii. 285 (1841). Liatris tenuifolia 
& Torr. & Gray, Fl. N. Am. ii. 70 (1841). Liatris tenuifolia 
var. laevigata (Nutt.) Robinson, Proc. Amer. Acad. Arts and Sci. 
xlvii. 201 (1911). Liatris tenuifolia var. quadriflora Chapm. Fl. 
S. U. S. ed. 2, Suppl. 626 (1883). Lacinzaria laevigata (Nutt.), 
Small, Fl. S. E. U. S. 1175 (1903). 

Found only in peninsular Florida, along the coast as far south 
as Big Pine Key.—FLORIDA. Without stated locality: Nov. 
26, J. Read (P, isotype). Duvar Co.: vicinity of Mayport and 
Jacksonville, 1870-76, H. D. Keeler (NY, ND); St. Nicholas, 
Oct. 1897, L. H. Lighthipe (N Y), dry pine barrens, near Jackson- 
ville, Oct., A. H. Curtiss, 1174 (US, 63074); near Jacksonville, 
1893, A. H. Curtiss, 4447 (NY); dry pine barrens, near Jackson- 
ville, Oct. 7, 1898, A. H. Curtiss, 6289 (F). Baker Co.: Glen 
Saint Mary, Oct. 1927, H. Hume (G). FrRanxKuINn Co.: Apa- 
lachicola, Biltmore Herb., 4116a (NY, Q). SaiNT Jouns Co.: 
flatwoods, Road 14A, near Spuds, Oct. 14, 1941, E. West & Miss 
L. Arnold (F). Cray Co.: cut-over pine woods, Keystone 
Heights, Oct. 11, 1945, H. Hume, Nevins & Miss L. Arnold (F). 
Putnam Co.: scrubby flatwoods, n. of Beecher Springs, Welaka, 
Sept. 28, 1940, A. M. Laessle (F). ArAcHvA Co.: south of 
Prairie, Alachua, Oct. 24, 1940, W. A. Murrill (F). FLAGLER 
Co.: flatwoods, 5 mis. e. of Co. line, Hwy. 28, Andalusia, Oct. 10, 
1940, E. West & Miss L. Arnold (F). Marton Co.: in a scrub, 
Ocala National Forest, Sept. 12, 1929, H. O'Neill (US, F). 
VoLUsiA Co.: pine wood, dry sandy soil, s. of New Smyrna, Oct. 
14, 1944, Mrs. H. T. Butts (OA). SeMiNoLE Co.: high pinelands, 
Sanford, Sept. 22, 1927, O. F. Burger & E. West (F). Laxe Co.: 
Eustis, Aug. 16-25, 1894, G. V. Nash, 1669 (G, NY, P, ND, US); 
Eustis, Sept. 10, 1895, G. V. Nash, 2599 (G, NY, ND, US); 
sandy soil, Eustis, Sept. 10, 1900, Biltmore Herb. 4116e (US); 
open sand, 5 mis. e. of Leesburg, Aug. 17, 1939, W. A. Murrill 
(F). OranGeE Co.: Clarcona, Sept. 25, 1899, Miss M. Meislahn, 
72a (US); high pineland, Wekiwa Springs, Sept. 25, 1929, H. 
O'Neill (US); flatwoods, Gotha, Aug. 30, 1929, F. Vasku (F); 
pineland, Windermere, Sept. 3, 1929, F. Vasku (F); sandhills, 
Windermere, Sept. 10, 1929, F. Vasku (F). BmEvAnD Co.: pine 
barrens, Indian River Region, Nov. 9, 1902, A. Fredholm, 5565 
(G); sandy soil, Cocoa, Sept. 5, 1936, A. S. Rhoads (F). OSCEOLA 
Co.: low pinelands, Deer Park, Sept. 24, 1927, O. F. Burger & E. 
West (F). HrinLsBonovuaGH Co.: flatwoods, w. of Plant City, 
Aug. 19, 1945, L. O. Gaiser, Mrs. H. T. Butts & Miss L. Arnold 


290 Rhodora [OcroBER 


(F). OkEECHOBEE Co.: Okeechobee, Sept. 26, 1903, A. Fredholm, 
6022 (G); pine woods, Okeechobee, Oct. 12, 1941, J. H. Davis 
(F). Manatee Co.: Bradentown, Aug. 10, 1900, S. M. Tracy, 
7100 (G, US). Marrin Co.: pine flatwoods, w. of Stuart, Nov. 
23, 1942, J. H. Davis (F). Sarasora Co.: airfields in flatwoods, 
Venice, Aug. 18, 1945, L. O. Gaiser, Mrs. H. T. Butts & Miss L. 
Arnold (F). CuaARLorTE Co.: flatwoods ditch, s. of Punta Gorda, 
Aug. 18, 1945, L. O. Gaiser, Mrs. H. T. Butts & Miss L. Arnold 
(F). Lee Co.: Marco, A. S. Hitchcock, 154 (G, NY, US). 
COLLIER Co.: golf course, Naples, Nov. 2, 1939, Miss E. Scull 
(F). Dave Co.: pinelands, s. of Miami R., Nov. 26, Dec. 20, 
1913, J. K. Small & G. K. Small, 4791 (NY); between Miami & 
Kendall Sta., Nov. 5, 1906, J. K. Small & J. J. Carter, 2752 (NY); 
pinelands, s. of Miami R., Nov. 20, 1912, J. K. Small, 3858 & 
3848 (NY); pinelands, Miami, Oct. 28-Nov. 28, 1903, J. K. Small 
& J. J. Carter, 534 (NY, P); between Cocoanut Grove & Cutler, 
Oct. 31, Nov. 4, 1903, J. K. Small & J. J. Carter, 1457 (NY); 
Cocoanut Grove, Nov. 2-5, 1901, J. K. Small & G. V. Nash, 184 
(NY); pinelands, Cutler, Mar. 27, 1904, N. L. Britton, 287 (NY); 
Homestead, Sept. 16, 1927, S. Hawkins (F); pinelands, Big Pine 
Key, Dec. 2, 1912, J. K. Small, 3966 (NY); Pine Key, J. L. 
Blodgett (N Y); Long Pine Key, Aug. 25, 1937, Miss E. Scull (F). 

Nuttall (Gen. ii. 131 (1818)) described L. tenuifolia as having 
“leaves almost like those of Pinus palustris but flat and linear", 
and as being 2-4 feet tall with a long raceme of 1-2 feet (a photo- 
graph of his type at the British Museum, received through Mr. 
Weatherby, shows a stem about 6 feet tall). He gave the habitat 
as sandy forests of North and South Carolina. By comparison 
of specimens in the herbaria with this photograph of the type, 
this species does not seem to have been collected in North Caro- 
lina, but from South Carolina to Florida and westward through 
Alabama. Chapman (Fl. S. U. S. ed. 2, Suppl. 626 (1883)) de- 
scribed L. tenuifolia var. quadriflora from the banks of the Caloosa 
River, S. Florida, as having rigid, involute leaves and those of 
the upper stem setaceous. 

In 1841, Nuttall (Trans. Amer. Phil. Soc. vii. 285 (1841)) 
described as a new species Liatris laevigata: “with almost filiform, 
subulate leaves; radical ones nearly a foot long, pungently acute 
and coriaceous”. Examination of a photograph of the type 
specimen in the British Museum shows a plant only a little more 
than one foot high and with few basal broader leaves, that seems 
to include the concept of Chapman’s var. quadiflora, and that is 
represented in the herbaria by specimens from along the Florida 


1946] Gaiser,—The Genus Liatris 291 


coast to Big Pine Key. Although Nuttall referred to L. tenui- 
folia as being ‘‘everywhere smooth" and L. laevigata as "in every 
part very smooth", close study of the photograph of the former 
type shows some scattered cilia at the base of the leaves. Such 
a presence of cilia, in varying degrees, seems to be a constant 
accompaniment to the finely linear leaves of specimens from 
South Carolina, Georgia and Alabama. Plants wholly glabrous, 
having glossy, involute, almost quill-like leaves varying in width 
from 2 to 8 mm., are found only in peninsular Florida. It seems 
possible too that these latter may be more moisture-loving as 
they occur often in low pinelands and in flatwoods where the 
water level may be high. 

Torrey & Gray (Fl. N. Am. ii. 70 (1841)) treated L. laevigata 
as a variety of L. tenuifolia and this was followed in Gray, Synop. 
Fl. i?. 112 (1884), and in Proc. Amer. Acad. Sci. xlvii. 201 (1911), 
where Robinson described it as a conspicuously more robust vari- 
ety and referred to specimens of Mr. G. V. Nash, no. 1669 and no. 
2599, both from Eustis, Florida, as examples, and to that of Prof. 
Hitchcock, no. 154, also from Marco, Florida, as representing an 
intergradation between the more slender and more robust forms. 
The diameters of the corms of these three specimens, represented 
in three herbaria visited (G, NY, US), vary slightly, but all range 
between 18 and 38 mm. in diameter, thus coming well within the 
general measurements for L. laevigata. In collections from 
Florida there is evidence that the two types occur side by side as 
selections from the lists of specimens will show: i 

MANATEE Co.: Bradentown, S. M. Tracy, 7100, ((G, US. 
resembles laevigata), (NY, T, resembles tenuifolia)); FRANKLIN 
Co.: Apalachicola, Biltmore Herb. 4116a ((G, US, resembles 
tenuifolia), (NY, Q, resembles laevigata)); Duvar Co.: Jackson- 
ville, A. H. Curtiss, 1174 ((US 3074, resembles laevigata), (US 
63073 & G, seem intermediate)). 

Other specimens from Florida give evidence of intergradation 
between the two, as Robinson stated, in having leaves of inter- 
mediate width, or having some cilia at the base of the leaves, as 
well as showing intermediacy of stoutness of above- and below- 
ground stems. Some of these are listed below: 

Without stated locality: Chapman (ex Herb. G. Thurber (G) 


& 449, 450 (US) & US 63076, US 968368); 1842-1844, F. Rugel 
(US). ManatEr Co.: Manatee, J. H. Simpson (US). DvvaAL 


292 Rhodora [OCTOBER 


Co.: Jacksonville, Nov. 1891, W. G. Farlow (G); Jacksonville, 
Oct., A. H. Curtiss, 1174 (G). Dape Co.: Miami, Nov. 28, 1933, 
F. Duckett, 242 (G). Guur Co.: St. Vincent Isl., Nov. 2, 1910, 
W. L. McAtee, 1810 (US). Voxrusra Co.: s. of New Smyrna, 
Oct. 14, 1944, Mrs. H. T. Butts (OA). 

However, the great interest lies in the fact that all the inter- 
mediates come from Florida where both species have their great- 
est (and L. laevigata its exclusive) distribution. Mrs. H. T. 
Butts has collected from one location, south of New Smyrna, 
both L. laevigata and tenuifolia (including an albino specimen), 
as well as what has been here interpreted as an intermediate. 
Yet in the dry scrub of South Carolina or the sandy hills of 
Georgia the more slender L. tenuifolia alone seems to be repre- 
sented. Thus because of the limitations in habitat of L. laevigata 
as well as the quite different appearance of leaf and plant, they 
are regarded as separate species that intergrade in Florida where 
their ranges overlap. 

X Lrarris Boyxini Torr. & Gray, emend. (L. elegans X tenui- 
folia). Stem nearly glabrous, slender, 3-6 dm. tall: leaves 
rather scattered, linear, lower elongated, upper short and seta- 
ceous: spike 1.5-2.5 dm. long, of rather crowded, subsessile or 
shortly pedicellate, 3-4-flowered heads; phyllaries glabrous, the 
outer ones short, lanceolate-subulate, the interior lanceolate or 
linear, with scarious margins and acuminate spreading summits, 
surpassing the flowers in length; flowers pale purple; corolla 9 
mm. long, tube as well as filaments without any hairs; pappus 7 
mm. long, plumose; achene ca. 4 mm. long, villous.—Fl. N. Amer. 
ii. 70 (1841). 

GEORGIA. Muscogee Co.: near Columbus, Dr. Boykin 
(G, NY, TYPE). SUMTER Co.: along the high, sandy bank of 
Flint River, Sept. 10, 1900, R. M. Harper, 635 (NY, US). 

L. Boykinii was described by Torrey & Gray from a plant col- 
lected near Coumbus, Georgia, by Dr. Boykin (G, NY). Later 
Gray (Synop. Fl. i'. 110 (1884)) again included it, referring only 
to the single collection and stating: “not since found". Heads of 
this species were said to be larger than in L. tenuifolia and rather 
smaller than those of L. secunda. Small, Man. S. E. Fl. 1333 
(1933), states: “perhaps a hybrid between L. elegans and L. 
tenuifolia”. In 1900, R. M. Harper collected specimens, no. 635, 
along the high sandy bank of the Flint River, Sumter Co., 
Georgia (NY, US), with L. elegans and L. tenuifolia, noting on 
one sheet that the plants were intermediates between the two 


1946] Gaiser,—The Genus Liatris 293 


species. Examination of these specimens shows the upper leaves 
very narrow and linear, quite like those of L. tenuifolia. The 
heads, 4-flowered, ca. 10 mm. long, have outer phyllaries that 
are narrowly lanceolate and inner ones with free spreading, pink, 
petaloid, prolonged tips suggesting the bracts of L. elegans. In 
numbers of flowers per head, in characters of corolla-tube, pappus 
and achene, the Harper collection bears a resemblance to Dr. 
Boykin’s plant and it seems probable that these two collections 
from Georgia, where both L. tenuifolia and L. elegans occur, repre- 
sent intermediates between the two species. During this inves- 
tigation no other specimens have been found. 


Series VI. Scanrosar. Plants with stiff, robust stalks of 
the inflorescence bearing few to numerous large, campanulate, 
hemispheric or subglobose heads in loose open spikes or panicles; 
leaves mostly lanceolatc, rarely oblanceolate; heads 15-70- 
flowered; phyllaries broad, orbicular, spatulate or obovate, mostly 
squarrose or bullate before the opening of the flowers; corolla- 
tube pilose within (except in L. ligulistylis) ; achene 3-7 mm. long. 
—From along the coast in New England, southward to Georgia, 
westward through Tennessee and Kentucky, to become wide- 
spread from Michigan and Wisconsin southward to Texas and 
Oklahoma and westward from the prairie provinces along the 
Rocky Mts. to New Mexico. 


a. Heads short-cylindrical to globose... .b. 
b. Heads subglobose, 25-50-flowered; phyllaries squarrose or 
bullate from before the time of opening of flowers (except 
in L. scariosa var. virginiana)... .c. 
c. Phyllaries herbaceous, ciliolate, with only very narrow 
üf any) searious margin, pubescent to rough; outer 
ones markedly and middle ones moderately squarrose; 
leaves and stem pubescent, basal leaves broadly 
obovate oi il ENG es va wie cece eure ge pee se 17. L. scariosa. 
c. Phyllaries thin, glabrous, broadly scarious, erose and 
colored, all markedly bullate; stem and leaves asperous 
or glabrous; leaves linear to linear-lanceolate........ 18. L. aspera. 
b. Heads short-cylindrical to subglobose, 25-40-flowered; 
phyllaries mostly erect, appressed and herbaceous, never 
bullate and erose, though outer ones sometimes recurved.19. L. scabra. 
a. Heads campanulate to hemispheric; phyllaries erect and 
loosely appressed through the maturing of the flowers. . . .d. 
d. Heads hemispheric, 25-70-flowered. . . .e. 
e. Leaves few, glabrous to densely pubescent but scabrous 
to the touch along the margin; basal leaves broadly 
lanceolate, reduced abruptly upwards to linear 
bracts; phyllaries broadly spatulate with deep scari- 
ous, erose and colored margins; inflorescence of few 
to 20 heads; corolla-tube non-pilose within.....20. L. ligulistylis. 


294 Rhodora [OcroBER 


e. Leaves numerous, glabrous or with but scattered pubes- 

cence on the lower surface and along the margin, 

often twisted, all linear-lanceolate; phyllaries oblong, 

herbaceous, hardly at all scarious but finely ciliolate 

on the margin; inflorescence of usually more than 20 
| MO MMC Des: 21. L. borealis. 

d. Heads campanulate, 15-25-flowered; leaves lanceolate, 

with upper ones sometimes linear, glabrous to asperous; 

phyllaries oblong to narrowly spatulate, mostly herba- 

ceous and light green, with very narrow (if any) mem- 
branous margins, outer ones sometimes squarrose....22. L. Earlei. 


17. Liarris scartosa (L.) Willd. Corm rounded, up to 5 cm. 
in diameter: stems one to several, usually with dense semi- 
appressed pubescence, 4-8 dm. high: leaves not numerous, from 
sparingly to densely pubescent, even to scabrous to the touch on 
both sides; basal leaves 0.8-1.5 dm. long, and 2-5 cm. wide, 
broadly oblanceolate, oblong to almost obovoid, narrowing to 
clasp the stem; upwards the bluntly oblanceolate leaves shorten- 
ing through narrower ones 4-7 em. long and 0.5-0.7 cm. wide, 
to braets subtending the heads: inflorescence of 15-30 almost 
globular heads in an open raceme or occasionally panicle: heads 
15-50-flowered, subglobose, 1.5-2.5 cm. in diameter, on short 
pedicels or occasionally longer ones forming a panicle-like 
inflorescence; phyllaries rather leathery and mostly recurved 
(except in var. virginiana), the outermost ones ovate and soon 
recurved, the middle and inner ones rounded at the tips, also 
somewhat recurved, mostly herbaceous, sometimes showing 
color, with a very narrow, thinner, ciliolate margin; corolla 
purple, pilose in the base of the tube, 7-11 mm. long; pappus 
6-9 mm. long; achene 4-5 mm. long.—Spec. Pl. iii. 1635 (1803). 
Serratula scariosa Linn. Sp. Pl. ii. 818 (1753). 

Mostly in the mountains from southern Pennsylvania to the 
border of South Carolina but one variety from the plains in the 
Carolinas, Georgia and Alabama. 


KEY TO VARIETIES 


a. Phyllaries herbaceous, the outer ones soon recurved, the mid- 
dle and inner ones somewhat recurved; basal leaves broadly 
obovate, the upper ones oblanceolate... .b. 
b. Heads of 25-50 flowers. ..........uuuuuuuuuuuuuuuueee. var. typica. 
b. Heads of 15-25 flowers.............. 0.00000 ccc cues var. squarrulosa. 
a. Phyllaries mostly herbaceous or with narrow membranous 
margin, the outer ones but slightly if at all squarrose, mid- 
dle and inner ones loosely erect; basal leaves broad- to long- 
mam EE TITIUQUSTTCRETM EET ccebns cave tenert var. virginiana. 


Var. typica. Stem, leaves and phyllaries as described for the 
species: heads large, 25-50-flowered: corolla-tube 10-11 mm. 
long; pappus 8-9 mm. long; achene ca. 5 mm. long.—Serratula 
scariosa L. Sp. Pl. 818 (1753), sens. strict., with plant of Linnaean 
herbarium as type, not that of Gronovius. 


1946] Gaiser,—The Genus Liatris 295 


In the Appalachian mountains from southern Pennsylvania 
through North Carolina —PENNSYLVANIA. Co. unde- 
termined: Mts. Alleghany, Rafinesque (P). Perry Co.: Upper 
Henry Valley, Sept. 5, 1920, W. L. Abbott (P). FRANKLIN Co.: 
Mercersburg, ex. Detwiler Herb., 17-1 (P); Blue Ridge Summit, 
1886, E. Tatnall (G). Futton Co.: McConnelsburg, Sept. 
1907, C. S. Williamson (P), Sept. 1907, E. B. Bartram, collected 
by W. S. (NY); Tonolloway Creek, Sept. 20, 1870, E. L. Ten- 
brook (P). MARYLAND. Battimore Co.: without stated 
locality, Aug. 1886, G. L. S., 1176 (G). ALLEGHANY Co.: Cum- 
berland, Sept. 12, 1910, J. E. Harned (US, 648416, -17); moun- 
tainside, near Cumberland, Sept. 1934, W. Rhoades (G); s. of 
Cumberland, Sept. 8, 1926, E. S. Steele, 97 (G). VIRGINIA. 
FREDERICK Co.: shale near Dehaven, Sept. 15, 1940, F. W. Hun- 
newell, 16947 (G). Loupon Co.: opposite Point of Rocks, Sept. 
12, 1935, W. R. Mazon, 10770 (US). Warren Co.: dry woods, 
near Bentonville, Sept. 7, 1938, F. W. Hunnewell, 15762 (G). 
Pace Co.: vicinity of Blue Ridge, Stony Man Mt. near Luray, 
Sept. 2, 1901, E. S. & Mrs. Steele, 224 (US, 418571). FAIRFAX 
Co.: Difficult Run, Sept. 29, 1904, W. Palmer (US), Sept. 18, 
1899, E. S. Steele (US); pike near Difficult Run, Sept. 30, 1911, 
E. S. Steele (G); pike near Difficult Run, Oct. 6, 1907, E. S. 
Steele (G, US); near mouth of Difficult Run, Sept. 25, 1909, F. W. 
Pennell (US); pike near Difficult Run, Great Falls, Oct. 2, 1910, 
F. W. Pennell (US). SuENANDoAH Co.: Massanutten Mts., s. 
end of Short Mt., steep dry shales near roadside, Aug. 19, 1938, 
H. A. Allard, 5466 (G); low hills, s. of hotel, vicinity of Orkney 
Springs, alt. 450 m., Sept. 11, 1911, E. S. Steele, 121 (US, 1521106) ; 
low hills s. of hotel, vicinity of Orkney Springs, alt. 480 m., 
Sept. 11, 1911, E. S. Steele, 126 (G); 127 (US); on true shale 
barrens, at foot of Pugh's Run, n. of Woodstock, Massanutten 
Mt., Sept. 22, 1940, H. A. Allard, 8228 (G). HiaHraAND Co.: 
shale barrens, Shenandoah Mt., Staunton-Monterey Road, 
Sept. 9, 1934, Miss E. S. Rawlinson 269 (US). Brann Co.: 
East River Mt., in rocky places, Sept. 1, 1931, E. L. Core, 6841 
(NY). RAPPAHANNOCK Co.: 6 mis. w. of Sperryville, Sept. 21, 
1905, Mrs. E. P. Miller (US590496, -7). WEST VIRGINIA. 
Moraan Co.: Largent, Aug. 25, 1933, E. J. Alexander, T. H. 
Everett, S. D. Pearson (NY). HaMrsninE Co.: dry sandy woods, 
Cold Spring Gap, North Mt., Sept. 6, 1936, F. W. Hunnewell, 
14430 (G). Harpy Co.: Bean Settlements, Sept. 27, 1930, 
W. M. Sharp (G). GREENBRIER Co.: White Sulphur Springs, 
Sept. 7, 1906, E. S. Steele (US 494223). NORTH CAROLINA. 
Durnam Co.: open rocky ground, near Eno R., H. L. Blomquist, 
10518 (P); edge of ditch, Duke Forest, Oct. 1, 1932, H. L. Blom- 
quist, 439 (US). 


296 Rhodora [OcroBER 


Var. SQUARRULOSA (Michx. Gray. Differing from var. 
typica in slenderer habit, with leaves and heads smaller: heads 
15-25-flowered, ca. 1 cm. high and 1-1.5 em. thick when the 
flowers are open; corolla-tube ca. 7 mm. long; pappus 5-6 mm. 
long; mature achene 4 mm. long.—Synopt. Fl. i?. 110 (1884). 
L. squarrulosa Michx. Fl. Bor.-Am. ii. 92 (1803); Shinners, Am. 
Midl. Nat. xxix. 33 (1943).  Laciniaria scariosa var. squarrulosa 
Small & Vail. Mem. Torr. Bot. Club, iv. 28 (1894), in part. 

Piedmont and Coastal plain of N. and 8. Carolina, Georgia and 
Alabama.—NORTH CAROLINA. Duruam Co.: Pont Rock, 
Aug., 1896, C. S. Williamson (P) ; open, rocky ground, north of Eno 
R., old Oxford Road, from Durham, Sept. 24, 1928, H. L. Blom- 
quist, 10578 (F, P). SOUTH CAROLINA. BxnkkLEY Co.: dry, 
rich soil, Santee Canal, H. W. Ravenel, 3 collections with slightly 
differing data (G). ArikEN Co.: Aiken, Ravenel (NY). GEOR- 
GIA. WiLKEs Co.: without locality, 1833, Herb. J. A. Lowell (G). 
McDvrrrg Co.: pine-barrens, vicinity of Thomson, Oct. 10, 
1910, H. H. Bartlett 2420 (G, US). Ricumonp Co.: dry barrens, 
Aug. 1876, and oak woods near pool, Oct. 10, 1898, Augusta, A. 
Cuthbert (F). JEFFERSON Co.: without locality, Sept. 25, 1897, 
H. Hopkins 40, 41 (NY). ALABAMA. Marssal Co.: 
rocky woodland, Albertsville, Oct. 9, 1900, Biltmore Herb. 
2670a (US). 

Var. virginiana (Lunell) comb. nov.—Similar in size, habit and 
floret-characters to var. typica but differing in the somewhat 
reduced more lanceolate leaves and the mostly erect membra- 
nous-margined phyllaries: basal leaves broadly lanceolate, 7-12 
em. long, 2-3 em. wide, narrowed to a winged petiole of about 
one third the length of the blade, upper ones gradually shortened 
and non-petiolate; heads 20-30-flowered (sometimes up to 50), 
ca. 1.5 cm. tall and wide, turbinate to hemispherical by reason of 
the erect phyllaries; outer phyllaries short, ovate, herbaceous 
with fine ciliolate margin, sometimes slightly recurved: middle 
ones longer, loosely erect, herbaceous and ciliolate-margined for 
the most part but with rounded tips, narrowly erose and usually 
purplish.—Laciniaria scariosa var. virginiana Lunell, Amer. Mid. 
Nat. ii. 172 (1912). Laciniaria scariosa var. borealis Lunell, 
ibid 264, probably Cirsium non ramosum . . . flores ferens 
pauciores majores . . . of Gron. Virg. i. 92 (1739). 

Mostly in the mountains from southern Pennsylvania to South 
Carolina.—PENNSYLVANIA. Centre Co.: without stated 
locality, Sept. 5, 1868, J. T. Rothrock (G). LxniaH Co.: on 
roadside embankment at edge of woods, 1 mi. s. w. of Schnecks- 
ville, Sept. 30, 1917, J. W. Pretz, 9171 (US). HuwTINGTON Co.: 
dry wooded shaley hillside, 2 mis. n. w. of Petersburg, Sept. 21, 
1941, H. A. Wahl, 1163 (G). CnuEesrER Co.: without stated 
locality, Aug. 1858-64, S. P. Sharples (G). Bxpromp Co.: 


1946] Gaiser,—The Genus Liatris 297 


wood road, 14% mis. n. w. of Breezewood, alt. 1060’, Aug. 3, 
1940, D. Berkheimer, 2178 (G). MARYLAND. ALLEGHANY 
Co.: Cumberland, Sept. 12, 1910, J. E. Harned (US 648418). 
GannETT Co.: Mountain Lake Park, Sept. 2, 1906, J. J. Carter 
(NY), Aug. 30, 1906, C. D. Lippincott (N Y), Aug. 26, 1928, E. S. 
Steele, 3 (US 14860-45, -49) ; vicinity of Oakland, Mountain Lake 
Park, E. S. Steele, Sept. 4, 1910 (No. 84 US 64857-1, -2), Sept. 7, 
1910 (No. 60 US 648566), Sept. 7, 1910 (No. 21 US 6485-61 to 
-70 excluding -66), Sept. 16, 1910 (No. 61 US 64857-3, -4); 
scrubby ground toward Deer Park, Aug. 25, 1921, E. S. Steele, 
150 (G); low open ground, Rwy. e. of Mountain Lake Park, Aug. 
9, 1921, E. S. Steele, 118 (G). WEST VIRGINIA. Preston 
Co.: grassy ground n. of Lake, Terra Alta, Sept. 4, 1920, E. S. 
Steele, 299 (US 1117671); flat ground n. of lake, Terra Alta, Aug. 
31, 1923, E. S. Steele, 166 (US 128604-1, -2); n. side of lake e. of 
camp, vicinity of Terra Alta, Aug. 18, 1924, E. S. Steele (US 
1326616); near camp n. of lake, vicinity of Terra Alta, Sept. 11, 
1924, E. S. Steele, 114 (US 13263-39, -40); roadside toward 
quarry, Terra Alta, Sept. 9, 1925, E. S. Steele, 78 (G, US 148603-6, 
-7, -8, -9, -40, -43, -44); Boys’ Camp n. of lake, vicinity of Terra 
Alta, Aug. 26, 1926, E. S. Steele, 68 (G). "TuckEn Co.: Canaan 
Valley, W. V. U. Biol. Exped. (G). GREENBRIER Co.: White 
Sulphur Springs, Aug. 27, 1903, K. K. Mackenzie 363 ((I. type), 
NY, US, (G, without no.)); mt. north of Springs (alt. 2000'— 
3000^, White Sulphur Springs, Sept. 7, 1906, E. S. Steele (G, US); 
Kate's Mt., White Sulphur Springs, Sept. 4, 1920, Miss M. S. 
Franklin (G). Monroe Co.: Peter's Mt., Aug. 31, 1903, E. S. 
Steele & Mrs. Steele (US, 490324); vicinity of Old Sweet Springs, 
E. S. Steele, Sept. 11, 1908 (US 648302), Sept. 9, 1905 (US, 
590189), Sept. 2, 1905 (US, 590187); ridge of Peter's Mt., on 
State line, Sept. 12, 1905, E. S. Steele (G, US, 590190); Peter's 
Mt., vicinity of Old Sweet Springs on Va. & W. Va. line, Sept. 
11, 1908, E. S. Steele (G, US 63531). VIRGINIA: Warren Co.: 
Little Passage Creek, Sept. 21, 1897, G. S. Miller (US). PAGE Co.: 
Stony Man Mt., Aug. 11, 1901, W. Palmer, 61 (US); crevices of 
rocks, Stony Man Mt., Aug. 18, 1901, W. Palmer & W. H. King, 61 
(US); exposed cliffs, Stony Man Mt., near Luray, Aug. 31, 1901, 
E. S. Steele & Mrs. Steele, 224 (G, NY, US); vicinity of Blue Ridge, 
Stony Man Mt., near Luray, Sept. 2, 1901, E. S. Steele & Mrs. 
Steele 224 (G, NY); near Luray, Sept. 18, 1905, G. S. Miller (US). 
SHENANDOAH Co.: brushy slope, Great North Mt., vicinity of Ork- 
ney Springs, Sept. 4, 1911, E. S. Steele, 64 (G), 62 (G, US); Great 
North Mountain, vicinity of Orkney Springs, Sept. 14, 1911, E. S. 
Steele, 141 (G (US, 1520733)) ; low hills s. of hotel, vicinity of Ork- 
ney Springs, Sept. 5, 1911, E. S. Steele, 74 (G); vicinity of Orkney 
Springs, Sept. 4, 1911, E. S. Steele, 69 (US 609901-2); on red 
sandstone barrens, on top of Great North Mt., Sept. 19, 1937, 


298 Rhodora [OcroBER 


H. A. Allard, 3765 (G). Barn Co.: sandstone soil, Mill Mt., 
vicinity of Millboro, Aug. 20, 1906, E. S. Steele (G); vicinity of 
Millboro (alt. 485 m.), E. S. Steele, Sept. 3, 1906 (G, US 648421), 
Sept. 11, 1906 (G, US 494573); on shale in vicinity of Millboro, 
Sept. 3, 1906, E. S. Steele (US, 494572, 648301). RockBRIDGE 
Co.: Mill Mt., vicinity of Millboro, Sept. 16, 1907, E. S. Steele 
(US, 494571). Cnaira Co.: s. of Peter's Mt., vicinity of Orkney 
Springs, Sept. 12, 1905, E. S. Steele (US, 590188); Peter's Mt., 
Sept. 1, 19083, E. S. Steele & Mrs. Steele (G, NY). NORTH 
CAROLINA. Yancey Co.: Wayah Bald Summit, Sept. 11, 
1933, E. J. Alexander, T. H. Everett & S. D. Pearson (NY). 
BuNCOMBE Co.: vicinity of Montreat, Sept. 9, 1913, P. C. 
Standley & H. C. Bolm (US, 10499, 10501-02, -11, -12, -13, -14). 
Haywoop Co.: on slopes of Pine Mt., vicinity of Eagle’s Nest, 
Sept. 6, 1910, P. C. Standley, 5552 (G, US); near Waynesville, 
near Highlands, Biltmore Herb., 2670v (US, 957890). Swain 
Co.: Great Smoky Mts., Aug. 28, 1891, E. C. Beardslee & C. A. 
Kofoid (G). Macon Co.: below Satula Mt., (alt. 2500’) High- 
lands, Sept. 2, 1902, E. E. Magee (G). SOUTH CAROLINA. 
OcoNEE Co.: summit of Thomas Bald Mt., near Georgia border 
(alt. 5200’), Aug. 19, 1893, J. K. Small (NY); eastern base of 
Tomassee Knob (alt. 1200’), Sept. 14, 1938, R. T. Clausen & 
H. Trapido, 3620 (NY). 

No species of Liatris has been so misunderstood and become 
the “catch-all” for as many different entities as Liatris scariosa. 
When describing Serratula scariosa, Linnaeus cited Gronovius 
(Gron. Virg. 92 (1739)), who described a plant observed and 
collected in Virginia by Clayton as follows: ‘Cirsium non ramo- 
sum foliis lateralibus flores ferens pauciores majores squamis 
hiantibus armatos pediculis curtis insidentibus, radice etiam 
tuberosa". The further references to descriptions by Banister 
(Plant. Virg. Ban. 1929 (1693)) and Plukenet (Pluk. Mant. 105 
(1749)) do not clearly identify the species, nor does the figure of 
Plukenet (Pluk. Phyto. t. 177 f. 4 (1696)) accompanying a de- 
scription (Pluk. Alm. 142 (1696)) that includes the phrase 
“floribus scariosis’’. 

Linnaeus added to the description ‘‘calycibus squarrosis pedun- 
culatis obtusis lateralibus" and, in the final note pointing out 
differences from Serratula squarrosa, he again referred to the 
same character: “‘calycibus squarrosis obtusis". The distinctive 
squarrose phyllaries point to a character from which many of the 
determinations of Liatris scariosa have varied. 

There is in the Linnaean herbarium a specimen labelled scar?- 


1946] Gaiser,—The Genus Liatris 299 


osa which Linnaeus had at the time of writing his description, a 
photograph of which Professor Fernald kindly allowed me to see. 
Though not a complete plant (the tip of the inflorescence and the 
base of the plant being lacking) this shows twelve large heads ca. 
1.5 cm. in diameter borne at the ends of pedicels about 3 em. long, 
and the phyllaries are squarrose. The outermost ones are dis- 
tinctly reflexed against the pedicels and the middle and inner ones 
to a slighter degree. All appear quite herbaceous and not at all 
scarious but rather slightly ciliolate on the margin. The leaves 
just below the lowest heads are oblanceolate, about 5 cm. long 
and 1.5 cm. wide, narrowing from the middle to a clasping base. 
Though the basal leaves are wanting on the specimen they would 
undoubtedly have been quite wide since a width of 1.5 cm. for an 
upper cauline leaf of Liatris is large. By use of a lens the rachis 
and leaves are seen to be pubescent. 

Willdenow (Sp. Pl. iii. 1635 (1803)), when transferring this 
species to the genus Liatris, again emphasized the obovate squar- 
rose phyllaries and added to the description of the inflorescence, 
a terminal leafy raceme, and to that of the leaves "utrinque 
attenuatis margine scabris". It is noteworthy that no mention 
of *scarious" phyllaries was made by any of the authors except 
Plukenet, whose phrase “‘floribus scariosis” is thus probably to 
blame for the name. Dr. H. K. Svenson,! who examined the 
Willdenow specimen and made drawings and notes, states: 
“bracts not particularly scarious, the scarious character perhaps 
referring to the fringe of hairs". 

Though I have not seen the Clayton plant nor a photograph 
of it, there is in the National Herbarium a tracing of it, accom- 
panying some correspondence by E. G. Baker in connection with 
determinations of specimens sent to him. When commenting on 
the plant from the Herb. Gronovius (British Museum Herb.) he 
noted: “‘(a) the bracts of the involucre are ciliate on the margin. 
(b) the bracts in the type are straight not folded. (c) the bracts 
do not tend to enlarge near the summit". The tracing shows 12 
heads, almost hemispherical with the phyllaries erect, not reflexed, 
and the upper stem-leaves quite as wide as those in Linnaeus’s 
plant. Thus, though the Gronovian plant does not perfectly 
match the Linnaean one and we are omitting it from the synony- 


1 Private communication. 


300 Rhodora . [OcroBER 


my of L. scariosa var. typica, we believe it may represent what oc- 
curs abundantly in the mountains of Pennsylvania, Maryland, 
West Virginia and Virginia. Specimens from there have been ex- 
amined that show the outer phyllaries to be herbaceous and 
spreading or but slightly reflexed, with the middle and inner ones 
a little scarious on the margin, less leathery than in typica, often 
colorful and also erect. The leaves vary from glabrous to asper- 
ous with basal ones very broad- to longer-oblanceolate and peti- 
olate. Lunell described a plant from West Virginia, Aug. 27, 
1908, K. K. Mackenzie, no. 368, White Sulphur Springs (I), as 
Laciniaria scariosa var. virginiana (Amer. Mid. Nat. ii. 172 
(1912)), which he later renamed var. borealis, ibid. 264. It would 
seem to represent such a derivative. That there are slight varia- 
tions in leaf- and especially in phyllary-characters can be seen by 
comparing the three specimens of the same date and number of 
the collector (I, NY, US), and a fourth (G) which has no number. 
While the type at the University of Indiana Herbarium has 
phyllaries quite ciliolate on the margins, the specimens at the 
National and Gray Herbaria have them hardly at all so, though 
in all three the phyllaries are erect. The specimen at the New 
York Botanical Garden, with rougher leaves and spreading 
phyllaries with only narrowly scarious, ciliolate margins, comes 
nearer L. scariosa var. typica. While other specimens have been 
seen that represent variations which perhaps may be combina- 
tions of parental characters of L. scariosa var. typica and L. 
aspera var. glabra, both of which occur in that range, there is 
nevertheless sufficient constancy found in these specimens of 
Mackenzie's to recognize their individuality and to relate them 
to L. scariosa. We have therefore retained Lunell's first varietal 
name and refer them to Liatris scariosa (L.) Willd. var. virginiana 
Lunell. 

Michaux (Fl. Bor.-Amer. ii. 92 (1803)) described a species 
from South Carolina with lanceolate leaves, rough on the margin, 
and with outer phyllaries squarrose, calling it Liatris squarrulosa 
and in a note following the Latin description he stated that it 
seemed to be like Serratula scariosa but he wondered why, as 
Plukenet held, it should be called scariosa. From photographs 
of Michaux's type specimen and of a duplicate and rather better 
specimen, obtained by Mr. Weatherby, it seems to have been a 


1946] Gaiser,—The Genus Liatris 301 


less robust, slender variety of the same species that is shown in 
the photograph of the Linnaean type. The heads are slightly 
smaller and the phyllaries more elongate than rounded but they 
are obtuse. The basal leaves are broadly, and the upper ones 
narrowly oblanceolate so that the whole plant strongly resembles 
the Linnaean one and we have no hesitancy in accepting its 
reduction to varietal ranking by Gray (Synopt. Fl. i?. 110 (1884)) 
as Liatris scariosa var. squarrulosa. However, we do not accept 
the appended synonymy of L. heterophylla R. Br. (Ait. Hort. 
Kew. ed. 2, iv. 503 (1812)) which was described as having acute, 
lanceolate phyllaries. A photograph of that type specimen from 
the British Museum, a plant cultivated by Mr. William Malcolm, 
shows a plant with a very different involucre. Most striking are 
the long, pointed phyllaries. From a packet at the Gray Her- 
barium, containing the phyllaries of a fragmentary head of this 
specimen, from the Banks Herbarium, they were found to be 
very thin and glabrous as well as linear. Pursh (Fl. ii. 508 
(1814)) when including a description of the species, after having 
seen the specimen, stated that the flowers were of the size of L. 
graminifolia, and he cited L. varia Herb. Banks ms. in synony- 
my. At the bottom of the sheet of L. heterophylla (Banks) there 
can be seen faintly written “L. varia? which undoubtedly ex- 
plains this reference. When Torrey & Gray (Fl. ii. 75 (1841)) 
described the species by use of the then recently received head 
and additional notes supplied by Mr. Bennett, they concluded 
that, though resembling L. scariosa in size and shape of heads, 
it did not match even depauperate specimens of that species 
because of the pointed scales. As in the time of those writers, 
"apparently the species has not been subsequently met with in 
this country", and we leave it among the doubtful species. 


18. LIATRIS ASPERA Michx. Corm rounded, irregular, sub- 
globose, 2-5 cm. in diameter: stems frequently single, sometimes 
several, quite stout, 4-11 dm. high, glabrous below with scattered 
hairs above on the rachis of the flowering spike, to asperous over 
the entire stem-length; leaves mostly linear-lanceolate though 
frequently almost linear; the basal broader, 1-1.5 dm. long, 1-2 
cm. wide, rhombic-lanceolate, narrowed into petioles of about 
half the total length, glabrous on both surfaces and lacking any 
marginal roughness, or asperous on one or both surfaces; upper 
leaves sessile, reduced to less than the length of the heads they 


302 Rhodora [OCTOBER 


subtend: inflorescence a long open spike of usually more than 20 
sessile to pedunculate somewhat globose, 25—40-flowered, heads 
1.5-2.5 cm. in diameter; phyllaries all glabrous with exposed tips 
rounded and with broad scarious margins, and slightly bullate, 
middle and inner ones oblong-spatulate, to rounded and strongly 
bullate, giving the globose head a puckered appearance; corolla 
usually purple, rarely white, pilose within the tube at the base of 
the stamens, tube 8-10 mm. long; pappus 7-8 mm. long; achene 
4-6 mm. long.—Fl. Bor.-Amer. ii. 92 (1803); Ell. Sk. ii. 276 
(1822(?)); DC. Prodr. v. 130 (1836). L. scariosa sensu Sims, 
Curtis's Bot. Mag. t. 1709 (1815) not (L.) Willd. L. sphaeroidea 
sensu Sweet Br. Fl. Gard. 1 ser., t. 87 (1824); Shinners, Amer. 
Mid. Nat. xxix. 34 (1943), not Michx. Laciniaria aspera Greene, 
Pittonia, iv. 318 (1901). Liatris scariosa Willd. f. Benkii Macb., 
Field Mus. Pub. Bot. iv. 127 (1927). Liatris sphaeroidea f. Benkii 
(Macb.) Shinners, Amer. Mid. Nat. xxix. 35 (1933). 

Var. typica. Stem rough-puberulent above or throughout 
with appressed or more or less incurved hairs: leaves asperous 
with dense, short, stiff hairs, linear-lanceolate, generally narrowly 
so.—Liatris aspera Michx. sens. strict. Laciniaria scariosa (L.) 
Hill vars. porrecta and obesa Lunell, Amer. Midl. Nat. ii. 159-162 
(1912). L. scariosa vars. virgata, strictissima and salutans 
Lunell, Amer. Midl. Nat. ii. 169-177 (1912). Liatris sphaeroidea 
var. salutans (Lunell) Shinners, Amer. Midl. Nat. xxix. 37 
(1943). Liatris sphaeroidea forma asperifolia Shinners, Amer. 
Midl. Nat. xxix. 36 (1943). Lacinaria indecidua and Lacinaria 
stratiotes Steele ex Winter, Analysis Flowering Plants of Nebras- 
ka, 143 (1936), Contrib. Bot. Surv. Neb. n. s. x. 143 (1936). 

Central States from Ohio, Wisconsin, Minnesota and North 
Dakota southward to Louisiana, Oklahoma and Texas.—OHIO. 
Lucas Co.: n. of Monclova, Aug. 8, 1924, Miss B. Garber (US). 
Scroro Co.: Friendship, Shawnee State Forest, D. Demaree, 11139 
(NY). INDIANA. Porter Co.: on the dunes, Dune Park, 
Sept. 17, 1909, E. S. Steele, 163a (G). JasPER Co.: open sandy 
places, Sept. 13, 1942, C. M. Ek (F). WISCONSIN. Co. un- 
determined: St. Helena, Aug. 1881, T. H. Bradwin (G). ONEIDA 
Co.: Manson, Sept. 1935, B. O. Dodge (NY). Pork Co.: dry 
ground, Rwy. right of way, St. Croix Falls, Sept. 4, 1927, N. C. 
Fassett & L. R. Wilson, 5453 (G). Brown Co.: Preble (plant to 
left), Aug. 21, 1878, J. H. Schuette (G). Pepin Co.: open dunes, 
lower terrace (albino), Aug. 27, 1927, N. C. Fassett, 4482 (G). 
JUNEAU Co.: abandoned field, 2 mis. s. of Matha, Aug. 27, 1937, 
J. W. Thomson (NY). Saux Co.: on the cliffs, on the s. side of 
Devil's Lake, Aug. 30, 1909, E. S. Steele, 80 (G). WAUKESHA 
Co.: Nashotah, Aug. 24, 1884, R. N. Larrabee (G). Grant Co.: 
Boscobel prairies (one plant), July 1886, C. H. Sylvester (NY). 
Racıne Co.: prairies, Sept. 8, 1882, H. E. Hasse (NY). Rock 


1946] Gaiser,—The Genus Liatris 303 


Co.: Clinton, along the Chicago & N. W. Rwy., Sept. 1, 1909, 
E. S. Steele, 96c (G), 96a (NY), 96b (US, 608836). MINNE- 
SOTA. Pine Co.: in sand at edge of forest, Highway 61, near 
Willow R., Aug. 10, 1938, Miss O. Lakela, 2726 (G). Topp Co.: 
dry sandy ground, Staples, Aug. 6, 1910, Z. L. Chandonnet (G); 
Staples, Z. L. Chandonnet, Aug. 18, 1914 (no. 307) (US), Aug. 9, 
1911 (no. 4) (M, type of Laciniaria scariosa var. porrecta Lunell) ; 
(no. 1) (M, type of Laciniaria scariosa var. obesa Lunell). Cui- 
saco Co.: Center City, July 1892, B. C. Taylor (G, plant 1). 
SrEARN Co.: St. Anthony, July 22, 1888, J. H. Schuette (NY). 
HENNEPIN Co.: Fort Snelling, Sept. 1888, E. A. Mearns, 141 
(US); prairies, Fort Snelling Reservation, Sept. 21, 1907, C. O. 
Rosendahl, 2066 (G); Richfield, Sept. 1875, N. H. Winchell (NY). 
WABASHA Co.: sand prairie, about 2 mis. n. of Weaver, Sept. 28, 
1930, N. Hotchkiss & P. Jones 4171 (US). GoopnuE Co.: Vasa, 
Aug. 1893, A. P. Anderson (US). LiNcorN Co.: Verdi, Aug. 
1891, E. P. Sheldon, S1344 (M). ILLINOIS. Without stated 
locality: July, 1846, S. B. Mead (G). McHenry Co.: Ringwood, 
G. Vasey (G). WiNNEBAGO Co.: Fountaindale, M. S. Bebb (G, 
281 US). Jo Daviess Co.: Hanover, Aug. 18, 1908, H. A. 
Gleason (G). Cook Co.: Riverside, Aug. 1912, J. M. Greenman, 
3831 (G); prairies, near Chicago City, Sept. 9-12, 1892, Ohlendorf 
(NY); vicinity of Chicago, Aug. 1909, Miss Reynolds, 2783 
(NY, US). Prora Co.: dry sandy ground, Peoria, Aug. 1904, 
F. E. Macdonald (M, type of Laciniaria scariosa var. strictissima 
Lunell); dry sandy soil, Peoria, Aug. 1904, F. E. Macdonald, 
1904 (G, NY); gravelly soil, Peoria, Sept. 3, 1908, F. E. Mac- 
donald (G). McLean Co.: prairieland, Hendrix, Aug. 31, 1904, 
B. L. Robinson (G). CnHaMPAriGN Co.: Champaign, by I. C. 
Rwy., Sept. 11, 1909, A. S. Pease, 12397 (G); Urbana, Sept. 25, 
1900, Miss M. L. Sheldon (G); original prairies, Rantoul, Oct. 5, 
1907, F. C. Gates, 2043 (US). Mason Co.: Topeka, Aug. 22, 
1904, H. A. Gleason (G); Decatur, Sept. 1, 1939, R. G. Mills 
(NY). Morgan Co.: vicinity of Concord, 2 mis. s. of Chapin, 
Sept. 1916, Miss S. Pratt (US). Prxe Co.: Rockport, Aug. 
1904, J. F. Clevenger (US). IOWA. Emmerr Co.: Armstrong, 
Aug. 1890, R. I. Cratty (G); prairie, slope of Four Mile Creek, 3 
mis. w. of Estherville, Aug. 13, 1934, Miss A. Hayden, 10535 
(NY). Fayerre Co.: prairies, Sept. 1894, B. Fink (G). CHERO- 
KEE Co.: upland prairie slopes, 3 mis. s. of Cherokee, Sept. 5, 
1937, Miss A. Hayden, 10537 (NY). HauirroN Co.: dry prairie, 
along C. & N. W. Rwy., 2 mis. w. of Webster City, Sept. 13, 
1933, Miss A. Hayden, 413 (NY). CARROLL Co.: Carroll, Aug. 
29, 1896, L. H. Pammel, 38 (G, NY). JoHNson Co.: prairies, 
Sept. 3, 1895, M. F. Fitzpatrick (I, type of Laciniaria scariosa 
var. virgata Lunell) ; Coralville, Sept. 3, 1909, M. P. Somes (US). 
PowESHIEK Co.: Grinnell, 1875, M. E. Jones (G, NY), Aug.-Sept. 


304 Rhodora [OcroBER 


1907, Miss R. Drew (G, NY, US 494671). Darras Co.: Red- 
field, Sept. 5, 1867, J. A. Allen (G). MISSOURI. Co. unde- 
termined: prairies, Sept. 1838, N. Riehl, 10 (NY). ATCHISON 
Co.: dry ground, Oct. 1893, B. F. Bush, 199 (NY). Anar Co.: 
Kirksville, Sept. 5, 1883, C. S. Sheldon, 3505 (NY). Macow Co.: 
Ethel, Sept. 22, 1915, B. F. Bush, 7802 (US). Jackson Co.: 
dry prairies, Martin City, Sept. 18, 1901, K. K. Mackenzie, 469 
(NY, US); barrens, Dodson, Oct. 4, 1906, B. F. Bush, 4155 (G), 
4159 (US); Dodson, Aug. 26, 1895, B. F. Bush, 242 (NY), Sept. 
27, 1915, W. W. Eggleston, 12046 (NY); Sheffield, Sept. 6, 1896, 
B. F. Bush, 909 (US); prairies, Lee's Summit, Sept. 5, 1906, 
B. F. Bush, 4098 (G), 4097 (US). Sr. Lours Co.: Allenton, Aug. 
30, 1894, G. W. Letterman (NY, US). JouwsoN Co.: near 
Warrensburg, Nov. 5, 1916, G. W. Stevens, 4424 (NY). BARTON 
Co.: dry prairies, Golden City, Oct. 7, 1913, E. J. Palmer, 4593 
(US). GREENE Co.: vicinity of Springfield, P. C. Standley, Aug. 
21, 1912 (no. 9172) (US), Aug. 31, 1911 (no. 8569) (G, US), Aug. 
1906 (US). Barry Co.: Hailey, J. W. Phillips (G; US, Sept. 
15,1915). ARKANSAS. BezwroN Co.: without stated locality, 
1899, E. N. Plank (NY). NORTH DAKOTA.  RicuraAND Co.: 
Hankinson, Aug. 25, 1902, P. A. Rydberg, 1151 (NY). SOUTH 
DAKOTA. RonEnrs Co.: hillsides, Aug. 1922, W. H. Over, 
14357 (US). ManmnsHaLnL Co.: mouth of St. Peter's River, 
Sept. 19, 1839, C. A. Geyer, Nicollet's N. W. Exped., 271 (US). 
Grant Co.: virgin prairie, Clear Lake near Big Stone Lake, Aug. 
1, 1940, P. Johnson, 69 (NY). Sprnx Co.: Doland, Sept. 10, 
1896, L. W. Carter (NY). Brooxines Co.: Lake Hendricks, 
Aug. 1906, Miss F. N. Vasey (G, US). Yanxrton Co.: high 
knolls, Jamesville, Aug. 24, 1899, L. A. Bruce, 55 (US). NE- 
BRASKA. Co. undetermined: prairies, Sept. 16, 1874, O. 
Kuntze, 2923 (NY). ANTELOPE Co.: Brunswick, Sept. 5, 1908, 
N. F. Peterson (US). SaAuNpEns Co.: Mead's Ranch, Aug. 24, 
1893, F. Clements, 2901 (G (US, type of Lacinaria indecidua 
Steele) N). Howarp Co.: St. Paul, Aug. 29, 1919, J. M. Bates 
(G). Cass Co.: Weeping Water, Aug., T. A. Williams (US, 
517526). LANCASTER Co.: Lincoln, 1906, C. E. Bessey (US), Aug. 
1886, H. J. Webber (NY); Lancaster, Sept. 10, 1873, S. Angley 
(N, type of Lacinaria stratiotes Steele). Satine Co.: Crete, C. E. 
Brown (US). Kearney Co.: Minden, Aug. 26, 1917, Dr. H. 
Hapeman (G). KANSAS. Co. undetermined: from Council 
Grove to Fort Leavenworth, Aug. 1847, Plant. Nov. Mex., A. 
Fendler, 302 (333)? (G). Ritey Co.: prairie, Sept. 12, 1895, J. B. 

1 From Senate Report intended to illustrate a map of the Hydrographical Basin of 
the upper Mississippi River made by I. N. Nicollet, Feb. 16, 1841, Washington 1843, 
the place of this collection would seem to be about Marshall Co. just a little north of 
Day Co. 

: In Plant. Fendl. A. Gray, 1849, page 63. specimen no, 302 is listed as L. scariosa 
Willd. and was collected from ‘‘Council Grove to Fort Leavenworth'', Aug. 1847. The 


1946] Gaiser,—The Genus Liatris 305 


Norton, 214 (G, US, NY); Manhattan, Sept. 12, 1892, J. B. Nor- 
ton (NY). Douvuaras Co.: Lawrence, W. C. Stevens (US). SHaw- 
NEE Co.: Topeka, Summer, 1897, Prof. Harshberger, 3469 (US). 
Lyon Co.: high prairies, 3 mis. n. of Madison, Sept. 18, 1941, H. 
A. Stephens (G). Linn Co.: grassy hillside, near Parker, Oct. 21, 
1916, G. W. Stevens, 4341 (NY). CowrEv Co.: without stated 
locality, Aug. 22, 1898, M. White (NY). OKLAHOMA. Orra- 
WA Co.: in dry pasture, Ottawa, Aug. 29, 1913, G. W. Stevens, 
2511 (G). Nowara Co.: grassy roadside, Lenapah, Aug. 19, 
1913, G. W. Stevens, 2176 (G). Rocers Co.: 4 mis. n. of Catale, 
Oct. 9, 1938, M. Hopkins & M. Van Valkenburgh, 3678 (O). TuL- 
sa Co.: without stated locality, Autumn, 1926, E. R. Force, 13026 
(O); dry railroad side, n. e. of Tulsa, Sept. 17, 1939, U. T. Water- 
fall, 1797 (O). Payne Co.: 6 mis. n. of Stillwater, Sept. 13, 1935, 
E. E. Richardson (O). Mvsxoakk Co.: Lot 4, Sept. 4, 1927, E. L. 
Little, 2227 (O). TEXAS. BasrnoP Co.: Bastrop, Nov. 24, 
1928, B. C. Tharp (NY), Oct. 1926, H. H. Duval (US). WALKER 
Co.: 14 mis. s. w. of Huntsville, Sept. 28, 1934, V. L. Cory, 10260 
(G). Harris Co.: sandy soil near Houston, Aug. 21, 1903, Bilt- 
more Herb. 2670e (US); Houston, Sept. 28, 1917, E. J. Palmer, 
12786 (US); near Houston, Sept. 22, 1903, Biltmore Herb., 2670j 
((I, type of Laciniaria scariosa var. salutans Lunell) (US)); 
Houston, G. L. Fisher, Sept. 24, 1937 (no. 37160) (US), Aug. 23, 
1915 (no. 1513) (US), Aug. 23, 1914 (no. 2032) (US), 1842, F. 
Lindheimer (G). GonzaLes Co.: Cottonwood Springs, Sept. 7, 
1933, H. B. Parks, 7682 (G). 

Var. intermedia (Lunell), comb. nov. Stem glabrous below, 
with appressed hairs above on the rachis of the flowering spike; 
leaves glabrous or with but a few scattered hairs and frequently 
broadly lanceolate.—Laciniaria scariosa (L.) Hill var. intermedia 
Lunell, Amer. Mid. Nat. ii. 173, 177 (1912) = var. media Lunell 
ibid. 264 (1912). | Laciniaria scariosa var. petiolata Lunell, Amer. 
Mid. Nat. ii. 172, 176 (1912). Liatris sphaeroidea of many 
authors, not Michx. Fl. Bor.-Amer. ii. 92 (1803). Probably L. 
sphaeroidea sensu Lodd. Bot. Cab. t. 1417 (1828), though de- 
scription insufficient. 

Occurring east of the Mississippi through North and South 
Carolina to Georgia and Florida, from Ontario through Indiana 
and Kentucky to Alabama and west of the Mississippi more 
generally in the southern central plains-states, Missouri, Arkan- 
sas, Louisiana, Texas and Oklahoma.—WEST VIRGINIA. 
RrircurE Co.: Berea, dry hillside-field, Aug. 21, 1922, L. F. & F. 
R. Randolph, 1376 (G). NORTH CAROLINA. RUTHERFORD 
Co.: Hickory Nut Gap, Salola Mt., to Chimney Rock, Oct. 3, 
1901, J. K. Small & A. M. Huger (NY). Henperson Co.: dry 


same applies to (333), which bracketed number represents the number under which it 
was distributed. 


