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ANNALS
OF THE
MISSOURI BOTANICAL GARDEN
b^. NECS "T E48
Annals
of the
Missouri Botanical
Garden
Volume XX
1933
With Twenty-three Plates and Forty-three Figures
Published quarterly at 8 West King Street, Lancaster, Pa., Kid the Board of Trustees
of the Missouri Botanical Garden, St. Louis, Mo
Entered as second-class matter at the post-office at Lancaster, Pennsylvania,
under the Act of March 3, 1879.
549006
Annals
of the
Missouri Botanical Garden
A Quarterly Journal containing Scientific Contributions
from the Missouri Botanical Garden and the Graduate Labora-
tory of the Henry Shaw School of Botany of Washington Uni-
versity in affiliation with the Missouri Botanical Garden.
Information
The Annals of the Missouri Botanical Garden appears four times during
the calendar year: February, April, September, and November. Four numbers
constitute a volume.
Subscription Price — — — $6.00 per volume
Single Numbers - — — — 1.50 each
Contents of previous issues of the Annals of the Missouri Botanical Garden
are listed in the Agricultural Index, published by the H. W. Wilson Co.
STAFF
OF THE MISSOURI BOTANICAL GARDEN
GEORGE T. Moore.
Assistant to the Director,
KATHERINE H. LEIGE.
HERMANN VON SCHRENE, CARROLL W. DoDGE,
Pathologist. Mycologist
Jesse M. GREENMAN, RoBERT E. WOODSON, JR.,
Curator of de Herbarium. Research Assistant.
EnNEsT S. REYNOLDs, NEeLL C. H
Physiologist. Con Ea dn "Editor of Publications.
BOARD OF TRUSTEES
OF THE MISSOURI BOTANICAL GARDEN
President,
GEORGE C. Hrrcncock.
Vice-President,
SAMUEL C. Davis.
Second Vice-President,
DANIEL K, CATLIN.
L. RAY CARTER. ALBERT T. PERKINS.
Tuomas S. Marrirr. EUGENE PETTUS.
GEORGE T. Moore. ETHAN A. H. SHEPLEY.
FRED G. ZEIBIG.
EX-OFFICIO MEMBERS:
GEORGE R. THR BERNARD F. Dic
Chancellor oe n: University. Mayor of the e p St. Louis,
WILLIAM SCARLETT ALBERT KUNTZ
Bishop of the Diodi of Missouri. President of The Academy of Sci-
ence of St. Louis.
Davip C. Topp
President of the Board of Education of St. Louis.
GEORGE F. HEFFERNAN, Secretary.
TABLE OF CONTENTS
A New Yellow Nymphaea from Tropical Africa
George H. Pring and Robert E. Woodson, Jr
ee om m n
Contribution to the Lichen Flora of North
Lo 0 oe oo kh Wes ee OR ale Veli J. P. B. Räsänen
Astranthium and Related Genera....Esther L. Larsen
Ferns and Fern Allies of Missouri...........
M. Hlizabeth Pinkerton
$9 wx € & «o9 WLW E Row Ee wow $e «9
Blastomycosis: Report of a Case, with a Study
of an Etiologie Factor and a Classification of
aue Organism. ........... cane eee Morris Moore
A Monograph of the American Species of the
Genus Halenis...........-22 ee Caroline K. Allen
The Sensitivity of Orchid Seedlings to Nutri-
tional Ions :.......4.66..5543 eee F. L. Wynd
A Revision of the North American Species of
Di 5.0 MMRMCPRENNRUNENEENRO (s son Lily M. Perry
Nutrient Solutions for Orchids. ........ F. Lyle Wynd
The Foliose and Fruticose Lichens of Costa
BE E Sees es on eS ERN Carroll W. Dodge
A Neutral (?) Strain of Mucor sphaerosporus
from Missouri.......... Sa Morris Moore
A Study of Hndomyces capsulatus Rewbridge,
Dodge and Ayers: A Causative Agent of
Fatal Cerebrospinal Meningitis....... Morris Moore
Sources of Carbohydrate for Germination and
Growth of Orchid Seedlings........ F. Lyle Wynd..
79-118
119-222
223—231
239-362
363-372
373—407
469
471—568
569-581
Super Optimal and Thermal Death Tempera-
tures of the Cotton Plant as Affected by
Variations in Relative Humidity..........
.... Dorothy Megowen Berkley and Earl E. Berkley
Studies in the Apocynaceae. IV. The Amer-
ican Genera of Echitoideae. . Robert E. Woodson, Jr.
Report of a Botanical Expedition into the
Mountains of Western Texas.............
nee Julian A. Steyermark and John Adam Moore
General Index to Volume XX...................
PAGE
583-604
605—790
791-8006
807—817
Annals
of the
Missouri Botanical Garden
Vor. 20 FEBRUARY, 1933 No. 1
A NEW YELLOW NYMPHAEA FROM TROPICAL AFRICA!
GEORGE H. PRING
Superintendent, Missouri Botanical Garden
ROBERT E. WOODSON, Jr.
Research Assistant, Missouri Botanical Garden
Instructor in Botany, Henry Shaw School of Botany of Washington University
Nymphaea ($ Brachyceras) Burttii Pring & Woodson, spec.
nov., foliis magnis longe-petiolatis orbiculare-sagittatis margine
plus minusve conspicue undulato-sinuatis apice late obtusis vel
rotundatis sinu profundo auriculis leviter divergentibus obtusis
25-35 cm. longis subcoriaceis utrinque viridibus vel saepius
purpureo-maculatis glaberrimis vel umbilico paulo papillato
subtus nervis manifestis sed vix prominentibus; floribus speci-
osissimis primulino-flavis 15-18 cm. diametro metientibus,
sepalis ovato-lanceolatis acutiusculis 5-8 em. longis 2-3 cm. latis
dilute viridibus immaculatis laevibus, petalis ca. 20-23 anguste
ellipticis apice acuminatis vel anguste acutis medio versus grada-
tim angustatis plerisque 5-nervatis exterioribus quam sepalis
paulo brevioribus ca. 4.5-7.0 em. longis 1.0-2.5 cm. latis, stami-
nibus 190-200 linearibus connectivo manifeste elongato basi
paulo ampliato exterioribus ca. 5 cm. longis dilute cadmio-flavis,
carpellis 28-30 stylo linearo profunde inclinato stigmate concavo;
fructu minore 3.5—4.0 cm. diametro metiente, seminibus ovoideo-
oblongoideis apice minutissime apiculatis longitudine leviter punc-
ticulato-striatis griseo-brunneis maximo ca. 0.075 cm. minimo
ca. 0.05 cm. diametro metientibus.—Cult. Missouri Botanical
Garden, Aug., 1930, G. H. Pring s. n. (Herb. Mo. Bot. Garden,
TYPE).
Leaves large, long-petiolate, orbicular-sagittate, margin more
or less conspicuously undulate-sinuate, apex broadly obtuse or
rotund, sinus relatively deep and narrow, auricles obtuse, slightly
1 Issued April 29, 1933.
ANN. Mo. Bor. Garb., Vor. 20, 1933. (1)
[Vor. 20
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN
divergent, 25-35 cm. long, subcoriaceous, either surface green, or
occasionally somewhat purplish-maculate, particularly when
young, glabrous, or the umbilicus slightly papillate, the nerves
manifest but not prominent beneath; flowers showy, primrose-
yellow, very fragrant, 15-18 cm. in diameter; sepals ovate-
lanceolate, acute, 5-8 cm. long, 2-3 cm. broad, pale green, without
purple spots, smooth; petals about 20-23, narrowly elliptic, apex
Fig. 1. Nymphaea Burttii Pring & Woodson: S., sepals; P., petals; St., stamens;
S.L., Au ee leaves; F. L., floating leaves.
acuminate to narrowly acute, the base gradually narrowed from
about the middle, 5-nerved, the exterior somewhat shorter than
the sepals, usually 4.5-7.0 em. long, 1.0-2.5 em. broad; stamens
190-200, linear, the exterior about 5 cm. long, cadmium-yellow,
the connective manifestly elongate, somewhat broadened at the
base; carpels 28-30, the style linear, sharply inclined, the stigma
concave; fruit relatively small, about 3.5—4.0 cm. in diameter;
seeds ovoid-oblongoid, minutely apiculate, rather inconspicuously
1933]
PRING AND WOODSON—A NEW YELLOW NYMPHAEA 3
puncticulate-striate longitudinally, about 0.075 cm. long, 0.05 cm.
broad, grayish-brown.—Cultivated at the Missouri Botanical
Garden, Aug., 1930, G. H. Pring s. n. (Herb. Mo. Bot. Garden,
TYPE).
Only two yellow tropical Nymphaeas have previously been
described from Africa: N. sulphurea Gilg and N. Stuhlmannit
Schwfth. & Gilg. The former differs from N. Burttii chiefly in
the smaller flowers (4-7 cm. in diam.) and leaves (4.5-5.5 cm.
long), which are almost exactly orbicular-cordate in outline,
purplish-maculate sepals, and more elongate tubers. N. Stuhl-
mannii is easily distinguishable from N. Burttit by the somewhat
smaller flowers (10-15 cm. in diam.), with broader, obovate,
obtuse or rounded petals, and smaller (21-25 cm. long), entire
leaves, which are broadly ovate-cordate in outline, with broad,
rounded, regular lobes. Furthermore, the venation of the leaves
of N. Stuhlmannii is extremely verrucose beneath. The type
specimen of N. Stuhlmannii has not been available to the writers
for personal examination, but has been compared with & duplicate
specimen of N. Burttit by Dr. H. Melchior, of the Botanical
Museum at Berlin-Dahlem, who kindly affirmed the distinction
of either species. A photograph of the type specimen of N.
Stuhlmannit (Stuhlmann 410 in Hb. Berol.), generously provided
by Dr. L. Diels, Director of the Botanical Garden and Museum
at Berlin-Dahlem, has been deposited in the herbarium of the
Missouri Botanical Garden.
After a search of over ten years, the director of the Missouri
Botanical Garden obtained in September, 1929, a seed-pod of
what was presumed to be Nymphaea Stuhlmannii through the
personal efforts of Mr. B. D. Burtt, Esq., botanist for the Tsetse
Research Bureau, Kondoa, Tanganyika Territory. An excerpt
from Mr. Burtt’s notes is quoted.
“The seed was collected from plants growing in a seasonal rain-pond in the
Sambala ‘Mbuga’ seasonal swamp, the plants having spectacular yellow flowers
8 inches in diameter and sweet scented. The flowers float on the surface of
the water and on examination were found to contain dead bees (Apis melifera
[?]) that were imprisoned by the anthers over the stigmatic surface of the flowers.
The plants were collected on May 19, 1929, the seed from the same locality on
July 15, 1929. Other plants were collected March 16, 1929, from a seasonal
rain pond near Salia, Kondoa Distr. I have observed the plant in seasonal
rain ponds at Magungila, Wembare Steppe in 1928, also near Lilbilin, Massai
Land, in 1927."
[Vor. 20, 1933]
4 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The single, small seed-pod, in somewhat immature condition,
was received at the Garden on September 19, 1929. It was im-
mediately cleaned, and the many seeds planted in the greenhouse
water-lily tanks. Three weeks later a single seed germinated
from the lot, producing its first primrose-yellow flower on June 17,
1930. On July 30, the plant was removed from its pot and plant-
ed in the pond out of doors where it continued to bloom until
frost.
'The successful introduction of a yellow tropical water-lily has
opened an entirely new field for the hybridist. For many years
the available colors in the Brachyceras group have been limited
to blue and pink, and more recently to white through the intro-
duction from the Missouri Botanical Garden of the hybrid
“Mrs. G. H. Pring.” During the past three seasons 250 pol-
linations have been made with the pollen of N. Burttii, resulting
in many variable forms of commercial interest. The first season
(1930) was devoted to selfing the yellow-flowered species, and
many fertile seed-pods resulted. The following year twenty
specimens were grown from seed and planted in the ponds out of
doors. No variation in the color of the flowers was observed, but
a variable character was evident in the peduncle and petioles,
some being pure green, while about an equal number were brownish.
An unusual character, which appears to be dominant in all the
plants, is a peculiar twisting of the peduncle, the day before the
flower opens, in such a manner as to submerge the bud. The
following morning, however, the peduncle straightens, holding
the open flower erect in a natural position. This action has also
been observed in several hybrids.
As a propagator, Nymphaea Burttii is by far the poorest of any
grown at the Garden. Of the fifty propagating tubers secured
from pot plants during the past two seasons, only one has pro-
duced growth up to the present, despite the fact that they have
been in the heated propagating tanks for a period of six months.
Since it is an extremely poor propagator from tubers and extreme
heat is necessary for growth, it is very doubtful if it will find a
place in garden pools. On the other hand, the hybrids derived
from it, particularly the “Saint Louis," are both readily propa-
gated from tubers and suitable for cooler water.
[Vor. 20, 1933]
ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 1
Leaf, bud, and flower of Nymphaea Burttii Pring & Woodson, X 2$.
del. A. A. Heinze
ANN. Mo. Bor. GARD., Vor. 20, 1933 PLATE d)
i
WOODSON—A NEW YELLOW NYMPHAEA
PRING 4
CONTRIBUTION TO THE LICHEN FLORA OF
NORTH AMERICA!
VELI J. P. B. RASANEN
Instructor in the Agricultural School, Kurkijoki, Finnland
The lichens here described have been collected in the northern
section of North America. Of these specimens, 18 are Alaskan,
60 from eastern Canada (New Brunswick, except one from
Quebec), approximately 150 from western Canada (British
Columbia and Alberta), and a few from the state of Wyoming.
The Alaskan lichens, 15 species in all, are largely the usual
arctic ground lichens. The New Brunswick lichens, which com-
prise 46 species, come from around Dalhousie and were collected
in the year 1930 by the agronomist Tapio Reijonen. Usnea
longissima was collected near Franklin, Quebec, by Dr. Viljo
Kujala in 1931 during his scientific journey. On this same
journey, he collected in western Canada (120 species, varieties,
and forms) and in Wyoming (6 species). The latter specimens
come from the sandstone in a cactus desert.
The lichens from western Canada were collected by Dr.
Viljo Kujala in various types of forest in British Columbia in the
following localities: Jackman, Longworth, Aleza Lake, Prince
George, Six Mile Lake (Trout Lake), Hazleton, Dorreen, Co-
wichan Lake (Vancouver Island), Kamloops, Fish Lake (near
Kamloops), Blue River, Valemount, Golden, Field, and in Al-
berta in the vicinity of Lake Louise. The desert lichens come
principally from the vicinity of Kamloops in British Columbia
and from Wainwright in Alberta.
Although the same species have been encountered in several
different localities, the total number of species, varieties, and
forms is 171 with 5 entirely new species, as follows: Usnea Kujalae,
Placodium xanthostigmoideum, Nephromium canadense, Diplo-
schistes canadensis, and Sphaerophorus Tuckermaniv.
The list further includes eleven new varieties and forms, as
follows: Usnea comosa var. stuppea, Alectoria sarmentosa var.
gigantea, A. Fremontu f. perfertilis, Cetraria nigricans var. pallida,
! Duplicates of most of the specimens have been deposited in the herbarium of the
Missouri Botanical Garden.
Issued April 29, 1933.
ANN. Mo. Bor. Garp., Vor. 20, 1933. (7)
[Vor. 20
8 ANNALS OF THE MISSOURI BOTANICAL GARDEN
C. tenuifolia var. reticulata, C. tenuifolia var. pseudoislandica f.
septentrionalis, C. juniperina var. canadensis, C. juniperina var.
crispata, Parmelia elegantula var. americana, Cyanisticta Hookeri
var. septentrionalis, M ycoblastus sanguinarius var. Dodgeanus.
Species new to North America or otherwise noteworthy include
the following: Usnea lapponica, U. dasypoga, U. caucasica, U.
prostrata, U. rugulosa, U. similis, Alectoria Fremontii subsp.
olivacea, A. jubata var. Vrangiana, Ramalina Roesleri, Ochrolechia
upsaliensis, Pertusaria xanthostoma, Physcia pyrithrocardia, P.
muscigena f. squarrosa, P. endophoenicea, Peltigera Maurizüi, P.
canina var. suomensis, P. lepidophora, P. Nylanderi, Cetraria
Merrillii, Parmelia Delavayi, Squamaria alphoplaca, Nephromium
subparile, Lobaria oregana, Diploschistes bryophiloides.
The lichens collected by Dr. Kujala are principally epiphytic
and soil-lichens from a variety of forest types, although a few of
the soil lichens come from fields or desert. It might be inter-
esting to note here that in regions as widely separated and isolated
as the North American prairies and the south Russian and Asiatic
steppes and deserts there appear many identical, or, at least
similar, lichen forms living on the ground. Such steppe lichens
are: Parmelia vagans (appearing also in south Russia and Asia),
Physcia muscigena (many forms, a few also growing in the
mountains), Acarospora Schleicheri (appears in the Asiatic, North-
African, and Russian steppes, in addition to the European Alps
and Pyrenees),! Psora decipiens (also in deserts and mountains),
Diploschistes canadensis (the similar Russian D. scruposus var.
terrestris), Ochrolechia upsaliensis (sporadically found outside of
the steppes), Cladonia cariosa var. cribrosa (particularly on other
mineral substrates, slightly calciphile).
As the lichens in this collection were not collected by a lichen-
ologist, the lichen flora of the various localities is very poorly
represented and poor in species. They are, however, of particular
interest to the European investigator on account of the many
similar forms. The typical European Cetraria juniperina and
C. islandica do not appear at all amongst the American lichen
specimens (the latter also being absent from the ''Cetraria
islandica" specimens collected in Tierra del Fuego by Professor
Auer's expedition in 1929).
! Magnusson, Monogr. Acarospora. p. 395, 1929.
1933]
RASANEN—CONTRIBUTION TO LICHEN FLORA OF AMERICA 9
As the lichens from the various localities have not been suffi-
ciently collected, conclusions as to the relationship between the
American and European species are still hypothetical, particularly
as the present systematic revision of lichens necessitates many
corrections of the earlier determinations. We can, however, state
that as far as our present knowledge extends a great number of
identical lichen forms appear both in North America and Eurasia,
although the quantitative relationships between the different
species differ considerably on the two sides of the Atlantic.
I wish to acknowledge with thanks Dr. C. W. Dodge, My-
cologist to the Missouri Botanical Garden, who has kindly trans-
lated my manuscript from German to English, and T. Hidén, of
Helsinski, Finnland, who has gone over the latin text of this work.
UsNEA DASYPOGA (Ach. Röhl. New Brunswick: Dalhousie; British Columbia:
Prince George and Six Mile Lake, on Picea.
USNEA DASYPOGA Var. SUBSCABRATA Vain. British Columbia: Hazleton, on
Betula and Pinus. Sterile.
SNEA CAUCASICA Vain. Thallus prostratus, glauco-viridis, medulla sublaxa,
K—.—British Columbia: Six Mile Lake, on Picea. Fertile.
USNEA PROSTRATA (Vain.) Ras. British Columbia: Aleza Lake, on Picea. Sterile.
UsNEA RUGULOSA Vain. British Columbia: Hazleton, on trunks of Pinus Murray-
ana. Sterile
USNEA LONGISSIMA Ach. Quebec: north shore of the St. Lawrence Bay, Franklin,
especially on branches of Picea in a mixed forest; British Columbia: Vancouver,
Catillano Canyon, on Thuja in a forest with abundant ferns. Sterile.
Usnea comosa (Ach.) Vain. Thallus erectus, parce sorediosus, soredia isidiosa;
medulla crebra, K —.—New Brunswick: Dalhousie, on pue British Columbia:
Hazleton, Xn on branches of Pinus Murrayana. Steri
SNEA COMOSA var. stuppea Räs., var. nov. Thallus erectus aut suberectus,
brevior fruticulosus, 5-7 cm. longus, laevigatus vel leviter verrucosus, sorediosus,
pallido-stramineus; soredia maculiformia, demum parce isidiosa. Medulla laxa,
stuppea, K—.—British Columbia: Hazleton, iba pin on small, half-dried
Pinus, Picea, and Betula, on a sandy heath. Steri
Usnea simitis Motyka
U. subcomosa Vain. in Kgl. Danske Vidensk. Selsk. Skrifter, Nat. og Math.
Afd. VIII. 6: 392 (110) 1924.
With the above species, occasional in Hazleton. Sterile
UsNEA LAPPONICA Vain. With the above species, Hazleton, on Pinus. Sterile.
UsNEA HIRTA (L.) Motyka. pae Columbia: Aleza Lake, occasional on Picea
canadensis and P. Engelmannii. i
UsNEA Kujalae! Räs., sp. nov. Thallus curtus, idum fruticulosus, 3-5 cm.
longus, bene ramosus, lacteo-stramineus vel albo-viridis, sorediosus. Rami laevigati,
sine verruculis nervisque. Medulla laxissima, K—. borate rara, terminalia,
2-3 mm. lata, albido-straminea, fibrillosa, pinni: Sporae 9-10 X 5.3-7 u, ovoideo-
! In honor of Dr. Viljo Kujala.
[Vor. 20
10 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ellipsoideae, 8 : nae, monostichae. Asci cylindrici. Hymenium 80 u crassum, I+,
caerulescens. Subsimilis Usneae sorediiferae Motyka, sed thallus albidus et parce
sorediosus.— British Columbia: Hazleton, abundant on branches of Pinus Murray-
ana. Rarely fertile.
ALECTORIA SARMENTOSA Ach. Thallus elongatus, pendulus vel prostratus, K =
K(Cl) + leviter roseus, demum ferrugineus, I= leviter caerulescens. Sporae
normaliter binae vel 4:nae, fuscae.—British Columbia: Aleza Lake, frequent on
branches of Abies lasiocarpa, Picea canadensis, and P. Engelmannii; Longworth,
frequent on branches of Thuja; Hazleton, rarely on Pinus. Fertile.
ALECTORIA SARMENTOSA var. gigantea Riis., var. nov. Thallus elongatus, pendu-
lus vel prostratus, I em. longus, albidovetrami neus, parce pseudocyphellatus,
esorediatus, K=, K(Cl)=, Cl, I= leviter caerulescens. Apothecia vulgaria,
plana vel concava, atro-fuece vel livido-fusca, nuda vel leviter pruinosa a
normaliter 3:nae, 25-31 X 14.5-20 y, ellipsoideae, fusco-nigrae. Hymenium 130 pu
crassum, I + UN —British Columbia: Longworth, frequent on Thuja;
Dorreen, on Pseudots
marron LAETA (Tayl) Linds.
A. japonica Tuck., A. osteina Nyl., A. lata DR. (Arkiv f. Bot. 20": 24. 1926).
Thallus singes vel prostratus, osteo-ochroleucus, apicem versus concolor,
K= passim obsolete intensive fulvescens, K(Cl)-r roseus.—British Columbia:
Hazleton, occasional on the ground; Alaska, occasional on the ground. Sterile.
ALECTORIA OCHROLEUCA (Ehrh.) N Thallus pro parte ochroleucus, majore
parte viridi-niger, K= vel basim versus obsolete + dilute fulvescens, K(Cl) =.
Alaska, on the ground. Sterile.
ALECTORIA IMPLEXA (Hoffm.) Nyl. f. FUscibuLA Arn. Thallus pallido-fuscescens,
K= lutescens, sorediosus. Soredia albida.—British Columbia: Hazleton, on
branches of Pinus, Aleza Lake, on branches of Abies and Picea i
y
E
c
on trunks of Abies; British Columbia: Kamloops, Fish Lake, Golden, and Aleza
Lake, very frequent on branches and trunks of Pseudotsuga and Pinus Murrayana.
Also on conifers near Hazleton. Sterile.
— JUBATA var. VRANGIANA (Gyeln.) Ras.
ngiana Gyeln. (Magyar Bot. Lap. 31: 46. 1932)
Thallus 1 parce sorediosus, vel fere esorediatus, olivaceo-viridis vel fusco-viridis,
K- oon .— British Columbia: Hazleton, on conifers, occasional. Sterile.
Aisoni A Fremonti Tuck. f. perfertilis Räs., forma nov. Thallus prostratus,
E RAO meti laevis, subnitidus, esorediatus, K=, Cl=, K(Cl)=.
Apothecia vulgaria numerosissima, lateralia, convexa, flavida. Sporae 5-8 X 5 y,
ovoideae, 8: nae, hyalinae. Hymenium I+ caerulescens.— British Colmibis:
Golden, on the trunks and branches of Pinus Murrayana.
ALECTORIA FREMONTI subsp. oLrvACEA Ris. Thallus olivaceo-castaneus,
prostratus, laevis, nitidus, esorediatus, K=, Cl=, K(Cl)=. Apothecia rarissima,
minora, flavida.—British Columbia: Golden, on iion Mies iE and Pseudotsuga,
Kamloops, Fish Lake, on Pinus Murrayana. Ste
The sorediose form, named by Du Rietz (Arkiv f. Bot. 20": 8. 1926) subsp. Eriks-
sonit, was not seen among the North American specimens. It would appear more
correct, as Du Rietz (l. c.) thinks, not to regard the sorediose form as the main
species. My subspecies olivacea (Rüsünen, Medd. Soc. Fauna Flora Fennica 43: 4.
1933]
RASANEN—CONTRIBUTION TO LICHEN FLORA OF AMERICA 11
1916) is much shinier than the North American bid and has very rare apothecia,
so that we have here three different Alectoria t
A THRAUSTA (Ach.) Nyl. British Cui: Hazleton, on branches of
posi Prince George and Six Mile Lake, on Picea. :
ALINA phan nox Nyl. British Columbia: Hazleton, on branches of Pinus
os rare. Ster
RAMALINA ROESLERI ae Nyl. New Brunswick: Dalhousie, on branches of
Picea, Sorbus, and on a wooden wall. Sterile.
RAMALINA CALICARIS (L.) Fr. New Brunswick: Dalhousie, on trunks of Populus.
Fertile.
LETHARIA VULPINA (L.) Vain. Thallus sorediosus. Medulla I+ caerulescens.—
British Columbia: Kamloops, Fish Lake, and Field (subalpine), on branches and
trunks of Pseudotsuga. Sterile.
LETHARIA VULPINA f. INCOMPTA Ach. Thallus sorediosus et bene isidiosus, me-
dulla I+ eaerulescens.—Found with the bci m near Fish Lake, Golden, and
Jackman, B. C., on Pinus Murrayana. Ste
CORNICULARIA DIVERGENS Ach. Alaska b on the ground. Sterile.
CETRARIA MERRILLII D
C. californica var. Tuckermanii Howe
Thallus 0.5-2 cm. longus, rigidus, erectus, divaricato-ramosus, opacus, olivaceo-
niger. Rami angulato-nervosi, foveolati. Apothecia vulgaria, 0.5-3 mm. lata,
plana vel convexa, terminalia, atra vel fusco-nigra, nuda, subnitida, subciliata.
Sporae 8:nae, 6-8 X 4-5.5 y, ovoideae, incoloratae.—British Columbia: Golden,
x ocu d Fish Lake, and Hazleton, on branches of Pinus Murrayana and other
conifer; ertile.
ARIA NIGRICANS Nyl. var. pallida Rüs., var. nov. I" superne pallidus
vel E a ia, quove albidus, basim versus san eo-lentus; medulla I+
caerulescens. Laciniae 1-2 mm. latae, subcanaliculatae, AME ap ope dicho-
tome ramosae.—Alaska, on the ground with Alectoria ochroleuca and Cornicularia
divergens. Sterile.
CETRARIA TENUIFOLIA (L.) Vain. Thallus canaliculatus, laevigatus, medulla
K —.—Alaska, rarely with the previous species. Sterile.
CETRARIA TENUIFOLIA var. reticulata Rüs. var. nov. Thallus subtus foveato-
reticulatus, castaneo-fuscus, in marginibus dense isidioso-dentatus. ^ Laciniae
angustae, SPSE EE UE A crispatae, marginem versus pseudocyphellatae.
Medulla K—.—British Columbia: Kamloops, on stones, Valemount and Aleza
Lake, on the ground. Sterile.
CETRARIA TENUIFOLIA Var. PSEUDOISLANDICA Ris. (Ann. Bot. Soc. Zool.-bot.
Fenn. Vanamo 2: 15. 1932) f. septentrionalis Räs., forma nov. Medulla I—.
Thallus similis C. islandicae, sed I—, et margo laciniarum fere sine spinis vel ciliis.
Color pallido-castaneus.—British Columbia: Golden, on trunks and branches of
dwarf shrubs near the ground. Sterile.
The true Cetraria islandica (L.) Ach., which gives the reaction ‘‘Medulla I lutes-
cens, demum ferrugineo-rubescens," has not been seen by me from either North or
South America.
TRARIA NIVALIS (L.) Ach. British Columbia: Kamloops, on the ground and
rocks in semi-desert; Alaska, on the ground. Sterile.
CETRARIA CUCULLATA (Bell) Ach. British Columbia: rage on the ground
with Arctostaphylos Uva-ursi; Alaska (alpine), on the ground. Steril
[Vor. 20
12 ANNALS OF THE MISSOURI BOTANICAL GARDEN
CETRARIA CAPERATA B. Vain.
C. pinastri (Scop.) R
British Columbia: eal a small specimen with other Cetraria species on
branches of Pinus Murrayana, Aleza Lake, a chiefly geoplese lichen (Rüsünen, Uber
Flechtenstand u. Fl. Veget. im Westl. Nordfinnl., p. 84. 1927), with Parmelia
ambigua and P. pallescens on the base of Abies lastocarpa. Sterile.
CETRARIA JUNIPERINA (L.) Fr. var. viRIDIS (Schweinitz) Riis.
C. viridis gprs apud Halsey, Ann. Lyceum Nat. Hist. New York 1: 16.
1824; C. juniperina var. virescens Tuck.
Thallus ae rigidus; laciniae obtusae, obscuro-virides; K=, Cl=, I=.
Apothecia liri ed AEN 3-8 mm. lata, fusco-nigra, subnitida, margo sub-
crenata, tenu Sporae 8 : nae, ovoida 6-8 X 3.5-6.5 u, incoloratae.—British
Columbia: Golden, common on branches of Pinus Murrayana. Fertile.