306 Rhodora [OCTOBER 


woods, East Flat Rock, Sept. 15, 1926, F. W. Hunnewell, 10014 
(G). - Macon Co.: Yellow Mt., ex Torrey Herb., Curtis (NY). 
SOUTH CAROLINA.  GREENVILLE Co.: Caesar's Head, Aug. 
2, 1881, J. D. Smith (G); rocky woods, Caesar's Head, Aug. 12, 
1881, J. D. Smith, 53 (US). GEORGIA.  Frorvp Co.: without 
stated locality, Chapman (US). McDurrir Co.: sandhills, 
vicinity of Thomson, Sept. 9, 1908, H. H. Bartlett, 1491 (G, US); 
vicinity of Thomson, Sept. 7, 1907, H. H. Bartlett (US). FLORI- 
DA. Jackson Co.: Sneads, Aug. 19, 1942, R. A. Knight (F). 
ALACHUA Co.: high thin woods, Warrens Cave, Gainesville, Oct. 
25, 1927, Louchs, Miss L. Arnold & E. West (F); dry roadside, 
Sugarfoot, Gainesville, Aug. 28, 1932, Miss L. Arnold (F). 
ONTARIO. Lampton Co.: Port Franks, Aug. 31, 1905, C. K. 
Dodge (G, US); sandy open ground near L. Huron, Port Franks, 
Sept. 2, 1929, E. J. Palmer, 36270 (G); near Sarnia, Aug. 18, 
1892, C. K. Dodge, 11587 (Ot); rather low grounds, Point Ed- 
wards, July, 1887, Burgess, 9854 (Ot); Sarnia, Aug. 16, 1901, J. 
M. Macoun, 22614 (Ot); among small oaks, near Sarnia, Sept. 8, 
1904, C. K. Dodge, 1 (Ot, US); 4 mis. s. of Grand Bend, Sept. 3, 
1934, H. H. Brown, 4661, 4667 (HB); Grand Bend, Aug. 6, 1932, 
Marie-Victorin, R. Germain & Jacques, 49246 (To). KENT Co.: 
Squirrel Is., Aug. 31, 1904, C. K. Dodge, 2 (Ot, US). Essex Co.: 
woods, Sandwich, July 27, 1901, J. M. Macoun, 26615 (Ot); 
Point Pelee, Aug. 24, 1931, H. H. Brown (HB). MICHIGAN. 
Co. undetermined: Union Pier, Sept. 19, 1934, Miss A. Fishman, 
231 (O). Kewrrnaw Co.: without stated locality, Aug. 1889, 
O. A. F. (G). Sr. Crarm Co.: Fort Gratiot, ex Torr. Herb. Dr. 
Pitcher, 1829 (NY); near Port Huron, Aug. 30, 1892, C. K. Dodge 
(US). Carmoun Co.: east of Albion, Aug. 9, 1906, C. E. Barr 
(G). Sr. JosEePH Co.: without stated locality, Sept. 2, 1838, 
Houghton (NY). OHIO. Without stated locality. Herb. Schw. 
(P). Co. undetermined: Margaretta Ridge, near Mt. Clemens, 
Sept. 25, 1904, E. L. Moseley (US). Errr Co.: sand dune, Cedar 
Point, R. J. Webb, 5491 (G). Lucas Co.: 10 mis. w. of Toledo, 
Sept. 2, 1897, E. L. Moseley (US). Woop Co.: Plain twsp., Sect. 
21, Aug. 10, 1937, R. E. Shanks, 2201 (NY). Ross Co.: Scioto 
Trails State Forest, dry wooded hills, Stoney Creek, Aug. 19, 
1935, D. Demaree, 11532 (US). INDIANA. SrEeunaEN Co.: 
14% mi. n. of Clear Lake, Sept. 11, 1904, C. C. Deam (NY); 
sandy woods, e. side of Clear Lake, Sept. 11, 1904, C. C. Deam 
(M, type of Laciniaria scariosa var. petiolata Lunell). La 
GnANGE Co.: high dry bank, e. side of Pretty Lake, Ava. 27, 
1914, C. C. Deam, 14876 (G); on the bank of the Pigeon R., 
about 2 mis. e. of Ontario, Aug. 30, 1914, C. C. Deam, 15050 (US). 
Porter Co.: sand dune, n. of Mineral Springs, Aug. 14, 1911, C. 
C. Deam, 9620 (M, type of Laciniaria scariosa var. intermedia 
Lunell); Dune Park, Sept. 4, 1906, L. M. Umbach, 1447 (US). 


1946] Gaiser,—The Genus Liatris 307 


LAKE Co.: open sandy woods, Miller’s, Sept. 20, 1914, F. W. 
Johnson, 1525 (NY); Buffington to Pine, old beaches “Lake 
Chicago", Sept. 20, 1901, E. S. Steele, 183 (US 609101); Mar- 
SHALL Co.: Indiana Harbour, Aug. 26, 1916, C. C. Deam, 21299 
(G); eopse near Lost Lake, Lake Maxinkuckee, Culver, Aug. 21, 
1926, J. R. Churchill (G); Lake Maxinkuckee, 1899, B. W. 
Evermann, 1118 (US); sandy open knoll, outlet of Lost Lake, 
Plymouth, Sept. 3, 1909, H. W. Clark (US). SmARKE Co.: 
sandy soil, along N. Y. C. Rwy., 1.3 mi. w. of North Judson, 
Sept. 6, 1941, C. M. Ek (G, NY). Harrison Co.: Barrens s. e. 
of Corydon (albino), Sept. 5, 1836, Mr. Clapp (G). KEN- 
TUCKY. WnurirLEYv Co.: dry sand bank along Rwy., Cumber- 
land Falls, Sept. 11, 1940, F. T. McFarland, 66 (G, NY). TEN- 
NESSEE. Roane Co.: Malden's Ridge near Harriman, Aug. 
20, 1903, Biltmore Herb., 2670g (US). SEQvATCHIE Co.: dry 
conglomerate rocks, Cagle, Aug. 17, 1938, H. K. Svenson, 9659 
(G). HaMwiLTOoN Co.: dry soil, Lookout Mt., Aug. 24, 1897, 
Biltmore Herb., 2670 (NY, G) (US 957898, 332418); Lookout 
Mt., Sept. 6, 1877, L. F. Ward (US 134395). ALABAMA. 
Without stated locality: Sept. 1841, ez Herb. G. Thurber, S. B. 
Buckley (G). WISCONSIN. Pork Co.: St. Croix Falls, Aug. 
13, 1900, C. F. Baker (G). Brown Co.: Preble (plant to right), 
Aug. 21, 1878, J. H. Schuette (G). WaAvsHARA Co.: Wild Rose, 
Aug. 8, 1919, W. L. McAtee, 3053 (US). JuNEAv Co.: 2 mis. s. 
of Mather, Aug. 27, 1937, J. W. Thomson (NY); Camp Douglas, 
Aug. 20, 1890, E. A. Mearns, 141 (NY); Camp Douglas (albino), 
Aug. 28, 1890, E. A. Mearns, 141 (US). Snurnovaaw Co.: 
sandy ridges, s. of Sheboygan, Aug. 11, 1904, L. H. Shinners, 
2554 (NY). CoruwmBia Co.: Dells of the Wisconsin, Aug. 19, 
1893, Amer. Ass. Adv. Sci. Meet. (NY). Saux Co.: vicinity of 
Kilbourn, on Wisconsin R., Aug. 25, 1909, E. S. Steele, 11d (NY); 
on the cliffs, Devil's Lake, Aug. 31, 1909, E. S. Steele, 90b (NY); 
rocky ground, about Devil's Lake, Baraboo, Sept. 7, 1925, E. J. 
Palmer, 28383 (G). Grant Co.: prairies, Boscobel (plant to 
left), July 1886, C. H. Sylvester (NY). MINNESOTA. ANOKA 
Co.: Moore Lakes, Sept. 5, 1926, P. A. Rydberg, 9673 (NY). 
ILLINOIS. Cook Co.: Lakeview, Sept. 7, 1882, W. Deane (G); 
prairie, near S. Chicago, Sept. 15, 1910, O. E. Lansing, 2858 
(G, US); Chicago, Dr. Scammon (NY); vicinity of Palos Park, 
Sept. 6—7, 1909, E. S. Steele, 132a (US 672726). Prorta Co.: 
exposed hillside, gravelly soil, Peoria, Sept. 3, 1908, F. E. Mc 
Donald (US 609908). MISSOURI. Prxe Co.: McCune, Aug. 
25, 1916, J. Davis, 250 (US). Jackson Co.: Dodson, Aug. 22, 
1895, B. F. Bush, 242 (US). Ozark Co.: rocky open ground, 
near top of Bald Jesse, near Gainesville, Oct. 10, 1927, E. J. 
Palmer, 33071 (G). GREENE Co.: vicinity of Strafford, Aug. 27, 
1912, P. C. Standley, 9478 (G). Taney Co.: open rocky ground, 


308 Rhodora [OCTOBER 


Malva, Sept. 17, 1924, E. J. Palmer, 26189 (G). ARKANSAS. 
Co. undetermined: Ozarks, on high banks along Frisco Rwy., 
Sept. 10, 1927, A. Ruth, 79 (US). FAULKNER Co.: dry hills, 
Quitman, Sept. 4, 1934, D. Demaree, 10950 (NY). Lonoke Co.: ~ 
Carlisle, July 31, 1938, D. Demaree, 18012 (O); Pressure Reser- 
voir, Hot Springs National Park, July 20, 1934, H. R. Gregg, 
269 (US). Yeu Co.: ridges, Mt. Nebo State Park, Aug. 30, 
1939, D. Demaree, 20593 (G). GARLAND Co.: along Rwy., near 
Gulph Bridge, July 19, 1935, F. J. Scully, 374a (G); near Hot 
Springs, 1928, R. Runyon, 1179 (NY); summit of hills, Hot 
Springs, July 20, 1931, R. Runyon, 1500 (US). Sevier Co.: 
Prairie de Queen Park, July 1937, P. W. Beck (O). Drew Co.: 
Monticello, Sept. 10, 1938, D. Demaree, 18297 (I); old fields 9 
mis. s. of Monticello, Aug. 30, 1936, D. Demaree, 13574 (G, O). 
Hempsteap Co.: Ozan, Aug. 25, 1937, D. Demaree, 15967 (0). 
MILLER Co.: Texarkana, Aug. 20, 1896, A. A. Heller & E. G. 
` Heller, 4125 ((G, NY, US)). LOUISIANA. Co. undetermined: 
W. Louisiana, 1839, Dr. Hale (US). Cappo Co.: open dry 
field, along low woods, Pine Hill Rd., ca. 5 mis. n. w. of Shreve- 
port, Aug. 6, 1938, D. S. Correll & H. B. Correll, 10090 (NY). 
OKLAHOMA. Mayes Co.: dry hillside pasture, Adair, Sept. 
2, 1913, G. W. Stevens, 2581 (G, NY). Tursa Co.: Tulsa, Oct. 
30, 1913, G. W. Stevens, 2089 (G). CmEEK Co.: Arkansas R., 
Aug. 21, 1895, J. W. Blankinship (G). Mvskoakk Co.: Lot 1, 
July 23, 1927, E. L. Little, 1903 (O); Lot 3, Sept. 7, 1927, E. L. 
Little, 3127 (O); 2 mis. w. of Muskogee, Aug. 8, 1926, E. L. 
Little, 257 (O); near Muskogee, Sept. 25, 1896, L. F. Ward, 10 
(US). Huaues Co.: prairie, Aug. 26, 1938, C. C. Smith (O). 
SEMINOLE Co.: Seminole (albino), July 21, 1936, M. Hopkins & 
D. Demaree, 48 (O). Le Fiore Co.: Stapp, Aug. 25, 1937, D. 
Demaree (O); in open woods, in mt. valley, Page, Sept. 9, 1913, 
G. W. Stevens, 2714 (G); on s. side of Rwy. embankment, near 
Page, Sept. 8, 1913, G. W. Stevens, 2627 (G, NY). LATIMER Co.: 
Wilburton, Aug. 10, 1930, O. M. Clark (O). Prrrssure Co.: 
McAlester, Aug. 8, 1894, C. S. Newhall (G). TEXAS. Gray- 
son Co.: Denison, Sept. 13, 1906, F. J. Tyler (US). Grece Co.: 
without stated locality, Autumn, 1941, C. L. York (G). Harris 
Co.: Houston, 1842, Lindheimer (G). WALLER Co.: Hempstead, 
Aug. 28, 1932, B. C. Tharp (G). 


This species is widely distributed but has not always been 
accepted, having been referred to L. scariosa by Torr. & Gray 
(Fl. N. Am. ii. 75 (1841)) and again by Gray (Synop. Fl. 1?. 110 
(1884)). Examination of a photograph at the Gray Herbarium 
of the type specimen collected by Michaux in the “Illinois 
meadows”, though not showing a complete plant, made it possible 


1946] Gaiser,—The Genus Liatris 309 


to determine the nature of the phyllaries. The outer ones are 
recurved, the middle and inner ones have rounded or spatulate 
tips with broad petaloid margins and are clearly crisped. In 
pressing, these have sometimes been folded back on themselves 
or left concave and cup-like. The upper cauline leaves are linear- 
lanceolate with acute tips. As Greene, who recognized this 
species under Laciniaria (Pittonia, iv. 318 (1901)) stated, it is 
difficult ‘‘to understand on what principle a plant so well marked 
as this could be confused . . . with L. scariosa". From the type 
of that species it is easily known by the middle and inner, gla- 
brous, broadly scarious, bullate phyllaries, rather than herba- 
ceous, squarrose ones, and by the acute lanceolate, rather than 
oblanceolate, leaves. 

Michaux had given Illinois as the type locality for this species. 
From the account in his journal (Proc. Amer. Phil. Soc. xxvi. 129 
(1888)) and the map of F. A. Michaux!, Illinois would be as far 
west as he had travelled and would refer to the region east of the 
Mississippi, there being no Indiana at that time. Plants with 
the familiar puckered heads of the aspera type are now known to 
occur from southern Ontario, Michigan and Ohio south to the 
Carolinas and southwestward to Louisiana, Texas and Okla- 
homa. When leaves of specimens from Illinois southward and 
westward are examined it is found that some are rough, some 
glabrous and some of varying degrees of intermediacy of pubes- 
cence. In contrast, the southeastern specimens, from the moun- 
tains of Tennessee to South Carolina, Alabama and Florida, are 
more constantly glabrous. To the author, therefore, it seemed 
best to broaden Michaux's conception of the species to allow for 
leaves from very rough to very smooth and to consider them as 
two varieties, the typical and the glabrous respectively. The 
lists of specimens so separated show the range of the former from 
Wisconsin, Indiana and Illinois westward while the latter extends 
eastward, as well, into Kentucky and the Carolinas and from 
there southward into Alabama and Florida. 

When examining the types of Lunell's numerous varieties of 
Laciniaria scariosa it was found that two of the seven (see also 
no. 20), namely var. intermedia (which was later called media) 
and var. petiolata, both from Indiana, come under the glabrous 


1 Travels to the westward of the Alleghany Mountains. J. Mawman (1805). 


310 Rhodora [OCTOBER 


variety of L. aspera Michx. as here interpreted. Lunell’s earlier 
name intermedia takes precedence in this new varietal combina- 
tion, which has been made on the basis of the characters of his 
type and not upon the characters described and classified by him. 
The glabrous Liatris sphaeroidea of Michaux, however, is not 
included in synonymy. 

Michaux (l. c.) described L. sphaeroidea as having smooth 
leaves, pedunculate flowers and oval erect bracts and gave as 
habitat the high mountains of Carolina as well as the meadows of 
Illinois. The identification and delimitation of L. sphaeroidea 
and L. aspera has been a perplexing matter. Comparison of the 
photographs of Michaux’s two plants seen at the Gray Herbarium, 
reveals similar inflorescences of large heads in loose racemes, 
though they are sessile in the latter and short-pedunculate in the 
former. Since within other species all conditions from sessile to 
pedunculate heads occur, that character is not diagnostic. The 
phyllaries, however, are of different shapes and quite differently . 
disposed in the two specimens. The outer ones of L. sphaeroidea 
are outspread but hardly recurved. The middle and inner ones 
are erect, elongate, oblong and slightly spatulate at the tip, with 
narrow scarious borders, some just a little crisped but none 
really concave as in L. aspera. As stated above, Michaux 
referred to smooth leaves in L. sphaeroidea in contrast to very 
rough ones in L. aspera and gave it a habitat in the high moun- 
tains of Carolina as well as in the meadows of Illinois. 

Careful search for specimens resembling Michaux’s type of 
sphaeroidea in the characters of the head and phyllaries, rather 
than just in smooth leaves or pedunculate heads, shows a great 
many from around the Great Lakes region. Since on the label 
of Michaux’s type-specimen occurs “Prairies vers Mississippi" 
it seems that there may be some justification for the interpreta- 
tion of his species being like many of those of the Great Lakes 
Basin. Where it occurs along with L. aspera, as it does among 
the pine-oak scrub on the sand-ridges of the southeastern shore 
of Lake Huron in Ontario, it is often difficult to be sure of the 
determination, especially when it has grown under particularly 
unfavorable circumstances or during unusually dry summers. 
To this author, L. sphaeroidea occurring as glabrous, semi- 
hirsute and hirsute plants, comes very close to and is perhaps 


1946] Gaiser,—The Genus Liatris 311 


some hybrid of L. aspera Michx. Whether it may have arisen 
originally in the more central northern region as an intermediate 
between L. aspera and L. ligulistylis we cannot say. 

L. aspera Michx. is undoubtedly the species of widest geo- 
graphic range of the Scar?osae series and is therefore responsible 
for many confusing mid-forms. At the northwestern limit of 
the range of L. aspera, as in Wisconsin, Minnesota and North 
Dakota, it meets L. ligulistylis. From this species, with fewer 
large heads and erect, lacerate phyllaries, it is clearly distinguish- 
able. The presence of pilosity in the corolla-tube of L. aspera and 
its absence in L. ligulistylis makes a very satisfactory additional 
character for confirmation of the separation of the two. On 
this basis seven of Lunell’s varieties of Laciniaria scariosa have 
been transferred to L. aspera (five under var. typica and two as 
var. intermedia), while nineteen others were included under L. 
ligulistylis. Intergradations between the two species, however, 
are numerous and varied and eight of these were recognized also 
among Lunell's scariosa varieties (see no. 20). With all the 
recombinations of leaf-, stem-, head-, phyllary- and corolla-tube- 
characters, these intermediate specimens, found mostly west of 
Lake Michigan, are nevertheless distinguishable by the generally 
larger and more hemispherical mature heads from the segregate 
that we have interpreted as X L. sphaeroidea Michx. of the 
Great Lakes region with smaller, more campanulate or globular 
heads. That they more nearly resemble L. aspera can be judged 
by the confusion of interpretation seen in various authors' lists 
of synonyms. By contrast, X L. Neiuwlandii (Lunell) Shinners 
(see no. 20) has a recognizably stronger relationship to L. liguli- 
stylis and may represent a more recent derivative. 

That no specimens of X L. sphaeroidea are here listed as occur- 
ring in the high mountains of Carolina, which were included along 
with the meadows of Illinois as the habitat, may perhaps be due 
to Michaux having casually matched his specimen by its glabros- 
ity with the variety of L. aspera to be found in that region (var. 
intermedia), without giving weight to the differences he noted in 
the phyllaries of the two species he described. There might also 
have been specimens of the L. scariosa var. virginiana that 
Michaux saw occurring in those mountains, with the phyllaries 
less reflexed and with slightly more scarious margins than in 


312 Rhodora [OcroBER 


variety typica, for he certainly thought of L. scariosa as having 
wholly herbaceous phyllaries; to wit his observation under L. 
squarrulosa ** Videtur SERRATULA scariosa L. non autem video cur, 
Pluknetio duce, scariosam dixerit." Examination of Michaux's 
herbarium for any further specimens from the Carolina region 
would certainly be interesting and might help final elucidation of 
this alliance, which has been so variously interpreted. 

Laciniaria Deamii Lunell (Amer. Mid. Nat. ii. 169 (1912)), 
described from a plant of C. C. Deam, no. 1747, from the base of 
open dunes, just south and east of Indiana Harbor, Lake Co., Ind. 
(I), can be included under this interpretation of X L. sphaeroidea 
because of the wholly glabrous phyllaries, petaloid on the margins 
and a little crisped, and the campanulate heads of about thirty 
flowers. Examination of N. C. Fassett, no. 21207, from Bear 
Lake, Wolf Lake, Noble Co., Ind., Sept. 11, 1941 (W), which by 
Shinners (Amer. Mid. Nat. xxix. 31 (1943)) was made the type 
of X L. Deamii (Lunell) Shinners f. albina, showed marked differ- 
ences from Lunell's type, aside from the flower-color, in the 
almost wholly herbaceous and somewhat pubescent phyllaries 
and the few (8 fully developed) large heads of 45-50 flowers. 
Like it, a plant of normal color, no. 21208, collected at the same 
time and place by N. C. Fassett, in the large terminal heads 
seems rather to show relationship to X L. Niewwlandii, the type 
of which was described from Indiana. 

That X L. sphaeroidea has a stable identity is judged from the 
number of times it has probably been one of the parents in other 
recognized hybrids. It has seemed to cross with species of two 
other series including one of each other section, as with L. spicata 
of the Spicatae series in the production of X L. Steelei (see no. 1) 
and with L. cylindracea of the Cylindraceae series and section 
Euliatris in X L. Gladewitzii Farwell (see no. 31). 

X LIATRIS SPHAEROIDEA Michx. Rootstock irregular, sub- 
globose, 2-5 cm. in diameter: stems one or few, 4-11 dm. high, 
from glabrous to asperous as in L. aspera: leaves glabrous, some- 
what pubescent or asperous, linear-lanceolate, the basal 1-1.5 
dm. long and ca. 1 em. wide, subpetiolate, reduced upwards to 
narrowly lanceolate ones not much longer at the base of the 
inflorescence than the heads subtended: inflorescence a raceme or 


panicle of numerous, 20-40, sessile or pedicellate heads of 25-40 
flowers: heads barely globose (when young cylindric-campanulate) 


1946] Gaiser,—The Genus Liatris 313 


with phyllaries only slightly bullate or crisped, all loosely erect, 
outer ones obovate or oblong with slightly spatulate and scarious- 
margined tips, middle ones more elongate and with the spatulate 
ends still scarious-margined and only slightly crisped; corolla 
usually purple, sometimes pale mauvish pink, rarely white; tube 
ca. 9 mm. long, pilose within; pappus 7-8 mm. long, barbellate; 
achene 4-5 mm. long.—F I. Bor.-Amer. ii. 92 (1803). Ell. Sk. ii. 
281 (1822?); DC. Prodr. v. 130 (1836); not L. sphaeroidea sensu 
Shinners, Amer. Mid. Nat. xxix. 35 (1943). Suprago sphaero- 
cephala Cass. Dict. li. p. 386 (1827). Laciniaria Deamit Lunell, 
Amer. Mid. Nat. ii. 169 (1912). Liatris scariosa var. Deamii 
Peattie, Amer. Mid. Nat. x. 132 (1926). X Liatris Deamii (Lunell) 
Shinners, Amer. Mid. Nat. xxix. 31 (1943), in part. 

Southern Ontario to Minnesota and Nebraska, south to 
Tennessee and Arkansas.—ON TARIO. Lampton Co.: dry poor 
soil, Walpole Is., Sept. 24, 1909, C. K. Dodge (US). Essex Co.: 
in open woods, Leamington, July 30, 1892, J. M. Macoun, 
22769 (plant to left) (Ot). MICHIGAN. Detra Co.: in jack 
pines, near Rapid R., Aug. 19, 1933, F. C. Gates, 17430 (US). 
ST. CLAIR Co.: dry sandy soil, near Port Huron, Aug. 10, 1896, 
C. K. Dodge (US); Lakeside Cemetery, Port Huron, Aug. 28, 1904, 
Aug. 23, 1911, C. K. Dodge (US). IwanuaM Co.: along roadside 
in sand, Haslet, Aug. 30, 1917, T. C. Yuncker, 723 (US). OHIO. 
Co. undetermined: top of dry cliff, Ross Hollow, Sept. 27, 1936, 
Bartley & Pontius, 152 (NY). Erie Co.: Cedar Point, Oct. 8, 
1904, E. L. Moseley (US). Lvcas Co.: Spencer Twsp., sandy 
soil, Sept. 26, 1921, E. L. Moseley (US); several mis. n. w. of 
Whitehouse, Aug. 28, 1927, E. L. Moseley (US). INDIANA. 
STEUBEN Co.: in sandy wood, e. side of Clear Lake, Aug. 21, 
1904, Sept. 9, 1904, C. C. Deam (G). Porter Co.: Dune Park, 
Sept. 2, 1898, L. M. Umbach (US 609933); Dune Park, Sept. 17, 
1909, E. S. Steele, 160 (G, US 609009); on and among dunes. 
Dune Park, Sept. 17, 1909, E. S. Steele, 162b (US 609017). 
Laxe Co.: near L. Michigan, about 14 mi. from lake-front, 1906, 
C. C. Deam, 1747 (US); sandy pine ridges, Pine, Sept. 28, 1910, 
O. E. Lansing, 2889 (G); Indiana Harbor, Sept. 15, 1909, E. S. 
Steele, 150b (US 608997) ; 153b (US 609044) ; base of open dunes, 
just s. & e. of Indiana Harbor, Sept. 23, 1906, C. C. Deam (I, 
type of Laciniaria Deamit Lunell); Hammond, old beaches, 
“Lake Chicago", Sept. 14, 1909, E. S. Steele, 143a, 146a, 146c 
(G); Sept. 18, 1909, E. S. Steele, 174a (US 609078) ; Buffington to 
Pine, old beaches “Lake Chicago", Sept. 20, 1909, E. S. Steele, 
184a (US 609014), 184c (G, US 609106), 151c (US 672715), 
180b (G), 180g (US 699097), 189e (G, US 609127), 189f (G), 
187a (G, US), 188b (G, US 609123. KENTUCKY. Rowan 
Co.: prairie patch, Clack Mt., Sept. 27, 1936, Miss E. L. Braun 
(G). TENNESSEE. Kwnox Co.: vicinity of Knoxville, Sept. 


314 Rhodora [OCTOBER 


18, 1890, F. Lamson-Scribner (US). WISCONSIN. BURNETT 
Co.: dry sandy ground n. of Danbury, Aug. 26, 1940, L. H. 
Shinners & J. Catenhuse 2836 (G). Brown Co.: Preble (plant 
to right), Aug. 21, 1878, J. H. Schuette (G). Saux Co.: rocky 
ground, about Devil’s Lake, Sept. 7, 1925, E. J. Palmer 28383 
(G); Devil’s Lake, Aug. 15, 1881, J. M. Holzinger (US); eastern 
range of cliffs, Devils Lake, Aug. 31, 1909, E. S. Steele, 90c 
(US 608814), 90e (US 608812), 90h (US 608817); vicinity of 
Kilbourn, Aug. 30, 1909, E. S. Steele, 79 (US); dry sandy bluffs 
of river, vicinity of Kilbourn, Aug. 25, 1909, E. S. Steele, 11b 
(US 619845), 11c (US 608703). MINNESOTA. Irasca Co.: 
sandy soil, Grand Rapids, Aug. 1891, J. H. Sandberg, 741 (US). 
OTTER Tatu: sandy soil, Richdale, Aug. 13, 1912, Z. L. Chandon- 
net, 0.1 (US); sandy hills, Richdale, Aug. 28, 1913 (no. 136), Aug. 
12, 1914 (no. 280) Z. L. Chandonnet (US); Topp Co.: dry sandy 
soil, Staples, Aug. 19, 1912, Z. L. Chandonnet, 19, 25, (US); 
sandy soil, Staples, Aug. 22, 1913, Z. L. Chandonnet, 130 (US). 
Cuisaco Co.: Center City, July 1892, B. C. Taylor, plant 2 (G). 
ILLINOIS. Coox Co.: pebbly clay of Valparaiso moraine, 
vicinity of Palos Park, Sept. 6-7, 1909, E. S. Steele, 132a (US 
608918), 133 (US 608929); Rogers Park, old beaches, “Lake 
Chicago’’, Sept. 4, 1909, E. S. Steele, 105 (G, US 608862), Sept. 
10-11, 1909, 141a (US 608960), 140 (G, US 608966); dry open 
woods, sand dunes, Miller's, Sept. 4, 1911, E. E. Sherff (G). 
IOWA. Fayette Co.: prairies, Sept. 5, 1894, B. Fink, 617 (US). 
CHEROKEE Co.: ca. 3 mis. s. of Cherokee, on upland slopes, Pilot 
Twsp., Sect. 23, Sept. 5, 1937, Miss A. Hayden, 10537 (G). 
JOHNSON Co.: without stated locality, Sept. 9, 1909, M. P. Somes, 
3873 (US). ARKANSAS. Cannon Co.: dry hillsides, Eureka 
Springs, Sept. 20, 1913, E. J. Palmer, 4404 (US). NEBRASKA. 
Pierce Co.: Plainview, Sept. 7, 1908, W. F. Petersen (US). 
KANSAS. SHaAwNEE Co.: Topeka, Aug. 22, 1877, E. A. 
Popinoe (US). 

X Liarris WrAvEnI Shinners (L. aspera X punctata). Stems 
slender, 12-24, from an elongate, penetrating rootstock ca. 10 
em. long, 4-5 dm. tall, mostly glabrous but with some whitish 
pubescence along the upper part of the flowering stalk and bearing 
many, punctate, linear to narrowly linear-lanceolate leaves as in 
L. punctata; lower ones 15 cm. long, 0.5-1 cm. wide, reduced 
gradually upwards to bracts exceeding the basal heads: inflores- 
cence 15-30 cm. long, dense and spike-like; heads numerous, 
campanulate to turbinate, 1.5-2 em. long and 1-1.5 em. wide, 
12-15-flowered; phyllaries linear-lanceolate, erect and moderately 
loose; outer ones herbaceous oblong or somewhat triangular with 
acute tips, 5-7 mm. long; middle and inner ones up to 1.5 cm. 
long and 3-5 mm. wide, oblong, blunt at tip, with a narrow 
scarious, erose, paler margin; corolla purple, 9-11 mm. long, 


1946] Gaiser,—The Genus Liatris 315 


quite pilose within; achenes ca. 5 mm. long; pappus 8 mm. long 
and plumose.—Amer. Mid. Nat. xxii. 38 (1943). 

The type of this hybrid was one plant (no. 16) of a seedling 
population grown from seeds (received in 1926, from Dr. E. J. 
Weaver and collected from uplands near Lincoln, Lancaster Co., 
Neb.) in 1927 at Crediton, Ontario by L. O. Gaiser. Plants no. 
4 and 17, also mentioned in the description by Shinners (l. c.), 
were of the same and of the 1928 population respectively grown 
likewise from seeds of the same package. Both Dr. Weaver and 
myself are at a loss to explain the location ‘“Locarina, Nebraska" 
given for the type specimen by Shinners. Other specimens from 
the same plants are now placed in the Gray Herbarium. 

The rest of the seedlings grown during the two successive 
years 1927 and 1928 were quite like the herbarium specimens of 
L. aspera Michx. var. typica received along with the seeds. 
Herbarium specimens and a package of seeds of L. punctata that 
were received from Dr. Weaver at the same time, from the same 
locality, proved to be variety nebraskana, having the characteris- 
tic narrow leaves, lacking prominent cilia on the margin and the 
slender heads with narrow, lanceolate phyllaries, also without 
marginal cilia. Thus it seems very probable that the three 
seedlings were the result of hybridization in the field of L. aspera 
var. typica and L. punctata var. nebraskana. As Shinners 
understood L. sphaeroidea in the same sense as L. aspera var. 
typica, as used here, the parentage given above is really the same 
as that given by him. 

These hybrids resemble L. punctata var. nebraskana in the 
tufted, numerous, slender stems, much shorter than those of L. 
aspera seedlings growing alongside, in the numerous punctate, 
linear leaves, the shorter, dense, spike-like inflorescence and the 
plumose pappus. In the pubescence along the upper part of the 
stem, the campanulate to turbinate heads of 12-15 flowers with 
phyllaries that are loosely erect, having the middle ones broader, 
narrowly scarious and blunt-tipped, and the length of the achene 
they resemble L. aspera. 

From viable seeds collected from each of the three seedling 
plants, F? populations were grown. "They were indeed à mixed 
lot showing sometimes greater resemblances to L. aspera and 
sometimes to L. punctata. 


316 Rhodora [OcroBER 


19. LIATRIS SCABRA (Greene) K. Schum. Stems straight, 
stiff, from a somewhat rounded corm 2 em. or more in diameter, 
covered with retrorse hairs and distinctly scabrous, generally 6-9 
dm. high, bearing about 20-30 usually short-peduncled heads in 
an open spike, rarely becoming paniculate: leaves scabrous on 
upper and lower surfaces, the lower 10-15 cm. long and 10-25 mm. 
wide, oblanceolate, narrowing to a winged petiole less than half 
the length of the blade; upper leaves bluntly lanceolate, 4—5 em. 
long and 5 mm. wide, non-petiolate and with clasping base, tend- 
ing to diverge from the stem at right angles: inflorescence of 
20-35-flowered heads, ca. 2 cm. long and 1.5-2 em. wide when 
flowers are open, somewhat cylindrical to turbinate, frequently 
all on short pedicels or with the basal becoming long-pedunculate; 
phyllaries entirely herbaceous and green, densely pubescent with 
short hairs or scabrous, margin ciliolate, the outer ovate to acute 
and sometimes recurved, the inner and middle oblong-obovate, 
spatulate with almost orbicular tips, generally erect and ap- 
pressed without any or with hardly any colored rim; corolla-tube 
pilose within, 10-15 mm. long; pappus 8-10 mm. long; achene ca. 
5 mm. long.—Just, Bot. Jahresb. xxix. 569 (1903).  Laciniaria 
scabra Greene, Pittonia, iv. 317 (1901). Laciniaria Shorti? Alex. 
in Small, Man. S. E. Fl. 1335 (1933) in part. Liatris aspera 
sensu Shinners, Amer. Mid. Nat. xxix. 34 (1943), not Michx.— 
Ohio and Illinois to Alabama and Mississippi, and westward 
into Arkansas and Oklahoma.—OHIO. Without stated locality, 
1842, C. W. Short (NY, type of Laciniaria Shortit Alex.). ERIE 
Co.: Castalia prairie, Sept. 19, 1909, E. L. Moseley (US). Orra- 
WA Co.: Port Clinton, Aug. 13, 1895, E. L. Moseley (G, US); 
Catawba Isl., Sept. 5, 1897, E. L. Moseley (US); between Port 
Clinton & Catawba Isl, Aug. 13, 1895, E. L. Moseley (US). 
FRANKLIN Co.: Georgesville, Aug. 29, 1892, W. C. Werner (NY). 
INDIANA. Cass Co.: along rwy., 1 mi. e. of Lake Cicott, 
Aug. 16, 1940, C. M. Ek (NY). Viao Co.: without stated locali- 
ty, W. S. Blatchley (US). Brown Co.: crest of open wooded 
knob, ca. 9 mis. s. e. of Nashville, Oct. 10, 1935, C. C. Deam, 
56936 (G). Jackson Co.: open chestnut oak ridge, 4 mis. n. w. 
of Medora, Sept. 4, 1934, R. M. Kriebel, 3022 (G). WASHING- 
TON Co.: s. slope of Quercus ridge, 12 mis. n. of Salem, Oct. 5, 15, 
1916, C. C. Deam, 22461 (US). Harrison Co.: sterile wooded 
slope, ca. 3 mis. n. of Elisabeth, Oct. 16, 1917, C. C. Deam, 
24374 (US); s. slope of Elisabeth Hill, 3 mis. e. of Elisabeth, Oct. 
13, 1916, C. C. Deam, 22429, 22432 (US). KENTUCKY. Co. 
undetermined: barrens of Kentucky, Sept. 1835, C. W. Short (G). 
Burtt Co.: Shepherdsville, Sept. 3, 1903, Biltmore Herb., 2670f 
(US). Epmonson Co.: dry woodlands, near Mammoth Cave, 
Sept. 21, 1903, Biltmore Herb., 2670i (US). MISSISSIPPI. 
OxTIBBEHA Co.: Starkville, Oct. 1, 1889, S. M. Tracy (US). 


1946] Gaiser,—The Genus Liatris 317 


ILLINOIS. Co. undetermined: pine hills, Sept. 23, 1890, F. S. 
Earle, ND, type). ARKANSAS. Puraskr Co.: north of White 
City Park, Little Rock, Sept. 19, 1931, D. Demaree, 8200, 8211 
(US); open dry woods, Pulaski Heights, Little Rock, Sept. 15, 
1931, D. Demaree, 8172 (G, NY, US); near White City Park, 
road to Quarry, Little Rock, Sept. 30, 1931, D. Demaree, 8333 
(US). GanraAND Co.: dry slopes of West Mt., Hot Springs, Oct. 
15, 1925, E. J. Palmer, 29229 (NY); dry woods, Sleepy Water 
Rd., Hot Springs, Sept. 8, 1935, F. J. Scully, 500 (G). JEFFER- 
son Co.: open pine-oak-hickory ridge woods, Pine Bluff, Oct. 1, 
1942, D. Demaree, 24099 (G). Crank Co.: high, dry, rocky 
wooded hill, Oct. 28, 1932, D. Demaree, 10001 (NY). ASHLEY 
Co.: knolls in open woods, Hamburg, Sept. 27, 1937, D. Demaree, 
16351 (NY); prairie-like regions, Fountain Hill, Oct. 11, 1937, D. 
Demaree, 13919 (O). LOUISIANA. NarcurirocHES Co.: dry 
open woods, Natchitoches, Oct. 3, 1915 (no. 8799), Oct. 7, 1917 
(no. 8888), E. J. Palmer (US). RaPrrpEs Co.: pinelands, Sept. 
10, 1900, Biltmore Herb., 2670n (US), Alexandria, Dr. Hale (NY), 
Alexandria, J. Hale (NY). OKLAHOMA. Le Frore Co.: 
severely cut pine-oak woods, Kiamichi Mts., July 17, 1930, E. L. 
Little & C. E. Olmstead, 608 (O). PusuMATAHA Co.: open areas 
in oak forest, Aug. 27, 1938, C. C. Smith, 918 (O). 


This species is quite unlike L. aspera Michx. in the densely 
scabrous stem and leaves, and the herbaceous, green, pubescent 
to scabrous phyllaries, non-scarious on the margin and mostly 
erect and appressed, or with only the outer ones squarrose, and 
none crisped or bullate. However, plants with involucral and 
leaf-characters intermediate between these two species have 
been seen from Indiana, Arkansas and Kansas, which are in the 
range of both; e. g. Oct. 10, 1935, C. C. Deam, no. 56936, from 9 
mis. e. of Nashville, Brown Co., Indiana (G); Sept. 12, 1895, J. B. 
Norton, no. 214, from prairie, Riley Co., Kansas (G), and Oct. 28, 
1932, D. Demaree, no. 10001, from Amity, Clark Co., Arkansas 
(G), though the specimen of the last collector's number and date 
at the New York Botanical Garden seems more nearly to re- 
semble L. scabra. 

L. scabra can be differentiated from L. scariosa by the arrange- 
ment of the phyllaries as well as the shape of the leaves; in L. 
scariosa of the Eastern States the basal leaves are broadly obo- 
vate. However, it is not difficult to think of L. scabra as a modi- 
fication of L. scariosa as it travelled westward. In fact it was 
rather startling to see how very much the heads and phyllaries 


318 Rhodora [OCTOBER 


of the Oklahoma plants from the Kiamichi Mts. resembled those 
of L. scariosa from the southern Appalachian Mts. 


20. LIATRIS LIGULISTYLIS (Nels.) K. Sch. Corm shallow, 
rounded, 1-4 cm. in diameter: stems single or several, 1-6 dm. 
high, glabrous below with white appressed pubescence on the 
usually reddish flowering stem or pubescent in most parts: 
leaves varying from glabrous to sparingly hispidulous along mid- 
vein of lower surface or to densely pubescent on both surfaces but 
always with cilia on the margin; basal leaves lanceolate-oblong or 
oblanceolate, 8-15 cm. long and 1-1.5 cm. wide, usually with a 
long margined petiole (the radical sometimes 20—40 cm. long); 
leaves reduced abruptly upwards, there more lanceolate, bract- 
like along a kind of strict spike: inflorescence of comparatively 
few heads (1-15) on short peduncles 1-5 cm. long, with the 
terminal head often much larger than the others, with varying 
conditions of moisture and soil the inflorescence bearing more 
heads with longer peduncles: heads broadly campanulate, be- 
coming hemispheric, 2-3 cm. wide, 40-70-flowered (terminal one 
sometimes twice that size); phyllaries glabrous, erect, with less 
rounded, more irregular, spatulate, broadly lacerate, scarious, 
usually colored tips, (sometimes the outer phyllaries of older 
heads appear to have been pressed out and backwards by the 
expansion of the maturing heads but in young heads are always 
erect); all phyllaries quite similar in texture though outer ones 
shorter, oblong, ovate or orbicular; middle and inner ones 
spatulate-oblong; corolla-tube nonpilose within, but occasionally 
a few hairs on the outside of base of tube, 9-11 mm. long; pappus 
8-10 mm. long, smoky purple when mature; achene 5-6 mm. 
long.—Just, Bot. Jahresb. xxix'. 569 (1903).  Laciniaria liguli- 
stylis Nels. Bot. Gaz. xxxi. 405 (1901). Liatris Rosendahlit 
Rydb. Brittonia, i. 100 (1931), not sensu Shinners, Amer. Midl. 
Nat. xxix. 40 (1943). Liatris Haywardii and L. Herrickii Rydb. 
(the latter a depauperate specimen) op. cit. i. 99 (1931).  Lacini- 
aria formosa Greene, Leaflets, i. 145 (1905). | Laciniaria scariosa 
(L.) Hill var. corymbulosa Sheld. Bull. Geol. & Nat. Hist. Surv. 
Minn. ix. 77, t. vi (1894), in part — forma corymbulosa Sheld. 
Quart. Bull. Univ. Minn. i. 27 (1892). Laciniaria scariosa vars. 
basilaris, supereminens, praeceps, praestans, multiplex, perusta, 
angustata and opima Lunell, Amer. Mid. Nat. ii. 92 (1911). 
Laciniaria scariosa var. scalaris Lunell, Amer. Mid. Nat. ii. 127 
(1911). Laciniaria scariosa var. subcorymbosa Lunell, Amer. 
Mid. Nat. ii. 158-9 (1912). Laciniaria scariosa forma uniflora 
Sheldon, Quart. Bull. Univ. Minn. i. 27 (1892), probably.  Lacin- 
taria scariosa var. uniflora Lunell, Amer. Mid. Nat. iii. 344 (1914). 
Laciniaria scariosa vars. exuberans, singularis, immanis, crista- 
galli, insolens, composita, annuens Lunell, Amer. Mid. Nat. v. 31- 


1946] Gaiser,—The Genus Liatris 319 


46 (1917). Laciniaria scariosa var. inconcinna Lunell, Amer. Mid. 
Nat. v. 241 (1919). Liatris ligulistylis f. leucantha Shinners, Amer. 
Mid. Nat. xxix. 39 (1943), albino, as was probably Laciniaria 
scariosa f. globosa Sheldon, Quart. Bull. Univ. Minn. i. 27 (1892) 
(this type not seen). 

Wisconsin, southern Manitoba, Saskatchewan, and Alberta!, 
Minnesota, North and South Dakota, Montana, Wyoming, 
Colorado, and Northern New Mexico.—WISCONSIN. Co. un- 
determined: Aug. 1844, S. B. Mead (NY). WavukEsHa Co.: 
Scuppernong Marsh, Aug. 4, 1941, H. C. Greene (G). JEFFER- 
SON Co.: C. M. & St. P. Rwy., embankment 24 mi. s. of Palmyra, 
July 29, 1940, L. H. Shinners, 2429 (G). MANITOBA. (bor- 
der of Ontario) Rainy Lake & River, Dr. Richardson, 9853 (Ot). 
62 F s. w.: Camp No. 7, South Antler Creek, Aug. 11, 1873, G. M. 
Dawson, Br. N. A. Boundary Comm. (G). 62 F 9: Souris, Aug. 
21, 1889, T. L. Walker (Q). 62 G 4: open prairie, n. of Killarney, 
Aug. 4, 1896, J. M. Macoun, 12435 (G, NY, Ot). 62 K 6: Bird- 
tail Creek, near Birtle, June 26, 1906, J. M. Macoun & W. 
Herriot 69884 (NY, Ot, US). 62 I1 3: border of marshes, Stoney 
Mt., Sept. 5, 1884, J. M. Macoun, 9855 (Ot). 62 K 1: open 
prairie, Rapid City, July 25, 1896, J. M. Macoun, 12241 (Ot). 
MINNESOTA. Betrrami Co.: Bemidge, July 27, 1925, L. H. 
Pammel, 876 (G). Hussar Co.: Benedict, July 31, 1914, H. J. 
Bergman, 2971 (G); Cass Lake, July 29, 1914, L. H. Pammel & 
H. E. Pammel, 671 (G); dry sterile sandy opening in jack pine, 
La Salle Springs, July 2, 1932, M. F. Buell, 503 (G); jack pine 
forest, 11 mis. s. of Hubbard Co. n. border, along Hwy. 71, Aug. 
9, 1941, J. W. Moore & D. L. Jacobs, 15099 (G). CLEAR WATER 
Co.: north boundary, Itasca Park, Aug. 18, 1929, M. L. Grant, 
3114 (G, NY, US); dry sandy soil, e. of La Salle Springs, Lake 
Itasca State Park, Aug. 17, 1918, C. O. Rosendahl, 3699 (M, 
type of Liatris Rosendahlii Rydb.). Becker Co.: brushland, 
Detroit, Aug. 14, 1914, Z. L. Chandonnet, 302 (US). OTTER 
Tait Co.: Silver Lake, Aug. 1892, E. P. Sheldon (G). HENNE- 
PIN Co.: Minneapolis, Aug. 1878, C. L. Herrick (M, type of 
Liatris Herrickit Rydb.). Wricur Co.: Cedar Lake, Sept. 1890, 
F. L. Holtz (M 211606 (type of Laciniaria scariosa forma uniflora 
Sheldon) and probably 211608 though now lacking head). 
Kanpiyou! Co.: Whitefield Twsp. Aug. 1, 1892, W. D. Frost 
(US 201918). LiwconN Co.: Verdi, Aug. 1891, E. P. Sheldon, 
81364 (M); Lake Benton, Aug. 1891, E. P. Sheldon, 81270 (M). 
NORTH DAKOTA. RorErTTE Co.: in dry soil, open borders 
of woodland, Turtle Mountains, near St. John, Aug. 30, 1909, 
J. Lunell, 1026 (M, type of Laciniaria scariosa var. perusta). 


1 Localities in Manitoba, Saskatchewan and Alberta are indicated by standard nota- 
tion with reference to sheets of the National Topographical Series, Dept. Mines & 
Resources, Ottawa, Canada. 


320 Rhodora [OCTOBER 


Ramsey Co.: in rich prairie soil, Devil’s Lake, Aug. 18, 1910, 
J. Lunell, 1021 ((M, type of Laciniaria scariosa var. supereminens 
Lunell) NY). McHenry Co.: Towner, Aug. 12, 1908, J. Lunell 
(NY), Aug. 12, 1920, J. Lunell, 1016 (M, type of Laciniaria 
scariosa var. basilaris Lunell); in rich soil, Towner, Aug. 12, 1908, 
J. Lunell, 1023 (M, type of Laciniaria scariosa var. praestans 
Lunell). Benson Co.: Leeds, Aug. 15, 1908, Aug. 6, 19, 1909, 
J. Lunell (G), Aug. 10, 1907, Aug. 6, 9, 10, 1909, J. Lunell (NY), 
Aug. 29, 1899, Aug. 10, 19, 1909, J. Lunell (US), Aug. 19, 1909, 
Aug. 20, 1915, J. Lunell (M); in rich soil, Leeds, Aug. 25, 1914, 
J. Lunell, 1018 (M, type of Laciniaria scariosa var. uniflora 
Lunell); dry soil, Leeds, Aug. 13, 1910, J. Lunell, 1022 (M, 
type of Laciniaria scariosa var. praeceps Lunell); moderately dry 
soil, Leeds, Aug. 15, 1909, J. Lunell, 1025 (M, type of Lacini- 
aria scariosa var.!multipler Lunell); in rich prairie loam, Leeds, 
Aug. 27, 1918, J. Lunell (M, type of Laciniaria scariosa var. 
inconcinna Lunell); in meadowland, along coulee, Leeds, Sept. 
8, 1908, J. Lunell, 1027 (M, type of Laciniaria scariosa var. 
angustata Lunell); in moderately moist, rich soil, Leeds, July 31, 
1909, J. Lunell, 1028 (M, type of Laciniaria scariosa var. scalaris 
Lunell); Leeds, Aug. 19, 1914, J. Lunell, 1031 (M, type of 
Laciniaria scariosa var. insolens Lunell); in rich meadowland, 
Leeds, Sept. 6, 1910, J. Lunell, 1035 (M, type of Laciniaria 
scariosa var. opima Lunell) ; Leeds, Sept. 9, 1916, J. Lunell, 1036 
(M, type of Laciniaria scariosa var. annuens Lunell); Butte, 
Aug. 9, 1908, Aug. 17, 1909, J. Lunell (G, US), Aug. 15, 1914, 
J. Lunell (D, Aug. 26, 1917, J. Lunell (M), Aug. 22, 1915, J. 
Lunell, 1019 (M, type of Laciniaria scariosa var. singularis 
Lunell), Sept. 3, 1916, J. Lunell, 1029 (M, type of Laciniaria 
scariosa var. immanis Lunell, July 29, 1906, J. Lunell, 1030 (M, 
type of Laciniaria scariosa var. crista-galli Lunell), Aug. 15, 1915, 
J. Lunell, 1032 (M, type of Laciniaria scariosa var. composita Lu- 
nell), Aug. 26, 1914, J. Lunell, 1034 (M, type of Laciniaria scari- 
osa var. subcorymbosa Lunell); Butte, Aug. 15, 1915, J. Lunell, 
1024 (M, type of Laciniaria scariosa var. exuberans Lunell). 
Kipperr Co.: Bird Lake, Dawson, Aug. 10, 1917, F. P. Metcalf, 
261 (US); Deer Lake, Aug. 15, 1917, F. P. Metcalf, 296 (US). 
Morton Co.: Mandan, Aug. 14, 1927, E. L. Larson, 157 (G). 
SOUTH DAKOTA. Co. undetermined: Black Hills, Miss Pratt, 
152 (NY). Grant Co.: virgin prairie, Clear Lake, near Big 
Stone Lake, Aug. 1, 1940, P. Johnson, 69 (G). Mane Co.: 
Black Hills near Fort Meade, July 28, 1887, W. H. Forwood (US). 
LAWRENCE Co.: limestone slope, Spearfish Canyon, H. E. Hay- 
ward, 166 (NY, type of Liatris Haywardii, Rydb.); Spearfish, 
Aug. 5, 1908, N. F. Petersen, 2 (US); high open woods, Spearfish 
Canyon, Aug. 7, 1908, N. F. Petersen, 3 (US); Iron Creek, Aug. 
7, 1908 (no. 3, 3a9, Aug. 14, 1908 (no. 3) N. F. Petersen (US); 


1946] Gaiser,—The Genus Liatris 321 


Black Hills, W. H. Forwood, 1447 (G); Deadwood to Sturgis Rd., 
T. 4 N. R. 3 E., Aug. 26, 1910, J. Murdock, 4325 (G); high hill- 
sides, Deadwood, July 31, 1913, W. P. Carr, 140 (G, NY, US); 
Deadwood, 1926, H. E. Hayward, 263 (NY); Nemo, 1912, S. S. 
Visher, 1560 (NY). Custer Co.: Custer, Black Hills (alt. 
5500’) Aug. 1, 1892, P. A. Rydberg, 753 (G, NY, US); Custer, 
Aug. 21, 1908, N. F. Petersen, 4 (US). SASKATCHEWAN. 
Without stated locality: Palliser’s Br. N. Amer. Exped., 1857-8, 
E. Bourgeau (G); Exact region undetermined: open prairies, near 
Hazel, July 11, 1906, J. M. Macoun & W. Herriot, 69885 (NY, 
US, Ot); prairies, bare hills (G. T. P. Rwy.), July 31, 1906, 
J. M. Macoun & W. Herriot, 69888 (NY, US, Ot); Round Valley 
Lake (G. T. P. Rwy.), Aug. 7, 1906, J. M. Macoun & W. Herriot, 
69889 (NY, US, Ot); prairies, 10 mis. w. of Round Valley Lake, 
Aug. 8, 1906, J. M. Macoun & W. Herriot, 69890 (US, Ot); dry 
gravelly soil, Old Wives Creek, July 25, 1880, J. M. Macoun, 
72719 (Ot). 72 P 9:30 mis. s. of Touchwood, July 18, 1906, J. 
M. Macoun & W. Herriot, 69886 (NY, Ot); borders of marshes, 
Touchwood Hills, Aug. 11, 1872, J. M. Macoun, 9856 (Ot). 
73 B 2: prairies w. of Saskatoon, July 29, 1906, J. M. Macoun & 
W. Herriot, 69887 (NY, US, Ot); 73 C 12-13: prairies, fresh 
water lake, e. of Lake Manito, Aug. 8, 1906, J. M. Macoun & W. 
Herriot, 69891 (NY, US, Ot); prairies, Lake Manito (G. T. P. 
Rwy.), Aug. 9, 1906, J. M. Macoun & W. Herriot, 69892 (NY, 
US, Ot). 73 D 15-16: Ribstone Creek (G. T. P. Rwy.), Aug. 
11, 1906, J. M. Macoun & W. Herriot, 69893 (NY, US, Ot); 
prairies, Ribstone Creek (G. T. P. Rwy.), Aug. 11, 1906, J. M. 
Macoun & W. Herriot, 69894 (NY, US, Ot); prairies, Ribstone 
Creek (G. T. P. Rwy.), Aug. 12, 1906, J. M. Macoun & W. 
Herriot, 69895 (G, NY, US, Ot); prairies, Ribstone Creek 
(G. T. P. Rwy.), Aug. 14, 1906, J. M. Macoun & W. Herriot, 
69897 (NY, US, Ot); 73 H 4: sandhills, n. of Prince Albert, July 
13, 1896, J. M. Macoun, 12749 (Ot); open sandy woodlands, 
Prince Albert, Aug. 15, 1913, F. W. Johnson, 1417 (US). 62E5: 
Weyburn prairie, A. B. Sanson, 135 (NY). 62 L 12: Indian 
Head, Aug. 14, 1895, W. Spreadborough, 10830 (Ot). 62 M 1: 
Yorkton, Aug. 19, 1908, W. Crawford (Q). 72 I 6: prairies, 
Thunder Creek, Moose Jaw, Aug. 27, 1895, W. Spreadborough, 
10831 (Ot). ALBERTA. $83H 11: dry prairie, edge of brush, 
Fort Saskatchewan, Aug. 15, 1935, G. H. Turner, 36 (G, NY); 
prairies, w. side of Beaver Hills, Aug. 23, 1906, J. M. Macoun & 
W. Herriot, 69898 (G, NY, US, Ot). 83 H 12: dry prairies, 
vicinity of Edmonton, Aug. 3, 1908, W. C. McCalla, 2527 (Ot). 
83 A 4: prairie, Red Deer, Aug. 1895, H. M. Gaetz, 10832 (Ot). 
MONTANA. SHERIDAN Co.: Westby, Aug. 9, 1928, Miss E. 
Larsen, 204 (US). WYOMING. Crook Co.: sandy pine 
woods, 6 mis. n. w. of Hulett, Sept. 14, 1937, M. Owenby, 1509 


322 Rhodora [OCTOBER 


(NY); open woods, Black Hills, July 23, 1910, A. Nelson, 9496 
(G, NY, US); Devil’s Tower, Aug. 9, 1897, L. W. Carter (US); 
Bear Lodge Mts., near Sundance, Aug. 9, 1897, L. W. Carter 
(US). ArnBANY Co.: Laramie Peak, Aug. 8, 1895, A. Nelson, 
1651 (G, NY, US, isotypes); meadows, Bacon’s Ranch, Aug. 15, 
1903, A. Nelson, 8925 (G, NY, US). COLORADO. Co. unde- 
termined: between Arkansas and South Platte River, July 29, 
1883, R. W. Woodward (G); Coyote Creek, Aug. 29, 1883, B. H. 
Smith (P); Arkansas Valley, Lieut. Wheeler’s Expedition, 1873, 
J. Wolf & J. T. Rothrock, 458 (G, US); latitude 39-41, 1862, 
Hall & Harbour (G); Colorado Terr. (latitude 39-41), 1864, 
C. C. Parry (O, US). Larimer Co.: foot of Longs Peak, 1862, 
C. C. Parry (G); Estes Park, Aug. 20, 1864, C. C. Parry (G); 
mountains (alt. 7500’), Aug. 19, 1895, C. S. Crandall (NY). 
Rovurr Co.: Trout Creek, 1873, J. Wolf, 458 (NY). Apams Co.: 
Denver, Aug. 1873, J. M. Coulter (P, US). AmaPAHOE Co.: 
South Park, 1871, W. M. Canby (G). BovurpER Co.: Sugar 
Loaf Mt. (alt. 8000’), 1902, F. Tweedy, 4935 (NY); JEFFERSON 
Co.: rocky hillsides, Aug. 30, 1888, N. G. Smith (US). Mesa 
Co.: Bridge Pass, 1856, H. Engelmann (G). Er Paso Co.: near 
Manitou, 1899, Mrs. C. N. S. Horner (G); Palmer Lake, 1896, 
Miss Hughes, 9 (G). Gunnison Co.: rich meadows, Parlin 
(alt. 8000’), Aug. 20, 1901, B. H. Smith, 110 (NY, P); Jack's 
Cabin, region of Gunnison watershed, July 26, 1901, C. F. Baker, 
610 (NY, US (ND type of Laciniaria formosa Greene)); Irwin 
Lakes, 1896, F. Clements, 378 (NY). Custer Co.: West Cliffe, 
1896, C. L. Shear, 3459 (NY). Hvuznrawo Co.: La Veta, 1896, 
C. L. Shear, 3561 (NY). ArAMosA Co.: Alamosa, 1896, F. 
Clements 122 (NY). ARcHULETA Co.: Pagosa Springs, Aug. 19, 
1893, B. H. Smith (P), Aug. 30, 1899, C. F. Baker, 691 (G, NY, 
US); grassy meadow near Dyke, Aug. 11, 1937, Marion Ownbey, 
1430 (NY). NEW MEXICO. Co. undetermined: Sierra 
Grande, Aug. 15, 1903, A. H. Howell, 212 (US). Corrax Co.: 
marshy hillside, vicinity of Ute Park, (alt. 2200-2900 m.), Sept. 
6, 1916, P. C. Standley, 14397 (G, NY, US); clay & shale in oak- 
pine woods, top of Raton Pass (alt. 7800’), Aug. 6, 1941, U. T. 
Waterfall, 3186 (G). San Miavzr Co.: Rociada, 1905, J. E. 
Dandelin (G). 