CETRARIA JUNIPERINA var. canadensis Räs., var. nov. Thallus erectus vel sub-
erectus, subrigidus; laciniae obtusae, puni Pelih vel interdum viridi-flavae, super
reticulato-nervosae, intus flavae. Apothecia terminalia, minora, 1-3 mm. lata,
castaneo-fusca, nitida; margo lacerato-crenata. Sporae 6.5-8 X 4-5.2 u, ovoideae,
8:nae, hyalinae.—British Columbia: Kamloops, Fish Lake, Aleza Lake, and
Hazleton, very frequent on twigs of conifers.
C. juniperina (the main species) has apothecia chiefly in the middle of the thallus
and the lobes are very divided. Also, the color of this species is not so bright a golden
yellow as in the variety canadensis, and the surface is furthermore much smoother in
the species.
TRARIA JUNIPERINA var. crispata Riis., var. nov. Sicut var. canadensis
thallus dense lacerato-crispatus vel mibpaoiliatas: Apothecia rara. Est Ripe
cum C. juniperina var. terrestris Schaer. (= C. Tilesii Ach. in Zahlbr. Cat. Lich.
Univ. 6: 341. 1930).— British Columbia: Golden, rarely on branches of Pinus
Murrayana with var. viridis.
CETRARIA ALEURITES (Ach.) Th. Fr. New Brunswick: Dalhousie, on trunks of
Abies and on wood. Sterile.
CETRARIA CHRYSANTHA Tuck.
Thallus stramineus, lacunoso-nervosus, REN subtus fusco-niger, K+
lutescens, Cl=, K(Cl)-r roseus, I=.—Alaska (alpine), on the ground. Sterile.
CETRARIA LAURON Ach. var. han DR. Thallus subtus fusco-niger, non
isidiosus, I =.—British Columbia: Kamloops, on stones in semi desert. Sterile.
CETRARIA NoRVEGICA (Lynge) DR. Thallus isidiosus, I -.—With the above
species in Kamloops, on stones. Sterile.
CETRARIA GLAUCA (L.) Ach. Thallus IF caerulescens.—With the above species
in Kamloops, on stones; Prince George, Six Mile c" abundant on Picea; Aleza
Lake and Vancouver, on branches of Picea.
CETRARIA GLAUCA V&r. STENOPHYLLA Tuck. (Syn. North Amer. Lich. 1: 36. 1882).
Thallus I F caerulescens, anguste laciniatus, glaucus, subtus niger.—British
Columbia: Vancouver Is, Lake Cowichan, occasional on the lower branches of
Picea in the forest. Sterile.
CETRARIA SCUTATA (Wulf.) Poetsch
C. chlorophylla (Willd.) Dalla Torre et Sarnth
British Columbia: Kamloops, Fish Lake, Golden, Hazleton, Aleza Lzke, on the
branches of Pinus Murrayana and Abies; Vancouver Is., Lake Cowichan, occasional
on Thuja gigantea. Sterile.
1933]
RASANEN—CONTRIBUTION TO LICHEN FLORA OF AMERICA 13
NEPHROMOPSIS PLATYPHYLLA (Tuck.) Herre
Cetraria er Lab Tuck.
Thallus 1-3 cm. latus, irregulariter laciniatus, super opacus, obscure fuscus,
tuberculosus, s pad pallide fuscus, reticulato-nervosus. Apothecia marginalia,
minora, 1-2 mm. lata, plana, fusca, tenuiter marginata; margo crenata. Sporae
sphaeroideae vel ‘eibiellipsoidene, 8:nae, incoloratae, 4-6.5 u. Hymenium ca. 50 p
crassum, I+ caerulescens.—British Columbia: Golden, on twigs of Pinus Murray-
ana.
NEPHROMOPSIS CILIARIS (Ach.) Hue. British Columbia: Aleza Lake, a small
specimen on Pinus Murrayana. Steri e.
PARMELIA PALLESCENS (Neck.) Ras
P. hyperopta
New Brunswick: Dalhousie, on trunk of Abies; E rs: Columbia: Aleza Lake, on
base of Abies lasiocarpa as & geoplese lichen. Ste
RMELIA AMBIGUA (Wulf.) Ach. New aga Dalhousie, on wood; British
Columbia: Aleza Lake, on base of Abies lasiocarpa and Pinus Murrayana. Sterile
PARMELIA PHYSODES (L.) Ach. f. LABROSA Ach. New Brunswick: xr mdi on
trunk of Abies; British Columbia: Aleza Lake, on trunks of Picea, Kamloops
Pinus Murrayana. rile.
PARMELIA PHYSODES f. viTTATA Mereshk. British Columbia: Aleza Lake and
Hazleton, on branches of Pinus Murrayana. Sterile
PARMELIA LOPHYREA Ach. Thallus cinereus, — Ir K+ eee foe
= roseus. Apothecia vulgaria, subtus ampullacea, demum
Sporae sphaeroideae, 2.5-5 u, 8 : nae, incoloratae. Hymenium 50 u eat T.
caerulescens.— British Columbia: Golden, on branches of Pinus Murrayana; Aleza
Lake, on branches of Picea
PARMELIA DELAVAYI (Hue) Nyl. Thallus parvus, esorediatus, obscure cinereus.
Apothecia vulgaria, demum 5 mm. lata, subtus non ampullacea, discus fuscus,
urceolatus vel deplanatus. Sporae 5-7 X 4-5 u, 8:nae, ovoideae. Hymenium
ca. 55 u crassum, I+ caerulescens.—British Columbia: Golden, on Pinus Murray-
ana.
PARMELIA ENTEROMORPHA Ach. Thallus esorediatus, ventricoso-inflatus, cavus,
K= lutescens, K(Cl) =.—British Columbia: Vancouver Is, Lake Cowichan, on
trunk of Thuja gigantea. Sterile.
PARMELIA VITTATA (Ach.) Róhl. British Columbia: Prince George, Six Mile
Lake, on trunk of Betula?. Sterile.
PARMELIA SULCATA Tayl. New Brunswick: Dalhousie, on trunks of Betula,
Sorbus, Picea, Abies, and on wood; British Columbia: Prince George, Six Mile Lake,
on Betula; Hazleton, on Betula; Golden, on Pinus Murrayana and Pseudotsuga;
Kamloops on arid ground. Sterile.
PARMELIA VAGANS Nyl.
P. molliuscula Tuck. (Syn. North Amer. Lich. 1: 64. 1882).
Thallus dichotome laciniatus, sine sorediis et isidiis, stramineo-virens, K F fulves-
cens.—British Columbia: Kamloops, on the ground between grasses and shrubs.
Sterile. The lichen is a typical desert lichen and appears also in such environment
in Russia and Asia.
PARMELIA SUBAURIFERA Nyl. New Brunswick: Dalhousie, on the trunk of
Sorbus; British Columbia: Hazleton, on the trunk of a Populus. Steri
PARMELIA PAPULOSA (Schaer.) Vain.
P. exasperatula Nyl
[Vor. 20
14 ANNALS OF THE MISSOURI BOTANICAL GARDEN
New Brunswick: Dalhousie, on the trunk of Sorbus. Sterile.
PARMELIA ELEGANTULA (Zahlbr.) Rüs. var. americana "e var. nov. Thallus
opacus, obscure olivaceo-fuscescens, isidiosus, K=, Cl=, K(Cl)=. Isidia sub-
cylindrica, curta, concoloria. Apothecia desunt.—British Sema Kamloops, on
a Pseudotsuga in semi-desert
PARMELIA PUBESCENS (L.) Vain. var. RETICULATA Cromb. British Columbia:
Kamloops, on a stone in semi-desert. Steri
THAMNOLIA VERMICULARIS (Sw.) Schaer. : ddl (regio alpina), on the ground
between tufts of Cornicularia divergens and Cladonia rangiferina
SIPHULA cERATITES (Wahlbg.) Fr. British Columbia: Doreen, on stones in
moist mountain forest. Sterile.
STEREOCAULON TOMENTOSUM Fr. British Columbia: Dorreen, on branches of
Tsuga; Alberta: Lake Louise, on mossy earth; ay m alpina), between Cornic-
ularia divergens and Cetraria chrysantha. Rarely fer
LECANIA DIMERA (Nyl.) Th. Fr. British ci diae - Aleza Lake, very common on
the trunks of Populus. Fertile.
SQUAMARIA ALPHOPLACA (Wahlbg.) Dub. "Thallus orbicularis, radiatus, griseus,
K+ partim rubescens. Medulla I—. Apothecia ca. 1 mm. lata, nigra, nuda vel
tenuiter pruinosa. Sporae 9, 6.5 m, 8 : nae.—U. S. A.: Cody, Wyoming, on sand-
stone in a cactus desert.
ASPICILIA CALCAREA T Mudd. With the previous on sandstone in cactus
desert, in Cody. Ferti
LECANORA COILOCARPA A (Ach. ) Nyl. New Brunswick: Dalhousie, on a rail fence.
Fertile.
LECANORA UMBRINA (Ehrh.) Róhl. New Brunswick: Dalhousie, on a stone in a
field. Fertile.
LECANORA 8YMMICTA Ach. "Thallus areolato-verruculosus, mox totus sorediosus,
virescens, K—, Cl4- aurantiaco-rubescens. Apothecia livida, mox convexa, fere
immarginata.—New Brunswick: Dalhousie, on a wood rail fence
mictera (Nyl.) Rüs.—in the summer of 1931, on a wooden fence in Petsamo, Finnland.
Except for this I have not seen this central European and North American species.
LECANORA SUBINTRICATA Nyl. British Columbia: Kamloops, Fish Lake, on
trunks and branches of Pinus Murrayana.
OCHROLECHIA UPSALIENSIS (L.) Mass. Thallus K-, Cl-, K(Cl)-. Epi-
thecium K(Cl)—. Apothecia 1-2 mm. lata, planiuscula vel urceolata, pruinosa.
Margo crassa, subrugosa.—British Columbia: on rotting vegetable remains in
somewhat shaded positions in the semi-desert.
PHLYCTIS ARGENA (Ach.) Flot. New Brunswick: Dalhousie, on trunk of Abies sp.
Sterile.
PERTUSARIA PERTUSA (L.) Tuck. New Brunswick: Dalhousie, together with
previous species on Abies. Fertile.
PERTUSARIA XANTHOSTOMA (Smrft.) Fr. Thallus verruculoso-inaequalis, K —,
K(Cl)—. Apothecia 0.5 mm. lata, punctiformia, pallido-lutescentia. Excipulum
K+, sanguineo-rubescens. Sporae 4:nae, ellipsoideae, incoloratae, 58-66 X 26-40 u.
Hymenium I+ caerulescens.—New Brunswick: Dalhousie, on a Thuja trunk.
PrERTUSARIA MULTIPUNCTA (Turn.) Nyl. British Columbia: Aleza Lake, on the
trunk of an Abies. Fertile.
PERTUSARIA FAGINEA UT CURES I IU stranthium
ce. ae E a nd unequally lacerated crown; ray-flowers in 2 or
Achaeto
88. bins d Mas ud subterete, or distinctly 4—5-angled.... Aphanostephus
KEY To SPECIES OF ASTRANTHIUM
A. Stems erect or suberect from a distinctly ligneous base.
s. Leaves glalgmus. 056054 605s RR SEMEL Aces y es 1. A. mimum
aa. Leaves more or less pubescent.
b. Stems usually unbranched above.
€. Leaves oblong-elliptic. .... Eat DEI eis ss 2. A. xylopodum
cc. Leaves oblanceolate to lineal spe. a renas roie 3. A. orthopodum
bb. Stems usually branched above.
6 Lenves dentate.. 5c er EN E oe ATTA e s 4. A, mexicanum
| Leaves ntie, 66s 1o" Re Ar aie estas 4a. Var. chihuahuense
AA. Seas erect or procumbent from an annual or slightly ligneous base.
a. Plants with long slender runners....................-. 5. A. xanthocomoides
aa. Plants without long slender runners.
bk Aches glabrous.: Loo EIER QU ess 6. A. purpurascens
bb. Achenes pubescent m glochidiate-tipped hairs.
c. Leaves not rosulat
d. Plants not nane UT a seeoose 7. A. integrifolium
dd. Plants conspicuously branched.................. 7a. Var. ciliatum
66. Leaves TORAO. oea reesi ee des ees 7b. Var. rosulatum
TAXONOMY
ASTRANTHIUM
Astranthium Nutt. Trans. Am. Phil. Soc. N. S. 7: 312. 1841;
Benth. & Hook. Gen. Pl. 2: 265. 1873, as to synonymy.
Bellis Michx. and Am. authors, Fl. Bor. Am. 2: 131. 1803,
not of Linnaeus; Hook. Bot. Mag. 52: pl. 3465. 1835; DC.
Prodr. 5: 304. 1836; Raf. New Fl. Am. 2: 24. 1836; Torr. &
[Vor. 20
30 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Gray, Fl. N. Am. 2:189. 1842; Gray in Hemsl. Biol. Cent.-Am.
Bot. 2: 118. 1881; Gray, Syn. Fl. N. Am. 12: 163. 1884, and
ed. 2, 163. 1886 and 1888; Britton & Brown, Ill. Fl. 3: 350.
1898, and ed. 2, 3: 402. 1913; Britton, Manual, 943. 1901,
and ed. 2, 1905; Rob. & Fern. in Gray, Manual, ed. 7, 799. 1908;
Small, Fl. Southeastern U. S. 1202. 1903, and ed. 2, 1913.
Herbaceous, caulescent, glabrate, or pubescent annuals or
perennials. Leaves alternate, sessile or petioled, linear-lanceo-
late to obovate-spathulate, entire or dentate. Involucre 2-3-
seriate, imbricated, appressed, lanceolate bracts usually with
lacerately ciliate, membranaceous margins. Heads few- to many-
flowered. Ray-flowers pistillate, rays 2-3-dentate. Disk-flowers
tubular, perfect, corolla 5-lobed. Style-branches linear-lanceo-
late, acute, papillose at the tip, stigmatic surfaces confined to
the lowermost margins of the style-branches and extending one
third of their length. Pappus entirely lacking or an inconspicu-
ous ring-like crown. Achenes obovate, compressed, narrowed
at the base, pubescent with glochidiate or emarginate hairs,
rarely glabrous.
Type species: Astranthium integrifolium (Michx.) Nutt. Trans.
Am. Phil. Soc. N. S. 7: 312. 1841, which was based on Bellis
integrifolia Michx. Fl. Bor. Am. 2: 131. 1803, ''ad ripas rivu-
lorum et in collibus umbrosis Tennassée.”’
1. Astranthium mimum (Blake) Larsen, n. comb.
Bellis mima Blake, Contr. U. 8. Nat. Herb. 22: 594. 1924.
Herbaceous perennial, 38-50 cm. high; stems several, simple,
erect, monocephalous, greenish, striate, glabrous or sparsely
pubescent with spreading hairs; basal leaves few, obovate-
oblong, apiculate, narrowed at the base into a slender petiole,
15-20 em. long, 2-3 em. wide, somewhat membranous in the
dried state; stem-leaves linear-lanceolate, hirsute-ciliate, lower
5-6 cm. long, 4-6 mm. wide, gradually reduced above, the
uppermost 1 cm. long, 1 mm. wide; peduncles terminal, mono-
cephalous, enlarged just below the head; involucre 2-2.5 cm. in
diameter; bracts 2-seriate, equal, 7 mm. high, linear, acute,
sparsely pilose, ciliate; ray-flowers about 60, white, fertile, 3-
dentate, about 12 mm. long; disk-flowers yellow, fertile, pappus
1933]
LARSEN—ASTRANTHIUM AND RELATED GENERA 31
none; achenes of ray and disk similar, oblong, compressed,
glabrous.
Distribution: State of Durango, Mexico. Known only from type specimen.
Specimens examined:
Duranco: from Sierra Madre, 30 miles north of Guanacevi, alt. 2440-2745 m.,
Aug. 18, 1898, Nelson 4786 (US, No. 332830, TYPE).
2. Astranthium xylopodum Larsen, n. name.
Keerlia mexicana Gray, Proc. Am. Acad. 22: 422. 1887, not
Bellis mexicana Gray, Smiths. Contr. [Pl. Wright. pt. 1] 3: 93.
1852.
Perennial from a woody base; stems several, simple, or slightly
branched near the top, striate, hirsute with spreading pubescence;
peduncles 10-14 cm. long, naked or bearing 1-2 small linear-
lanceolate bracts near the inflorescence, hirsute-pubescent;
leaves oblong-elliptic, evenly pubescent with appressed hirsute
hairs, margins strongly ciliate; involucre 1-2 cm. in diameter;
bracts linear-lanceolate, pubescent, margins lacerately ciliate,
somewhat membranaceous; ray-flowers fertile, about 20; pappus
lacking; achenes 4-nerved, glabrous.
Distribution: known only from the State of Jalisco, Mexico.
Specimens examined:
JALISCO: shaded hillsides near Guadalajara, June 27-July 14, 1893, Pringle 4412
(M, G); Rio Blanco, July 1886, Edward Palmer 146 (G, TYPE).
3. Astranthium orthopodum (Rob. & Fern.) Larsen, n. comb.
Bellis orthopoda Rob. & Fern. Proc. Am. Acad. 30: 117. 1894.
Perennial from a short, thick rootstalk; stems several, de-
cumbent or suberect, 10-20 cm. high, simple or somewhat
branched, appressed-pubescent, monocephalous; leaves thick,
entire, appressed-pubescent, basal leaves oblong-spathulate, 3—4
cm. long, 3-5 mm. broad, stem-leaves linear-lanceolate, erect,
gradually reduced toward the inflorescence; peduncles 3-4.5 cm.
long, appressed-pubescent; involucre 10-12 mm. in diameter;
bracts 2-seriate, linear-lanceolate, acute, purple-margined, sparse-
ly and somewhat appressed-pubescent, 4-5 mm. long; ray-
flowers about 30, 1 cm. long; achenes sparsely pubescent with
glochidiate or straight and slightly emarginate hairs; pappus an
inconspicuous whitish ring-like crown.
Distribution: western Chihuahua, southward along mountains in the state of
Durango, Mexico.
[Vor. 20
32 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Specimens examined:
Cuimuanva: Guachachie, June 25, 1892, Hartman 523 (G, TYPE); vicinity of
Madera, alt. 2250 m., May 27-June 3, 1908, Edward Palmer 287 (
DvnaNao: City of Durango and vicinity, Apr.-Nov. 1896, Edward Palmer 163
(M); Otinapa, July 25-Aug. 5, 1906, Edward Palmer 425 (M).
4. Astranthium mexicanum (Gray) Larsen, n. comb.
Bellis mexicana Gray, Smiths. Contr. [Pl. Wright. pt. 1] 3: 93.
1852; Gray in Hemsl. Biol. Cent.-Am. Bot. 2: 118. 1881.
Perennial from a branched ligneous base; stems 15-60 cm.
high, several, branched above, striate, pubescent with spreading
hairs; leaves appressed-pubescent, those of the stem sessile,
entire, or saliently dentate, oblong-linear, gradually reduced
toward the inflorescence, basal leaves ellipsoid-spathulate,
crenate-dentate, 5-20 cm. long, .8-2.5 em. broad; involucre .8-
1.4 em. in diameter; bracts 2-3-seriate, cinereous-pubescent,
linear-lanceolate, purple-tipped; ray-flowers white, numerous,
.5-1.0 em. long; pappus an inconspicuous ring-like crown or
almost obsolete; achenes compressed, pubescent.
Distribution: mountains of south-central Mexico.
Specimens examin
HripALao: Sierra de Pachuca, alt. 3076 m., Aug. 26, 1902, Pringle 9857 (M).
Mexico: Apr. 26, 1849, Gregg 701 (M, even); along brooks, Ixtaccihuatl, March-
July 1903, Purpus 159 (M); moist open woods and meadows about timber line,
Ixtaccihuatl, Oct. 1905, Purpus 1575 (M); “above timber line,” Popocatepetl, Oct.
1908, Purpus 3640 (M).
FEpERAL District: Cima, Aug. 24, 1910, Orcutt 3768 (M); Serrania de Ajusco,
alt. 3076 m., Aug. 8, 1896, Pringle 6442 (M).
More tos: Toro, alt. 3015 m., Aug. 5, el Fisher (M, No. 914805).
Oaxaca: Sierra de San Felipe, alt. 3076 m., June 28, 1894, pee 4719 (M);
Sierra de San Felipe, alt. 3076 m., Sept. 1894, ym 261 (M).
4a. Var. chihuahuense Larsen, n. var.” Plate 3.
Resembles the species in habit but differs from it in that it
is much less pubescent; the leaf margins are essentially entire,
ciliate; the pappus is more conspicuous than that of the species
and consists of a narrow fluted crown; the achenes are glabrous,
or glabrate.
? Astranthium mexicanum (Gray) Larsen var. chihuahuense Larsen, n. var.,
planta herbacea perennis, 6-7 dm. alta, supra ramosa; marginibus foliorum integris
vel subintegris; pappo coroniforme, margine irregulariter sinuato-dentato; achaeniis
glabris vel glabratis—Mexico: canyons of Sierra Madre, State of Chihuahua, Oct.
4, 1888, Pringle 2015 (M, No. 122921, TYPE).
1933]
LARSEN—ASTRANTHIUM AND RELATED GENERA 33
Distribution: State of Chihuahua, Mexico, known only from the type.
Specimens examined:
CurmvuanHva: canyons of Sierra Madre, Oct. 4, 1888, Pringle 2015 (M, No. 122921,
TYPE).
5. Astranthium xanthocomoides (Less.) Larsen, n. comb.
Brachycome xanthocomoides Less. Syn. 192. 1832, nomen sub-
nudum; Schlecht. in Linnaea 9: 265. 1835.
Brachycome xeranthemoides Steud. Nomencl. Bot. ed. 2, 220.
1840, in part (typographical error).
Keerlia linearifolia DC. Prodr. 5: 310. 1836, in part as to
synon.
Bellis xanthocomoides (Less.) Gray in Hemsl. Biol. Cent.-Am.
Bot. 2:118. 1881
Annual; stems several, striate, procumbent or ascending,
branching, leafy throughout, producing slender runners; leaves
spathulate, apiculate, sparsely pubescent, gradually reduced
toward the peduncles, 1.5-2.5 cm. long, .3—.8 cm. broad; pe-
duncles 1-6 cm. long, densely pubescent at the base of the invo-
lucre; involucre .8-1.2 cm. in diameter; bracts 2-seriate, lanceo-
late, acute, pubescent along the lower portion of the main axis,
membranaceous-margined; ray-flowers white, .5-.8 cm. long,
about 16; pappus practically lacking; achenes compressed,
pubescent, becoming more or less glabrate.
Distribution: xe meadows and open woods of east-central Mexico.
Specimens exam
Vera Cruz: prope la Jaya (La Hoya), June 29, Schiede 206 (M, TYPE).
HripALGO: meadows of Sierra de Pachuca, alt. 2760 m., July 17-28, 1898, Pringle
6888 (M); alpine meadows, Sierra de Pachuca, alt. 2023 m., Aug. 26, 1902, Pringle
9858 (M); moist meadows and open woods, Pachuca, July 1905, Purpus 1844 (M).
6. Astranthium purpurascens (Rob.) Larsen, n. comb.
Bellis purpurascens Rob. Proc. Am. Acad. 27: 172. 1892.
Perennial; roots of numerous fibers; stems erect, branched, 4—
45 cm. high, striate, pubescent with spreading hairs; leaves
apiculate, pubescent with spreading hairs, margins Bewteshat
ciliate; lower leaves oblong-ovate, 2.5—4.0 cm. long, 1.5-6 cm.
broad; upper leaves linear to linear-lanceolate, gradually reduced,
1 cm. or less in length at the base of the peduncles; peduncles
3-8 cm. long; involucre .8-1.0 cm. in diameter; involucral bracts
about 14-16, lanceolate, acute, scarious-margined, only slightly
[Vor. 20
34 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ciliate near the apex, pubescent along the main axis with a few
upwardly appressed hairs; rays 10-15, whitish-purple, .5—8 cm.
long; pappus lacking; achenes glabrous, golden-brown at matu-
rity.
This species is closely allied to A. integrifolium (Michx.) Nutt. ;
the chief difference is to be found in achenial characters. Astran-
thium integrifolium has achenes which are pubescent with glo-
chidiate-tipped hairs whereas A. purpurascens (Rob.) Larsen
has glabrous achenes.
Distribution: known -— from the type locality and Chiapas, Mexico.
Specimens examine
San Luis Porosi: shaded grassy slopes, barranca of Las Canoas, Aug. 18, 1891,
Pringle 3819 (M, coTYPE).
Curapas: without definite locality, coll. of 1864—70, Ghiesbreght 548 (M).
7. Astranthium integrifolium (Michx.) Nutt. Trans. Am.
Phil. Soc. N. 8. 7: 312. 1841.
Bellis integrifolia Michx. Fl. Bor. Am. 2: 131. 1803; Hook.
Bot. Mag. 52: pl. 3455. 1835, in part, exclusive of Brachycome
zanthocomoides Less.; DC. Prodr. 5: 304. 1836; Raf. New FI.
Am. 2: 24. 1836; Torr. & Gray, Fl. N. Am. 2: 189. 1842;
Gray, Syn. Fl. N. Am. 1?: 163. 1884, and ed. 2, 1886 and 1888;
Britton & Brown, Ill. Fl. 3: 350. 1898, and ed. 2, 3: 402. 1913;
Britton, Manual, 943. 1901, and ed. 2, 1905; Rob. & Fern. in
Gray, Manual, ed. 7, 799. 1908; Small, Fl. Southeastern U. S.
1202. 1903, and ed. 2, 1913.
Bellis nutans Raf. New Fl. Am. 2: 23. 1830.
Bellis parviflora Raf. Ibid.
Eclipta integrifolia Spreng. Syst. Veg. 3: 602. 1826.
Annual, openly branched from near the base, 8-45 cm. high,
sparsely pubescent with spreading or subappressed hairs, fre-
quently conspicuously pubescent near the base, leafy throughout;
lower leaves oblong-spathulate, 2.5-4.0 cm. long, .8-1.4 cm.
broad; stem-leaves oblong-ovate, sparsely pubescent, margins
ciliate, 3.5 em. long, 1.5 em. broad, gradually reduced toward the
base of the peduncle; peduncles 4-9 em. long, densely pubescent
at the base of the involucre; involucre 2-seriate, about .6-1.2 cm.
in diameter, bracts lanceolate, acuminate, acute, membranous
margins only slightly lacerate, sparsely pubescent along the main
1933]
LARSEN—ASTRANTHIUM AND RELATED GENERA 35
axis with a few spreading hairs; ray-flowers fertile, 16-22, pur-
plish-blue; achenes compressed, pubescent with glochidiate-
tipped hairs.
Distribution: sandy soil, south-central Kentucky to northwestern Georgia, west-
ward to southeastern Kansas and eastern Oklahoma.
Specimens examin
GroraIa: dry ra Ringold Road, Chickamauga, May 27, 1911, Churchill (M,
No. 839094).
Mississippi: Tchula, Holmes Co., Apr. 18, 1927, ves is Anderson 1555 (M).
Kentucky: Bowling Green, May 6, 1892, Price (M, No. 122882
TENNESSEE: West Nashville, May 26-27, 1909, 2 4422 (M); moist field,
Joelton, Davidson Co., July 16, 1922, Svenson 118 (G); copses around Nashville,
May-June, Gattinger 1297 (M); rocky open hillsides, near Erin, Houston Co., May
24, 1920, E. J. Palmer 17623 (M, G
Missouri: gravelly barrens, Noel, May 10, 1915, Bush 7534 (M, G); gravelly
barrens, Noel, May 10, 1915, Bush 7534 A (M); rocky open ground, near Jane,
MacDonald Co., May 23, 1931, E. J. Palmer 39297 (M); rocky Mà r bald knobs,
along creek, near Oasis, Taney Co., June 3, 1931, E. J. Palmer 8 M).
ARKANSAS: river bottoms near Fayettotille, June 1835, nci 129 [607] M);
alluvial soil, waste places, cultivated fields, Fayetteville, May, Harvey 45 (M
Fayetteville, 1880, Harvey (M, No. 122880); Hot Springs, May, 1879, Soulard (M,
No. 122885); sandy soil, Oik, Marion Co., June 15, 1914, E. J. Palmer 5992 (M).
Kansas: rocky soil, Chautauqua Co., Mar 7, 1897, Hitchcock 1056 (M).
OKLAHOMA: in woods and BREL. common near Tishomingo, Johnston Co.,
Apr. 27, 1916, Houghton 3549 (G); clay washes, near Ardmore, Carter Co., Apr. 17,
1913, Stevens 77 (M); Arbuckle Mts., Crusher, May 12, 1916, Emig 606 (M); Catoosa,
May 8, 1895, Bush 897 (M); Ind. Terr., 1877, Butler (M, No. 122886); upland
prairies, sandy soil, near Howe, Le Flore Co., May 25, 1931, E. J. Palmer 39343 (M).
7a. Var. ciliatum (Raf.) Larsen, n. comb. Pl. 4, fig. 1.
Bellis ciliata Raf. New. Fl. Am. 2: 24. 1836.
Bellis integrifolia Gray, Smiths. Contr. [Pl. Wright. pt. 2]
5:78. 1853, in part.
Stems several, diffusely branched from near the base, 1-3 dm.
high, terminated by long slender peduncles; leaves oblong-
spathulate to linear-oblong, reduced to small bract-like leaves
at the base of the peduncles, .8-3.0 cm. long, .2-.8 cm. broad,
sparsely pubescent with subappressed hairs, margins ciliate;
peduncles 4-6 cm. long; involucre .5-.8 cm. in diameter; in-
volucral bracts lance-elliptic, acute, the narrow Tina scous
margins lacerately ciliate; ray-flowers about 9-15; achenes pubes-
cent with glochidiate-tipped hairs.
Distribution: from southeastern Oklahoma and southwestern Arkansas to south-
central Texas, west to the Pecos
[Vor. 20
36 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Specimens examined:
ARKANSAS: moist open ground, Fayetteville, Washington Co., July 6, 1915, E.
J. Palmer 8176 (M); Coal Bank, May 14, 1895, Bush 895 (M); Redfork, May 14,
1895, Bush 896 (M).
OKLAHOMA: open woods near Idabel, McCurtain Co., May 18, 1916, Houghton 3645
G).
Texas: Apr.-May, 1844, Lindheimer 251 (M); “Pine’s Island," May 5, 1903, Rever-
chon (M, No. 122889); Reverchon 440 in part (M, No. 122890); clay barrens, Bryan,
Brazos Co., May 27, 1915, E. J. Palmer 7785 (M); Pecos and Limpio, June 1851-2,
Wright 1176 in part (M, No. 123093); dry banks, Hempstead, Apr. 24, 1872, Hall
3806 (M).