The description of Laciniaria ligulistylis differentiated this 
species clearly from other western ones. However, since Nelson 
described the leaves as being glabrous, subsequent writers in 
referring to plants of the northern mid-west region have dis- 
tinguished from this species similar plants that had pubescent 
leaves, as Rydberg in L. Rosendahlii and Lunell in the numerous 
varieties of Laciniaria scariosa cited above: When describing 


1946] Gaiser,—The Genus Liatris 323 


the plants of that species from North Dakota, Lunell (Amer. 
Mid. Nat. ii. 90 (1911)) stated generally: ‘‘The leaves . . . 
sometimes glabrate, but never perfectly glabrous" and again: 
“The Rocky Mountain forms as described by Prof. Aven Nelson 
(Liatris ligulistylis) are single-stemmed with glabrous leaves, else 
they appear in general characters to be near relatives of our 
plants". Dr. Nelson has kindly examined for the writer speci- 
mens received from Saskatchewan of which the leaves had not 
only a ciliate margin but also some pubescence. These he iden- 
tified as L. ligulistylis and in correspondence he stated: “I find 
that I have placed three of Lunell’s specimens under L. ligul- 
stylis. He distributed them as varieties of L. scariosa. I am 
wondering if these three may not have been included in Dr. 
Rydberg's L. Rosendahlii. I see no sufficient reason for keeping 
of any of these out of L. ligulistylis". The discovery that the 
corolla-tube of Liatris ligulistylis lacks any pilosity within, as 
does the type specimen of L. Rosendahlii Rydb. (C. O. Rosendahl, 
no. 3699, Itasca State Park, Minn. (M)) has given a fine test- 
character to accompany determinations, especially as all other 
members of the Scariosae series do have hairs in the corolla-tube 
near the base of the filaments. 

Examination of the type specimens of Lunell's varieties of 
Laciniaria scariosa kindly loaned me by Prof. Rosendahl of the 
University of Minnesota, has made it seem advisable to add to 
the synonymy of L. ligulistylis many of those varieties. Most 
of them were based on variations of leaf-shape and -arrangement, 
but excepting the character of pubecsent leaves, were said by 
Lunell to be “near relatives" of L. ligulistylis. On examination 
of the corolla-tubes, 19 of the type specimens of Lunell’s varieties 
from North Dakota and Minnesota were found to have no hairs 
within, and by other leaf-, head- and phyllary-characters to fall 
correctly under L. ligulistylis. These are found in the list of 
synonyms. Eight of the others (varieties brachiata, Chandonnetii 
nictitans, praecellens, propinqua, ramea, superans and superscan- 
dens) showed some pilosity and other characters giving evidence 
that they are intermediates between this species and L. aspera. 

Although the type specimen of Sheldon's Laciniaria scariosa 
var. corymbulosa (Leiberg, no. 27 from Mankato, Blue Earth Co., 
Minn. (M)) has unfortunately not been located, from the splendid 


324 Rhodora [OcroBER 


plate given of it (Bull. Geol. & Nat. Hist. Surv. Minn. ix. 77. t. 6 
(1894)) when raising it from formal to varietal rank, and from the 
examination of two of three specimens then cited (Aug. 1891, E. 
P. Sheldon, nos. 1270 and 1364 from Lakes Benton and Verdi 
respectively, Lincoln Co., Minn. (M)) which proved to be L. 
ligulistylis (Nels.) K. Sch., we are here including it in the synony- 
my of that species. However, one specimen cited (Aug. 1891, E. 
P. Sheldon, no. 1586 from Lake Benton, Lincoln Co., Minn. (M)) 
proved to be an intermediate, as are the specimens of Aug. 16, 
1901, L. R. Waldron and T. F. Manns from the vicinity of Fargo, 
N. Dak. (G, US). Successful hybridization between these two 
species has resulted in a wide variety of combinations of parental 
characters too inconstant to be typified in one hybrid description, 
yet easily recognizable (see no. 18). 

In the description of Liatris Haywardii Rydb. (Brittonia i. 99 
(1931)) from South Dakota there seems to be a close parallelism 
to L. Rosendahlii and L. ligulistylis in the size of the corm, 
leaves, bracts, achene and pappus. The outer phyllaries were 
described as elliptical and erose on the margin and the inner ones 
oblong. Rydberg has stated that it differed from L. aspera 
"jin the big heads often 2 cm. broad, and in the outer bracts 
which are twice as long as broad". Examination of the type 
specimen (H. E. Hayward, no. 166, Spearfish Canyon, Lawrence 
Co., South Dakota, (NY)) showed that the leaves were hirsute 
on the upper and lower surfaces, and the long phyllaries were 
loose and erect, as in the involucre of L. ligulistylis. By compari- 
son of the flowers it was found that Hayward's plant, as well as 
that of Miss Pratt (no. 153, from the Black Hills, South Dakota 
(NY)), also mentioned by Rydberg, lacked pilosity inside the 
corolla-tube and this gave further confirmation of the relation- 
ship to L. ligulistylis. Thus again, as in L. aspera (see no. 18), 
there seems to be a range of plants varying in all degrees of pu- 
bescence from glabrous to quite hirsute, but always roughened , 
by the cilia on the margin and without any clear-cut geographical 
limits, so that they can hardly be sharply separated into two 
varieties, as in that species, and it has seemed best to place them 
all under one species, L. ligulistylis. Sometimes plants are less 
robust and the heads do not acquire the usual size or there are 
other features of aberrancy. Such a specimen was described as 
Liatris Herrickii Rydb. (Brittonia, i. 99 (1931)). 


1946] Gaiser,—The Genus Liatris 325 


An interspecific hybrid of L. ligulistylis and L. punctata, 
X L. fallacior (Lunell) Rydb., will be discussed under L. punctata 
(see no. 24) but one of less certain parentage is here included. 


X Liatris Nieuwlandii (Lunell), stat. nov. Stems stiff, 6-10 
dm. tall, glabrous or sparingly pubescent below and quite pubes- 
cent above or densely pubescent throughout; leaves sometimes 
glabrous or quite pubescent on lower surface and sparingly so 
above, usually with a rough margin; basal ones broadly lanceo- 
late, 8-10 cm. long, gradually reduced upwards: inflorescence of 
few (9-20), sometimes of more numerous, large, sessile or pedi- 
cellate heads of 40-70 flowers; phyllaries erect, herbaceous, some- 
what cinereous, mostly green to the margins though sometimes 
quite purplish in color, broadly obovate or somewhat spatulate, 
almost non-petaloid or with but narrow petaloid or finely ciliolate 
margins; corolla 9-11 mm. long, generally non-pilose within the 
tube, though sometimes with scattered hairs; pappus 8-10 mm. 
long; achene 5-6 mm. long.—Laciniaria scariosa var. Nieuwlandii 
Lunell (including f. versicolor, f. gracillima and f. borealis (= f. 
septentrionalis ibid. p. 264)) and var. praesignis Lunell, Amer. 
Mid. Nat. ii. 169-177 (1912). Liatris novaeangliae var. Nieuw- 
landii (Lunell) Shinners, Amer. Mid. Nat. xxix. 31 (1943) 
including f. alba Shinners, ibid. 

From Michigan and Wisconsin southward into Ohio, Indiana, 
Illinois and Missouri.—MICHIGAN. Without stated locality: 
Dr. Crossman (G). CRAWFORD Co.: sandy jack-pine plains, 634 
mis. n. n. w. of Grayling, Sept. 14, 1935, F. J. Hermann, 7324 
(US); vicinity of Grayling, July 122, C. V. Piper (US). GRAND 
TRAVERSE Co.: Boardman Plains, Sept. 2, 1919, W. L. McAtee, 
3114 (US). LAPEER Co.: s. side of Lapeer, Aug. 17, 1911, C. K. 
Dodge (US 690400-1-2). INcHam Co.: college grounds, Agric. 
College, Sept. 20, 1890, C. F. Wheeler (US 63528). Wayne Co. 
7 Mile Road, Detroit, Nov. 13, 1911, J. A. Niewwland, 1680 
(ND type of Laciniaria scariosa (L.) Hill var. Niewwlandii f. 
septentrionalis Lunell). WasmTENAW Co.: Ypsilanti, Sept. 26, 
1857, W. Booit (G); steep grassy bank, 2.3 mis. n. e. of Ann 
Arbor, Sept. 5, 1937, E. J. Hermann, 9196 (NY). OHIO. 
Erie Co.: Castalia Prairie, Sept. 19, 1909, E. L. Moseley (US), 
Lucas Co.: near Toledo, Aug. 1879, J. A. Sanford, 1176 (NY). 
STARK Co.: Canton South Swamp, Sept. 1, 1912, Mrs. Case (G). 
FRANKLIN Co.: Columbus, W. S., 64 (G). INDIANA. Srev- 
BEN Co.: in a prairie condition, 44 mi. n. of Clear Lake, Aug. 21, 
1904, C. C. Deam (M, type of Laciniaria scariosa (L.) Hill var. 
Nieuwlandii f. versicolor Lunell) ((G, 12 sheets) NY); Sept. 11, 
1904 (M, type of Lacinzaria scariosa (L.) Hill var. Niewwlandii 
f. gracillima Lunell) ((G, 2 sheets) NY); without stated locality, 
Aug. 21, 1904, C. C. Deam (US 494351); in sandy woods, near 


326 Rhodora [OCTOBER 


Cedar Lake, Sept. 9, 1903, C. C. Deam (US 440343); without 
stated locality, Sept. 11, 1904, C. C. Deam (US 494350); gravelly 
roadside, on the e. side of Tamarack Lake, Aug. 19, 1916, C. C. 
Deam, 20918 (US); n. e. of Clear Lake, Aug. 18, 1916, C. C. Deam, 
20900 (US); e. of Hog Bog Lake, C. C. Deam, 20934 (US). La 
GRANGE Co.: on high bank of Pigeon R., 2 mis. e. of Ontario, 
Aug. 30, 1914, C. C. Deam, 15075 (US). Porter Co.: sandy soil 
along Erie Rwy. just w. of Kouts, Sept. 12, 1942, R. C. Friesner, 
17471 (G). Noste Co.: dry white-black oak woods, 14% mis. 
s. w. of Rome City, Aug. 26, 1914, C. C. Deam, 14776 (G, US). 
Kosciusko Co.: in a gravel pit, n. side of Winona Assembly 
Grounds, Aug. 17, 1906, C. C. Deam, 1496 (G); in sandy woods, 
w. side of Pike Lake, near Warsaw, Aug. 17, 1906, C. C. Deam, 
1524 (NY, US); w. side of Pike Lake, Sept. 16, 1906, C. C. Deam 
(US). MansHALL Co.: outlet of Lake Maxinkuckee, Sept. 5, 
1909, H. W. Clark (US). ALLEN Co.: prairie, 34 mi. s. of Fort 
Wayne, Aug. 19, 1900, C. C. Deam (M, type of Laciniaria scariosa 
(L. Hill var. N?ewwland?i Lunell). Cass Co.: sandy prairie 
patches along rwy. L$ mi. e. of L. Cicott, Sept. 26, 1936, R. 
Friesner, 10136 (NY). Warren Co.: stone bluffs of Pine Creek, 
2 mis. above Mudlavia, Sept. 11, 1911, C. C. Deam, 9986 (M, 
type of Laciniaria scariosa (L.) Hill var. praesignis Lunell). 
WISCONSIN. Watworts Co.: Lauderdale Lakes, Aug. 1890. 
M. S. Bebb (G): without stated locality, 1872, M. S. Bebb (US). 
Rock Co.: oak openings along the Chicago N. W. Rwy., Clinton, 
Sept. 1, 1909, E. S. Steele, 97a (G, US) 97b (G, US) 97e, 97d 
(US). ILLINOIS. Coox Co.: pebbly clay of Valparaioso 
moraine, vicinity of Palos Park, Sept. 6-7, 1909, E. S. Steele, 134a 
(G) 135 (G, US) 135e (G, US); dry hills, Palos Park, Sept. 6, 
1900, L. M. Umbach (US); thickets, Palos Park, Sept. 17, 1907, 
L. M. Umbach, 2119 (US). Hancock Co.: Augusta, Aug. 1844, 
S. B. Mead (NY). Mernarp Co.: without stated locality, Aug. 
1872, M. M. Milligan (US). MISSOURI. Prix Co.: McCune, 
Aug. 28, 1915, J. Davis, 249 (US). 


(To be continued) 


A SECOND STATION FOR CORYDALIS FLAVULA IN CONNECTICUT. 
—Corydalis flavula (syn. Capnoides flavulum) is not recorded in 
Gray's Manual, 7th Edition, or in Britton & Brown's Illustrated 
Flora, 2nd Edition, as found in New England. In 1925 Mr. 
Hugh S. Clark of Lancaster, Massachusetts, then a student at 
Wesleyan, found the plant on a stony ledge on Higby Mountain, 
a trap ridge in the western part of Middletown and Middlefield, 
Connecticut. He recorded the find in a note published in 


1946]  Fernald,— Does Bartonia verna grow in Virginia? 327 


Ruopona, 28: 68. The present writer was for a number of years 
active in trail work on Higby Mountain and other trap ridges in 
central Connecticut, and as an amateur botanist became fairly 
familiar with the plants on them. In or about 1932, while work- 
ing on the Mattabessett Trail on Lamentation Mountain, which 
is the next ridge north of Higby Mountain and which lies partly 
in the towns of Meriden, Berlin, and Middletown, but chiefly in 
Meriden, he discovered a specimen of the plant. This fact was 
noted in his flower-book and mentioned to Mr. Arthur E. 
Blewitt of Waterbury, Connecticut. Subsequent search for the 
plant was unsuccessful until April 28, 1946, when it was redis- 
covered by the writer when on a hike of the Connecticut Chapter 
of the Appalachian Mountain Club. It was found in several 
places, but especially in a large patch near the south end of the 
main ridge at an elevation of about 720 ft. Unlike C. semper- 
virens, which grows on and about exposed ledges and blossoms 
all summer and which grows in this same area, C. flavula is found 
in rich leaf-mold in the woods, which grow quite close to the edge 
of the escarpment. The plant is weak and sprawling and its 
blossoming period is from April into June. As its name indicates, 
the flowers are pale yellow. The species is only sparingly found 
in New York State. The counties listed for it in House's Anno- 
tated List of the Ferns and Flowering Plants of New York State 
are Ulster, Dutchess, Orange, Westchester, and Rockland. It 
has also been found on Manhasset Neck, Long Island.—FnEp- 
ERICK W. KILBOURNE, Cheshire, Connecticut. 


Dors BARTONIA VERNA GROW IN VrRGINIA?—[In 1803 Michaux 
described and illustrated his CENTAURELLA VERNA Michx. Fl. 
Bor.-Am. i. 98, t. 12, fig. 2 (1803), from sphagnous habitats in 
South Carolina (in sphagnosis Carolinae inferioris). In 1814 
Pursh, Fl. Am. Sept. i. 99 (1814), changed the name to C. vernalis 
and on p. 100 described C. vernalis 8. uniflora "caule unifloro’’, 
giving for the two plants the range “In mossy swamps: Virginia 
to Georgia. © May-July . . . Var. B. v. s. in Herb. Lyon." 
Bartonia verna (Michx.) Muhl., based on Centaurella verna 
Michx., has subsequently been regularly treated as coming north 
into Virginia. Nevertheless, painstaking search (always with 


328 Rhodora [OcroBER 


this “spook” in mind) through 14 seasons in hundreds of proper 
habitats in southeastern Virginia has failed to bring the plant to 
light, although B. virginica (L.) BSP. is there abundant and the 
often larger-flowered B. paniculata (Michx.) Muhl. especially so. 
In the Gray Herbarium the most northern definite station 
represented for B. verna is Wilmington in southeastern North 
Carolina. The flowering dates on the labels of B. verna range 
from December 26 (Florida) to April 3 (South Carolina), which 
would lead one to expect it in southeastern Virginia in late March 
and April. Small says *"Winter-spring". 

It is, consequently, significant that Pursh said for his Cent- 
aurella vernalis *May-July" and that he did not differentiate 
the very characteristic C. paniculata Michx. (Bartonia paniculata 
(Michx.) Muhl.) from B. virginica (L.) BSP. (Sagina virginica L., 
Bartonia tenella Muhl.), Pursh merging these two species as his 
Centaurella autumnalis. In view of the abundance “In mossy 
swamps" of southeastern Virginia of Bartonia paniculata and B. 
virginica, which begin flowering in July, it seems not improbable 
that Pursh's “July” and his Virginia record were not based on 
actual B. verna. If any actual evidence of the truly vernal B. 
verna in Virginia exists I shall welcome the information.—M. L. 
FERNALD. 


Volume 48, no. 573, including pages 201-264 and plates 1049 and 1050, was 
issued 12 September, 1946. 


NOV 13 1946 


Hovova 


JOURNAL OF THE 


NEW ENGLAND BOTANICAL CLUB 


Conducted and published for the Club, by 
MERRITT LYNDON FERNALD, Editor-in-Chief 


CHARLES ALFRED WEATHERBY 
ALBERT FREDERICK HILL > Associate Editors 
STUART KIMBALL HARRIS 


Vol. 48. November, 1946. No. 575. 


CONTENTS: 


Vegetation of Artificial Lakes in Northwestern Arkansas. 


J.B. Moyle, E. L. Nielsen and O. R. Younge. ...... ........ 329 
Senecio tomentosus, forma alabamensis. M. L. Fernald. ....... 330 
The Genus Liatris. L.O. Gaiser (continued). .............. wee 33l 
Presumable Identity of Cheilanthes lanosa. M. L. Fernald. .... 383 
Some trivial American Forms of the Lady-fern. M. L. Fernald. 389 
Centaurea maculosa in Indiana. Edwin D. Hull. .............. 391 
Setaria Faberii in North Carolina. Carroll E. Wood, Jr. ...... 301 


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JOURNAL OF 


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Vol. 48 November, 1946. No. 575. 


VEGETATION OF ARTIFICIAL LAKES IN 
NORTHWESTERN ARKANSAS 


J. B. Movrz, E. L. Nersen AND O. R. Younce! 


During recent years several artificial lakes or water reservoirs 
have been created in northwestern Arkansas. Aquatic and 
marsh vegetation has rapidly invaded these newly flooded areas 
which were formerly farm and forest land. In 1939, extensive 
collecting was done at four of these reservoirs to determine the 
nature and extent of natural revegetation. The reservoirs con- 
sidered include Lake Wedington, near Fayetteville in Washington 
County; Lake Atalanta, near Rogers in Benton County; the 
reservoir at Cave Springs in Benton County and that in Devil’s 
Den State Park, Washington County. Lake Wedington and 
Lake Atalanta each cover about 80 acres and have a maximum 
depth of about 40 feet. The reservoirs at Cave Springs and 
Devil’s Den have areas of about 3 and 5 acres, respectively, and 
a depth of about 15 feet. 

Two years after Lake Wedington began to fill, 39 species of 
aquatic and wet soil plants had established themselves. The 
commonest submerged species were Chara Braunii Gmel., Najas 
guadalupensis (Spreng. Morong and Potamogeton Berchtold: 
Fieber var. tenuissimus (Mert. & Koch) Fern. The most com- 

1 Aquatic Biologist, Bureau Fisheries Research, Minnesota Department of Con- 
servation, St. Paul; Associate Agronomist, Division of Forage Crops and Diseases, 
U. S. Department of Agriculture, Soils and Agricultural Engineering, Madison, Wis. 
and Lancaster, Ky., respectively. The collections were made by Nielsen and Younge 


and the specific determinations by Moyle. Specimens are deposited in the Herbarium 
of the University of Minnesota. 


330 Rhodora [NOVEMBER 


mon shoreline species was Eleocharis obtusa (Willd.) Schult. 
Other species collected include Typha latifolia L., Potamogeton 
nodosus Poiret, P. natans L., Alisma Plantago-aquatica L., Lopho- 
tocarpus calycinus (Engelm.) J. G. Smith, Sagittaria ambigua J. G. 
Smith, S. graminea Michx., S. latifolia Willd., S. rigida Pursh, 
Anacharis occidentalis (Pursh) Victorin, Eragrostis hypnoides 
(Lam.) BSP., Echinochloa pungens (Poiret) Rydberg, E. colonum 
(L.) Link, Leersia oryzoides (L.) Swartz, Carex gynandra Schwein., 
Cyperus acuminatus Torr. & Hook., C. esculentus L., C. ovularis 
(Michx.) Torr., C. odoratus L., Commelina hirtella Vahl, Juncus 
diffusissimus Buckley, J. effusus L., J. interior Wiegand, J. 
nodatus Coville, Polygonum coccineum Muhl., P. Persicaria L., 
P. punctatum Ell., Ceratophyllum demersum L., Nelumbo penta- 
petala Walt., Ludwigia alternifolia L., Nymphoides peltatum 
(Gmel.) Britten & Rendle, Heliotropium indicum L., Lindernia 
anagallidea (Michx.) Pennell, Utricularia biflora Lam. and 
Bidens frondosa L. 

A year after the filling of Lake Atalanta the following six 
species of aquatic and shoreline plants were present: Potamogeton 
foliosus Raf. var. genuinus Fern., Kyllinga pumila Michx., 
Cyperus esculentus L., C. strigosus L. Leersia oryzoides (L.) 
Swartz and Nasturtium officinale R. Br. The Devil’s Den reser- 
voir, two years after filling, had a sparse flora of Chara sp. and 
Juncus effusus L. The exact age of the reservoir at Cave Springs, 
which is somewhat older, was not ascertained. Eight species 
were collected here. These included Chara fragilis Desvaux, 
Typha latifolia L., Potamogeton foliosus Raf. var. genuinus Fern, 
Leersia oryzoides (L.) Swartz, Cyperus strigosus L., Scirpus pal- 
lidus (Britton) Fern., Lycopus rubellus Moench and Mentha 
canadensis L. 


SENECIO TOMENTOSUS Michx., forma alabamensis (Britton) 
stat. nov. S. alabamensis Britton ex Small, Fl. Se. U. S. 1305 
(1903). 

Several collections from eastern Virginia, such as Grimes, no. 
3552 from James City County, Heller, no. 850 from Isle of Wight 
County, and Fernald & Moore, nos. 15,162 and 15,163 from 
Nansemond County, show that S. alabamensis is at best an ex- 
treme glabrous or glabrescent form of the generally tomentose 


1946] Gaiser,— The Genus Liatris 331 


S. tomentosus. Its stout caudex, coarse fistulous and soon very 
brittle and fractured stems, its leaf-outline and toothing, inflores- 
cence, and hirtellous columnar achenes are inseparable from those 
of the much commoner typical S. tomentosus.—M. L. FERNALD. 


THE GENUS LIATRIS 
L. O. GAISER 
(Continued from page 263) 


With L. ligulistylis (Nels.) K. Sch. abundant in the prairie 
provinces and the states immediately adjacent (Wisconsin, 
Minnesota, North and South Dakota) and thence southwestward 
through Wyoming, Colorado and northern New Mexico, there 
are found in Michigan, Illinois, Indiana and Ohio, (a geographi- 
cally central region for the series Scariosae), what may be 
intermediates between L. ligulistylis and another species. Lunell 
described Laciniaria scariosa var. Niewwlandii as having stem 
not unusually 1 m. high, simple or even branched (then with 
few-headed branches), with a few-headed, short-pedicelled, 
narrow short inflorescence to ample many-headed, long-pedi- 
celled or long-branched one with an ample green foliage . 
involueral braets bright green over the whole surface or with 
narrow almost entire purple margins. Examination of the type 
sbowed a striking similarity in the nine large heads to a robust 
form of L. ligulistylis with herbaceous phyllaries. That modifica- 
tions of the type are easily found was recognized by Lunell since 
he described 3 forms: f. borealis (renamed septentrionalis (Amer. 
Mid. Nat. ii. 264 (1912)), f. versicolor and f. gracillima. In the 
note regarding f. borealis, Lunell made this remark: the “short 
stem and short, few-headed raceme bespeak its northern origin". 
The type of forma borealis Nov. 13, 1911, J. H. Nieuwland, no. 
1680, from 7 Mile Road, Detroit, Michigan (N. D.) has 10 heads, 
and a few buds, with erect, suborbicular, only slightly scarious- 
margined phyllaries, and again resembles L. ligulistylis in general 
habit, though the leaves are numerous and linear-lanceolate, sug- 
gesting possibly a blending with L. borealis of the New England 
States. Other specimens seen have given a similar suggestion, 
as Dr. Crossman, without locality, Michigan (G). However, 


332 Rhodora . [NOVEMBER 


mostly the specimens, especially those from Michigan, have 
broadly lanceolate basal leaves and large few-headed inflores- 
cences with erect phyllaries that at least associate them with L. 
ligulistylis. We regard them as a hybrid of L. ligulistylis and 
possibly L. borealis. Examination of the corolla-tubes of the 
type specimens of all three formae showed occasional hairs inside 
rather than the very definite non-pilose condition in L. liguli- 
stylis and thus too gave evidence of hybridity. 

In the two complete plants on the type sheet of Lunell’s 
Laciniaria scariosa var. praesignis (of C. C. Deam, no. 9986, 
from stone bluffs of Pine Creek above Mudlavia, Warren Co., 
Ind. (M)) there seem to be clearly combined the leaf-characters 
and few heads in an almost strict spike of L. ligulistylis with 
erect phyllaries having finely ciliolate margins and though they 
appear to be somewhat depauperate specimens we place this 
variety here in synonymy. A third incomplete plant on the 
sheet with an inflorescence of more numerous (19) heads, having 
phyllaries that are pubescent on the surface could be an inter- 
mediate of L. scabra and L. aspera (see no. 19). 


21. LIATRIS BOREALIS Nutt. ex MacNab. Stem glabrous or 
sparingly pubescent with appressed hairs, somewhat striate, 4-6 
dm. high from a rounded corm up to 5 em. in diameter: leaves 
numerous, often twisted, glabrous or with sparse distribution of 
hairs along the midrib beneath or on the lower surface and ciliate 
along the margin, the basal lanceolate, 1-1.5 dm. long and 1 em. 
wide, merely narrowed to a clasping base, hardly petiolate; upper 
cauline leaves numerous, linear-lanceolate, sessile, gradually 
reduced in length to bracts 2-3 cm. long, subtending the flower- 
heads: inflorescence with rachis pubescent, 1-3 dm. long, of 
5-30 short- or longer-pedicelled heads, of 35-60 flowers, broadly 
campanulate to hemispherical in form because of the loosely 
appressed but erect, non-recurved, rounded to oblong phyllaries; 
basal herbaceous outer phyllaries ovate to rounded, sometimes 
slightly pubescent like the pedicels, with ciliolate, rarely colored 
margins; middle and inner phyllaries rounded to linear, thinner, 
often marginally colored, though varying from narrowly to 
hardly at all petaloid, with a finely ciliolate margin (where the 
phyllaries are more petaloid the fine ciliolae practically disap- 
pear); corolla purple, rarely white, tube scantily pilose within, 
9-10 mm. long; pappus ca. 8 mm. long; achene 6-7 mm. long.— 
L. borealis Nutt. ex MacNab, Edinb. New Phil. Jour. xix. 60 
(1835). L. scariosa Q intermedia Ell. Sk. ii. 281 (1824). L. 
scariosa of Edwards, Bot. Reg. t. 590 (1821) and Lindley, Bot. 


1946] Gaiser,— The Genus Liatris 333 


Reg. t. 156 (1835), not (L.) Willd. Liatris squarrosa sensu 
Sweet, Br. Fl. Gard. t. 44 (1824), not (L.) Willd. Laciniaria 
scariosa var. novae-angliae Lunell, Amer. Mid. Nat. ii. 172, 177 
(1912). Liatris novae-angliae (Lunell) Shinners, Amer. Mid. 
Nat. xxix. 29 (1943). L. novae-angliae f. albiflora Shinners, 
Amer. Mid. Nat. xxix. 29 (1943). 

Southwestern Maine to Pennsylvania.—M AINE. Without 
stated locality, Aug. 21, 1927, E. E. Perkins (US). Yorx Co.: 
North Berwick, Sept. 5, 1891, J. C. Parlin (G). NEW HAMP- 
SHIRE. Without stated locality, Aug.-Sept., Little (P). MAS- 
SACHUSETTS. Essex Co.: Ipswich, Oakes (G, NY, P); in the 
fields, Ipswich, 1816, Dr. Boott, 1353 (US); Salem, Nuttall (P, 
type); roadside, Boxford, Sept. 2, 1887, G. G. Kennedy (G, P); 
Boxford, Aug. 1893, F. H. Peabody (G); Boxford (albino), Sept. 
10, 1909, A. L. Page (G). MippLEsEx Co.: Tewksbury, Sept. 24, 
1899, E. F. Williams, 2 (G); Westford, E. F. Fletcher (G); Con- 
cord, Aug. 30, 1888, C. E. Faxon (G); old graveyard, Malden, 
C. E. Faxon (G). Worcester Co.: border of Wachusett 
Reservoir, Boylston, 1930, E. W. Bemis (G); Rutland, Sept. 6, 
1926, R. H. Piper, 283 (O). FRANKLIN Co.: New Salem, Aug. 
21, 1929, A. S. Goodale, W. Markert & R. H. Piper, 55731 (G). 
NonroLk Co.: Dedham, Islington Junction, Aug. 22, 1897, E. F. 
Williams, 1 (G). HaAMPrsHinE Co.: Prescott, Aug. 26, 1929, R. 
H. Piper, 517 (O). HaMPDEN Co.: sparsely wooded, sandy 
plain, Southwick, Sept. 14, 1914, F. C. Seymour, 279 (NY). 
BARNSTABLE Co.: dry, sandy field,s. of Blueberry Pond, Brewster, 
Sept. 12, 1927, M. L. Fernald, 490 (G, NY, P, O); dry soil, near 
mouth of Red River, Harwich, Aug. 25, 1918, M. L. Fernald & 
B. Long, 17455 (G, P); dry argillaceous fields, n. of No Bottom 
Pond, Brewster, Sept. 7, 1918, M. L. Fernald & B. Long, 17456 
(G, NY, P); Chatham, M. L. Fernald & B. Long, 19175, 10505 
(P); Quissett Harbour, Sept. 6, 1923, M. L. Fernald, B. Long & 
J. M. Fogg (P), Sept. 20, 1918, M. L. Fernald & C. A. Weatherby, 
17457 (G); Oyster Pond, Falmouth, F. W. Pennell, 3497 (P); 
s. w. of Barnstable, Sept. 16-17, 1916, R. C. Bean, F. W. Bird & 
C. H. Knowlton (P); North Falmouth, F. W. Pennell, L. A. 
Kenoyer, 3466 (P); dry soil, W. Falmouth, Aug. 27, 1906, J. A. 
Cushman (Q); Hyannis Point, Sept. 5, 1898, J. M. Greenman, 
441 (G). PrLvMwourH Co.: sandy woods and thickets, near 
Rickard's Pond, Carver, Aug. 30, 1913, M. L. Fernald, F. W. 
Hunnewell & B. Long, 10503 (G, P); Plymouth, Sept. 17, 1889, 
J. H. Redfied (P), Mr. Gilbert (G); Marion, Sept. 1888, Miss A. 
M. Vail (NY); Brockton, Sept. 27, 1901, C. B. Blomberg (US). 
NANTUCKET Co.: Wauwinet, Sept. 8, 1894, E. F. Williams (P); 
moorland, F. W. Pennell, 11002 (P); dry roadside, Sept. 4, 1913, 
Miss E. M. Moody (G); roadsides, Aug. 30, 1897, Miss F. N. 
Vasey (US); without stated locality, Sept. 14, 1899, E. P. Bicknell 


334 Rhodora [NOVEMBER 


(NY). Duxe Co.: Chappaquiddick Isl., Aug. 1898, A. Hollick 
(G, NY). RHODE ISLAND. Proviprencr Co.: Cumberland 
(Manville), 1880, G. Hunt & W. W. Bailey, 175 (US); Warwick 
Twsp., Sept. 5, 1875, Amer. Mus. Nat. Hist. (NY). Kent Co.: 
Johnston Twsp., Aug. 8, 1878, G. C. Capron (NY); Coventry, 
Aug. 1895, J. F. Smith (US). Newport Co.: dry hillside, near 
New Shoreham Center, Block Isl., Sept. 15, 1913, M. L. Fernald, 
B. Long & G. S. Torrey, 10506 (G, NY, P); Block Isl., 1916, 
Gravatt (US). CONNECTICUT. | WiNpHAM Co.: dry grav- 
elly soil, Putnam, Sept. 5, 1908, C. H. Bissell & C. A. Weatherby 
(G). New Lonpon Co.: sandy soil, s. of Westchester Sta., 
Colchester, Sept. 23, 1904, C. B. Graves (G). New Haven Co.: 
dry soil, near coast, Guilford, Sept. 16, 1906, G. H. Bartlett (G); 
Stratford, Sept. 9, 1895, E. H. Eames (G); Stratford (albino), 
Oct. 5, 1927, E. H. Eames, 10402 (G); New Haven, Sept., 1874, 
A. H. Young, 23 (ND, type of Laciniaria scariosa var. novae- 
angliae Lunell); in sandy waste, near New Haven, Nov. 9, 1879, 
J. A. Allen (P). FarnrrELD Co.: Fairfield, Sept. 9, 1893, E. H. 
Eames (US). NEW JERSEY. Monmouts Co.: Middletown, 
Sept. 1838, Torr. & Gray (G). NEW YORK. Co. undeter- 
mined: Long Isl., J. Torrey (P). ArnBANY Co.: sandy soil, near 
Londonville, Sept. 15, 1932, H. D. House, 20173 (G, NY). 
QuEENS Co.: Old Mill Yacht Club (albino), Sept. 8, 1940, H. N. 
Moldenke, 11606 (NY). New York Co.: Harlem River, Sept. 
23, 1865, W. H. Leggett (NY). Surronk Co.: Greenport, Sept. 
1870, W. G. Farlow (G); Aquebogue, Sept. 1873, H. M. Young 
(US); in wet pockets, among sand dunes, Easthampton, Long 
Isl., Aug. 18, 1938, W. C. Muenscher & O. F. Curtis, 6603 (G, 
NY); sandy pine woods, East Islip, Aug. 31, 1938, W. C. Muen- 
scher & O. F. Curtis, 6602 (US); near MacKay Radio Sta., Aug. 
18, 1938, W. C. Muenscher & O. F. Curtis, 6603 (G), 6602 (P). 
Nassau Co.: near Flushing, Sept. 17, 1936, J. Monachino, 175 
(O); Locust Grove, Sept. 3, 1936, S. A. Cain (NY). PENNSYL- 
VANIA. Centre Co.: dry, rocky, grassy bank in Half Moon 
Valley, 2 mis. n. of State College, Aug. 18, 1938, H. A. Wahl, 
226 (G). 

MacNab (Edin. New Phil. Jour. xix. 60 (1835)) in publishing 
Liatris borealis stated: ** While looking through the herbarium of 
Nuttall when at Philadelphia, I found that the species was 
marked L. borealis but no description of it has yet been pub- 
lished”. 

Presumably the type specimen is at the British Museum in 
London where Nuttall’s Herbarium is, but Mr. Weatherby 
did not find it when obtaining photographs of others of Nuttall’s 
types in the summer of 1939. There is a specimen in the Her- 


1946] Gaiser,—The Genus Liatris 335 


barium of the Philadelphia Academy of Sciences from Salem, 
Mass., named Liatris borealis in Nuttall’s handwriting. The 
name borealis has been crossed out, but Dr. Pennell' stated: 
* the width of the line made, makes me think it was not Nuttall 
himself". This specimen may then be the type. 

The plant in Philadelphia, though incomplete, shows there had 
been 7 heads on pedicels about equal in length to the one remain- 
ing head, 2 cm. long, with generally erect phyllaries. The outer 
ones are hardly at all scarious, finely ciliolate; the inner linear- 
oblong ones becoming a little more scarious but still having a 
fine ciliolate margin. The few remaining cauline leaves are 
narrowly linear and sessile having a few hairs scattered over the 
surface. 

From MacNab's description, Liatris borealis had a simple 
pubescent stem; ovate-lanceolate, acuminate, long-petiolate 
radical leaves; stem-leaves sessile, narrowed toward both ends 
and rough on the margins; outer phyllaries obovate with colored 
margins; inner ones linear, equal in length to the heads; heads on 
rather long pedicels, remote, alternating in two rows. ‘Habitat: 
dry woods and rocky places throughout the Alleghany moun- 
tains". Having omitted mention of the character of the phyl- 
laries in this description, he however adds the note: ‘‘ This species 
is readily distinguished from L. scar/osa (which by some is 
supposed to be only a variety), by its being very much smoother, 
having fewer leaves and flowers and reflected bracteas; also by 
its short styles which scarcely exceed the corolla, whereas in L. 
scariosa the styles are long and the leaves, flowers and bracts are 
all upright”. The phyllaries (“bracteas”) of Nuttall’s plant from 
Salem are clearly not reflexed, whereas those of Liatris scariosa, 
as seen in the Linnaean type (see no. 17), certainly are. Paxton, 
(Paxt. Mag. v. t. 27 (1838)) so briefly described Liatris borealis, 
accompanied by a plate, from a plant cultivated in a nursery at 
Epsom, Scotland, received in 1836 from the Glasgow Botanical 
Garden, where it was presumed to have been introduced by Mr. 
Drummond, that it is not determinative. Torr. & Gray (Fl. N. 
Am. ii. 75 (1841)) referred Liatris borealis of Paxton to L. scariosa, 
and Gray (Synop. Fl. 1?, 110 (1884)) did the same. I do not 
find any reference in either work to L. borealis MacNab. ‘That 


1 By private communication, 


336 Rhodora [NOVEMBER 


Paxton’s plant was identical with Nuttall’s is doubtful. His de- 
scription and accompanying plate seem to be nearer L. scariosa 
var. typica, but in any case MacNab’s description is earlier and 
hence must have precedence. Whether MacNab wrote his de- 
scription from brief notes made on Nuttall’s plant or whether 
there was another plant is not known, but the essential character 
which he emphasizes in his discussion of L. borealis, that differs 
from the plant we have seen in the Philadelphia Academy, 
labelled in Nuttall’s handwriting, is the reflexed rather than erect 
phyllaries. The description above embodies the writer’s ideas of 
Liatris borealis Nutt. Examination of the type specimen of La- 
ciniaria scariosa var. novae-angliae Lunell (Amer. Mid. Nat. ii. 
172, 177 (1912)) showed it to be the same species as Nuttall’s 
plant from Salem, Mass. 


22. Liatris EARLEI (Greene) K. Sch. Corm small, subglo- 
bose, ca. 2 cm. in diameter: stem usually single, 4-9 dm. high, 
often somewhat virgate, softly and densely pubescent, or asperous 
with short white hairs, or even almost glabrous: leaves glabrous, 
softly pubescent and rough on the margins only, or more rarely 
scabrous, the basal sharply lanceolate, 10-20 cm. long, 5-15 cm. 
wide, subpetiolate, narrowing to a short- or longer-winged petiole 
from one third to one half the length of the blade, the upper 
rigid, sharp-pointed, narrowly lanceolate or linear, reduced from 
6-8 cm. long to narrow bracts less than 1 cm. subtending the 
heads: inflorescence narrowly racemose, of 20-50 subsessile heads 
on erect or depressed pedicels about as long as the heads, or more 
rarely becoming branched and paniculate by the elongation of 
the pedicels into slender peduncles bearing several heads each; 
heads of 15-25 flowers, somewhat turbinate or campanulate, 
1-1.5 cm. long and ca. 1 em. wide when flowers are open: phyl- 
laries appressed or sometimes recurved, herbaceous, green and 
softly pubescent, with ciliolate but almost non-scarious margins; 
outer phyllaries subovate; middle and inner ones oblong-spatu- 
late, 7-8 mm. long and 2-3 mm. wide, herbaceous and usually 
finely pubescent, sometimes . with purplish ciliolate margin; 
corolla-tube 7-9 mm. long, scantily to moderately pilose at the 
base of the tube: mature achene 3-4 mm. long; pappus about 6 
mm. long, barbellate.—Just, Bot. Jahresb. xxix!. 569 (1903). 
Laciniaria Earlei Greene, Pittonia, iv. 316 (1901). Laciniaria 
Tracyi Alex. ex Small, Man. S. E. Fl. 1335 (1933). Laciniaria 
Ruthii Alex. l. c., not Lacinaria Ruthii Bush, Amer. Midl. Nat. 
xii. 316 (1931). Liatris squarrulosa sensu Shinners, Amer. Mid]. 
Nat. xxix. 33 (1943), not Michx. 

Occasional from North Carolina to Florida, abundant from 


1946] Gaiser,—The Genus Liatris 337 


Tennessee to Alabama, and again occasional westward to 
Indiana, Louisiana and Texas—NORTH CAROLINA. Co. 
undetermined: s. slopes of Joanna Bald, Sept. 20, 1900, 22 (NY). 
Haywoop Co.: Waynesville, 1897, T. G. Harbison (G). CHERO- 
KEE Co.: 3-4 mis. e. of Andrews, Sept. 20, 1900, 21 (NY). 
SOUTH CAROLINA. BravronT Co.: sandy soil, Sept. 6, 
1904, Biltmore Herb., 2670k (US). GEORGIA. RICHMOND 
Co.: oakwoods, Augusta, Sept. 25, Oct. 10, 1898, A. Cuthbert 
(NY). FLORIDA. Without stated locality: in rich soil, 
Chapman (G). INDIANA. Harrison Co.: s. slope of Elisa- 
beth Hill, 3 mis. e. of Elisabeth, Oct. 13, 1916, C. C. Deam, 
22429 (US). KENTUCKY. Logan Co.: rocky hill near Rus- 
sellville, Sept. 17, 1903, Biltmore Herb., 2670h (US). TENNES- 
SEE. Co. undetermined: Hiwassee Valley, Oct. 1896, A. Ruth, 
34 (NY, type of Laciniaria Ruthii Alex.); sandy ground, Hiwas- 
see Valley, Oct. 1896, A. Ruth (US); mts., E. Tennessee, Smoky 
Range, Sept. 1897, A. Ruth, 3767 (NY); dry soil, White Cliff 
Springs, E. Tennessee, Aug. 1879, G. Andrews (US); sandy soil, 
Higdon, E. Tennessee, Oct. 1895, A. Ruth (US). Knox Co.: 
thickets, Knoxville, June 1897 (no. 3766), Aug. 1896 (no. 36), 
A. Ruth (NY); copses, Knoxville, June 1898, A. Ruth, 657 (US); 
open woods, Knoxville, July 1891, A. Ruth (US, 694398); dry 
woods, 5 mis. from Knoxville, Oct. 5, 1897, A. Ruth, 205 (US). 
Cocks Co.: dry rock banks, Wolf Creek, Sept. 24, 1897, Biltmore 
Herb., 2670a (G, US). BrouwT Co.: mountainsides, July 8, 
1898, A. Ruth 674 (NY). Mownok Co.: 10 mis. n. w. of Tapoco, 
Sept. 21, 1933, E. J. Alexander, T. H. Everett & S. D. Pearson 
(NY); Tapoco, Sept. 21, 1933, E. J. Alexander, T. H. Everett & 
S. D. Pearson (NY). Grunpy Co.: near Monteagle, Sept. 23, 
1933, E. J. Alexander, T. H. Everett & S. D. Pearson (NY). 
Pork Co.: near Archville, Sept. 25, 1933, E. J. Alexander, T. H. 
Everett & S. D. Pearson (NY). HawirroN Co.: Lookout Mt., 
1878, G. R. Vasey (US, 63440). ALABAMA. Mapison Co.: 
on Monte Sano, near Huntsville, Autumn, 1937, Mrs. N. G. 
Stevens (NY). CuLLMAN Co.: without stated locality, Oct. 5, 
1901, Biltmore Herb., 2670m (US). TALLADEGA Co.: slopes of 
Lookout Mt., near Childersburg, Oct. 2, 1899, Biltmore Herb., 
9537 (US). JEFFERSON Co.: dry hills, Avondale, Oct. 9, 1900, 
Biltmore Herb., 3402 (G, US). Tuscatoosa Co.: dry hills, 
Tuscaloosa, Oct. 4, 1898, W. M. Canby, 67 (G). GREENE Co.: 
shalk prairie, 2 mis. n. w. of Greene, Nov. 5, 1933, R. M. Harper, 
3137 (G, NY, US). Lez Co.: Auburn, Sept. 27, 1896, F. S. 
Earle (ND, type). BarpwiN Co.: Gateswood, Oct. 31, 1903, 
S. M. Tracy, 8558 (G (NY, type of Laciniaria Tracyi Alex.) US, 
ND). Mosie Co.: Mobile, Sept. 24, 1912, H. H. Bartlett, 3217 
(G, US); mixed woods, s. of Mobile, Sept. 24, 1912; H. H. Bart- 
lett, 3223 (US); dry woods, Mobile, Oct. 14, 1898, C. Mohr (US); 


338 Rhodora [NOVEMBER 


Mobile, Aug. C. Mohr (US); dry pine woods, Mobile, Oct. 20, 
1896, Herb. Geol. Surv. Ala. (US); pine woods, n. e. of Whistler, 
Oct. 1919, E. W. Graves, 1370 (US); barrens, w. of Spring Hill, 
Oct. 1918, E. W. Graves, 587 (US); sandy scrubby pineland, 
Hollander's Isl., Sept. 3, 1912, F. W. Pennell, 4505 (US). MIS- 
SISSIPPI. ArrALA Co.: Ethel, W. W. McBride (US). LAUDER- 
DALE Co.: Meridian, Oct. 10, 1896, C. Schuchert (US). JACKSON 
Co.: Ocean Springs, Oct. 17, 1898, S. M. Tracy 4871 (US). 
Harrison Co.: Biloxi, Aug. 23, 1898, S. M. Tracy, 4334 (NY); 
grassy barrens, near Mississippi City, Sept. 14, 1885, J. D. Smith, 
425 (US). LOUISIANA. CarcasrEU Co.: Jacksonville, T. 
Drummond, 66 (G). TEXAS. Without stated locality: C. 
Wrignt (G). Harris Co.: Houston, Sept. 24, 1937, G. L. 
Fisher (US). 

Since L. squarrulosa came to be recognized as a variety of L. 
scariosa with small heads, all such specimens were watched for 
and carefully compared. Though we have found a near match 
for Michaux’s type in H. W. Ravenel’s specimens from Santee 
Canal, S. C. (G), and those of H. H. Bartlett, no. 2420, from pine 
barrens near Thomson, McDuffie Co., Ga. (G, US), they are 
nevertheless comparatively rare in herbarium collections. 

Greene (Pittonia, iv. 316 (1901)) described a plant collected 
Sept. 27, 1896, by F. S. Earle, at Auburn, Lee Co., Ala. (ND), as 
Laciniaria Earlei, with a racemose inflorescence of small cam- 
panulate heads on a tall, somewhat virgate stem, having ap- 
pressed, non-scarious, purple, ciliolate-margined phyllaries, and 
narrowly lanceolate and linear leaves “glabrous . . . except for 
a few bristly marginal hairs at bases of some leaves”.  Laciniaria 
Tracyi Alex. (Man. S. E. Fl. 1335 (1933)) was also described from 
an Alabama plant (Oct. 31, 1903, S. M. Tracy, no. 8558, from 
Gateswood, Baldwin Co. (NY)) as having somewhat rounded, 
spatulate, pubescent phyllaries and linear, very scabrous lower 
leaves. Examination of the two type specimens showed no 
further marked differences in heads, flowers or leaf-shape and, as 
in other species of the genus there occurs the entire range from 
glabrous to scabrous leaves. it seemed these two were very simi- 
lar, especially, since in a specimen like E. W. Graves, no. 587, 
from the barrens w. of Spring Hill, Mobile Co., Ala. Oct., 1918 
(US), there is to be seen a very close parallel to Tracy’s plant 
with, in addition, two very basal leaves that are long and lanceo- 
late. The description of L. Ruthii Alex. for a plant collected 


1946] Gaiser,—The Genus Liatris 339 


Oct. 1896, A. Ruth, no. 34, from the Hiwassee Valley, E. Tennes- 
see (NY), with glabrous linear-lanceolate leaves and slightly 
larger heads, would seem to suggest a plant slightly more robust, 
but with heads having similarly somewhat spatulate, pubescent 
(though longer) phyllaries and glabrous rather than scabrous 
leaves. Examination of the types and comparison of a number 
of specimens, having smaller heads, from North Carolina to 
Florida and from Tennessee to Mississippi show that it is an easy 
step from Michaux’s South Carolina plant (the type of L. squar- 
rulosa) to the Alabama plants of Earle and Tracy and another 
easy step to the glabrous one of Ruth from Tennessee. All have 
15-25-flowered heads with the narrower herbaceous phyllaries 
and, though Michaux described those of his plant as squarrose 
and in the others they were stated to be erect, it is found that 
they are sometimes squarrose in heads of the narrow-leaved 
plants from Alabama. Thus, though it 1s difficult to draw sharp 
lines, the prevalence of linear or sharply pointed, narrowly 
lanceolate-linear, upper cauline leaves with lanceolate lower ones 
in the type plants of Earle, Tracy and Ruth, represents the 
trend in these smaller-headed relatives of L. scariosa from the 
region around the Gulf of Mexico in Alabama and Mississippi 
and northward. Specimens from the middle and northern coun- 
ties of Alabama, as Talladega, Jefferson and Cullman, foretell by 
their lanceolate basal leaves the material from Tennessee. 
Since none of these specimens have the broadly obovate basal 
and narrower but still oblanceolate upper leaves of Michaux's 
plants of L. squarrulosa we are including them under Liatris 
Earlei. 

As in all species of this series, plants of intermediate characters 
between two species of the same geographic range can be 
recognized. In Tennessee, from which the lanceolate-leaved 
specimen of Ruth came, other specimens show, in combination 
with broadly lanceolate basal leaves quite scarious-margined and 
slightly crisped elongate phyllaries, thus suggesting a blending 
with L. aspera var. intermedia. Biltmore Herb., no. 2670, from 
dry soil, Lookout Mt., Hamilton Co., Aug. 24, 1897 (US, 332418) 
is such a specimen while another sheet, also Biltmore Herb., 2670 
(US, 957890) of the same date and locality, is clearly L. aspera 
var. intermedia. Similar to the above intermediate are Biltmore 


340 Rhodora [NOVEMBER 


Herb. No. 2670b, from dry soil, near Hendersonville, Henderson 
Co., N. C., Sept. 14, 1898 (US) and Biltmore Herb. No. 26701, 
from Chickamauga Park, Ga., Sept. 21, 1899 (US). So also 
collections from Illinois of Sept. 13, 14, and 23, 1914, R. Ridgway, 
nos. 90, 88, and 87 respectively, from Calhoun prairie, from near 
Olney and from Sugar Creek prairie, seem to present intermedi- 
ates between the very asperous form of L. Earlei and L. scabra. 


Series VII. ELEGANTES. Distinctive species with plume- 
like inflorescence due to long heads (2.5-3 cm. long) in loose 
spicate to paniculate arrangement with prolonged, colorful, 
scarious, reflexed phyllaries; corolla as frequently white as purple. 
nonpilose within; achene 4-6 mm. long; pappus 9-11 mm. long 
and manifestly plumose. 

From South Carolina and Florida and westward into Texas 
and south-west Arkansas. 


23. LIATRIS ELEGANS (Walt.) Michx. Rootstock commonly 
relatively small, globose, 1-3 cm. in diameter, though in one 
variety elongate, tapering, up to 3 cm. in diameter and 15 cm. 
long; stems one or two, 3-12 dm. high, finely pubescent and leafy: 
leaves sessile, glabrous, punctate, linear to linear-lanceolate, 
reduced upwards from basal ones not more than 10 em. long 
and 5 mm. wide to bracts subtending the long heads; upper 
leaves commonly soon deflexed; inflorescence 3-5 dm. high, 
cylindrical to pyramidal according to amount of elongation 
basally of usually short pedicels of subsessile heads into longer 
sub-branched peduncles; heads usually 5-flowered, 2.5-3 cm. long; 
outer phyllaries short, lanceolate, herbaceous: inner ones pro- 
longed into dilated, lanceolate and reflexed or truncate and 
rounded, petaloid, serrulate tips, phlox-pink or white and sur- 
passing the flowers and pappus; corolla 9-11 mm. long, as fre- 
quently white as purple, or of intermediate mauve shades 
(yellow in the singular f. Fisher’), nonpilose within; stamens 
usually purple even within the white flowers, though sometimes 
white also; achene 4-6 mm. long; pappus long-plumose, 9-11 mm. 
long.—Fl. Bor.-Amer. ii. 91 (1803); Willd. Sp. Pl. iii. 1635 
(1803)!. Staehelina elegans Walt. Fl. Car. 202 (1788). Serra- 
tula speciosa Ait. Hort. Kew. iii. 138 (1789). Eupatorium speci- 
osum Vent. Hort. Cels. 79 (1802). Liatris radians Bertol. Misc. 
v. 9. t. 1 (1846). 


1 See Schubert in Ruopora xliv. 147—150 (1942) for comparative dates of publication. 


1946] Gaiser,—The Genus Liatris 341 


KEY TO VARIETIES 
a. Inner phyllaries dilated into lanceolate, reflexed serrulate 
petaloid tips... .b. 
b. Rootstock small, rounded; common over the range of the 


SDecles te pee esr COMM Ere a Meee Serie .var. typica. 
b. Rootstock elongate, tapering; found only in the Carizzo 
BANCO hexagon MN uc i ak NE gs a! var. carizzana, 


a. Inner phyllaries dilated into truncate, rounded, irregularly 
serrate, scarious tips; found only on St. Helena Isl., South 
Garoliria etre) ne oe E i, ee Oc tt var. flabellata. 


Var. typica. Rootstock small, globose, as found commonly 
over the range of the species: inner phyllaries with lanceclate- 
acuminate, dilated, petaloid apices showing the midrib extended 
at the tip, ca. 2 cm. long; corolla-tube 9-11 mm. long; achene 
4-5 mm. long; pappus 8-11 mm. long.—Liatris elegans (Walt.) 
Michx. Fl. Bor. Amer. ii. 91 (1803) sens. strict.; Willd. Sp. Pl. 
iii. 1635 (1803); Edwards Bot. Reg. t. 267 (1818); DC. Prodr. v. 
129 (1836); Torr. & Gray Fl. N. Am. ii. 68 (1841); Gray Synop. 
Fl. i. 109 (1884). Laciniaria elegans O. Ktze. Rev. Gen. i. 349 
(1891); Small, Man. 8. E. Fl. 1332, with fig. (1933). 