7b. Var. rosulatum Larsen, n. var.? Pl. 4, fig. 2.
Bellis integrifolia Gray, Smiths. Contr. [Pl. Wright. pt. 2] 5:
78. 1853, in part.
Stems several, sparsely branched near the base, pubescent with
spreading hairs, .5-1.5 dm. high; leaves sparsely pubescent with
subvillose hairs; basal leaves numerous, forming a rosette, oblong-
spathulate, 2-7 em. long, .5-1 cm. broad; stem-leaves oblong-
spathulate to linear-lanceolate, gradually reduced to the base of
the peduncle; peduncle 1.5-5 cm. long; involucre about .7 em. in
diameter; bracts lance-elliptic, acute, the narrow membranaceous
margins somewhat lacerately ciliate near the apex; ray-flowers
about 16; achenes densely pubescent with glochidiate-tipped
hairs.
Distribution: — Oklahoma and eastern 'Texas west to Presidio County.
— exa
OKLAHOMA: open sepas near Mannsville, Johnston Co., May 16, 1916, Griffith 3455
(M, G); pnis cu Gap, Apr. 16, 1877, Butler 63 (M); Sapulpa, Apr. 28, 1895,
Bush 927 (M); * Arkansas," May 20, 1895, Bush 929 (M).
Texas: sandy prairies, Columbia, Brazoria Co., March 25, 1914, E. J. Palmer 5028
(M); Vietoria, Vietoria Co., March 5, 1916, E. J. Palmer 9070 (M); sandy open
ground, Larissa, Cherokee Oó., Apr. 7, 1916, E. J. Palmer 9388 (M); sandy prairies,
Matagorda, Matagorda Co., March 5, 1914, E. J. Palmer 4856 (M, No. 753171, TYPE);
on plains, Handley, Apr. 15, 1913, Ruth 74 (M, G); along railroads near Houston,
Apr. 23, 1899, Eggert (M, Nos. 122893 & 122895); Terrell, Apr. 6, 1903, Reverchon
4006 (M); prairies near Victoria, Apr. 7 & 10, 1900, Eggert (M, No. 122899); Pecos
and Limpio, June 1851-2, Wright 1176 in part (M, No. 123091); near San Felipe,
233 Astranthium integrifolium (Michx.) Nutt. var. rosulatum Larsen, n. var.,
caule simplice vel basi ramoso, 0.5-1.5 dm. alto, subvilloso; foliis basalibus rosulatis,
oblongo-spathulatis, 2-7 cm. longis, usque ad 1 cm. latis, integris, ad apicem rotun-
datis vel obtusis, sparse pubescentibus; foliis caulinis oblongo-spathulatis vel lineari-
—— integris, superne sensim minoribus; pedunculis 1-6 em. longis.— TExas:
sandy prairies, Matagorda, Matagorda County, March 5, 1914, E. J. Palmer
4856 (M, 753171, TYPE).
1933]
LARSEN——ASTRANTHIUM AND RELATED GENERA 37
Apr. 1839, Lindheimer (M, No. 122904); along river, Columbia, Apr. 19, 1899, Bush
195 (M); on prairie, Chenango Junction, Apr. 18, 1900, Bush 60 (M); Reverchon 440
in part (M, No. 122898); open ground, along railway grade near Marfa, Presidio Co.,
June 18, 1926, E. J. Palmer 31036 (M).
KEERLIA
Keerlia Gray, Bost. Jour. Nat. Hist. [Pl. Lindh. pt. 2] 6: 221,
222. 1850, in part as to K. bellidifolia and K. effusa, not DC.;
Smiths. Contr. [Pl. Wright. pt. 1] 3: 92. 1852, not DC.; Syn.
Fl. N. Am. 1?: 164. 1884, and ed. 2, 164. 1886 and 1888.
Bourdonia Greene, Erythea 1: 207. 1893.
Herbaceous, caulescent, pubescent annuals or perennials.
Leaves alternate, sessile or petioled, oblong to obovate-spathulate,
entire. Involucre of imbricated lanceolate bracts, with mem-
branaceous margins. Heads few-flowered. Ray-flowers in a
single series, pistillate, fertile. Disk-flowers tubular, five-lobed,
frequently sterile. Branches of the style lanceolate, acutish,
hairy toward the tip. Pappus a thickened ring-like or slightly
lacerated crown. Achenes compressed, glabrate or hirsute-
pubescent.
Type species: K. bellidifolia Gray & Engelm. Proc. Am.
Acad. 1: 47. 1848, which was based on No. 415 of Lindheimer,
collected above Guadeloupe, Texas, 1845-1846.
KEY TO THE SPECIES
Stems branched from the base... .....ccds ccc dnioe cccscece 8. K. Bigs od pa
Stems simple to the inflorescence................0 cee cece ceeneues 9. K. effusa
8. Keerlia bellidifolia Gray and Engelm. Proc. Am. Acad. 1:
47. 1848; Gray, Bost. Jour. Nat. Hist. [Pl. Lindh. pt. 2] 6: 220.
1850; Smiths. Contr. [Pl. Wright. pt. 1] 3: 92. 1852; Syn. FI.
N. o 1:: 164. 1884, and ed. 2,164. 1886 and 1888.
Bourdonia bellidifolia (Gray & Engelm.) Greene, Erythea 1:
207. 1893.
Annual; the caudex giving rise to much-branched stems 9—30
em. high, leafy throughout, striate, pubescent with multicellular
spreading hairs; leaves thin, sparsely pubescent, apiculate, lower
leaves obovate-spathulate, uppermost somewhat linear, gradually
reduced toward the inflorescence, 1—4 cm. long, .4-1.0 em. broad,
narrowed below to a subpetiolate base; heads small; involucre
[Vor. 20
38 ANNALS OF THE MISSOURI BOTANICAL GARDEN
.3-.5 em. in diameter, bracts 2-3-seriate, glabrous, linear-lanceo-
late, acute, with membranaceous margins; ray-flowers 4-15, blue,
fertile; disk-flowers 15-20, frequently sterile, style branches
shorter than those of the ray-flowers; pappus a thickened ring-
like crown; achenes nerved, compressed, glabrate.
Distribution: southern Texas; doubtless also in adjacent Mexico.
Texas: along Nueces Bay, Nueces Co., alt. 6 m., March 12, 1894, Heller
1436 (M); Rock Springs, Apr. 17, 1930, Marcus E. Jones 26575 (M); above Guada-
lupe, 1845-46, Lindheimer 415 (M, co-rvrE); near New Braunfels, May 1848,
Lindheimer 628 (M); Comanche Spring, New Braunfels, Apr. 1850, Lindheimer 932
(M); rocky calcareous ground, Del Rio, Valverde Co., March 26, 1917, E. J. Palmer
11384 (M); dry limestone hills, Concan, Uvalde Co., Apr. 13, 1917, E. J. Palmer
11550 (M); vicinity of San Antonio, Apr. 1919, von Schrenk (M, No. 88063); near
Austin, March 17, 1908, York 410 (M).
9. Keerlia effusa Gray, Bost. Jour. Nat. Hist. [Pl. Lindh. pt.
2]6:222. 1850; Smiths. Contr. [Pl. Wright. pt. 1] 3: 93. 1852;
Syn. Fl. N. Am. 1?: 165. 1884, and ed. 2, 165. 1886 and 1888.
Bourdonia effusa (Gray) Greene, Erythea 1: 207. 1893.
Perennial, 15-60 cm. high; stem simple below, diffusely branched
above into an open glabrous panicle; stem and leaves hispid-
pubescent; leaves thin, lowermost leaves obovate-spathulate,
3-9 cm. long, .5-1.3 cm. broad, narrowed at the base into a
petiole, those of the stem oblong with a broad, sessile, somewhat
amplexicaul base, gradually reduced toward the inflorescence;
heads numerous, small; involucre turbinate, 2-3 cm. in diameter;
involucral bracts glabrous, 3-4-seriate, linear-lanceolate, acute,
membranaceous-margined; ray-flowers 4-7, white, fertile; disk-
flowers 6-9, sterile; pappus consisting of a thickened irregularly
lacerate crown; achenes compressed, hispid-pubescent.
Distribution: south-central Texas.
Specimens examined:
Texas: Berlandier 499 (M); White Oak Creek, Gillespie Co., Jermy 815 (M);
shady banks of the Upper Guadalupe, 50 miles above New Braunfels, 1847-48,
Lindheimer 629 (M, co-ryre); dry hillside thickets, Leakey, Edwards Co., June 10,
1916, E. J. Palmer 10165 (M); dry hillside thickets, Spanish Pass, Kendall Co., Sept.
28, 1916, E. J. Palmer 10839 (M); San Geronimo Creek, June 1884, Reverchon 1534
(M); Comanche Spring, New Braunfels, 1849, Lindheimer 933 (M).
1933]
LARSEN—ASTRANTHIUM AND RELATED GENERA 39
List or ExsiCCATAE
The distribution numbers are dii in italics. The number in parenthesis is
the species number used in this revisio
Berlandier, J. L. 499
Bush, B. F. 897, HA 754A (7); 896,
896 (7a); 60, - xdi 7b).
Churehill, J J. R. -— ji
Eggert, H. — (7b).
Eggleston, W. M 4422 (7).
Emig, W. H. 606 (7).
Engelmann, " p [607] (7).
Fisher, G. L. — (4
Gattinger, A. 1297 (7).
Griffith, F. 3455 (7b).
Hall, Elihu. 306 (7a).
me C. V. 623 (3).
Harvey, F. L. —, 45 (7).
Heller, A. A. 1436 M.
Hitchcock, A. S. 1055
Houghton, H. W. 3549 na 3645 (Ta).
Jermy, Gustav. 815 (9).
Jones, Marcus E. 26575 (8).
Lindheimer, F. 251 (7a);
628, 932 (8); T 933 (9).
Nelson, E. W. 4786 (1).
Orcutt, C. R. bos (4).
—, (7b); 416,
Palmer, E. J. 5992, 17623, 39297, 39343,
39483 (7); 7785, 8176 (Ta); 4855, 5088,
9070, 9388, 31036 (Tb); 11384, 11550
(8); 10165, 10839 (9).
Palmer, Edward, 146 (2); 163, 287, 425
(3).
Price, Sadie F. —
Pringle, C. G. 2); 4719, 6442,
9857 nt 2016 AS 6888, 9858 (5);
3819
Purpus, A. 159, 1575, 8640 (4); 1344
Reventhaty J. —, 440 in t (7a); —
(7b); 4006, 440 in part (7b); ; 1584 (9).
Ruth, Albert. ?4 (7b).
Schiede, G. 206 (5).
Smith, Charles L. 261 (4).
Svenson, H. K. 118 (7).
von Schrenk, H. (8).
Woodson, R. E. Jr., and Anderson, E. 8.
1666 (7).
e Charles. 1176 in part (7a); 1176
in
rt (7b
Yo H. H. 410 (8).
[Vor. 20, 1933]
ANNALS OF THE MISSOURI BOTANICAL GARDEN
PLATE 2
Astranthium — (Michx.) Nutt. from Eggleston 4422 in the Missouri
Botanical Garden Herbar
918135
Involucre d receptacle x 5.
Style-branches of the disk-flower, X 10.
Style-branch greatly enlarged.
Bellis perennis L. from Pring, in the Missouri Botanical Garden Herbarium No.
Involucre and receptacle, X 5.
Ray-flower, X 10.
Disk-flower, X 10.
Stamen, X 10.
Style-branches of the disk-flower, X 10.
Style-branch greatly enlarged.
ANN. Mo. Bor. Garb., Vor. 20, 1933 PLAYE 2
Sys
he .
ce e.
LA
nm
^ "js "^s o
y ae 9 Tu 7,
E31 Nu NÉ
T^ Pal
4 |
gu |
d X9 ff i” + : / / /
j IH pH
LARSEN—ASTRANTHIUM AND RELATED GENERA
[Vor. 20, 1933]
42 ANNALS OF THE MISSOURI BOTANICAL GARDEN
PLATE 3
Astranthium mexicanum (Gray) Larsen var. chihuahuense Larsen. From the type
specimen, Pringle 2015, in the Missouri Botanical Garden Herbarium.
ANN. Mo. Bor. GARD., Vor. 20, 1933
^
PLATE 3
\
CEE
Uu «Ira?
uate,
a4 4°
Yen ate
3 C. G. PRINGLE,
PLANTA MEXICAN Æ.
1858,
TATE CP CEIRTANA | a
2015 Aphantostephus Arkansanus, Gray.
Canyons of Seren Waites
4, fhteter
LARSEN—ASTRANTHIUM AND RELATED GENERA
[Vor. 20, 1933]
44 ANNALS OF THE MISSOURI BOTANICAL GARDEN
PLATE 4
Fig. 1. Astranthium integrifolium (Michx.) Nutt. var. ciliatum (Raf.) Larsen.
From Lindheimer 251, in the Missouri Botanical Garden Herbarium.
Fig. 2. Astranthium integrifolium (Michx.) Nutt. var. rosulatum Larsen. From
the type specimen, Palmer 4855, in the Missouri Botanical Garden Herbarium.
PLATE 4
)
D
0, 193:
2
ANN. Mo, Bor. GARD., Vor.
d
Au
on
fie yer Pacman imd Sadun 1marta iia
A gat, b
ge tats *
Pa re
tan ^x
* *
Flora Texana exsiceata.
$251.
Foe 4) Jag V. Lidicinse, 1541
LARSEN—ASTRANTHIUM AND RELATED GENERA
FERNS AND FERN ALLIES OF MISSOURI!
M. ELIZABETH PINKERTON
Assistant in Botany, Henry Shaw School of Botany of Washington University
This paper is based primarily upon material in the herbarium
of the Missouri Botanical Garden, the University of Missouri,
and the private collections of Mr. John H. Kellogg and of the
author. After each specific name reference is given to the
original publication; this is followed by reference to the two
standard manuals and to an illustrated monograph covering the
fern-flora of the region concerned, where the species or variety
is treated under the same or a different name. These are:
Gray, ‘New Manual of Botany,’ ed. 7, 1908; Britton and Brown,
‘Illustrated Flora of the Northern States and Canada,’ ed. 1,
1896, and ed. 2, 1913; and Eaton, ‘Ferns of North America,’
vol. 1, 1879 (plates 1-45) and vol. 2, 1880 (plates 46-81). Time
of fruiting, the general distribution for North America, and the
habitat for Missouri in particular precede the specific description.
The names of the principal collectors whose material was ex-
amined are listed, and definite citations made when the plant is
rare.
Key To FAMILIES
1. Plants with short vertical stems or rootstocks. .............. eee 2
1. Plants with horizontally spreading stems or rootstocks...............-00+065 4
2. Leaves onion-like, producing sporangia at their bases............... Isoetaceae
2. Leaves neither onion-like nor producing sporangia at their bases...............
3. Sporangia borne on under sides of leaves................ sese. Polypodiaceae
3. Sporangia borne terminally in special fruiting clusters.......... Ophioglossaceae
4. Sterile leaves large, usually compound or variously divided.................. 5
4. Sterile leaves small, often acale-lik®, 5.2.02. ccc c ccc ce cece ecb oeseseberseues 6
5. Fertile se leaf-like or pod-like; if the latter, sporangia covered with
Inque sla ei P MPA r ei ice ara 8 (Sie Polypodiaceae
5. Fertile segments 1 not leaf-like; sporangia naked................ Osmundaceae
6. Plants: truly quaie......... REA EREAMERA REA e rrr RR Salviniaceae
6. Plants terrestria " aba do dre ot 0 ono 6g c Cor oU Coo yp cL Sin oe 7
T: Stems jointed; sporangia clustered underneath the scales of terminal cone-
(ee E T ET Equisetaceae
7. Stems not jointed; sporangia borne on upper surfaces of leaves...............
! This article is a portion only of a thesis submitted to the Board of Graduate
Studies of livor aed Ho dide in partial fulfillment for the requirements of the
degree of master of sci
Issued April 29, 1933.
ANN. Mo. Bor. GARD., Vor. 20, 1933. (45)
[Vor. 20
46 ANNALS OF THE MISSOURI BOTANICAL GARDEN
8. Bpores all of to mine Mid. i... eoe error Lycopodiaceae
8. Spores of two kinds (larger macrospores and much smaller microspores)... . .
edith eS ON ERA DONORS ET A Wee E Rhode E Selaginellaceae
OPHIOGLOSSACEAE
1. Veins free; sporangia separate in compound inflorescences.......... Botrychium
1. Veins reticulate; sporangia disposed in a solitary spike.......... Ophioglossum
Fig. 1. Key map to distribution chart (fig. 2) showing location of counties in
Missouri.
OPHIOGLOSSUM
1. Sterile segments obtuse, usually solitary; middle areoles long and narrow,
outer hexagonal and containing a few included veinlets...... 1. O. vulgatum
1. Sterile segments cuspidate, 2-5; areoles wide and containing many anasto-
mosing TARNEN 5 isi i443 EAR RAAOAOALARXo EE EAE 0455058785 2. O. Engelmanni
1. Ophioglossum vulgatum L. Sp. Pl.2:1062. 1753. Gray, p.
47; Britton & Brown, ed. 1 and 2, 1: 2.
ATCHISON
HOLT
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) Gheilanthes alabamensis Pieces T 3 4 Lt t
AE t ml i + + LAUS + +
: lanosa + b+ + + ttt + Js: oae $ FEEF 4
5 oystopt eris bulbifera + +44 ++ + + + fe + T fe ark + + ide +
i fragili ttt. B, ET ++ tto +44 ++ + woe rS Ax + + +++ + Xx
j Donnøtaoatsa” puneti lobule +
26 Ea uis etum EFFEEF "m ++ + TRIN + T"
27 incid 1 + + =- + a +
28 praealtum Aat" d ++ +4 ++ ee grits Ms +
29 isoetes Butleri +
30 rae pits ge ; i Š
E melan poda :
32 Lrgopoaiin luein ++ + +
33 broken iiber £ ve ^
34 Ni io A A
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36 Ophioglossum mentenek mao he + F + + F 4-
37 ep huir + =
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$9 Claytonian ps T ++ +
40 galis spectabilis t + + + t t tt + —
41 Pellsos atropurpur Tt tkt + err 6s ee + je t —++ d
42 labella Eq. Rt Ne d + + tE + + + HH
43 Polypodium Le rei Ttt e + + i ortt ++ +++ iia
4 rginianum +++ + - +
45 Polys blend ror on PE H--oc-FEFATLFLGMA + + + + +t + +t+tttt ++4+ + Ta ab +44 +
46 Pteretis nodulosa + ;
47 Pteridium letiusculum NN. + It e t
48 Len X
3elaginella rupestris EA EER kp ; s r FT. i. + 4 [ z
50 Thelypteris Da seRoppu M : 4- Vu
51 arginalis Bas + * ENT ae + t + + a
$2.4, n palustris. ubescens TU + : +
59. sp nulosa 2 :
54 in t di 3
55 Wo MEA dede sn -tCkRR TEM + + ^ P bees. + a + — x
56 woodwardia areolata WAR E ;
Fig. 2. Chart showing distribution of posi by counties: — signifies reported only; +, specimens seen.
vo
t+ + +
+
Me
++
+ ++ +4 tt
+
1933]
PINKERTON—FERNS OF MISSOURI 47
Rootstock short, oblique or erect; common stalk half or more
above ground, constituting 1/3 to 2/5 the length of the plant;
sterile segments sessile, ovate, 1-5 inches long, 0.5-2 inches
broad; basal veins about 9; sporophyll 0.75-2 inches long, apicu-
late. May to August.
Distribution: Nova Scotia to momi bentii to Missouri and Texas. South-
eastern Missouri—swamps or moist me
Specimens examined: Palmer 14716, 1 ges 1 4748, 14766.
2. Ophioglossum Engelmanni Prantl in Ber. Deut. Bot. Ges. 1:
351. 1883. Gray, p. 47; Britton & Brown, ed. 1 and 2, 1: 2.
Rootstock cylindrical; common stalk mostly below ground,
sheathed by persistent leaf-bases; sterile segments sessile, ellip-
tic to ovate, 1-3.5 inches long, 0.5-2 inches broad; basal veins
13 or more, transverse ones oblique; sporophyll 0.5-1 inch long,
apiculate. March to October.
Distribution: South Carolina to Florida, westward to Arizona. Limestone glades,
especially where there is a thin layer of fine rich and damp soil.
Specimens examined: Palmer, Kellogg, Letterman, Engelmann, Bush, Pinkerton,
Eggert, Greenman.
BOTRYCHIUM
. Bud wholly enclosed; cells of epidermis straight; sterile blade petiolate and
ever over 7 inches broad; common stalk mostly underground............
_
1. Bud exposed along one side at base of rootstock; cells of epidermis flexuous;
sterile blade sessile, 2-16 inches broad and nearly as long; common stalk
above ground, half the length of plant................. ec cece teen eens 4
2. Ultimate segments deeply laciniate................. 2. B. obliquum var. dissectum
2. Ultimate segments serrulate-dentate. ......... ccc cece cece cece nnn 3
3. Plant coarse; common stalk very shave underground; frond E ee
aaiae 655 os 6 5 osc s o's 0 0 0 ohh eee 1o 11 RS 1. B. obliquum
3. Plant slender; common stalk up to 2 inches above ground; frond coriaceous
on drying. iesnas sear cine t EERERRES 8. B. obliquum var. Weit
4. Ultimate segments T ripe sporangia mtraw-colored, openi
but slightly in dehiscence.............. 5. B. virginianum var. i A
4. Ultimate segments acute; pa sporangia brown, opening widely in dehiscence.
TC TUMOR CES a Re cee 4. B. virginianum
1. Botrychium obliquum Mühl. in Willd. Sp. Pl. 5: 63. 1810.
Gray, p. 48; Britton & Brown, ed. 1, 1: 3, ed. 2, 1: 5; Eaton,
pl. 20, fig. 2.
B. ternatum Sw. in Schrad. Jour. für die Bot. 2 (1800): 111.
1801
B. ternatum var. obliquum D. C. Eaton, Ferns N. Am. 1:
149. 1879
[Vor. 20
48 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Plant 5-15 inches tall; bud pilose; sterile blade long-stalked,
2-5 inches broad, tripinnatifid or tripinnate below; ultimate
segments obliquely ovate or oblong-lanceolate, acutish terminal
ones elongate; sporophyll long-stalked, usually stout, several-
pinnate. August to October.
Distribution: Maine to Alabama, westward to Michigan and Texas. Moist
woods, neutral soil.
Specimens examined: Palmer, Bush, Kellogg, Letterman, Eggert, Trelease.
2. Botrychium obliquum var. dissectum Prantl in K. Bot.
Gard. Berlin, Jahrb. 3: 342. 1884. Gray, p. 49; Britton &
Brown, ed. 1, 1:3, ed. 2, 1: 5; Eaton, pl. 20, fig. 1.
B. dissectum Spreng. Anleit. 3: 172. 1804.
B. ternatum var. dissectum D. C. Eaton, Ferns N. Am. 1:
150. 1879.
Character of rootstock, fruiting stalks, and texture of the plant
all similar to the species; frond subternately divided, basal
divisions unequally and broadly deltoid, decompound; the upper
and secondary pinnae deltoid-lanceolate, pinnate, with laciniate
or deeply cut pinnules; the ultimate divisions divergent, narrow,
and incised.
Distribution: Maine to Florida, westward to Illinois and Missouri. Rich moist
woods, deeply sha
Specimens examined: Eggert, Letterman, Pinkerton.
3. Botrychium obliquum var. tenuifolium (Underw.) Gilbert
in Fern Bull. 11:99. 1903. Gray, p. 49. Pl. 5, fig. 2.
Plant small, up to 12 inches high; common stalk up to 2 inches
high, not all underground, slender; blade ternate with few divi-
sions, 5-10 em. broad and about 5 em. long; leaves coriaceous;
ultimate segments broad, about 1.5 cm. long, ovate, acutish,
conspicuously serrulate. September and October.
Distribution: Virginia to Arkansas. Southern Missouri—swampy land.
Specimens examined: Dunklin Co., Bush 134, and coll. of Sept. 28 and Oct. 28,
1897, T'release; Butler Co., Bush 3110.
4. Botrychium virginianum (L.) Sw. in Schrad. Jour. für die
Bot. 2 (1800): 111. 1801. Gray, p. 49; Britton & Brown, ed. 1,
1: 4, ed. 2, 1: 6; Eaton, pl. 33.
Common stalk slender, above ground 1/2-2/3 the length of
1933]
PINKERTON—FERNS OF MISSOURI 49
plant; blade sessile or nearly so, membranaceous, ultimate
segments toothed; sporophyll long-stalked, bi-tripinnate. June
and July.
Distribution: New Brunswick to Alabama, westward to Idaho and Texas. Rich
mois woods.
mens examined: Davis, Engelmann, Letterman, Palmer, Bush, Pinkerton,
Eggert, "Kellogg, Trelease, Larsen, Daniels, Emig, Blankenship.
5. Botrychium virginianum var. intermedium Butters in Rho-
dora 19: 207. 1917.
Ultimate segments of the sterile frond spatulate, penultimate
ones ovate, not crowded; segments of the fertile frond opening
wide in dehiscence; sporangia straw-colored, up to 0.8 mm. long.
Specimens examined: Monteer, Shannon Co., Bush 4724; Whiteside, Lincoln Co.,
Davis, coll. of 1910. The writer feels that the species is so generally variable that
this variety is not very distinct.
POLYPODIACEAE
I. Indum lacking. o.oo si eu i serva ng DD rie rx Re GO ee i à 2
1. Indusia present, at least in carly stages, ee cog oka ss os) shen cose tenons 4
2. Fruit dots near margin, becoming panne somewhat protected by over-
lapping leaf margins; lower surfaces of leaves covered with a white powder.
"PUE rn ee Gaye otholaena
2. Fruit dots separate, era absolutely bores COWPOUPNFP Iur 3
3. Fronds linear, once-plunatid. ......... DERI Polypodium
3. Fronds triangular, more Ph once- pinnata e. qua sns ss enda ria Thelypteris
4. Fronds dimorphic E- fronds not MG eu aes + > she vi a eee R PA 5
4; Fronds monomorgnhie « . se- «5.6. s.s-ccdeualege en SO Bees 2 5.6 5s 6 na AE eens 6
5. Sterile fronds pin innatiid; veins anastomosed; fertile segments bipinnate.Onoclea
5. Sterile fronds bipinnatifid; veins free and unbranched; fertile segments
pinnate .eiscooesiocosv-XARRPTTRIBRSRENE vies C++ dys eo Pteretis
6. Indusa MARINA: . «6.6 ns ness 0 ears ne Eade ARES er ERR Fen ke
6. Indusia not marginal. ...... a ETRAS IU. Las cha Sees ens 11
7. Indusia not formed of revolute margins, but cup-shaped and opening ter-
Pcie | socero co ede eR IS EE cen ennstaedtia
7. Indusia formed in part at least by revolute margins. .......... eese 8
8. Indusia definitely interrupted, occurring in is little rounded flaps...
DUMMIES EE ur ddinnium
8. Indusia continuous or only slightly broken. .......... leen
9. Indusia double... ... 60620. vs oss ss sen eee. esos es) E Pteridium
9. Indusia single... cscs cap REA ees RR
10. Blades with a few large, relatively smooth segments.............. s. Pellaea
10. Blades with many small, usually tomentose or hairy segments. ..... m
I1. Bori elongated... eode eorr oie 8 RRS Exod Rn ERA rrr
IL Bori rounded... 50s eas vs cdo + 05605 eee eer RD eee ES.
[Vor. 20
50 ANNALS OF THE MISSOURI BOTANICAL GARDEN
12. Sori dispersed irregularly on lateral veins... 13
JE vM MM Fri Feria. U E HDADUE ORA CR CES RO Camptosorus
IM TE DNE Re T ee en en we ee ee yor eS 14
a ee X Aa ACE ee: eee ee A Asplenium
14. Sori mostly curved over the end of veins..................0000 000s Athyrium
rr ene as Pe ree ee ree ta ae re es 16
15. Indusia attached at base or at one side... 2.1.6... 17
16. Indusia orbicular, attached centrally.............00....0000 0000. Polystichum
16. Indusia reniform, attached at sinus... eese Thelypteris
17. Plants never glandular, but scaly on stipes and rootstocks; segments more
or - hein indusia tapering, attached at one side and becoming obscure
M MAN LrrruaeQureetkgtrrCOra a REAREXSAEXA4 xS ORIREERM PES Cystopteris
17. Plants ver d also scaly; segments obtuse; indusia entire when young,
splitting at top into several stellate segments.................... Wo
ADIANTUM
1. Main rachis unbranched..............0. 00.000 cee eee eee 1. A. Capillus-Veneris
1. Main rachis dichotomously branched with 4-5 pinnae on each side. .2. A. pedatum
1. Adiantum Capillus-Veneris L. Sp. Pl. 2: 1096. 1753.
Gray, p. 36; Britton & Brown, ed. 1, 1: 27, ed. 2, 2: 31; Eaton,
pl. 87. Venus hair.
Rootstock creeping, chaffy; stipe very slender, 3-12 inches
long, black or brownish, slightly scaly; frond ovate-lanceolate,
6-12 inches long, 4-12 inches wide at base; pinnules wedge-
obovate or rhomboid, long-stalked, glabrous, membranaceous,
margins variously incised, veinlets flabellately forking from base;
fruit dots lunate or transversely oblong. June to August.
Distribution: New Jersey to Florida, westward to South Dakota and California.
Moist rocky places, ravines, wet limestone cliffs.
Specimens examined: Kellogg, Daniels, Bush, Palmer, Trelease, Emig, Shepard.
2. Adiantum pedatum L. Sp. Pl. 2: 1095. 1753. Gray, p.
35; Britton & Brown, ed. 1, 1: 27, ed. 2, 2: 31; Eaton, pl. 18.
Maidenhair.
Rootstock long, creeping, chaffy; stipe 9-18 inches long,
shining, dark brown to black, slightly scaly at base, once-forked
at summit, each division bearing on one side only several pinnate
divisions (occasionally tri-forked); blade reniform-orbicular,
8-18 inches broad, membranaceous, glabrous; pinnules short-
stalked, oblong, triangular or end ones fan-shaped; lower margin
entire, all veins proceeding from it, upper margin lobed. June
to August.