From South Carolina, south to Florida and west through 
Alabama, Louisiana, into Texas and again northward into south- 
west Arkansas.—SOU TH CAROLINA. Without stated locali- 
ty: Chapman (G). Co. undetermined: low country of South 
Carolina, L. R. Gibbes (NY). GEORGETOWN Co.: pine barrens, 5 
mis. s. of Georgetown, Sept. 9, 1939, R. K. Godfrey, 8135 (G, NY). 
BEAUFORT Co.: Beaufort District, 1882 (no. 377), 1888, Dr. J. H. 
Mellichamp (US); Bluffton, Dr. J. H. Mellichamp (NY); in 
woods, St. Helena Isl., Sept., A. Cuthbert (NY); dry woods, St. 
Helena Isl., Sept. 1883, A. Cuthbert (F); dry barrens, St. Helena 
Isl., Sept. 1884, Sept. 1894, A. Cuthbert (F); dry woods, St. 
Helena Isl., Sept. 1900, A. Cuthbert (NY, US); St. Helena Isl., 
Sept. 1900, A. Cuthbert (US); dry open woods, St. Helena Isl., 
Sept. 1894, A. Cuthbert (US); Sept. 21, 1902, Oct. 12, 1902, A. 
Cuthbert (F). GEORGIA. Without stated locality: 1839, Dr. 
Torrey (G, NY); Dr. Boykin (NY). Co. undetermined: along 
Flint River, Ez. Herb. Chapman (US). RicuMmonp Co.: barrens, 
Augusta, Sept. 1896, A. Cuthbert (NY). Briss Co.: sandy ridges, 
5 mis. e. of Macon, Sept. 4, 1883, J. D. Smith, 2004 (US). Tay- 
LoR Co.: dry sand hills, Anthony’s Mill, 4 mis. w. of Reynolds, 
Sept. 14, 1936, J. H. Pyron & R. McVaugh, 1300 (US). CHATTA- 
HOOCHEE Co.: dry soil, near Columbus, Sept. 17, 1902, ex Biltmore 
Herb., 2668h (NY); sandy banks, 4 mis. e. of Columbus, Sept. 7, 
1883, J. D. Smith, 1981 (US); Chattahoochee, Aug. 23, 1897, 
S. M. Tracy 3423 (NY). Sumter Co.: dry sandy soil, south of 
Americus, Aug. 29, 1900, R. M. Harper, 532 (G, NY, US). 
CHARLTON Co.: St. Mary’s River Swamp, below Trader’s Hill, 
July 24-26, 1895, J. K. Small (NY). FLORIDA. Without 


342 Rhodora [NOVEMBER 


stated locality: 1842-49, F. Rugel, 471 (US). Co. UNDETER- 
MINED: southern Florida, ex Chapman Herb. (NY); in dry, grav- 
elly oakwoods, ex Chapman Herb. (G). Duvar Co.: dry pine 
barrens, Jacksonville, Oct. 22, 1894, A. H. Curtiss, 5309 (G, NY, 
US); vicinity of Jacksonville, A. H. Curtiss, 1877 (ND); near 
Jacksonville, Oct. 21, 1893, A. H. Curtiss, 4445 (US); dry pine 
barrens, near Jacksonville, Oct. 1888, A. H. Curtiss, 1170 (G, 
NY, US, ND); St. Nicholas, July 28, 1896, L. H. Lighthipe, 
335 (NY). CoruwnBiA Co.: in high pine-turkey-oak woods n. 
of Camp Oleno, Oct. 6, 1940, W. A. Murrill (F). HAMILTON 
Co.: banks of Suwannee R., White Springs, Sept. 30, 1941, 
E. West & Miss L. Arnold (F). GapspEN Co.: Quincy, Sept. 
4, 1895, G. V. Nash, 2570 (G, NY, US, ND, F); Tallahassee, 
N. K. Berg (NY); open pinelands, western part of county, Aug. 
30, 1936, H. Foster, 116 (F). WakULLA Co.: without locality, 
Nov. 1891, W. G. Farlow (G). Jackson Co.: Sneads, Aug. 19, 
1942, R. A. Knight (F). WasuiNGTON Co.: s. of Chipley, Aug. 
23, 1942, M. Senner (F). OkaLoosA Co.: Crestview, Oct. 21, 
1936, Miss M. Hodges (F). Satnt JonNs Co.: dry pine barrens, 
St. Augustine, Oct. 1875, Miss M. C. Reynolds (NY, US, ND); 
pine barrens, Oct. Miss M. C. Reynolds, 306 (US). ArAcnHuA Co.: 
roadside, Hawthorne Rd., Oct. 31, 1931, Miss L. Arnold (F). 
'TAvLoR Co.: high pine-oak woods, 9 mis. s. of Perry, Oct. 8, 
1940, W. A. Murrill (F). Marion Co.: Citra, Nov. 10, 1941, R. 
A. Knight (F); high pineland, Belleview, Sept. 15, 1927, O. F. 
Burger & E. West (F). Cirrus Co.: sandy dry oak-pine woods, 
on U. S. Hwy. 41, 2 mis. s. of Holder, Oct. 16, 1945, H. H. Hume 
(F). Pasco Co.: dry woods, Lacoochee, Sept. 15, 1927, O. F. 
Burger & E. West (F). HinrsBonovGH Co.: Tampa Bay, G. 
Thurber (G). ALABAMA. Without stated locality: Mr. Lea 
(G); 1840, ex Herb. Meisner (NY); Gates (G, NY, B, isotypes of 
L. radians Bertol.); Oct. 1820, S. B. Buckley (G). Co. undeter- 
mined: pine woods, Sept., S. B. Buckley (NY, US). Lee Co.: 
open pine woods, 6 mis. s. of Auburn, Sept. 23, 1899, F. S. 
Earle & E. S. Earle, 94 (G, NY, US, ND); Auburn, Sept. 11, 
1898, F. S. Earle & C. F. Baker (NY, US), C. F. Baker (NY), 
Sept. 9, 1897, F. S. Earle & C. F. Baker, 1341 (N Y), Sept. 1900, 
F. E. Lloyd & F. S. Earle (NY). AvmAvGA Co.: sandy old 
field, between Autaugaville and Booth, Sept. 24, 1934, R. M. 
Harper, 3264 (G, NY, US). Barsour Co.: sandy roadside, 
between Cottonton & Eufaula, Aug. 12, 1927, K. M. Wiegand & 
W. E. Manning, 3172 (G); around Eufaula, Aug. 1888, G. Mc- 
Carthy (US). Escamsra Co.: in high pineland, Canoe, Oct. 25, 
1932, F. S. Blanton, 7071 (US); in low pineland, Atmore, Aug. 3, 
1933, O. Blanton, 152 (G); in high hammock, 3 mis. n. of Canoe, 
Oct. 17, 1929, H. O'Neill, 6171 (US, F). BarpwiN Co.: Gates- 
wood, Oct. 30, 1903, S. M. Tracy, 8568 (G, NY, US, T). Mo- 


1946] Gaiser,—The Genus Liatris 343 


BILE Co.: dry exposed banks, Mobile, Oct. 16, 1896, C. Mohr 
(US); dry pine woods, Mobile, Sept. 28, 1878 (US); Mobile, Oct., 
C. Mohr (US); Mobile, 1868, C. Mohr (US); dry close soil, 
copses and thickets, Mobile, Oct. 16, 1886, C. Mohr (US); dry 
pine barrens, Mobile, Sept. 1, 1862, C. Mohr (Q); pine barrens, w. 
of Spring Hill, Aug. 1918, E. W. Graves, 691 (US). ARKAN- 
SAS. Without stated locality: Drummond, 141 (G); Leaven- 
worth (NY). GARLAND Co.: dry woods, Glenwood Rd., 8 mis. 
from Hot Springs, Aug. 11, 1935, F. J. Scully, 434 (G); open 
field, Glenwood Rd., 8 mis. from Hot Springs, Aug. 8, 1935, F. J. 
Scully, 499 (G); Hot Springs, Aug. 1879, G. W. Letterman (US). 
Hor Sprines Co.: dry sandy ground, Malvern, Sept. 4, 1915, 
E. J. Palmer, 8453 (US). Prge Co.: dry, rocky, oak-hickory 
woods, between New Hope & Langley, Oct. 5, 1932, D. Demaree, 
9501 (G, NY). Crank Co.: Gurdon, C. Mohr (US). BRADLEY 
Co.: rich open wood, Jersey, Sept. 18, 1938, D. Demaree, 18325 
(O). HEMPsTEAD Co.: banks of Yellow Creek, near McNab, 
Oct. 5, 1923, J. M. Greenman, 4427 (G); sandy pine woods, 
Fulton, Oct. 15, 1914, E. J. Palmer, 6844 (US). MILLER Co.: 
near Texarkana, Aug. 20, 1898, A. A. Heller & E. G. Heller, 4124 
(G, NY, US). LOUISIANA. Without stated locality: ez. 
Torrey Herb., C. W. Short (NY). Co. undetermined: Red River, 
no collector, no date (G). NarcnirocHes Co.: sandy open 
woods, Natchitoches, Sept. 30, 1915, E. J. Palmer, 8753 (US). 
RarrpES Co.: Levins, Alexandria, Nov. 8, 1893, C. Mohr (NY); 
Alexandria, 1842, Hale ((G, without stated locality), NY, US); 
Glenmora, Sept. 30, 1913, F. W. Pennell, 5634 (NY); open dry 
hillside, n. edge of Pineville, July 30, 1938, D. S. Correll & H. B. 
Correll, 9931 (G, NY). TEXAS. Without stated locality: 
Sept. 1850, G. Thurber (G); Wright (G). 1843, Lindheimer, 71 
(G). Co. undetermined: Shawnee villages, Canadian R.,! Aug. 
1853, Lieut. A. W. Whipple Exped., J. M. Bigelow, 1853-4, 
(NY, US); between Indianola & San Antonio, Oct. 9, 1891, A. 
Schott (NY); without stated locality: Mexican boundary survey 
of Major Emory, C. C. Parry, J. M. Bigelow, C. Wright & A. 
Schott, 451 (NY). Bower Co.: sandy soil, near Texarkana, Oct. 
7, 1903, ex Biltmore Herb. 2668k (NY); Texarkana, G. Letterman, 
Oct. 15, 1894 (NY, US), Oct. 19, 1894 (US), Sept. 27, 1927, B. 
C. Tharp (G); oak-hickory w. of Texarkana, Sept. 27, 1927, B. C. 
Tharp, 2519b (G); Texarkana, 1896, E. N. Plank (NY), Sept. 21, 
1877, L. F. Ward (US); dry open woods, Texarkana, Oct. 27, 
1925, E. J. Palmer, 29412 (G). Rep River Co.: sandy loam, 
near Detroit, Sept. 15, 1937, E. Adams, 35 (US). Grayson Co.: 
Denison, Sept. 12, 13, 1906, F. L. Tyler (US). Cass Co.: Gallo- 

1 According to Bigelow, Report on Lieut. Whipple's Expedition, p. 96, Sec. 4. L. 


elegans was collected on this trip from Shawnee villages along the Canadian River, in 
August. 


344 Rhodora [NovEMBER 


way, near Atlanta, Oct. 9, 1895, E. Seler, 1007 (G); Bivins, Oct. 
15, 1940, O. McGinnis, 28011 (G). Dexron Co.: prairies, on 
Houston clay, between Benton & Aubrey, Sept. 22, 1937, W. L. 
McCart (US). Upsuur Co.: open ground, deep fine sands, Big 
Sandy, Sept. 27, 1926, E. J. Palmer, 31749 (G, NY). DALLAS 
Co.: Dallas, Aug. 1876, J. Reverchon (NY). Tarrant Co.: in 
deep woods, Riverside, Sept. 12, 1926, A. Ruth, 1435 (US). 
GnEGG Co.: without stated locality, Autumn, 1939, C. L. York 
(G). SwrrH Co.: sandy fields and open places, in pine-oak 
woods, s. e. of Camp Fannin, 8 mis. n. e. of Tyler, Oct. 12, 1943, 
H. E. Moore, 556 (G). San AvausriNE Co.: San Augustine, 
G. L. Crockett (US). HovsroN Co.: deep sands, open woods, 
Grapeland, Sept. 22, 1917, E. J. Palmer, 12842 (G). WALKER 
Co.: 834 mis. n. of Huntsville, Sept. 29, 1934, V. L. Cory, 10469 
(G); in an old graveyard, vicinity of Huntsville, July 9-12, 1909, 
R. A. Dixon, 332 (G, NY); Timberline, 1414 mis. s. w. of Hunts- 
ville, Sept. 28, 1934, H. B. Parks & V. L. Cory, 10254 (T). Mr- 
LAM Co.: sandy woods, 4 mis. east of Cameron, Aug. 4, 1929, S. E. 
Wolff, 11026 (US). Harprn Co.: 5 mi. south of Silsbee, Oct. 15, 
1936, V. L. Cory, 20035 (G). Jnrrerson Co.: Nome, Sept. 2, 
1937, G. L. Fisher, 37159 (US). Harris Co.: 10144 mis. west of 
Laporte, Oct. 8, 1934, Cory, 11415 (G), Houston, Aug. 19, 1923, 
G. L. Fisher, 56 (US). Austin Co.: Industry, 1894, H. Wurzlow, 
30 (US). GowzaLEs Co.: Palmetto State Park, Aug. 1, 1941, 
B. C. Tharp, 145 (G); without locality, Oct. 10, 1940, Tharp (G). 
Victoria Co.: Aloe, Sept. 18, 1913, F. W. Pennell (NY). ATas- 
cosa Co.: sandy open ground, Pleasanton, Sept. 23, 1916, E. J. 
Palmer 10794 (US). Nuerces Co.: Flour Bluff, 12 mis. south of 
Corpus Christi, Sept. 27, 1936, S. G. & J. A. Drushel, 10396, 
(NY). Brooxs Co.: Santa Fe Ranch, June 26, 1941, Tharp 
(G). KLEBERG or KENNEDY Co.: shell bank on Padre Island, 
Sept. 3, 1927, Tharp (G). OKLAHOMA. McCurtain Co.: 
deep, rich coastal plain woods, near entrance to cypress swamp 5 
(a southeast of Broken Bow, Oct. 2, 1940, M. Hopkins, 5566 

). 

Var. TypicA forma FisuERi Standl. Differing from var. 
typica only in having the corolla, as well as the tips of the phyl- 
laries, lemon-yellow.—Field Mus. Pub. Bot. xi. 276 (1936).— 
Known only from the type specimen, collected Aug. 26, 1934 by 
G. L. Fisher on hillsides near and on the highway, Copperas 
Cove, Coryell, Texas (Chicago). 

Var. carizzana, var. nov., planta robusta ad 9 dm. alta, a var. 
typica rhizomate elongato conico 2.5 cm. diametro ad 15 cm. 
longo, phyllariis paullo longioribus (2.5 cm.), achaeniis 6 mm. 
longis, pappo 11-12 mm. longo diversa.—Medina, Wilson, 
Atascosa and Hudspeth counties, Texas.— TEXAS. MEDINA 
Co.: 3 mi. southwest of Devine, Oct. 12, 1934, V. L. Cory, 11726 


1946] Gaiser,— The Genus Liatris 345 


(G, type); 234 mi. southwest of Derby, May 8, 1935, H. B. Parks 
& Cory, 12778 (T). WirsoN Co.: Kicaster School, Oct. 22, 
1934, Parks & Cory, 11877 (T), June 24, 1935, Parks & Cory, 
15136 (T); post-oak woods, July 21, 1925, Tharp (G). Atascosa 
Co.:9 mi. east of Poteet, Nov. 12, 1934, Cory, 11721 (G). Houps- 
PETH Co.: Sierra Blanca, Sept. 7, 1925, Tharp, 3772 (US). 

Var. flabellata (Small), n. comb. Differing from var. typica 
only in having inner phyllaries with rounded, dilated, toothed, 
petaloid tips.—Laciniaria flabellata Small, Bull. Torr. Bot. Club, 
xxv. 472 (1898). Liatris flabellata (Small) K. Sch. Just, Bot. 
Jahresb. xxvi. pt. 1, 378 (1900). SOUTH CAROLINA. Beav- 
FORT Co.: dry, open woods, St. Helena Isl., Sept. 1894, A. Cuth- 
bert (NY, type, US). 

. As stated in the introduction, plants growing [in the out- 

cropping of the Eocene] in the Carrizzo sands of Medina, Bexar 
and Wilson counties of Texas are all said to have deeply pene- 
trating root systems. As all specimens of Liatris elegans I have 
seen from these counties, as well as one from Poteet, Atascosa 
Co., where there is the same formation, have such very long 
underground systems, really remarkable in a species which has 
usually a relatively small globose corm, it has seemed advisable 
to make of these a variety, here called var. carizzana. The only 
other specimen of L. elegans, not coming from this region, with 
an elongate rootstock is one of Sept. 7, 1925, B. C. Tharp, no. 
3772, from Sierra Blanca, Hudspeth Co., Texas (US), and con- 
cerning the soil in which it grew, Prof. B. C. Tharp! has written 
“deep sand and in that respect it resembles the Carizzo sand very 
much". 

The type collection of Laciniaria flabellata Small provides the 
only examples of specimens with dilated, truncate, petaloid, 
inner phyllaries. Several other collections in successive years 
by the same collector from that island are quite like Liatris 
elegans var. typica, although the acuminate phyllaries are always 
quite conspicuous and somewhat serrate. On the sheet of a col- 
lection made on Sept. 21, 1902 (F), is the collector's observation 
“note gradation from typical acuminate bracts of elegans to- 
wards flabellata.” Though the modification has seemingly been 
a very rare one, it is here given varietal rank under L. elegans. 

A hybrid of this species and L. tenuifolia Nutt. was discussed 
under that species (see no. 15). 


1 By private communication. 


346 Rhodora [NOVEMBER 


Series VIII. Puncrarar. Plants mostly glabrous, with 
many-stemmed, generally erect, though sometimes recumbent 
spikes from an elongate, branching rootstock in two species and 
a globular one in other species; leaves numerous, mostly con- 
spicuously punctate, linear and narrow; inflorescences generally 
densely spicate (rarely loose); heads slenderly cylindrical, 8-20 
mm. long, 4-8-flowered (10-14 flowers in one species) with 
herbaceous appressed phyllaries; corolla-tube quite pilose within; 
achene 5-12 mm. long; pappus distinctly plumose, 8-15 mm. 
long.—From northern Mexico and Texas to the prairies of south- 
ern Canada and west to the east side of the Rocky Mts. 


a. Plants with a long underground rootstock. . . . b. 
b. Stems moderately stiff; leaves 1.5-5 mm. wide, conspicu- 
ously punctate, frequently ciliate; rootstock deeply 
penetrating; plants of wide range.................. 24. L. punctata. 
b. Stems more slender and flexible; leaves 1-2 mm. wide, 
punctate, not ciliate; rootstock horizontal; in sandy 
prairies in Minnesota... .......0...00.000 00000. 25. L. densispicata. 
a. Plants with a somewhat globular corm... .c. 
c. Heads 4-6-flowered. . . .d. 
d. Leaves 1.5-5 mm. wide, punctate, stiff; phyllaries 
mucronate-tipped; heads numerous or rarely few and 


distant... es 26. L. mucronata. 
d. Leaves 1-3 mm. wide, almost epunctate, soft; phyllaries 
acuminate; heads numerous.................. 27. L. angustifolia, 


c. Heads 10-14-flowered, few, very distant on few-branched 
stems; leaves narrow, rather channelled and bract-like.28. L. bracteata. 


24. Liarris PUNCTATA Hook. Stems numerous, glabrous, 
striate, 1.5-8 dm. long, from a crown above an elongate, mostly 
branched rootstock, often penetrating long distances: leaves 
numerous, glabrous, rigid, linear, conspicuously punctate (but 
not to be distinguished from other species by this punctate 
character which is common to most); basal leaves 8-15 cm. long, 
1.5-6 mm. wide, with cutinous margin bearing prominent cilia 
or only scantily so provided, gradually diminishing, below the 
beginning of a short spike or through the spike in the case of a 
longer one, to bracts shorter than the heads they subtend: 
inflorescence generally densely, sometimes loosely spicate, 6-30 
em. in length and 2-3 cm. in width when flowers are open, of 
usually crowded heads 1.5-2 cm. long, 8-10 mm. wide, containing 
4-8 flowers; phyllaries herbaceous, thick, punctate and closely 
appressed except for the free tips, the outer short, rigid, ovate- 
acuminate or cuspidate, the inner oblong with acute or mucronate 
to lanceolate-acuminate tips, 10-14 mm. long and 1.5-2 mm. 
wide, prominently ciliate-margined in individuals having cilia 
on the margin of the leaves and correspondingly less ciliate to 
merely membranous-margined in others; corolla purple (rarely 
white) 9-12 mm. long, with the tube inside and filaments of 
stamens quite pilose; pappus distinctly plumose, 9-11 mm. long, 


1946] Gaiser,— The Genus Liatris 347 


usually slightly exceeded in length by the corolla; achene 6-7 
mm. long, ribbed and hairy.— Fl. Bor.-Amer. 1. 306, t. 305 (1834). 
Laciniaria punctata (Hook.) O. Ktze. Rev. Gen. i. 349 (1891). 
L. cylindrica Torr., Ann. N. Y. Lyceum, ii. 210 (1824), as to 
plant described. Laciniaria punctata var. turgida Lunell, Amer. 
Mid. Nat. v. 241 (1918). Laciniaria punctata f. corymbosa 
Sheldon, Quart. Bull. Univ. Minn. i. 27 (1892) and f. albiflora 
Sheldon ibid. 26. 
KEY TO VARIETIES 


a. Inflorescence dense... .b. 
b. Stems stiff and short, 1.5-3 dm. tall; leaves 3-5 mm. wide, 
ciliate-margined; phyllaries prominently ciliate; of more 
northern plains and western mountains................. var. typica. 
b. Stems more slender, 4-8 dm. tall; leaves narrower, 2-3 mm. 
wide, and longer than in var. typica; leaves and phyllaries 
without prominent ciliate margins; Minnesota and Dako- 
ta, southward to the plains of Texas................ var. nebraskana. 
a. Inflorescence lax; stems tall; leaves few and larger than in var. 
typica, 12.5-15 cm. long and 4-6 mm. wide, almost glabrous; 
heads fewer and scattered; mountains of Trans-Pecos Texas 
and northern Coahuila, Mexico....................... var. mexicana. 


Var. typica. Stems 1.5-3 dm. long and stiff: basal leaves 
8-10 em. long and 3-5 mm. wide with prominent cilia along the 
cutinized margin: inflorescence a short spike 6-20 cm. in length, 
of crowded heads ca. 2 em. long, containing 5-8 flowers: phyl- 
laries prominently margined by long white cilia, inner ones 
lanceolate-acuminate or oblong with acute or mucronate tips.— 
L. punctata Hook. Fl. Bor.-Amer. 1. 306 t. 305 (1834), sens. strict. 
Liatris punctata var. y, Torr. & Gray Fl. N. Am. ii. 69 (1841). 

From Manitoba and Saskatchewan south to Iowa and Kansas, 
and from Alberta southward along the eastern Rockies into New 
Mexico and Trans-Pecos Texas.—MANITOBA. 62 H 14: dry 
gravelly knolls, Fort Garry (Winnipeg), Aug. 5, 1872, J. Macoun, - 
9851 (stem to right) (Ot). 62 G 3: on the open prairie, Chater, 
July 25, 1896, J. Macoun, 12227 (Ot). 62 G 4: on the hill s. of 
the lake, Killarney, Aug. 3, 1896, J. Macoun, 12196 (Ot). 62 G 
16: Portage la-Prairie, Aug., 1896, J. K. McMorine (Q); 62 K 1: 
Rivers, Aug. 7, 1933, H. H. Brown (HB). MINNESOTA. 
WASHINGTON Co.: St. Paul & St. Anthony, 1861, 7. J. Hale (G). 
OTTER Tart Co.: dry prairies, Perham, Aug. 14, 1910, Z. L. 
Chandonnet (US). HENNEPIN Co.: dry rocky hillside, Fort 
Snelling, Sept. 1, 1888, E. A. Mearns, 140 (N Y, US); Fort Snelling 
to rivers of north, Dr. Jarvis (G). Noses Co.: dry hills, Adrian, 
Aug. 22, 1895, F. W. Hunnewell (GE IOWA. PLYMOUTH Co.: 
Akron, Sept. 1909, Mrs. E. Bredall (US). Woopsury Co.: Sioux 
City, Aug. 29, 1896, L. H. Pammel, 39 (NY, US). NORTH 

1 Localities in Manitoba, Saskatchewan and Alberta are indicated by standard nota- 


tion with reference to sheets of the National Topographical Series, Dept. Mines & 
Resources, Ottawa, Canada. 


348 Rhodora [NOVEMBER 


DAKOTA. BorriNEAU Co.: Bottineau, Aug. 24, 1890, C. B. Wal- 
dron (NY). Warp Co.: on roadside, Minot, Aug. 18, 1931, Miss 
R. Weikert, 1 (NY). Benson Co.: prairies, Leeds, Aug. 28, 1899, 
J. Lunell (G); in dry prairies, Leeds, Sept. 8, 1909, J. Lunell (US); 
Butte, J. Lunell, Oct. 21, 1908 (US), Oct. 2, 1908 (NY). Bur- 
LEIGH Co.: along Missouri R., near Bismarck, Aug. 26, 1917, F. P. 
Metcalf, 369 (G). Starx Co.: Dickinson, Sept. 8, 1908, W. R. Hol- 
gate (G). SOUTH DAKOTA. Co. undetermined: western part of 
State, Aug. 1891, T. A. Williams (US). RosBEnrS Co.: clay banks, 
Big Stone Lake, Aug. 1922, W. H. Over, 14351 (US). Grant Co.: 

virgin prairie, "Troy, near Big Stone Lake, Aug. 4, 1940, P. 
Johnson, 68 (G, NY). Day Co.!: sandy, arid prairie heights, 
around the Sioux, lower St. Peter R., near Fort Pierre, Missouri 
Hills, dividing ridges between Missouri & Mississippi waters, 
Aug. 20, 1839, Nicollet’s N. W. Exped., 272 (*var. y”) (US); 
(without stated locality), Nicollet’s N. W. Exped., C. A. Geyer 
(G “var. Y" (NY “var.’’)); probably Day Co.: Oak Gulch, 
Sept. 5, 1896, L. W. Carter (NY). Burre Co.: Belle Fourche, 
July 30, 1926, H. E. Hayward, 31 (NY). Brooxines Co.: 
White, Aug. 29, 1906, Miss F. N. Vasey (US). KiwasBunRY Co.: 
Iroquois, Aug. 9, 1894, J. J. Thornber (G). WasHABAUGH Co.: 
Pine Ridge Reservation, Aug. 1911, S. S. Visher, 2340 (NY). 
SHANNON Co.: Wolf Creek, near Pine Ridge Agency, Aug. 9, 
1911, S. S. Vasher, 2207 (NY). Farı River Co.: Hot Springs, 
Sept. 1893, Lieut. W. E. Safford (US); Hot Springs (alt. 3500’), 
Aug. 3, 1892, P. A. Rydberg, 754 (US). NEBRASKA. "TnuoMas 
Co.: Thedford (excepting tall stem to right), Aug. 7, 1889, 
H. I. Webber (NY); on sandhills, on Middle Loup R., near 
Thedford Aug. 19, 1893, P. A. Rydberg, 1706 (US). Apams Co.: 
Hastings, Sept. 16, 1886, Prof. Thompson (G). KANSAS. 
CHEYENNE Co.: near Catholic Church, St. Francis, Sept. 7, 1936, 
Mrs. J. M. Steller, 91 (NY). Doveras Co.: Lawrence, W. C. 
Stevens (US). WarLAcE Co.: Wallace, Aug. 22, 1884, G. W. 
Letterman (NY). HawirTOÓN Co.: vicinity of Syracuse, Sept. 14, 
1912, J. N. Rose & W. R. Fitch, 17013 (NY, US). STEVENS 
Co.: "Moonlight, Aug. 15, 1893, C. H. Thompson, 174 (NY, US). 

Morton Co.: without stated locality, July 28, 1891, M. A. 
Carleton, 353 (US). SASKATCHEWAN. Without stated lo- 
cality: 1858, Palliser’s Br. N. Amer. Expl. Exped., E. Bourgeau 
(G, NY); ex Herb. Musei Brit. Douglas, 9849 (Ot). Exact 


! From Senate Report intended to illustrate a map of the H ydrographical Basin of the 
Upper Mississippi River made by I. N. Nicollet, Feb. 16, 1841, Washington, 1843, p. 53: 
“I was at St. Peter's when I made up my mind to visit the sources of the Mississippi''— 
"Left St. Peter's on July 26, 1836''—‘‘on July 29, I was ascending the Mississippi’’. 
No dates are given in the further description of travel but from the map they were, 
on Aug. 19, west of Big Stone Lake near what is called Cold Spring Lake—mouth of 
the Sioux River. 'This would seem to be about Day County, S. Dakota (see also 
following discussion). 


1946] Gaiser,—The Genus Liatris 349 


region undetermined, Saskatchewan plains, Aug. 3, 1872, J. 
Macoun (US without no.), 821 (NY). 72 P 14-13: Little Lake 
Manito, July 21, 1906, J. Macoun & W. Herriot, 77087 (G, Ot). 
73 B 2: along the G. T. Rwy., 15 mis. w. of Saskatoon, July 29, 
1906, J. Macoun & W. Herriot, 77086 (NY, Ot). 62 L 12: gravelly 
knolls in prairies, Indian Head, Sept. 16, 1891, J. Macoun, 9848 
(Ot); 72 I 6: prairies, Moosejaw, July 31, 1904, F. W. Johnson, 
972 (US). 72J 16: prairies, Brownlee, Aug. 7, 1913, F. W. John- 
son, 1259 (NY). 62 E 16: Moose Mountain, Aug. 1884, J. 
Macoun (US). 72 K 2: prairies, Sidewood, Assiniboine, Aug. 
26, 1892, F. E. Floyd (NY). 62 F 12: Redvers, Aug. 19, 1909, 
G. E. Copeland (Q). 62 M I: Yorkton, Aug. 19, 1908, W. Crawford 
(Q. ALBERTA. Without stated locality: prairies, Aug. 21, 
1913, F. W. Johnson, 1065 (US). 83 A 4: Red Deer, Aug. 2, 
1917, C. H. Young, 92730 (Ot). 74 M 11: rwy. track, Scotfield, 
Aug. 10, 1926, A. H. Brinkman 2541 (US); 82 P: dry soil, prairie 
hills (alt. 2200’-2500’), Rosedale Coulee, Aug. 6, 1915, Miss M. 
E. Moodie, 1170 (G, NY, US). 82 O 1: dry hills, Calgary, 
Aug. 19, 1913, Miss M. E. Moodie (US); dry ground & prairies, 
vic. of Calgary, Elbow R. Valley (alt. 3400'-3600') Aug. 6, 1913, 
Miss M. E. Moodie, 28 (NY); grassy hills ca. 13 mis. s. of Cal- 
gary, Aug. 1, 1941, C. L. Hitchcock & J. S. Martin, 7853 (NY). 
ca. 82 H 11: middle fork of Old Man R., Rocky Mts., Aug. 14, 
1883, Dawson, 9850 (Ot). 82 H 11: near irrigation ditch, s. of 
Standoff (s. of MacLeod), Aug. 15, 1929, E. H. Moss, 410 (G). 
82 H 12: in the valley of Old Man R., Fort MeLeod, Aug. 24, 
1897, J. Macoun, 22766 (Ot); prairies, Fort McLeod, Aug. 5, 
1895, J. Macoun, 10829 (Ot); MacLeod, Aug. 1923, R. H. Dixon, 
432 (NY); dry slope of gravelly ridge, w. of Pincher Creek, July 
26, 1939, E. H. Moss, 62 (G); Oldham R., n. of Pincher, Aug. 2, 
1939, E. H. Moss, 250 (G). MONTANA. Without stated 
locality: 1883, L. F. Ward (US). Co. undetermined: prairie, 
upper Missouri, Aug. 19, 1864, T. M. Rothammer, 477 (US); 
Mountain Sheep Buttes, Aug. 11, 1909, V. L. Bailey (US); 
without stated locality, Aug. 10, 1890, J. W. Blankinship, 131 
(US); upper Missouri, Aug. 27, 1864, T. M. Rothammer, 490 
(US). SurnipAN Co.: Westby, Aug. 12, 1928, Miss E. Larsen, 
211 (US). Dawson Co.: Colgate, near Glendive, Sept. 6, 1892, 
J. H. Sandberg, 1020 (US); Seven. Mile Creek, 15 mis. above 
Glendive, Aug. 11, 1883, L. F. Ward (US). CascapE Co.: 
Belt Creek, July 1883, F. W. Anderson (US); (albino) Great 
Falls, Aug. 24, 1891, R. S. Williams, 76 (US). Lewis & CLARK 
Co,: in vacant lots, Helena, Sept. 2, 1908, B. T. Butler, 649 (NY); 
White's Gulch, near Helena, Aug. 22, 1882, W. M. Canby (G); 
6 mis. s. e. of Helena, Aug. 12, 1931, J. T. Howell, 7897 (NY). 
TREASURE Co.: dry ground near Custer, Aug. 9, 1912, H. D. 
House, 4900 (US). MussELSHELL Co.: plains of Musselshell R., 


350 Rhodora [NOVEMBER 


Sept. 2, 1896, J. H. Flodman, 817 (NY, US). Jrrrerson Co.: 
rocky slopes, above Jefferson R., 10 mis. e. of Whitehall, Aug. 5, 
1941, J. F. Brenckle & L. H. Shinners, 41-061 (G). Pank Co.: 
Livingston, Yellowstone National Park, Mrs. E. W. Scheuber, 
Aug. 2, 1901 (US), 1901, 20 (NY). GarrArIN Co.: Bozeman, 
Aug. 17, 1900, E. J. Moore (G), Sept. 2, 1902, W. W. Jones (US); 
dry uplands, Bozeman, Aug. 24, 1905, J. W. Blankinship, 284 
(US); Cinnabar, Yellowstone River, July, 1884, F. Tweedy (US). 
WYOMING. Without stated locality: July 25-30, 1870, 
Hayden’s U. S. Geol. Survey (US). Co. undetermined: sources 
of the Platte, Dr. James! (NY); Aurora, July 30, 1900, W. 
Granger (NY). SHERIDAN Co.: Dayton (alt. 4000’), Sept. 1899, 
F. Tweedy 2072 (NY). Park Co.: Clark's Fork R., Aug. 1881, 
W. H. Forwood, 21 (G). Weston Co.: Cambria Canon, July 22, 
1896, A. Nelson, 2529 (NY). JomuwsoN Co.: dry plains near 
Rock Creek (alt. 6500’), Aug. 1, 1934, R. C. Rollins, 712 (G, NY); 
Buffalo (alt. 4000'-5000^), Sept. 1900, F. Tweedy, 3146 (NY). 
NrioBRARA Co.: dry plains, near Kirtly (alt. 5000’), Aug. 3, 
1931, R. C. Rollins, 36 (NY). PrATTE Co.: Cottonwood Canon, 
Aug. 4, 1895, A. Nelson, 1564 (G, NY, US); North Fork of 
the Platte Hiver, 1843-44, Fremont's Exped. to California 
(G, NY, US). ArnaaxY Co.: Dale Creek, Aug. 24, 1908, Mrs. J. 
Clemens (G); 15 mis. s. w. of Laramie, Aug. 22, 1901, E. D. 
Merrill & E. N. Wilcox, 1169 (G, US); dry bench lands, Centen- 
nial, Aug. 5, 1900, A. Nelson, 7952 (G, NY, US); Bear Creek, ca. 
2 mis. from Eagle Rock & 4 mis. from Laramie Peak, Aug. 22, 
1891, C. Schuchert (US). COLORADO. Without stated lo- 
cality: 1870, E. L. Greene (G), 1878, P. J. Mohr (US 720072), 
L. F. Ward (US 134488). Co. undetermined: meadow at foot- 
hills of Colorado (alt. 6500’), Aug. 9, 1890, C. S. Crandall, 272 
(US); Stanley's Ranch, Mrs. C. N. S. Horner (G). LARIMER 
Co.: Estes Park, Aug. 11, 1906, E. L. Johnston, 218 (US); 
Livermore, Aug. 22, 1900, G. E. Osterhout, 2231 (NY). Bour- 
DER Co.: mts., Boulder, Sept. 3, 1895, C. L. Shear, 4456 (US); 
near Boulder, Oct. 1, 1900, F. Ramaley, A201 (US); Boulder, 
Aug. 1891, Dr. E. Penard, 265 (NY); mts. between Sunshine 
and Ward (alt. 8000'-9500^), Aug. 1902, F. Tweedy, 4936 (NY). 
ARAPAHOE Co.: North Denver (plant to left), Aug. 16, 1910, 


1! This specimen, though without locality of collection on the Herbarium sheet, 
would seem to be that cited as Liatris cylindrica Torr. Ann. N. Y. Lyceum ii. 210 (1827) 
No. 204, for which is given, ‘‘sources of the Platte". (See also following discussion.) 

? From report of Fremont's Expedition, Washington, 1845, the date is Aug. 29, 
and place of collection, Black Hills of the Platte. Following the account of the 
itinerary “Aug. 27, on their return they halted where they had taken dinner on July 
27". By the map this was where Deer Creek enters the Platte, and is where Glenrock, 
Converse Co., Wyo. now is. Since they reached Fort Laramie on the last day of 
August the collection of L. punctata would be between Glenrock and Fort Laramie, 
possibly Platte Co.. 


1946] Gaiser,—The Genus Liatris 351 


Miss A. Eastwood, 44 (US); near Denver, Dec. 1, 1874, C. Mohr, ` 
(US); dry plains, n. w. of Denver (alt. 1500 m.), Sept. 17, 1901, L. 
M. Underwood & A. D. Selby, 529 (NY). JEFFERSON Co.: 
Golden, Oct. 8, 1882, N. L. Britton (NY); Golden (alt. 5500^), 
Aug. 21, 1884, C. S. Sheldon, 297 (NY); Morrison (alt. 6500’), 
Aug. 7, 1878, M. E. Jones, 546 (NY, US). Dovucras Co.: 
Buffalo Creek Canon, Sept. 14, 1909, H. H. Rusby (NY). EL 
Paso Co.: near Colorado Springs (alt. 6000^, Sept. 22, 1895, S. L. 
Clarke, 7 (NY); Colorado Springs (alt. 6000’), Sept. 19-21, 
1895, Mrs. S. L. Clarke (NY). Gunnison Co.: Gunnison, 1896, 
F. Clements, 229 (NY). CusrER Co.: West Cliff, Aug. 13, 1896, 
C. L. Shear, 3458 (NY). Orero Co.: vicinity of La Junta, Sept. 
16, 1912, J. N. Rose & W. R. Fitch, 17064 (NY, US). Hurrrano 
Co.: Cucharas Valley, near La Veta (alt. 7000’), Sept. 26, 1900, 
F. K. Vreeland, 683 (NY). Las Animas Co.: Raton Mts., 
Sept. 1, 1898, E. O. Wooton (US); Wootton, Sept. 11, 1909, H. H. 
Rusby (NY). NEW MEXICO. |. UuroN Co.: Folsom, Aug. 30, 
1903, A. H. Howell, 165 (US). Corrax Co.: Colfax, Aug. 13, 
1910, E. O. Wooton (US); open, pine-oak woods top of Raton 
Pass, alt. 7800 ft., Aug. 18, 1941, U. T. Waterfall, 3502 (G); 
vicinity of Raton, Oct. 27, 1913, J. N. Rose & W. R. Fitch, 17546 
(NY, US); Bartlett Ranch, Aug. 31, 1913, Wooton (NY); Vermejo 
Park (alt. 7600 Ft.), Aug. 31, 1913, Wooton (US), July-Aug., 
1894, Mrs. O. St. John, 100 (G); vicinity of Ute Park, alt. 2200- 
2900 m., Sept. 21, 1916, P. C. Standley, 14240 (G). San MIGUEL 
Co.: 15 mi. northwest of Las Vegas, Aug. 30, 1934, G. J. Good- 
man, 2322 (G, NY, O); near Bernal, alt. 6300 ft., Aug. 27, 1903, 
V. L. Bailey, 546 (US). BERNALILLO Co.: among rocks and on 
hillsides, Canyon Media, Sandia Mts., Miss C. C. Ellis, 284 (US); 
dry flats, east slope Sandia Mts., alt. 7500 ft., Ellis, 284 (US). 
LixcoLN Co.: Block Ranch, north of Capitans, Aug. 31, 1900, 
F. S. & E. S. Earle, 384 (NY, US); El Capitan Mts., alt. 8000 
ft., July 7, 1928, Earle (NY). 

Var. TYPICA forma coloradensis, f. nov., phyllariis mucronatis 
purpurascentibus, foliis plerumque angustioribus.—Chiefly Colo- 
rado and New Mexico.—COLORADO. Without stated locality: 
1862, E. Hall (US), C. Mohr (US, 783573). Co. undetermined: 
sterile plains (alt. 5700’), Aug. 20, 1884, C. S. Sheldon, 297 (US). 
Warp Co.: New Windsor, Aug. 20, 1901, G. E. Osterhout, 2339 
(NY, US); Greeley, Aug. 11, 1881, L. F. Ward (US); Dent, Aug. 
31, 1917, W. W. Jones, 535 (G). Larimer Co.: Fort Collins 
(alt. 5000’), July 29, 1891, J. M. Cowan (NY); Fort Collins, Aug. 
25, 1896, C. F. Baker (NY); College Farm, Col. Agric. College, 
July 29, 1892, C. S. Crandall (US); along rwy., s. of Agric. 
College, Aug. 23, 1898, State Agric. College, 2975 (NY, US); 
Estes Park, Aug. 1894, Mrs. J. M. Milligan (US). BourpER 
Co.: without stated locality, Aug. 1882, H. N. Patterson & F. 


352 Rhodora [NOVEMBER 


‘Beatty, 58 (US). ARapaHoE Co.: Melvin, Aug. 13, 1890, E. L. 
Hughes, 13 (G); plains, Denver, Sept. 12, 1888, W. G. Smith (US); 
North Denver, Aug. 16, 1910, Miss A. Eastwood, 44 (G, US 
plant to right); dry soil, Denver, Aug. 29, 1916, I. W. Clokey, 
2682 (G, NY). JxrrEnsoN Co.: Golden, road up to Lookout 
Mt., Aug. 28, 1916, E. L. Johnston, 218B (G). Lincoun Co.: 
Hugo, July 28, C. D. Marsh (US). Er Paso Co.: moist meadow 
among hills, 4 mis. s. of Palmer Lake, Aug. 7, 1941, U. T. Water- 
fall, 3203 (G); Colorado Springs, Aug. 20, 1889, B. W. Evermann 
(US, 310558 & 617864), Oct. 15, 1903, W. C. Sturgis (G, US), 
Oet. 1903, Mrs. J. M. Milligan (US); Colorado Springs (alt. 
1900 m.), Sept. 22, 1915, W. W. Eggleston, 12031 (US); dry 
plains, Colorado Springs, Aug. 6, 1892, C. S. Sheldon, 473 (NY, 
US); Pike's Peak, Cheyenne Mt., north slope, July 28, 1896, E. A. 
Bessey (NY); Pike's Peak, Aug. 23, 1887, S. M. Tracy & Evans, 
801 (stem to right) (NY); Garden of the Gods, Aug. 8, 1908, C. 
E. Bessey (US); dry hillsides, Colorado Springs, Aug. 3, 1939, 
J. H. Ehlers, 7785 (G); dry plains, near foothills, Aug. 11, 1897, 
C. S. Crandall, 2974 (G); “The Mesa", 3.5 mis. n. w. of Colorado 
Springs, Aug. 7, 1924, R. Bacigalupi, 678 (G); Manitou (alt. 
2100 m.), Aug. 16, 1901, F. E. Clements & E. S. Clements 72 (G, 
NY, US). Cuarrer Co.: Salida, Aug. 28, 1892, Miss I. Mulford 
(G, NY). PuzrzLo Co.: on plains about Pueblo, 1883, R. W. 
Woodward (G). HvrmraNo Co.: St. Mary's, Mrs. C. N. S. 
Horner (G). OKLAHOMA. Cimarron Co. (near New Mexi- 
co line): s. of Black Mesa 114 mis. w. of Kenton, Aug. 6, 1941, 
U. T. Waterfall, 3176 (G). NEW MEXICO: Co. undetermined: 
from Cottonwood Creek! of Ark. to San Miguel, Aug. 1847, A. 
Fendler, 299 (G). San Dovar Co.: hard clay soil, Nara Visa, 
Sept. 8, 1910, G. L. Fisher, 68 (US); dry open grassy ridges, 
Jemez Springs area, Aug. 24, 1931, A. Nelson, 11704 (G). SAN 
MIGvuEL Co.: Las Vegas, June, 1920, Bro. Anect, 1 (NY); Pecos, 
S. S. Holman (US); near Pecos (alt. 6700’), Aug. 15, 1908, P. C. 
Standley, 4958 (US). Quay Co.: Llano Estacado, Sept. 16, 
1853, Whipple Exped. J. M. Bigelow (NY (US 2 stems to left)); 
Guadalupe Mts. (alt. 6000'-7000^, Sept. 3-6, 1915, A. S. Hitch- 
cock (US); hard clay, Nara Visa, Sept. 8, 1910, G. L. Fisher, 68 
(US). Dona Ana Co.*: valley of Rio Grande, Mexican Boundary 
Survey, 450 (NY, US). 


1 Plant. Fendl. gives this region for the collection of the specimen no. 299, L. 
punctata. 

2 According to Bigelow,—Report on Lieut. Whipple's Expedition (1857), p. 96, 
Sec. 4, L. punctata was collected in rocky prairies from the Canadian River, Aug. 26, 
to the Llano Estacado, August to September. On p. 3, Llano Estacado is described 
as "the space in our route between Valley River and Fossil Creek, near Tucumari 
hills. This is a dry and generally timberless tract of country extending over a distance 
of about one hundred and ninety miles". The region now would probably be included 
in Quay Co., New Mexico. 

3 Plant. Nov. Thurb. Gray, Mem. Amer. Acad. Ser. 2, v. 297-328 (1855), ex- 
plains the route followed. 


1946] Gaiser,—The Genus Liatris 353 


Var. nebraskana, var. nov., varietatem typicam simulans 
caudice subterraneo ad 4 dm. longo caulibus multis, plerumque 
minus rigidis et altioribus (2.5-5 dm. maturitate, 1-5 dm. 
juventute); foliis plerumque angustioribus, 2-3 mm. latis, 
flexilioribus marginibus aut haud aut leviter ciliatis; inflores- 
centia 10-25 cm. longa; capitulis densis ca. 1.5 cm. longis 4-6 
floris; phyllariis plerumque submembranaceis rariter marginibus 
breviciliatis, interioribus lanceolato-acuminatis vel oblongis 
apice acuto.—L. punctata Hook. var. 6. Torr. & Gray Fl. N. Am. 
ii. 69 (1841). L. resinosa sensu DC. Prodr. v. 129 (1836), not 
Nutt. 

Type: on north hillsides, south of Chadron, Dawes Co., 
Nebraska, Sept. 1, 1936, W. L. Tolstead (G). 

Wisconsin, Illinois and Arkansas, west to South Dakota, 
Colorado, and New Mexico.—WISCONSIN. Sr. Crorx Co.: 
prairie hillside, 2 mis. w. of Robert, Aug. 6, 1934, N. C. Fassett, 
16969 (G). ILLINOIS. DuPace Co.: Rwy. Lisle, July 8, 
1925, A. J. Prisc, 26 (US). MINNESOTA. Pope Co.: dry 
prairies, Montevideo, Aug. 11, 1897, L. R. Moyer (NY). HENNE- 
PIN Co.: St. Paul, 1861, T. J. Hale (G); sandy soil, Aug. 1890, J. 
H. Sandberg (NY); Minneapolis, Aug. 1868, W. M. Canby, 542 
(US), Sept. 15, 1891, J. H. Sandberg (US). Martin Co.: 
without stated locality, Sept. 10, 1892, R. I. Cratty (G). IOWA. 
Without stated locality: R. I. Cratty (US). Emmer Co.: dry 
sandy knolls, Aug.-Sept. R. I. Cratty (NY). Cra Co.: gravelly 
prairie, cemetery in Dickens, Freeman T wsp., Sect. 17, Aug. 4, 
1934, Miss A. Hayden, 10528 (NY). WoopnpBunv Co.: Sioux 
City, Aug. 29, 1896, L. H. Pammel, 39 (G). MISSOURI. 
SHANNON Co.: broad slopes & bald tops of loess hills, near 
Montier, Sept. 3, 1920, E. J. Palmer, 18944 (G). Taney Co.: 
limestone ledge, Malva, Sept. 17, 1924, E. J. Palmer, 26190 
(NY). ARKANSAS. Without stated locality: 1839, Latrobe 
(NY). CannorL Co.: limestone ledges, Eureka Springs, Sept. 
28, 1913, E. J. Palmer, 4514 (US). SOUTH DAKOTA. With- 
out stated locality :! Nicollet’s N. W. Exped. C. S. Geyer “var. B" 
(NY). Custer Co.: Custer (alt. 5500’), Aug. 1, 1892, P. A. 
Rydberg, 754 (NY). Probably BENNETT oR WASHABAUGH Co.: 
Bear Creek (albino), Aug. 1891, T. A. Williams (US). NE- 
BRASKA. Cuerry Co.: on hardlands, Fish Hatcheries, Valen- 
tine, Aug. 15, 1936, W. L. Tolstead, 702 (G). Dawes Co.:on north 
hillsides, s. of Chadron, Sept. 1,1936, W. L. Tolstead, 800 (G, type). 
Tuomas Co.: on sandhills, on Middle Loup R., near Thedford, 
Sept. 13, 1893, P. A. Rydberg, 1761 (NY); Thedford (1 tall stem 
to right), Aug. 7, 1889, H. I. Webber (NY). BurrALo Co.: without 
stated locality, Aug. 1922, W. E. B. (O). Apams Co.: Ayr, Aug. 
31, 1905, J. M. Bates (G). KANSAS. Without stated locality: 


1 See also following discussion. 


354 Rhodora [NOVEMBER 


latitude 39°, 1868, E. Hall (US). RuürEv Co.: prairie, Sept. 12, 
1895, J. B. Norton, 213 (G, NY, US); prairie, Sept. 12, 1895, 
J. B. Norton, 212 (G, US). Jouwsow Co.: s. of Holliday, Aug. 
23, 1895, C. Rowe, 980 (NY). LiNN Co.: prairie, Parker, Oct. 
21, 1916, G. W. Stevens, 4346 (NY). SEpawick Co.: Wichita, 
Sept. 1905, S. F. Poole, 226 (G); high prairies, Wichita, Sept. 9, 
1890, B. B. Smyth, 254 (US). Nrosno Co.: near Chanute, Sept. 
23, 1896, L. F. Ward (US); Brookville, Sept. 1888, R. G. Eccles 
(NY). Prarr Co.: vicinity of Pratt, Sept. 23, 1912, J. N. Rose 
& W. R. Fitch, 17149 (US). MowTraoMEny Co.: dry prairies, 
Cherryvale, Sept. 16, 1900, F. W. Johnson (NY). CowrkEv Co.: 
Winfield, Aug. 1911, Miss F. N. Vasey (US). BARBER Co.: 
Medicine Lodge, 1906, C. R. Ball, 1081 (US). OKLAHOMA. 
Co. undetermined: open rocky hills, Sansbois Mts., Aug. 1891, 
C. S. Sheldon, 293 (US). CREEK Co.: Sapulpa, Sept. 19, 1894, 
B. F. Bush, 216 (G, NY, US). Loaaw Co.: near Guthrie, Oct. 
21, 1916, R. Keyser, 6087 (NY). KiwarisHER Co.: Huntsville, 
Sept. 20, 1895, Miss L. A. Blankinship (US). OkranoMa Co.: 
sandstone hillside, Oklahoma City, Aug. 29, 1937, U. T. Water- 
fall (NY). CrEvELAND Co.: Norman, Sept. 25, 1914, W. H. 
Emig, 347 (US); 10 mis. e. of Norman, Aug. 20, 1903, A. H. Van 
Fleet (US); 2 mis. e. of Norman, W. E. Bruner (US). Murray 
Co.: Platt National Park, Sulphur, July 24, 1935, G. M. Merrill, 
979 (NY). COLORADO. Co. undetermined: dry plains, C. 
Mohr (US, 783573). Larimer Co.: Estes Park, Aug. 11, 1905, 
E. L. Johnston, 218A (NY). BovurpEn Co.: Boulder, Sept. 
1895, P. A. Rydberg (NY). Denver Co.: Denver, Sept. 23, 
1887, S. M. Tracy & Evans, 902 (NY). Er Paso Co.: Colorado 
Springs (alt. 1900 m), Sept. 22, 1915, W. W. Eggleston, 19027 
(US); Colorado Springs, 1903, H. L. Shantz, 585 (US); Pike’s 
Peak (plant to left), Aug. 23, 1887, S. M. Tracy & Evans, 80 
(NY). NEW MEXICO. Co. undetermined: from Cotton- 
wood Creek of the Arkansas to San Miguel, Aug. 1847, A. Fendler 
(328)! (G). Corrax Co.: Johnson's Mesa, Aug. 16, 1910, E. O. 
Wooton (US). SAN Miauet Co.: vicinity of Las Vegas, Sept. 11, 
1895, E. N. Plank (NY). Quay Co.: Llano Estacado, Sept. 16, 
1853, Whipple Exped., J. M. Bigelow (US, stem to right). 

Var. mexicana, var. nov. a varietate typica differt plantis 
altissimis 5-8.5 dm. altis, plerumque caulibus paucis e caudice 
crasso subterraneo vel ovato vel subelongato; foliis paucis 
plerumque omnino glabrescentibus nitidis distantibus et maiori- 
bus 12.5-15 em. longis 4-6 mm. latis; inflorescentia laxa, inter- 
nodiis 1-2 cm. longis; capitulis 1.5-2 cm. altis, 5-6-floris; phyl- 
lariis vel ovato-acuminatis vel mucronatis, marginibus ciliatis aut 
sine ciliis. 

! Plant Fendl. gives this date and region for the collection of L. punctata (328), 
which bracketed number is the one under which it was distributed. 


1946] Gaiser,— The Genus Liatris 355 


Type: dry, gravelly places, among clumps of scrub oak, end 
of road from T. Armendaiz north into Sierra del Pino, Coahuila, 
Aug. 20-26, 1940, I. M. Johnston & C. H. Mueller, 457 (G). 

Known from southwestern Texas, and from Coahuila, San 
Luis Potosí and Tamaulipas, Mexico.— TEXAS. SCHLEICHER 
Co.: s. of Eldorado, Nov. 1, 1942, H. R. Reed, 40902, 40903 
(G). Epwarps Co.: Ranch Expt. Sta., pasture G, V. L. Cory, 
5010 (G), Oct. 16, 1932, V. L. Cory 5011! (G). Er Paso Co.: 
Guadalupe Mts., Sept. 1881, V. Havard (US, 134404), 1881, 
V. Havard, 56 (US 220338). Jerr Davis Co.: side of hills, pass 
of the Limpia,? Aug. 24, 1849, Exped. from West Texas to El Paso, 
C. Wright, 246 (G, US); dry exposed slopes (alt. 1800 m.) Mt. 
Locke, Davis Mts., Sept. 1936, L. C. Hinckley (G); foothill on 
s. w. of Mt. Locke, Davis Mts., on road up to Observatory 
(alt. 1850 m.), Sept. 1936, L. C. Hinckley, 884 (NY). BREWSTER 
Co.: Pine Canyon (5000’), Chisos Mts., Sept. 5, 1937, E. Marsh, 
330 (G); Chisos Mts., July 28, 1931, C. H. Mueller, 8176 (G, 
NY); Blue Creek, Chisos Mts., June 19, 1931, C. H. Mueller, 
8176 (G); dry soil, in wide valleys, Chisos Mts., Aug. 20, 1915, 
M. S. Young (G); grassy slope of College Hill, Alpine, Sept. 20, 
1935, O. E. Sperry, T. 354 (US); frequent on College Hill, Alpine, 
Aug. 20, 1936, B. A. Warnock, 1179 (G). NEW MEXICO. 
BERNALILLO Co.: dry flats, w. slope of Sandia Mts., near Allen's 
Ranch (alt. 7000^), Aug. 10, 1914, Miss C. C. Ellis, 284 (NY, US). 
MEXICO. CoaHviILA: dry gravelly places among clumps of 
scrub oak, end of road from T. Armendaiz, north into Sierra del 
Pino, Aug. 20-26, 1940, I. M. Johnston & C. H. Mueller, 457 
(G, type); Monclova, Feb.-Oct. 1880, E. Palmer, 419 (G); 100 
mis. n. of Monclova, Feb.-Oct. 1880, E. Palmer, 420 (G, US); 
valle de los Guajes, e. of the Sierra de la Encantada, 10 mis. s. 
of Rancho Buena Vista, Sept. 3, 1941, R. M. Stewart, 1354 (G). 
TAMAULIPAS: in dry broad arroyo, 19 km. s. w. of Misquihuana, 
rd. to Palmillas, Aug. 11, 1941, L. R. Stanford, K. L. Rether- 
ford & R. D. Northcroft, 900 (G (3 stems to left), NY). SAN 
Luis Porosi: Sierra Mts., en route San Luis Potosi to Tampico, 
Dec. 1878-Feb. 1879, E. Palmer, 1085 (G); Bagre, Minas de 
San Rafael, July 1911, C. A. Purpus, 5144 (G, US). 


Liatris punctata has probably the widest geographic range of 
any species of the genus, extending from about the 53rd parallel 
of latitude in the western Canadian provinces, south ward into 
northern Mexico, though not east of the Mississippi. Hooker's 
description covers Drummond's specimen from the plains of 
Saskatchewan and that of Douglas from the Red River and 


1 This specimen is fasciated, probably as the result of some injury. 
? In Gray, Plant. Wright. p. 83 (1852), no. 246, is stated to be from Pass of the 
Limpia, which is in Jeff Davis Co., Texas. 


356 Rhodora [NovEMBER 


Eagle Hills, Sask., Aug. 4, 1827 (Kew), of which photographs 
have been seen. This plant, as it occurs in Canada from Mani- 
toba to Alberta and for the most part in those states just south 
of the border (North Dakota, Montana and Wyoming) and 
southward along the eastern Rockies to New Mexico, closely re- 
sembles the type and is here called var. typica. A variety of 
more slender and often taller habit, with narrower leaves, and 
slightly smaller heads, occurs more generally in the plains states 
from the southern border of the Dakotas to the Oklahoma-Texas 
border-region, and is here separated as var. nebraskana. 

Torrey and Gray (Fl. N. Amer. ii. 69 (1841)) recognized two 
varieties of which y, described as having "leaves conspicuously 
ciliate with hispid hairs; inner scales of the involucre purplish 
above", probably belongs with var. typica, though the involucre 
is not necessarily colored. In contrast to var. y, they recognized 
as var. 8, plants with “leaves nearly all very narrowly linear; the 
margins remotely ciliate or naked; scales of the involucre nar- 
rower, tapering somewhat gradually into a cuspidate-acuminate 
point, at least the inner ones; spike usually short; stem often 
slender". Since the writers referred to collections of the Nicollet 
N. W. Expedition, by Charles A. Geyer (NY “var. 8"), it is pos- 
sible to recognize as this latter variety the one that occurs more 
generally in the central plains-states and we are calling it var. 
nebraskana. In the National Herbarium are two specimens of 
which one, of Aug. 20, 1839, Nicollet's N. W. Exped., no. 272, 
*var. Y", from the sandy and prairie heights, around the Sioux, 
lower St. Peter R., near “Fort Pierre" is apparently var. typica, as 
are also those at the Gray and New York Herbaria, without local- 
ity, labelled “var. y” and “var. " respectively. Another speci- 
men, with the rootstock 10 cm. long (of Aug. 20, 1839, Nicollet's 
N. W. Exped., no. 272, "fere var. @ T. & G.", from sandy arid 
prairie heights, around the Sioux, Upper St. Peter R., near Fort 
Pierre (US)), as well as one “fere 6” (G), represent the more 
slender variety in the character of the narrow leaves. However, 
by the size of the heads, and ciliate-margined phyllaries, they 
match var. typica of the western Canadian prairies. Thus the 
"fere var. Q", may be translated as an intermediate between the 
two varieties. By the report and maps of this expedition (see 
footnote to citation), these specimens would have been collected 


1946] Gaiser,—The Genus Liatris 357 


about Day Co., South Dakota. Thus it seems evident that both 
var. typica and var. nebraskana are represented in the collections 
of that expedition which reached a region where the range of the 
varieties overlap, and they had also apparently collected at that 
time an intermediate that was recognized by Torrey and Gray as 
"fere B”. Certainly many specimens now justify considering 
variations from the type as a second variety. 

In numerous specimens examined, wherever the subterranean 
stem is present, it is a long, often divided rootstock. Though not 
preserved on the specimens of Drummond and Douglas and there- 
fore not mentioned by Hooker, it is quite characteristic of L. 
punctata from western Canada and southward, whether variety 
typica or nebraskana. This we believe helps to distinguish the 
species from the more southern ones with which it has been con- 
fused, namely L. mucronata and L. angustifolia, for they have 
somewhat globular corms. 

In the prairie provinces of Canada specimens are readily identi- 
fied as var. typica as they agree well with the type specimens of 
Hooker who described the phyllaries ‘margine ciliato, lanatis 
mucronato-acuminatis". Specimens from near the south end of 
the mountain range, as in Colorado and New Mexico, have very 
frequently phyllaries that are markedly mucronate and might, 
from observation of this character alone, suggest relationship to 
L. mucronata. However, some of them have leaves 2-4 mm. 
wide, with ciliate margin, and an elongate ramifying rootstock, 
belong to L. punctata and are definitely var. typica. Others are 
more nearly like var. typica than var. nebraskana, but are dis- 
tinctive in having mucronate and short-ciliate, colorful phyllaries. 
Since specimen for specimen seen, punctata var. typica from 
western Canada shows lanceolate-acuminate phyllaries and from 
Colorado and New Mexico more frequently these mucronate ones, 
we are calling these latter, forma coloradensis. 

In the central plains-states of Nebraska, Kansas, and Okla- 
homa, the great majority of specimens seen are var. nebraskana. 
For example, only three specimens of var. typica have been seen 
from Nebraska, and in Kansas only from the counties on the 
western border along Colorado. The counties listed for speci- 
mens examined from this state are curiously enough classified as 
follows: 6 (of which 5 are from the western border) for var. typica, 


358 Rhodora [NOVEMBER 


9 (all from the central to the eastern part of the state) for var. 
nebraskana, and 5 (Riley, Graham, Ellis, Ellsworth, and Kiowa) 
rather central, for questionable intermediates. J. N. Rose and 
W. R. Fitch, crossing the state, seem to have collected on Sept. 
14, 1912, in the vicinity of Syracuse, Hamilton Co. (US) (i. e. on 
the Colorado border), var. typica and on Sept. 23, 1912, in the 
vicinity of Pratt, Pratt Co. (US) (i. e. along the central southern 
border of the state) var. nebraskana, or an intermediate. 

There are, however, many specimens which are not readily placed 
varietally. As frequently happens where the range of two species 
or varieties overlap, there are apparent intermediates. Thus in 
the states marginal to the range of each variety of L. punctata 
the number of doubtful specimens increases, and a list of these, 
A, is given below. Frequently it is difficult also to allocate speci- 
mens because they seem to be just intermediate between two 
species. From a region like Oklamona, where the range of L. 
mucronata overlaps that of L. punctata, and the plains variety 
nebraskana and forma coloradensis of var. typica of the latter 
species meet, come more specimens that are puzzling intermedi- 
ates. Often specimens of the same collector’s number and date 
in different herbaria will favor different interpretations. While 
it is impossible to make definite decisions when the rootstock is 
not present, as is often the case in herbarium specimens, still, 
though lacking the association of rounded or elongate rootstock 
with other recognized specific characters, there is evidence that 
this is a region of much hybridization. List A & B below give 
some of the intergradations. 


A. Intermediates of Liatris punctata Hook var. typica & var. 
nebraskana. MICHIGAN. Katamazoo Co.: prairie roadside, 
L6 mi. n. e. of Schoolcraft, Aug. 14, 1936, C. R. Hanes, 3646 (G); 
V6 mi. n. e. of Schoolcraft, Aug. 7, 1937, C. R. Hanes, 3887 (NY); 
prairie along roadside, 14 mi. n. e. of Schoolcraft, Aug. 1, 1938, 
C. R. Hanes, 308 (NY). MINNESOTA. Cuay Co.: Moorhead, 
Aug. 15, 1901, C. A. Ballard, 3154 (G). Orrer Tatu Co.: 
Battle Lake, Aug. 1892, E. P. Sheldon (G, US); dry hills and 
sandy soils, Fish Lake, Sept. 2, 1907, Z. L. Chandonnet (US). 
HENNEPIN Co.: Fort Snelling, Aug. 22, 1883, W. H. Manning 
(G), Sept. 21, 1890, E. A. Mearns, 140 (US). Wasasna Co.: 
Lake City, Aug. 29, 1883, W. H. Manning (G); Liberty, Aug. 29, 
1883, W. H. Manning (G). IOWA. Co. undetermined: western 
Iowa, 1872, H. H. Babcock (US). Emer Co.: prairies, Herb. 


1946] Gaiser,—The Genus Liatris 359 


M. Bebb, R. I. Cratty (G). Paro Auro Co.: gravelly prairie 
knoll, Lost Island Twsp., Sect. 20, Aug. 27, 1934, Miss A. Hayden, 
10530 (G, NY); gravelly prairie in cemetery at Dickens, Aug. 4, 
1934, Miss A. Hayden, 10528a (G). NORTH DAKOTA. 
Benson Co.: Butte, Aug. 26, Sept. 29, 1907, J. Lunell (NY). 
SrARK Co.: Dickinson, Sept. 18, 1908, W. R. Holgate (NY). 
Morton Co.: Mandan, 1915, J. T. Jarvis, 160 (US). Ricn- 
LAND Co.: prairie, Hankinson, Aug. 25, 1902, C. O. Rosendahl, 
1147 (NY). SOUTH DAKOTA. Without stated locality, 
probably Day Co.: sandy, arid prairie heights, around the Sioux, 
upper St. Peter R., near Fort Pierre, dividing ridges between 
Missouri & Mississippi waters, 1839, Nicollet’s N. W. Exped. 
“fere 8" (G); Aug. 20, 1839, Nicollet’s N. W. Exped. C. A. Geyer, 
272 ("fere 8 T & G” (US)). Meane Co.: Black Hills, near Fort 
Meade, Aug. 10, 1887, W. J. Forwood (US); Park, near mouth of 
Gimlet Creek, Aug. 4, 1910, J. Murdock (G, NY). FALL 
River Co.: on prairie in canyon bottom, n. w. of Hot Springs, 
Aug. 6, 1941, G. J. Goodman, 3300 (G, NY). .Yanxron Co.: on 
high unbroken prairies, Jamesville, Aug. 24, 1899, L. A. Bruce 
(US). NEBRASKA. Keyapana Co.: Carns, Aug. 24, 1893, 
F. Clements, 2902 (G, US). Tuomas Co.: on sandhills, Middle 
Loup R., near Thedford, Sept. 13, 1893, P. A. Rydberg, 1761 
(G, P); near Plummer Ford, Dismal R., Aug. 22, 1893, P. A. 
Rydberg, 1761 (US). Deve. Co.: without stated locality, Aug. 
1890, P. A. Rydberg, 138 (NY). Kearney Co.: Minden, Sept. 
5, 1920, Dr. Hapeman (G). KANSAS. Ruivey Co.: Manhattan, 
Sept. 1, 1892, S. Norton (NY); Fort Riley, Sept. 1892, E. E. Gayle, 
603 (NY). Granam Co.: Bogue, Sept. 8, R. H. Imler (US). 
Euuis Co.: rocky prairie soil, w. of Hays, June 20, 1927, E. Bondy 
(G,US,O). ErrswonTH Co.: Kanapolis, Aug. 27, 1891, E.O. Woo- 
ton (US). Kriowa Co.: Belvedere, Sept. 5, 1898, M. White (US). 

B. Intermediates of Liatris punctata Hook & Liatris mucronata 
DC. KANSAS. McPuerson Co.: McPherson, Sept. 5, 1890, 
W. A. Kellerman (US). OKLAHOMA. Woops Co.: Alva, 
Oct. 1913, G. W. Stevens, 2866 (G); prairie, Alva, Sept. 28, 1913, 
G. W. Stevens, 2851 (G, NY, US, O); Augusta, Oct. 12, 1896, 
U. S. Geol. Survey, L. F. Ward, 51 (US). Tursa Co.: prairie, 
near Tulsa, Oct. 30, 1913, G. W. Stevens, 2082 (G).  PAwNEE Co.: 
Pawnee, Aug. 30, 1895, J. W. Blankinship (G). OKLAHOMA Co.: 
near Oklahoma City, Sept. 16-17, 1938, S. S. White, 1156 (G). 
BrecxuaM Co.: 6 mis. s. of Elk City, Oct. 17, 1936, C. T. Eskew, 
1505 (US). Comancue Co.: Fort Sill, Aug. 28, 1916, Mrs. J. 
Clemens (G); prairie, w. of Wichita National Forest, Oct. 10, 
1936, C. T. Eskew, 1429 (G, NY, US, O). Harmon Co.: bad 
lands, eroded sand-stone & gypsum desert, Oct. 24, 1936, M. 
Hopkins, 1055 (G, NY, O). TEXAS. HUTCHINSON Co.: 
5.9 mis. n. of Borger, Sept. 29, 1936, H. B. Parks & V. L. Cory, 


360 Rhodora [NOVEMBER 


16338 (T). Cray Co.: 13 mis. n. e. of Henrietta, Oct. 20, 1940, 
V. L. Cory, 40761 (G, US). Lusszock Co.: vicinity of Lubbock, 
E. L. Reed, 3162 (US); vicinity of Lubbock, E. L. Reed, 3178 
(US). Er Paso Co.: Guadalupe Mts., Aug. 22, 1901, V. Bailey, 
700 (US). 

Allowing for the modification to mucronate phyllaries in south- 
ern L. punctata var. typica forma coloradensis, specimens from the 
mountains of New Mexico are predominantly typica. Of all the 
specimens I have seen from New Mexico, none show a markedly 
interrupted spike with long internodes between larger heads, al- 
though there is a suggestion of it in one collection by Miss C. C. 
Ellis, no. 284, from the Sandia mountains (NY, US). This 
character is noticeable in specimens from the Guadalupe and 
Davis mountains of the Trans-Pecos region of western Texas 
where slightly varying plants of L. punctata are of tall habit with 
lax spike of fewer distant heads and with fewer leaves. Gray 
(Pl. Wright. 83 (1852)) refers to one of Wright’s specimens as 
follows: “no. 246 Liatris punctata Hook . . . This is one of the 
narrower-leaved forms with the scales of the involucre very ob- 
tuse and abruptly mucronate. L. mucronata DC. (founded on 
Berlandier’s no. 1926) is a similar but more depauperate state of 
the same species." Examination of this collection, from Pass of 
the Limpia, Jeff Davis Co., Texas (G), shows one rather better 
specimen to have about 25 heads distributed over 30 cm. giving 
quite an interrupted spike. From the Chisos mountains of 
Brewster county and the adjoining northwestern region of Co- 
ahuila, Mexico, come specimens with interrupted spike and 
broader, almost glabrous leaves and an obese, oval rootstock. 
Until further collections have been made we group these speci- 
mens from the southernmost mountain localities as L. punctata, 
var. mexicana. 

Not to be confused with these, however, though also showing 
loose interrupted spikes, are plants from the more eastern plains 
of Mexico, in Tamaulipas, and northeastern Coahuila. These are 
tall, more slender plants with narrow leaves and mucronate phyl- 
laries and shorter corolla-tubes and pappus that associate them 
with L. mucronata, as it is seen from Frio, San Patricio and Duval 
counties of the Rio Grande plains of Texas. The rootstock is 
generally globular as seen in typical L. mucronata DC. but may 
be ovoid, suggesting that there may here too be intermediates be- 


1946] Gaiser,—The Genus Liatris 361 


tween the two species reaching their southern limit of range. We 
have placed this material as L. mucronata var. interrupta. 

L. punctata has established as good a foothold, if not a better 
one, in the northern and mountainous latitudes as L. ligulistylis, 
the only other species in Western Canada. Then, abounding 
along the eastern Rockies, it occurs southward in New Mexico, 
E] Paso, Jeff Davis and Brewster counties, Texas, at a height of 
5000 feet or more and ''peters out" in northern Mexico in the 
varietal form (var. mexicana). What Kearney and Peebles state 
in Arizona Flowering Plants and Ferns (1942, p. 9) would per- 
haps be expected of Liatris punctata: *Many species of this cate- 
gory [i. e. of Rocky Mt. distribution] range from the Canadian 
Rockies to the Sierra Madre in northern Mexico. . . . The 
higber mountains of Arizona and the elevated plateaus in the 
northern parts of the state offer congenial habitats for the charac- 
teristic plants of this category". Yet strangely no collections of 
this or any other species of Liatris from the state of Arizona have 
been found in any herbaria examined. Liatris punctata, though 
not crossing the Rockies, made a very successful eneroachment 
on their eastern slopes. 

Inatris cylindrica Torr. (Ann. N. Y. Lyceum ii. 210 (1827)) 
obviously equals Liatris cylindracea Michx. as to name (as shown 
by reference to Flor. Bor. Amer. and to the correct page of it) but 
not as to plant. The specimen of Dr. James, of Long's Ist Ex- 
pedition (NY), to which Torrey referred, proves to be L. punctata 
var. typica, but is without locality of collection. Its source then 
is doubtful; in the list it is given as ‘sources of the Platte?’’. 
This would be too far west for L. cylindracea. Torrey made the 
observation “Dr. James’ specimen resembles those collected by 
Captain Douglas in Cass' expedition except in not being above 
ten inches high with the leaves hairy on the margin"; that collec- 
tion Torrey (Am. Jour. Sci. & Arts iv. 66 (1822)) referred to L. 
squarrosa. This should, however, not prove too confusing since 
Captain Douglas! material came from the west shore of Lake 
Michigan and may very well have been L. cylindracea. Elliott, 
indeed (Sk. ii. 276 (1824)), refers to specimens of that species 
from that locality received from Torrey. It therefore seems 
clear that Torrey intended to call the James specimen L. cylin- 
dracea, but the locality given in the text is unlikely for that species 


362 Rhodora [NOVEMBER 


and it does suit L. punctata, which the specimen actually is. 
Gray, Synop. Fl. i?. 110 (1884), also makes L. cylindrica Torr. 
synonymous with L. punctata. 

That L. punctata crosses with species of other series, as of the 
Scariosae, is seen in X L. Weaveri Shinners (see no. 18) and in the 
following hybrid. 


X LIATRIS FALLACIOR (Lunell) Rydb. emend. Shinners. Stems 
single from an elongate (over 5 cm. long), horizontal rootstock, 
3.5-6 dm. tall, mostly green with scattered pubescence basally, 
which becomes more dense above, of long white appressed hairs; 
leaves linear, also softly pubescent with margins asperous due to 
short cilia about as in L. ligulistylis; basal leaves 8 cm. long and 
3 mm. wide, gradually narrowing to become bracts subtending the 
heads: inflorescence an irregular spike, 8-20 cm. long, or in slender 
specimens reduced to but a few (2-5) heads with the terminal 
one in all specimens markedly larger than the axillary ones 
(usually 2-3 times, but in one specimen, that of Lunell no. 1016, 
4—5 times as large): heads generally 14—18-flowered, and slightly 
turbinate, ca. 1.5 cm. wide and high, at the time of flowering; 
phyllaries narrow, oblong and obtuse, mostly green with narrow 
colorful petaloid margins fringed with distinct cilia (in the co- 
type there is slight evidence of a somewhat acute cusp disappear- 
ing behind the blunt tip); corolla 8-9 mm. long, moderately pilose 
within ; achene ca. 5 mm. long; pappus 7-8 mm. long and plumose 
though less conspicuously so than L. punctata.—Fl. Prairies and 
Plains, 780 (1932); Shinners, Amer. Midl. Nat. xxix. 40 (1943). 
Laciniaria fallacior Lunell, Amer. Midl. Nat. v. 38 (1917). 
Laciniaria fallacior var. celosioides Lunell, 1. c.—NORTH DA- 
KOTA. Benson Co.: on sunny, dry prairie, Leeds, Sept. 15, 
1916, Lunell, 1015 (TYPE of Laciniaria fallacior), 1016 (TYPE of 
Laciniaria fallacior var. celosioides). 


Known only from the type (2 plants on sheet) and cotype (3 
plants on sheet) of Lunell’s collection in North Dakota. These 
few specimens clearly show an intermediacy between Liatris 
punctata and L. ligulistylis. In the elongate, horizontal root- 
stock, narrow linear leaves, general spike-like inflorescence, the 
narrow-oblong shape of the phyllaries and the plumose (though 
not markedly so) pappus they resemble the former parent. In 
contrast the general pubescence of leaves and stem (especially 
that of the inflorescence-rachis), the large terminal heads and 
their development almost to the exclusion of others in several 
specimens, the obtuse tip and the slightly petaloid margin of the 
phyllaries and the length of the achene and pappus are more 


1946] Gaiser,—The Genus Liatris 363 


similar to L. ligulistylis. The complete effect is that of a very 
marked blending of the characters of two species, belonging in the 
two sections Suprago and Euliatris, with barbellate and plumose 
pappus respectively and this is interestingly enough seen in an 
intermediacy in the extent of plumosity in the resultant hybrid. 

The type specimen of Lunell's var. celos?oides differs in no way 
except that it shows an unusually large congested terminal head, 
a result of some fasciation. 

25. Liarris densispicata (Bush), comb. nov. Stems many, 
slender, glabrous, 3-6 dm. high, from an elongate rootstock that 
runs horizontally in the sand, giving off clusters of aerial stems; 
leaves numerous, punctate, glabrous, narrowly linear, soft and 
ascending, 5-10 cm. long, 1-2 mm. wide, gradually shortening 
to the bracts of the flowering spike: inflorescence 10-30 cm. long, 
usually of densely crowded, sessile, narrow cylindrical heads of 
3-8 flowers and 8-12 mm. long; phyllaries herbaceous, oblong- 
lanceolate, acute- or acuminate-tipped, without or with a few 
marginal cilia; flowers purple, corolla 10-11 mm. long, tube 
pilose within; pappus 9 mm. long, plumose; achenes 8 mm. long.— 
Lacinaria densisipicata Bush, Amer. Mid. Nat. xii. 313 (1931). 
Lacinaria arenicola Bush, Amer. Mid. Nat. xii. 314 (1931). 

Type from Bunker prairie, Anoka Co., Minnesota. Known 
only from that state.—ANoOKA Co.: sand-dunes, Bunker Prairie, 
Aug. 26, 1927, C. O. Rosendahl, 5421b (M, TYPE), 5420 (M, type 
of Lacinaria arenicola). HENNEPIN Co.: Minneapolis, Aug., 
1868, W. M. Canby, 541, 542 (US), Sept. 15, 1891, J. H. Sand- 
berg, 933 (US). 

This species seems distinct from L. punctata, var. typica and 
var. nebraskana, which are found in Minnesota, by the generally 
finer and more slender structure of stem, spike and leaves. In 
these characters it recalls L. angustifolia, which is mostly confined 
to Texas, though rarely collected in southern Missouri, Nebraska 
and Kansas. Its long, horizontal rhizome, however, indicates 
that the Minnesota locale is not just an isolated northern station 
of the latter species. 

Lacinaria arenicola Bush, described from plants from the same 
type region, collected also by C. O. Rosendahl, seems indistin- 
guishable in any marked character and represents probably a less 
vigorous and poorly nourished plant in which some heads have 
not developed, thus giving the irregular spike shown in the type 
sheet. Dr. Rosendahl, who also collected plants for the writer 
from the same region, was unable to make any distinctions. 


364 Rhodora [NOVEMBER 


26. LIATRIS MUCRONATA DC. Corm round, about 2-4 cm. in 
diameter, bearing a number of stems, mostly glabrous, rarely 
hirsute, 3-7 dm. in height, often reddish in color, with numerous 
narrowly linear, punctate leaves, mostly without marginal cilia 
(occasionally hirsute when accompanying a hirsute stem); basal 
leaves 5-8 cm. long and 1.5-5 mm. wide, diminishing upwards 
to short bracts often exceeding the heads of the lower portion of 
the spike-like inflorescence of usually crowded, sometimes scat- 
tered, heads, 8-30 cm. long; heads cylindrical, 1.2-1.8 cm. long 
(measured to the end of the pappus), of 4-6 flowers; phyllaries 
herbaceous, outermost ones short, ovate-lanceolate with mucro- 
nate tips, becoming longer inwards, the. innermost 9-11 mm. 
long by 2-3 mm. wide, oblong, with midvein prominent, abruptly 
mucronate to cuspidate and with margin finely ciliolate or but 
membranaceous; corolla purple, 9-10 mm. in length, only 
moderately pilose in the base of the tube; filaments sometimes 
pilose; pappus 6-7 mm. long, plumose; achene 5.5-7 mm. long, 
ribbed and hairy.—Prodr. v. 129 (1836). 

Var. typica. Inflorescence a dense spike of closely crowded 
heads 1.2-1.5 cm. long; more general throughout the species- 
range. L. mucronata DC. Prodr. v. 129 (1836) sens. strict.; 
Engelm. & Gray, Pl. Lindh. i. 10 (1845); Torr. & Gray, Fl. N. 
Amer. ii. 69 (1841); not sensu Gray, Plant. Wright. 83 (1852). 
Liatris acidota var. mucronata Gray, Synop. Fl. i?. 110 (1884). 
ated Lacinaria leptostachya Bush, Amer. Mid. Nat. xii. 314 

1931). 

Chiefly in Texas, from the eastern hardwood forest region 
westward through the Edwards plateau, with a few stations 
scattered northward through the plains region of Texas into 
Oklahoma, Kansas and  Missouri.—MISSOURI. ATCHISON 
Co.: dry ground, Oct. 1, 1893, B. F. Bush, 198 (G, US). Barry 
Co.: Eagle Rock, June 22, 1897, B. F. Bush, 118 (US). KANSAS. 
SEWARD Co.: vicinity of Liberal, Sept. 23, 1912, J. N. Rose & 
W. R. Fitch, 17149 (2 stems) (NY). OKLAHOMA. kKiwac- 
FISHER Co.: Huntsville, Sept. 20, 1895, Miss L. A. Blankinship 
(G). OxLAHOMA Co.: sandy clay on sandstone hillside, 215 mi. 
w., % mi. n. of Spencer, Aug. 6, 1939, U. T. Waterfall, 1552 
(G). BEckHaAM Co.: prairie, Cedar Twsp., Oct. 18, 1936, C. 
T. Eskew, 1509 (US, O). TEXAS. Co. undetermined: rocky 
prairies, Colorado to Guadaloupe, July, 1845, Lindheimer (G). 
Potrer Co.: Templin, 1927, V. L. Cory, 2521 (G). HARDE- 
MAN Co.: abundant on prairies, Chillicothe, Sept. 27, 1906, 
C. R. Ball, 1151 (NY, US). BainEv Co.: grassy hillsides, 
2 mis. s. of Muleshoe, Aug. 24, 1921, R. S. Ferris & C. D. Dun- 
can, 3428 (NY). Grayson Co.: Denison, Oct. 15, 1932, Savage 
(NY); Bonham to Sherman, Aug. 28, 1939, B. C. Tharp (G). 
Denton Co.: prairie on Houston clay, between Denton & 


1946] Gaiser,—The Genus Liatris 365 


Aubrey, Oct. 6, 1937, W. L. McCart, 674 (US). Kaurman Co.: 
vicinity of Terrell, May 8, 1904, F. J. Tyler (US). DALLAS Co.: 
Dallas, Aug. 15, 1875, J. Reverchon (G); dry soil, Dallas, Sept. 
1877, J . Reverchon (NY); rocky bluffs, Dallas, Aug.-Sept. vi 
Reverchon, ex Herb. Curtiss, 1180 (G, NY); Dallas, 1875, J. 
Reverchon, 15 (US); Dallas, Sept. 26, 1900, B. F. Bush, 1112 (G). 
TARRANT Co.: without stated locality, Oct. 10, 1925, A. Ruth, 
78 (NY). Parker Co.: Weatherford, Oct. 18, 1902, S. M. 
Tracy, 8143! (US). Hoop Co.: dry calcareous soil, Comanche 
Park, Sept. 15, 1914, E. J. Palmer, 6541 (US). CarrAHAN Co.: 
rocky open ground, Baird, Sept. 30, 1918, E. J. Palmer, 14555 
(US). ANwpnEnsoN Co.: alt. 300’, Sept. 20, 1937, G. L. Fisher, 
37168 (US). Brown Co.: Brownwood, Oct. 23, 1916, E. J. 
Palmer, 11106 (US). Lampasas Co.: 8 mis. n. w. of Lampasas, 
Sept. 23, 1935, H. B. Parks & V. L. Cory, 15692 (T). REAGAN 
Co.: 11 mi. n. w. of Stiles, Nov. 18, 1942, V. L. Cory, 40954 (G). 
WILLIAMSON Co.: Georgetown, Aug. 1929, M. D. Cody (F). 
Luano Co.: Enchanted Rock, Oct. 12, 1930, Miss E. Whitehouse 
(G). Travis Co.: dry calcareous soil, open ground, Austin, 
Sept. 21, 1916, E. J. Palmer, 10766 (US); Austin, Sept. 11, 1877, 
L. F. Ward (US); near Austin, Oct. 11, 1940, R. R. Innes, 105 (G). 
Bianco Co.: 614 mis. n. of Blanco, Sept. 23, 1935, V. L. Cory, 
15691 (G). Surron Co.: Roy Hudspeth’s, Sept. 3, 1938, V. L. 
Cory, 2520 (G); Schoolhouse Hill, Sonora, Sept. 12, 1942 (no. 
40175), Oct. 27, 1942 (no. 40828) V. L. Cory (G). Austin Co.: 
San Felipe (Austin) Jan. 1835, T. Drummond, 122 (G, NY). 
Hays Co.: San Marcos & vicinity, 1898, S. W. Stanfield (NY). 
Kerr Co.: Kerrville, Oct. 4, 1916, E. J. Palmer, 10896 (US). 
Comancheries (KERR & KENDALL Co. probably) from Boerne 
to Comfort, Nov.-Dec. 1828, Berlandier, 1926 (G, isotype). Ex 
prov. Mex. 1839, ex Herb. Musei Britanici (US). Comat Co.: 
Comanche Spring, New Braunfels, Sept., 1849, Lindheimer 940 
(G, NY, US, P), Sept., 1850, 941 (G, NY, US, P, O), 1849-53, 
942 (G, NY, US), Sept., 1850, 943 (G, NY, US, O). Bexar Co.: 
3 mi. southwest of Smithson Valley, Sept. 23, 1935, H. B. Parks 
& V. L. Cory, 15687 (T); vicinity of San Antonio, Oct. 19, 1912, 
J. N. Rose, 18016 (US), Sept. 3, 1921, Miss E. D. Schultz, 610 
(US), G. Jeremy, 39 (G), Aug., 1937, Sister Mary C. Metz, 3030 
(NY). 

Var. interrupta, var. nov., a varietate typica differt spicis 
interruptis, capitibus 1-3 cm. inter se distantibus 1.5-1.8 cm. 
longis.—TvPE from south of Mathis, San Patricio Co., Texas, 
Oct. 20, 1927, J. N. Rose & P. G. Russell, 24156 (G). 

Tamaulipas, San Luis Potosí, and northeastern Coahuila, 
Mexico, and some of the bordering counties of Texas.— TEXAS. 


1 The specimens at G, NY & T. might be considered intermediates between mucro- 
nata and angustifolia. 


366 Rhodora [NOVEMBER 


Without stated locality: Drummond (G). OrpnaAM Co.: Bravo, 
Oct. 7, 1909, C. R. Ball, 1593 (US). RaANparL Co.: Palo Duro 
Canyon, Sept. 2, 1907, C. R. Ball, 1217 (NY, US). Bexar Co.: 
San Antonio, Sept. 3, 1921, Miss E. D. Shultz, 610 (US). Uvar- 
DE Co.: Uvalde to Hondo, Oct. 10, 1936, B. C. Tharp (G). Frio 
Co.: 14 mis. s. of Pearsall, Sept. 18, 1939, F. Shreve, 9442 (G). 
San Parricio Co.: s. of Mathis, Oct. 20, 1927, J. N. Rose & R. 
G. Russell, 24156 (G, type, NY, US). Duvar Co.: San Diego, 
1884-1888, Miss B. Croft, 21 (NY, US). MEXICO. Coa- 
HUILA: Sierra de Santa Rosa, s. of Musquiz, July 13, 1938, E. G. 
Marsh, 1290 (G); Palm Canyon, Musquiz, Sept. 19, 1936, E. G. 
Marsh, 965 (G); Del Carmen Mts., Sept. 7, 1936, E. G. Marsh, 
792 (G); Rancho Aqua Dulce, Musquiz, July 1, 1936, F. L. Wynd 
& C. H. Mueller, 399 (G, US). TaAMaurrPAS: sierra near San 
Lucas, Jaumave, July, 1932, H. W. Von Rozynski, 520 (3 stems 
to right) (NY). 


Liatris mucronata has been passing as Liatris punctata or, as 
explained (see no. 4), has been erroneously appended to Liatris 
acidota as a variety. The differences between L. mucronata 
and L. acidota have already been noted under the latter species. 

The type of L. mucronata was collected Nov.—Dec. 1828, by 
Berlandier, no. 1926, from the Comancheries orientales, Texas 
(Geneva) while that of L. punctata (Kew) was collected on the 
plains of Saskatchewan. Though the type plant in Geneva has 
not been seen, a photograph of it and an isotype in the Gray 
Herbarium have been compared. In neither is there any root- 
stock present. Engelm. & Gray (PI. Lindh. i. 10 (1845)) clearly 
recognized the species, when describing the new species L. aci- 
dota, and referred to Lindheimer’s collections of both species 
They included in the description of L. mucronata ‘‘caudice glo- 
boso." Other specimens collected from Texas, similar in other 
characters to L. mucronata, show that a globose or rounded corm 
is characteristic of this species in contrast to the elongate and 
often branched rootstock of L. punctata. The two species in 
their type regions differ also in the nature of the phyllaries which 
are abruptly mucronate and cuspidate in this species, rather than 
lanceolate-acuminate as in L. punctata from the western Canadian 
provinces. L. mucronata has also a shorter pappus, a character 
in which, as in the globular rootstock, it shows resemblance to L. 
angustifolia. 

However, variations occurring in L. punctata over its great 
north-south range, undoubtedly help to explain some of the con- 


1946] Gaiser,—The Genus Liatris 367 


fusion of these two species. As stated under L. punctata (see 
no. 24), mucronate phyllaries, rather than long lanceolate-acumi- 
nate ones are common in f. coloradensis from the southern end of 
the range in Colorado and New Mexico. In the regions between 
these states and Texas, there are plenty of specimens giving 
evidence of a blending of the characters of L. punctata and L. 
mucronata. "Thus from the northern plains of Texas, from Okla- 
homa, Nebraska and Kansas come perplexing specimens which 
may represent intermediates between the two. Certainly when 
lacking rootstocks, as frequently happens, specimens are difficult 
to determine. So far, in southern Texas from the eastern border 
through the counties of the hardwood forest, blackland prairie, 
Edwards plateau and Rio Grande Valley, where L. mucronata 
abounds, no specimens with a long ramifying rootstock have been 
seen. From the northern plains of Texas, only one specimen has 
been seen, H. B. Parks and V. L. Cory from 5.9 mi. n. of Borger, 
Hutchison Co. (which is close to the Oklahoma border), Sept. 29, 
1935, no. 16338 (T), which has an elongated rootstock along with 
mucronate phyllaries that suggest L. punctata f. coloradensis. 
For other specimens, as in the list of intermediates, it has been 
less easy to make specific determinations. 

As quoted, in the discussion of L. punctata, Gray compared 
what was probably the only specimen he had from the El Paso 
region of Texas (Chas. Wright's no. 246 (G)) with L. mucronata 
because of its obtuse and abruptly mucronate bracts. No root- 
stock is present on the specimen. Yet it shows few heads sepa- 
rated by long internodes, in contrast to the quite dense spike-like 
arrangement in the type of L. mucronata, which Gray considered 
a ‘more depauperate state" of L. punctata. Gray’s opinion may 
have contributed further to the uncertainty of the determination 
of this species. Sufficient specimens have now been seen to re- 
fute the idea that L. mucronata DC. is just à depauperate form 
of L. punctata. Also such an interrupted spike as is seen in 
Wright's specimen of L. punctata is found in the most southern 
phase of L. punctata—var. mexicana. 

A plant that has so far been collected very rarely from the 
coastal plain region of Texas, with few large heads of 8-10 
flowers, often singly disposed along stems that may branch, and 
having longer achenes and pappus, seems a greater variant, and 
is here given specific rank as L. bracteata. 


368 Rhodora [NOVEMBER 


L. mucronata is to be distinguished from L. angustifolia, which 
occurs with it through Texas and casually resembles it, by the 
smaller heads with glabrous bracts and the softer smooth leaves, 
though here again there may be found intermediates, e. g., S. M. 
Tracy, No. 8143 from Weatherford, Parker Co., Texas, Oct. 18, 
1902 (NY and T) (see discussion under L. angustifolia). 


27. Liarris angustifolia (Bush), comb. nov. Stems slender, 
glabrous, 6-8 dm. high, often reddish in color, from a globose 
corm usually 2-4 cm. in diameter: leaves narrowly linear, soft, 
epunctate or hardly at all punctate; basal ones 5-10 cm. long, 
1-3 mm. wide, reduced progressively in length below a short 
spike to short bracts subtending the heads, but in longer spikes 
only so shortened throughout its length: inflorescence slender, 
20-60 cm. long, of densely crowded heads, 8-16 mm. long, 
sessile or rarely becoming pedunculate, as following injury: 
heads slender, cylindrical, 3—6-flowered, ca. 1.5 cm. in length; 
phyllaries lanceolate, outer ones broader, inner ones narrower, 
acuminate, thin-papery, glabrous with merely membranous or 
slightly ciliolate margins in exceptional cases, somewhat glutin- 
ous; flowers purple, corolla 9-10 mm. long, slightly pilose within 
on the tube and noticeably so on the filaments of the stamens; 
pappus ca. 7 mm. long with lateral cilia more than fifteen times 
the diameter of the seta though not long-plumose; achenes 5-7 
mm. long.—Lacinaria angustifolia Bush, Amer. Mid. Nat. xii. 
315 (1931). 

From the inner margin of the coastal plain region of Texas, 
northward and westward through the plains region into Okla- 
homa and occasionally Kansas, Nebraska, Arkansas and Mis- 
souri.—MISSOURI. Barry Co.: rocky slopes, bald knobs, 
along Mo.-Ark. line, Eagle Rock, July 27, 1926, E. J. Palmer, 
31455 (G). NEBRASKA. Dawes Co.: Crawford, Aug. 13, 
1896, E. N. Plank (NY). Cass Co.: prairies, Weeping Water, 
1888, T. A. Williams (US). KANSAS. Ritey Co.: Manhat- 
tan, Aug. 30, 1892, S. Norton (NY). Miawr Co.: Paola, Aug. 
1883, Aug. 1885, Dr. Oyster, 3503 (NY). Sumner Co.: high 
prairies, Caldwell, Sept. 10, 1890, B. B. Smyth, 279 (NY, US). 
OKLAHOMA. Creek Co.: along rwy. w. of Sapulpa, Sept. 6, 
1913, F. W. Pennell, 5378 (NY). CLEVELAND Co.: 10 mis. e. of 
Norman, Aug. 20, 1903, A. H. Van Fleet (US); Denver, Little 
River, Aug. 21, 1903, A. H. Van Fleet (O). PrrrsBURG Co.: 
meadow, Sept. 22, 1934, J. E. McClary (O). Murray Co.: 
Platt National Park, G. M. Merrill & W. A. Hagan, July 16, 
1935, 924 (NY, US, O); Sept. 30, 1935, 1535 (NY, O); near 
Sulphur, Aug. 5, 1939, H. Broadbent (O). CHocraw Co.: 
prairies, Fort Towson (Arkansas. Dr. Leavenworth) (G, NY). 


1946] Gaiser,—The Genus Liatris 369 


TEXAS. Fannin Co.: (plant to right), Bonham, Sept. 21, 
1877, L. F. Ward (US). Grayson Co.: Denison, Sept. 13, 1906, 
F. J. Tyler (US). Tarrant Co.: dry soil, Fort Worth, Sept. 5, 
1912, A. Ruth, 78 (M, 211347 type); rocky lands, Fort Worth, 
July 30, 1909, A. Ruth, 78 (US); without stated locality, Sept. 
1898, A. Ruth, 78 (US). Darras Co.: hills, Dallas, Sept. 26, 
1900, B. F. Bush, 1112 (NY, US); vicinity of Dallas, June 20, 
1929, Miss M. R. Stephenson, 129 (US); rocky bluffs, Dallas, 
1880, J. Reverchon (US). Kaurman Co.: Preston Bend, Sept. 
16, 1906, F. L. Tyler (US). SoMERvELL Co.: Paluxy valley, 
6 mis. above Glen Rose, Oct. 11, 1891, L. F. Ward (US). Cory- 
ELL Co.: rocky hillside, near Elige, Aug. 14, 1931, S. E. Wolff, 
3269 (US). San AvausrINE Co.: San Augustine, G. L. Crockett 
(US). Branco Co.: 10 mis. s. of Blanco, Sept. 23, 1935, V. L. 
Cory, 15683 (G). GirLEsPIE Co.: Half Moon, G. Jermy, 806 
(US). Harris Co.: Houston, Nov. 2, 1913, G. L. Fisher, 501 
(US). KENDALL Co.: Spanish Pass, Aug. 26, 1936, V. L. Cory, 
19382 (G); 6 mis. n. w. of Boerne, Sept. 22, 1936, V. L. Cory, 
20703 (G). Comat Co.: New Braunfels, July 1846, F. Lind- 
heimer (US). 

This species can be distinguished from L. acidota (with long 
basal leaves, looser, narrower spike lacking noticeable subtending 
bracts because of the abrupt reduction of the leaves basally to 
setaceous bracts leaving a strict almost naked stalk) by its spike 
of densely crowded heads with bracts progressively shortened 
from the uppermost of a gradually diminishing series of leaves. 
Also, the range of L. acidota is limited to the coastal plains region 
of Texas, whereas this species occurs back of the coastal plain. 
As Bush states of Lacinaria angustifolia, “This is the species, I 
have no doubt, that has been the basis of Missouri, Kansas, 
Oklahoma and Arkansas being cited as part of the range of L. 
acidota.”’ 

Bush described Lacinaria angustifolia as having “involucral 
bracts ovate-lanceolate, pointed or cuspidate, densely pubescent 
on the backs, ciliate on the margins, not punctate’. Examina- 
tion of the type specimen (no. 211347 Minn.) from a cultivated 
individual, grown in the greenhouse from seeds of Texas plants, 
failed to show that the phyllaries are pubescent; but as they are 
lanceolate-acuminate and as the specimen otherwise compares 
well with the species here in mind, Bush’s name has been taken 
up. 

Among the citations given by Bush 1. c. for this species is A. 


370 Rhodora [NOVEMBER 


Ruth, no. 78, Tarrant Co., Texas, Sept. 5, 1912 (Ruth Her- 
barium). A specimen of this collection at the Herbarium of the 
University of Minnesota, sheet no. 211348, is the given type of 
Bush’s Lacinaria Ruthii (Amer. Mid. Nat. xii. 316-17 (1931)). 
The description of that species is not so very different from that 
of angustifolia though the involucral bracts are given as ‘oblong, 
cuspidate or pointed, slightly ciliate on the margins, glabrous". 
Examinations of the type specimen and duplicates thereof (G, 
NY) as well as other specimens seen from Texas (e. g. Oct. 25, 
1917, A. Ruth, no. 736, from sandy woods, near Fort Worth, 
Tarrant Co. (NY), Oct. 18, 1902, S. M. Tracy, no. 8143 from 
Weatherford, Parke Co. (G, NY, T)) suggest an intermediate 
condition between Liatris mucronata and L. angustifolia. The 
former covers much the same range as L. angustifolia. In the 
characters of spike, corolla, pappus and achene it also resembles 
it but the phyllaries have a prominent midrib that becomes a 
distinct cusp.. Because the leaves of L. angustifolia are not dis- 
tinetly punctate and are soft and pliable, they are usually dis- 
tinguishable from those of equally narrow-leaved specimens of 
L. mucronata, which varies from a narrowly to a rather broadly 
(15 mm. wide) linear leaf and is occasionally hirsute. However, 
as the centre of range for both of these species lies in central 
Texas there are found confusing intermediates between them. 
From L. punctata var. nebraskana, L. angustifolia is especially 
distinguishable by the rounded rather than elongate rootstock, 
the shorter corolla and pappus and the shorter lateral setae of the 
pappus making it seem short-plumose by comparison. The dif- 
ference between the two is well shown on one sheet (NY) having 
a specimen with rootstock from Manhattan, Kansas, Aug. 30, 
1892, S. Norton (NY), representing L. angustifolia, and a fasciated 
stem only (without rootstock) Sept. 1, 1892, representing L. 
punctata var. nebraskana. That the two intergrade seems quite 
possible. When examining a number of collections of B. F. Bush 
of different dates, from an isolated area like that of the mounds, 
Holt Co., Mo. (see list below) it seemed impossible to make 
definite determinations. Some, as no. 12026, and no. 12356, with 
their narrow leaves and reddish stems, resemble L. angustifolia, 
while others, as no. 12006 and no. 12009, are more like L. punctata 
var. nebraskana. Thus it seems possible that in this zone inter- 
gradations had occurred and had been successfully propagating. 


1946] Gaiser,—The Genus Liatris 371 


Probable intermediates between L. punctata Hook. var. ne- 
braskana and L. angustifolia Bush.—MISSOURI. Horr Co.: 
on the mounds, Watson, Sept. 3, 1920, B. F. Bush, 9185, 9185a 
(NY), Sept. 30, 1930, Bush, 12006, 12009 (NY); high mounds, 
Mound City, Oct. 14, 1930, Bush, 12026, 12035 (NY), Sept. 29, 
1931, Bush, 12356 (NY). OKLAHOMA. CuocraAw Co: 
limestone prairies near Hugo, Oct. 6, 1923, E. J. Palmer, 24046 
(G). 

28. Liarris bracteata, sp. nov., herba pauci- vel multiramosa, 
laesa vel culta ramosior; caulibus glabris e cormo subgloboso ca. 
4 em. diametro; foliis linearibus glabris punctatis rigidis leviter 
canalieulatis, basalibus 7-12 cm. longis 2-3 mm. latis demum 
rigidis, superioribus ad bracteas subinde capitula superantes 
reductis; spica laxissima; capitulis paucis magnis plerumque 
8-10-floris subturbinatis 2 cm. longis summo pappo 1.5 cm. latis 
inter se 1-3 cm. distantibus, vel saepe capitulo solo terminali 
10—14-flore; phyllariis exterioribus ovato-acuminatis, interioribus 
late lanceolatis plerumque cuspidatis 10-12 mm. longis 3-4 mm. 
latis saepe purpurascentibus marginibusque longe ciliatis; 
corollis purpureis 9-11 mm. longis, tubo intus sparse piloso; 
pappo 10-15 mm. longo plumoso; achaeniis 8-12 mm. longis.— 
Known only from Texas.—Harris Co.: 15 mi. north of Houston, 
along Hwy. no. 290, Oct. 1936, R. G. Reeves (G, TYPE). TEXAS. 
Without stated locality: 1857, Leybold, ex Mus. Bot. Berol. 
(US, 616780); C. Wright (G). Harris Co.: without stated 
locality, Sept. 1850!, G. Thurber (G); ca. 14 mis. n. of Houston, 
along Hwy. no. 290, Aug. 29, 1936, R. G. Reeves, L. O. Gaiser & P. 
Snure, 48 (G); road from Cypress to Houston, Oct. 11, 1897, F. 
W. Thurow, 1 (US); Houston, 1842, F. Lindheimer (G); vicinity 
of Houston, Oct. 28, 1913, J. N. Rose, 18130 (US); Houston, Oct. 
9, 1918, Œ. L. Fisher, 193 (US). GarvEsTON Co.: without stated 
locality, Sept. 12, 1941, Mrs. A. F. Nelson (G). Maraacorpa Co.: 
north edge of town of Gulf, Oct. 10, 1934, V. L. Cory, 11566 (G); 
altitude 20', Matagorda, Oct. 14, 1936, G. L. Fisher, 3682 (US). 


Plants of the same population as the type had been seen and 
collected before the time of flowering, on Aug. 29, 1936, by the 
writer, along with Dr. R. G. Reeves who had seen it growing 
along Highway no. 290 for several years. Reports for November, 
1943, are that the stand has been temporarily destroyed by fire 
but it is hoped that the sturdy underground rootstock may still 
have survived. 

The species has seemingly been rarely collected. It is closely 

! Plant. Nov. Thurb. Gray (1855) begins only with an account of collections made in 


Oct.—Nov. 1850, and we are therefore unable to give the exact locality for this speci- 
men, dated Sept. 1850, but it is probably from between Houston and Austin. 


372 Rhodora [NOVEMBER 


matched by a specimen of V. L. Cory from the vicinity of Gulf, 
Matagorda Co., Texas (G), which he described under another 
name in an unpublished note seen at the Gray Herbarium, in 
which he called attention to the terminal heads. Because the 
heads are not merely terminal and axial ones do occur and de- 
velop, even though separated by long internodes, the name 
bracteata is here being used. 

The species most closely resembles L. mucronata var. interrupta 
in having few heads distributed in a lax spike but is distinctive in 
having heads of more and larger flowers, longer achenes and 
pappus (which is also long-plumose) than are found in that 
species. Cytological evidence too has borne out the need of con- 
sidering it a new species. 


SUMMARY OF THE PUNCTATAE 


It seems to the writer that plants of the Punctatae series, which 
undoubtedly spread westward and northward from some center, 
as Texas, had their origin in a species with a rounded corm, such 
as is common to most of the genus. L. mucronata, abundant 
throughout most of Texas, and represented in the Rio Grande 
plains and a little southward in Mexico by a variety showing a 
lax and elongated inflorescence, could have been such a species. 
However there also occurs quite abundantly in Texas L. angusti- 
folia, having soft fine leaves and acuminate rather than mucro- 
nate bracts. An ancestral form of either species might have given 
rise to L. punctata. 

Plants spreading westward, on reaching the extreme south- 
western region of Texas and the interior of New Mexico, where 
the mountains make a foothold more difficult, seem to have de- 
veloped elongate ramifying rootstocks. This characteristic, 
evident in L. punctata of the Trans-Pecos region, was apparently 
successful enough for the species to have followed along the 
mountains northward, even to the Canadian prairies. In har- 
mony with the suggestion of such migration is the observation 
that, in going northward, one finds a change from the predomi- 
nantly mucronate phyllaries in specimens from the southern 
mountains to generally lanceolate ones in specimens from 
Canada. Or the development of a branching and spreading root- 
stock might have come along with a direct northward extension 


1946] Gaiser,—The Genus Liatris 373 


from the warmer central region of Texas of such an angustifolia- 
like form. 

There are no sharp lines of demarcation between any two 
species except the marked characteristic of two types of root- 
stock. There is a general appearance common to the three 
species of broader and the two of more localized range which 
makes them, as a group, distinguishable from all other species, 
though it has often brought them all under the common name of 
L. punctata. L. acidota, when recognized, cannot be grouped 
with this alliance though it was wrongly associated with members 
of it, perhaps because some do show a rounded rootstock. 


SERIES IX. CYLINDRACEAE. Plants of intermediate height, 
3-9 dm. tall, from loosely cymose, branched and bushy to single- 
stalked; heads few, 15-60-flowered, cylindrical to turbinate, 
approaching 2 cm. in height and 1-2 cm. in diameter; phyllaries 
herbaceous, appressed, obtuse to mucronate, sometimes spread- 
ing at the apices but never recurved; achenes 5-9 mm. long.— 
From southern Ontario westward to Minnesota and south to 
Missouri, with one species each in Florida and Texas. 


a. Inflorescences cymosely branched; corolla-lobes non-pilose 
within....b. 

b. Heads compact, cylindrical; phyllaries closely imbricated, 
Uchennneronesimucronateg d oc nes 29. L. cymosa. 

b. Heads loose, turbinate to hemispheric; phyllaries loosely 
imbricated, the inner ones obtuse..............-. 30. L. Ohlingerae. 

a. Inflorescence generally a short cymose raceme; corolla-lobes 

manifestly pilose within; heads compact, cylindrical; phyl- 
laries closely appressed, obtuse to mucronate........ 31. L. cylindracea. 


29. Lratris cymosa (H. Ness) K. Sch. Corm rounded, up to 
3 em. in diameter: stems stiff, upright, 2.5-6 dm. tall, dichoto- 
mously cymosely branched above: leaves mostly glabrous, punc- 
tate, linear-lanceolate, the radical 15-20 cm. long and 5-15 mm. 
wide, tapering to a clasping petiole, the cauline linear, gradually 
reduced upwards: inflorescence finely pubescent and bearing heads 
in a distributed manner in either a simple or compound cyme; 
heads 2-2.5 cm. high and 7-10 mm. broad, of about 20 flowers; 
phyllaries appressed, closely imbricated, in about 6 series, slightly 
hirsute, ciliate-margined; the outer almost orbicular with rounded 
or truncate apices, the inner oblong with mucronate tips, often 
colored; flowers purple; corolla 15 mm. long, inner surface of 
lobes and tube smooth; pappus ca. 8-10 mm. long, plumose; 
achenes 8 mm. long, hispid on the ribs.—Just, Bot. Jahresb. 
xxvii. pt. i, 528 (1901). Laciniaria cymosa H. Ness, Bull. Torr. 
Bot. Club, xxvi. 21 (1899). 


374 Rhodora [NOVEMBER 


In Brazos and Washington counties, Texas.— Without locality, 
1897, A. M. Hildebrandt (US). Brazos Co.: 1 mi. southeast of 
A. & M. College, Oct., 1896, H. Ness (NY, type); College Station, 
June, 1897, R. H. Price, 9 (US), 1935, H. B. Parks, 14538 (G), 
14543 (T). WasuiNGTON Co.: without locality, Aug. 15, 1938, 
Miss E. Brackett (G); B. C. Tharp (G). 


Until seeing the two collections from Washington Co., Texas 
(G), which, as learned from Prof. B. C. Tharp, were collected 
twenty to thirty miles from the type-locality, I had known this 
species only from the one region where it was first discovered in 
1896, by Dr. H. Ness, one mile south of the Agricultural and 
Mechanical College of Texas, growing on stiff clayey soil so poor 
that only a few species of grass dispute the ground with it. 
“Differing strikingly from other species in the scattered heads 
borne in a loose compound cyme", it nevertheless has heads of 
approximate size, flower-content and phyllaries that seem to re- 
late it to L. cylindricacea. However, the inner surface of the 
corolla-lobes is glabrous rather than pilose as in that species. 
Thus we have again a parallel to the variation in the Scariosae 
series, where one species, L. ligulistylis, lacks the pilosity in the 
corolla-throat which other species of the series have. 


30. LIATRIS OHLINGERAE (Blake) B. L. Robinson. Root 
elongate, tuberous, somewhat segmented or lobed, fleshy, 2.5-4.5 
em. long, and about 0.7 cm. in diameter: stems 1-3, 6-9 dm. high, 
simple, stiff and puberulous: leaves numerous, narrow, linear, 
punctate, glabrous and sessile, the basal up to 5 em. long and 2 
mm. wide, gradually diminishing upwards: heads several, on 
slender bracteolate pedicels 1-5 em. long, or more numerous 
(8-18) in a cymose panicle, 2.5-3 em. high and, when flowers are 
open, hemispherical though containing only about 25 flowers; 
phyllaries long, erect, loosely appressed and not greatly over- 
lapping laterally; the outer suborbicular to obovate, the inner up 
to 2.5 em. long, lanceolate, obtuse-tipped, all herbaceous with 
narrow scarious or ciliolate usually purplish margins; phyllaries 
drop away almost completely leaving an exposed honeycombed 
receptacle; flowers ca. 1.5-2 cm. long; corolla purple, without 
pilosity, with long, spreading, almost rotate lobes; pappus a 
little shorter than the corolla-tube, ca. 13 mm. long and plumose; 
achenes 7-9 mm. long.—Contrib. Gray Herb. civ. 49 (1934). 
Lacinaria Ohlingerae Blake, Bull. Torr. Bot. Club 1. 203 t. 9 
(1923). Ammopursus Ohlingerae Small, Bull. Torr. Bot. Club, 
li. 393 (1924). 

In Polk and Highlands counties, Florida.—Pouk Co.: in scrub 


1946] Gaiser,—The Genus Liatris 375 


formation, 9 mi. southeast of Frostproof, Nov. 1, 1922, Mrs. F. 
E. Ohlinger (G, US, Type); in white-sand scrub, Lake Clinch, 
Frostproof, Sept. 23, 1934, (G, US), Sept., 1936 (US). Ohlinger. 
HicHuanps Co.: sandhills near De Soto City, Aug. 31, 1922, 
J. K. Small, J. W. Small & J. B. DeWinkeler, 10681 (G, F); dry 
pineland, Lake Stearns, Nov. 4, 1927, O. F. Burger & E. West 
(F); serub, Avon Park, Sept. 4, 1934, J. K. Small & E. West (F); 
scrub, Sebring, Sept. 5, 1934, Small & West (F); palmetto scrub, 
8 mi. southeast of Childs, Aug. 15, 1945, L. O. Gaiser, Mrs. E. H. 
Butts & Miss L. Arnold (F). 


'The rare Liatris Ohlingerae, known from but few stations in the 
adjoining counties of Polk and Highlands, Florida, was considered 
by Small to constitute a new genus, Ammopursus. It is indeed a 
very distinctive species by reason of its somewhat segmented and 
lobed root, its very large heads with loose phyllaries that overlap 
laterally very little and long flowers having an almost rotate 
corolla and also by an exposed somewhat honeycombed receptacle 
after the phyllaries have dropped away. However, because of 
the similarity of the achenes, pappus and punctate leaves, as well 
as general flower-characters, it is here considered a species of the 
genus Liatris. As it seems to approach L. cymosa more closely 
than L. scariosa in leaves, size and number of flowers per head, 
as well as in lack of pilosity in the corolla-tube, it is placed next 
to it in the Cylindraceae series. Like L. cymosa, it has so far 
been found only in a very limited region. 


31. LIATRIS CYLINDRACEA Michx. Stems one to several, 3-6 
dm. high, mostly glabrous though sometimes scantily hirsutulous, 
from a rounded corm up to 3 cm. in diameter: leaves linear, 
rigid, punctate, mostly glossy, glabrous, sometimes ciliate at base 
or along margin and rarely with the lower or both surfaces covered 
with short white hairs; radical leaves often 20 cm. long, 4-5 mm. 
wide, the cauline few and shorter along a lax inflorescence of 
usually 5-20 elongate heads of 30-60 flowers (sometimes only a 
single terminal head develops but undeveloped buds in lower 
axils appear in such plants); involucre herbaceous, appressed, 
making the large heads, 2-3 cm. long, and 1-1.5 em. in diameter, 
appear ovoid, before anthesis; phyllaries mostly glossy, rigid, 
ovate and rounded at summit with abrupt mucronate to acumi- 
nate tips, the outer sometimes spreading in older heads but 
never recurved or squarrose; corolla purple or rarely white, 12-14 
mm. long, with the inner surface of the spreading lobes distinctly 
hairy; achene 5-6 mm. long; pappus 1 em. long and plumose.— 
Fl. Bor.-Amer. ii. 93 (1803); Ell. Sk. ii. 275 (1822?); DC. Prodr. 


376 Rhodora [NOVEMBER 


30 (1836); Torr. & Gray Fl. N. Am. ii. 69 (1841), excl. syn. Willd. ; 
Gray Synop. Fl. i?. 109-110 (1884). L. intermedia Lindl. Bot. 
Reg. t. 948 (1825). Liatris stricta Macnab, Edinb. New Phil. 
Jour. xix. 60 (1835). Liatris flexuosa Thomas, Amer. Jour. Sci. 
xxvii. 338 (1839). Liatris squarrosa 8 intermedia DC., Torr. & 
Gray Fl. N. Am. ii. 68 (1841); Gray Synop. Fl. i?. 109 (1884). 
Liatris squarrosa sensu Torr. in Am. Jour. Sci. iv. 66 (1827), 
non (L.) Willd. Laciniaria squarrosa var. intermedia Mac 
Millan, Metasp. Minn. Valley, 506 (1892). Liatris marginata 
Cass. Dict. Sci. Nat. xxvi. 236 (1823). Liatris monocephala 
Cass. ibid. 237. Laciniaria nervata Greene, Pittonia iv. 317 
(1901). Liatris cylindracea var. solitaria MacM. Bot. Gaz. xv. 
333 (1890). Laciniaria cylindracea f. solitaria MacM. Metasp. 
Minn. Valley, 506 (1892). Laciniaria cylindracea var. solitaria 
Farwell, Rep. Mich. Acad. Sci. xvii. 171 (1916). 

From southern Ontario and western New York westward to 
Minnesota and Missouri.—ON'TARIO. | MaurrOoULIN Co.: limy 
soil, La Cloche Peninsula, July 11, 1937, A. S. Pease & R. C. 
Bean, 26205 (G); thin soil on limestone, Cloche Bluff, MeGregor 
Bay, July 25, 1915, F. V. Coville (US); limestone flats, Cloche 
Peninsula, Aug. 20, 1932, N. C. Fassett, 14885 (G); Strawberry 
Island, Aug. 9, 1892, W. Herriot (WH). WATERLOO Co.: dry 
ground near Galt, Sept. 5, 1905, J. E. Kerr (G, WH). YORK 
Co.: High Park, Toronto, Aug. 5, 1901, J. M. Macoun (G, NY); 
east Toronto, Aug. 7, 1892, C. W. Armstrong (US); on vacant 
lot, Babys Point, Toronto, Aug. 14, 1942, W. C. Mansell, 6453 
(HB); Humber plains, near Toronto, July 28, 1898, W. Scott, 
22770 (Ot); sandy soil, High Park, Toronto, Aug. 26, 1938, G. D. 
Darker (G); High Park, Toronto, Aug. 24, 1927, Sept. 15, 1929, 
Aug. 9, 1930, H. H. Brown (HB). PEEL Co.: sandy soil, Port 
Credit, Aug. 11, 1891, J. White, 965 (Ot). Brant Co.: moor- 
land ground e. of Brantford, Aug. 11, 1834, J. MacNab (P, 
probably isotype of Liatris stricta MacNab). Nomrorkx Co.: 
moist sandy meadow, Long Point, L. Erie, Aug. 25, 1938, H. A. 
Senn & J. H. Soper, 552 (G, NY); Turkey Point, Sept. 4, 1927, 
F. L. Davis (NY); Silver Hill, Aug. 3, 1936, H. H. Brown, 5261 
(HB). Muippiesex Co.: dry sandy woods, London, July 29, 
1879, Burgess (Ot); sandy woodland, London, July 29, 1879, 
Sept. 30, 1882, T. Millman (To). Lampton Co.: near Sarnia, 
C. K. Dodge, Aug. 22, 1892, 11586 (Ot), 1896 (To); Port Francis, 
Sauble R., Sept. 1, 1883, Burgess (Ot): sandy dunes along L. 
Huron near Port Franks, Sept. 2, 1929, E. J. Palmer, 36258 (G); 
Grand Bend, Aug. 22, 1932, H. H. Brown (HB). Kent Co.: 
dry soil in open woods, Rondeau Park, Aug. 14, 1934, R. F. Cain, 
1246 (To. NEW YORK. NracanaA Co.: whirlpool woods, 
July 1877, D. F. Day (NY); Niagara Falls, G. W. Clinton (NY); 
bluffs of Niagara R., Aug. 19, 1875, Morong (NY). Erw Co.: 


1946] Gaiser,—The Genus Liatris 377 


Buffalo, ex Herb. G. W. Clinton (Q). MICHIGAN. Cuer- 
BOYGAN Co.: jack-pine plains, s. of Burt Lake, Aug. 22, 1920, 
J. H. Ehlers, 1263 (G), July 30, 1928, F. C. Gates, 15388 (US); s. 
of Indian R., Aug. 21, 1981, H. A. Gleason (NY). CRAWFORD 
Co.: vic. of Grayling, July 1922, C. V. Piper (G, US). Iosco 
Co.: Oscoda, Aug. 23, 1906, H. H. Rusby (NY). Sm. Cuarr Co.: 
Fort Gratiot (Port Huron), 1829, Dr. Pitcher (N Y). WASHTENAW 
Co.: bank of Huron R., 14 mi. n. e. of Cedar Bend, Aug. 19, 1935, 
F. J. Hermann, 6989 (NY); open oak slope, Cedar Bend, 1 mi. 
n. e. of Ann Arbor, Aug. 5, 1937, F. J. Hermann, 9131 (G, US); 
dry bank, Ypsilanti, Sept. 16, 1917, B. F. Chandler (US). Kara- 
MAZOO Co.: n. w. part of Texas Twsp., July 16, 1938, C. R. Hanes, 
288 (NY). OHIO. FRANKLIN Co.: Georgesville, Aug. 29, 1892, 
W.C. Werner (NY). Apams Co.: Buzzard Rock, Sept. 11, 1937, 
F. Bartley & L. L. Pontius, 611 (NY). INDIANA. La 
PonTrE Co.: sandhills, s. shore of L. Michigan, Michigan City, 
Aug. 12, 1909, G. L. Fisher, 14 (US). PonrEÉR Co.: Dune Park, 
Sept. 25, 1914, F. W. Johnson, 1567 (NY); open sandy woods, 
Portchester, Aug. 29, 1915, F. W. Johnson, 2122 (US). LAKE 
Co.: sandy knolls, Edgmoor, Aug. 31, 1889, L. N. Johnson (G); 
Whiting, Aug. 29, 1893, N. L. Britton (NY); Pine, Aug. 22, 1915, 
F. W. Pennell (NY); Gibson, O. E. Lansing, Jr., 3914 (G). 
STARKE Co.: dry gravelly hillside, e. of Bass Lake, July 24, 1906, 
C. C. Deam, 1243 (NY). Nerwron Co.: prairie area along Pa. 
Rwy. & road 24, 1 mi. w. of Goodland, Aug. 14, 1943, R. C. 
Friesner, 17895 (G). Crank Co.; dry sands, Clarke, Aug. 4, 
1896, L. M. Umbach (US). WISCONSIN. Co. undetermined: 
sandy pine barrens around Thunder Lake, Aug. 1884, D. H. Hasse 
(NY). Price Co.: bluffs of Mississippi R., Aug. 9, 1926, P. A. 
Rydberg, 9671 (NY). 'TREMPEAULEAU Co.: Blair, 1889, F. F. 
Wood (US). Apams Co.: July, J. R. (US). Juneau Co.: 
Camp Douglas, E. A. Mearns, Aug. 22, 1890, 139 (US), July 16, 
1890, 130 (G). CorvuwnBia Co.: Dells of Wisconsin, D. H. Hasse 
(NY). Savk Co.: pine sand barrens, Delton, Aug. 6, 1892, R. H. 
True (G); sandy bluffs of Wisconsin R., vic. of Kilbourn, Aug. 
25-6, 1909 (no. 12), Aug. 28, 1909 (no. 56) E. S. Steele (US). 
MILWAUKEE Co.: Milwaukee, J. A. Lapham (G, NY). JEFFER- 
son Co.: Busseyville (F. Atkinson) 1872, T. Kumlien (Q). 
Dane Co.: s. of Madison, Aug. 30, 1893, J. R. Churchill (G). 
WarwonTH Co.: Delavan, Sept. 1887, Mrs. J. M. Milligan (US). 
Rock Co.: Beloit, Sept. 1882, L. H. Bailey, Jr. (G); along the 
Chicago & N. W. Rwy., Clinton, Sept. 1, 1909, E. S. Steele, 101 
(US). LaravETTE Co.: Fayette, Aug. 21, 1889, L. S. Cheney 
(G). ILLINOIS. Without stated locality: Chapman (NY), 
Babcock (G), Aug. 1872, H. H. Babcock (US), Mead, ex Herb. 
H. P. Sartwell (G), Dr. Mead (G), Vasey, 1873 (US), Vasey (G). 
Co. undetermined: prairies, Sept. 1837, C. W. Short (G, US). 