1933]
PINKERTON—FERNS OF MISSOURI 51
Distribution: Nova Scotia and Quebec to Georgia, westward to Alaska and
California. Moist rich woods.
Specimens examined: Bush, Davis, Merge Palmer, Trelease, Pinkerton,
Woodson, Kellogg, Broadhead, Emig, Macken:
ASPLENIUM
1. Blades pinnatifid, or only lower segments pinnate; apices long-attenuate........ 2
1. Blades 1-3-pinnate; apices not long-attenuate. ......... selle 3
2. Blades membranaceous; lower midribs black and shining on under side;
apices crenate, sometimes proliferous..................000005 8. A. ebenoides
2. p E ei xs green, herbaceous; prolongations sinuous-
BL NOt OOM NOUN es hx i cance EEEN Xt E uo 4. nnatifidum
3. Blades Pak only; pee of regular shape; margins not deeply Fahey
CCC HBC H HCCC ETO HHS SPO HHH CTH FE HHC OCHO HH EHH HOH HHC CHE HHH HH ETHEH HHH HAE OEE
Sousa os kk ic kN WEEE E 64 nd + err OU Oe RR TL
4. Pinnae mostly roundish, not auriculate...................04. 7. A. Trichomanes
4. Pinnae oblong or lanceolate, auriculate.... 2.2.0... ec ce ee cece ee ees 5
5. E and rachises shiny black, slender; pinnae mostly opposite; sori few and
om 000 than COBE. DEEP DES ^ . A. resiliens
5. Stives pe rachises shiny "reddish brown, coarse; pinnae m sitéciinte;
RH numerous HORT COSlAO, . oos or rn RENS . A. platyneuron
6. Stipes and rachises green throughout; ultimate segments ew cuneate;
mendo inbrisie....... oce ono tae eee ere De xs A. cryptolepis
6. Stipes and rachises shining chestnut-brown; lower pinnae bd into obtuse
sumentes; margins cronate. .. s. S. QUITE s erede 1. A. Bradleyi
1. Asplenium Bradleyi D. C. Eaton, Bull. Torr. Bot. Club 4:
11. 1873. Gray, p. 39; Britton & Brown, ed. 1, 1: 26, ed. 2, 1:
30; Eaton, pl. 51, figs, 4-8.
Rootstock short, covered with narrow acuminate blackish-
fuscous scales; finds 4-7 inches high, oblong-lanceolate; stipes
2-3.5 inches long, dark chestnut and shining, tufted, slender;
rachises brown, or green above; pinnae numerous, lower ones
no larger than the middle ones, obtuse or acutish, toothed, in
the largest fronds pinnatifid into oblong lobes which are toothed
at apices; sori short, borne near midveins, becoming confluent;
indusia membranaceous, persistent. July to September.
Distribution: New York to Georgia, westward to Missouri and Arkansas. On
sandstone or chert outcrops—comparatively rare and local
Specimens examined: Palmer, Bush, Trelease, Mackenzie, Greene, Pinkerton, Van
Dugen, Shepard
2. Asplenium cryptolepis (L.) Fernald in Rhodora 30: 37.
[Vor. 20
52 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1928. Gray, p. 39; Britton & Brown, ed. 1, 1: 25, ed. 2, 1: 29;
Eaton, pl. 15, fig. 1.
Asplenium Ruta-muraria L. Sp. Pl. 2: 1081. 1753.
Rootstock short, creeping, entangled, tufted; fronds 1-6
inches tall; stipes and rachises entirely green or slightly brown
at base; blades deltoid-ovate, smooth, subcoriaceous, bi-tri-
pinnate; ultimate segments few, stalked, 3-14 mm. long, narrowly
cuneate to roundish obovate; margins deeply fimbriate; veins
flabellate, no midveins; sori few (2-4 per pinna), oblong, covering
whole segment when mature; indusia delicate with ciliated margin.
July to September.
Distribution: Vermont to Georgia, westward to Illinois and Missouri. Shaded
limestone cliffs—scarce.
Specimens examined: T'release, Palmer, Bush, Russell.
3. Asplenium ebenoides R. R. Scott, Berkeley in Roy. Hort.
Soc. Jour. N. S. 1: 137. 1866. Gray, p. 38; Britton & Brown,
ed. 1, 1: 23, ed. 2, 1: 26; Eaton, pl. 4, fig. 2.
Rootstock short, creeping, chaffy; fronds up to a foot high;
stipes tufted, 4-9 inches long, young ones reddish brown, older
ones black, shining, slender; lower rachises dark and shining
underneath; blades firm-membranaceous, triangular-lanceolate,
variable, 3-12 inches long, 1-3 inches at base, tapering to a long-
acuminate apex which may become proliferous, lowest divisions
distinct, shorter; sori numerous throughout, mostly single and
opening obliquely upwards. August and September.
Distribution: Vermont to Alabama, westward to Missouri. Limestone—rare.
Specimens examined: T'release, Russell.
4. Asplenium pinnatifidum Nutt. Gen. 2: 251. 1818. Gray,
p. 38; Britton & Brown, ed. 1, 1: 22, ed. 2, 1: 27; Eaton, pl. 8,
. 2.
Rootstock short, creeping, branched, chaffy; fronds 6-9 inches
high; stipes brownish near base and green above, clustered;
blades 2-5 inches long, subcoriaceous, herbaceous, lanceolate-
acuminate from broad and sub-hastate base, pinnatifid; the
basal pinnae sometimes long-attenuate, lower lobes of pinnae
roundish-ovate, margin crenate, upper pinnae gradually smaller
and more adnate to winged rachises; sori straight, many, be-
1933]
PINKERTON—FERNS OF MISSOURI 53
coming confluent with age, mostly solitary, occurring also on the
slender prolongation. July to October.
Distribution: Massachusetts to Alabama, westward to Missouri; recorded from
Georgia and Arkansas. On La Motte sandstone only, in shaded crevices of cliffs.
Specimens examined: T'release, Engelmann, Russell, Pinkerton.
5. Asplenium platyneuron? (L.) Oakes in D. C. Eaton, Ferns
N. Am. 1: 24. 1879. Gray, p. 39; Britton & Brown, ed. 1, 1:
23, ed. 2, 1: 27; Eaton, pl. 4, fig. 1. Ebony spleenwort.
Asplenium ebeneum Ait. Hort. Kew. 3: 462. 1789.
Fronds 4-20 inches high, fertile ones tall and upright, sterile
ones short and spreading; stipes and rachises reddish brown,
shining, rather thick; blades linear-oblanceolate, tapering at base,
once-pinnate; pinnae 20—40 pairs, lanceolate, 0.5-1.5 inches long,
alternate, sessile, auricled on upper or both sides of base, and
more or less overlapping rachis; sori 8-15 in number, nearer
midveins than margins, becoming confluent. July to September.
Distribution: Vermont to Alabama, westward to Texas and Oklahoma; recorded
from Ontario and Colorado. Rocky open woods, preferring alkaline soil.
Specimens examined: Davis, Kellogg, Engelmann, Eggert, Trelease, Bush, Palmer,
Daniels, Pinkerton, Woodson, Russell, Mann, Dewart, Emig, Krause, Meek.
6. Asplenium resiliens Kunze, Linnaea 18: 331. 1844. Gray,
p. 39; Britton & Brown, ed. 1, 1: 23, ed. 2, 1:27; Eaton, pl. 36, figs.
5&6
Asplenium parvulum Mart. & Gal. in Mém. Acad. Brux. 15:
60. 1842.
Rootstocks with black scales; fronds 4-12.5 inches long;
stipes and rachises black and shining, slender; blades normally
linear-oblanceolate, pinnate; pinnae 4-12 mm. long, mostly
opposite, nearly sessile, upper edges auricled and on lower pinnae
both edges auricled, deflexed; blades widest in middle; margins
mostly entire or slightly crenate, tendency to incurve slightly;
fruit dots nearer outer margins than midribs, nearly parallel to
the midribs, oblong, few, sometimes becoming confluent. June
to October.
2 Var. incisum (E. C. Howe) Robinson in Rhodora 10: 29. 1908, has very brittle
stipes and the pinnae deeply pinnatifid. This appears to be merely an ecological
variation
OL. 20
54 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: Massachusetts to Florida, westward to New Mexico. Limestone
cliffs.
Specimens examined: T'release, Bush, Palmer, Russell, Pinkerton, Drouet.
7. Asplenium Trichomanes L. Sp. Pl. 2: 1080. 1753. Gray,
p. 39; Britton & Brown, ed. 1, 1: 24, ed. 2, 1: 28; Eaton, pl. 36,
fig. 1. Maidenhair spleenwort.
Rootstocks nearly erect, inconspicuously chaffy with narrow
black scales; fronds 3-6 inches high; stipes slender, densely
tufted, brownish-purple, polished, rachis similar to tip; blades
once-pinnate, linear; pinnae 3-7 mm. long, herbaceous, mostly
opposite, roundish, crenate margins, obliquely wedge-truncate
at base, attached by narrow points; sori medial or nearer the
midveins than margins, 3-6 pairs on outer sides of veins, be-
coming confluent; indusia membranaceous. July to September.
Distribution: Ontario to Alabama, westward to British Columbia and California;
widely distributed but local. Sandstone rocks where plenty of water is available.
Specimens examined: Kellogg, Letterman, Eggert, Engelmann, Palmer, Russell,
Pinkerton, Trelease, Morrison, Rickett, Mackenzie, Blankenshi ip.
ATHYRIUM
1. Fronds simply pinnate...............0000 s s sss esses 1. A. angustifolium
1; Fronds more NB DEIN. Loose Ayo eese ea ovo Pa ETSI 2
2. Fronds deeply bipinnatifid, margins lightly serrate-crenate. ... . » A. acrostichoides
2. Fronds usually tripinnatifid, margins deeply and irregularly incised...........
3. Rhizomes creeping, not densely covered with persistent feres fronds
widest near base; indusia with glandular cilia; spores nigrescent, wrinkled.
E nh oor eee eRe eT Rad da qoa Ro OE d 3. A. asplenioides
3. Rhizomes horizontal, completely concealed by thick fleshy bases of old
fronds; fronds widest near middle; indusia toothed or short-ciliate, never
glandular; spores yellow, slightly papillate.................. 4. A. angustum
1. Athyrium angustifolium (Michx.) Milde in Bot. Zeit. 48:
376. 1886. Gray, p. 39; Britton & Brown, ed. 1, 1: 24, ed. 2, 1:
28; Eaton, pl. 56, fig. 1. Narrow-leaved spleenwort.
Asplenium pycnocarpon Spreng. Anleit. 3: 112. 1804.
Asplenium angustifolium Michx. Fl. Bor. Am. 2: 265. 1803.
Athyrium pycnocarpon Tidestrom, Elys. Marianum, p. 36.
1906.
Rootstocks stout, creeping, with many long, branched rootlets;
stipes green except for brown base; fronds 1-2.5 feet long, mem-
branaceous, herbaceous, pinnate; pinnae 2-5 inches long, 20-30
pairs, short-stalked, linear-oblong, attenuate, margins slightly
1933]
PINKERTON—FERNS OF MISSOURI 55
wavy; fertile pinnae near top, narrower and shorter; sori 20-30
pairs, linear, slightly curved, lying along outer of bifurcated
veins; indusia firm, convex, concealed by strongly confluent sori
at maturity. August and September.
Distribution: Quebec to Georgia, westward to Michigan and Missouri; recorded
from Kansas and Minnesota. Moist woods and shaded ravines, reported occasion-
ally on sandy soil.
ecimens examined: Davis, Kellogg, Palmer, Eggert, Trelease, Bush, Pinkerton,
Daniels, Emig, Glatfelter.
2. Athyrium acrostichoides (Sw.) Diels in Engl. & Prantl,
Nat. Pfl. 14: 223. 1899. Gray, p. 39; Britton & Brown, ed. 1, 1:
26, ed. 2, 1: 80; Eaton, pl. 50. Silvery spleenwort.
Asplenium acrostichoides Sw. in Schrad. Jour. fiir die Bot.
2: (1800): 54. 1801.
Asplenium thelypteroides Michx. Fl. Bor. Am. 2: 265. 1803.
Athyrium thelypteroides (Michx.) Desv. in Mem. Soc. Linn.
Paris [Prodr. p. 266] 6: 266. 1827.
Rootstocks creeping, horizontal; stipes 8-16 inches long,
straw-colored, herbaceous, with a few scales on lower portion;
blades lanceolate to ovate-oblong, 1-3 feet long and 6-12 inches
broad, narrowed to base, deeply bipinnatifid; ultimate segments
distinct, obtuse; margins slightly serrate-crenate; sori 3-6 pairs
per segment, arranged more or less evenly along lateral veins,
mostly straight, oblong, some double. August to October.
Distribution: New Brunswick to Georgia, westward to Missouri; recorded north-
ward to Minnesota. Rich moist woods, or moist sandy soil.
Specimens examined: Palmer, Eggert, Davis, Letterman, Broadhead.
3. Athyrium asplenioides? (Michx.) Desv. in Mem. Soc. Linn.
Paris [Prodr. p. 266] 6: 266. 1827. Gray, p. 40; Britton &
Brown, ed. 1, 1: 26, ed. 2, 1: 30; Butters in Rhodora 19: 169.
1917.
Asplenium Athyrium Spreng. Anleit. 3: 113. 1804.
Athyrium Filix-foemina (L.) Roth in Rómer's Arch. f. Bot.
21:106. 1799.
Rhizomes horizontally creeping, partially covered by short
persistent leaf bases, the whole structure 1-1.5 cm. in diameter,
*'lhis species and A. angustum are very difficult to distinguish. It is often
necessary to have the whole plant, fruiting and not too mature, to be absolutely
certain. I have taken the character of the spore as my ultimate criterion.
[Vor. 20
56 ANNALS OF THE MISSOURI BOTANICAL GARDEN
with conspicuous projections of new growths before fronds of the
current season; stipes long, about equal to the deltoid lanceolate
fronds; young erowtha covered with scales soon deciduous, these
small and light tan-colored, cell walls thin and inconspicuous;
fronds bipinnate to tripinnatifid, second pair of pinnae commonly
the longest, the lowest only slightly shorter; pinnae narrower at
base; pinnules variously incised but apex more or less obtuse
due to the venation, since two veins usually end on the same
level; sori longer and narrower than in A. angustum, on the
anterior side of anterior vein of each lobe of pinnule, and some-
times on the lower veins, of typical athyroid type, rarely double;
young indusia ciliate with multicellular glandular-tipped hairs
nearly disappearing at maturity, leaving quite even margins;
sporangia stalk frequently supplied with a glandular hair; spores
furnished with a somewhat nigrescent, wrinkled exospore. July
to October.
Distribution: pe stent to Florida, westward to Missouri and Texas. Shaded
rich woods or cliffs, sandy so
Specimens examined: Kellogg, Trelease, Palmer, Bush, Soulard, Blankenship.
4. Athyrium angustum* (Willd.) Presl in Rel. Haenk. 1: 39.
1825. Gray, p. 40; Britton & Brown, ed. 1, 1: 26, ed. 2, 1: 30;
Butters in Rhodora 19: 169. 1917.
Asplenium Michauzii Spreng. Syst. 4: 88. 1827.
Asplenium Filix-foemina (L.) Roth in Rómer's Arch. f. Bot.
2': 106. 1799.
Rhizomes horizontal and somewhat oblique, condensed and
completely covered by thick fleshy bases, 2-5 cm. in diameter;
stipes up to half as long as the frond, a moderate number of
scales persistent, 1 x 1.5 mm., dark opaque with thick darker
cell walls and narrow cells; fronds bi-tripinnatifid, middle of
fronds widest and lower pinnae much shorter and often deflexed,
often large forms and polymorphic; pinnae not narrower at base;
pinnules and segments with acute apices, due to one vein ending
considerably beyond any of its neighbors; fertile fronds consider-
! Butters describes two varieties, namely: var. elatius and var. rubellum, distin-
guished chiefly by the dimorphism and thicker texture of the former (a sun form
and the monomorphism and thinner texture of the latter (a shade form). He claims
that the latter has a more northerly distribution; but in Missouri the two seem to be
purely ecological variations and scarcely worth varietal rank.
1933]
PINKERTON—FERNS OF MISSOURI 57
ably narrower and more acute than sterile ones; sori short and
wide, of typical athyroid type; indusia never glandular but
persistently short-ciliate; sporangia stalk rarely bearing glandular
hair; spores bright yellow and slightly papillate, no exospore and
not wrinkled. June to October.
Distribution: Labrador to Pennsylvania, westward to South Dakota and Missouri.
Rich woods.
Specimens examined: Bush, Davis, Palmer, Engelmann, Kellogg, Eggert, Russell.
CAMPTOSORUS
1. Camptosorus rhizophyllus (L.) Link, Hort. Berol. 2: 69.
1833. Gray, p. 40; Britton & Brown, ed. 1, 1: 21, ed. 2, 1: 26;
Eaton, pl. 8, fig. 1. Walking fern.
Rootstocks short, creeping; stipes grouped in tufts, spreading,
green, fleshy, 1-6 inches long; blades evergreen, subcoriaceous,
4-12 inches long, base auriculate, cordate or hastate, apex
attenuate and filiform, rooting at tips (or from auricles); sori
numerous, straight or slightly curved, single and on inside of
and parallel to veins, near midribs, on both sides of outer veins
and becoming confluent at exterior tips; indusia membranaceous.
All year.
Distribution: Ontario to Alabama, westward to oe and Oklahoma. Lime-
stone rocks, usually associated with Entodon (a m
Specimens examined: Bush, Davis, Engelmann, aie. Kellogg, Daniels, Palmer,
Pinkerton, Dewart, Russell, Rickett, Weller.
CHEILANTHES
I Fronds relatively amootih. i... 4o 444 ae scan eee 1. C. alabamensis
1. Fronds hairy or tomentose "ie
2. Plants small, 2-6 inches tall, matted; mature stipes nearly glabrous. A s
2. Plants taller, 4-20 inches, not matted; stipes hirsute.............. bibe
s» e**?289€829wó9À€9€v90466$o.0e*9»9e0e999259226»0698&€9484€9"9»2929252299
1. Cheilanthes alabamensis (Buckl. Kunze in Linnaea 20: 4.
1847. Gray, p. 36; Britton & Brown, ed. 1, 1: 30, ed. 2, 1: 34;
Eaton, pl. 57, fig. 7.
Rootstocks creeping, short, slender, with dark ferruginous
scales; stipes black, wiry, slightly villous at base due to fine
rusty scales 3-7 mm. long; blades scabrous to smooth, 2-10 inches
long, lanceolate, bipinnate; pinnae acuminate, lower ones shorter
than those above; pinnules often auriculate; indusia broad, pale
but firm, frequently broken by incision of pinnules. August to
October.
[Vor. 20
58 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: Florida, westward to Missouri and Arizona. Limestone cliffs.
Specimens examined: Kellogg, Palmer, Bush.
2. Cheilanthes Feei Moore, Ind. Fil. Gen. 38. 1857. Gray,
p. 36; Britton & Brown, ed. 2, 1: 34; Eaton, pl. 6, fig. 1.
Cheilanthes lanuginosa Nutt. in Hk. Sp. Fil. 2: 99. 1852.
Rootstocks short, clothed with narrow scales of black centers
and thin brown edges; fronds 2-6 inches tall; stipes densely
tufted, black or brown, originally woolly, becoming glabrous
when mature; blades bi-tripinnate, slightly tomentose above
and woolly below ; pinnules divided into minute rounded segments,
densely crowded; indusia herbaceous, continuous. July to Oc-
tober.
Distribution: Wisconsin to Texas, westward to Nevada and Arizona. Limestone
cliffs in dry and exposed localities.
Specimens examined: Davis, Kellogg, Engelmann, Trelease, Bush, Palmer, Daniels,
Pinkerton, Morrison, U phof.
3. Cheilanthes lanosa (Michx.) Watt in Jour. Bot. 12: 48.
1874. Gray, p. 36; Britton & Brown, ed. 1, 1: 31, ed. 2, 1: 34;
Eaton, pl. 2, fig. 2.
Cheilanthes vestita Sw. Syn. Fil. 128. 1806.
Rootstocks short, creeping, with pale brown scales; fronds
4-16 inches long; stipes wiry, dark brown, hirsute; blades her-
baceous, bipinnate; pinnae hirsute and somewhat glandular; in-
dusia inconspicuous, discontinuous. July to September.
Distribution: New York to Alabama, westward to Oklahoma; recorded from
— and Texas. Sandstone rocks, usually in dry and exposed places
Specimens examined: Kellogg, Eggert, Letterman, Palmer, Broadhead, Pinkerion,
Muller, Glatfelter, Mackenzie, Swallow, Emig, Blankenship, Link
CYSTOPTERIS
1. Pinnae short-stalked on rachises; pinnules at least narrowed at points of
attachment; segments ovate, acute, usually variously incised; indusia
ne err rr er CR e i cee 1. C. fragilis
1. Pinnae sessile on rachises; pinnules oblong, obtuse, regularly toothed; indusia
round or perona E AEE E E EE N E ID LE RA 2 Xo T
2. Fronds T oo EEEE "T 2. C. bulbifera
2. Fronds not estes NILUS Ead RR 8. C. bulbifera var. horizontalis
1. Cystopteris fragilis’ (L.) Bernh. in Schrad. Neues Jour.
5 An exceedingly variable species. Hybridization with C. bulbifera might account
for some of the aberration. Forms bearing bulblets have been included under this
group, as that character does not seem to be a constant feature for C. bulbifera alone.
1933]
PINKERTON—FERNS OF MISSOURI 59
Bot. 1*: 27. 1806. Gray, p. 43; Britton & Brown, ed. Leas MA
ed. 2, 1: 15; Eaton, pl. 53, fig. 1
Filiz Pots Under. Nat. Tesis ed. 6, 119. 1900.
Rootstocks elongated, often 4-5 inches long, or shorter and
condensed, slender, but covered with persistent leaf-bases, chaffy
at apex, scales delicate, ovate, acuminate, ferruginous; stipes
in a dense cluster, slender, brittle, 4-6 inches long; blades 6-8
inches long and about half as wide, thin and membranaceous,
ovate-lanceolate; basal pinnae commonly narrower than the
second and third pairs, apparently bipinnate but segments
usually connected by narrowly winged midribs, segments round-
ish-oval to ovate to rhomboid-ovate to ovate-lanceolate, toothed,
dentate or irregularly laciniate; veinlets pinnately arranged on
midveins, lower ones forked; sori small, roundish, seated on
middle of veins nearest midrib; indusia delicate, rounded, ovate,
or occasionally with narrow beak-like points, concealed by mature
sporangia; lower pinnae often sterile. May to July.
Distribution: Cosmopolitan. Rocky soil, moist woods, preferring alkaline soil.
ecimens examined: Davis, Jnosiptana: Trelease, Bush, Palmer, Pinkerton,
Eggert, Kellogg, Woodson, Harrison, Mann, Daniels, Blankenship, Duncan, Williams.
2. Cystopteris bulbifera (L.) Bernh. in Schrad. Neues Jour.
Bot. 1*: 10. 1806. Gray, p. 43; Britton & Brown, ed. 1, 1: 12,
ed. 2, 1: 15; Eaton, pl. 58, fig. 13.
Filix bulbifera Underw. Nat. Ferns, ed. 6, 119. 1900.
Rootstocks seldom over one inch long, chaffy at apex and
covered with persistent leaf-bases; stipes slender, rather brittle,
6-10 inches long; blades submembranaceous but of a brittle
rigidity, triangular-attenuate, 1—4 feet long, 3-5 inches broad
at base; pinnae attenuated upward, bipinnate at base, upper
pinnules attached by winged rachises; pinnae numerous (up to
40 pairs), oblong; pinnules oblong, obtuse, pinnately lobed; sori
numerous, all pinnae fertile, arranged in rows along each side
of midveins of pinnules, placed on the lowest superior veinlet of
each group near its middle and so near the midvein; indusia
truncate and fragile, covered by mature sporangia; bulblets some-
times present on the under side of the frond attached near the
base of or on the pinnae. July to August.
Distribution: Newfoundland to Georgia, westward to Michigan and Arkansas;
[Vor. 20
60 ANNALS OF THE MISSOURI BOTANICAL GARDEN
recorded pm Utah and Arizona. Rocky soil, preferring limestone, in moist shady
situation
nel examined: Bush, Davis, Kellogg, Trelease, Palmer, Pinkerton, Daniels,
Letterman
3. Cystopteris bulbifera var. horizontalis Lawson in Bot. Soc.
Edinb. Trans. 8:40. 1866. Pl. 5, fig. 1.
Fronds triangular-lanceolate, broad at base, not more than
three or four times longer than broad; pinnae horizontal; lowest
pinnules often quite broad with irregularly cut lobes and bearing
numerous medium-sized sori along the lateral veins, sometimes
almost tripinnate, or pinnules irregularly lobed, merely ovate-
lanceolate. May to October.
Distribution: southern Missouri and northern Arkansas. Damp limestone bluffs.
Specimens examined: Ilasco, Ralls Co., Davis 2663; Sulphur Springs, Jefferson Co.,
coll. of Oct. 23, 1898, T'release; Terre Bleue Cr., Ste. Genevieve Co., coll. of Aug. 29,
30, 1898, T'release; Lesterville, Reynolds Co., coll. of June 5, 1929, Kellogg; Tecumseh,
Ozark Co., Palmer 32896.
DENNSTAEDTIA
1. Dennstaedtia punctilobula (Michx.) Moore, Ind. Fil. Gen. 97.
1857. Gray, p. 45; Britton & Brown, ed. 1, 1: 12, ed. 2, 1: 14;
Eaton, pl. 44.
Dicksonia pilosiuscula Willd. Enum. 1076. 1809.
Dicksonia punctilobula Hk. Sp. Fil. 1: 79. 1846.
Rootstocks extensively creeping, slender, scaleless but finely
hairy at tips, irregularly branching with many long slender root-
lets; stipes rather stout, light brown, chaffless, slightly puberulent ;
blades 1-3 feet long, ovate-lanceolate, acuminate, delicately
herbaceous, hairy and minutely glandular, tripinnatifid; pinnae
numerous, lanceolate, pointed, second pair a little longer than
first; pinnules adnate to rachis and usually decurrent on it,
rhomboid-ovate, pinnatifid into oblong and obtuse cut-toothed
lobes; sori minute, on upper margins of the lobes of the pinnules;
indusia cup-like, delicate. August.
Distribution: New Brunswick to Georgia, westward to Illinois and Missouri.
Crevices of La Motte sandstone, rare.
Specimens examined: T'release, Russell, Eggert, Pinkerton.
NOTHOLAENA
1. Notholaena dealbata (Pursh) Kunze in Amer. Jour. Sci. II.
6: 82. 1848, as Nothochlaena. Gray, p. 35; Britton & Brown,
ed. 1, 1: 32, ed. 2, 1: 35; Eaton, pl. 9, fig. 2
1933]
PINKERTON—FERNS OF MISSOURI 61
Cheilanthes dealbata Pursh, Fl. Am. Sept. 2: 671. 1814.
Nothochlaena pulchella Kunze in Bot. Zeit. 1: 633. 1848.
Pellaea dealbata (Pursh) Prantl in Engler's Bot. Jahrb. 3:
417. 1882.
Notholaena nivea var. dealbata Davenp. in Cat. Davenp.
Herb. Suppl. 44. 1883.
Rootstocks short, creeping, with narrow brown chaffy scales;
stipes 1—4 inches long, tufted, wiry, slender, copper-brown, as are
the rachises; blades 2-4 inches long, broadly deltoid-ovate, 4—5-
pinnate, all but ultimate segments alternate, those sometimes
opposite; segments obovate-oval and entire or several-lobed, 1-2
mm. broad, upper surfaces pale green, coriaceous, lower white
and powdery, giving a silvery appearance; sporangia seated on
upper portions of the veins; no indusia, but a protection afforded
by the slightly turned-back margins. June to September.
Distribution: Missouri and Kansas to central Texas; recorded from Nebraska.
Dry calcareous rocks.
Specimens examined: Daniels, Bush, Palmer, Blankenship.
ONOCLEA
1. Onoclea sensibilis L. Sp. Pl. 2: 1062. 1753. Gray, p. 45;
Britton & Brown, ed. 1, 1: 9, ed. 2, 1: 11; Eaton, pl. 72, fig. 1.
Sensitive fern.
Rootstocks slender, creeping, rooting freely and often forking;
stipes coarse, straw-like, hollow, flattened, light brown when dry;
sterile blades triangular or triovate, midribs winged, widening
toward the apex, sinuses rounded; lowest segments broadly
lanceolate, herbaceous, sensitive to frost; veins conspicuous, re-
ticulate; margins variously rounded, lobed, toothed, serrate or
acute; fertile fronds 12-18 inches long, pinnate, contracted; each
segment a pouch filled with several sporangia; delicate hood-like
indusia. August to November.
Distribution: Newfoundland to Florida, westward to Kansas. Swamps or damp
rich soil.
Specimens examined: Bush, Davis, Eggert, Palmer, Morrison, M uller, Daniels,
Blankenship, Mackenzie, Williams.
PELLAEA
1. Pinnae dichotomously branched at apex........ 3. P. atropurpurea var. cristata
L. Pinnae not branched at apex......«4 4 4WANAAE ee nese a error 2
[Vor. 20
62 ANNALS OF THE MISSOURI BOTANICAL GARDEN
2. Stipes M: reddish-brown; pinnae usually membranaceous, pale green,
ore or s^ pec at both corners and tending to i at bases
into i Lo ee Os ba ks pad. new a kaw aden wad ees . P. glabella
2. Stipes scabrous, Sale scale sis. pinnae coriaceous, blue-green, ‘knee
And seldom PVN. 6555s 5.60504 ocn iw ook eh 2. P. atropurpurea
1. Pellaea glabella Mett., Kuhn in Linnaea 36: 87. 1869.
Gray, p. 37; Britton & Brown, ed. 2, 1: 33.
Pellaea atropurpurea var. Bushii Mackenzie, Flora Jackson
County, Mo., p. 5. 1902
Stipes and rachises brownish-red, smooth or but slightly hairy;
fronds simply pinnate above, the lower ternate or rarely quinate
(the entire pinnule seems to be breaking at the base—a possible
tendency toward compound pinnules) ; pinnae membranaceous to
coriaceous but always pale green. April to October.