378 Rhodora [NOVEMBER 


Laxe Co.: dry sandy ridges, along lake shore, Waukegan, 1906, 
H.A. Gleason & F. D. Shobe, 376 (G); Waukegan, Sept. 3, 1877, 
R. E. Earle (US). McHenry Co.: Cary, H. C. Benke, Nov. 18, 
1932, 5615 (G, US), Aug. 11, 1928, 4808 (US). WiNNEBAGO Co.: 
prairies, Aug. 1859, M. S. Bebb (G); Fountaindale, M. S. Bebb 
(NY, US). SmEPHENSON Co.: hilltops, along cliffs, Freeport, 
Sept. 1, 1901, C. F. Johnson (US). Cook Co.: Riverside, Aug. 
1911, J. M. Greenman (G); Chicago, H. H. Babcock (G, NY, US), 
Nov. 1887, ex Herb. W. Boott (G). Dv Pace Co.: Banks, Aug. 
11, 1891, W. S. Moffatt, 350 (US); Pine Hills, Aug. 16, 1878, F. S. 
Earle (ND). Kane Co.: Aurora, Sept. 1884, T. E. Boyce, 1172 
(G). Henry Co.: Galva, 1883, Mrs. C. N. S. Horner (G). 
Peoria Co.: dry prairies, Peoria, F. E. Macdonald, Sept. 1903 
(G), Aug. 1903 (NY). HkNpnERsoN Co.: prairies near Oquawka, 
Sept. 1875 & 1880, H. N. Patterson (NY); rocky hillsides, near 
Oquawka, 1883 (no. 15), Sept. & Oct. (no. 533), Sept. 1887 
(no. 534) H. N. Patterson (US); Oquawka, Sept. 1876, H. N. 
Patterson (ND, Q). CnaAMPArGN Co.: dry ground, Champaign, 
Aug. 2, 1899, H. A. Gleason, 886 (G); cutting on I. C. R. R. 
Sept. 11, 1909, A. S. Pease, 12402 (G). Cass Co.: dry prairies, 
Beardstown, Aug. 1842, C. A. Geyer (G, NY). MapnisoN Co.: 
meadows e. of St. Louis, Aug. 1836, Diehl (NY). MINNESOTA. 
Porr Co.: Glenwood, Aug. 1891, B. C. Taylor (NY, US). Henv- 
NEPIN Co.: Fort Snelling, Aug. 7, 1909, C. O. Rosendahl, 2346 
(G); copses, Aug. 1889, J. H. Sandberg (US). Kanpryonr Co.: 
Spicer, Aug. 1892, W. D. Frost (US). WABASHA Co.: Lake City, 
Aug. 1883 (NY), Sept. 1883 (G), W. H. Manning. Goopnur Co.: 
Featherstone, Aug. 1893, A. P. Anderson (G, US); Zumbrota, 
Aug. 1892, C. A. Ballard (US). WiNoNA Co.: dry open bluffs, 
Sept. 1905, J. M. Holzinger (NY); Winona, Sept. 1889, J. M. 
Holzinger (US). Nostes Co.: dry hills, Adrian, Aug. 29, 1895, 
Miss J. B. Patten (G). IOWA. Co. undetermined: prairies, 
Aug. 1815, W. W. Denslow (NY). FavETTE Co.: dry prairies, 
Aug. 1894, B. Fink (G), Aug. 3, 1894, B. Fink, 328 (US), Aug. 26, 
1897, J. R. Garraner, 727 (NY). DusuaqurE Co.: prairies, 
Dubuque, Aug. A. Morr (G). Harpin Co.: vicinity of Iowa 
Falls, Aug. 1928, M. E. Peck, 132 (G). MISSOURI. Sr. Lovis 
Co.: Meramec Highlands, Aug. 22, 1917, J. M. Greenman, 3833 
(G, NY); St. Louis, Drummond (G); St. Louis prairies, July 
1833, Engelmann (G); rocky hills, St. Louis, Aug. 30, 1875, H. 
Eggert, 143 (US); St. Louis, Aug. 30, 1875, H. Eggert (G, NY, 
US); Allenton, Aug. 25, 1883, G. W. Letterman (NY, US). 
SHANNON Co.: dry rocky woods, near Montier, Oct. 5, 1920, 
E. J. Palmer, 19295 (G); rocky woods, Montier, B. F. Bush, 
Oct. 8, 1905, 3596 (G, US), Sept. 10, 1908, 5127 (G, US), Aug. 5, 
1910, 6109 (US); common in woods, Montier, Aug. 2, 1899, 
B. F. Bush, 221 (NY, ND, type of Laciniaria nervata Greene); 


1946] Gaiser,—The Genus Liatris 379 


rocky woods, Montier, Aug. 4, 1927, B. F. Bush, 11421 (US). 
Wriaut Co.: 2 mis. s. e. of Cedar Gap, July 28, 1937, J. A. Steyer- 
mark, 23737 (NY). GREENE Co.: dry ground, Sept. 4, 1893, 
B. F. Bush, 197 (G); thin woods, vicinity of Strafford, Aug. 27, 
1912, P. C. Standley, 9497. (US). Ozank Co.: rocky glades above 
river bluffs, near Tecumseh, Oct. 8, 1927, E. J. Palmer, 32958 
(G); Tecumseh, Oct. 9, 1927, E. J. Palmer, 33036 (NY). AR- 
KANSAS. Baxter Co.: limestone ridges, hillsides, Cotter, 
Oct. 28, 1913, E. J. Palmer, 4770 (US); dry rocky woods, Cotter, 
Aug. 31, 1915, E. J. Palmer, 8398 (US). 


Liatris cylindracea was described by Michaux, from the mead- 
ows of Illinois as “tota hirsutula". Elliott (Sk. ii. 275-6 (1822?)) 
though including this species in his list of plants for South Caro- 
lina and Georgia “on the somewhat questionable authority of 
Pursh . . . questionable as regards the habitat of his species", de- 
scribed a glabrous plant which he had received from Dr. Torrey 
and which had been collected on the shores of Lake Michigan. 
He pointed out that: “although by a many flowered involucrum, 
and the want of pubescence, it varies from the description of 
Michaux, it yet resembles his plant in too many respects to be 
easily separated from it". By 1841, Torrey and Gray (Fl. ii. 69), 
having seen more specimens, described it as: “glabrous or slightly 
hairy", and reported it from Missouri to Upper Canada and 
Michigan. 

Specimens at present available from various herbaria indicate 
that this species 1s to be found mostly in the states lying along the 
upper part of the Mississippi River: on the west side in those ap- 
proximately north of the Missouri-Arkansas state-line, and on 
the east side in those north of the Ohio River, including Indiana, 
Michigan and the southern peninsula of Ontario, and in Niagara 
County of New York. The specimens from Manitoulin Island in 
Georgian Bay, which is north of its general range in Ontario, 
suggest that the species had come there by way of the narrow 
straits from northern Michigan since it is found in Cheboygan 
County. It appears northward from about where L. squarrosa 
leaves off. Where the range of the two overlap, as in Missouri 
and southern Illinois, specimens are more frequently hirsute 
than in Ontario where they are generally glabrous, and this would 
explain Michaux’s descriptive term “hirsutula’’, since he did not 
travel farther west or north than southern Illinois (see no. 6). 
It is also probable that intermediate forms showing a blending of 


380 Rhodora [NOVEMBER 


the hirsute character of L. squarrosa and the nonrecurved bracts 
of cylindracea occur. The two species have a common character 
in the pilosity on the inside of the corolla-lobes which was first 
noted by Nuttall, under his description of L. squarrosa (Gen. ii. 
132 (1818)). Intermediates between the two varieties glabrata 
and hirsuta of L. squarrosa have been mistaken for L. cylindracea 
but such specimens can be recognized by the recurved bracts. 
Throughout the range of L. cylindracea there is a diversity in 
the outer phyllaries, which may be less mucronate-tipped, ap- 
pearing acute and more elongate. This additional varying char- 
acter may have been responsible for the description by Lindley 
of a new species, Liatris intermedia, grown from roots collected 
in Ontario by Mr. Goldie (see discussion under no. 32), since 
Michaux’s description of L. cylindracea as being “tota hirsutula” 
and few-flowered would indicate a different plant. From the 
plate, showing cylindrical buds with appressed phyllaries, there 
seems little doubt that L. intermedia is correctly placed, with 
humerous other Ontario collections, under L. cylindracea. The 
fact that MacNab (Edin. Phil. Jour. xix. 60 (1835)) also described 
under the new name L. stricta a plant collected in Ontario from 
the roadsides for several miles through the pine barrens east of 
Brantford, Aug., 1834 (an isotype, if not the type, was seen at 
Philadelphia), and that again, Thomas (Amer. Jour. Sci. xxxvii. 
338 (1839)) gave a new name, L. flexuosa, to a specimen from the 
east bank of the Niagara River, seems to indicate that Michaux’s 
phrase had limited extremely the application of his description 
by other authors. By comparison with a photograph of Mi- 
chaux’s type, seen at the Gray Herbarium, the descriptions and 
figures of all of these Ontario plants fit into the concept of his 
species as seen more frequently at the northern end of its range. 
Being a species of shorter inflorescence with fewer large heads 
(as is L. squarrosa in the Squarrosae series and L. ligulistylis in 
the Scariosae series), specimens are frequently found having but 
one well developed terminal head. In the axils of the upper 
leaves of such plants, however, one or more small aborted buds 
are often discovered, evidence perhaps of some unsatisfactory 
conditions for full development of the plant and a consequent 
reduction in the number of heads developing, for the corm in 
some such specimens has been found to be partly rotted, as in 


1946] Gaiser,—The Genus Liatris 381 


H. C. Benke, no. 5616, from Cary, McHenry Co., Ill., Nov. 18, 
1932 (G, US). Also in a plant of several stems a single stem 
might be one-headed. Thus it seems hardly necessary to recog- 
nize the plant with a single terminal head as a variety, as did 
Farwell in Lacinaria cylindracea var. solitaria (Rep. Mich. Acad. 
Sci. xvii. 171 (1916)), or even as a form, as did MacMillan in 
Laciniaria cylindracea forma solitaria (Metasp. Minn. Valley 506 
(1892)). 

Though no report in the literature has been found of white- 
flowered plants of this species, one large plant was found, Aug. 
21, 1940, by the writer, no. 213, at Turkey Point, Oxford Co., 
Ont. (G), adding one more species to the list in which albinos 
exist. As was frequently experienced after transplantation, the 
albino died at the end of the next flowering season, though other 
individuals persisted several years longer. 


X Liarris Guapewirzi (Farwell) Shinners (L. aspera or 
sphaeroidea X cylindracea). Corm irregular or globose, 3-4 cm. 
in diameter: stems to a dozen, 4-9 dm. high, glabrous and striate 
below and pubescent above: leaves linear-lanceolate, punctate, 
glabrous to but scantily pubescent beneath, the lower up to 25 
em. long and 0.7-1.4 em. wide, the upper reduced, narrowly 
linear and acute: inflorescence an open raceme, 10-30 cm. long, 
of ca. 6-9 heads 1—4 cm. distant: heads 20-30-flowered, on short 
pedicels 0.5-1 cm. long, cylindrical to slightly turbinate, 1.8 cm. 
long and 1-1.2 cm. wide; phyllaries glabrous, punctate, obtuse 
and quite closely appressed, herbaceous with a scarious purplish 
margin, the outer ca. 0.3 cm. in diameter, almost orbicular or 
ovate, erose on the margin, the middle and inner oblong, 1-1.3 
em. long and 0.5 cm. wide with more colorful and more erose 
margins but not becoming crisped; corolla purple, 1-1.3 cm. long, 
with the lobes conspicuously hairy and the tube pilose within; 
pappus 9-10 mm. long, short-plumose, thus rather intermediate 
between barbellate and plumose; achene 6-7 mm. long and 
blackish Amer. Mid. Nat. xxix. 37 (1943) (L. cylindracea X 
sphaeroidea). Lacinaria Gladewitzii Farwell, Amer. Mid. Nat. 
x. 48 (1926) (Lacinaria cylindracea X scariosa var. sphaeroidea). 

Reported only from southern Ontario, Michigan and Wiscon- 
sin. —ONTARIO. Kenr Co.: sandy ground, Rondeau, Sept. 5, 
1931, N. C. Fassett, 15026 (W). LAMBTON Co.: sandy ground, 
pine-oak scrub, ca. 5 mi. south of Grand Bend, Sept. 6, 1942, 
L. O. & W. H. Gaiser, 243 (G). MICHIGAN. Oak.Lanp Co.: 
dry hills, Rochester, July 26, 1926, O. A. Farwell, 7584 (Herb. 
Cranbrook Inst., type). WISCONSIN. Crawrorp Co.: dry 


382 Rhodora [NOVEMBER 


summit of limestone bluff, Prairie du Chien, Aug. 20, 1927, 
N. C. Fassett, 4478 (W). 

Farwell found his type of Lacinaria Gladewitzii growing in 
association with what he called “L. scariosa and its variety 
sphaeroidea and L. cylindracea” and he stated it was an “exact 
intermediate between the last two”. 

Though neither the type specimen of L. Gladewitzii nor any 
material of what Farwell called Lacinaria scariosa var. sphaero- 
idea are available for examination at this time, two other collec- 
tions by Dr. N. C. Fassett, referred to Liatris Gladewitzii by 
Shinners have been seen. One of these, from sandy ground at 
Rondeau Park, on Lake Erie in Southern Ontario (W), was re- 
ported to be growing with L. sphaeroidea and the other came from 
limestone bluffs, in Crawford Co., Wise. (W). Also the writer 
has found one plant growing with a population of L. cylindracea 
south of Grand Bend, on the south-east shore of Lake Huron, 
where along the sandy stretch of pine-oak woods are also to be 
found L. aspera Michx. and X L. sphaeroidea Michx. 

These different collections seem to bring confirmation to the 
occurrence of this hybrid. The specimens resemble L. cylindra- 
cea in the narrow, rigid, almost glabrous leaves, the open raceme- 
like inflorescence of comparatively few short-pedicelled heads, 
the appressed phyllaries and especially the conspicuous hairs on 
the inside of the corolla-lobes. In the characters of the stem, 
pubescent above, and the phyllaries, narrowly scarious and a 
little colorful but not at all bullate, and the pilose corolla-tube, 
they bear resemblances to L. aspera or X L. sphaeroidea. The 
latter, as here interpreted and abounding around the Great 
Lakes, has become a very stabilized hybrid showing less crisped 
phyllaries and thus less globular and more campanulate heads 
than L. aspera (see no. 18). 


(To be continued) 


1946] Fernald,—Presumable Identity of Cheilanthes lanosa 383 


THE PRESUMABLE IDENTITY OF CHEILANTHES 
LANOSA 


M. L. FERNALD 


In 1803 Michaux, Fl. Bor.-Am. ii. 270 (1803), described from 
the mountains of Tennessee and North Carolina his Nephrodium 
lanosum: 


LANOSUM. N. parvulum; elegans; totum lanosis- 
simum: fronde oblonga, bipinnatifida; 
pinnis distantibus; pinnulis pinnatifi- 
dis; lobulis subrotundo-ovalibus, integris: 
punctis demum contiguis. 

OBS. Habitus quodammodo Po typ. fontani; 
paulo majus. 

HAB. in montibus saxosis Tennassée et 
Carolinae septentrionalis. 


Certainly Michaux's “totum lanosissimum”’ and his description 
of the frond and especially its pinnules are promptly matched by 
the extremely lanate fern of the mountains of North Carolina and 
Tennessee which was described thirty years later as Cheilanthes 
tomentosa Link, Hort. Berol. ii. 42 (1833), . . . "stipes tomento- 
sus... , pinnae . . infra dense tomentosae", which in his full 
description D. C. Eaton, Ferns N. Am. i. 346, 347, rendered, 
"stalks . . . covered with . . . soft woolly hairs . . . The 
fronds . . . of a grayish color from the abundance of fine en- 
tangled tomentum." Are not these descriptions very close to 
Michaux's “totum lanosissimum’’? 

Michaux saw in his Nephrodium lanosum the habit of Poly- 
podium fontanum of Europe, i. e. Asplenium fontanum (L.) 
Bernh., but the new American species was “a little larger". 
Hegi describes Asplenium fontanum as “Bis 25 cm hoch", 4. e. 
10 inches. Eaton, l. c., says of C. tomentosa: “The fronds vary 
from a few inches to over a foot in length". 

On the mountains of North Carolina and Tennessee there is 
another species of Cheilanthes, which was described only one year 
after Nephrodium lanosum of Michaux. This is Adiantum 
vestitum of Sprengel, Anleitung, iii. 122 (1804). 


Adiantum vestitum nenne ich eine Art, die Bose d’Antic in 
Karolina fand. Sie hat einen dreyfach gefiederten Wedel, der iiber 
und über mit feinen woltichten Haaren bedeckt ist. Die Blattchen 


384 Rhodora [NOVEMBER 


der ersten und zweyten Ordnung sind ey-lanzetförmig; die der letzten 

Ordnung sind linienförmig, gekerbt und schlagen sich um die Saamen- 

häufchen zurück. Bosc nannte dies Farrenkraut Acrostichum 

hispidum. 

Adiantum vestitum Spreng. (1804) from Carolina soon became 
Cheilanthes vestita (Spreng.) Swartz, Syn. Fil. 128 (1806) and 
under this name it was generally recognized in practically all 
works up to and through the 6th edition of Gray’s Manual (1890), 
the Pteridophyta by D. C. Eaton, there (p. 681) properly described 
as “hirsute” and with the same characteristic illustration (pl. 
xvii) as had appeared in earlier editions, although these treat- 
ments may have had other species mixed with C. vestita. That 
Cheilanthes vestita and C. tomentosa are wholly distinct species no 
one questions; but that C. vestita is at all the plant clearly 
described as Nephrodium lanosum Michx., “totum lanosissimum", 
I can not believe. Neither did the earliest students of the group, 
who had the Michaux material before them. Thus, in 1804, 
Poiret, writing at Paris with Michaux’s herbarium at hand, gave 
a more detailed account of the Michaux plant in Lamarck’s 
Encyclopédie, v. 538 (1804), as Polypodium lanosum, although 
Poiret, with true French courtesy, ascribed P. lanosum to Mi- 
chaux, a natural enough treatment since at the beginning of his 
long treatment of Nephrodium Michaux had entered *'Porx- 
PODIUM. L.". It is not necessary here to repeat Poiret’s tran- 
scription of the text of Michaux (already quoted); Poiret’s own 
additions, based obviously on the material before him, were as 
follows: 


105. Potypope laineux. Polypodium lanosum. Michaux. 


Polypodium pumilum, lanuginosum, fronde bi- 
pinnatá; foliolis oblongis; pinnulis linearibus, loba- 
tis, obtusis; stipite subcylindrico, ruffo. (N.) 


ave es. 48 e's ee R & 4 4- 9 4.9 y» M E. 9 - W 8. €-9 * 9* 5 5.9 ‘l 5. €.» AR. 


C'est une plant peu élevée, d'un port agréa- 
ble, lanugineuse sur toutes ses parties, dont les 
pétioles sont droits, roides, d'un brun foncé, 
cylindrique; un peu comprimés, garnis de folio- 
les alternes, distantes les unes des autres, munies 
de pinnules opposées, presque pinnatifides, fort 
petites, linéaires, divisées en lobes ovales, arron- 
dis, trés-entiers. La fructification consiste en pe- 
tits points épars, trés-rapprochés. 


1946] Fernald,—Presumable Identity of Cheilanthes lanosa 385 


Cette plante croft sur les rochers pierreux de 
la Caroline & dans quelques autres endroits de 
l'Amérique septentrionale. (V. s. Comm. Bosc.) 
Elle a beaucoup de rapports avec le polypodium 
fragrans, Desfont. 


It can hardly be affirmed that Poiret was describing something 
different and imagining characters not stated by Michaux, 
especially when the Michaux Herbarium was at his elbow. "The 
facts that Swartz, Syn. Fil. 58 (1806), in transferring Nephro- 
dium lanosum to Aspidium as A. lanosum, said "fronde tota 
lanosissima", while, on p. 128, in changing Adiantum vestitum 
Spreng. to Cheilanthes vestita, he said “frond . . . hispidulis", 
were corroborative, although they were somewhat literal tran- 
scripts of the original diagnoses. But, after many experiences 
with Michaux's species, my faith in the accuracy of André 
Michaux, his editor, L. C. Richard, and Poiret, who more fully 
described many of Michaux's plants, is so great that I place far 
more weight upon their descriptions than upon the confusions 
apparently made in the probably subsequent placing of loose 
labels upon the much-handled old specimens by a presumably 
non-botanical mounter. 

A sheet in Michaux's Herbarium at Paris (a beautiful photo- 
graph taken by Mrs. Weatherby before me) containes 6 broken- 
off fronds which very clearly belong to Cheilanthes vestita, the 
plant with hispid or hirsute (not tomentose or lanate) fronds. 
This sheet has pasted on (presumably at a later date) the label 
of “HERB. MUS. PARIS” bearing at the bottom “Herbier de 
l'Amérique septentrionale d’ANDRK Mricnavx", and below that 
the label in the handwriting of André Michaux of Polypodium 
lanosum, with the “Hab. in excelsis montibus saxosis Tennessee 
et Carolina septentrionalis 291." "The label is that of Nephro- 
dium lanosum, Michaux having removed his genus Nephrodium 
from the inclusive Polypodium of Linnaeus subsequent to writing 
the label; but the sheet of 6 specimens to which it became at- 
tached is not at all of plants "totum lanosissimum". Whether 
in the Michaux Herbarium or in those of Lamarck or of Poiret 
there is a Michaux sheet with very lanate fronds is an academic 
question which may sometime be settled. Certainly the sheet of 
specimens with the labels does not contain the plants which 
Michaux (or L. C. Richard) and, afterward, Poiret described. 


386 Rhodora [NovEMBER 


Such mixtures of labels, added to sheets which had apparently 
earlier been mounted, are occasional through the Michaux and 
other old Herbaria. In my own work with other groups I háve 
sometimes noted them.! Since the sheet which now bears 
Michaux's label “Polypodium lanosum’’, etc. has 6 fronds, it is 
significant that when D. C. Eaton studied Michaux's material in 
1866, he stated that there were “five medium-sized fronds" 
(D. C. Eaton in Canadian Naturalist, v. 26 (1870)). It is evident 
that Michaux’s Herbarium has had more than a single sheet 
which has passed as Nephrodium lanosum. I can not subscribe 
to the argument that, when Michaux and then Poiret described a 
plant as “totum lanosissimum" they really meant one which is 
merely hispid and not at all lanate. 

The first few species described from eastern North America of 
what is now the genus Cheilanthes were hopelessly misunderstood. 
'These confusions were specially concentrated in the late 50's of 
the last century. Thus, in his Species Filicum, ii. 98 and 99 
(1852), Hooker described as “Cheilanthes ‘vestita, Sw.’?” (the 
interrogation indicating his doubt) and illustrated (his t. CVIII. 
B) a fern with “stipites . . . as well as the main rachis... 
laxly woolly, fronds . . . at the margins beneath and on the 
partial rachis, densely woolly the wool more or less tawny”. 
As synonyms he gave (1) Cheilanthes lanuginosa Nutt. (an 
herbarium-name which was later, through Nuttall's material, 
identified with the western C. Feei Moore, based on Myriopteris 
gracilis Fée, not Cheilanthes gracilis Kaulf. (1824); (2) Nephro- 
dium lanosum Michx. (1803), with the derived binomial As- 
pidium lanosum Sw., just as he had included the original Adian- 
tum vestitum “‘ ‘Spreng. Anleit. iii. p. 122". The latter reference 
was quoted and the identity of the “densely woolly” “Cheilanthes 
vestita" was doubted by Hooker because he had not seen and 
seems rather to have doubted Swartz's correct description and 
Schkuhr’s accurate illustration of it. Hooker said (p. 99): 
“What we here describe and figure as Cheilanthes vestita is . . . 
no doubt the Nephrodium lanosum of Michaux, Fl. Bor. Am. 
(1803), and he properly describes the fronds as ‘lanosissimae’. 
Swartz, however, who adopts Sprengel's (prior?) specific name, 


1 For example see plate 1045 in Ruopona, xlviii, (1946) with the label over, instead 
of beneath the base of the leaf. 


1946] Fernald,—Presumable Identity of Cheilanthes lanosa 387 


vestita, given in a work to which I have no immediate access, 
describes the fronds as hispidulous. Schkuhr adopts the same 
term, and figures a plant, the under side of which gives no idea 
of the really woolly nature of the frond; having, moreover, 
entire oblong pinnules, with a solitary terminal involucre’’. 
Michaux was correct in describing one plant, the lanate C. 
lanosa (Michx.) D. C. Eaton; Sprengel and Schkuhr were as 
vividly (even to the involucre as shown by Schkuhr) correct in 
describing another species, C. vestita (Spreng.) Swartz! 

It is needless for one who is not a pteridologist and who is 
helpless in trying to understand all those who are, to follow all the 
subsequent confusions, but at least one other must be noted. 
This was D. C. Eaton's abbreviated and rather confused para- 
graph, without a word of description, in Torrey's Botany of the 
Mexican Boundary, 234 (1859), where the combination Cheilan- 
thes lanosa was based on a doubted basonym, with at least two 
other synonyms involved, thus giving us the now supposedly 
sacrosanct combination which has been in vogue for half a cen- 
tury, C. lanosa, for a plant which is not lanate! Here is Eaton's 
paragraph: 

CHEILANTHES LANOSA. C. vestita, Hook. l. c. p. 98, t. 108, B. 
Nephrodium lanosum, Michz. Fl. Bor.-Am. 2, p. 270? Myriopteris 
gracilis, Fée, l. c. p. 150, t. 29, f. 6. Along the Rio Grande; 
Wright. 'The name of C. vestita unquestionably belongs to the 


fern described and figured by Professor Gray under that name 
in the Manual, (2d. ed.) p. 592, t. 10. 


Embarrassingly enough, the last item is the only one that was 
well founded. Eaton, although making the transfer, doubted the 
identity with the others of the Michaux plant; Myriopteris 
gracilis is by all students now considered a separate species, 
Cheilanthes Feei Moore, while the plant of Charles Wright, which 
inspired the paragraph, was later identified by Eaton, in his 
Ferns of N. Am. i. 41 (1878) as C. lanuginosa Nutt. (originally a 
synonym only of Hooker's confused C. vestita, but validated in 
1863 by D. C. Eaton, although this was later than C. Fee; Moore 
(1857) which, I am told, is the same species). 

In his Ferns of N. Am. l. c. 13-15, Eaton got identities more 
straightened out, for he correctly took up the name Cheilanthes 
vestita for the plant with “fronds . . . hirsute . . . ; . . . the 


388 Rhodora [NOVEMBER 


ends of the roundish or oblong lobes reflexed, and forming sep- 
arate herbaceous involucres". But he still kept in its synonymy 
the perpetually misinterpreted Nephrodium lanosum Michx. 
(“totum lanosissimum") with the first unequivocal name for it 
under Cheilanthes, ‘‘Cheilanthes lanosa, D. A. WATT, in Journal of 
Botany, February, 1874, p. 48: not of Moore, Index Fil., p. 245, 
nor of Eaton, Mex. Boundary Botany, p. 234, which synonyms 
belong to Ch. lanuginosa, Nuttall”. On page 15, arguing for 
the retention of “well known" names, he said ‘‘Michaux’s name, 
Nephrodium lanosum, is undoubtedly the first published of the 
various names for this fern [C. vestita, with the characters as 
originally given by Sprengel and by Swartz] . . . Usually it is 
well to keep the oldest specific name when it is known; but . . . 
to endeavor to replace well-known specific names by older, but 
obscurer ones, is surely reprehensible”. 

I have been told that it is “reprehensible” to displace the name 
Cheilanthes tomentosa Link (1833) by C. lanosa (Michx.) D. C. 
Eaton (1859), based on Nephrodium lanosum Michx. (1803), for 
the former name is “established”, while the name C. lanosa has 
(erroneously) “become established" for C. vestita which is not 
lanose! I can hardly subscribe to this philosophy, even though 
Michaux's label got affixed, probably after his death, to a sheet of 
specimens which lacks the characters given by him, and in more 
detail by Poiret, from the original and perhaps now lost lanate 
specimens. For quite as long a period, through the 6th edition 
of Gray's Manual, the name C. vestita was correctly used for the 
hirtellous species. When it was “reprehensibly’’ displaced, 
through error, by the name C. lanosa established usage of that 
period was certainly (and unjustifiably) upset. It seems to me 
that the name CHEILANTHES LANOSA (Michx.) D. C. Eaton 
(1859), based nomenclaturall on Nephrodium lanosum Michx. 
(1803), should replace C. tomentosa Link (1833), if the original 
descriptions mean anything. 


1946] Fernald,—Some American Forms of the Lady-fern 389 


SOME TRIVIAL AMERICAN FORMS OF THE 
LADY-FERN 


M. L. FERNALD 


Most of the so-called “varieties” of Athyrium Filix-femina 
(including A. angustum (Willd. Presl and A. asplenioides 
(Michx.) Desv.) have already been properly transferred to 
formal rank but the few which follow seem to require such trans- 
fer. My apology for so treating them will be found in the dis- 
cussion below. 


ATHYRIUM FILIx-FEMINA (L.) Roth, var. MicHauxir (Spreng.) 
Farwell, forma laurentianum (Butters), stat. nov. A. angustum 
(Willd.) Presl, var. laurentianwm Butters in RHODORA, xix. 194 
(1917). 

A. FILIX-FEMINA, var. MicHauxu, forma confertum (Butters), 
comb. nov. A. angustum, forma confertum Butters, l. c. 195 (1917). 
A. angustum, var. confertum (Butters) C. S. & W. F. Lewis in 
Am. Fern. Journ. xi. 83 (1921). 

A. FILIx-FEMINA, var. MicHAUXII, forma laciniatum (Butters), 
comb. nov. A. angustum, forma laciniatum Butters, l. c. (1917). 

A. FILIX-FEMINA, Var. ASPLENIOIDES (Michx.) Farwell, forma 
subtripinnatum (Butters), comb. nov. A. asplenioides (Michx.) 
Desv., forma subtripinnatum Butters, l. c. 190 (1917). 


For once I fully agree with the late O. A. Farwell, who reduced 
Athyrium angustum and A. asplenioides to varietal rank under A. 
Filiz-femina. It seems to me quite impossible to keep apart on 
any stable morphological characters the circumboreal Athyrium 
Filix-femina and its two commoner eastern American repre- 
sentatives, A. angustum and A. asplenioides. Attempts to keep 
apart the two latter in the area where their ranges overlap are 
futile, this difficulty already noted by Weatherby in Am. Fern. 
Journ. xxvi. 134 (1936), where, considerately maintaining the 
three species recognized by Butters (A. Filix-femina, angustum 
aud asplenioides), he wrote as follows: 


But his three species are very closely related, often much alike 
in general aspect and not always easy to distinguish. The two of 
eastern North America [i. e. A. angustum and A. asplentoides, for 
A. Filiz-femina, either typical or as var. sitchense Rupr. grows on 
the Gaspé Peninsula and in Newfoundland as well as elsewhere in the 
East] are comparatively readily recognizable in the north and in the 
south where each is the exclusive occupant of its area; but in the region 
from southern New England to about the Potomac valley, where both 


390 Rhodora [NOVEMBER 


occur, they intergrade freely and in most perplexing fashion. All three 
would perhaps be better treated as geographic varieties of a single 
species, roughly analogous to those of Cystopteris fragilis. 


In this connection it is significant that the indusia shown by 
Butters, l. c., in his plate 123 and in his figs. 3C and 5A and B, 
for Athyrium Filix-femina are rather extreme. The figures of 
European indusia given by Schkuhr in his Kryptogamische 
Gewüchse, t. 58, by Hooker fil. in his British Ferns, t. 35, by 
Britten in his European Ferns (colored plate opp. p. 131) and by 
many others are reminiscent of indusia of A. angustum. Further- 
more, when he made his study of the group, or later, in checking 
identifications in 1932, Butters clearly wrote on a few sheets from 
eastern Canada and Maine such notes as ‘‘This appears to be 
true European A. Filix-femina—F. K. B.", such embarrassing 
specimens coming from the Mingan Islands and Lake St. John, 
Quebec, and from Mt. Desert Island, Maine. The Mingan 
sheet (Victorin & Rolland, no. 24,616) had been distributed as A. 
angustum, var. laurentianum. The latter anomalous plant in its 
compact and often strongly ascending rhizome and its very short 
lower pinnae is superficially so close an imitation of small Euro- 
pean plants that it is most difficult to view it as belonging to a 
separate species. In fact, when real pteridologists (to which 
highly specialized brotherhood I hardly belong) got hold of this 
form they wrote (Weatherby & Adams, List Vasc. Pl. Grand 
Manan—Contrib. Gray Herb. no. clviii. 21 (1945)) under 
Athyrium angustum: Most of the specimens seen belong to the 
form distinguished as var. LAURENTIANUM Butters. Here they 
tend to be rather strongly cespitose, the stipes short and the 
blades conspicuously narrowed toward the base, thus approx- 
imating in appearance true A. Filix-femina of Europe." I fully 
concur, and only on very plastic characters can such plants be 
kept apart from European material. In 1932 Butters evidently 
thought so. 

In Europe many scores of trivial forms have been treated as 
“varieties”, but in Hedwigia, xlv. 119-123 (1906), Goldschmidt 
treated them as subvarieties and forms. This sane course is 
approved by Bergdolt in Hegi, Ill. Fl. Mitteleur. i. 48, he saying 
“Alle diese Varietäten werden noch in zahlreiche Subvarietáten 
und Formen aufgeteilt". In view of the great diversity of forms 


1946] Wood,—Setaria Faberii in North Carolina 391 


assumed by the Lady-fern, the suggestion of the late James 
Britten (a crusty bachelor) that, in transferring the name Filiz- 
femina from the bracken to the present species, Linnaeus perhaps 
had in mind the French proverb “Souvent femme varie”, etc., 
is pertinent (or impertinent)! 


CENTAUREA MACULOSA IN INDIANA.—According to Deam in 
his “Flora of Indiana", A. A. Hansen reported this species as a 
weed near Atlanta in Hamilton Co., near the center of the state 
(Proc. Ind. Acad. Sci. 36: 251. 1927). Deam, although listing 
the plant in his ‘‘excluded species", states that it may become 
established soon if it has not already done so. Later, E. E. 
Sherff (RHoporA 48: 98. 1946) takes note of Deam's remarks, 
and states that this species seems thoroughly established in 
adjacent southern Michigan. 

September 22, 1941 I found a solitary plant on Route 20 in 
Porter Co., approximately 20 miles from the Michigan boundary. 
In a subsequent year I found another single plant on the same 
road, same county, about a mile west from the first. This region 
I looked over thoroughly in 1946, but no plants were found. 
August 27, 1946 I found several plants in the locality where the 
first was seen, apparently well established and able to compete 
successfully with any roadside weeds. 

Specimens have been sent to the Gray Herbarium.—EnpwiN 
D. Hutu, Gary, Indiana. 


SETARIA Faspert IN Norru Carouina.—In the interest of 
tracing the rather rapid spread of the Asiatic grass, Setaria Faberi? 
Herrm., in North America, it may be worth while to record this 
species for the first time in North Carolina. On August 5, 1946, 
while driving northwestward across the piedmont of North 
Carolina, I happened to glimpse the characteristic panicles of S. 
Faberii along Route 54 at Cane Creek in Orange County. Stop- 
ping to collect material, I found many scattered plants of the 
grass in a weedy roadside field with abundant Lespedeza stipu- 
lacea, the latter presumably planted intentionally and the former 
possibly introduced at that time. 


392 Rhodora [NOVEMBER 


Dr. H. L. Blomquist, to whom I sent material of this collection, 
upon examination of the specimens recalled collecting a robust 
Setaria, the arching panicles of which had attracted his attention, 
along U. S. Route 70 at Cherry Point in Craven County on Au- 
gust 23, 1945. Since at that time he was busy with other matters, 
the specimens had been stored away for future study. Examina- 
tion proves this material also to represent S. Faberii. 

At least these two collections, 150 miles apart, represent the 
grass in North Carolina. Whether or not it is permanently 
established in the state remains to be seen. Dr. Blomquist 
writes me that he did not see the species at Cherry Point in the 
summer of 1946. However, if it behaves in North Carolina in 
the same way as it has in western Virginia and in southeastern 
Pennsylvania and adjacent New Jersey it will soon become a 
well-established weed in recently disturbed soils. As H. A. 
Allard pointed out in reporting the grass from northern Virginia,! 
it does not seem to be able to compete with other species in 
ground which is not repeatedly disturbed, becoming then dwarf 
or disappearing altogether. In favorable places, however, as in 
rich bottomland corn fields, the plants may often reach a height 
of more than six feet. 

The Craven County collection, Blomquist 13722, and that from 
Orange County, Wood 6532, are represented at the Gray Her- 
barium, the U. S. National Herbarium, the University of Penn- 
sylvania and Duke University.—Cannorr E. Woop, JR., Gray 
Herbarium. 


! Va. Journ. of Science 2: 119. 1941. 


Volume 48, no. 574, containing pages 265—328, was issued 19 October, 1946. 


DEC 28 1946 


Dodora 


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Vol. 48. December, 1946, No. 576. 
CONTENTS: 

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No. IV. The Myrtaceous Genus Syzygium Gaertner in Borneo, by E. D. 
Merrill and L. M. Perry. 68 pp. 1939. $1.50. 


No. V. The Old World Species of the Celastraceous Genus Microtropis 
Wallich, by E. D. Merrill and F. L. Freeman. 40 pp. 1940. $1.00. 


Gray Herbarium of Harvard University, Cambridge 38, Mass. 


Rhodora 


JOURNAL OF 


THE NEW ENGLAND BOTANICAL CLUB 


Vol. 48. December, 1946. No. 576. 


THE GENUS LIATRIS 
L. O. GAISER 
(Continued from page 383) 


SERIES X. SQvuannosaxk. Plants of intermediate height, 3—6 
dm. tall, with from few racemiform to more branched cymiform 
inflorescence-stalks; heads large, 25-40-flowered, broadly cylin- 
drical to isodiametric, often with foliaceous phyllaries loosely 
erect and out-spreading, becoming squarrose-tipped in the 
majority of the varieties; achenes 5-6 mm. long. 32. L. squarrosa. 

From Delaware to Tennessee, south to the Gulf of Mexico and 
west of the Mississippi through the central and western plains 
states to the Kansas-Colorado border and northward into South 
Dakota. 

32. LIATRIS sqQUARROSA (L.) Michx. Plants 3-6 dm. high 
from a rounded corm up to 4 cm. in diameter, with several to 
numerous stems, glabrous, soft-pubescent or hairy; leaves linear 
and rigid, punctate, glabrous or hirsute the radical sometimes 
very long, 15-25 cm. long, 4-7 mm. wide, the cauline 10-15 cm. 
long; inflorescence with one to few heads and raceme-like, or 
with many heads, branched and paniculate: heads almost iso- 
diametric or elongate, 1.5-3 cm. long, 25-40-flowered (terminal 
ones frequently greatly exceeding the rest, up to 60-flowered), 
varying in width according to the spread of the phyllaries, sessile 
or on short pedicels; phyllaries foliaceous, glabrous or hirsute, 
ciliate-membranous or callose-margined, the inner in all varieties 
narrowly linear with acute tips, 1.5-2 cm. long and 1-2 mm. 
wide; outer ones elongate, triangular-lanceolate, often like re- 
duced leaves, ciliate-margined; middle ones similar, acuminate 
and spreading, or broader, more closely appressed basally, 
mucronate and squarrose at the tip; corolla phlox-purple, 9-15 
mm. long; inner surface of corolla-lobes conspicuously hairy; 
outer flowers of the head tending to be bent outwards approxi- 
mately at mid-corolla length, thus giving the heads a broad, flat 


394 Rhodora [DECEMBER 


top; stigmas sometimes white; achene 5-6 mm. long; pappus 
7-12 cm. long.—Fl. Bor.-Am. ii. 92 (1803); Willd. Sp. Pl. iii. 1634 
(1803); Pursh, Fl. Am. Sept. ii. 509 (1814); Torr. & Gray, FI. 
N. Am. ii. 68 (1841), excl. var. 3; Gray, Synopt. Fl. i?. 109 (1884), 
excl. var. intermedia. Serratula squarrosa L. Sp. Pl. 818 (1753). 
Pteronia caroliniana Walt. Fl. Carol. 202 (1788). Laciniaria 
squarrosa (L.) Hill, Veg. Syst. iv. 49, t. 46, fig. 1 (1762). 


KEY To VARIETIES 


a. Stem, leaves and phyllaries with varying degrees of pubescence. . . . b. 
b. Phyllaries long... .c. 

c. Outer phyllaries acuminate, ciliate-margined, squarrose 
or spreading and leafy, middle and inner ones mem- 
branous-margined and squarrose or merely spreading; 

lants hispidulous with short, whitish hairs....d. 
d. Plants stout and much-branched, with numerous large, 
almost isodiametric, thick heads; phyllaries mostly 


SQUAITOBB. . linh rernm dua nin a var. typica. 
d. Plants of one or a few slender stems, bearing few, 
longer, more cylindrical heads.................. var. gracilenta. 


b. Phyllaries short... .e. 
e. Outer phyllaries acute, with or without ciliate margins, 
middle and inner ones acute, usually callose-margined 
and squarrose; heads cylindrical and variable in size 
var. alabamensis, 
e. Outer phyllaries a few, short, acute bracts, not foliaceous, 
middle ones appressed and abruptly mucronate into a 
definite cusp, squarrose and long-ciliate; heads small, 
neatly cylindrical; plants hirsute with longer tawny 
hairs; leaves narrowly linear......0....0..0.0..00 0000 var. hirsuta. 
a. Stem, leaves, and phyllaries glabrous. . . .f. 
f. Head surrounded. by spreading, non-recurved, leaf-like 
bracts exceeding the length of the head; outer phyllaries 
acuminate, erect, middle and inner ones similarly elon- 
gate, linear, non-cuspidate, erect or spreading; heads 
sessile, frequently only a terminal one developing on a 
"lg RNC M" (acc re var. compacta. 
f. Heads lacking conspicuous, long, foliaceous outer bracts; 
outer phyllaries acute, squarrose or spreading, middle 
and inner ones narrow, callose-margined, cuspidate and 
recurved; heads on short pedicels and numerous....... . var. glabrata. 


Var. typica. Plants with dense, short, white hairs covering 
the stem and frequently both or only the lower surface of the 
generally 3-8-veined leaves, though leaves sometimes are quite 
glabrous; numerous heads on peduncles of various lengths giving 
the appearance often of a much-branched and *'floribund" 
plant; outermost phyllaries leaf-like and spreading, narrowly lan- 
ceolate and hardly overlapping, frequently covered by and always 
fringed with white hairs; middle phyllaries lanceolate, pubescent 
or glabrous, sometimes membranous-margined, more frequently 
ciliate, closely imbricated, the long acuminate or mucronate tips 
(that in the outer ones decrease from one half to one quarter the 
length of the bract) sometimes becoming somewhat involute and 


1946] Gaiser,—The Genus Liatris 395 


squarrose so that the over-all measurements of a head are almost 
the dimensions of a square; heads average 1.8 x 1.6 cm., 25-40- 
flowered; corolla ca. 15 mm. long, pappus 12 mm. long.—4z atris 
squarrosa Q floribunda Torr. & Gray Fl. N. Am. ii. 68 (1841). 
Liatris squarrosa R. Sweet, Br. Fl. Gard. 2 ser. t. 44 (1823). 
Vernonia hirsutiflora Poiret, Encyc. Meth. Bot. viii. 502 (1808). 
Liatris hirsutiflora Kostel. Allg. Med. Pharm. Fl. ii. 651 (1833). 

Delaware to northern Florida, westward to Alabama, Illinois 
and Missouri.—SOUTHERN STATES. Without stated lo- 
cality: ex Michx. Herb. (NY). DELAWARE. NEWCASTLE 
Co.: Wilmington, W. M. Canby (G). DISTRICT OF COLUM- 
BIA. Without stated locality: July 29, 1893, L. F. Ward (US). 
MARYLAND. QukEN ANNEs Co.: by roadside, near Centre- 
ville, Aug. 11, 1935, H. A. Allard, 884 (G, NY). Prince 
GEonGES Co.: Brooks Sta., Aug. 2, 1885, Prof. Ward (US). 
CnHanLES Co.: Stump Neck, Aug. 6, 1942, J. M. Turpin, 475 
(US). VIRGINIA. Without stated locality: 1878, Dr. Fore- 
man (US). Farrrax Co.: in a field, n. side of Accotink Bay, 
Aug. 8, 1893, L. F. Ward (US); wooded hills of New Alexandria, 
Aug. 10, 1919, E. C. Leonard, 776 (US). Prince WirLiAM Co.: 
Manassas, Sept. 18, 1907, E. S. Steele (US). FAUQUIER Co.: 
Catletts, Aug. 1, 1908, Mrs. E. S. Steele (US). Hrnrico Co.: 
dry sandy field, Westover Hills, July 21, 1986, M. L. Fernald & 
B. Long, 6415 (G, NY, US); near Richmond, July 19, 1935, E. 
Luttrell, 5679 (G); dry open pine woods, 14 mi. w. of Elko, Aug. 
8, 1921 (G), Aug. 16, 1921 (NY) E. J. Grimes, 4195 (G, NY); 
sandy oak and pine lands, Elko, Aug. 30, 1925, E. T. Wherry & 
F. W. Pennell, 12528 (NY); Whiteoak Swamp, w. of Elko, July 
23, 1938, M. L. Fernald & B. Long, 8872 (G). PRINCE GEORGE 
Co.: s. of Petersburg, July 22, 1936, M. L. Fernald & B. Long, 
6416 (NY). CAMPBELL Co.: without stated locality, July 1893, 
A. A. Heller (G); Lynchburg, July 19, 1925, W. A. Murrill (F). 
BEpronp Co.: roadside near Huddleston, Aug. 9, 1914, Miss J. 
Fauntleroy, 653 (US), Aug. 4, 1871, A. H. Curtiss ((G, plant to 
left), NY, Q). DiNwrippniE Co.: dry, open, border of argillaceous 
woods, s. of Petersburg, July 15, 1938, M. L. Fernald & B. Long, 
8871 (G). Prince Epwarp Co.: without stated locality, Aug. 
25, 1880, J. D. Smith (US), Sept. 1885, Prof. Blair, 317 (US). 
PITTSYLVANIA Co.: Fall Creek, July 20, 1893, A. A. Heller, 1111 
(G, NY, US). GnEENSVILLE Co.: old fields, vicinity of Belfield, 
July 1, 1904, O. M. Meyncke (US); Sprouse's Corners, Aug. 17, 
1936, Bartley & Pontius, 540 (NY). MECKLENBURG Co.: 
Finchley, Aug. 20-22, 1908, W. W. Eggleston, 4010 (G, NY, US). 
WEST VIRGINIA. CABELL Co.: Roland Park, near Hunting- 
ton, July 23, 1937, F. A. Gilbert, 988 (G, NY); dry open field near 
Huntington, Sept. 20, 1931, F. A. Gilbert, 198 (G); dry field in 
clay soil, Roland Park, near Huntington, July 24, 1940, F. A. 


396 Rhodora [DECEMBER 


Gilbert, 132 & 156 (NY, F without no.). NORTH CAROLINA. 
GRANVILLE Co.: Hester, July 23, 1937, R. K. Godfrey (G). 
CASWELL Co.: roadside, 10 mis. n. of Yanceyville, July 31, 1938, 
R. K. Godfrey, 5592 (G). Gurirronp Co.: Greensboro, Aug. 14, 
1903, Biltmore Herb., 580j (US). FomsvrH Co.: open woods, 
Winston Salem, Aug. 1, 1921, P. O. Schallert (G). MoonE 
Co.: Pinehurst, Aug. 1897, O. Katzenstein (G). Gaston Co.: 
Crowder’s Mt., July 29, 1902, Biltmore Herb., 580h (US). CLEVE- 
LAND Co.: King's Mt., Aug. 27-30, 1884, J. K. Small (NY): 
slopes of King's Mt., July 1896, J. K. Small (NY). Scornanp 
Co.: sandhill, 12 mis. north of Laurinburg, July 14, 1938, R. K. 
Godfrey, 5040 (G). New Hanover Co.: sand ridge at Carolina 
Beach, Aug. 7, 1938, R. K. Godfrey, 5906 (G). SOUTH CARO- 
LINA. LkxiNGTON Co.: Columbia, Gibbes (G). GEORGETOWN 
Co.: sandy woods, 31% mis. s. of Georgetown, July 21, 1939, R. K. 
Godfrey & R. M. Tryon, 794 (G, NY). Aiken Co.: sand hills, 
Aiken, July 17, 1900, Biltmore Herb. 580d (US) and July 22, 
1905, Biltmore Herb., 580p (US). CHARLESTON Co.: sandy place, 
McClellanville, Sept. 2, 1940, P. O. Schallert (G). BEAUFORT 
Co.: Beaufort district, 1883, J. H. Mellichamp (US). GEOR- 
GIA. Watxker Co.: dry soil, open woodlands, Chickamauga 
Park, July 14, 1899, Biltmore Herb., 580c (US). Biss Co.: dry 
soil, Macon, July 31, 1901, Biltmore Herb., 580g (US). Ricn- 
MOND Co.: dry oak woods, Augusta, July 31, 1898, A. Cuthbert 
(NY, US, F). FLORIDA. GanpspEn Co.: pinelands, River 
Junction, Aug. 31, 1901, Biltmore Herb., 580f (US). OHIO. 
Without stated locality: 1834, Riddell (G, NY). Erte Co.: 
cemetery, Castalia, July 21, 1895, E. L. Moseley (G, US). Lucas 
Co.: oak openings, n. e. of Neapolis, Aug. 26, 1927, E. L. Moseley 
(US). Apams Co.: without stated locality, July 27, 1940, D. M. 
Brown (NY). INDIANA. CrankE Co.: dry open knobs, 
July 30, 1909, C. C. Deam, 5408 (NY). KENTUCKY. MEADE 
Co.: near Brandenburg, Aug. 9, 1928, W. A. Anderson (G). 
CaLLoway Co.: dry field, between Murray and New Concord, 
July 20, 1937, L. B. Smith & A. R. Hodgdon, 4089 (G, NY, US). 
TENNESSEE. Without stated locality: 1870, G. R. Vasey 
(US). GRAINGER Co.: Clinch Mt., Aug. 12, 1880, J. D. Smith 
(US). CnmukATHAM Co.: dry shale banks, Pegram, Aug. 20, 1940, 
H. K. Svenson, 10607 (B). Knox Co.: open ground, near 
Knoxville, July 12, 1900, A. Ruth, 41 (G); dry sandy soil, Knox- 
ville, June 1896, A. Ruth, 37 (NY); sandy ground, Knoxville, 
June 1897, A. Ruth, 3772 (NY); mt. sides, Smoky Mt. Range, 
Sept. 1897, A. Ruth, 3773 (NY); Knoxville, June 1898, A. Ruth, 
659 (US); dry ground, Aug. 1898, A. Ruth, 676 (NY); near 
Young's High School, Knoxville, Aug. 2, 1934, M. B. Wilson, 
2968 (NY). Roane Co.: Harriman, Aug. 18, 1909, Biltmore 
Herb., 580i (US). Grunpy Co.: near Monteagle, Sept. 23, 


1946] Gaiser,—The Genus Liatris 397 


1933, E. J. Alexander, T. H. Everett, S. D. Pearson (NY). ALA- 
BAMA. Co. undetermined: dry sandy pine ridges (two plants 
to right), Sept. 1878, C. Mohr (US). Barsour Co.: dry 
oakwoods-border, 8 mis. s. of Eufaula, Aug. 11, 1927, K. M. 
Wiegand & W. E. Manning, 3177 (G). Escampra Co.: high 
pineland, Atmore, July 11, 1934, O. Blanton, 193 (US). MOBILE 
Co.: pine barrens, w. of Spring Hill, Aug. 1918, E. W. Graves, 
989 (G); pine barrens, July 1918, E. W. Graves, 558b (US). 
ILLINOIS. Co. undetermined: south Illinois, G. Vasey, 1210 
(G). RicnuraNp Co.: Olney, July 29, 1914, R. Ridgway, 91 (G); 
near Olney, Sept. 6, 1914, R. Ridgway (G). Marion Co.: 
without stated locality, M. S. Bebb (Q). Perry Co.: Du 
Quoin, Aug. 4, 1893, H. Eggert (NY, US); prairies, Aug. 4, 1893, 
H. Eggert (US). MISSOURI. Sropparp Co.: Bernie, Aug. 2, 
1895, B. F. Bush, 235 (NY). 

Var. gracilenta, var. nov., molliter pubescens vel rarius fere 
glabra gracilis; caulibus singulis vel paucis raro ramosis capitula 
sessilia vel subsessilia plerumque 1-5 gerentibus; foliis plerumque 
longis angustisque, radicalibus ad 25 cm. longis 4 mm. latis; 
capitulis elongatis cylindricis plerumque 1.9 cm. longis 1.2 cm. 
latis ca. 25-floris; phyllariis longe lanceolato-acuminatis apice 
laxe patentibus nec recurvatis, exterioribus 1-1.5 cm. longis 
2 mm. latis, medianis 2 cm. longis basi apicis subspathulati 4—5 
mm. latis; corollae tubo ca. 15 mm. longo; pappo 12 mm. longo.— 
Verisimiliter L. squarrosa sensu Ell. Sk. ii. 282 (1822?). 

Tyre from Ocean Springs, Jackson Co., Mississippi, Aug. 11, 
1895, J. S. Skehan (G). 

Mostly from south Georgia, Florida, Alabama, Mississippi 
and Louisiana.—Without stated locality: Drummond, 172 
“A. & L.” (G). VIRGINIA. Beprorp Co.: without stated lo- 
cality: Aug. 4, 1871, A. H. Curtiss (plant to right) (G). NORTH 
CAROLINA. Mapison Co.: Marshall, French Broad River, 
Aug. 7, 1880, J. D. Smith (US); Hot Springs, Aug. 13, 1888, B. 
W. Evermann (US, 617885 and 310560). Buncomse Co.: dry 
soil, Biltmore, July 30, 1897, Biltmore Herb., 580a (G, NY, US 
332424 and 959080); Biltmore, Aug. 1894, Biltmore Herb., 580 
(US, 959081); Cedar Cliff Mt., Aug. 17, 1904, Biltmore Herb. 
580m (US). GEORGIA. Watker Co.: dry soil, open wood- 
lands, Chickamauga Park, July 14, 1899, Biltmore Herb., 580c 
(US). FLovp Co.: Lavender Mt., Rome, Aug. Herb. Chapman 
(US). GwiNNETT Co.: Banks of Yellow R., July 20, 1893, J. K. 
Small (G). FLORIDA. Without stated locality: Chapman 
(NY, US). WasnuiNaTON Co.: dry rolling pine barrens, near 
Chipley, July 28, 1901, A. H. Curtiss 6849 (G, NY, US, Q). 
KENTUCKY. Co. undetermined: barrens of Kentucky, July 


1 Though probably from Covington, La. (see footnote under discussion of this 
species). 


398 Rhodora [DECEMBER 


1837, C. W. Short (G). ALABAMA. Co. undetermined: dry 
sandy pine ridges (one plant to left), Sept. 1878, C. Mohr (G). 
Co. undetermined: dry sandy pine ridges (one plant to left), 
Sept. 1878, C. Mohr (US). WasniNGTON Co.: low pine barrens 
(one plant to right) June 15, 1895, C. Mohr (US). MISSIS- 
SIPPI. Corran Co.: 2 mis. s. of Hazelhurst, July 14, 1925, F. 
Cook (US). Jackson Co.: Ocean Springs, Aug. 11, 1895, J. 
Skehan 1358 (G, type); Ocean Springs, July 27, 1896, C. L. 
Pollard, 1082 (US); Biloxi, Sept. 4, 1900, S. M. Tracy, 487 (NY). 
Harrison Co.: sandy pineland, 1 mi. e. of Mississippi City, Aug. 
27, 1912, F. W. Pennell, 4347 (NY, US); grassy pine barrens, near 
Mississippi City, Sept. 15, 1885, J. D. Smith 424 (US). MIS- 
SOURI. Buruer Co.: Malden, Aug. 21, 1894, B. F. Bush 386 
(G, NY). GnEENE Co.: dry rocky woods, Aug. 6, 1895, J. W. 
Blankinship (G). LOUISIANA. Without stated locality: ex 
Herb. C. W. Short, Mr. Steinhauer (G). Narcurrocuks Co.: 
Natchez, C. Wright (G). RarripEs Co.: pinelands, Sept. 10, 
1900, Biltmore Herb., 580e (two outermost plants) (US). TANGI- 
PAHOA Co.: dry soil, s. of Hammond, June 20, 1938, D. S. Correll 
& H. B. Correll, 9255 (G, NY). Sr. Tammany Co.: vicinity of 
Covington, Aug. 20, 1920, Bro. G. Arséne, 12614 (US); vicinity 
of Covington, July 1919, Bro. G. Arséne, 11665 (US); Covington, 
1832, Drummond (NY). TEXAS. Jackson Co.: 4 mis. n. of 
Edna, July 1935, J. A. Drushel, 10339 (NY). NeEwron Co.: 
without stated locality, July 23, 1939, B. C. Tharp (G). 

Var. alabamensis (Alex.), comb. nov. Plants that appear to 
be intermediate between varieties gracilenta and glabrata, with 
stems and leaves and outer phyllaries glabrous or softly pubes- 
cent; outer phyllaries spreading or slightly squarrose, triangular- 
lanceolate-acuminate 5-8 mm. long and 3-5 mm. wide and 
ciliate-margined; middle phyllaries glabrous, more sharply 
pointed and cellose-margined, 1-1.5 cm. long and 5 mm. wide; 
heads from 1.5-2 cm. long and 1-1.5 cm. wide, of about 30 
flowers; corolla 10-12 mm. long; pappus ca. 1 cm. long.—La- 
ciniaria squarrosa alabamensis Alex. ex Small Man. S. E. FI. 
1333 (1933) without indication of status. 

Alabama, Mississippi, Louisiana and eastern Texas.—ALA- 
BAMA. Harz Co.: dry chalk prairies, 1⁄4 mi. w. n. w. of Rose- 
mary, Aug. 23, 1934, R. M. Harper, 3252 (G, NY, US); chalk 
prairies, 5 mis. s. of Newbern, Aug. 1, 1938, H. K. Svenson & R. 
M. Harper 9801 (G). LEE Co.: Auburn, Aug. 11, 1897, F. S. 
Earle & C. F. Baker, 1344 (NY, type). EscAMBrIA Co.: high 
pineland, Atmore, July 11, 1934, O. Blanton, 183 (G). MISSIS- 
SIPPI. OxrrBBEHA Co.: Agric. College, 1896, C. L. Pollard, 
1318 (G, NY (US, plant to left)). Jackson Co.: Ocean Springs, 
July 27, 1896, C. L. Pollard, 1082 (NY, US); Sept. 1, 1891, A. B. 
Seymour, 193 (G); Biloxi, S. M. Tracy, July 31, 1900, 4334 (G), 


1946) Gaiser,— The Genus Liatris 399 


Aug. 31, 1900, 9734 (US); Aug. 23, 1898, 4335 (NY, US). LOUI- 
SIANA. Without stated locality: ex Herb. G. Thurber, S. T. 
Olney (G). RaPrrpEs Co.: pinelands (centre plant), Sept. 10, 
1900, Biltmore Herb., 580e (US). Jerrerson Davis Co.: low 
prairies, Welsh, Sept. 10, 1915, E. J. Palmer, 8484 (US). TEXAS. 
Pork Co.: dry upland woods, Livingston, Sept. 19, 1917, E. J. 
Palmer, 12802 (US). WarkreR Co.: without stated locality, 
Aug. 1, 1937, B. C. Tharp (G). Brazos Co.: Bryan, Aug. 17, 
1903, Biltmore Herb., 580i (US); College Station, 1888, Herb. 
Boston, Soc. Nat. Hist., Mr. P (G). WasniNGTON Co.: without 
stated locality, Aug. 1, 1938, Miss E. Brackett (G). BASTROP 
Co.: clay banks, Bestrop, 1930, H. H. Duval, 24 (G); without 
stated locality, Aug. 22, 1941, B. C. Tharp, 188 (G). Harris 
Co.: Houston, 1842, Lindheimer (G). DrWirr Co.: without 
stated locality, July 5, 1942, B. C. Tharp (G); western DeWitt, 
July 25, 1941, Miss M. Riedel (G).  FAvETTE Co.: near Schulen- 
berg, Aug. 2, 1935, H. B. Parks, 14559 (G). GarvEsTON Co.: 
Kemah, Sept. 6, 1926, G. L. Fisher, 261 (US). 

Var. hirsuta Rydb. Plants with hirsute stems and hirsute, 
narrow, 1-3-veined leaves and long-ciliate-margined phyllaries; 
few outermost phyllaries may be narrowly lanceolate but most 
are broadly oblong, narrowing abruptly into long cusp-like tips 
that alone are strongly reflexed giving the head of ca. 20 flowers 
a more compact, narrow, cylindrical appearance, averaging 1.3 
x l em.; outer phyllaries 0.7-1 cm. long and 4-5 mm. wide; 
middle ones 1-1.3 cm. long and 3-4 mm. wide; corolla ca. 1 cm. 
long; pappus 8 mm. long.— Liatris hirsuta Rydb. Brittonia i. 98 
(1931). L. squarrosa var. hirsuta Rydb. l. c. 99 in synon.; here 
accepted and validated. L. squarrosa hirsuta (Rydb.) F. C. 
Gates, Trans. Kans. Acad. Sci. xlii. 138 (1940), without indica- 
tion of status. 

Chiefly west of the Mississippi in the Ozarks of Missouri 
and the western mountainous regions of Arkansas; also in eastern 
Kansas and Oklahoma.—KENTUCKY. Co. undetermined: 
barrens of Kentucky, July 1871, W. M. Canby (G). MISSIS- 
SIPPI. Horwrs Co.: without stated locality, Aug. 10, 1893, 
S. M. Tracy (NY). IOWA. Pato Arro Co.: high prairie hill- 
top, s. of Virgin Lake, Highland Twsp., Sect. 30, Sept. 16, 1937, 
Miss A. Hayden, 10533 (G, NY). MISSOURI. Mercer Co.: 
dry open woods, 5 mis. s. w. of Linesville, July 5, 1933, E. J. 
Palmer & J. A. Steyermark, 41312 (NY, US). Anar Co.: dry 
banks, open woods, 4 mis. n. of Kirksville, July 3, 1933, E. J. 
Palmer & J. A. Steyermark, 41174 (NY). Jacxson Co.: 
prairies, Lee's Summit, July 25, 1905, B. F. Bush, 3096a (G, NY, 
US); Little Blue, July 22, 1896, B. F. Bush, 454 (US); dry hills, 
e. of Dodson, July 26, 1896, K. K. Mackenzie (NY). Cass Co.: 
barrens, west of Belton, Aug. 4, 1902, K. K. MacKenzie, 97 (US, 


400 Rhodora [DECEMBER 


NY). Vernon Co.: dry banks, border of woods, along creek 
near Nevada, Aug. 6, 1933, E. J. Palmer & J. A. Steyermark 
42138 (US, NY). Pork Co.: vicinity of Graydon Springs, 
Sept. 7, 1912, P. C. Standley, 9909 (US); on top of cherty lime 
barren, on steep slopes along Pomme de Terre R., 2 mis. n. e. of 
Rondo, Aug. 1, 1937, J. A. Steyermark, 24097 (NY). GREENE 
Co.: vieinity of Willard, Aug. 30, 1912, P. C. Standley, 9665 
(US); dry ground, Sept. 4, 1893, B. F. Bush, 197 (NY); Spring- 
field, 1888, J. W. Blankinship (US); along Rwy., n. e. of town, 
vicinity of Springfield, Sept. 1, 1921, P. C. Standley, 8622 (G, 
US). JasPER Co.: rocky open ground, n. fork of Spring River, 
near Neck City, Sept. 21, 1923, E. J. Palmer, 23863 (NY); high 
prairies, Sarcoxie, July 15, 1914, E. J. Palmer, 6261 (US). 
Barry Co.: Purdy, Aug. 17, 1905, B. F. Bush, 3254 (G, NY); 
Shell Mound, July 11, 1927, E. J. Palmer, 3244 (NY); Hailey, 
Sept. 15, 1905, J. W. Phillips (US); rocky slopes, open woods, 
near Seligman, Oct. 24, 1925, E. J. Palmer, 29370 (Q); woods, 
Eagle Rock, Aug. 11, 1905, B. F. Bush, 3217 (G, NY, US). 
Newton Co.: along chert bluffs of Shoal Creek, near Redding’s 
Mill, July 14, 1927, E. J. Palmer, 32505 (NY, type); Joplin, 
Sept. 8, 1912, E. J. Palmer, 3836 (NY, US). TANEY Co.: rocky 
woods, Swan, Sept. 27, 1905, B. F. Bush, 3467 (G). McDoNarp 
Co.: dry ground, B. F. Bush, July 24, 1893, 200A (G), Sept. 1, 
1893, 200B (US). ARKANSAS. Co. undetermined: Red 
River, ex Torrey Herb., Dr. Pitcher (NY); n. w. Arkansas, Sept., 
F. L. Harvey, 14 (G). Carrom Co.: dry hillsides, Eureka 
Springs, Sept. 30, 1913, E. J. Palmer, 4531 (US). Purasxr Co.: 
dry ledges, open woods, w. of Pulaski Heights, Little Rock, 
Sept. 21, 1931, D. Demaree, 8219 (G, NY); near Little Rock, G. 
W. Letterman (NY). Yxrr Co.: dry soil, Mt. Nebo, Aug. 4, 
1903, Biltmore Herb., 580k (US). Garrano Co.: open dry woods, 
end of Whitington Ave., Hot Springs, Aug. 25, 1935, F. J. Scully, 
461, 462 (G); open woods, Glenwood Road, 8 mis. from Hot 
Springs, July 21, 1935, F. J. Scully, 386 (G); near Hot Springs, 
1928, R. Runyon, 1179a (NY). LOUISIANA. NATCHITOCHES 
Co.: dry sandy hills, 3 mis. s. of Coldwater, July 26, 1938, D. S. 
Correll & H. B. Correll 9811 (G, NY); sandy open woods, Oct. 9, 
1915, E. J. Palmer, 8800 (US). NEBRASKA. Co. unde-. 
termined: prairies, Aug. 4, 1853, T. E. Hayden (NY). KANSAS. 
Co. undetermined: Wolf Creek to Independence Creek, July 30, 
1849, A. Fendler (G). Atcuison Co.: prairie, 12 mis. w. of 
Atchison, July 18, 1936, Sister Jeanette, 150 (NY). RirEY Co.: 
Manhattan, Aug. 9, 1892, J. B. S. Norton (US). Dovaras Co.: 
without stated locality, Aug. 1870, L. H. Hoysradt (NY). 
MoNTGoMERY Co.: Caney, open dry banks, July 21, 1933, E. J. 
Palmer, 41795 (NY). OKLAHOMA. Co. undetermined: Sans 


1946] Gaiser,—The Genus Liatris 401 


Bois Mts., Aug. 1891, C. S. Sheldon, 304 (US); prairies,! Aug. 26, 
1853, Lieut. A. W. Whipple's Exped., J. M. Bigelow (US); Camp 
no. 4, N. W. Agency!, July 28, 1853, Bigelow (NY); Camp no. 4!, 
July 28, 1853, Bigelow, 357 (NY). Nowara Co.: on dry hill- 
top, Lenapah, Aug. 19, 1913, G. W. Stevens, 2175 (G, US). 
CHEROKEE Co.: dry rocky woodlands, Aug. 18, 1895, J. W. 
Blankinship (G, US). Le Fiore Co.: Cavanal Mt., Poteau, 
Oct. 28, 1915, E. J. Palmer, 9059 (US). PusHMaTAHA Co.: 
sandy meadow, Tuskahoma, June 20, 1937, U. T. Waterfall (NY). 

Var. compacta Torr. & Gray. Glabrous plants with shining 
narrowly linear, leaves 2-3 mm. (occasionally up to 1 em.) wide; 
heads of 20-40 flowers, generally sessile, generally borne singly 
and terminally on the stems, surrounded by long foliaceous 
bracts twice the length of the head, 5 cm. long, 2-4 mm. wide; 
phyllaries all completely glabrous, leaf-like, elongate, acuminate 
to cuspidate and spreading, but not squarrose at any time: 
outer ones 1-2 cm. long and 4 mm. wide; middle ones linear, 
1.5-2 em. long and 2-3 mm. wide; corolla ca. 1 em. long; pappus 
7-8 mm. long.—Fl. N. Am. ii. 68 (1841). Liatris compacta 
Rydb., Brittonia i. 98 (1931). 

Observed from Arkansas only.—CouNTY UNDETERMINED: Dr. 
Leavenworth (G, NY, type. Perhaps from Louisiana). PULASKI 
Co.: Little Rock, Sept. 30, 1931, D. Demaree, 8338 (US). GAR- 
LAND Co.: Hot Springs, novaculite outcrops, slopes of Hot 
Springs Mt., Oct. 9, 1925, E. J. Palmer, 29057 (NY), rocky top 
of West Mt., June 4, 1923, Palmer 23061, grassy patches on 
novaculite rubble slope, June 18, 1933, H. R. Gregg, 38 (US), 
1931, R. Runyon, 1442 (US), dry sandstone hills, July, 1880, 
F. L. Harvey, 42 (G), dry, rocky ground, slopes of West Mt., 
Oct. 15, 1925, Palmer, 29230 (NY); rocky bluff, Camp Charlton, 
Crystal Springs, June 14, 1942, Demaree, 23215 (G). Hor 
Sprina Co.: rocky bluffs, Magnet Cove, May 1, 1938, Demaree, 
17252 (G, NY). 

Var. glabrata (Rydb.), comb. nov. Plants entirely glabrous 
with rigid narrow 1—3-veined leaves and phyllaries narrower than 
in var. hirsuta, short, acuminate, or cuspidate, callose-margined, 
and recurved after flowers have opened though they may be 
quite erect before that time; outer ones triangular-lanceolate, 
0.7-1 cm. long; middle ones 1-1.5 cm. long and 4 mm. wide: 
heads trim, without the long foliaceous outermost bracts of vars. 
compacta and typica, averaging 1.3x 1 cm., of 20-30 flowers; 
corolla ca. 1 cm. long; pappus 7-8 mm. long.—Liatris glabrata 
Rydb. Brittonia i. 98 (1931). L. squarrosa glabrata (Rydb.) F. C. 


1 According to Bigelow—Report on Lieut. Whipple's Expedition (1857) p. 96, Sect. 
4, L. squarrosa was said to have been found on the prairies, August 26. From the 
map and notes the party followed the Canadian River, practically all of which lies 
in Oklahoma. 


402 Rhodora [DECEMBER 


Gates, Trans. Kans. Sci. xlii. 138 (1940) without indication of 
status. 

In the plains-states, South Dakota to Texas, west to the 
Colorado border of Kansas and east to Missouri—MISSOURI. 
GREENE Co.: dry rocky woods, Aug. 7, 1895, J. W. Blankinship 
(G); prairies, n. e. of Springfield, Aug. 21, 1912, P. C. Standley, 
9113 (US). ARKANSAS. Barron Co.: Plum Buttes on Ar- 
kansas R., near Great Bend,! 1846, Lieut. Abert's Exped. (NY). 
SOUTH DAKOTA. SrmaNLEY Co.: banks of Mo., below Fort 
Pierre, 1853, F. V. Hayden (NY). Topp Co.: high land, Rosebud 
Creek, July 2, 1896, E. J. Wallace (NY); Rosebud, E. J. Wallace 
(US). Brnnerr Co.: sandhills, near La Creek P. O., Aug. 12, 
1911, S. S. Visher, 2232 (NY). NEBRASKA. Co. unde- 
termined: Platte R., near the Forks, Herb. G. Thurber, H. Engel- 
mann (G); Platte R., Lieut. Fremont (G, NY). Knox Co.: 
Pisheville, Aug. 18, 1893, F. Clements, 2734 (G, NY, US); Fort 
Niobrara, June 25, 1888, T. E. Wilcox (NY); Niobrara Game 
Reserve, near Valentine, July 22, 1936, W. L. Tolstead, 527 (Q). 
KeryapaHa Co.: Cuba, 1908, J. S. Haller (US). Rock Co.: 
without stated locality, Aug. 26, 1890, C. Rutter (US). Brown 
Co.: Long Pine, G. D. Swezey, 52 (NY). Tuomas Co.: near 
Plummer Ford, Dismal River, Aug. 23, 1893, P. A. Rydberg, 
1505 (G, (NY, type), US)). Grant Co.: lake region, 4 mis. n. e. 
of Whitman, 1893, P. A. Rydberg, 1505 (G, NY). Custer Co.: 
Callaway, Aug. 17, 1902, J. M. Bates (G). Burrato Co.: Fort 
Kearney, Platte R., 1856, H. Engelmann (G). KANSAS. 
PorrAwATOMIE Co.: without stated locality, Sept. 1, 1892, S. 
Norton (NY). RinEv Co.: low woods, Aug. 9, 1895, J. B. 
Norton, 278 (G); prairie, July 18, 1895, J. B. Norton, 212 (US). 
Geary Co.: Fort Riley, July 20, 1892, E. E. Gayle, 542 (NY). 
ErLswonTH Co.: near Ellsworth, Aug. 2, 1930, E. J. Palmer, 
38167 (G, US). Finney Co.: 2 mis. s. e. of Garden City, 
Kansas Nat. Forest, Aug. 1912, Kellerman (US). Reno Co.: 
without stated locality, July 1891, M. A. Carlton, 290 (US). 
Kiowa Co.: Greensburg, Aug. 5, 1890, B. B. Smyth, 74 (US). 
OKLAHOMA. Co. undetermined: Sans Bois Mts., Aug. 21, 
1891, C. S. Sheldon (NY); tributaries of Washita, 1852, Marcy’s 
Exped., ex Herb. G. Thurber (G). Osace Co.: open woods, near 
Ponca, Aug. 6, 1913, G. W. Stevens, 1924 (G). Woopwarp Co.: 
sandy soil, Woodward, Aug. 21, 1931, L. F. Locke, 11 (US). 
Payne Co.: Stillwater, July 5, 1896, E. E. Bogue (G), 1918, 


! From notes of à military reconnaissance from Fort Leavenworth in Missouri to 
San Diego in California, including part of the Arkansas, Del Norte & Gila Rivers, by 
Lieut. Abert, appendix 6, p. 386, and map, this collection would come from the Bend 
of the Arkansas R., just where Walnut Creek enters it, in Barton County. 

2 The Report Fremont's Expedition, Washington, 1845, p. 90 of Catalogue of 
Plants, refers without date to “L. squarrosa var. intermedia DC. a small form of the 
plant on the Platte"— which probably refers to these specimens. 


1946] Gaiser,—The Genus Liatris 403 


F. A. Waugh, 284 (US). Dxwzx Co.: dry open ground, plains 
near Taloga, July 26, 1933, E. J. Palmer, 41957 (P). LINCOLN 
Co.: Fonts, Aug. 28, 1895, J. W. Blankinship (G, US). Okra- 
Homa Co.: sandy clay, sandstone hills, n. w. of Spencer, Aug. 6, 
1939, U. T. Waterfall, 1551 (G); sandy soil, 2 mis. w. of Nicoma 
Park, Aug. 9, 1940, U. T. Waterfall, 2343 (G). Cappo Co.: 
Anadarko, July 17, 1891, C. S. Sheldon, 168 (US); without 
stated locality, June 27, 1903, A. H. Van Fleet (US); between 
Fort Cobb & Fort Arbuckle, 1868, E. J. Palmer, 469 (US). 
Aroxa Co.: gravelly hillside, Stringtown, June 25, 1891, C. S5. 
Sheldon, 68 (US). Bryan Co.: vicinity of Durant, 1931, W. L. 
Blain, 220 (US). TEXAS. Darras Co.: sandy soil, near 
Dallas, June 1882, J. Reverchon, Herb. Curtiss, 1171a (G, US); 
Dallas, July 1874, J. Reverchon (G); sandy woods, Dallas, June 
1880, J. Reverchon, 409 (US); dry uplands, Dallas, Sept. 1877, 
J. Reverchon, 412 (US). Tarrant Co.: in sandy soil, along 
right of way, T. & P. Rwy., near Handley, July 28, 1913, A. Ruth, 
404 (US 504842); along T. & P. Rwy., near Polytechnic, July 
28, 1913, A. Ruth, 404 (US, 910626); meadows, July 1913, A. 
Ruth, 404 (US, 587685); without stated locality, May 1928, A. 
Ruth, 403 (US). WaLKER Co.: on blackland prairie, vicinity 
of Huntsville, July 9-12, 1909, R. A. Dixon, 398 (G). COLORA- 
DO. Co.undetermined: prairies on the Arkansas! below Turkey 
Creek, Sept. 1847, A. Fendler, 298 (G). Yuma Co.: Wray, 
Aug. 11, 1913, W. W. Eggleston, 15534 (G, NY), Aug. 5, 1909, 
G. E. Osterhout, 3991 (NY), July 1-4, 1919, W. W. Eggleston, 
15173 (US), July 2, 1910, H. L. Shantz (US). 


The type of Liatris squarrosa is the Linnaean specimen, à 
photograph of which was seen at the Gray Herbarium, as was also 
one of the specimen back of the reference to the Hortus Cliffor- 
tianus. These specimens represent a generally leafy plant, of a 
branched appearance, with pubescent stems and leaves and 
squarrose or spreading bracts (var. floribunda of Torr. & Gray). 
From this typical variety, occurring chiefly in Virginia and North 
and South Carolina, can be distinguished a more slender, fewer- 
and longer-leaved, generally slightly pubescent plant having 
fewer heads (1-5) with erect lanceolate rather than squarrose 
phyllaries, which predominates in the region just north of the 
Gulf of Mexico, in southern Georgia, Alabama, Mississippi and 
Louisiana, which we are calling var. gracilenta. The wholly 
glabrous plant with long, acuminate phyllaries is Torrey & 

1 From Plant. Fendl. Mem. Amer. Acad. Ser. 2, iv. 1-116 (1849), this is the place 


and time of collection of no. 298, and the route followed through September was from 
Bent's Fort, now in Otero Co., to the Raton Mts. in Las Animas Co. 


404 Rhodora [DECEMBER 


Gray’s var. compacta, later reduced by Gray to var. intermedia, 
a variety open to question. 

The more common variety of the species in the states just west 
of the Mississippi is a hirsute plant, described by Rydberg l. c. 
as Liatris hirsuta. It is generally less branching and by reason 
of its narrower leaves, less foliaceous in appearance than var. 
typica. ‘The distinctly long-ciliate-margined phyllaries, except 
for a few outermost spreading ones, are tightly appressed with 
only the long cuspidate tips recurved. Extending from Texas 
farther westward and northward in the plains states is a slender 
glabrous variety, without the conspicuous long acuminate bracts 
of var. compacta, but with squarrose callose-margined phyllaries 
that Rydberg l. c. described as Liatris glabrata. In synonymy 
with this he placed var. intermedia of Gray, Synopt. Fl. i2. 109 
(1884) in part, as to description. 

It is obvious that much confusion in this species has arisen 
from the failure properly to place Liatris intermedia as described 
by Lindley (Bot. Reg. t. 948 (1825)) from a plant brought to the 
Horticultural Society in 1824 by David Douglas and grown from 
roots collected in Canada by Mr. Goldie who also had pressed 
specimens of it. From the Journal kept by Douglas during his 
travels in North America, 1823-27 (Wm. Wesley & Son, 364 
pages), it is found that he collected in the vicinity of Amherst- 
burg and Sandwich, now in Essex county, in the southwestern 
part of the Ontario peninsula. From the examination of Lind- 
ley's plate, the plant collected by Douglas strongly resembles 
Liatris cylindracea. Especially do the typical cylindrical younger 
heads before opening of the flowers place it with L. cylindracea 
rather than L. squarrosa, which would show recurved or spreading - 
bracts at the same age. Examination of various herbaria for 
specimens collected in Ontario has not disclosed any L. squarrosa 
found in the province, though L. cylindracea has been collected 
along the shores of Lake Erie and Lake Huron north to Mani- 
toulin Island, and from the northwest shore of Lake Ontario. 
The matter was further confused when Hooker, in Flor. Bor.- 
Amer. i. 306 (1834), listed only L. scariosa, L. punctata and L. 
squarrosa, giving Canada as habitat for the last species and re- 
ferring to Mr. Goldie as the collector. In the note under L. 
squarrosa he states: "I fear the L. intermedia of the Bot. Reg. 


1946] Gaiser,—The Genus Liatris 405 


can only be considered a luxuriant state of the present. Mr. 
Goldie’s original specimens in my Herbarium do not at all differ 
from the true squarrosa; and the cultivated individual only ex- 
hibits longer flower-stalks". Macoun (Cat. Can. Plants i. 542 
(1886)) included L. squarrosa, but gave ‘‘western Ontario. 
(Gray.)" as the only reference, adding “not noticed by any of the 
late collectors" and for var. intermedia he stated "same situation 
as the type. (Gray.) Not lately collected". Though neither 
Lindley's plant nor the original herbarium specimen from 
Canada has been seen, by the plate, description and locality of 
collection, L. intermedia has been placed in the synonymy of L. 
cylindracea. The range of L. squarrosa does not include Canada. 
Though we believe six varieties of L. squarrosa can be distin- 
guished, many intermediates are found, especially where two 
varieties coincide geographically. So, in the Southern Appala- 
chian region, as in Georgia, where one finds var. typica and also 
var. gracilenta, of more southern range, there are more plants of 
5-9 heads that are clearly intermediates between the two varie- 
ties. In numerous cases specimens show combined characters of 
both varieties and some of these intergradations have been listed 
below. Sometimes, specimens of the same number in various 
herbaria suggest two different varieties, as R. M. Harper, no. 273, 
from Rocky Face Mt., Whitfield Co., Ga., July 21, 1900, of which 
those at Gray and the New York Botanical Garden resemble var. 
typica and that at Washington, more nearly var. gracilenta. 


Intermediates between L. squarrosa var. typica and var. 
gracilenta. NORTH CAROLINA. IREDELL Co.: Statesville, 
July, 1878, M. E. Hyams (US). GEORGIA: WnirFIELD Co.: 
dry sandy field, s. end of Rocky Face Mt., July 21, 1900, R. M. 
Harper, 273 (G, NY, US). Harr Co.: dry oak woods, 4 mis. s. 
of Hartwell, Aug. 19, 1927, K. M. Wiegand & W. E. Manning, 
3178 (G). GwiNNETT Co.: banks of Yellow River, July 20, 
1893, J. K. Small (NY, US). CnarrAHoocHEE Co.: Chatta- 
hoochee River, Fort Benning Military Reserve, July 7, 1926, 
J. O. Andes, 2910 (NY). Sumrer Co.: dry pine barrens s. of 
Leslie, July 13, 1901, R. M. Harper, 1066 (G, NY, US). FLOR- 
IDA. Without stated locality: Chapman ez Torrey Herb. (NY), 
Chapman ex. Herb. Thurber (G). LrsERTY Co.: damp woods, 
on the road from Quincy to Aspalaga and Chattahoochee, Chap- 
man. ex. Herb. J. A. Lowell (G). KENTUCKY. Without 
stated locality: 1842, C. W. Short (NY). Co. undetermined: 


406 Rhodora [DECEMBER 


Green River region, Aug. 21, 1895, J. N. Rose (US); WHITLEY 
Co.: cliffs of the waterfall, Cumberland Falls, Aug. 21, 1883, J. 
D. Smith (US). ALABAMA.  Cray Co.: Elders, July 29, 1896, 
C. Mohr (US). WasnuiNGTON Co.: low pine barrens, June 15, 
1895, C. Mohr (one plant to left) (US). Moste Co.: in woods, 
Spring Hill, Aug. 20, 1897, B. F. Bush, 155 (NY, US). MISSIS- 
SIPPI. OxkrrBBEHA Co.: Agricultural College, Aug. 11-17, 
1896, C. L. Pollard, 1318 (plant to right, US). 

In Mississippi and Louisiana, var. gracilenta is the dominant 
variety (though again there are some intermediates) and gener- 
ally it is softly pubescent. However, sometimes it is an almost 
glabrous, tall slender plant, e. g. C. L. Pollard, no. 1082, from 
Ocean Springs, Jackson Co., Miss. (US), or Arséne, no. 12614, 
from the vicinity of Covington, St. Tammany Parish, La. (US). 
Yet for the distinctly glabrous, long-foliaceous-bracted plant de- 
scribed by Torrey and Gray l. c. as var. compacta from a plant 
collected by Leavenworth, in “Arkansas or Louisiana" (G, NY), 
I have found no exact match. The description, “glabrous; 
leaves crowded, very narrow, heads several, closely sessile, ap- 
proximate”, excludes these Louisiana and Mississippi specimens, 
since most of them are not wholly glabrous and have heads quite 
distant, even sometimes pedunculate as seen also in the Drum- 
mond specimens (G, NY), and referred to by Hooker! (1835). 
But in a limited range, in Arkansas, I have found glabrous speci- 
mens varying from the compacta type only in having slightly 
wider leaves and phyllaries. As these seem very distinctive and 
nearest to the Leavenworth specimens, I have included them with 
it in var. compacta; the Louisiana and Mississippi plants seem 
more adequately placed with var. gracilenta. "The other glabrous 
variety, with shorter, recurved phyllaries, is the only prevalent 
one in the western plains-states, and in going from Louisiana 
westward through Texas, one finds the glabrous form with the 
shorter phyllaries becoming increasingly abundant. In south- 
eastern Texas, where var. glabrata finds its eastern limit, one often 
sees specimens approaching it in the character of the phyllaries, 
yet perhaps with softly pubescent stem and sometimes much- 
branched inflorescence as in E. J. Palmer, no. 12802, from 

1 W. J. Hooker—Companion to the Botanical Magazine. Nov. 1835, p. 95, “no. 
410 Liatris squarrosa Willd.—Covington. $ floribus longius pedicellatis involucri 


squamis appressis vix squarrosis, . . . This is a very remarkable variety but I do 
not think it can be separated from L, squarrosa”. 


1946] Gaiser,—The Genus Liatris 407 


Livingston, Polk Co. (US), or G. L. Fisher, no. 261, from Kemah, 
Galveston Co., Texas (US). These seem understandable as 
intermediates between the western var. glabrata and the southern 
var. gracilenta where the two varieties have met. In specimens 
from the Mississippi and Alabama region, varying degrees of 
pubescence are found in combination with a number of modifica- 
tions of spreading or recurved, callose-margined phyllaries and 
in intermediacy in the size of the heads. Such phases we believe 
were described by Alexander as Laciniaria squarrosa alabamensis. 
Though it seems rather difficult to limit this variety with such 
varying intermediate expressions, we are retaining it here. 
Further to the north and west, in Kansas and Oklahoma, some 
specimens are again intermediate between var. glabrata and var. 
compacta, as can be seen in the specimens: M. A. Carleton no. 290, 
without locality, Reno Co., Kans. (US), or U. T. Waterfall, no. 
107 from poor sandy soil, 4% mi. e. of Yale, Payne Co., Okla. (G) 
or G. M. Merrill no. 886, Buffalo Springs, Platt National Park, 
Murray Co., Okla. (NY). Asingle specimen, A. Ruth, no. 739, 
in sandy woods, Tarrant Co., Texas (NY), shows a combination 
of the tawny hairs of var. hirsuta with general habit and involu- 
eral characters of var. gracilenta. 

Collections from Kentucky and Tennessee have shown, grow- 
ing in close populations, quite an assortment of variations. Some 
less hirsute plants compare well with var. typica, others resemble 
that variety though the phyllaries are shorter and more acute 
(as those listed for Knox Co., Tenn.). The latter condition was 
also noted in the few specimens from farther north in Ohio and 
Illinois and these are sometimes incorrectly found in herbaria 
under the name var. intermedia. Some Tennessee specimens 
appear to be intermediates between var. typica and var. gracilenta 
` while others appear to have blended characters of var. typica and 
var. hirsuta, abundant to the west across the Mississippi. In 
Iowa, which is just north of the range of this latter variety, the 
specimens generally showed very scant pubescence and approach 
the intergrades between var. hirsuta and var. glabrata, the latter 
predominating in the northern and western plains states (S. 
Dakota, Nebraska, Kansas). Thus there seem to be definite 
geographic entities which intergrade where their ranges overlap 
and some of these various probable intergradations have been 
listed below. 


408 Rhodora [DECEMBER 


Intermediates between L. squarrosa var. hirsuta and var. 
glabrata. IOWA. Co. undetermined: central Iowa, H. H. 
Babcock (US). DickiNsoN Co.: upland prairie, n. of Milford, 
Aug. 2, 1912, B. Shimek (G, US). Osckora Co.: prairies, near 
Sibley, Sept. 18, 1879, E. L. Greene (G); prairie, n. of Ocheyedan 
mound, Aug. 1912, B. Shimek (G, US). Piymoutx Co.: Akron, 
Sept. 1909, Mrs. E. Bredall (US). Decatur Co.: prairies, Aug. 
25, 1897, T. J. Fitzpatrick (NY, US). MISSOURI. Lara- 
YETTE Co.: dry prairies, Emma, Aug. 10, 1897, C. H. Demetrio, 
116 (G). GREENE Co.: dry woods, Aug. 7, 1895, J. W. Blankin- 
ship (G). ARKANSAS. Putasxr Co.: roadside by Quarry 
Country Club, Pulaski Heights, Little Rock, Sept. 23, 1931, 
D. Demaree, 8219 (US). Yer Co.: rocky flats, Mount Nebo 
State Park, D. Demaree, Aug. 16, 1939 (no. 20060), Aug. 30, 1939 
(no. 20590) (G). KANSAS. Without stated locality: 1876 
E. A. Popinoe (US). OKLAHOMA. Cmnmxrx Co.: Sapulpa, 
July 30, 1894, B. F. Bush, 215 (G, NY, US). Cappo Co.: 
False Washita, between Fort Cobb & Fort Arbuckle, 1868, E. 
Palmer, 454 (US). Le Fiore Co.: in dry creek-channel, base of 
Rich Mt., Page, Sept. 8, 1913, G. W. Stevens, 2691 (G, NY, US); 
grassy rwy. right of way, Page, July 11, 1914, O. W. Blakeley, 
1493 (G). PrrrsBuRG Co.: McAlester, July 8, 1894, C. S. 
Newhall (G). 

It would seem as though from approximately the Mississippi 
westward the indifferently somewhat glabrous or hirsute typical 
L. squarrosa became divided into two strains, one distinctly 
hirsute (var. hirsuta), the other distinctly glabrous (var. glabrata, 
and var. compacta). The first two of these are easily compre- 
hended as modifications of the squarrose-bracted var. typica to 
the east. As to the third, var. compacta, the transition from the 
condition in var. typica to open, erect, somewhat spreading bracts 
as in var. gracilenta, growing somewhat farther south, passed on 
into the Mississippi-Louisiana region where it stands out against 
the squarrose var. glabrata. It may be merely its persistence . 
further westward in Arkansas that we recognize as var. compacta. 

While the species L. squarrosa shows a range of varieties and 
intermediates between those varieties, it is nevertheless clearly 
distinguishable from L. cylindracea Michx., which is most closely 
related to it. 'The phyllaries of the latter are obtuse to mu- 
cronate-tipped and though the outer ones may spread slightly, 
are mostly appressed, not squarrose. Variation in the shape of 
the phyllaries from rounded at the apex, as described by Michaux, 
to more lanceolate ones, was probably responsible for the proposal 


1946] Gaiser,—The Genus Liatris 409 


of Lindley's L. intermedia, but there is complete gradation from 
one state to the other within L. cylindracea. 

That L. squarrosa hybridizes with other species is known from 
the reported finding of X L. Ridgway Standl. (see no. 6) along 
with plants of it and of L. pycnostachya of the Pycnostachyae 
series belonging to the other section Suprago. A second hybrid 
is here described, likewise with a species in the other section, but 
of the Scariosae series. 


x LiaTRIS creditonensis, hybr. nov. (L. ligulistylis X squarrosa 
var. glabrata), planta fruticem simulans; caulibus 12 vel plures 
e cormo ca. 2.5 diametro ca. 50 cm. altis basem versus glabris 
superne sparse pubescentibus, inflorescentiae rachi pilis albis 
adpressis strigosa; foliis glabris, superioribus ad bracteas capitula 
inferiora duplo superantibus capitula superiora aequantes re- 
ductis; inflorescentia racemosa; capitulis 40—50-floris subtur- 
binatis 1.6 cm. longis 1.8 cm. latis, phyllariis laxe erectis oblongis 
apice subarcuato acuto fere omnino herbaceis marginibus scario- 
sis angustis leviter revolutis, exterioribus ca. 5 mm. longis 3 mm. 
latis, medianis interioribusque ca. 1 cm. longis 5 mm. latis sub- 
acutis; corollis ca. 9 mm. longis pallide purpureis, tubo intus 
glabro, lobis pagina interiore conspicue pilosis; stigmatibus albis; 
achaeniis 4-5 mm. longis; pappo ca. 6 mm. longo manifeste 
barbellato.— The TYPE appeared in the garden at Crediton, 
Huron Co., Ontario, in the summer of 1931 (G). 


A single plant of this hybrid appeared in a plot among plants 
of L. squarrosa var. glabrata, seeds of which had been collected 
Oct. 4, 1927, from a high cliff, n. w. of Royal, Antelope Co., 
Nebraska by Dr. Wernicke, and were received through Prof. 
R. A. Harper. Growing next at the time were plants of L. 
ligulistylis collected near Erskine, Red Lake Co., Minn. by the 
writer in 1929. The plant resembled the former species in pro- 
ducing a number of stems with a raceme of ca. 20 uniformly sized 
heads (without an unusually large terminal one), in the glabrous 
leaves and their gradual diminution so that they projected as 
bracts subtending the heads through at least two-thirds of the 
length of the inflorescence and in the pilosity on the inside of the 
corolla-lobes. It resembled L. ligulistylis in the pubescence of 
the rachis of the inflorescence, in the many-flowered (40-50) 
heads, in the involucre of loose erect phyllaries and in the barbel- 
late rather than plumose pappus. It showed intermediate char- 
acters in the shape and width of the leaves, in the acute but 


410 Rhodora [DECEMBER 


broader phyllaries that were almost wholly herbaceous except for 
the narrow margins but on the whole was a true intermediate 
between the two species. 

As the plant was under cultivation, it was possible to collect 
seeds and 15 varied seedlings gave proof of their viability. Of 
the progeny two plants appeared almost identical with the parent 
and seven others varied only slightly in the width of leaves or 
phyllaries from it. Of the other seedlings two were very like the 
parental L. squarrosa var. glabrata: three were more like it than 
L. ligulistylis in the pointed phyllaries and one other in having 
narrowly linear leaves. All of the seedlings had hairs within the 
corolla-lobes, thus showing a much stronger relationship to L. 
squarrosa var. glabrata. 


EXCLUDED NAMES AND SPECIES 


Liatris amplexicaulis Raf. New Fl. N. Amer. iv. 76 (1838)—prob- 
ably Trilisa odoratissima. 

Liatris baicalensis Adams in Mem. Soc. Nat. Mosc. v. 115 (1817) 
—obviously not a Liatris. 

Liatris bellidifolia Michx. Fl. Bor.-Amer. ii. 92 (1803) = Carphe- 
phorus bellidifolius. 

Liatris brasiliensis Gardn. Hook. London Jour. Bot. v. 461 (1846) 
= Leptoclinium brasiliense Benth. & Hook. Gen. Plant. ii. 244 
(1873). 

Liatris cordata D. Don ex Royle Illust. Bot. Himal. 247 (1835) 
—nomen nudum. 

Liatris cordata Royle ex C. B. Clarke, Compos. Indicae, 247 (1876) 
= Ainsliaea aptera. 

Liatris corymbosa Nutt. Gen. Amer. ii. 132 (1818) = Carphe- 
phorus corymbosus. 

Liatris fruticosa Nutt. Amer. Jour. Sci. v. 299 (1822) = Garberia 
fruticosa. 

Liatris lanata Spreng. ex. DC. Prodr. vi. 540 (1837) obviously 
not a Liatris. 

Liatris latifolia Don, Prodr. Fl. Nep. 169 (1825) obviously not a 
Liatris. 

Liatris lobelioides Wall. ex. C. B. Clarke, Compos. Indicae, 246 
(1876) = Ainsliaea pteropoda. 

Liatris odoratissima Willd. Sp. Pl. iii. 1637 (1803) = Trilisa 
odoratissima. 

Liatris oppositifolia Nutt. in Amer. Jour. Sci. v. 299 (1822) = 
Eupatorium ivaefolium. 

Liatris paniculata Michx. Fl. Bor. Amer. ii. 93 (1803) = Trilisa 
paniculata. 


1946] Gaiser,—The Genus Liatris 411 


Liatris squamosa Nutt. Jour. Acad. Nat. Sci. Phil. vii. 73 (1834) 
= Carphephorus Pseudo-Liatris. 

Liatris tomentosa Michx. Fl. Bor. Amer. ii. 93 (1803) = Carphe- 
phorus tomentosus. 

Liatris trichotoma Gardn. Hook. London Jour. Bot. vi. 435 (1847) 
— Leptoclinium trichotomum Benth. ex. Baker in Martius Fl. 
Brasil. vi. (2) 272 (1875-6). 

Liatris umbellata Bertol. Misc. Bot. v. 13, t. 4 (1846) = Vernonia 
angustifolia. ; 

Liatris Walteri Ell. Sk. ii. 285 (1821-24) = Carphephorus tomen- 
tosus var. Walteri (Ell.) Fern. 


DOUBTFUL SPECIES 


Liatris alata (Nelson) K. Sch. Just, Bot. Jahresb. xxxix. 569 
(1903)—with “reflected lower leaves, long foliar-bracted spike 
and alate bracts of the involucre”; the type of the proposed 
species (Aug. 1895, J. H. Kimmons, Creek Nation, Indian 
Territory of Oklahoma (Rocky Mt. Herb.) ‘‘still represented 
only by the type number from Oklahoma."! 

Liatris botrys Raf. New Fl. N. Amer. iv. 76 (1838)—remains a 
question. 

Liatris heterophylla R. Br. in Ait. Hort. Kew. ed. 2 iv. 503 (1812)— 
Though the type specimen has not been seen, from examination 
of a photograph of it and the phyllaries of a head of it in a small 
packet at the Gray Herbarium, this still seems a doubtful and 
rare species (see discussion no. 17). 

Liatris hirsutissima Poir. ex Steudel, Nom. Bot. ed. 2, ii. 40 (1841) 
—published in synonymy of Liatris pycnostachya Michx. 

Liatris linaria Raf. Fl. Ludov. 61. (1817)—descr. Cynarocephale 
élégante sans épines Rob. Voy. Louis. iii. 429 (1807). Prob- 
ably L. pycnostachya Michx. 

Laciniaria platylepis Small, Bull. Torr. Bot. Club xxvii. 281 
(1900); Fl. S. E. U. S. 1174 (1903).— Type specimen of Hale 
from Louisiana (N. Y.) said not to be like aczdota because of 
inner rounded bracts. 

Inatris picta Barton ex Greene Herb.—Published in synonymy 
of L. spicata L. in DC. Prodr. v.! 130 (1836). The specimen 
has not been located.? 

Liatris pumila Lodd. Cab. t. 147 (1817)—Insufficient description 
to identify; by plate probably L. spicata (L.) Willd.  Lacini- 
ps p var. pumila Porter, Mem. Torr. Bot. Club v. 314 

1894). 

Liatris rigida Raf. New Fl. Amer. iv. 76 (1838)—insufficient 

description. 


1 By private communication from Prof. A. Nelson. 
? By private communication from Dr. F. W. Pennell and Mr. C. A. Weatherby. 


412 Rhodora [DECEMBER 


Laciniaria scariosa var. trilisioides Farwell, Rep. Mich. Acad. Sci. 
xvii. 170-171 (1916)—Liatris novae-angliae var. Nieuwlandii f. 
trilisioides (Farwell) Shinners, Amer. Mid. Nat. xxix. 31 (1943) 
—was reported as a sweet-scented form, but the type has not 
been available. 

Laciniaria serotina Greene, Pitt. iv. 315 (1901)—probably an 
intermediate between Liatris spicata (L.) Willd. and L. 
pycnostachya Michx. (See discussion no. 1). 

Liatris turbinata Sweet, Hort. Brit. ed. i. 220 (1836) = nomen. 

Liatris uniflora Raf. New Fl. Amer. iv. 76 (1836)—by description 
and region of collection may very well apply to specimens of 
L. squarrosa (L.) Willd. var. gracilenta, bearing one head only. 

Liatris varia Herb. Banks, Pursh, Fl. ii. 508 (1814)— published in 
synonymy of L. heterophylla R. Br. 


McMasTER UNIVERSITY, 
Hamilton, Ontario. 


IMPATIENS ROYLEI VERSUS I. GLANDULIFERA 
C. A. WEATHERBY 


The handsome Himalayan species of Impatiens commonly 
known as J. Roylei has now escaped from cultivation and estab- 
lished itself so successfully at so many localities, from Nova 
Scotia to Washington and British Columbia, that it has earned a 
place in our manuals. Its correct name therefore becomes a 
matter of some importance. 

I. Roylei is a substitute name given by Walpers, Repert. 1. 475 
(1842), to I. glandulifera Royle because of 7. glandulifera Arnott, 
which Walpers either supposed to be earlier or, at a time when 
priority was more lightly regarded than at present, thought 
preferable. Since the title-page of Royle’s Illustrations bears 
the date 1839 and that of the first volume of the Companion to 
the Botanical Magazine, in which Arnott’s species was published, 
1835, Walpers may well have thought he was observing strict 
priority. In any case, his name was adopted by Sir Joseph 
Hooker in the Flora of British India and the Epitome of the 
British Indian Species of Impatiens, and by nearly everyone else 
who has had occasion to refer to the species, though J. glanduli- 
fera has persisted to some extent in horticultural literature and 
has been regularly cited as a synonym by L. H. Bailey (e. g. in 
Hortus Second). 


1946] Weatherby,—Impatiens Roylei versus I. glandulifera 413 


In 1900, however, in a note which seems to have been generally 
overlooked (Journ. Bot. xxxviii. 87-88), Hiern pointed out that 
both Royle's work and the Companion were published in parts 
and that the part of the former which contains 7. glandulifera 
was issued more than a year before the part of the Companion in 
which Arnott's homonym appeared. Sprague in 1933 (Kew 
Bull. 362-364; 378-390) went into the matter in detail, estab- 
lishing dates of issue for all parts of both works and confirming 
Hiern's conclusions. "They were further confirmed by Stearn in 
1943 (Journ. Arnold Arb. xxiv. 484—487) from a copy of Royle's 
Illustrations still in the original fascicles which he discovered. 

The correct name for the species is, then, I. glandulifera Royle. 
Bibliography (the dates for Royle and Arnott taken from 
Sprague) is as follows. 


IMPATIENS GLANDULIFERA Royle, Ill. Bot. Himalaya Mts. t. 
28, fig. 2, plate without analyses of flower, (March, 1834) and p. 
151 (Jan., 1835); Lindl. in Bot. Reg. xxvi. t. 22 (1840), as glandu- 
ligera; Hook. in Bot. Mag. t. 4020 (1843), as glanduligera; 
Sprague in Kew Bull. 1933: 386. Not J. glandulifera Arn. in 
Comp. Bot. Mag. i. 322 (June, 1836).—1. Roylei Walp. Repert. 
i. 475 (1842); Hook. f. & Thomps. in Journ. Linn. Soc. iv. 127 
(1860); Hook. f. Fl. Brit. India, i. 468 (1872) and in Rec. Bot. 
Survey India, iv. 7 (1904); E. Loew in Engl. Bot. Jahrb. xiv. 
167, t. 1, 2 (1892); Bailey, Standard Cycl. Hort. 1644, fig. 1954 
(1915); Toppin in Kew Bull. 1920: 347. 


The flowers of 7. glandulifera are typically, except for the 
short green spur, of a uniform dark magenta-red without spots, 
and, with a single exception, all the American material I have 
seen is of this typical form, or at least indistinguishable in the 
dried state. At St. John, New Brunswick, however, where the 
species crowds several small areas of moist waste ground, many 
individuals have pale pink corollas with brownish or reddish 
spots on the sac, yellowish spur and a narrow green band running 
along the upper side of the sac to the spur. This is apparently 
I. Roylei var. pallidiflora Hook. f. in Bot. Mag. t. 7647 (1899). 
Since Hooker gives no differential characters except color of 
flower and size of plant for his variety and since the St. John 
material seems to show no others, this may appropriately be 
treated as a color-form and, at some risk of repeating a combina- 
tion already made in recent European literature, be called 


414 Rhodora [DECEMBER 


I. GLANDULIFERA Royle, forma pallidiflora (Hook. f.) n. comb. 
I. Roylei var. pallidiflora Hook. f. in Bot. Mag. t. 7647 (1899). 

Hegi, Ill. Fl. Mittel-Eur. v. 314 (1924), describes a pure albino 
as I. Roylei forma albida Hort. This is very likely the same as 
I. Roylei var. candida (Lindl. Hook. f. Fl. Brit. India, i. 469 
(1872); I. candida Lindl. in Bot. Reg. xxvi. Misc. Not. 85 (1840) 
and xxvii. t. 20 (1841). Isaw no pure white flowers at St. John.! 


Gray HERBARIUM. 


1 Since the above was in type I have, through the kindness of Prof, G., N. Jones, 
seen further specimens of two collections by him—Mukilteo, Snohomish Co., Wash- 
ington, Aug. 4, 1937, no. 10545 &nd Port Moody, British Columbia, Aug. 14, 1935, 
s.n. These show that the plants at both the above stations were of forma pallidi- 
flora. I have seen no other western material. 


Volume 48, no. 575, containing pages 329—392, was issued ? November, 1946. 


ERRATA 


Page 14, line 28; for 1006 read. 1000. 

Page 38, line 5; for transfer read transfers. 

Page 44, line 23; for aments, with read aments with. 
Page 142, line 1; for S. U. Se. read Se. U.S. 

Page 173, line 23; for Willd. read Michx. 

Page 254, line 4; for 16028 read 10028. 

Page 262, line 31; for Jones read Jonas. 


1946] Index to Volume 48 i 


INDEX TO VOLUME 48 


New scientific names are printed in full-face type 


Abies balsamea, 163 

Acerates, 209; lanuginosa, 116; 
viridiflora, var. linearis, 208, 209 

Achillea lanulosa, 116 

Achras bahamensis, 164; zapotilla, 
var. parviflora, 164, 8. parvifolia, 
164 

Acrostichum hispidum, 384 

Adiantum vestitum, 383-386 

Agrimonia, 71; gryposepala, 71; 
microcarpa, 71; pubescens, 71; 
rostellata, 71 

Agropyron, 22; pungens, 20; repens, 
20, 22, f. aristatum, 20, f. pilosum, 
20, var. subulatum, 20, 25, f. 
heberhachis, 20, f. setiferum, 20, 
f. Vaillantianum, 20; repens, f. 
trichorrhachis, 20; Smithii, 20, 
100; trachycaulum, var. glaucum, 
20, 23, var. majus, 20, 23, var. 
novae-angliae, 20, 23 

Ainsliaea aptera, 410; pteropoda, 
410 

Alders, 205 

Alisma brevipes, 86, 87; parviflorum, 
87, 88; Plantago, 87, e. ameri- 
canum, 87, 6. parviflorum, 88, 
var. triviale, 87; Plantago-aqua- 
tica, 86, 87, 330, North American 
Representatives of, 86, subsp. 
brevipes, 86, 87, var. brevipes, 87, 
var. parviflorum, 88, var. triviale, 
87; subcordatum, 87, 88; superb- 
um, 87; triviale, 87 

Allium canadense, 115; tricoccum, 
61, 62, in Kalamazoo County, 
Michigan, Observations on two 
Ecological Races of, 61; tri- 
florum, 62 

Altitude in New Hampshire, Betula 
glandulosa at a low, 183 

Amblystegium Juratzkanum, 2; 
varium, var. ovatum, 2 

Ambrosia, 73, 74 

Amelanchier, 125, 128-131, 133, 
(Review), A Monograph of, 129; 
amabilis, 132; austromontana, 
126, 127, 131, 132; Bartramiana, 
130, 132; canadensis, 126, 128- 
130; X grandiflora, 130; humilis, 
126, 127, 131-133; huronensis, 
132; laevis, 130; neglecta, 130, 
132, 133; oblongifolia, 126, 128; 


ovalis, 125, 128, 129, 131, 132, 
pl. 1028; pallida, 132; rotundi- 
folia, 128, 131; sanguinea, 132; 
spicata, 125-129, 131-133, pl. 
1027, Not an American Species, 
125; stolonifera, 126-129, 131- 
133, pls. 1029, 1030; vulgaris, 
128, 131 

American Forms of the Lady-fern, 
Some Trivial, 389 

American Species, Amelanchier spi- 
cata not an, 125 

Ammi capillaceum, 162 

Ammopursus, 375; Ohlingerae, 374 

Amos Eaton Herbarium, The, 201 

Anacharis occidentalis, 330 

Anemone canadensis, 116 

Anonymos graminifolia, 247, 248, 
255; graminifolius, 222 

Another later Homonym, 163 

Antennaria, 116-118, 121; alpina, 
119, 121, var. canescens, 119; 
aprica, 120; Brainerdii, 118; dio- 
ica, 119; fallax, 118; microphylla, 
119, 120; neglecta, 118, 119, 121; 
neodioica, 118; Parlinii, 118; 
parvifolia, 119-121; petaloidea, 
118; plantaginifolia, 119, 121; 
rosea, 120; solitaria, 121; sub- 
viscosa, 119; umbrinella, 119, 
121; vexillifera, 119; ,virginica, 
118, var. argillicola, 118 

Appearance of Epipactis Helle- 
borine, Sporadic, 88 

Application of the Name Euphorbia 
maculata L., 197 

Aquatic and Prairie Vegetation in 
Southwestern Minnesota, Notes 
on, 113 

Arabis dentata, 208, var. phala- 
crocarpa, 208; perstellata, 208, 
var. phalacrocarpa, 208, var. 
Shortii, 208 

Araceae, 267 

Arethusa divaricata, 186 

Aristida tuberculosa, 37 

Arkansas, Vegetation of Artificial 
Lakes in Northwestern, 329 

Arnica fulgens, 101, 103 

Artificial Lakes in Northwestern 
Arkansas, Vegetation of, 329 

Asclepias, 209; viridiflora, 209, var. 
lanceolata, 209, var. linearis, 208 


il Rhodora 


Ash, Red, 61 
Asperella Hystrix, var. Bigeloviana, 
22 


Asplenium fontanum, 383 

Aspidium, 385; lanosum, 385, 386 

Aster anticostensis, 271 

Astomum Muhlenbergianum, 3, 4 

Astragalus bisulcatus, 99; caryo- 
carpus Fruits, Size, Shape and 
Number in, 111; Drummondii, 
100; pectinatus, 99, 100; tenellus, 
102 


Athyrium angustum, 389, 390, var. 
confertum, 389; angustum, f. 
laciniatum, 389, var. laurenti- 
anum, 389, 390; asplenioides, 389, 
f. subtripinnatum, 389; Filix- 
femina, 389, 390, var. aspleni- 
oides, f. subtripinnatum, 389, 
var. Michauxii, f. confertum, 
389, f. laciniatum, 389, f. laur- 
entianum, 389, var. sitchense, 
389 

Aulacomnium palustre, var. imbri- 
catum, 1 


Back to Carex rostrata, 145 

Baldwin, J. T., Jr., Spiraea latifolia, 
var. septentrionalis in Virginia, 
112 

Balsam Poplar, 103-109 

Bartonia paniculata, 328; tenella, 
328; verna, 327, 328, grow in 
Virginia?, Does, 327; virginica, 
328 

Bean, R. C., Ninth Report of the 
Committee on Plant Distribu- 
tion, 17 

Beech, 61 

Bermudiana graminea, flore, etc., 
153-158, 215; Grass-leaved, 154 

Betula glandulosa, 183, at low Alti- 
tude in New Hampshire, 183 

Bidens, 83; frondosa, 330; hyper- 
borea, 82, 83, var. laurentiana, 
83, var. typica, 82, 83 

Birch, swamp, 61 

Black Poplar, 106 

Blewitt, Arthur E., Edgar Burton 
Harger, 263 

Blue-eyed Grass, 153 

Boreal Plants in Southeastern Min- 
nesota, Relict, 163 

Botanical Visits to Forts Clark, 
Mandan and Union in North 
Dakota, 98, pls. 1025, 1026 

Botrychium minganense, 271 

Bouteloua 27, curtipendula, 17; 


[DECEMBER 


gracilis, 17, 100; radicosa, 17; 
rigidiseta, 17; simplex, 17 

Brachythecium acutum, 2; Bestii, 
2; campestre, 2; flexicaule, 2; 
populeum, 2; rivulare, 2; velu- 
tinum, 2 

Briza media, 18; minor, 18 

Bromus, 21; arvensis, 18; brizae- 
formis, 18; ciliatus, 18, 23, 25, 
var. intonsus, 18, 23; commu- 
tatus, 18; Dudleyi, 18, 25; 
erectus, 18; inermis, 18, f. ari- 
status, 18; Japonicus, 18; Kalmii, 
18, 24; latiglumis, 18, 25, 26, f. 
incanus, 18; marginatus, 18, var. 
seminudus, 18; mollis, 18, f. 
leiostachys, 18; purgans, 18, 24, 
f. glabriflorus, 18, f. laevivagin- 
atus, 18; racemosus, 18; rigidus, 
var. Gussonii, 18; rubens, 18; 
secalinus, 18; sterilis, 18; squar- 
rosus, 18; tectorum, 18, 26 

Bryhnia Hultenii, 2 

Bryophytes, 1 

Buckwheat, 97 

Buddleia, 175 

Bursera Simaruba, 105 


Calceolus parviflorus, 4 

Calipogon parviflorum, 11 

Callitriche heterophylla, 114 

Calopogon barbatus, 11, var. multi- 
florus, 11; multiflorus, 11; pal- 
lidus, 11; parviflorus, 11; pul- 
chellus, 189, f. latifolius, 188, var. 
latifolius, 188, 189 

Camelina microcarpa, 99 

Campanula, 209, 212 

Capnoides flavulum, 326; Halei, 207 

Cardamine pulcherrima, 12 

Carex 70, 82, 145, 146; ampullacea, 
146; crinita, 55, var. brevicrinis, 
54-50, var. gynandra, 55, 56; 
Eleocharis, 115; gynandra, 330; 
inflata, 145, 146, var. ambigens, 
146, var. anticostensis, 146, var. 
utriculata, 146; laevigata, 145, 
146; Mitchelliana, 55, 56; Pseudo- 
Cyperus, 55; rostrata, 145, 146, 
Back to, 145, var. ambigens, 146, 
var. anticostensis, 146, var. 
utriculata, 146; utriculata, 146, 
var. minor, 146; vesicaria, 145 

Carices, 145 

Carphephorus  bellidifolius, 410; 
corymbosus, 410; Pseudo-Liatris, 
411; tomentosus, 411, var. 
Walteri, 411 


1946] Index to Volume 48 iii 


Carya cordiformis, 206; ovalis, 206; 
ovata, 206 

Cathea pulchella, f. latifolia, 188 

Centaurea maculosa, 98, in Indiana, 
391 

Centaurella autumnalis, 328; pani- 
culata, 328; verna, 327, 328; 


Chara sp., 330; Braunii, 329; 
fragilis, 330 

Cheilanthes 383, 386, 388; Feei, 386, 
387; gracilis, 386; lanosa, 387, 
388, Presumable Identity of, 383; 
lanuginosa, 386-388; siliquosa, 
272; tomentosa, 383, 384, 388; 
vestita, 384—388 

Chenopodium, 100; album, 100; 
Berlandieri, 100 

Chloris cucullata, 17, 27; elegans, 
27; virgata, 17 

Chrysobalanus, New Combination 
in, 136; pallidus, 136 

Chrysocoma affinis, 255 

Chrysopsis mariana, f. efulgens, 69 

Cicuta Victorinii, 272 

Cirsium minganense, 271; non 
ramosum, etc., 296, 298 

Cleistes divaricata, 186-188, 215, 
pl. 1047, var. bifaria, 186-188, 
215, pl. 1048, var. typica, 187 

Cliff-brake, Oregon, 272 

Clubmosses, 267 

Collomia linearis, 99 

Combination in  Chrysobalanus, 
New, 136 

Commelina hirtella, 330 

Committee on Plant Distribution, 
Ninth Report of the, 17 

Compositae, 170; of the North- 
eastern United States, III. Inu- 
leae and Senecioneae, Notes on, 
116; subtribe Ambrosinae, 73 

Connecticut, Second Station for 
Corydalis flavula in, 326 

Conringia, 99 

Contributions from the Gray Her- 
barium of Harvard University— 
No. CLX, 5-16, 27-40, 41-60, 
65-81, pls. 993-1020; No. CLXII, 
137-162, 184-197, 207-216, pls. 
1031-1050 

Coral-roots, 193 

Corallorhiza, 193, 194, 197; Coral- 


lorhiza subsp. coloradensis, 194, 
196; ericetorum, 196; innata, 193, 
195, var. virescens, 196; macu- 
lata, 197, var. punicea, 197; 
montana, 12; odontorhiza, 12, 
197, B. verna, 196; striata, 197, f. 
fulva, 197; trifida, 193-196, var. 
verna, 196, var. virescens, 196; 
verna, 194—196 

Corallorrhiza, 193, 194 

Cornus canadensis, 163 

Corona solis minor, etc., 75 

Correction, Helianthus—a, 112 

Cory, V. L., Genus Palafoxia in 
Texas, 84 

Corydalis aurea, var. australis, 207; 
flavula, 326, 327, in Connecticut, 
Second Station for, 326; Halei, 
207; micrantha, 207, var. diffusa, 
207, var. pachysiliquosa, 207; 
sempervirens, 327 

Crataegus, 95, 129; racemosa, 127, 
128; rotundifolia, 128, 131; spi- 
cata, 125-129, 131, 132, pl. 1027 

Crepis, 117; acuminata, 117 

Cronquist, Arthur, Notes on Com- 
positae of the Northeastern Unit- 
ed States, III. Inuleae and 
Senecioneae, 116 

Cymbidium odontorhizon, 12 

Cynarocephale, 411 

Cynodon dactylon, 17 

Cynosurus, 26; cristatus, 18 

Cyperus acuminatus, 330; escu- 
lentus, 330; odoratus, 330; ovu- 
laris, 330; strigosus, 330 

Cypripedium bulbosum, var. parvi- 
florum, 4; Calceolus, 8., 4, var. 
parviflorum, 4; Calceolus L., 
var. parviflorum, 4, var. pubes- 
cens, 4; flavescens, B., C. parvi- 
florum, 4; hirsutum, var. parvi- 
florum, 4; luteum, var. parvi- 
florum, 4; parviflorum, 4 

Cystopteris fragilis, 390 


Dactylis, 21; glomerata, 18, var. 
ciliata, 18, var. detonsa, 18, var. 
vivipara, 18 

Dactyloctenium aegyptium, 17 

Dactylogramma cinnoides, 64 

Danthonia, 212 

Darlington Oak, 144 

Delphinium virescens, 115 

Dentaria grandiflora, 12; macro- 
carpa, 12, var. pulcherrima, 12; 
tenella, var. pulcherrima, 12 

Descurainia Sophia, 99 


iv Rhodora 


a eximia, 272, in Vermont, 

272 

Dicranum fulvum, 1; fuscescens, 2; 
scoparium, var. orthophyllum, 2 

Difficulties in North American 
Salix, 13-16, 27-40, 41-49, pls. 
995-1006 

Dilly, Wild, 164 

Diplachne maritima, 18, 25; uni- 
nervia, 18 

Distichlis spicata, 18, 25 

Ditrichum pallidum, 2 

Does Bartonia verna grow in Vir- 
ginia?, 327; Habenaria cristata 
still grow in New England?, 64 

Dog’s-tooth-violet, 266 

Downy False Foxglove, 205, 206 

Drepanocladus aduncus, var. capil- 
lifolius, 3, var. Kneiffi, 2 


Eastern United States, Stenanthium 
in the, 148-152, pls. 1037-1041 
Eaton, Richard J., Dicentra eximia 
in Vermont, 272 

Echinacea, 99; angustifolia, 99 

Echinochloa colonum, 330; pungens, 
330 

Ecological Races of Allium tricoc- 
cum in Kalamazoo County, 
Michigan, Observations on two, 
61 

Elatine triandra, 114, f. intermedia, 
114, f. terrestris, 116 

Eleocharis caribaea, 37; geniculata, 
37; melanocarpa, 37; obtusa, 330 

Eleusine indica, 17, 26 

Elm, white, 61 

Elymus, 22; arenarius, var. villosus, 
20, 25; canadensis, 20, 24, f. glau- 
cifolius, 20; caput-medusae, 20; 
riparius, 20, 23, 24; villosus, 20, 
24, f. arkansanus, 20; virginicus, 
20, 22, f. australis, 21, var. 
glabriflorus, 21, var. halophilus, 
21, 25, f. hirsutiglumis, 21, var. 
jejunus, 21, var. submuticus, 21; 
Wiegandii, 21, f. calvescens, 21 

Epipactis Helleborine, 88, Sporadic 
Appearance of, 88; latifolia, 88 

Equisetum arvense, f. campestre, 82 

Eragrostis capillaris, 18, 24; Frankii, 
18, 24; hypnoides, 18, 24, 330; 
intermedia, 18; megastachya, 18; 
multicaulis, 18, 21; pectinacea, 
18, 24; pilosa, 18; poaeoides, 18; 
spectabilis, 18, 24, var. sparsi- 
hirsuta, 19, 24 

Erechtites hieraciifolia, 122, var. 