Distribution: Ontario and Vermont to Pennsylvania, westward to South Dakota
and northern Arkansas. Exposed high places on limestone cliffs
Specimens examined: Davis, Eggert, Kellogg, Bush, Palmer, Pinkerton, Trelease.
2. Pellaea atropurpurea (L.) Link, Fil. Sp. in Hort. Berol. 59.
1841. Gray, p. 37; Britton & Brown, ed. 1, 1: 29, ed. 2, 1: 33;
Eaton, pl. 54, fig. 4.
Rootstoek short, densely covered with rusty scales about 2
mm. in length; fronds 4-12 inches long, pinnate or below bipinnate,
coriaceous; fertile segments linear, more or less pointed at apex;
sterile segments approaching oval, shortly stalked; veins obscure;
continuous indusia of reflexed margins. June to September.
Distribution: Connecticut to Florida, westward to South Dakota and Texas.
Pinkerton, Kellogg, Daniels.
3. Pellaea atropurpurea var. cristata Trel. in Rept. Mo. Bot.
Gard. 12: 77. 1901. Gray, p. 37.
Pinnae dichotomously forked.
Distribution: known only from Eureka, Missouri. Limestone.
Specimens examined: Eureka, 1899, Pauls.
POLYPODIUM
. Blades smooth, green... sess 1. P. virginianum
: Blades densely hun a | ERETETITIETTCTT TT TT TOIT S.P. elige
1933]
PINKERTON—FERNS OF MISSOURI 63
1. Polypodium virginianum L. Sp. Pl. 2: 1085. 1753. Gray,
p. 34; Britton & Brown, ed. 1, 1: 32, ed. 2, 1: 36; Eaton, pl. 31,
fig. 1. Common polypody.
Polypodium vulgare L. Sp. Pl. 2: 1085. 1753.
Rootstocks close to surface of soil, covered with chaffy, red-
brown scales; stipes smooth, herbaceous, light green; blades ovate-
oblong or narrowly oblong, subcoriaceous or chartaceous, ever-
green, simple, deeply pinnatifid, smooth; segments linear-oblong,
obtuse or slightly acute, crenulate and serrate, sinuses rounded,
alternate, margins obscurely dentate; sorilarge, naked. July.
pde Newfoundland to Florida, westward to Minnesota and Arkansas.
sandstone or sandy soil.
[OE HA examined: Eggert, Russell, Engelmann, Trelease, Pinkerton, Letterman,
Greene
2. Polypodium polypodioides (L.) Hitche. in Rept. Mo. Bot.
Gard. 4:156. 1893. Gray, p. 34; Britton & Brown, ed. 1, 1:33,
ed. 2, 1: 36; Eaton, pl. 26, fig. 2. Gray polypody.
Polypodium incanum Sw. Fl. Ind. Occ. 3: 1645. 1806.
Rootstocks woody, covered with small dark brown scales; stipes
1-4 inches long, slender, bearing peltate ovate scales with dark
brown centers; blades oblong-lanceolate, pinnate; segments ob-
long, obtuse, entire, sessile, separated by rounded sinuses, alter-
nate, upper surfaces smooth or with few scales, lower densely scaly;
fruit dots small and naked. July to September.
Distribution: Massachusetts to Florida, westward to Missouri and Texas. Usually
an epiphyte on swamp trees, particularly Taxodium distichum; also on sandstone
rocks.
Specimens examined: Eggert, Trelease, Bush, Pinkerton, Rickett, Kellogg, Mac-
kenzie, Meek.
POLYSTICHUM
1. Margins serrulate; fertile pinnae contracted; sori confluent... . 1. P. acrostichoides
1. Margins deeply toothed or pinnatifid; fertile pinnae scarcely contracted; sori
not confluent and appearing only on tips of lower pinnae..............
a seti cule YR e ir e onus hi ERE 2. P. acrostichoides var. incisum
1. Polystichum acrostichoides (Michx.) Schott, Gen. Fil. 17.
1834. Gray, p. 40; Britton & Brown, ed. 1, 1: 14, ed. 2, 1: 16;
Eaton, pl. 34. Christmas fern.
Aspidium acrostichoides Sw. Syn. Fil. 44. 1806.
Dryopteris acrostichoides Kuntze, Rev. Gen. Pl.2:812. 1891.
[Vor. 20
64 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Rootstocks stout, creeping, with persistent leaf-bases; stipes
densely tufted, with chaff of large golden-brown scales, 5-7 mm.
wide; blades lanceolate, pinnate, 1-2 feet long, 3-5 inches wide,
rigid, evergreen, subcoriaceous; pinnae numerous, 1-3 inches
long, oblong-lanceolate, short-stalked, upwardly falcate or lowest
slightly deflexed, apex acutish, upper sides auriculate; margins
serrulate to incised with incurved bristle-pointed teeth; veins free,
branching three to four times; upper pinnae of fertile fronds more
or less contracted and heavily soriferous; sori terminal on lower
veinlets in 2-4 rows, becoming confluent with age; indusia round,
indurated, not glandular, persistent. July to August.
Distribution: Maine to Florida, westward to Michigan and Texas; recorded from
Nova Scotia, Shady hillsides of ravines, in rich soil which is interspersed with
on.
Specimens examined: Bush, — Eggert, Trelease, Palmer, Pinkerton, Emig,
Kellogg, Daniels, Thomas, Mackenzie
2. Polystichum acrostichoides var. incisum Gray, Man. Bot.
ed. 1, 632. 1848. Gray, p. 40; Britton & Brown, ed. 1, 1: 14,
ed. 2, 1: 16.
Polystichum acrostichoides var. Schweinitzii (Beck) Small in
Bull. Torr. Bot. Club 20: 464. i
Aspidium schweinitzii Beck, Bot. North. & Mid. States, ed.
1, 449.
Aspidium acrostichoides var. incisum D. C. Eaton, Ferns N.
Am. 1: 258. 1879.
Segments few and distant, large, irregularly incised; upper
pinnae covered by confluent sori, lower ones fertile at tips only,
sori large; veins numerous, frequently overlapping to form ir-
regular areoles.
Distribution: This seems to be nearly as common as the type in southern Missouri
and northern Arkansas; but since there appears to be a series of intermediate forms
between the two I have not attempted to separate the individual ranges.
PTERETIS
1. Pteretis nodulosa (Michx.) Nieuwl. in Am. Midl. Nat. 4:
334. 1916. Gray, p. 45; Britton & Brown, ed. 1, 1: 9, ed. 2, 1:
11; Eaton, pl. 73. Ostrich fern.
Osmunda Struthiopteris L. Sp. Pl. 2: 1066. 1753.
Onoclea Struthiopteris Hoffm. Deutsch. Fl. 2:11. 1795.
1933]
PINKERTON—FERNS OF MISSOURI 65
Struthiopteris germanica Willd. Enum. 1071. 1809.
Matteuccia Struthiopteris (L.) Todaro in Syn. Pl. Acot. Vasc.
Sicilia, p. 30. 1866.
Matteuccia nodulosa (Michx.) Fernald in Rhodora 17: 164.
1915.
Rootstocks stout and ascending, with slender underground
stolons; sterile blades 2-7 feet high, 6-15 inches broad, short-
stalked, broadly oblanceolate, abruptly short-acuminate, gradual-
ly narrowed below middle, lower pinnae reduced; pinnae narrow,
deeply pinnatifid; segments oblong, obtuse, entire; veins simple,
fertile blades 1-7.5 feet high, with pod-like brown pinnae, in-
cluded sori confluent. July.
Distribution: Newfoundland to New York, westward to Michigan and Missouri;
recorded from Virginia. Alluvial soil.
Specimens examined: Livonia, Putnam Co., Bush 7780, 7780A, 7780B.
PTERIDIUM
1. l hel ted, seldom redivided. . .2. P. latiusculum var. Paa
1. Pinnules oblong-lanceolate, at least a few vinnatifid ida d NN RE . P. latiusculum
1. Pteridium latiusculum (Desv.) Hieron. Wissenschaftl.
Ergeb. d. Schwed. Rhodesia-Kongo-Exp. 1911-12, Heft 1, p. 7.
1914; Maxon in Am. Fern Jour. 9: 43. 1919. Gray, p. 36;
Britton & Brown, ed. 1, 1: 28, ed. 2, 1: 32; Eaton, pl. 35.
Pteris aquilina L. Sp. Pl. 2: 1075. 1753.
Pteridium aquilinum Kuhn in Decken, Reisen in Ost-Afrika
33:11. 1879.
Rootstocks cord-like, creeping, blackish, deeply buried; stipes
solitary, erect, naked, 1-3 feet high, swollen at base, brownish;
blades 2-4 feet long, 1-3 feet broad, triangular-ovate, rigidly
subcoriaceous, subternate; the long-stalked basal pinnae and
middle ones bipinnate, those above lobed or simple; segments
oblong-lanceolate, under surfaces glabrous or pubescent; veins
close-packed, free; indusia continuous around the edge of the
pinnules, double. July to September.
Distribution: Cosmopolitan, open woods, preferring sandy soil.
Specimens examined: Kellogg, Eggert, Palmer, Bush.
2. Pteridium latiusculum var. pseudocaudatum Maxon in Am.
Fern Jour. 9: 44. 1919. Gray, p. 36; Britton & Brown, ed. 2,
1: 32.
[Vor. 20
66 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The variety occurs in this region but does not seem to have a
distribution different from that of the species. It is distinguished
by its very narrow elongated pinnules.
THELYPTERIS
Dryopteris Adans.
Polystichum Roth
Aspidium Swartz
Nephrodium Richard
Lastrea Bory
Phegopteris (Presl) Fée
L INQUE DNE s 5: 65:5.65 6050000 Sra eR OSA EX SEA 5. T. heragonoptera
b 30 MEE LS EE 05055 0 ek EEEEREVEYREAAVCWXTEPS ECEERSERERESDIRARI 2
2. Blades bipinnatifid or bipinnate; segments not spinulose.................... 3
2. Blades tripinnatifid or tripinnate; segments spinulose....................... 4
3. Sori medial, small; blades membranaceous, not evergreen.................
COSERKRLEREXENAATCRERARWARDEREKENRE SREZSATLATAA 1. T. palustris var. pubescens
3. Sori near margins, large; blades subcoriaceous, evergreen...... 2. T. marginalis
4. Indusia glandless; pinnae decidedly oblique to rachises, scales of stipe pale
UNI Hoe d Eee eee Tey: avesECNICO EI I 8. T. spinulosa
4. Indusia glandular; pinnae more or less at right angles to rachises; scales of
stipe brown with a dark center.............. 4. T. spinulosa var. intermedia
1. Thelypteris palustris var. pubescens’ (Lawson) Fernald in
Rhodora 31: 34. 1929. Gray, p. 41; Britton & Brown, ed. 1,
1: 15, ed. 2, 1: 18; Eaton, pl. 30. Marsh shield-fern.
Thelypteris Thelypteris Nieuwl. in Am. Midl. Nat. 1: 226.
1910
Rootstocks slender, creeping, blackish, nearly naked; stipes
as long or longer than the blades, blackish at base, sparingly
chaffy; blades oblong-lanceolate, pinnate, 1-3 feet long, 4-6
inches wide, membranaceous, scarcely narrowed at base, short-
acuminate; pinnae 20-30 pairs, alternate, short-stalked, approxi-
mately at right angles to stalks, linear-lanceolate, broadest at
base, deeply pinnatifid; segments oblong-obtuse, mostly entire;
veins pinnate, usually once-forked; fertile fronds usually on
longer stalks and of narrower segments than the former; sori
nearly medial, crowded; indusia glabrous. August.
* Eaton mentions an unusual form—‘the lower two or three pairs are usually
but little shorter than those above them; but fronds are occasionally found in which
they are conspicuously reduced." Such & form is one from Iron Lake, Iron Co.,
Kellogg 1634.
1933] :
PINKERTON—FERNS OF MISSOURI 07
Distribution: New Brunswick to Florida, westward to Texas. Wet woods.
Specimens examined: Eggert, Kellogg, Bush, Pinkerton.
2. Thelypteris marginalis (L.) Nieuwl. in Am. Midl. Nat. 1:
226. 1910. Gray, p. 42; Britton & Brown, ed. 1, 1: 17, ed. 2,
1: 20; Eaton, pl. 55. Evergreen wood fern.
Rootstocks stout, ascending, covered with long chaffy brown
scales; stipes several inches to a foot long, light tan, somewhat
chaffy; blades 6-30 inches long, evergreen, subcoriaceous, ovate-
lanceolate, scarcely narrowed at base, deeply bipinnatifid; pinnae
numerous, practically sessile, lanceolate, acuminate, slightly
broader above the base; pinnules adnate to narrowly winged sec-
ondary rachis, oblong to oblong-lanceolate, faintly crenately-
toothed; veins free, forked or pinnately branched; sori large, near
margins of segments; indusia hard, orbicular-reniform, glabrous,
dark brown. July to August.
Distribution: Nova Scotia to Georgia, westward to Kansas and Oklahoma. Sand-
Stone ledges where it is moderately moist.
Specimens examined: Letterman, Engelmann, Eggert, Kellogg, Palmer, Pinkerton,
Muller, Rickett, Mackenzie, Broadhead, Link, Blankenship, Trelease.
3. Thelypteris spinulosa (Retz.) Nieuwl. in Am. Midl. Nat. 1:
226. 1910. Gray, p. 43; Britton & Brown, ed. 1, 1: 18, ed. 2,
1: 21; Eaton, pl. 68. Spinulose shield-fern.
Rootstocks stout, creeping, chaffy; stipes 4-14 inches long,
chaffy ; blades 0.5-3 feet long, ovate-lanceolate to oblong, acumi-
nate, bi-tripinnate, firmly membranaceous; primary pinnae short-
stalked, lower pairs triangular-ovate to triangular-lanceolate, re-
maining pinnae gradually narrower in outline; secondary rachises
narrowly wing-margined; pinnules oblong, subacute, incised with
spinulose-serrate lobes; sori small, sub-marginal, terminal on vein-
lets; indusia flat, round-reniform, glandless.
Distribution: Labrador to Virginia, westward to Idaho. Moist woods, alluvial
soil.
Specimens examined: Neeleyville, Butler Co., coll. of Oct. 30, 1899, Russell.
4. Thelypteris spinulosa var. intermedia (Retz.) Nieuwl. in
Am. Midl. Nat. 2: 278. 1912. Gray, p. 43; Britton & Brown,
ed. 2, 1: 22.
Similar to the species except for the glandular indusia and
right-angled relation of pinnae to rachis.
[Vor. 20
68 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: In Missouri—more northern, sandy so
Specimens examined: Pickle Springs, Ste. ced Co., Pinkerton 1; and coll.
of May 21, 1930, Kellogg.
5. Thelypteris hexagonoptera (Michx.) Weatherby in Rhodora
21: 179. 1919. Gray, p. 35; Britton & Brown, ed. 1, 1: 19,
ed. 2, 1: 23; Eaton, pl. 65. Broad beech-fern.
Phegopteris hexagonoptera (Michx.) Fée, Gen. Fil. 243.
Rootstocks elongated, slender, creeping, chaffy with gold scales;
stipes 8-20 inches long, sender: greenish, or straw-colored;
blades triangular, 7-12 inches long, 7-15 inches wide, thinly
herbaceous, deep green, slightly hairy or granular, bipinnatifid;
pinnae adnate to winged rachis, the lowermost ones broadest
and largely ovate to ovate-lanceolate, and others lanceolate; seg-
ments usually bluntly acuminate and crenate; veins pinnate and
free, branched or not; sori borne over whole frond, small, near
end of veins and so near margins; no indusia. August.
a New Brunswick to Delaware, westward to Oklahoma. Moist woods
and rav
"iens examined: Davis, Eggert, Palmer, Kellogg, Bush, Pinkerton, Rickett,
Link.
WOODSIA
1. Woodsia obtusa (Spreng.) Torr. Cat. Pl. in Geol. Rept. N.
Y. 195. 1840. Gray, p. 44; Britton & Brown, ed. 1, 1: 11, ed.
2, 1: 14; Eaton, pl. 71.
Rootstocks short, creeping, chaffy with narrow light brown
scales; stipes 2-6 inches long, green with darkish base in living
plants and drying to a bright brownish straw color; fronds 8-15
inches long, broadly lanceolate, narrower at base than in middle,
abruptly terminating at tip, membranaceo-herbaceous, minutely
glandular, deep blue-green in color, nearly bipinnate; pinnae
remote, short-stalked, obtuse, triangular, ovate to oblong, mostly
opposite; segments oblong, obtuse, crenately toothed; sori sub-
terminal on veins, nearer margin than midveins; young indusia
subglobose, splitting into several irregular lobes which extend
out beyond the sporangia, difficult to detect. September.
Distribution: Vermont to Alabama, westward to Wisconsin and Texas; recorded
from Nova Scotia. Moist calcareous or acid soil.
Specimens examined: Davis, Kellogg, Palmer, Pinkerton.
1933]
PINKERTON—FERNS OF MISSOURI 69
WOODWARDIA
1. Woodwardia areolata (L.) Moore, Ind. Fil. Gen. 45. 1857.
Gray, p. 38; Britton & Brown, ed. 1, 1: 20, ed. 2, 1: 25; Eaton,
pl. 22, fig. 2.
Lorinseria areolata (L.) Presl in Epim. Bot. 72. 1849.
Acrostichum areolatum L. Sp. Pl. 2: 1069. 1753.
Woodwardia angustifolia J. E. Smith in Mem. Acad. Turin
Se Ap
Rootstocks creeping, several to 12 inches long, often branched,
less than 1/4 inch thick, with some scales near apex; stipes dark
at base, paler above, bearing a few scales, dimorphic; sterile
fronds 9-10 inches long, oblong-ovate, pinnate, bright green
above, paler below; rachis winged from tip to just below base of
blades; sinuses rounded, segments acute, 1—4 inches long, 0.5-1
inch wide, finely serrate, membranaceous; veins finely reticulated,
with a longitudinal row of narrow areoles along each side of midribs
and midveins, and several rows of hexagonal aeroles and free
veins running outwards to serrated edges; fertile fronds taller,
with a darker stalk; segments and wing of rachis much narrowed;
one row of areoles on each side of midribs, each covered by a
brown involucre attached to the outer enclosing veins and open
along midrib; sporangia also from enclosing veinlets; sterile
fronds appearing in May and fertile ones later. August to
October.
Distribution: Massachusetts to Florida, westward to Missouri and Texas; re-
corded from Maine. Swamps and moist soil.
Specimens examined: Poplar Bluff, Butler Co., July, 1898, Eby.
EQUISETACEAE
EQUISETUM
1. Stems annual; dir&orphie......... Vet sees 1. E. arvense
1. Stems perennial; monomorphic. o. -graer hae eesi e nn heh 2
2. Sheaths cylindrical, green, turning gray, with black lines at bases and tops,
vue and undilated, splitting with age; ridges almost smooth; often very
plant... roD Oe rr rn 2. E. praealtum
2. Sheathe ae eae, green with narrow black limbs, elongate, not splitting;
ridges with one row of tubercules; medium-sized plants....... . E. laevigatum
1. Equisetum arvense L. Sp. Pl. 2: 1061. 1753. Gray, p. 52;
Britton & Brown, ed. 1, 1: 36, ed. 2, 1:39.
[Vor. 20
70 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Rootstock tuberiferous, felted with brown wool, extensively
creeping; fronds dimorphic; sterile ones annual, prostrate or
erect, green, rather slender, 12-24 inches high, 6-19 furrowed,
with scattered stomata; sheaths whitish, tipped with about 12
acuminate, brown, separate teeth; branches whorled, simple or
compound, not drooping, the 3-4 angled sheaths of branches
consisting usually of 4 teeth, or often 3, rarely 5, long and acumin-
ate; fertile fronds annual, appearing in early spring before sterile
ones, usually unbranched, succulent, and withering after spores
are ripe, 4-10 inches high, light brown, sheaths conspicuous,
long, flaring and pointed, of 8-12 teeth; spikes not apiculate;
variable. May.
Distribution: Greenland to Alabama, westward to Alaska and California. Sandy
soil, in waste places, along streams, etc.
Specimens examined: ^ Kellogg, Bush, Daniels, Palmer, Letterman, Eggert,
Trelease, Daniels, Mackenzie
2. Equisetum praealtum’ Raf. Fl. Ludovic. 13. 1817. Gray,
p. 53; Britton & Brown, ed. 1, 1: 38, ed. 2, 1: 41.
Equisetum hyemale var. robustum A. A. Eaton in Fern Bull.
11: 74. i
Hippochaete prealta (Raf.) Farwell in Mem. N. Y. Bot.
Gard. 6: 467. 1916.
Equisetum robustum A. Br., Engelm. in Amer. Jour. Sci. 46
88. 1844
Fronds perennial, evergreen, 3-11 feet high, erect; stems rough,
20-48 ridges bearing silica in single rows; sheaths cylindrical,
short-appressed, not dilated or only slightly when young,* at
first green but soon turning black or gray with black bands above
and below, splitting with age; sheath segments normally tri-
carinate; teeth dark and caducous; cones pointed. May.
T Schaffner (Am. Fern Jour. 13: 33-41. 1923), says: "Although P E.
ipsas usually bears cones on shoots of the season. Shoots ster firs
year may bear cones the second, both terminal and on lateral branches. Branching
is rare the first season unless the shoot is injured, but the second year branching i is
common even on uninjured shoots. E. praealtum is an exceedingly le species,
some forms recognized probably being genetie and some ecological, but none of
these forms passes out of the specific limits as usually drawn. Some are short and
robust; some tall and massive; some very slender."
* Young shoots are often very difficult to distinguish from E. laevigatum.
1933]
PINKERTON—FERNS OF MISSOURI 71
Distribution: Quebec to Georgia, westward to British Columbia and New Mexico.
Wet sandy places.
Specimens examined: Davis, Engelmann, Bush, Palmer, Pinkerton, Eggert, Trelease,
Daniels, Throuse, Demetrio.
3. Equisetum laevigatum A. Br., Engelm. in Amer. Jour. Sci.
46:87. 1844. Gray, p. 53; Britton & Brown, ed. 1, 1: 38, ed.
ATIS
Fronds perennial, erect, mostly simple, pale green, 1-5 feet
high, 14-30 ridged, almost smooth, with stomata in two rows on
each side of depressions; sheaths funnel-shaped, elongated, and
green with usually a narrow black band at thetop; white-margined
teeth soon deciduous; cones pointed. May to June.
Distribution: New York to North Carolina, westward to Washington and Cali-
fornia. Along streams, especially in san il.
Specimens examined: Engelmann, Bush, Daniels, Eggert, Palmer.
OSMUNDACEAE
OSMUNDA
1. Sterile fronds truly bipinnate; pinnules stalked and bend separated.......
P EA Nen Sah 4a SER FP PUR be n ASS . O. regalis var. spectabilis
i, Sterile fronds bipintattd. ....2. «4.56 don nena ou Eee BU EN 2
2. Fronds dimorphic; sterile pinnae with tufts of m hairs at base; apices of
fronds and pinnae tapering; veins inconspicuous............ . O. cinnamomea
2. Fronds monomorphic but fertile part of fioul ia is die middle several pairs of
pinnae; no tufts of hair at base of pinnae; apices of fronds and pinnae
abruptly narrowed and scarcely acute; veins dark-colored, conspicuous.
dala bac E EG e riu d ME M Ae A E 3. O. Claytoniana
1. Osmunda regalis var. spectabilis Fernald in Rhodora 32:
72. 1930. Gray, p. 46; Britton & Brown, ed. 1, 1:5, ed. 2, 1: 7.
Rootstocks stout, creeping, covered with persistent leaf-bases;
fronds 2-6 feet high, stipes never chaffy, green or yellow, rounded
on back, flattened on front; blades ovate-oblong, bipinnate;
pinnae iios opposite, ovate; pinnules unequal, 6-12 pairs plus
the terminal one, ES Corno short-petioled, distant, oval-
oblong or oblong-lanceolate, vidoe ones often auriculate on
lower side, margins crenulate-serrate, apex obtuse or subacute;
apical pinnae fertile, bipinnate, ultimate divisions thread-like,
containing no chlorophyll, entirely covered with sporangia. May
to July.
[Vor. 20
72 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: Newfoundland to Florida, westward to Saskatchewan and Missis-
sippi. Lowlands, swamps, marshes, and wet woods.
Specimens examined: Eggert, Palmer, Kellogg, Bush, Pinkerton, Engelmann,
Trelease, Mackenzie.
2. Osmunda cinnamomea L. Sp. Pl. 2: 1066. 1753. Gray, p.
47; Britton & Brown, ed. 1, 1: 5, ed. 2, 1: 7.
Rootstocks creeping, massive, bearing circular clusters of
sterile leaves with one or more fertile ones within; fronds dimor-
phic, sterile ones 1 foot or more high, oblong-lanceolate, acu-
minate, tapering, deeply pinnatifid; pinnae oblong-lanceolate,
acute, tapering, tomentose tuft at base of each pinna; pinnules
obtuse, subcoriaceous, green; veins inconspicuous, veinlets once-
forked near midvein; margins entire or obscurely crenulate;
fertile fronds about as tall as the sterile, bipinnate, and covered
with cinnamon-colored sporangia, arising early in the spring pre-
ceding the sterile ones. May an
Distribution: Newfoundland, westward to Minnesota and New Mexico. Sand-
Specimens examined: Eggert, T'release, Kellogg, Pinkerton, Russell.
3. Osmunda Claytoniana L. Sp. Pl. 2: 1066. 1758. Gray, p
46; Britton & Brown, ed. 1, 1: 6, ed. 2, 1: 8.
Rootstocks creeping, stout, with imbricated leaf-bases; stipes
several inches to 2 feet long, woolly when young but never chaffy ;
fertile fronds taller than the sterile and in the midst of the crown
formation, oblong-lanceolate, 1—4 feet long; lowest pinnae about
half as long as middle ones, acute and often rounded; pinnae
barely acute, never acuminate, short-stalked, lütioeclste from a
broad base; pinnules close; 2—6 pairs of fertile pinnae near middle
of frond, shorter than sterile pinnae and deflexed in a mature speci-
men, closely bipinnate, woolly, covered with bivalvular reticulated
sporangia. May to July.
mto Adr Newfoundland to North Carolina, westward to Minnesota and
ri.
mps and moist woods, moist sandstone ledges.
el thence Bush, Eggert, Davis, Palmer.
LYCOPODIACEAE
LYCOPODIUM
1. Sporophylls segregated into slender cones; habit of plants fan-like.........
els Hise EGET ELS Ae heed ve hee oo ae 3. L. complanatum var. flabelliforme
1933]
PINKERTON—FERNS OF MISSOURI 73
1. Sporophylls not differing from vegetative leaves; habit of plants rope-like...... 2
2. All the leaves broadest above the middle; margins jagged...... 1. L. lucidulum
2. Shorter leaves broadest at base; margins entire or slightly denticulate......
OPE PUDRS M RODA c E 2. L. lucidulum var. porophilum
1. Lycopodium lucidulum Michx. Fl. Bor. Am. 2: 284. 1803.
Gray, p. 55; Britton & Brown, ed. 1, 1: 40, ed. 2, 1: 44.
Stems assurgent from decumbent persistent bases giving rise
to a few vertical stems; leaves dark green and shining, widespread
or becoming deflexed, acute, broadest above middle, erose-
denticulate, arranged in alternating series of long and short
members, the latter often entire and usually bearing the sporangia;
gemmiferous. August to October.
Distribution: Newfoundland to Delaware, westward to Alaska and Washington.
On sandstone only, usually associated with Sphagnum.
Specimens examined: Kellogg, Eggert, Pinkerton.
2. Lycopodium lucidulum var. porophilum (Lloyd & Underw.)
Clute, Fern Allies, p. 3. 1905. Gray, p. 55; Britton & Brown,
ed. 2, 1: 44.
Lycopodium porophilum Lloyd & Underw. in Bull. Torr. Bot.
Club 27:150. 1900.
Essentially like L. lucidulum except that the shorter leaves are
broadest at the base and the margins are nearly smooth, and are
not deflexed.
Distribution: in the same places as the species. Sandstone
Specimens examined: Terre Bleue Cr., Ste. Genevieve Co., coll. of Aug. 29, 1898,
Trelease, and Pinkerton 81.
3. Lycopodium complanatum var. flabelliforme Fernald in
Rhodora 3: 280. 1901. Gray, p. 57; Britton & Brown, ed. 1,
1: 43, ed. 2, 1: 48.
Rhizomes slender, creeping, with numerous erect stems which
branch irregularly, giving rise to a flattened fan-shaped vege-
tative structure, about a foot high with 4-ranked imbricated
scale-leaves, those of the two lateral rows broad, with spreading
tips, of the upper row narrow and incurved, and of the lower
row minute deltoid-cuspidate; peduncles long, dichotomously
branched at tips and bearing a number of slender cones about
an inch long. August and September.
[Vor. 20
74 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: — to West Virginia, westward to Alaska and Idaho. Open
pine woods on sandy so
Specimens examined: "Pickle Springs, Ste. Genevieve Co., Kellogg 8718.
SELAGINELLACEAE
SELAGINELLA
1. Plants bearing ill-defined strobili; leaves dimorphic, 4-ranked....... 1. S. apoda
1. Plants bearing distinct strobili; leaves of one kind, spirally arranged.2. S. rupestris
l. Selaginella apoda (L.) Fernald in Rhodora 17: 68. 1915.
Gray, p. 58; Britton & Brown, ed. 1, 1: 45, ed. 2, 1: 49.
Selaginella apus (L.) Spring in Mart. Fl. Bras. 12:119. 1840.
Stems prostrate and creeping, 1-4 inches long, pale green,
delicate; leaves of two kinds, four-ranked and spreading, the
smaller pointed and appressed to the stem; no distinct cones;
fertile leaves near tip of branches, those containing macrospores
conspicuously bulged. July to September.
Distribution: Massachusetts to Florida, westward to Michigan and Louisiana.
Moist shaded places, among grasses.
Specimens examined: Eggert, Bush, Palmer, Mackenzie.
2. Selaginella rupestris (L.) Spring in Mart. Fl. Bras. 12: 118.
1840. Gray, p. 57; Britton & Brown, ed. 1, 1: 44, ed. 2, 1: 49.
Stems densely tufted, bearing occasional sterile runners; all
leaves alike, narrow, appressed, and imbricated, bristle-tipped,
gray-green; strobili four-sided. August to October.
Distribution: Quebec to Alabama, westward to Minnesota and Oklahoma. In
dry exposed places where there is a little soil, sandstone, or chert.
Specimens examined: Eggert, Russell, Bush, Palmer, Pinkerton, Greene, Broadhead,
VanIngen, Shepard.
ISOETACEAE
ISOETES
1. Megaspores reticulate; sporangia unmarked................. 1. I. Engelmanni
. Megaspores tuberculate; sporangia marked in some w 2
. Megaspores less than 480 in diameter; sporangia ae BE with brown spots.
Fleck 92 UR RD PAR RUE RR ERE RUE OR ala aaa Galeua 4s Gee EORR 2. I. melanopoda
2. Megaspores more than 480 y in diameter; sporangia marked with brown lines.
ee E re a ere er re ES 8. I. Butleri
t2 m=
1. Isoetes Engelmanni A. Br. in Flora 29: 178. 1846. Gray,
p. 61; Britton & Brown, ed. 1, 1: 48, ed. 2, 1: 53.
Corms 2-lobed; leaves 15-60, 13-50 cm. long, light green; sto-
1933]
PINKERTON—FERNS OF MISSOURI 75
mata numerous; peripheral strands variable in number or none;
sporangia oblong, unspotted, with narrow velum; megaspores
white, 400-570 y in diameter, distinctly marked with honey-
comb network of narrow ridges; microspores 21-30 y, seldom
33 y, in length, smooth to minutely roughened.
Distribution: eastern border to rom valley. Near ponds.
Specimens examined: Engelma
2. Isoetes melanopoda Gay and Dur. in Bull. Soc. Bot. Fr.
11:102. 1864. Gray, p. 61; Britton & Brown, ed. 1, 1: 48, ed.
21:54.
Corms 2-lobed; leaves 15-60, 15—40 cm. long, slender, erect,
firm, bright green, usually black and shining at base, with Mat
pale membranaceous border, little (2-3 em.) extended above spo-
rangium level; stomata present; peripheral strands 4—6 cardinal,
plus as many as 14 accessory groups; ligule subulate, triangular;
sporangia oblong, 0.5-3 cm. long, marked by numerous brown
spots; velum variable, from very narrow to covering half of
sporangium; megaspores 280—440 y. in diameter, marked with low
tubercules, frequently confluent into short low wrinkles; micro-
spores frequently ashy-gray, 20-30 u long, fine-spinulose.
Distribution: Illinois to Texas. Wet prairies.
Specimens examined: Pfeiffer, Bush, Palmer.
3. Isoetes Butleri Engelm. in Bot. Gaz. 3: 1. 1878. Gray,
p. 61; Britton & Brown, ed. 1, 1: 48, ed. 2, 1: 54.
Corms 2-lobed; leaves 8-30, 8-15 cm. long, more slender and
rigid than J. melanopoda, tapering to apex; stomata numerous;
peripheral strands usually 4, sometimes more in number; ligule
elongated, cordate at base; sporangium oblong, 6-7 mm. long,
marked with brown lines; velum very narrow; megaspores vari-
able, commonly 480-650 u in diameter, marked with numerous
tubercules, usually distinct, occasionally confluent; microspores
27-37 y long, papillose.
Distribution: Tennessee, westward to Kansas and Oklahoma. Limestone barrens.
Specimens examined: Eggert, Bush, Palmer.
SALVINIACEAE
AZOLLA
1. Azolla caroliniana Willd. Sp. Pl. 5: 541. 1810. Gray, p
50; Britton & Brown, ed. 1, 1: 35, ed. 2, 1: 38.
[Vor. 20
76 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Plants floating on surface of water, often covering large areas,
deltoid or triangular-ovate in outline, 6—25 mm. broad, pinnately
branched; lobes ovate, lower lobe reddish, upper greenish with a
reddish border; megaspores minutely granulate with three acces-
sory corpuscles; masses of microspores armed with rigid septate
processes.
Distribution: Lake Ontario to Florida, westward to Washington and California.
On surface of still waters.
Specimens examined: Eggert, Engelmann, T'release, Bush, Mackenzie.
INDEX TO SPECIES
Acrostichum areolatum............
Adiantum
CCC eC cece ere se eeenernesees
? Names in italics are synonyms.
asplenioides................... 55
—— ERETT x4 doit 55
pycnocarpom... cce 54
irterethi DET OTT ER Fat 55
D — Pc 75
earolinianga...............Lses 75
Botrycbium.............. sees 47
OÉSasclum. 6 5.6 60 65.5 E ark a 48
OU 5 israr a TII 47
var. dissectum.............. 48
var. tenuifolium............. 48
ternatum... 6... ccc ee eee ee 47
var. dissectum............04. 48
var. a cease eevee TEM 47
bp colpo oeer ike sin anaes 48
var. peas a A REE 49
CDU. reda icine 57
rhizophyllus.................. 57
Cheilanthes..............2...... 57
alabamensis................... 57
dealbata.. 2.0.0... ce cee eee 61
b i v5 45-3 oe eee 58
IF c RE 58
lanuginosa... 0.0.06... 58
Vesttia. wb we cede nh 58
es o e s PEETTTTOURT TTC TET. 58
bulbifera. oo... 0.000.000.0000... 59
var. horizontalis............. 60
His... ETA eee eee 58
Dennstaedtia................... 60
punctilobula.................. 60
EURO 5k iiss eR REFER 60
gilostuscula ........... esee. 60
1933
PINKERTON—FERNS OF MISSOURI 77
punciilobula. ...... e cae eee 60. PD.» teen ERES 61
y penc cor mots or 66 AUODUPDUINA.,.... Feed cane oo 62
ocrosticoMel . Vers 6 val. DUME LS io CONSU 62
acq DA Re eae tess Mews 6 Var. Gh.) oe ie) 62
EWR Lolo aS Fascists codes 69 2s os taeda ane E 61
voies FOE o e eek 70 gabela: sos cv Xs oe Phan 62
BV EUER. eerte I PROG. ica CR eR Ee 66
WRI PAPERS RO EE TETTE EE T 70 hezagonoptera. ooo ss Bc esee 68
Mg, RTT CET TT eee E E E 70 einem DEDERIS aia 62
XR UMEN EAE LL rex 59 TM cec
ee a ae eer anne 59 eae TOWN IU eee €3
BEL slicers tas 2 rb 59 hay Tp umet 63
Hippochacie POG ooe ra aea 70 ee Ae Sey ee 63
Tr c crc 74 Posen, 204 BORNE 63
push BEocerua ipu s M e E RE 75 acrostichoides. . ....1.... 2... 63
Ingelmannic cs ose 74 var. méisum. anru Bik 6.0035 64
verei NEC 75 var. Schweinitsli. ods oos 64
SEEN eso vies SAR E 66: . Polystichum.. c. en E 66
anaes aa: Peer Perey Terre 600 Preeti ccc Ss 64
DOPU ea ean a a 72 ici e VETUS E T. 64
bce flabelliforme. ..... 78 Piendium..... o CBE 65
lucidulum..................... 73 aquilinum. .... ves nee nn 65
var. porophilum............. 73 latiusculum. 2:5 Es 65
porophilum.......... lessen. 73 var. pseudocaudatum......... 65
o MINER TERT TT TOT eee ree 65
NEN cocos odit eee eee 65 lire us PORE Ee "or 7
Birulhiopteris. ............ LL... 00. "dE Iesse rex 74
EDO. Leo eod nese R 66 uis TOM E Pe: ee ene 74
BODDOlBBUA. o erae rex 60 oc MEME VETERES 74
so dio lí MEME ERU NI, 60 Struthiopteris germanica........... 65
nivea dealbata. ........... Lulu. OMNEM 0o. s aer a ae 66
Nothochlaena pulchella............ 6l hexagonoptera TONER SUA 68
UBND ech ewes a be aes 61 a, TET T 67
os pl MEME NEMINEM dead 61 palustris pubescens............ 66
gg en Tasse ees 64 a Pee RA 67
CDI PRETI TEES et 46 var. ^ NU MES 67
PERDERE, (Coe d xe d Core 47 AA TAn a P Sees TE 66
FUSRLUIM. ou adien aaa cedit (AN T R E T rus 68
MUNN cios REESE T 71 | PME INE MT. n 68
MUDMBODN. ric o ON ada 4 xh r3. o AMEMMMERÉ 69
Olaytoniana o Lond Eo an we ees 72 TO ....... dero DS 69
regalis spectabilis. ............. 71 ae, mee” pe 69
Struthioptreris.................. 64
[Vor. 20, 1933]
78 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 5
Fig. 1. Cystopteris bulbifera var. horizontalis Lawson. From E. J. Palmer No.
32896, in the Herbarium of the Missouri Botanical Garden.
Fig. 2. Botrychium obliquum var. tenuifolium (Underw.) Gilbert. From Dr.
William — coll. of October 28, 1897, in the Herbarium of the Missouri Bo-
tanical Garden
INNOSSIN AO SNUAA—NOLUAMNId
Ann. Mo. Bor. GARD., Vor. 20, 1933
PLATE 5
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BLASTOMYCOSIS: REPORT OF A CASE, WITH
A STUDY OF AN ETIOLOGIC FACTOR AND
A CLASSIFICATION OF THE ORGANISM
MORRIS MOORE
Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany
of Washington University
INTRODUCTION
It is the purpose of this paper to describe briefly the disease
known clinically as blastomycosis, and to try to clarify the rec-
ognition of the organism involved. The literature of the field
is at present too extensive for an entire review, and since numer-
ous workers have already given excellent discussions on the
clinical aspects of the infection, as to its gross pathology, micro-
scopic histo-pathology or cellular reactions, and the biological
or rather immunological phenomena, a review would be un-
necessary here. However numerous such papers may be, there
is still much work to be done on the subject.
The author has attempted to clear up, at least in his own mind,
several undecided points in the disease: first, the establishment
of the etiological agent of blastomycosis; second, the determina-
tion of the exact classification of the organism. In the past,
and even at the present, medical men have grouped under one
general heading all organisms which were responsible for the
same clinical condition. Good as this system may be for general
diagnosis, much difficulty is encountered because of the fact that
physicians are inclined to devote very little time to a study of
the organism, thus rendering any therapeutic measures, if avail-
able, indefinite, inasmuch as several of the fungi present varying
degrees of pathogenicity and require different therapeutie meas-
ures.
Thus we find that numerous species of the genera Saccharo-
myces, Monilia, Cryptococcus, Endomyces, Sporotrichum, and
others have, at one time or another, been considered etiological
agents of blastomycosis. A review of the history will illustrate
these facts.
HisTORY
Years before the first case was definitely described as blastomy-
cosis, investigators had performed a certain amount of work on
ANN. Mo. Garp., Vor. 20, 1933 (79)
[Vor. 20
80 ANNALS OF THE MISSOURI BOTANICAL GARDEN
fungi involved in cases of infection and had established these
organisms as etiological factors, particularly the yeast and yeast-
like groups.
Chronologically, the list is quite long, but it is worthy of note.
According to Hufschmitt, Sartory, Sartory, and Meyer (731), we
find that in 1845 Remak, and in 1853 Robin, in his ‘Histoire
Naturelle des Végétaux Parasites de l'Homme et des Animaux,’
discovered the normal existence of the yeast Cryptococcus guttu-
latus in the rabbit intestine. A few years previously, Hannover
(cited in Buschke and Joseph, '28) had found yeast in the urine
of diabetie patients. Investigations then tended to turn to the
parasitism of these organisms in animals, with the result that
Bernard during the course of his work on fermentations attempted
the first animal experiment by injecting beer yeast into these
subjects. Following this work, Popoff, Grohe, Roussy, and
several others showed the pathogenic actions of the yeasts on
mammals, and Rivolta in 1873 in his ‘Parasiti Vegetali’ dem-
onstrated for the first time a yeast infection in a horse. In
the meantime, Metchnikoff and Weismann showed the parasitism
of the Saccharomycetaceae in the lower animals. In 1892
Wernicke showed the first mycosis in man and named it ** maladie
protozoique de la peau." The following year Troisier and
Achalme (793) definitely established the relation between yeasts
and man. In the meantime, several workers attempted to show
the destructive ability of these organisms on the animal tissues.
Popoff in his work had used dogs as his subjects, but impure
cultures. Raum ('91) inoculated animals with large amounts of
yeasts, and a rise in their temperature, shortness of breath, and
death resulted. Neumayer (91) fed animals with cultures and
also inoculated them subepidermally. His feeding developed a
gastro-enteritis which he believed due to fermentation, since the
skin inoculations were of no value. The yeasts of these workers
were probably of the non-pathogenic types, for L. Rabinowitsch
(96), a few years later, showed fifty various yeast-like organisms
with seven pathogenic for animals. Nesczadimenko (’99) made
peritoneal injections of yeasts in a physiological saline solution
into rats, mice, guinea-pigs, and dogs, with death ensuing from
eight to twelve days. He concluded, however, that these
organisms were not so deadly, although causing this mortality.
1933]
MOORE—BLASTOMYCOSIS 81
The first actual case of blastomycosis, so-called, was reported
by Gilchrist at the June, 1894, session of the American Dermato-
logical Association. His paper resulted from the finding of
peculiar yeast-like bodies in the diseased tissue of a patient. The
doctor attending the patient had given the diagnosis as a typical
case of chronic scrofuloderma. Several months after Gilchrist’s
report, Busse (’94) brought to light the extraordinary case to
which he later gave the name “‘Saccharomycosis hominis." The
patient was a woman thirty-one years of age who had suffered
from a localized subperiosteal inflammation of the left tibia. An
examination of the abscess, which opened spontaneously, re-
vealed ‘‘numerous doubly contoured, very refractive, roundish
and ovoid bodies,” and these were found to be situated both
intracellularly and extracellularly in the pus and abscess wall.
These organisms when isolated in pure culture and then inocu-
lated experimentally in animals proved to be what were later
known to be blastomycetes. The patient later developed super-
ficial ulcers on the face, subperiosteal swellings on the right ulna
and the left sixth rib near the axillary line, with death ensuing.
Busse cultured the yeast from the ulnar swellings and from the
bottom of the ulcers.
Approximately two years later, the first case reported by
Gilchrist was published in detail in the Johns Hopkins Hospital
Reports of 1896. In the meantime, however, several others had
noticed similar cases among guinea-pigs, horses, mice, and other
lower animals (Sanfelice, 795, 796, 96a, Roncali, '95, Corselli
and Frisco, '95, Tokishige, '96, and others).
In 1896, Curtis isolated a fungus similar to that described by
the former writers from a myxomatous tumor of theleg. In the
same year, Gilchrist, in conjunction with Stokes, published a
short paper on a second case of blastomycosis, and this was
published in detail two years later (Gilchrist and Stokes, ’98).
Simoni (’97), working on the diseased tonsils of patients, found
budding yeast-like cells in twenty tonsils. Maffuci and Sirleo
(98) examined numerous tumors and found budding cells in a
great number of tissues. Many other reports followed, as that
of Hyde, Hektoen, and Bevan (’99) with a supplement by Hek-
toen (’99) later in the year, Hessler (’99) with a case report, and
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82 ANNALS OF THE MISSOURI BOTANICAL GARDEN
several during the same year and 1900. In the following year
appeared the elaborate work of Ricketts (01), with a study of
the organism from a case of systemic blastomycosis by Otis and
Evans (03). Eisendrath and Ormsby (705) described a systemic
infection, and Irons and Graham (706) reported a severe gener-
alized systemic disorder. Hektoen (’07) gave a comprehensive
review of the literature, and from that time on the medical
journals have published too great a number of cases of infection
due to yeast-like organisms, under the heading of blastomycosis,
to render a complete survey of literature a matter for a paper of
this length.
ETIOLOGY AND CLINICAL MANIFESTATIONS
The disease known clinically as blastomycosis is very likely
due to a plurality of organisms and not species of the same genus
as indicated by previous writers. This is clearly evident as seen
by the great number of papers published and the cases reported,
involving such fungi as Saccharomyces, Oidium, Monilia, En-
domyces, Cryptococcus, Coccidioides, and even such a form as
Sporotrichum. The clinicians have referred to the category of
blastomycosis any clinico-pathological condition which may be
due to yeast-like or budding fungi. It must be understood,
therefore, that the term as here used refers only to the clinical
aspect of the condition.
In America, clinicians and medical men, more especially
medical mycologists, are inclined to class as the cause of blas-
tomycosis only that organism which was described originally by
Gilehrist and Stokes in 1894 and in this view the author is greatly
in accord. On the other hand, European workers consider only
that organism which was reported by Busse (94) and so elaborated
on by Buschke ('98). However, by reason of priority, the organism
of the former workers should hold the position so designated and
be established as such. Further remarks on the Gilchrist organ-
ism will be found in the discussion.
Blastomycosis presents numerous clinical manifestations and
in this respect it is protean, being found in practically every organ
of the human body either in biopsy or autopsy material. No
immunity towards the invading organism is established by any
1933]
MOORE —BLASTOMYCOSIS 83
of the anatomical structures. Clinically, the condition presents
lesions which are alike both for the cutaneous type of the disease,
that is that group of infections which may be found occurring
superficially, or for the systemic type of the disorder, occurring
in the lungs, bones, meninges, liver, or other viscera. This
division is based on the work of Jacobson and his associates
(32), who further separate the cutaneous type into that of
primary in character, as occurring in the epidermic layers or the
cutis, as shown by Hagiwara (22) and Hashimoto (722), whose
organisms, although not of the Gilchrist type yet coming under
the general heading of blastomycosis as known clinically, also
the work of Grschebin (727) ; and secondary, due to an infection of
the deeper tissues, internal viscera, or bony structures, as shown
by Irons and Graham (706), Ryerson (709) for bones, and many
others.
The primary form or the cutis infection Jacobson further
designates as presenting one of three varied appearances: papulo-
ulcerative; verrucous or papillomatous; gummatous.
The papulo-ulcerative type Jacobson designates as being initial
lesions which are papulo-pustular in character and of epidermic
origin (shown by Hessler, '99, Hektoen, ’99, Ricketts, '01,
Engelhardt, '24, and Fabry, ’27, '28). These lesions rupture in
the course of time and empty out the purulent exudate on the
surface of the skin, with the probable ultimate formation of
crusts. The process may be proliferative and involve a great
area of the immediate vicinity. The lesions usually show a
violaceous border with the involution of the peripheral surfaces
and perhaps consequent scarring and atrophy.
The verrucous or papillomatous type (Froilano De Mello and
Rodrigues, '29) is characterized as being nodular or papular in
character and present on a normal or deep-red, infiltrated skin.
Several of the lesions may coalesce to form papillomatous patches
which resemble verrucous tuberculosis. These lesions may break
up into healing areas which upon drying present irregular scars.
The characteristic color as noted above is found here too, as well
as the sloping periphery.
The gummatous type develops from the subcutaneous layers of
the tissue of the deeper portion of the cutis in the form of small,
[Vor. 20
84 ANNALS OF THE MISSOURI BOTANICAL GARDEN
slightly elevated, somewhat tender, reddish, deep-seated, soft
nodules situated on the characteristic violaceous-red surface of
the skin. There is a diffusion of the color with subsequent
establishment of new nodules in the vicinity. The nodules en-
large, become soft and gummatous, and then break down to form
masses of ulcerative, proliferative materials bordered as in the
other two types, and contain numerous abscesses.
The secondary cutaneous form consists chiefly of variously
formed ulcers which give off a purulent or sanguino-purulent
discharge from a soft, granulating floor. Some may develop
crusts with raised edges, while some may assume hyperplastic
functions with papillomatous characteristics, and usually there
may be a metastatic action on the part of the ulcers represented
by the formation of new lesions which are surrounded by a dark
red or purplish zone. Healing may be spontaneous with indurated
scars, as noted in some cases, or infection may persist but may
finally succumb to treatment with iodides as was noted in the case
reported here.
A study of the ulcers formed in blastomycosis shows them to
originate in abscesses which from a clinical point of view can be
divided into the superficial and the deep types. Secondary,
cutaneous, superficial ulcers arise usually in the subcutaneous
tissues as nodules of varying size as shown by Stober (14),
Engelhardt (24), Ferguson (’28), and Montgomery and Ormsby
(08). These ulcers usually enlarge, rupture, and spread the
material over the surface of the skin, setting up new foci, or in
some cases they have been found to dry up and disappear.
The deep type of secondary cutaneous blastomycosis (Grsche-
bin, '27, ’28), characterized by smaller number and deep-
seatedness, is by far the most serious of the two, involving de-
structive processes of the bone, muscle, and deep tissues and
organs. It rarely shows any inflammatory reaction, but can be
distinguished by the purulent character of the abscesses as con-
trasted with the mucoid or mucopurulent nature of the super-
ficial abscesses.
The above types represent the typical forms occurring in a
clinic. However, Weidman and Douglas (’21) reported the
occurrence of a sarcoma-like tumor on the leg of a patient, which
1933]
MOORE—BLASTOMYCOSIS 85
looked like lupus vulgaris and yielded blastomycetoid bodies on
histological sectioning. Then, about six years later, Cleland
(27) reported a case with the formation of a myxomatous-looking
tumor mass which also showed typical cells on sectioning. These,
however, are rare and until more cases are reported cannot be
placed in the definite clinical types.
Under the heading of cutaneous blastomycosis, Castellani has
also established principal types of blastomycosis of the cutis from
a clinical point of view.
1. Blastomycosis verrucosa (Synonym: Blastomycosis, Gilchrist t s
Etiology: Cryptococcus dermatitidis Gilchrist and Stokes, 1896 (Synonym: Cryp-
tococcus gilchristi Vuillemin).
Castellani here creates the genus Blastomycoides.
2. Blastomycosis ulcerativa profunda sen mutilans (Synonym: Blastomycosis,
Wernicke-Ophüls type or Blastomycosis coccidioides type).
Etiology: Coccidioides immitis Rixford and Gilchrist, 1896.
3. Blastomycosis purulenta profunda (Synonym: Blastomycosis, Busse type;
Blastomycosis subcutanea purulenta).
Etiology: Cryptococcus hominis Vuillemin, 1901 (probably covers many species).
4. Blastomycosis glutealis (Synonym: Blastomycosis, Kartulis type
Etiology: mycological investigations iis dis completed. The fungi seem to
belong to the genera M onilia and Cryptoco
astomycetica Pil eee (Folliculitis decalvans cryptococcica,
pro parte, Castellani type). (Synonyms: Furunculosis cryptococcica; Pseudo-
furunculosis blastomycetica; Furunculosis mycetica; Folliculitis decalvans sacchar-
omycetica J shape decalvans moniliaca).
Etiology: yeast-like fungi either of genus Cryptococcus or Monilia (No asci or
ascospores ects to Castellani).
In addition to the above types, Castellani adds the following,
although they are in no way connected with the blastomycosis
organism: Blastomycosis epidermica; Intertrigo blastomycetica;
Dermatitis blastomycetica interdigitalis.
For clinical purposes in diagnosing skin infections such a
classification is good, but for correct determination of the etio-
logical agent, it is essential that each type of involvement be
named with the infective agent designated as such. For example,
if the organism be a saccharomycete, the disease should be called
saccharomycosis; if a monilia, then moniliomycosis; if an endomy-
cete, then endomycosis. Thus, when the organism is correctly
Fr the amount of time necessary to determine the right sort
of curative measurefor that type of infection may be taken, and the
amount of time necessary for healing reduced.
ó [Vor. 20
86 ANNALS OF THE MISSOURI BOTANICAL GARDEN
SYSTEMIC BLASTOMYCOSIS
As mentioned previously, the disease is protean in its clinical
manifestations, with the result that practically every organ in
the body has been shown to be infected either in the living, by
various measures, such as X-ray, or in autopsy material. No
vital organ is immune, and this in itself is sufficient to cause a
careful physician to give more attention to therapeutic measures.
This universal infectivity of the agent was especially shown by
such writers as Otis and Evans ('03), Eisendrath and Ormsby
(05), Le Count and Myers (’05), Irons and Graham (06),
Montgomery and Ormsby ('08), Wade and Bell (16), Garr (725),
Panja (25), Toepel (29), and Maner and Hammack (30).
There is, however, a difference in frequency with which the vari-
ous organs show their susceptibility.
'The portion of the anatomy that shows the greatest amount of
infection is the skin, either primary or secondary, having about
95 per cent of all cases recorded. This phase of blastomycosis
has received the greatest amount of attention principally because
it is so prevalent, but also because it is usually a manifestation of
a metastasis from the deeper organs, and this helps bring forth the
diagnosis of blastomycosis.
The pulmonary system, including the lungs and bronchi,
constitutes the second most frequent and the most common sys-
temic form, being present to the extent of approximately 95 percent
in systemie infection in available autopsy records, and about 35 per
cent in primary cases. This was shown by such menas Stober (14),
Wade and Bell (16), Wade (18), Dennis (18), Miller (27),
Medlar (27), and Mazza and Niño (28). The disease probably
is primary in the bronchi and from there spreads to thelungs. If
secondary in the lungs, as in systemic disorders, the process may
be slow and chronic; if primary, however, the spread may be
rapid and fatal.
The kidneys are next in frequency of infection. The genito-
urinary involvement is usually secondary by way of metastatie
foci through the blood. The disease in these organs manifests
itself in the form of nephritis, showing casts and albumen in the
urine. The culturability of the organism from samples, however,
1933]
MOORE—BLASTOMYCOSIS 87
cannot be demonstrated unless the kidneys are accompanied by
infected bladder or prostates.
The spleen follows next in order, but this organ is usually
easily susceptible so that a great amount of infection is to be
expected in any systemic disorder of this sort.
The complication of the bones and joints seems next in the
amount of infection. This form of the syndrome is very common
in systemic disorders, and may even be a primary infection if
the diagnosis in the patient here described was correct seven
years previous to his entry at the Barnard Skin and Cancer
Hospital. This type of the disease may manifest itself in the
form of arthritis, osteitis, osteomyelitis, or periostitis, according
to Ryerson ('08-'09) and Stober (’14). The process usually
results in a suppuration, formation of sequestra, and abscesses
which break down intervening cell walls and coalesce, causing
great damage.
The liver appears to be a rather frequent subject to the in-
fection, coming next in the order of frequency. This is to be
expected in systemic disorders where the blood plays an impor-
tant part. Metastases through the blood vessels are fairly com-
mon, and yeast cells are easily transported to the main organs in
this manner.
The pleura too are susceptible to a great extent, and here the
proximity to the lungs is a great factor in their thfedtion.
The lymph glands follow along rather closely, as shown by
Wanamaker (728), especially for the cervical lymph glands.
Cerebrospinal involvement, including the brain, meninges,
spinal cord, and skull bones, occurs fairly often as a secondary
metastatic process in generalized systemic blastomycosis, ac-
cording to studies made (J. T. Moore, ’20, Freeman and Weid-
man, '23, Greenfield, '24, Wilhelmj, '25, and Gáspár, '29), being
found in at least 12 per cent of the cases. When the disease is
secondary to a general systemic infection, there may be osteo-
myelitis of the skull bones with destructive processes. The
diagnosis rests not upon any clinical entities which may be present,
because the inflammatory reaction simulates many other condi-
tions, but upon the actual laboratory finding of the organism
either in the spinal fluid or in sections of the diseased brain tissue.
[Vor. 20
88 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Wilhelmj (725) states that in those cases where there is no patho-
logical symptomatology or clinical manifestation on any other
part of the body, and when the meninges are infected during the
primary stages of the invasion, death may occur without the
initial appearance of general metastatic foci, and such a condition
he calls primary cerebrospinal blastomycosis. The spine may be
involved in the process in a suppurating condition (Parker, ’23),
but this condition is relatively rare.
Jacobson lists the vertebrae as being next to the brain in
susceptibility to attack. This condition has been noted on
several occasions. Roentgenographic studies usually reveal an
infection of bodies of the vertebrae, and lamina and posterior
ligaments may show an involvement which simulates greatly
tuberculous Pott’s disease.
Prostatic infection in blastomycosis is often noticed (Parmenter
and Simpson, '19). Usually it is associated with a genito-urinary
complication and involves the urinary bladder (Rhamy, ’26).
In these cases acute urinary urgency and pyuria are well-defined
symptoms.
The heart lesions in blastomycosis are shown first in the
pericardium and then in the myocardium in the form of an in-
flammatory reaction (Hurley, 716).
Pancreatic involvement follows in frequency.
Infection in the peritoneum is the next most common. Jacob-
son reports finding the disease in the abdominal viscera in the
following decreasing order of frequency: kidneys, spleen, liver,
lymph glands, pancreas, peritoneum, adrenals, and gastro-
intestinal tract. These organs, as pointed out previously, become
involved usually through metastasis by way of the blood stream
or by direct transmission from tissue to tissue. In this manner,
testicular blastomycosis is usually developed. Blanchard, Swartz
and Binot (’03), as early as 1908, noted an intraperitoneal in-
volvement.
The eyes may also be involved. The infection here is very
painful and often dangerous, leading to blindness with perhaps a
complication of the nervous system and the brain. McKee (26)
and Ferguson (’28) noted cases of the eye which were secondary
infections due to a metastasis from the pulmonary apparatus.
1933]
MOORE—BLASTOMYCOSIS 89
Laryngeal and tracheal blastomycosis are rare infections.
Jacobson records four in America and one in Europe. All the
patients were adult males working either with the soil or its
products, one being a clerk in a general store (Dennis,’ 18,
Downing, '18, C. Jackson, '26, and New, '28). The larynx
showed a “chronic inflammatory mucosa with a grayish, minutely
nodular surface in some portion of the lesion, with a few minute,
isolated, yellowish nodules." ‘There was often a reddish, raw
portion of the larynx due to the ease with which the superficial
layers came off with coughing. The process resembled very
closely tuberculosis.
Involvement of the tongue is perhaps a rare occurrence, but
cases are not reported in great numbers purely because sputa
smears usually show a variety of yeasts and thus no definite
etiological significance is attached to those obtained. The first
case was reported by Copelli (13), a second one by New (17)
from the Mayo Clinie. Since that time, however, Mazza and
Canevari (29) reported a case from Argentina, and Niño (29)
reported an infection of the lower lip with the involvement of the
tongue. Such an infection usually hinders respiration, inasmuch
as a tumor-like growth, as evidenced by Copelli and New, de-
veloped which enlarged in the back portion of the tongue and
practically filled the entire larynx.
SYMPTOMATOLOGY
Blastomycosis when of the primary cutaneous type presents
no clinieal symptomatology of discomfort or pain except for that
expressed because of the lesions. When of the systemic type,
however, the condition is very different. There are numerous
clinical factors to make a picture which might easily be confused
with a number of diseases.