[DECEMBER 


Demon 122; megalocarpa, 

2 

Erigeron glabellus, 102, var. asper, 
102; strigosus, 116 

Eriocaulon decangulare, 58 

Eriogonum flavum, 101; multiceps, 
101; pauciflorum, 101 

Eupatorium ivaefolium, 410; speci- 
osum, 340 

Euphorbia glyptosperma, 100; hy- 
pericifolia, 200; maculata, 197— 
200; maculata L., Application of 
the Name, 197; nutans, 197; 
Preslii, 197; supina, 189—200 

Eurhynchium hians, 3; strigosum, 
var. robustum, 3; Swartzii, 3 


Fagopyrum esculentum, 97 

False Foxglove, Downy, 205, 206 

Fassett, Norman C., Bidens hy- 
perborea, var. typica, 82 

Fernald, M. L., A Monograph of 
Amelanchier (Review), 129; Ame- 
lanchier spicata not an American 
Species, 125; Cypripedium Cal- 
ceolus, var. parviflorum, 4; Dif- 
ficulties in North American Salix, 
13-16, 27-40, 41-49, pls. 995— 
1006; Does Bartonia verna grow 
in Virginia?, 327; Does Habenaria 
cristata still grow in New Eng- 
land?, 64; Helianthus—a Correc- 
tion, 112; Identifications and 
Reidentifications of North Ameri- 
can Plants, 137-162, 184-197, 
207—216, pls. 1031-1050; Nomen- 
clatural Transfers in Polygonum, 
49, 54; North American Repre- 
sentatives of Alisma Plantago- 
aquatica, 86; Novelties in Our 
Flora, 54-60, 65-80, pls. 1007- 
1020; Presumable Identity of 
Cheilanthes lanosa, 383; Senecio 
tomentosus, forma alabamensis, 
330; Some Species in Rafinesque's 
“Herbarium — Rafinesquianum", 
5-13, pls. 993, 994; Some trivial 
American Forms of the Lady- 
fern, 389; Sporadic Appearance of 
Epipactis Helleborine, 88; Tech- 
nical Studies on North American 
Plants, 5-16, 27-40, 41-60, 65- 
81, pls. 993-1020; Triodanis 
versus Specularia, 209—214, pls. 
1049, 1050; Varieties of Lyco- 
podium inundatum, 134 

Ferraria pulchella, 158 

Festuca, 21, 70; capillata, 19; 
elatior, 19; obtusa, 19, 23, 24; 


1946] Index to Volume 48 v 


ovina, 19, var. duriuscula, 19, f. 
hispidula, 19; prolifera, 19, 23; 
rubra, 19, 25, var. commutata, 
19, f. megastachys, 19, var. mul- 
tiflora, 19; f. squarrosa, 19; var. 
trichophylla, 19; saximontana, 
19, 23 

Festucas, 26 

Firs, 163 

Floating knotweed, 52 

Flora, Novelties in Our, 54-60, 65- 
81, pls. 1007-1020 

Forms of the Lady-fern, Some 
Trivial American, 389 

Forts Clark, Mandan and Union in 
North Dakota, Botanical Visits 
to, 98, pls. 1025, 1026 

Fosberg, F. R., Application of the 
Name Euphorbia maculata L., 
197 

Frére Marie-Victorin, 265 

Fringed-orchid, 185 


Gaiser, L. O., The Genus Liatris, 
165-183, 216—263, 273-326, 331- 
382, 393-412 

Garberia fruticosa, 410 

Gentiana Victorinii, 272 

Genus Liatris, The, 165-183, 216- 
263, 273-326, 331-382, 393-412 

Genus Palafoxia in Texas, 84 

Geobalanus pallidus, 136 

Gerardia, 206; virginica, 205 

Gilly, Charles L., Another later 
Homonym, 163 

Glyceria, 21; acutiflora, 19, 24; 
borealis, 19, 23; canadensis, 19, 
22; Fernaldii, 19, 22; fluitans, 19; 
grandis, 19, 22, f. pallescens, 19; 
laxa, 19, 25, 26; melicaria, 19, 23; 
obtusa, 19, 25, 26; pallida, 19, 24; 
septentrionalis, 19, 24; striata, 
19, 22, var. stricta, 19, 23 

Gnaphalium obtusifolium, 121, var. 
saxicola, 121; purpureum, 121, 
bere purpureum, 121; saxicola, 
121 

Goodyera decipiens, 11; Menziesii, 
11, 12; oblongifolia, 11, 12 

Gramineae, 205, Tribe Chlorideae, 
17, Tribe Festuceae, 17, Tribe 
Hordeae, 17 

Grass, Blue-eyed, 153 

Grass-leaved Bermudiana, 154 

Gratiola neglecta, 115 

Gray Herbarium of Harvard Uni- 
versity, Contributions from the, 
—No. CLX, 5-16, 27-40, 41-60, 
65-81, pls. 993-1020; No. CLXII, 


137-162, 184-197, 207-216, pls. 
1031-1050 

Gray's Manual Range, Notes on 
certain Plants in the, 89, pls. 
1021-1024 

Grimmia alpicola, 2; apocarpa, 2 

Gymnosperms, 267 

Gyrostachys Beckii, 6; Grayi, 6; 
parviflora, 5; simplex, 6 


Habeeb, Herbert, Some Mosses 
from Windsor, Nova Scotia, 1 
Habenaria X Andrewsii, 184, 185; 
clavellata, 10, 161, 162, 215, pl. 
1045, var. ophioglossoides, 161, 
162, 215, pl. 1046; cristata, 64, 
still grow in New England?, 
Does, 64; elegans, var. maritima, 
10; fimbriata, 184-186, f. albi- 
flora, 185, f. mentotonsa, 184, 
185; lacera, 184—186, var. terrae- 
novae, 184, 185; maritima, 10, 11; 
psycodes, 184—186, f. ecalcarata, 
184, f. varians, 184, var. varians, 
184; psycodes X lacera, 185 

Hanes, Clarence R., Observations 
on two Ecological Races of Al- 
lium tricoccum in Kalamazoo 
County, Michigan, 61 

Harger, Edgar Burton, 263 

Harpalium rigidum, 79 

Harvard University, Contributions 
from the Gray Herbarium of— 
No. CLX, 5-16, 27-40, 41-60, 
65-81, pls. 993-1020; No. CLXII, 
137-162, 184-197, 207-216, pls. 
1031-1050 

Hedysarum boreale, 102 

Helenium nudiflorum, 82 

Helianthus, 112;—a Correction, 112; 
annuus, 79; atrorubens, 75-79, 
var. alsodes, 74-76, 81, pl. 1020, 
var. normalis, 75, 76, var. pubes- 
cens, 77, 78; foliis ovatis, 75; 
dies: 76, 77; heterophyllus, 77; 

entuckiensis, 77, 78; laetiflorus, 

79, 80, 112, var. rigidus, 79, 80, 
112; var. subrhomboideus, 79, 
rigidus, 79, 80, f. flavus, 79, 112; 
scaberrimus, 78, 79, var. subrhom- 
boideus, 79, 80; silphioides, 77— 
79; sparsifolius, 76; subrhomboi- 
deus, 79, 80 


'Heliotropium indicum, 330 


Helleborine Lilii folio caulem, etc., 
186 

Helonias graminea, 148, 149, 151 

Herbarium, The Amos Eaton, 201 

“Herbarium Rafinesquianum”, 


vi Rhodora 


Some Species in Rafinesque’s, 5- 
13, pls. 993, 994 

Hermann, F. J., Muhlenbergia 
setosa an Untenable Name, 63 

Heteranthera limosa, 114 

Heterocodon, 209, 211, 212 

Hickory, 206 

Hill, A. F., Ninth Report of the 
Committee on Plant Distribu- 
tion, 17 

Hodgdon, Albion R., Notes on 
New Hampshire Plants, 205 

Homonym, Another later, 163 

Hordeum aegiceras, 21; jubatum, 
21, 26, 116; marinum, 21; muri- 
num, 21; nodosum, 21; pusillum, 
21, 27, 115; vulgare, 21, var. tri- 
furcatum, 21 

Horsetails, 267 

Hull, Edwin D., Centaurea macu- 
losa in Indiana, 391 

Hydnum chrysorhizum, 202, 203 

Hydranthelium rotundifolium, 116 

-i „remar maae m irriguum, 3 

Hylocomium brevirostre, 3 

Hypericum canadense, 58 

Hypnum curvifolium, 3 

Hystrix, 24; patula, 21, 23, 24, var. 
Bigeloviana, 21-23 


Ibidium Beckii, 6; ovale, 5; parvi- 
florum, 6 

Identity of Cheilanthes lanosa, 
Presumable, 383; Quercus lauri- 
folia, The, 137-145, pls. 1031- 
1036; of Sisyrinchium angusti- 
folium, The, 152-160, pls. 1042- 
1044 

Iliamna, 89, 91, 92, 94; latibracteata, 
95; remota, 89-93, pls. 1021, 
1022, var. Corei, 96, pl. 1024, 
var. typica, 93-95, pls. 1021- 
1023 

Impatiens, 412, candida, 414; glan- 

ulifera, 412, 413, I. Roylei 

versus, 412, f. pallidiflora, 414; 
glanduligera, 413; Roylei, 412, 
413, versus I. glandulifera, 412, 
f. albida, 414; var. candida, 414, 
var. pallidiflora, 413, 414 

Indiana, Centaurea maculosa in, 
391 

Inuleae and Senecioneae, Notes on 
Compositae of the Northeastern 
United States, III., 116 

Todanthus dentatus, 208 

Iris virginica, 15 

Isoetes melanopoda, 114, 115 

Iva xanthifolia, 100, 101 


[DECEMBER 


Juncus, $82; diffusissimus, 330; 
Dudleyi, 115; effusus, 330; in- 
terior, 330; nodatus, 330 

Juniperus horizontalis, 102 


Kalamazoo County, Michigan, Ob- 
servations on two Ecological 
Races of Allium tricoccum in, 61 

Kilbourne, Frederick W., Second 
Station for Corydalis flavula in 
Connecticut, 326 

Knotweed, floating, 52; swimming, 

2 


Knowlton, C. H., Ninth Report of 
the Committee on Plant Distribu- 
tion, 17 

Koeleria cristata, 115 

Kucyniak, James, Frère Marie- 
Victorin, 265 

Kyllinga pumila, 330 


Lacinaria, 167; acidota, 232, 235; 
angustifolia, 368-370; arenicola, 
363; brachyphylla, 232; Chap- 
manii, var. longifolia, 280; cy- 
lindracea, var. solitaria, 381; 
cylindracea X scariosa, var. 
sphaeroidea, 381; Gladewitzii, 
381, 382; Halei, 232, 233, 235; 
Helleri, 260; indecidua, 302, 304; 
kansana, 228, 229; leptostachya, 
364; microcephala, 230, 231; Ò - 
lingerae, 374; regimontis, 277; 
Ruthii, 336, 370; scariosa, var. 
trilisioides, 175; secunda, 278, 
282; spicata, var. albiflora, 179, 
var. foliacea, 179, 180, 227; 
stratiotes, 302, 304 

Laciniaria, 167, 309; aspera, 302; 
carinata, 277, 278; Chapmanii, 
280; chlorolepis, 236, 237; cylin- 
dracea, f. solitaria, 376, 381, var. 
solitaria, 376; cymosa, 373; 
Deamiae, 380, 381, Deamii, 312, 
313, f. albina, 312; densispicata, 
363; Earlei, 336, 338; elegans, 
341; elegantula, 254, 255, 259; 
elongata, 220, 224; fallacior, 362, 
var. celosioides, 362, 363; flabel- 
lata, 345; formosa, 318, 322; 
Garberi, 236; gracilis, 273; laevi- 
gata, 289; lancifolia, 228, 229; 
Langloisii, 238, 244; laxa, 273, 
276, 277; ligulistylis, 318, 322, 
323; macilenta, 238; Nashii, 236, 
237; nervata, 376, 378; pauciflora, 
281, 284; platylepis, 411; poly- 
phylla, 230; Pana 347, f. 
albiflora, 347, f. corymbosa, 347, 


1946] 


var. turgida, 347; Ruthii, 336- 
338; scabra, 316, 317; scariosa, 
309, 311, 323, f. corymbulosa, 
318, f. globosa, 319, f. uniflora, 
318, 319, var. angustata, 318, 
320, var. annuens, 318, 320, var. 
basilaris, 318, 320, var. borealis, 
296, 300, var. brachiata, 323, var. 
Chandonnetii, 323, var. com- 
posita, 318, 320, var. corymbu- 
losa, 318, 323, var. crista-galli, 
318, 320, var. exuberans, 318, 320, 
var. immanis, 318, 320, var. in- 
concinna, 319, 320, var. insolens, 
318, 320, var. intermedia, 305, 
306, 309, 310, var. media, 305, 
309, var. multiplex, 318, 320, var. 
nictitans, 323, var. Nieuwlandii, 
324, 326, 331, f. borealis, 325, 331, 
f. gracillima, 325, 331, f. septent- 
rionalis, 325, 331, f. versicolor, 
325, 331, var. novae-angliae, 333, 
334, 336, var. obesa, 302, 303, 
var. opima, 318, 320, var. perusta, 
318, 319, var. petiolata, 305, 306, 
309, var. porrecta, 302, 303, var. 
praecellens, 323, var. praeceps, 
318, 320, var. praesignis, 325, 
326, 332, var. praestans, 318, 320, 
var. propinqua, 323, var. ramea, 
323, var. salutans, 302, 305, var. 
scalaris, 318, 320, var. singularis, 
318, 320, var. squarrulosa, 296, 
var. strictissima, 302, 303, var. 
subeorymbosa, 318, 320, var. 
superans, 323, var. supereminens, 
318, 320, var. superscandens, 323, 
var. trilisioides, 412, var. uniflora, 
318, 320, var. virgata, 302, 303, 
var. virginiana, 296, 300; serotina, 
225, 226, 412; Shortii, 316; 
Smallii, 253, 258, 259; spicata, f. 
albiflora, 179, var. pumila, 411; 
squarrosa, 394, squarrosa ala- 
bamensis, 398, 407, var. inter- 
media, 376; tenuifolia, 287; 
Tracyi, 336-338; vittata, 217, 


Index to Volume 48 vii 


Lemna abbreviata, 146, 147; ango- 
lensis, 147; arhiza, 147; cy- 
clostasa, 146, 147; gibba, 147; 
minima, 147; minor, 147, var. ? 
Cyclostasa, 146, 147; polyrrhiza, 
147; Torreyi, 146; trisulca, 147; 
valdiviana, 146, 147, 205, not L. 
cyclostasa, 146, var. abbreviata, 
147, var. ? platyclados, 147 

Lepidium densiflorum, 116 

Leptochloa filiformis, 17 

Leptoclinium brasiliense, 410; tri- 
chotomum, 411 

Leptodictyum riparium, 3, f. longi- 
folium, 3 

Lepturus paniculatus, 114 

Lespedeza stipulacea, 391 

Liatris, The Genus, 165-183, 216- 
263, 273-326, 331-382, 393-412; 
sect. Euliatris, 167, 176, 177, 235, 
312, 363, sect. Suprago, 167, 176, 
235, 363, 409; ser. Cylindraceae, 
177, 312, 373, 375, ser. Elegantes, 
177, 340, ser. Graminifoliae, 172, 
177, 226, 232, 237, 246, 260, 279, 
285, 286, ser. Pauciflorae, 177, 
279, 285, 286, ser. Punctatae, 
169-171, 177, 235, 237, 346, 372, 
ser. Pycnostachyae, 169, 173, 177, 
237, 244, 246, 409, ser. Scariosae, 
171, 177, 228, 259, 293, 311, 323, 
331, 362, 374, 380, 409, ser. 
Spicatae, 168-170, 173, 177, 225, 
229, 232, 236, 237, 244, 246, 312, 
ser. Squarrosae, 173, 174, 177, 
380, 393, ser. Tenuifoliae, 174, 
177, 286; acidota, 178, 232-236, 
366, 369, 373, 411, var. mucro- 
nata, 364, var. vernalis, 232, 234; 
alata, 411; amplexicaulis, 410; 
angustifolia, 170, 235, 346, 357, 
363, 365, 366, 368, 370-372; 
aspera, 168, 170, 173-175, 246, 
293, 301, 302, 310-312, 315-317, 
323, 324, 332, 382, var. glabra, 
300, var. intermedia, 305, 311, 
339, var. typica, 302, 315; aspera 


220, 223 

Lactuca ludoviciana, 102; pulchella, 
100; seariola, 100, 102 

Lady-fern, 391; Some trivial Ameri- 
can Forms of the, 389 

Lakela, Olga, Previously unreported 
Plants from Minnesota, 81 

Lakes in Northwestern Arkansas, 
Vegetation of Artificial, 329 

Laurel Oak, 138, 145 

Lechea villosa, 206 

Leersia oryzoides, 330 


X punctata, 314; aspera X 
cylindracea, 381;faspera X pyc- 
nostachya, 245; baicalensis, 410; 
Bebbiana, 238, 239, 243, 244; 
bellidifolia, 410; borealis, 168, 
173, 294, 331, 332, 334-336; 
botrys, 411; X Boykinii, 292; 
bracteata, 346, 367, 371; bra- 
chystachya, 238, 243, brasiliensis, 
410; carinata, 277, 285; Chap- 
manii, 279, 281; compacta, 401; 
cordata, 410; corymbosa, 410; X 


viii 


creditonensis, 409; cylindracea, 
168, 171, 172, 174, 175, 312, 361, 
373-375, 379, 380, 382, 404, 405, 
408, 409, var. solitaria, 376; cylin- 
dracea X sphaeroidea, 381; cylin- 
drica, 347, 350, 361, 362; cymosa, 
171, 373, 375; X Deamii, 313; 
densispicata, 170, 237, 346, 363; 
dubia, 250, 255, 258; Earlei, 294, 
336, 339, 340; elegans, 169, 173, 
174, 292, 293, 340, 341, 343, 345, 
var. carizzana, 169, 341, 344, 
345; elegans, f. Fisheri, 174, 340, 
344, var. flabellata, 341, 345, 
var. typica, 341, 344, 345; ele- 
ans X tenuifolia, 292; X fal- 
acior, 325, 362; flabellata, 345; 
flexuosa, 376, 380; X Frostii, 245, 
246; fruticosa, 410; Garberi, 169, 
178, 225, 236, 237; glabrata, 401, 
404; Gladewitzii, 312; X Glade- 
witzii, 381, 382; gracilis, 176, 247, 
259, 273, 276, 277, 284; gramini- 
folia, 176, 221, 226, 232, 247, 248, 
255, 258-261, 263, 301, var. 
dubia, 247, 250, 253, 256-202, 
278, var. pe 248, 254, 
259, 277, var. lasia, 248, 253, 258, 
var. Smallii, 248, 253, 259, 261, 
263, var. typica, 247, 248, 255, 
256, 261, 278; Haywardii, 318, 
320, 324; Helleri, 176, 247, 259, 
260, 263; Herrickii, 318, 319, 324; 
heterophylla, 301, 411, 412; hir- 
suta, 399, 404; hirsutiflora, 395; 
hirsutissima, 411; intermedia, 376, 
380, 404, 405, 409; kansana, 228, 
229; laevigata, 170, 286, 288, 290- 
292; lanata, 410; lanceolata, 273, 
276, 277; lancifolia, 177, 228-230, 
243, 244; Langloisii, 238, 240, 
244; latifolia, 410; laxa, 273; lig- 
ulistylis, 168, 171-174, 176, 177, 
246, 293, 311, 318, 324, 325, 331, 
332, 361-363, 374, 380, 409, 410, 
f. leucantha, 319; ligulistylis X 
squarrosa, var. glabrata, 409; 
linaria, 411; lobelioides, 410; ma- 
erostachya, 178, 221, 222, 242; 
magnifica, 178; marginata, 376; 
microcephala, 168, 172, 178, 230, 
232, 236, 279; monocephala, 376; 
mucronata, 170, 232-235, 346, 
357-360, 364-368, 370, 372, var. 
interrupta, 361, 365, 372, var. 
typica, 364; X Nieuwlandii, 311, 
312, 325; novae-angliae, 333, f. 
albiflora, 333, var. Nieuwlandii, 
325, f. alba, 325, f. trilisioides, 


Rhodora 


[DECEMBER 


412; odoratissima, 410; Ohlinger- 
ae, 373-375; oppositifolia, 410; 
paniculata, 410; pauciflora, 279, 
281, 284-286; pauciflosculosa, 
273, 276; picta, 411; pilosa, 250, 
253, 257-259, 263; propinqua, 
250; pumila, 411; punctata, 168, 
170, 172, 175, 176, 233, 237, 285, 
314, 315, 325, 346, 347, 350, 352, 
354, 355, 357-362, 366, 367, 372, 
404, var. B., 353, 356, 357, var. 
Y., 347, 348, 356, var. mexicana, 
347, 364, 360, 361, 367, var. ne- 
braskana, 314, 347, 353, 356- 
358, 363, 370, 371, var. turgida, 
172, var. typica, 347, 356-358, 
360, 361, 363, f. coloradensis, 
351, 357, 358, 360, 367; pycnosta- 
chya, 170, 171, 173, 226, 229, 230, 
232, 242-246, 409, 411, 412, a., 
238, 8., 238, 243, f. Hubrichti, 
238; pycnostachya X squarrosa, 
245; radians, 340; regimontis, 227, 
247, 271, 278, 285, 286; resinosa, 
217, 222, 223, 353; x Ridgwayi, 
245, 409; rigida, 411; Rosen- 
dahlii, 318, 319, 322-324; scabra, 
293, 316, 332, 340; scariosa, 166, 
173, 258, 293, 294, 298, 300—302, 
304, 308, 309, 311, 317, 318, 322, 
323, 332, 335, 338, 339, 375, 382, 
404, f. Benkii, 302, var. Deamii, 
313, 8. intermedia, 332, var. 
sphaeroidea, 382, var. squar- 
rulosa, 294, 296, 301, var. typica, . 
294, 296, 300, 312, 336, var. vir- 
giniana, 293, 294, 296, 300, 311; 
secunda, 279, 282, 284-286, 292; 
sessiliflora, 217, 219, 223, 225; 
sphaeroidea, 302, 305, 309, 313, 
315, 382, f. asperifolia, 302, f. 
Benkii, 302, var. salutans, 302; 
sphaeroidea X cylindracea, 381; 
X sphaeroidea, 168, 228, 311, 
312, 382; spicata, 168, 171, 174, 
175, 177, 178, 220-222, 224—220, 
228, 229, 232, 236, 242-245, 256, 
261—203, 286, 312, 411, 412, var. 
B., 217, 222, f. albiflora, 179, 227, 
B. macrostachya, 178, var. ma- 
crostachya, 222, var. montana, 
216, 222, y. racemosa, 250, var. 
resinosa, 216, 223-225, 227, 286, 
var. typica, 178, 216, 217, 223- 
225, 227-229, f. montana, 216, 
223, 229; spicata X sphaeroidea, 
227; squamosa, 411; squarrosa, 
171-174, 220, 245, 333, 361, 376, 
379, 380, 393, 395, 401, 403-406, 


1946} Index to Volume 48 ix 


408, 409, 8., 406, var. è., 394, 
var. alabamensis, 394, 398, var. 
compacta, 394, 401, 406-408, g. 
floribunda, 395, var. floribunda, 
403, var. glabrata, 380, 394, 398, 
401, 406—410, squarrosa glabrata, 
401, var. gracilenta, 394, 397, 
398, 403, 405—408, 412, var. hir- 
suta, 380, 394, 399, 401, 407, 408, 
squarrosa hirsuta, 399, 8. inter- 
media, 376, var. intermedia, 394, 
402, 404, 405, 407, var. typica, 
394, 401, 404, 405, 407, 408; 
squarrulosa, 296, 300, 312, 336, 
338, 339; X Steelei, 227, 312; 
stricta, 376, 380; tenuifolia, 175, 
286, 288, 290-293, 345, 8., 289, 
var. laevigata, 289, var. quadri- 
flora, 289, 290; tomentosa, 411; 
trichotoma, 411; turbinata, 412; 
turgida, 168, 247, 258, 259, 261; 
umbellata, 411; uniflora, 412; 
varia, 301, 412; virgata, 250, 253; 
Walteri, 411; X Weaveri, 314, 
362 i 

Liliacea, 266 

Lilies, 102 

Liliiflorae, 267 

Limodorum praecox, 190, 191; tu- 
berosum, f. latifolium, 188, var. 
nanum, 189 

Limosella aquatica, 115, 116 

Lindernia anagallidea, 330 

Linum sulcatum, 115 

Liparis, 189 

Listera auriculata, 82 

Lolium, 26; multiflorum, 21, var. 
diminutum, 21; perenne, 21; 
temulentum, 21, var. leptochae- 
tum, 21 

Lonicera hirsuta, 202, 203 

Lophotocarpus calycinus, 330 

Ludwigia alternifolia, 330 

Luzula nemorosa, 82 

Lycopodium adpressum, 135, 136, 
f. polyclavatum, 136; alopecur- 
oides, 134, 135, ssp. adpressum, 
136, f. adpressum, 136, var. ad- 
pressum, 136, f. polyclavatum, 
136; Bigelovii, 134; carolinianum, 
134-136; Chapmani, 135, 136; 
inundatum, 134-136, Varieties 
of, 134, var. adpressum, 135, 136, 
f. polyclavatum, 136, y. alope- 
curoides, 134, var. appressum, 
136, 8. Bigelovii, 134, 135, var. 
Bigelovii, 135, 136, f. furcatum, 
136, f. polyclavatum, 136, var. 


robustum, 135, 136, f. furcatum, 
136 

Lycopus amplectens, 37; rubellus, 
330; sessilifolius, 37 

Lythrum alatum, 40, in Maine, 40 


Maine, Lythrum alatum in, 40 

Malus, 132 

Manilkara, 164; bahamensis, 164; 
emarginata, 164; Jaimiqui, 164; 
parvifolia, 164 

Manual Range, Notes on certain 
Plants in the Gray's, 89, pls. 
1021-1024; Some Orchids of the, 
161-162, 184-197, pls. 1045-1048 

Maple, 61 

Marsilea vestita, 115 

Mentha canadensis, 330 

Merrill, E. D., The Amos Eaton 
Herbarium, 201 

Mertensia paniculata, 163 

Mespilus Amelanchier, 128, 131, 
132; canadensis, 128 

Michigan, Observations on two 
Ecological Races of Allium tri- 
coccum in Kalamazoo County, 61 

Milium effusum, 81, 82 

Mimusops bahamensis, 164; emar- 
ginata, 164; floridana, 164; Jai- 
miqui, 164; parviflora, 164; par- 
vifolia, 164; Eieberi, 164 

Minnesota, Previously unreported 
Plants from, 81; Relict Boreal 
Plants in Southeastern, 163 

Molinia caerulea, 19 

Monarda fistulosa, var. typica, f. 
albescens, 97 

Monograph of Amelanchier (Re- 
view), A, 129 

Moore, J. W., Notes on Aquatic and 
Prairie Vegetation in Southwest- 
ern Minnesota, 113 

Mosses, 1; from Windsor, Nova 
Scotia, Some, 1 

Moyle, J. B., Relict Boreal Plants 
in Southeastern Minnesota, 163; 
Vegetation of Artificial Lakes in 
Northwestern Arkansas, 329 

Muhlenbergia, 63; glomerata, 64, 
var. cinnoides, 64; microsperma, 
64; racemosa, 63; setosa, 63, 64, 
an Untenable Name, 63, var. cin- 
noides, 64 

Myosurus minimus, 115 

Myriopteris gracilis, 386, 387 


Name Euphorbia maculata L., Ap- 
eas of the, 197; Muhlen- 
ergia setosa an Untenable, 63 


x Rhodora 


Najas guadalupensis, 329 

Nardosmia, 123; corymbosa, 124; 
frigida, 124, var. corymbosa, 124, 
var. genuina, 123, var. palmata, 
124; Hookeriana, 124; palmata, 
124; speciosa, 124 

Nardus stricta, 21 

Narrow-leaved wintergreen oak, 143 

Nasturtium officinale, 330 

Nelumbo pentapetala, 330 

Neottia gracilis, 6, 7, 190, 191, var. 
B. secunda, 5, 7, 9; lacera, 5-7; 
plantaginea, 6; Slender, 6; tor- 
tilis, 190, 191 

Nephrodium, 384, 385; lanosum, 
383-388 

New Combination in Chrysobala- 
nus, 136 

New England?, Does Habenaria 
cristata still grow in, 64 

New Hampshire, Betula glandulosa 
at a low Altitude in, 183; Plants, 
Notes on, 205 

Nielsen, E. L., Vegetation of Arti- 
ficial Lakes in Northwestern 
Arkansas, 329 

Ninth Report of the Committee on 
Plant Distribution, 17 

Nomen ambiguum, Populus bal- 
samifera of Linnaeus not a, 103 

Nomenclatural Transfers in Poly- 
gonum, 49-54 

North American Plants, Technical 
Studies on, 5-16, 27-40, 41-60, 
65-81, pls. 993-1020; Representa- 
tives of Alisma Plantago-aqua- 
tica, 86; Salix, Difficulties in, 13— 
16, 27—40, 41-49, pls. 995-1006 

North Carolina, Setaria Faberii in, 
391 

North Dakota, Botanical Visits to 
Forts Clark, Mandan and Union 
in, 98, pls. 1025, 1026 

Northeastern United States, III. 
Inuleae and Senecioneae, Notes 
on Compositae of the, 116 

Notes on Aquatic and Prairie Vege- 
tation in Southwestern Minne- 
sota, 113; certain Plants in the 
Gray's Manual Range, 89, pls. 
1021-1024; Compositae of the 
Northeastern United States, III. 
Inuleae and Senecioneae, 116; 
New Hampshire Plants, 205 

Nova Seotia, Some Mosses from 
Windsor, 1 

Novelties in Our Flora, 54—60, 65- 
81, pls. 1007-1020 


[DECEMBER . 


Number in Astragalus caryocarpus 
Fruits, Size, Shape and, 111 
Nymphoides peltatum, 330 


Oak, Darlington, 144; laurel, 138, 
145; narrow-leaved wintergreen, 
143; water, 143; white, 206 

Observations on two Ecological 
Races of Allium tricoccum in 
Kalamazoo County, Michigan, 61 

Ophrys aestivalis, 191; barbata, 11; 
Corallorhiza, 193 

Orchidaceae, 9, 193 

Orchids, 7; of the Manual Range, 
Some, 161-162, 184-197, pls. 
1045-1048 

Orchioides decipiens, 11; Menziesii, 
1 


1 

Orchis clavellata, 10, 161, 162, 215; 
tridentata, 162 

Oregon cliff-brake, 272 

Orthotrichum elegans, 3; sordidum, 
3; speciosum, 3 

Othake, 85; macrolepis, 84-86; 
Reverchonii, 86; robustum, 86; 
roseum, 86; texanum, 86 

Ownbey, Marion, Observations on 
two Ecological Races of Allium 
tricoccum in Kalamazoo County, 
Michigan, 61 


Palafoxia, 85; in Texas, Genus, 84; 
linearis, 84, 85; macrolepis, 86; 
Reverchonii, 86; riogranden- 
sis, 84, 85; rosea, 86, var. ro- 
busta, 86; sphacelata, 86 

Panicum, 212; subg. Dicanthelium, 
212; auburne, 37; verrucosum, 58 

Pappophorum mucronulatum, 19 

Parlin, John C., Lythrum alatum 
in Maine, 40 

Pedicularis canadensis, 59, 60, 80, 
pl. 1009, var. Dobbsii, 59, 60, 80, 
pls. 1009, 1010, var. fluviatilis, 60 

Penstemon albidus, 99 

Peramium decipiens, 11; Menziesii, 
11 

Persicaria fluitans, 49, 52; meso- 
chora, 52; oregana, 52; purpurata, 
52 


Petasites, 123; alaskanus, 123; 
corymbosus, 124; dentatus, 123; 
frigidus, 122-124, var. corymbo- 
sus, 123, 125, var. genuinus, 
123, 125, var. hyperboreoides, 
124, var. palmatus, 124, 125; 
gracilis, 123; Hookerianus, 124; 
hyperboreus, 124; nivalis, 124; 
palmatus. 122, 125, var. frigidus, 


1946] Index to Volume 48 xi 


124; sagittatus, 122, 123; speci- 
osus, 124, 125, var. frigidus, 124; 
trigonophyllus, 124; vitifolius, 
122, 124; Warrenii, 123 

Phascum cuspidatum, 3, 4, var. 
americanum, 3 

Phlox alyssifolia, 100 

Phragmites, 21; communis, var. 
Berlandieri, 19, 25, 26, 205 

Phymosia remota, 89-91, 94 

Pinus Banksiana, 37 

Piperia maritima, 10 

Plagiobotrys scopulorum, 115 

Plagiothecium laetum, 3 

Plant Distribution, Ninth Report 
of the Committee on, 17 

Plantago elongata, 116 

Plants from Minnesota, Previously 
unreported, 81; in the Gray's 
Manual Range, Notes on certain, 
89, pls. 1021-1024; in Southeast- 
ern Minnesota, Relict Boreal, 
163; Notes on New Hampshire, 
205; Technical Studies on North 
American, 5-16, 27-40, 41-60, 
65-81, pls. 993-1020 

Pleuridium subulatum, 2 

Poa alpigena, 19, 22, 23; alsodes, 19; 
angustifolia, 19, 22; annua, 19; 
Chaixii, 82; Chapmaniana, 19; 
compressa, 19; glauca, 19, 23; 
languida, 19, 24; laxa, 19, 23; 
nemoralis, 19, 23, 82, var. glau- 
cantha, 19; palustris, 19, 21, 22; 
pratensis, 19, 22; saltuensis, 19, 
21, 23, var. microlepis, 19, 23; 
sylvestris, 20, 81; trivialis, 20, 81, 
82 


Podosaemum setosum, 63 
Polygonum, Nomenclatural Trans- 
fers in, 49-54; amphibium, 49-51, 
var. aquaticum, 49, 50, var. 
Hartwrightii, 51, var. margina- 
tum, f. hirtuosum, 49, var. natans, 
50, 52, var. a. natans, 49, 50, var. 
stipulaceum, 50, 51, f. fluitans, 
49, f. hirtuosum, 49, f. simile, 
49, 51; amphibium, f. terrestre, 
51, var. terrestre, 51; arifolium, 
53, 54, var. lentiforme, 53, 54, var. 
pubescens, 53; cilinode, var. 
breve, 54; cilinode, f. erectum, 
54, var. erectum, 54; coccineum, 
330; fluitans, 49—52, 202; Hart- 
wrightii, 51; natans, 49, 50, 52, 
202, 203, f. genuinum, 51, f. 
Hartwrightii, 51; orientale, 51; 
Persicaria, 330; punctatum, 330; 


sagittatum, 53, 54, var. pubescens, 


Polypodium, 384, 385; fontanum, 
383; fragrans, 385; lanosum, 384— 
386 

Polypteris, 85 

Polytrichum formosum, 3; juni- 
perinum, var. alpestre, 3 

Poplar, Balsam, 103-109; Black, 106 

Populus, 42, 43; sect. Aigeiros, 106, 
sect. Tacamahaca, 104, 107; bal- 
samifera, 103-105, 107-109, of 
Linnaeus not à Nomen ambigu- 
um, 103; candicans, 104; delt- 
oides, 103, 104, 106, 107; foliis 
cordatis, etc., 104, 106, 108; foliis 
ovatis acutis, etc., 104, 108; foliis 
subcordatis, etc., 107; hetero- 
phylla, 106, 107; nigra, folio 
maximo, etc. 104, 106, 108; 
suaveolens, 107; Tacamahacca, 
103, 104, 107-109, var. lanceo- 
lata, 104 

Potamogeton, 70; Berchtoldi, var. 
tenuissimus, 329; foliosus, var. 
genuinus, 330; natans, 330; no- 
dosus, 330° 

Potentilla pennsylvanica, var. 
strigosa, 115 

Poterium sitchense, 12 

Pottia truncata, 4 

Prairie Vegetation in Southwestern 
Minnesota, Notes on Aquatic 
and, 113 

Presumable Identity of Cheilanthes 
lanosa, 383 

Previously unreported Plants from 
Minnesota, 81 

Psilocarya nitens, 37, 58 

Psilosanthus, 167 

Psoralea esculenta, 99 

Pteridophyta, 384 

Pteronia caroliniana, 394 

Ptilimnium capillaceum, 162 

Puccinellia, 22; distans, 20, var. 
angustifolia, 20; fasciculata, 20, 
25; maritima, 20, 25; Nuttalliana, 
20; paupercula, var. alaskana, 20, 
25 


Pyrola chlorantha, 163; secunda, 
163 
Pyrus, 132 


Quercus alba, 206; aquatica, 142, 
var., 214, var. hybrida, 142, g. 
laurifolia, 141, 8. nana, 144, var. 
nana, 144; dentata, 142, 143; 
hemisphaerica, 138-140, 142-144, 
214, pls. 1035, 1036, var. nana, 


xii Rhodora 


144; hybrida, 141, 142; laurifolia, 
137, 139-144, 214, pls. 1031-1034, 
The Identity of, 137-145, pls. 
1031-1036, a. acuta, 141, hybrida, 
140-142, var. hybrida, 139, 141, 
142, 214, pl. 1031, 8. obtusa, 140, 
141; laurina, 141; nana, 144; 
nigra, 141-144; obtusa, 140-142, 
214, var. obovatifolia, 140, 214; 
Phellos, 138, 144, 214, var. lauri- 
folia, 141; Prinus, 206; rhombica, 
138-141, 214, pls. 1031-1033, var. 
acuta, 139, var. obovatifolia, 138, 
140, 214, pl. 1034 


Races of Allium tricoccum in Kala- 
mazoo County, Michigan, Obser- 
vations on two Ecological, 61 

Rafinesque’s *Herbarium Rafines- 
quianum", Some Species in, 5-13, 
pls. 993, 994 

Red ash, 61 

Relict Boreal Plants in Southeast- 
ern Minnesota, 163 

Report of the Committee on Plant 

istribution, Ninth, 17 

Representatives of Alisma Plantago- 
aquatica, North American, 86 

(Review), A Monograph of Amelan- 
chier, 129 

Rhamnus alnifolia, 163, 205 

Rhus Vernix, 205 

Rhynchospora filifolia, 58 

Rosa “arkansana”’, 100; arkansana, 
116; blanda, 100; Woodsii, 100 

Rosaceae, 129, subfam. Pomoideae, 
130 

Roses, 100 

Rouleau, Ernest, Populus balsami- 
fera of Linnaeus not a Nomen 
ambiguum, 103 

Rubus, 129; pubescens, 163 

Rudbeckia, 79; fulgida, 75 


Sabatia difformis, 58 

Sagina virginica, 328 

Sagittaria ambigua, 330; graminea, 
330; latifolia, 330; rigida, 330 

Salices, 42 

Salix, Difficulties in North Ameri- 
can, 13-16, 27-40, 41-49, pls. 
995-1006; Adenophyllae, 15, 34, 
37,38; $ Árgyrocarpae, 44; Bon- 
plandianae, 43; Cordatae, 15, 34, 
37, 38; Herbaceae, 44; Nigrae, 43; 
Pentandrae, 43; Reticulatae, 41— 
43; $ Uva-ursi, 44; adenophylla, 
14, 28, 31-38, 48; alva, var. pam- 
eachiana, 38, var. vitellina X S. 


[DECEMBER 


fragilis, 38; alba X lucida, 38; 
anglorum, var. antiplasta, 44; 
angustata, 38; arctica, var. anti- 
plasta, 44; argyrocarpa, 44; Beb- 
biana, 43; Bonplandiana, 43; 
candida, 28; caroliniana, 28-31; 
conifera, 14; cordata, 14, 27, 28, 
30-38, 48, pls. 997-1000, var. 
&brasa, 34, 8. angustata, 38; cor- 
data angustata, 31, 1? forma dis- 
color, 31; cordata, f. mollis, 38; 
decipiens, 38; discolor, 27, 43, 
var. eriocephala, 27; eriocephala, 
27, 28, 31; fragilis, 38; glauco- 
phylla, 45, var. albovestita, 45; 
glaucophylloides, 45, var. albo- 
vestita, 45, var. glaucophylla, 45, 
f. lasioclada, 45; gracilis, 46-48, 
var. textoris, 46, 47; herbacea, 
44; Humboldtiana, 43; humilis, 
43, var. hyporhysa, 45, var. 
longifolia f. rigidiuscula, 45, var. 
microphylla, 46, f. curtifolia, 
46, f. festiva, 46, f. tortifolia, 
46, var. rigidiuscula, 45, var. 
tristis, 46; incana, 28; interior, 39, 
var. exterior, 38, 39; interior, f. 
Wheeleri, 39; jejuna, 41, 43, 49, 
pl. 1006; x. Jesupi, 38; leiolepis, 
41, 43, 49, pl. 1005; longifolia, 39; 
longipes, 29-31, var. Wardi, 29; 
longirostris, 28; lucida, 43; mis- 
souriensis, 27, 28; nigra, 28-31, 
43, var. Wardi, 30; nivalis, 43; 
occidentalis, 30; pameachiana, 
38; pentandra, 28, 29, 43; petio- 
laris, 46-48, var. angustifolia, 46, 
var. rosmarinoides, 46; Pitcheri- 
ana, 31; reticulata, 39, 41-43, 49, 
pls. 1003, 1004, var. semicalva, 
39, 41-43, 49, pls. 1003, 1004; 
rigida, 32-34, 38, 43, 48, pls. 995, 
996, var. angustata, 38; rigida, f. 
mollis, 38; syrticola, 14, 31, 34- 
37, 48, pls. 1001, 1002; triandra, 
29, 30; tristis, 14, 28, 46, f. curti- 
folia, 46, f. festiva, 46; tristis mi- 
crophylla, 46; Uva-ursi, 44; ves- 
tita, 40-43, f. mensalis, 42, var. 
silophylla, 41; vitellina, 38; 
ardi, 29-31 
Sanguisorba canadensis, £8. lati- : 
folia, 12; latifolia, 12; officinalis, 
12; sitchensis, 12; stipulata, 12 
Sapotaceae, 163 
Sarracenia heterophylla, 202, 203 


Schedonnardus, 114; paniculatus, 
114; texanus, 114 


1946] - 


Schizachne purpurascens, 20, f. al- 
bicans, 20 

Scirpus pallidus, 330; Rollandii, 271 

Scleropoa rigida, 20 

Scutellaria parvula, 116 

Secale cereale, 21 

Second Station for Corydalis flavula 
in Connecticut, 326 

Senecio alabamensis, 330; tomen- 
tosus, 331, forma alabamensis, 
330 

Senecioneae, Notes on Compositae 
of the Northeastern nited 
States, III., Inuleae and, 116 

Serapias radicibus palmato-fibrosis, 
etc., 186 

Serratula, 242, 284; pilosa, 250, 257; 
seariosa, 294, 298, 300, 312; 
speciosa, 340; spicata, 178, 220, 
221, 250, 256; squarrosa, 298, 394 

Serviceberry, 127 

Setaria, 392; Faberii, 391, in North 
Carolina, 391 

Shape and Number in Astragalus 
caryocarpus Fruits, Size, 111 

Sherff, Earl Edward, Notes on cer- 
tain Plants in the Gray’s Manual 
Range, 89, pls. 1021-1024 

Shortia dentata, 208 

Sisymbrium dentatum, 208 

Sisyrinchium, 152, 153, 156; anceps, 
152, 153, 157, 158, 215; angusti- 
folium, 152-158, 215, pls. 1042, 
1043, The Identity of, 152-160, 
pls. 1042-1044; Bermudiana, 152- 
154, 156, 158, var. œ., 155, 156, 
var. anceps, 152, var. mucro- 
natum, 152; caeruleum parvum, 
etc., 155-157, 215; capillare, 156; 
foliis lineari-gladiolatis, ete., 154, 
158; gramineum, 152, 153, 155, 
156, 158, 215; graminoides, 155, 
157, 158, 215; minus, 157; mon- 
tanum, 152, 157, 158, 215, pl. 
1044, var. crebrum, 159, 215, 
pls. 1043, 1044; mucronatum, 
152, 155, 150, 215, pl. 1042; 
septentrionale, 158 

Size, Shape and Number in Astra- 
galus caryocarpus Fruits, 111 

Slender Neottia, 6 

Sloanea emarginata, 164 

Smith, Lyman B., New Combina- 
tion in Chrysobalanus, 136 

Solidago, $ Euthamia, 65; gramini- 
folia, var. Nuttallii, 66, 81, pl. 
1012, X S. microcephala, 65; X 
hirtipes, 65, 66, 81, pl. 1011; mi- 


Index to Volume 48 xiii 


crocephala, 65, 66, 81, pl. 1012; 
Victorinii, 271 

Some Mosses from Windsor, Nova 
Scotia, 1; Orchids of the Manual 
Range, 161-162, 184-197, pls. 
1045-1048; Species in Rafines- 
que’s “Herbarium Rafinesqui- 
anum", 5-13, pls. 993, 994; 
Trivial American Forms of the 
Lady-fern, 389 

Southeastern Minnesota, Relict 
Boreal Plants in, 163 

Spartina, 21; alterniflora, 17, 25, 
var. pilosa, 17, 25; caespitosa, 17, 
25; cynosuroides, 17, 25; patens, 
17, 25, var. juncea, 17, 25; pec- 
tinata, 18, 25, 26, var. Suttiei, 18 

Species, Amelanchier spicata not an 
American, 125 

Specularia, 209-214; coloradoensis, 
216, pl. 1050; falcata, 211; 
Holzingeri, 214; hybrida, 210, 
212, 213, 216, pl. 1049; lampro- 
sperma, 214; speculum, 210; 
Speculum, 8. libanensis, 213, 8. 
racemosa, 211, var. racemosa, 
211; Speculum-Veneris, 210, 211, 
213, 215, pl. 1049; texana, 214 

Sphaeralcea acerifolia, 89; coccinea, 
99; remota, 90 

Spiraea latifolia, 112, var. septen- 
trionalis, 112, in Virginia, 112 

Spiranthes, 6; aestivalis, 191; Beckii, 
6, 9, 10, 189-191; cernua, 191, 
var. parviflora, 5; decipiens, 11; 
gracilis, 6-9, 13, 190, 191, pl. 994; 
Grayi, 6, 189, 192; lacera, 5-9, 13, 
190, 191, pl. 993; montana, 5, 9; 
ovalis, 5, 9; parviflora, 5; planta- 
ginea, 6; praecox, 190, 191; sim- 
plex, 6, 10, 189, 192; Smallii, 6; 
tuberosa, 6, 10, 189, 191, 192, var. 
Grayi, 189, 192; vernalis, 191 

Sporadic Appearance of Epipactis 
Helleborine, 88 

Stachys hyssopifolia, 37 

Staehelina elegans, 340 

Station for Corydalis flavula in Con- 
necticut, Second, 326 

Stenanthium, 148, in the Eastern 
United States, 148-152, pls. 1037— 
1041; angustifolium, 149, 151, *S. 
gramineum, 151; gramineum, 148- 
151, 214, pls. 1037, 1038, var. 
micranthum, 148, 150-152, 215, 
pl. 1041, var. robustum, 148, 151, 
152, 214, 215, pls. 1039, 1040, var. 
typicum, 151, pls. 1037, 1038; 
robustum, 148-150, 152, 214 


xiv Rhodora 


Stevens, O. A. Botanical Visits to 
Forts Clark, Mandan and Union 
in North Dakota, 98, pls. 1025, 
1026; Size, Shape and Number in 
Astragalus caryocarpus Fruits, 
111 

Stevia sphacelata, 86 

Stipa spartea, 116 

Stomoisia virgatula, 60 

Studies on North American Plants, 
Technical, 5-16, 27-40, 41-60, 
65-81, pls. 993-1020 

Suprago, 167; sphaerocephala, 313 

Swamp birch, 61 

Swimming knotweed, 52 

Symphoricarpos albus, 133 


Tacamahaca foliis crenatis, 107 

'Tamarack, 61 

Technical Studies on North Ameri- 
can Plants, 5-16, 27—40, 41-60, 
65-81, pls. 993-1020 

Texas, Genus Palafoxia in, 84 

Thaspia trifoliata, 162 

Thaspium trifoliatum, 162 

Thermopsis rhombifolia, 102 

Thuja occidentalis, 197 

Tillaea aquatica, 116 

Trilisa, 176; odoratissima, 410; 
paniculata, 410 

Triodanis, 209-213, versus Specu- 
laria, 209—214, pls. 1049, 1050; 
biflora, 212; coloradoensis, 209- 
213, 216, pl. 1050; falcata, 213; 
Holzingeri, 212-214; lampro- 
sperma, 214; perfoliata, 209, 213; 
texana, 212, 214 

Triodia flava, 20, 24 

Triorchis Beckii, 6; Grayi, 6; ovalis, 
6 


Triplasis purpurea, 20, 25 

Triticum aestivum, 21 

Trivial American Forms of the 
Lady-fern, Some, 389 

Tryon, Jr., R. M., Notes on Aqua- 
tic and Prairie Vegetation in 
Southwestern Minnesota, 113 

Tussaca oblongifolia, 11 

Tussilago corymbosa, 123, 124; 
frigida, 124; palmata, 124 

Two Ecological Races of Allium 
tricoccum in Kalamazoo County, 
Michigan, Observations on, 61 

Typha latifolia, 330 


Umbelliferae, 162 

United States, III. Inuleae and 
Senecioneae, Notes on Composi- 
tae of the Northeastern, 116; 


[DECEMBER 


Stenanthium in the Eastern, 148- 
152, pls. 1037-1041 

Unreported Plants from Minnesota, 
Previously, 81 

Untenable Name, Muhlenbergia 
setosa an, 63 

Utricularia, 211, 212; biflora, 330, 
juncea, 60, f. minima, 60, f. vir- 
gatula, 60; subulata, 60, f. 
cleistogama, 60; virgatula, 60 


Valerianella Chenopodifolia, 96 

Varieties of Lycopodium inunda- 
tum, 134 

Vegetation in Southwestern Min- 
nesota, Notes on Aquatic and 
Prairie, 113; of Artificial Lakes 
in Northwestern Arkansas, 329 

Veratrum angustifolium, 149, 151 

Verbascum Phlomoides, 97; Thap- 
sus, 97 

Verbena simplex, 115, 116 

Vermont, Dicentra eximia in, 272 

Vernonia angustifolia, 411; hirsuti- 
flora, 395 

Victorin, Frére Marie-, 265 

Viola, 212 

Virginia, Does Bartonia verna grow 
in?, 327; Spiraea latifolia, var. 
septentrionalis in, 112 

Visits to Forts Clark, Mandan and 
Union in North Dakota, Botani- 
cal, 98, pls. 1025, 1026 

Vulpia 21, megalura, 20; myurus, 
20; octoflora, var. tenella, 20, 24 


Water Oak, 143 
Weatherby, C. A., Impatiens Roylei 
versus I. glandulifera, 412 
Weisia microstoma, 3, 4; viridula, 3 
White elm, 61; oak, 206 
Wild dilly, 164 
Wilde, Charlotte Endicott, Betula 
landulosa at a low Altitude in 
ew Hampshire, 183 
Willows, 14, 29, 42, 47 
Windsor, Nova Scotia, Some Mosses 
from, 1 
Witchgrass, 22 
Wood, Carroll E., Jr., Setaria 
Faberii in North Carolina, 391 


Xanthium, § Acanthoxanthium, 73; 
§ Euxanthium, 73; ambrosioides, 
74; canadense, 72; Chasei, 66-70, 
72, 73, 81, pls. 1013, 1014; chin- 
ense, 67, 68, 70, 72, 81, pl. 1015; 
curvescens, 68, 69, 81, pl. 1016; 
echinatum, 67, 69, 71, 73, 81, pl. 


3 1753 00341 3694 
1946] Index to Volume 48 XV 


1018; globosum, 66, 67, 72, 81, pl. 
1015; inaequilaterum, 73; in- 
dicum, 73; inflexum, 68, 81, pl. 


Xerophyllum gramineum, 151 
Xylosteum solonis, 202, 203 
Xyris, 58; § Brevifoliae, 56, 57; am- 


1015; italicum, 67, 70, 81, pl. 
1015; leptocarpum, 68, 69, 81, 
pl. 1016; macrocarpum, 81; ori- 
entale, 68-70, 81, pl. 1017; ovi- 
forme, 67; pensylvanicum, 70, 72; 
Roxburghii, 73; speciosum, 70, 
73; spinosum, 71-74; strumarium, 
66, 67, 70, 71, 73, 74, 81, pl. 
1013; varians, 69, 81, pl. 1019; 
Wootoni, 70 


bigua, 58; Bayardi, 56-58, 80, 
pl. 1007; brevifolia, 56, 57, 80, 
pl. 1008; caroliniana, 58; Drum- 
mondii, 57; flabelliformis, 57, 80, 
pl. 1008 


Younge, O. R., Vegetation of Arti- 


fieial Lakes in Northwestern 
Arkansas, 329