According to numerous investigators, the onset of the disease
varies with the person and amount of infection. It may be
intense and acute, leading to death in a short time, or insidious
and mild, with a prolonged chronic condition, death finally
occurring as a result of a secondary complication. There is a
typieal set of symptoms once the disease is well established.
This consists of typical malaise, recurrent chills, loss of weight,
[Vor. 20
90 ANNALS OF THE MISSOURI BOTANICAL GARDEN
as evidenced in the present case, loss of strength leading to general
emaciation, night sweats, although morning sweats may be
present too, irregular fever, pain in affected parts, and a rapid
pulse.
The disease, as noted before, may be primary in the skin with
subsequent spread systemically or it may be systemic with the
formation in the later stages of nodular growths on the skin.
Unless the patient is well taken care of, systemic infection re-
sults, leading ultimately to death.
DIFFERENTIAL DIAGNOSIS
The final diagnosis of blastomycosis rests on the finding of
the organism either culturally in a lesion of the patient or, if
that be unavailable as in systemic disorders, the identification of
the fungus in biopsy or autopsy material.
As pointed out previously, the disease is protean in character,
with the result that a careful examination must be made to
diagnose it blastomycosis, comparing it with the several well-
known clinico-pathological entities which it may simulate. The
most noteworthy of these complicating diseases are as follows:
(a) The dermic lesions described previously, developing necrotic
and papillomatous growths or ulcers, resemble very closely
epitheliomas, differing only in the rapidity of evolution and the
absence of deep induration, verrucas, tuberculosis in its various
forms, and syphilis. Its resemblance to sporotrichosis (Lewis,
17) has often been noted, but it differs in being less sluggish. It
differs from syphilis only by the softness of the lesions, the red-
dish-blue ring around the lesion, and by a negative Wassermann,
with, of course, the presence of the organism in the blastomycosis
infection; (b) The systemic infections of blastomycosis must be
distinguished through laboratory methods from a great many
complications, particularly coccidioidal granuloma described in a
previous paper by the author (M. Moore, ’32). The organism of
coccidioidal granuloma, Coccidioides immitis, reproduces by en-
dosporulation, and the blastomycosis organism, through budding.
Furthermore, the lesions in the so-called ‘‘California disease"
are more rapid in evolution than in blastomycosis; (c) Infection
of the glands which is quite rare in blastomycosis often suggests
1933]
MOORE—BLASTOMYCOSIS 91
lymphatic leukemia, Hodgkin’s disease, and possibly lympho-
sarcoma; (d) abre i testinal lesions may often resemble typhoid
and in some cases the isolation of the organism is necessary to
rule out this disease; (e) Osseous infection quite often resembles
tuberculosis, petticularly as reported by Ryerson (708-09); (f)
Pulmonary blastomycosis very often presents the same clinical,
histological, and pathological pictures as tuberculosis, as noted
by Medlar (27) and Miller (27); (g) Infections of the brain often
confuse the pathologist or clinician with its similarity to torula
(Freeman and Weidman, ’23), epidemic meningitis, and even
tumors of the spinal cord; (h) Sutejew, Utenkow, and Zeitlin
(29) find that the use of bromides and iodides evidently causes
an allergy which in its reactions presents lesions similar to
those caused by the infective agent of blastomycosis and is often
confused with it, the latter differing in their more rapid evo-
lution.
It would seem, therefore, that the recognition of blastomycosis
is not a very easy matter. The really important fact concerned
with this work is to find the organism, which usually requires
laboratory technique, and to verify its pathogenicity by animal
inoculation in order to comply with Koch’s postulates.
Predisposing factors.—The infective agent shows no particular
preference as to sex, although more cases have been males than
females, of the industrial classes, chiefly the workers of the soil
and its products, a fact well exemplified in the present case.
There is no discrimination as to race or color, all peoples being
affected in like fashion. Stober ('14) correlates the type and
amount of infection with the habitat and environment of the
patient. As far as is known, all ages are susceptible, a similar
condition existing for many other fungi which tend to become
systemic, such as coccidioidal granuloma.
Treatment.—The successful therapeutic measures in blastomy-
cosis are few and limited. The best treatment or cure for the
disease rests of course on the skill of the physician in detecting it
at an early stage before it has seized a definite foothold, when it
can be kept from becoming systemic. If the disease is definitely
located in a particular section, surgery may be used to eliminate
it, as suggested by several authors (Wade and Bell, ’16, R. H.
[Vor. 20
92 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Jackson, '26). Cautery has also been used with beneficial
results. Hedge ('28) has employed carbon dioxide snow in
freezing cutaneous lesions with measurable success. X-ray
treatment has also been used frequently, being combined with
the administration of iodides. On the whole, primary cutaneous
lesions yield fairly readily to iodides and even the application of
crystal violet and gentian violet, although their use if at all
successful is empirical because dye therapy does not rest on any
scientific basis (Spring, '29).
In systemic infections, however, one has to contend with
complications. The extent of the infection cannot be determined
easily. The only cases to recover from an infection of this sort
are those to which special care and attention coupled with a
change to a clean, pure atmosphere had been made, with the
administration of large amounts of saturated iodide solution,
either potassium or sodium, although the former has been used
more extensively. The dyes mentioned above have proven
worthless to the systemic type of disorder. Several workers have
advocated copper sulphate, but many others have found this to
be useless. Roentgenotherapy has, as yet, no really therapeutic
importance. Stober has applied immuno-therapy in the form of
a vaccine of suspended blastomycotic cells, heated to 110° C., but
no definite results can be shown.
It would seem, therefore, that therapeusis is greatly in need of
investigation.
Immunological reactions.—Immunology in blastomycosis has
not reached any definite point as yet. Agglutinins have been
reported by some, and negative results by others. Precipitins
have also had the same reaction, as well as complement fixation.
On the whole, results are indefinite and a good deal more in-
formation is needed. The main difficulty seems to be that the
toxins of the blastomycosis fungus are difficult to demonstrate.
So far, immunology is an open book with only attempts proving
nothing written on its pages, and it is to be hoped that more
work can be done along this line for the benefit of those who may
be inflicted with this syndrome-complex.
Mortality.—The number of deaths resulting from blastomycosis
is a factor worthy of note, since systemic disorders due to the
1933]
MOORE—BLASTOMYCOSIS 93
typical blastomycotic organism usually prove fatal. This is so
because systemic infections are rarely diagnosed as such until
there have been cutaneous outbreaks, with the result that thera-
peutic measures are given too late for beneficial results. It is
difficult to quote figures because there are many cases occurring
which have rapid recovery and the physician does not report
them. Moreover, true blastomycosis is difficult to diagnose un-
less smears and cultures are made from the abscesses. In many
instances, several attempts are necessary before any fungous
growth is obtained, and unless the investigator is well trained in
mycological technique and in the recognition of such forms typical
of the Gilchrist organism the application of Koch’s postulates
would be essential, particularly with mice and guinea-pigs.
REPORT OF CASE
Clinical History—M. H. L. (Hospital No. 50095), a white, widowed male, farmer
by occupation, 43 years of age entered the Barnard Free Skin and Cancer Hospital,
at Saint Louis, Missouri, April 5, 1932, with an ulcerating, proliferating infection of
the left hand and forearm, which the patient claims to have had for 5 years.
Family History.—Father died as a result of high blood pressure, hypertension, at
age of 56 years. Mother dead due to throat trouble, at age of 30 years. Three
brothers and one sister dead due to an infancy cause. Patient has 7 children, all
living and well. Wife dead due to double pneumonia at age of 38 years. No
history of diabetes, nephritis, syphilis, or cancer in fami
Past History.—Past health has been good; usual obildliood diseases. He had
pneumonia at 15 years of age, bronchial pneumonia at 18 years of age, and influenza in
1918. He has had no operations other than those to be mentioned in the present
illness. Right leg injured 7 years ago
Personal History.—Patient denied any venereal disease.
Present Illness.—Patient stated that 7 years previously, a limb had fallen on his
right lower leg, causing a knot to form. The physician who examined his leg thought
he had a periostitis and opened the lesion on the leg. About 2 years later, or 5 years
previous to entering the hospital, a mossy, verrucous-like lesion similar to that on the
leg appeared on the back of the left hand. Both the lesions on the right lower leg
and on the hand continued to spread. The patient went to the Vanderbilt Univer-
sity Hospital, at Nashville, Tennessee, about a year later, at which time the eruption
had spread over most of the anterior and lateral sides of the right lower leg with two
lesions of the right thigh above the right knee and with involvement with most of
the back of the left hand and part of the forearm. He remained in the Vanderbilt
Hospital for about 3 months, during which time the lesions on the right leg and
arm were curetted and treated with X-ray. At the time of his discharge, the lesions
on his right leg had healed, as had most of the hand except that portion near the
thumb and the wrist. Another currettement was performed, but the lesion per-
sisted. He received two X-ray treatments each on the leg, arm, and hand. Six
years previous to entering the Barnard, he had lesions on the right elbow region;
[Vor. 20
94 ANNALS OF THE MISSOURI BOTANICAL GARDEN
these lesions healed. About 4 to 6 years before his entry he had lost some weight,
but at the present time complains of no recent loss of weight. He gives no histo
of hemaptysis, night sweats, fever, or frequent colds. About 10 days previous to
entry, a reddish lump developed on the flexor surface of the left lower arm near the
elbow. This lump is not very tender or painful. He has been applying potash on
the ulcerated area of the left hand
Physical Examination.—Patient is a thin, coóperative, moderately active, white
male 43 years of age, who has had the condition herein to be described as blastomy-
cosis of the left arm and hand for approximately 6 years and who had the same con-
dition on the leg for 3 years until it was curetted and X-rayed 4 years previously.
The right leg and left arm show scarring which will be described later.
Head.—Normal size and shape
Eyes.—Eyes react to light and accommodation. Reactions normal.
Ears.—No discharge, — normal.
Nose.—Septum intact, no ulce
rales —Slight redness.
Mouth.—Several teeth mi
Neck.—No stiffness or rigidity, tonsillar and cervical — pee
Thorax.—Thorax hairy, thin, and symmetrical; expansion fair.
Lungs.—Breath sounds a little harsh and rough over both | E principally the
right and right apex. Voice sounds normal, but louder on the right than on the left.
No persistent or moist ráles or ráles heard after coughing. Expansion and resonance
normal.
Cardiac.—Cardiac sounds a little slow and distant, but of a normal quality. No
enlargement or pathological murmur. Blood pressure, 106 systolic and 66 diastolic.
Abdomen.—No masses or tenderness. Inguinal glands a little enlarged.
Genitalia.—Normal male; no discharge or penile sore
Reflexes.—Superficial—present and active. Babinski negative. Deep—present
and active
Extremities.—Left arm and hand—The lower third of the left forearm, back of
the hand and extending 2 or 3 cm. on to the palm, is involved in an atrophic process
sharply defined superiorly and inferiorly with some scaling, no telangiectasis. On
the external surface of the forearm extending on to adjoining parts of the hand and
thumb is an ulcerative process which has been covered with a black crust. Ul-
cerated area of the left thumb and the flexor surface of the left forearm consists of
warty-like and cone-like, multiple abscesses with some elevation of the borders.
There is a deep scar on the right elbow region. The anterior, medial, and lateral
aspects of the right lower leg are covered with a thin, smooth scar about 12 to 14
inches long and covering two-thirds of the right lower leg. There are two scars
just above the right knee, about 4 inches, and 8 X 4 inches in diameter, respectively.
On the flexor surface of the left forearm, just below the elbow, is an abscess which
is red in color, oval in shape, semifluctuant, practically non-tender and not hot.
Extending up from and about the abscess is a chin of firm lymph nodes. The lymph
nodes above the left elbow and epitrochlear region are a little enlarged.
Laboratory Findings.—Urine negative, being pale straw in color and clear. Blood
tests showed 4,600,000 red blood cells and 10,200 white blood cells, with 84 per cent
hemoglobin. The differential blood count showed 26 lymphocytes, 8 large mononu-
clears and transitionals, 156 polynuclear neutrophiles, 2 polymorphonuclear eosino-
philes, and 2 basophiles. Wassermann negative. Smears from the left hand and
1933]
MOORE—BLASTOMYCOSIS 95
arm showed budding forms of yeast cells on April 7, 1932. Blood, urine, and pus
from left hand and lower arm and pus withdrawn from abscess of left arm were
inoculated on glucose-glycerine agar on April 8. Spinal puncture on April 12
showed a clear fluid with a normal pressure. Glucose-glycerine agar and blood agar
were inoculated with spinal fluid, with no growth occurring
April 9, 1932.—The patient was started on potassium iodide with dosage up till
signs of intoxication, and an ointment saturated with gentian violet was applied
locally. This treatment was followed with 10 per cent sodium iodide intravenously
for several days when patient showed an improvement
April 16, 1932.—X-ray of left forearm and wrist, right femur, and right leg showed
no abnormal bone changes. The hilar and bronchial structures of both lungs showed
considerable thickening, inflammatory in character, but the parenchymatous
portion of both lungs appeared free from any active pathology
7, 1932.—Patient showed & very marked alone AM with still some
evidence of trouble around the left thumb.
Clinical Diagnosis.—Blastomycosis of left hand and a
The agar inoculated from the left arm developed a ania which went through the
three stages typical of the organism of blastomycosis: the yeast-like growth; the
prickly type of growth; and the cottony type of growth. This culture was used for
the studies carried out in this paper.
ANIMAL INOCULATION
A suspension of a 10-day-old culture of the organism in sterile
saline was inoculated in a mouse, intratesticularly. An orchitis
developed and the mouse showed typical malaise, emaciation,
loss of appetite, loss of weight, rise in temperature, with death
ensuing in three weeks. The internal viscera showed numerous,
pin-point lesions at autopsy, which when squeezed exuded a
muco-purulent material from which the yeast cells were isolated.
This was in accordance with Koch’s postulates.
TECHNIQUE
In order to ascertain the morphological characteristics of the
fungus, the organism was suspended in hanging-drop cultures,
allowed to grow, and observations made from time to time. For
finer detail and structure, several transfers were made to slides on
which had been placed a drop of a mixture of glycerine (Merck
C. P.) and a 1 per cent solution of crystal violet. The fungus
was allowed to remain for a period of one-half to one hour to
allow for a clearing of the material and sufficient staining to render
satisfactory results. This method proved adequate for the work
here described. Another method was used also, whereby material
was fixed on a slide smeared with albumin and then stained with
[Vor. 20
96 ANNALS OF THE MISSOURI BOTANICAL GARDEN
methylene blue and eosine. The first method, however, was
sufficient.
DESCRIPTION
A study of the organism in lesions reveals a yeast-like growth
of spherical or ovoid, budding and branching cells with no evidence
of any filamentous hyphae. These cells, measuring approxi-
mately 7-12 y in diameter and sometimes as much as 20 y in
length, may occur singly, in groups of two's, three's or four's, as
individual colonies. On closer examination microscopically,
these yeast-like cells (pl. 6, figs. 1-9) show a heavy, reticulated,
granular, and in many cases, vacuolated, protoplasm, and a very
definite nucleus with emanating streams of cytoplasm. In many
cells the nucleus is barely distinguishable, being a mass of cyto-
plasmie structures, but further study after several subcultures
shows up this part of the cell very frequently. Many of the yeasts
of this group may show a double-contoured, highly refractile
membrane, and this feature is of little diagnostic use unless dem-
onstrated in tissue sections, as by Rewbridge, Dodge, and Ayers
(29) for Endomyces capsulatus, and by Moore (McBride and
Thompson, '33) for E. capsulatus var. isabellinus. Thus Wade
(16) ascribes to the fungus structure in tissues: (1) an inner
delicate capsula vera; and (2) an outer applied capsula sclerotica.
In any case, the capsule is lost on repeated cultivation on arti-
ficial media.
With agar as a substrate, the yeast-like cells elongate (pl. 6,
figs. 10-12, 14-17) and on acid media become thin-walled and
long with a diameter of 2-214 u. These hyphae become inter-
twined and are composed chiefly of isodiametric cells. On neutral
or slightly alkaline media, with protein as the chief source of
nitrogen, the hyphae are thicker and shorter, with a diameter of
3-4 u. This condition is especially true of media with an excess
of carbohydrate as found in glycerine agar.
Budding cells are numerous on slightly acid media, being about
9 u in diameter (pl. 6, figs. 26 and 28).
The hyphae, at first clear, become granular, and at various
points along the sides develop numerous small, knob-like pro-
jections of the limiting membrane, which enlarge, round out or
become pyriform and sessile, measure approximately 5 y in
1933]
MOORE—BLASTOMYCOSIS 97
diameter, and occur usually near a septum. These are known as
conidia and may remain attached, at times to small stems or
sterigmata, or may become free and develop in the media by
budding (pl. 6, figs. 24, 26, 28-29).
Racquet mycelium (pl. 6, figs. 23-24), a phenomenon char-
acteristic of the fungi of this group and especially of the Tri-
chophytons and various other Ascomycetes, is common here,
having the swollen portion 5-6 y in diameter and the narrow
section 3-314 y in diameter. Chlamydospores may be found
arising in the hyphae or terminal as hypnospores, varying in
size and shape from round cells 7-81% y in diameter, to elongated,
widened cells 5144-714 u wide and 12-15 y long (pl. 6, figs. 22,
25, 27—29, 31, 32, and 37), or they may arise as sessile cells from
the hyphae (pl. 6, fig. 36). Round, thick cells with a coarse,
granular cytoplasm are particularly evident on cornmeal agar
(pl. 6, figs. 26 and 28).
When first examined in a hanging-drop preparation, one may
see oil droplets on the hyphae which are strongly suggestive of
endospores. These disappear, however, when the organism is
stained. In some cases, as in pl. 6, fig. 25, a hollow sphere or
vacuole surrounded by a hyaline, gelatinous substance may be
present in the filament.
The development in tissue, as has been noted, is chiefly by
budding or gemmation. ‘The process begins with the projection
of the inner layer of the cell or endosporium, following Hektoen,
which pushes the transparent zone and outer membrane in front
of it. The bud becomes enlarged and surrounded by the same
walls as surround the mother cell, and division takes place by the
presence of a cross-wall which is formed by the pinching in of
the cell (pl. 6, figs. 1-3, 6-7). On artificial media, on the other
hand, proliferation of the fungus is brought about through sexual
reproduction which is heterogamous. Two terminal cells may
fuse (pl. 6, fig. 19) or two hyphae growing side by side may send
out lateral cells which copulate (pl. 6, fig. 18). In either case,
a spherical ascus results which may be terminal on a long filament
or lateral on a short peduncle (pl. 6, fig. 33) and has a thick
capsule (pl. 6, figs. 21 and 33), sometimes surrounded by a sheath
as in pl. 6, fig. 35. The latter case, however, is rare and was
[Vor. 20
98 ANNALS OF THE MISSOURI BOTANICAL GARDEN
observed only three times. There are formed 8 spherical to
ovoid, smooth spores which are hyaline when young and held in
a gelatinous substance and when older become chamois-colored
and granular, and have at maturity a diameter of 2-3 y, varying
on different media (pl. 6, figs. 38 and 33).
This organism therefore agrees with that described by Gilchrist
and Stokes as Blastomyces dermatitidis, but further observation
has here been made on the sexual development.
CULTURAL DESCRIPTIONS
The culture in this case was growing on glucose-glycerine agar,
being an inoculum from an abscess on the left arm. All cultures
used in the cultural determinations were taken from the above
tube and grown at room temperature, approximately 22° C.
Having the two stages so common with yeast-like organisms
and characteristic of several members of the Endomycetales, it
was thought desirable to transfer the fungus on a wide variety of
media and pH range. Possessed with saccharomycetous prop-
erties, on the one hand, and filamentous fungous affinities, on
the other, the above method of culturing proved to be satis-
factory in this work.
The following media used are arranged in the order of decreas-
ing hydrogen-ion concentration.
Raulin's Solution (pH 4.1).—Culture shows a thin, smooth
suspension of yeast-like cells, budding and branching, varying in
size from 414-514 u x 7-815 u, with several showing a change to
filamentous formation.
Richards’ Solution Agar (Media consisting of Richards’ solution
with the addition of 1.5 per cent agar. pH 4.4).—Growth sparse, of
fine filaments. Colony 3V$ cm. in diameter at end of 30 days.
Culture shows long hyphae projecting from edge of growth, 214-
3 u in diameter, with numerous budding cells. Filaments
branching, with cross-walls, numerous chlamydospores, and
swellings. Characteristics of the group present. Color of
colony isabella to cinnamon, strongly suggestive of chamois, due
to the spores and asci which are in abundance.
Czapek's Agar (pH 4.4).—Color of colony white, becoming
chamois with age. Growth very sparse and cottony, with much
1933]
MOORE—BLASTOMYCOSIS 99
of the mycelium submerged in the agar. Colony 4 cm. in diam-
eter at end of 24 days. Hyphae long and thin, 112-2 y in
diameter, with swellings approximately 4 x 12 y, several thick-
walled chlamydospores 714 x 14 y, and numerous terminal
hypnospores. Several 8-spored asci seen, as well as many
conidia.
Malt Extract Agar (pH 5.8).—Growth slow and cultural
characteristics insufficiently different to be taken into discussion.
Sabouraud's Agar (pH 5.6).— Growth rapid, profuse, obtaining
a diameter of 714 cm. at end of 30 days. Culturally the colony
simulates very much that of Microsporon audouini of Ota and
Langeron in the presence of several radiating ridges from the round
center, the inoculum, and the several concentric rings of growth
of decreasing abundance, just outside the ridges. Color of
colony white when young and becoming the characteristic
chamois when older. Like M. audouini, it has racquet mycelium,
chains of round cells on a hypha measuring 314 y in diameter,
numerous chlamydospores 8 x 12 y, terminal hypnospores 5 x
11 u, and many conidia, characters found also in the Tricho-
phytons and peculiar to Zndomyces capsulatus. Unlike M.
audouini, however, this organism reproduces by the formation
of asci which are numerous here, measuring from 10 to 13 y iu
diameter, containing 8 spores.
Sabouraud's Broth (Sabouraud's medium minus the agar. pH
5.6).—Culture consists of submerged mycelium of large flakes,
each measuring approximately 2 cm. in diameter at end of 24
days. Mycelium floating on surface, dry, and chamois colored,
with white region, presumably the young hyphal elements. In
general, growth is good. Microscopically, the culture shows
long, narrow hyphae 214 y in diameter, branching and inter-
twining. Submerged mycelium shows almost no swellings,
chlamydospores, terminal hypnospores, nor thick-walled cells, as
compared with the great number found in that on the surface:
The several that are present, however, show a great reduction in
size and form from the exposed, the measurements of which are
similar to those on agar.
Potato-dextrose Agar (pH 5.6).—Growth profuse and cottony,
covering the surface of the agar completely. Diameter of colony
[Vor. 20
100 ANNALS OF THE MISSOURI BOTANICAL GARDEN
71% cm. after 24 days. Color cinnamon, with colony showing
concentrie circles of color alternating with white, and the cinna-
mon very pronounced, due perhaps to the medium constituents.
Hyphae 3 y. in diameter, with numerous, thick-walled cells 744 u
in diameter, budding cells, swellings 5 x 12 u, and chlamydospores
varying in size from 4-7 x 9-14 u. Asci numerous, measuring
approximately 13 y in diameter.
Corn-meal Agar (product of Digestive Ferments Co. pH 6.0).—
Growth poor, colony being 114 cm. in diameter at end of 24 days.
Color white. Growth around inoculum loose and cottony.
Hyphae short, thick-walled, 214 » in diameter, with numerous
budding cells approximately 7 » in diameter. Chlamydospores
numerous, 7 x 13 y, terminal hypnospores several, 5 x 12 u; a few
asci seen, 11 y in diameter. Conidia abundant, 5 y in diameter.
June-beetle Agar (medium consisting of a 4 per cent extract of
June beetles, Lachnosterna fusca, plus 1.5 per cent agar, sterilized
at 20 pounds pressure for 20 minutes, with a final pH 6.1).—
Growth of loose, flat, cottony mycelium, forming concentric
circles of decreasing abundance until a ring of fine filaments
surrounds the culture. Colony 514 cm. in diameter at end of
24 days. Hyphae 2-21% y in diameter, with many conidia 5 y
in diameter. Asci 12-13 y in diameter, thick-walled, enclosed in
a sheath. Abundance of racquet mycelium.
June-beetle Dextrose Agar (above medium plus 2 per cent dextrose).
—Growth fair, attaining a diameter of 3 cm. at end of 30 days.
Colony bright chamois in color, cerebriform, and cottony. Many
conidia, 415-5 y in diameter. Hyphae 215-3 y in diameter and
fairly short. Chlamydospores 8 x 16 y and numerous, as well as
terminal hypnosp 5x12u. Asciround, 12-14 y in diameter.
Lactose Agar (product of Digestive Ferments Co., lactose broth
plus 1.5 per cent agar. pH 6.8).—Growth good, reaching a diam-
eter of 6 cm. at end of 24 days. Colony chamois-color, profuse
and cottony, with a region of very fine mycelium surrounding it.
Hyphae 31⁄4 y. in diameter with numerous conidia 5 y. in diameter,
budding off. Many thick-walled resting cells 7 » in diameter.
Characteristic racquet mycelium, chlamydospores, terminal
hypnospores, with properties similar to those on Sabouraud’s
agar.
1933]
MOORE—BLASTOMYCOSIS 101
Lactose Broth (product of Digestive Ferments Co. pH 6.8).—
Growth good, large white flakes being formed in the solution which
later become intertwined, forming a mat of mycelium 7 y in
diameter. Hyphae slightly reduced, 3 u in diameter, 114 y in the
younger filaments. Preponderance of budding cells 7 u in di-
ameter, with thick-walled chlamydospores, asci, and terminal hyp-
nospores, but reduced in size as compared with the growth on
agar.
Eosine-methylene-blue Agar (agar used as one of a routine, prod-
uct of Digestive Ferments Co. pH 7.0).—Growth good, with a
diameter of 514 cm. at end of 24 days. Culture compact, due to
the hyphae having absorbed the stain from the substrate and
turning the mycelium pink. Colony appears powdery with age.
Hyphae characteristic, with swellings, 3 » in diameter. Many
conidia, 5 x in diameter, and hypnospores with several chlamy-
dospores.
Glycerine Agar (nutrient agar as prepared by the Digestive Fer-
ments Co. plus 6 per cent glycerine, Merck C. P. pH 7.1).—Growth
fair, having a diameter of 5 cm. at end of 24 days. Culture shows
a crinkled, moist region of budding yeast-like cells and a dry
filamentous, cottony, chamois-colored region which has changed
to the mycelial form characteristic on agar. Filamentous hy-
phae 314 y in diameter, characteristic swellings being present
which are slightly larger than those found on lactose agar.
Racquet mycelium also present.
Nutrient Agar (product of Digestive Ferments Co. pH 7.2).—
Growth rapid, covering a region 7 cm. in diameter at end of 30
days. Colony filamentous, cottony, brown, with concentric rings
of growth, the outermost being white. Hyphae 214 u in diam-
eter. Growth similar to that on Sabouraud’s agar micro-
scopically, with numerous conidia 5 y in diameter and asci 13 y
in diameter.
Nutrient Broth (pH 7.2).—Culture forms a mat of intertwining
mycelium of long hyphae 2-214 y in diameter, with swellings,
asci, and chlamydospores. Very few conidia. Terminal hyp-
nospores several, but reduced in size, 4 x 9 y.
Endo's Agar (product of Digestive Ferments Co. pH 7.5).—
Growth fair, colony having a diameter of 314 cm. at end of 30
[Vor. 20
102 ANNALS OF THE MISSOURI BOTANICAL GARDEN
days. Culture shows radiating ridges from center of inoculum,
with growth becoming flat due to the stain in the medium which
is absorbed by the hyphae, as in the case of the eosine-methylene-
blue agar, giving the mycelium a pink color. Microscopically,
the hyphae have a diameter of 24% y. Numerous conidia 5 y
in diameter. Culture otherwise similar to that on eosine-
methylene-blue agar.
Gelatine (nutrient agar plus 1.5 per cent gelatine).—Slow lique-
faction beginning after 30 days.
Culturally, the fungus is very characteristic of the organism
of blastomycosis in that it passes through the three typical stages:
the moist, yeast-like stage with a flat growth; the prickly culture
with the colonies simulating greatly small burrs (coremia); and
the final, cottony growth present on agar after extended growth.
DISCUSSION
As stated in the introduction of this paper, it would seem that
the syndrome-complex, commonly known as blastomycosis, has
an innumerable list of etiological factors, each causing a condition
so much like the other that clinicians have grouped them under
one head. However, should one encounter any of these in a
clinic one would find that therapeutic measures are so vastly
different, varying with the organism, that a direct and accurate
knowledge of the causative agent in each particular patient is
absolutely essential.
In the past, medical men, not particularly trained in mycologi-
eal taxonomy, were inclined to class together all fungi presenting
ascomycetous characters under one name, Blastomyces. So
great is the confusion to-day that it is necessary to pick out these
pathogenic fungi and classify each one separately.
The organism isolated in the first case was termed Blastomyces
dermatitidis by Gilchrist in 1894 because of its budding properties
in the lesion. In a case of dermatitis reported by Gilchrist and
Stokes (’96) the organism, which was evidently of the type termed
Blastomyces, was called an Oidium. In a following paper (Gil-
christ and Stokes, '98), it was made known that the organism
described in the previous paper was called an Oidium because it
did not ferment glucose, saccharose, or lactose, and although
1933]
MOORE—BLASTOMYCOSIS 103
developing by gemmation or budding in the tissues, human and
animal, developed mycelia with the formation of conidia upon
artificial media. Ricketts (01) made an extensive study of the
organism, distinguishing it from several of the yeasts but failing
to consider several of the yeast-like fungi, and proposed definitely
the name Oidium for the genus of the Gilchrist fungus. After
this work, several terms were applied to the disease. Busse (’94)
described his case a short time after Gilchrist reported his and he
named the organism Saccharomyces hominis. Vuillemin in a
later publication assigned the organism to the genus Cryptococcus
and called it C. gilchristi. However, he failed to make a careful
study of the organism culturally on artificial media, and a
classification which places a great emphasis on the yeast-like
appearance of the fungus in lesions is not exactly justifiable.
Brumpt (’27) places the organism in the genus Mycoderma,
calling it M. dermatitis. This terminology, however, is synony-
mous with Oidium, and in that case is likewise useless.
For a great number of years, no great work was done to establish
definitely the position of Gilchrist’s organism, and the name
Blastomyces as created by him still held sway. The term pre-
sents a lot of difficulties. In the first place, the Blastomycetes,
according to Buschke, are that group which develops through
budding, provided à mycelium is formed on agar, while to the
group of Blastomycetes, as Naegeli names budding, would belong
the genera Endomyces, Saccharomyces, Cryptococcus, Monilia, and
Oidium. Now the question arises as to what the actual meaning
of the word blastomycete is. According to Vuillemin (’01), it
does not designate a natural group, a botanical family based on
genealogical affinities. There is in existence a genus Blastomyces,
but these organisms are not budding fungi in the sense of Buschke.
They are filamentous fungi whose spore-bearing elements, whether
terminal, lateral, or intercalary, can be isolated by disarticulation,
following Costantin and Rolland (quoted by Vuillemin). Frank
(quoted by Vuillemin) established the Blastomycetes as an order
to include such fungi as the beer yeasts whose elements are iso-
lated by budding and not by disarticulation. In this respect, by
virtue of the law of priority, it would seem that the name Blas-
tomycetes, as designated by Frank, should remain. However,
[Vor. 20
104 ANNALS OF THE MISSOURI BOTANICAL GARDEN
if by general agreement the name of a genus could replace that of
an order, then, according to the rules of nomenclature, the genus
of Costantin and Rolland is legal, and the name as designated
here is not legitimate by reason of the lack of distinct characters
which have no generic value.
Castellani recently proposed a new classification of yeast-like
or budding fungi based on the presence or absence of ascospores,
which includes families of both the Ascomycetes and Fungi Im-
perfecti.
1. Saccharomycetaceae: budding cells, asci, and ascospores,
but no mycelium in culture.
2. Endomycetaceae: budding cells, asci, and ascospores, with
mycelium in culture.
3. Cryptococcaceae: budding cells (blastospores), no asci and
no mycelium in culture.
4. Oosporaceae: budding cells, no asci, but mycelium in culture.
In addition to this family classification, he created a new genus
which he calls Blastomycoides, to which he assigns three species,
and places it in the family Oosporaceae: 1. Blastomycoides der-
matitidis, synonym Blastomyces dermatitidis Gilchrist and Stokes;
2. Blastomycoides immitis, synonym Coccidioides immitis Rixford
and Gilchrist; 3. Blastomycoides tulanensis Castellani. He de-
fines the genus Blastomycoides as: ‘‘Oosporaceae appearing in the
lesions as large roundish cells from eight to twenty microns in
diameter, or larger, with the protoplasm containing a number of
well-marked granules or spherules, and with a membrane showing
a well-defined double contour; in dextrose agar cultures a large
amount of mycelium is present." He bases further differentiation
of the three species on their cultural differences when grown on
mannitol, lactose, glucose, and galactose agar.
The second species that he names, Blastomycoides immitis, has
already been discussed and classified by the author in a previous
paper (M. Moore, ’32). The author has made no pretext of
studying the third species, so that nothing can be said about that.
The first species, however, Blastomycoides dermatitidis, is alto-
gether misplaced, simply because there are asci present in the
mycelium in culture. This of course would refer the genus to the
family Endomycetaceae, in which group the writer definitely
establishes the organism.
1933]
MOORE—BLASTOMYCOSIS 105
Observations on the growth, development, reproduction, and
further evolution of the fungus show that there are budding cells
in the lesions, mycelium formed on agar with an intermediate
stage showing the change from the yeast-like to the filamentous
form. In accordance with this, Mellon (724, ’26, ’26a) has
recorded the fact that asci do occur particularly ‘‘in the so-called
secondary colonies of the cultures which also contained ‘dau-
ernzellen’ and pigmented oidia." The author wishes to affirm
Mellon’s findings as to the presence of asci, but suggests that
these structures are present in the third stage, whereas the second
step would consist of the intermediary forms which have an
appearance very much like greatly enlarged oidia. Furthermore,
it would seem that Mellon’s description was indefinite, inasmuch as
he refers to an ascus as an ascospore, and oil droplets are suggested
by him as being chromatin indicators and forerunners of the future
spores. Such factors as these are very important in the taxonomy
of this type of fungi and should not be dealt with so promiscu-
ously. Furthermore, Mellon has not paid much attention to the
fact that no matter how old the lesion may be the blastomycosis
organism found there does not change from its yeast-like, budding
growth until it has been transferred to artificial media, where the
change is an adaptation to the mode of life it must lead; in other
words, the change from active parasitism to one of saprophytism.
It is to be understood, however, that a change such as suggested
here will not necessarily reduce its viability, at least for the time
being.
A study of the evolution of the organism has repeatedly shown,
in several hanging-drop cultures, that reproduction is hetero-
gamous, as given in the description, with the final formation in
the series of a large eight-spored ascus. Mellon in his papers
consistently shows a four-spored ascus. It would seem, therefore,
that he either has an organism unlike the one here described for
the blastomycosis parasite, or else he has taken for granted as
spores the four oil droplets which may and often have been found
to occur on a mature eight-spored ascus, as was evidenced by
the author on another ascomycete, Endomyces capsulatus var.
isabellinus Moore, which was described in a case in another
paper (McBride and Thompson, ’33) and also in Endomyces
[Vor. 20
106 ANNALS OF THE MISSOURI BOTANICAL GARDEN
capsulatus Rewbridge, Dodge and Ayers (29). Furthermore, it
is quite possible that Mellon has observed the ascus just previous
to the division of the nuclei, in the formation of the eight-spored
ascus. "This latter statement is only a conjecture on the part of
the writer, but in any case the cultural descriptions do not agree
with those given by the early investigators and with which the
organism here described does agree.
In view of such criteria, it would seem that the organism
formerly described as Blastomyces dermatitidis Gilchrist 1894, is
not strictly a member of that genus, the name of which, on
account of its etymological derivation, is essentially a misnomer.
Because of its morphological characteristics, Blastomyces derma-
titidis does not present those affinities entirely but simply as one
phase of its life cycle. However, particularly because of its asco-
mycetous attributes, it should belong to the class Ascomycetes,
order Endomycetales, family Endomycetaceae, and because of
its similarity in morphology and reproduction (perfect stage) to
that of Endomyces capsulatus and its variety, it should belong to
the genus Endomyces. If taxonomic position in this family and
genus be dependent on the number of spores in the ascus, it
would seem, according to Whitman ('13), that this organism
should belong in the genus Oleina. However, the genus Endomy-
ces contains a number of pathogenic species with eight-spored asci,
whereas Oleina has no pathogenic species, and until a classifica-
tion better than the one now in existence be established, dermati-
tidis should be placed with Endommyces.
From the above statement it would appear that the organism
should now be known as:
Endomyces dermatitidis (Gilchrist 1894), M. Moore, n. comb.
Mycelium in lesions of budding yeast-like cells 7-12 y in diam-
eter and sometimes as much as 20 y in length, occurring singly,
in groups of two's, three's, or four's. Growth on agar of isodia-
metric cells 2-214 y in diameter on acid media and 3-4 y in diam-
eter on slightly alkaline media. Hyphae septate, with conidia
pyriform or round, pedunculate or sessile, 5 v in diameter.
Racquet mycelium present, 5-6 y in diameter at swollen portion
and 3-31 y in diameter at narrow portion. Chlamydospores
terminal or lateral or intercalary, 514—714 x 12-15 u, or sometimes
1933]
MOORE—BLASTOMYCOSIS 107
round, 7 y. in diameter. Copulation heterogamous, asci spherical,
8-13 y in diameter, with 8 spherical to ovoid, smooth, hyaline
to light chamois-colored spores 2-3 y in diameter, at maturity.
Colony white in color, becoming cinnamon to brown with age.
SUMMARY
1. The history of blastomycosis is given, with a review of the
early work on yeast-like, fungous pathogenicity, and a report of
the first case published.
2. The etiology and clinical manifestations represent a number
of conditions due to several yeast or yeast-like organisms: Sac-
charomyces, Oidium, Monilia, Endomyces, Cryptococcus, and
Coccidioides, which have been placed in one category to constitute
the agents responsible for the syndrome-complex, blastomycosis.
3. The disease is shown to simulate several conditions, in
which cases the diagnosis must be arrived at through the isolation
of the organism and the application of Koch’s postulates.
4. Immunological reactions and therapeutic measures are as
yet indefinite as to specific results, although beneficial results
have been reported by the use of iodides.
5. A case of blastomycosis of the arm and hand is reported.
6. There is a description of the organism, culturally and mor-
phologically, showing its relationship to the class Ascomycetes.
7. The fungus is definitely established as Endomyces dermati-
tidis of the family Endomycetaceae.
ACKNOWLEDGMENTS
The author wishes to express his sincere gratitude to the
following: Dr. Carroll W. Dodge, Professor of Botany in the
Henry Shaw School of Botany of Washington University, for his
interest, criticisms, and helpful suggestions; Dr. George T. Moore,
Director of the Missouri Botanical Garden, for the courtesies
extended; Dr. Martin F. Engman, Dermatologist to the Wash-
ington University Hospitals, for the use of the data with respect
to the case reported; and Miss Nell C. Horner, librarian of the
Missouri Botanical Garden, for her assistance.
[Vor. 20
108 ANNALS OF THE MISSOURI BOTANICAL GARDEN
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[Vor. 20
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—— ———, (29a). Onixis y perionixis de origen blastomicósico (Estudio clinico y
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—, Palant (30). Nuevas observaciones de onixis y peri-
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Ormsby, 2 S. (21). Blastomycosis. A practical treatise on diseases of the skin.
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03
1903.
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[Vor. 20
114 ANNALS OF THE MISSOURI BOTANICAL GARDEN
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MOORE—BLASTOMYCOSIS 115
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[Vor. 20, 1933]
116 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 6
All drawings made with camera lucida at a magnification of 800.
Figs. 1-9. Yeast-like cells
Figs. 1, 6, 7. On Raulin’s solution.
Figs. 2-5, 9. On Sabouraud's agar.
Fig. 8. On glycerine agar
Figs. 10-12, 14-17. Yeast-like cells showing a change to mycelial formation.
Figs. 10, 12. On potato-dextrose agar.
Figs. 11, 14. On glycerine agar.
Figs. 15-17. On Sabouraud’s agar.
Fig. 13. Germinating spores on Richards’ solution agar.
Fig. 18. Heterogamous copulation of lateral cells on Sabouraud's agar.
Fig. 19. Heterogamous copulation of terminal cells on Richards’ solution agar.
Fig. 20. Copulating branch on corn-meal agar
Fig. 21. Maturing ascus on Richards’ solution agar.
Fig. 22. Terminal hypnospore on Sabouraud’s agar.
Fig. 23. Racquet mycelium on June-beetle agar.
Fig. 24. Mycelium showing conidia on Richards’ solution agar.
Fig. 25. AN showing round terminal chlamydospores and swollen hypha
on nutrient agar
Figs. 26, - Myoslium showing conidia, oidia-like cells, and resting cells on
corn-meal ag
Fig. 27. Chesed on Czapek’s a
Fig. 29. Mycelium showing conidia on oue agar.
Fig. 30. Racquet formation on Czapek's agar
Fig. 31. Terminal chlamydospore on — ager.
Fig. 32. Terminal hypnospore on Endo’
Fig. 33. Maturing lateral ascus on cedi agar
Fig. 34. Racquet mycelium on June-beetle dextrose agar.
Fig. 35. Ascus — with a third sheath in proximity to a round resting cell,
on potato-dextrose a
Lateral na on — agar.
Fig. 37. Resting cell on Sabouraud's a
Fig. 38. Mature ascus on sor oca agar.
Ann. Mo. Bor. Ganp., Vor. 20, 1933 PLATE 6
MOORE—BLASTOMYCOSIS
[Vor. 20, 1933]
118 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 7
Fig. 1. Photograph of hand of patient on day of entry, April 8, 1932, showing
lesion involving portion of thum
ig. 2. Photograph taken on Apel 8, 1932, showing abscess on flexor surface of
left a arm.
g. 3. Photograph showing marked improvement after treatment with sodium
iodide intravenously.
Photograph showing almost complete healing.
ANN. Mo. Bor. Ganp., Vor. 20, 1933
9017
50099
PLATE 7
ee
F3312?
D50095
MOORE—BLASTOM Y COSIS
A MONOGRAPH OF THE AMERICAN SPECIES
OF THE GENUS HALENIA!
CAROLINE K. ALLEN
Formerly Missouri Botanical Garden Special Fellow in Botany,
Henry Shaw School of Botany of Washington University
INTRODUCTION
Frequent attempts to determine recent collections of Halenia,
especially from Central and South America, have revealed the
need for a comprehensive taxonomic treatment of the American
species of the genus. Incidental determination of isolated
species in herbaria, which furnish scanty material, has led to
error and the duplication of species has resulted. Few of these
fragmentary treatments are provided with adequate descriptions,
and still fewer are accompanied by illustrations. The present
paper is a monographie study of the American species of Halenia.
The first portion is devoted to the North and Central American
representatives of the genus and the second to those of South
America.
The writer at this time wishes to express her appreciation to
Dr. George T. Moore, Director of the Missouri Botanical Garden,
where this work has been carried on, for the privilege of using
the facilities afforded by the herbarium, library, and laboratory.
It is with pleasure, indeed, that she also acknowledges the courte-
ous assistance and helpful criticism offered by Dr. J. M. Green-
man, under whose personal supervision the study was begun
and completed.
For the loan of specimens for study the writer is especially
indebted to the curators of the herbaria of the following institu-
tions: Gray Herbarium of Harvard University, New York
Botanical Garden, Philadelphia Academy of Natural Sciences,
United States National Herbarium, Field Museum of Natural
History, the Dudley Herbarium of Stanford University, Brooklyn
Botanic Garden, Iowa Agricultural College, Natural History Mu-
1 An investigation carried out at the Missouri Botanical Garden in the Graduate
Laboratory of the Henry Shaw School of Botany of Washington University, and
submitted as a thesis in partial fulfillment of the requirements for the degree of
doctor of philosophy in the Henry Shaw School of Botany of Washington University.
Issued April 29, 1933.
Ann. Mo. Bor. Ganp., Vor. 20, 1933. (119)
[Vor. 20
120 ANNALS OF THE MISSOURI BOTANICAL GARDEN
seum of Vienna, Botanical Museum of Stockholm, the Delessert
Herbarium at Geneva, Royal Botanie Gardens, Kew, and the
British Museum of Natural History.
Particularly does the writer wish to express her gratitude to
Mr. R. I. Cratty, of the Iowa State College, for the loan of
specimens from the Parry Herbarium deposited there; to the
curator of the Herbarium of the Botanical Garden of Madrid,
for supplying a photograph of the type of Swertia cucullata;
and to the curator of the Herbarium of the Jardin des Plantes,
Paris, for photographs of the types of Humboldt, Bonpland and
Kunth.
In connection with visits to various herbaria, the author
desires to acknowledge the courtesy and kindly assistance of Dr.
Ernst Gilg and Dr. Robert Pilger, of the Botanical Garden at
Berlin-Dahlem; Dr. Walter Robyns of the Botanical Garden,
Brussels; and Mr. Spencer Savage, in charge of the Linnaean
Herbarium, of the Linnaean Society of London. "Thanks are due
also to Dr. George E. Nichols, Director of the Marsh Botanical
Garden, Yale University, for his kindness in procuring seeds of
Halenia; to Miss Nell C. Horner, Librarian of the Missouri
Botanical Garden, and Dr. John H. Barnhart, of the New
York Botanical Garden, for assistance in bibliography; to Dr.
Roland V. La Garde, of the Missouri Botanical Garden, for
preparation of photographs; and to the George F. Cram Com-
pany, for permission to use the copyright outline maps of North
and South America.
HISTORY OF THE GENUS
Linnaeus in the ‘Amoenitates Academicae," which appeared
in 1751, published short descriptions of two genera of the Genti-
anaceae, namely, Swertia, consisting of five species, and Gentiana,
of twenty-three. Under Swertia Linnaeus listed the spurred
gentian with the following description.
“4. Swertia corollis quadrifidis quadricornibus.
Amoen. acad. 2. p. 344.
Habitat in Sibiria, Gmelin; Canada, Kalm.”
This was apparently the only species of spurred gentian he had
! Linnaeus, Amoen. Acad. 2: 344. 1751.
1933]
ALLEN—THE GENUS HALENIA 121
ever seen, and, it being closely related to the Swertia he knew, he
placed it in that group, giving it the specific name corniculata or
"^ horn-tipped," as distinct from the others.
Gmelin, in his ‘Flora Sibirica,? published in 1769, referred to
this as being synonymous with his genus T'etragonanthus, which
he had described or mentioned in a previous book or manuscript,
and which he evidently based upon specimens collected by G. H.
Stellar. 'This work contains a brief description, but a good
illustration of Swertia.
On account of the presence of spurs on the corolla, Bórck-
hausen,’ in 1796, segregated Swertia corniculata from the Linnaean
genus Swertia and called it Halenia after Jonas Halen. Although
the latter had included a short description of it in a previously
published dissertation on Kamtchatka plants, Bórckhausen
must be considered the author of the genus. He cited as a
synonym Swertia corniculata Linnaeus, but changed the binomial
name to Halenia sibirica.
The name Swertia, however, persisted for some time in litera-
ture, the generic descriptions becoming more elaborate and
detailed with each publication. Ruiz and Pavon described
and illustrated Swertia umbellata from Peru‘ in 1802. Michaux
included the genus in his ‘Flora Boreali-Americana'5 appearing
the following year, and Humboldt, Bonpland and Kunth,’
described six new species from Mexico and South America in
1818. A few species, together with a new variety, were published
in ‘Linnaea’ by Schlectendal and Chamisso? in 1830. But it
was not until Grisebach began his extensive study of the Gentian-
aceae that any attempt was made to bring together the species
of the world. As a result, when his ‘Observationes’! appeared
in 1836, followed in 1839 by the 'Genera et Species Gentian-
earum,'? many of the existing names fell to synonymy.
Grisebach divided Halenia into two main divisions: the first,
2 Gmelin, Fl. Sib. 4: 114. 1769.
3 Bórckhausen in Roemer, Arkiv für Botanik 11: 25. 1796.
* Ruiz & Pavon, Fl. Peruv. 3:21. pl. 242. 1802.
5 Michaux, Fl. Bor.-Am. 1: 97. 1803.
$ Humboldt, Bonpland & Kunth, Nov. Gen. & Sp. Pl. 3: 174. 1818.
1 Schlectendal & Chamisso in Linnaea 5: 122.
$ Grisebach, Obs. Gent. 36.
? Grisebach, Gen. & Sp. Gent. 322-398, 1839.
[Vor. 20
ev. ANNALS OF THE MISSOURI BOTANICAL GARDEN
with spurs ascending and spreading; the second, with spurs
pendulous to ineurved. He devoted careful attention to the
descriptions, synonymy, affinities, differentiating characters,
and habitat of each species mentioned. The collector and type
were given in each case. He also separated from the Swertia of
Humboldt, Bonpland and Kunth two species, brevicornis and
parviflora, and founded on them a new genus Hxadenus distin-
guished by the presence of pits at the base of the corolla, instead
of spurs, and by central placentation. This abolished the
Swertia of Humboldt, Bonpland, and Kunth, which was based
on Swertia corniculata Linnaeus, and left the non-spurred Swertia
originally described by Linnaeus a genus entirely distinct from
our present Halenia.
Hooker's *Flora,'!? published in 1840, contained good illustra-
tions of Halenia deflexa with the varieties Brentoniana and
heterantha.
Nearly a decade elapsed before any significant study was done
on the genus as a whole. Bentham," in 1839-1840, in describing
Hartweg’s plants from Mexico added two new species, multiflora
and decumbens. Martens and Galeotti in 1844 in their *'Enu-
meration of Mexican plants collected by Galeotti"!? described,
along with other new Halenia species, two new species of Exa-
denus. Walpers’ ' Repertorium '? contains reference to Exadenus,
but Weddell in 18591 merged the two genera. He considered
the group as a whole to consist of two main subdivisions, the
first being that group with spurs, and the second the spurless
species. ‘The former he subdivided into section l—ovary uni-
locular; section 2—ovary bilocular, and the corolla having small
spurs. Here he placed the two original species of Hzxadenus
Grisebach which were based on the species of Swertia Humboldt,
Bonpland & Kunth, namely, parviflora and brevicornis, but
neglected to transfer the species described by Martens and
Galeotti. The second subdivision, without spurs, he also divided
into two sections, on the presence of a uni- or bilocular ovary.
10 Hooker, Fl. Bor.-Am. 2: 67. pl. 155-6. 1840.
11 Bentham, Pl. Hartw. 24. 1839; 67. 1840.
? Martens & Galeotti in Bull. Acad. Brux. 11!: 370. 1844.
18 Walpers, Rep. Bot. Syst. 6: 508. 1846-47.
^ Weddell, Chlor. And. 2: 74. 1859.
1933]
ALLEN—THE GENUS HALENIA 123
His argument for combining the two genera was based on the
fact that the two characters, the bilocular ovary and the short
spurs do not always coincide. In this work, he described five
new South American species based on collections of Triana,
Funck & Schlim, Goudot, Purdie, etc.
At the time of the publication of Bentham and Hooker's
‘Genera Plantarum,’ 1876, there were about twenty-five rec-
ognized species of Halenia from the whole world.
Hemsley, in ‘Biologia Centrali-Americana,'5* which appeared
in 1882, referred the Exadenus Martens and Galeotti to Halenia.
Gilg in Engler and Prantl’s ‘Natiirlichen Pflanzenfamilien '?
recognized the validity of the generic name Halenia and included
a careful description and good illustrations.
Sessé & Mocifio's 'Flora* contains a description of Swertia
cucullata which has escaped the notice of succeeding monog-
raphers, and has not been transferred to Halenia. A photo-
graph of the type specimen kindly furnished by Dr. E. Balguerias,
Curator of the Botanical Garden at Madrid, reveals the fact
that it is without question Halenia brevicornis Griseb., though
it is rather difficult to determine whether or not it is the species
proper or one of its many forms.
From this time until the monographic treatment of South
American species was published by Gilg!? in 1916, several species
and varieties of Mexican and Central American Halenia were
published. Halenia Rothrockit Gray,?° now Halenia recurva, in
1876 was based on plants collected in Arizona by Dr. J. T.
Rothrock. G. Don, in 1838, in the ‘General History of the
Dichlamydeous Plants’?! included a description of the existing
species of Halenia, making several new combinations from the
species of Humboldt, Bonpland and Kunth.? Some of his species
were taken from the manuscript of D. Don, thus appearing in
% Bentham & Hooker, Gen. Pl. 2: 817. 1876.
16 Hemsley, Biol. Cent.-Am. Bot. 2: 351. 1882.
17 Gilg in Engler & Prantl, Nat. Pflanzenfam. 42: 89. 1895.
18 Sessé & Mocifio, Flora Mexicana, 73. 1894.
19 Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, pp. 93-122. 1916.
20 Gray in Proc. Am. Acad. 11: 84. 1876.
21 Don, G. Gen. Hist. 4: 177. 1838.
2 Humboldt, Bonpland & Kunth, Nov. Gen. & Sp. Pl. 3: 174. 1818.
[Vor. 20
124 ANNALS OF THE MISSOURI BOTANICAL GARDEN
publication for the first time. Kuntze,” in 1891, revived the
generic name T'eiragonanthus Gmelin. He has been followed by
a few later taxonomists, among whom are Britton, Small,”
Rydberg,” and others. However, Halenia was placed on the list
of nomina conservanda, and thus must be regarded as the correct
name in accordance with the International Rules of Botanical
Nomenclature. Halenia crassiuscula Robinson and Seaton?’ ap-
peared in 1893. Britton,?* in 1894, reduced H. Brentoniana
Grisebach to a variety of Tetragonanthus deflexus, while in 1899
Fernald?* placed H. heterantha Grisebach under Halenia deflexa
var. heterantha. Other new species and new combinations ap-
peared in rapid succession, due to the extensive collecting done
during the period from 1890 to about 1920, by Conzatti, Pringle,
Purpus, and others in Mexico, and Weberbauer in South America.
Among these newly published species were: Halenia candida
Ramirez,*° 1895; H. chlorantha Greenman,* 1905; H. bella, H.
caespitosa Gilg, 1906; H. Conzattii Greenman,* 1912; H. guate-
malensis Loesener, and H. plantaginea var. latifolia Loesener*
(now H. guatemalensis var. latifolia (Loesener) Allen) 1913.
The type species up to this time had been called H. sibirica,
the name given it by Börckhausen. However, according to
the rules of priority, sibirica should give way to the older name
corniculata used by Linnaeus. Accordingly, Druce% in 1914
revived corniculata which is at present the accepted name for
the type species of Halenia. Britton and Brown in the 'Illus-
trated Flora, reduced Grisebach’s species Brentoniana and
heterantha to varieties of deflexa.
* Kuntze, Rev. Gen. Pl. 2: 431. 1891.
^ Britton, Manual, 734. 1901.
*5 Small, Fl. Southeastern U. S. 931. 1913.
* Rydberg, Fl. Rocky Mts. 666. 1922.
*7 Robinson & Seaton in Proc. Am. Acad. 28: 113. 1893.
*8 Britton in Mem. Torrey Bot. Club 5: 261. 1894.
?? Fernald in Rhodora 1: 37.
3 Ramirez in Inform. Secret. Foment. Mexic. "ed Mont. Ajusco). 34. 1895.
3l Greenman in Proc. Am. Acad. 41: 240. 1
2 Gilg in Fedde, Rep. Spec. Nov. 2: 52. D.
% Greenman in Publ. Field Mus. Bot. 2: 335. 1912.
^ Loesener in Verh. Bot. Ver. Brandenb. 55: 182. 1913.
% Druce in Rept. Bot. Exch. Cl. Brit. Isles 3: 419. 1914.
* Britton & Brown, Ill. Fl. 3:15. 1913.
1933]
ALLEN—THE GENUS HALENIA 125
Gilg,?" in 1916, monographed the South American representa-
tives of the genus, adding several new species. At that time, the
herbarium at Berlin contained as complete a series of South
American plants as were to be found any where, but, in some cases
at least, the material was too inadequate to determine the limits
of variation of certain of the species proposed. Hence, some of
the species recognized, studied in the light of subsequent collec-
tions, have fallen into synonymy. Gilg created three main
divisions:
A. Nectaria parva vel obsoleta, rarius extrinsecus breviter
semigloboso-prominentia. Folia manifeste carnoso-sub-
coriacea.
Nectaria haud calcariformia, sed extrinsecus ad basin
corollae alte globoso- vel coniformi-prominentia. Folia
semper tenuiter herbacea.
C. Nectaria extrinsecus calcaria manifeste evoluta formantia.
Division A contains the largest known Halenia, native of South
America only, which in inflorescence, flower, and leaf habit,
shows affinities with the western species of Swertia and Frasera
as well. Plants in division A are very distinct and occur in
Colombia and Venezuela. These Gilg apparently considered
the most primitive.
In division B we find the brevicornis-parviflora complex. Gilg
considered H. brevicornis (HBK.) Don a valid species known only
from South America, and parviflora a native of Mexico; he fur-
ther described a new species erythraeoides from Venezuela which
agrees in every respect with the parviflora type from Mexico.
Division C is subdivided according to the length of spurs; here
again, confusion has occurred, as is apt to be the case when
herbarium material is scanty and field work impossible. From a
limited experience in the field with a North American species,
the author can state with conviction that it is possible for two
plants belonging to the same species, growing side by side, to
vary not only in form but in size as well. It has been found also
that the spur character is inconstant. Axillary flowers and those
blooming late in the season frequently possess shorter spurs or
no spurs at all. If this character is so variable in one species, it
3 Gilg in Engl. Bot Jahrb. 54: Beibl. 118, p. 98. 1916.
w
[Vor. 20
126 ANNALS OF THE MISSOURI BOTANICAL GARDEN
is reasonable to suppose that the same situation may obtain in
other species. Only close attention to ecological detail and a
wealth of material not yet available from South America can
enable the monographer to delimit species from these little-
known regions even with a fair degree of accuracy.
Briquet,?* in 1931, described several new species which will be
treated below.
Gross MORPHOLOGY
Habit.—The American species of the genus Halenia are gla-
brous, strictly herbaceous, or somewhat ligneous annuals or per-
ennials. They may be of caespitose habit, as is illustrated by H.
caespitosa, or coarse, fleshy, foliose plants with single stems, 5 or
more dm. high, as in H. hygrophila and related species, or scapose
with the basal leaves disposed in a rosette, as in H. plantaginea,
or slender, graceful, simple or branching plants 1.5-6 dm. high, as
in H. brevicornis and its varieties. As a whole, the genus is not
colorful, the flowers being various shades of yellow and yellow-
green, except in the northernmost species, which has purple
owers.
Hoots.— The root system of the North American species is
fibrous, with a persistent, slender tap-root, frequently more or less
woody in texture, though typically that of an annual or perennial
in cross-section. In the South American species the root is, for
the most part, ligneous and thick.
Stems.—The stems are simple or branched, usually erect,
though they may be decumbent, as in Halenia decumbens and H.
Weddelliana. They are of two types, mostly foliose as in H.
deflexa and Schiedeana, or scapose as in H. plantaginea. The
stems often continue underground for a short distance, sending
out erect, flowering stems at irregular intervals. These may be
angular or terete; if the former they are often slightly winged, due
to the decurrence of the leaves, and usually faintly striate.
Leaves.—The leaves are opposite and decussate, or infrequently
whorled as in H. verticillata, either entirely cauline or disposed in
a rosette, sessile or petiolate. The petioles exhibit the same
characteristics as the stem, the decurrence of the calyx-lobes
* Briquet in Candollea 4: 317. 1931.
1933]
ALLEN—THE GENUS HALENIA 127
frequently being apparent. Where both petiolate and sessile
leaves occur on the same plant, the basal leaves have petioles
often equal to or longer than the blade, and the petioles become
decreasingly shorter toward the summit of the stem, the upper-
most leaves being sessile. The leaves are entire, 1-, 3- or 5-
nerved, and range from ovate to obovate or spatulate, or
lanceolate to linear, varying from .5 to 8 or 10 cm. in length
and up to 6 em. in width. The leaves of the North American
species are, on the whole, thin and herbaceous, whereas in the
South American species they are frequently coarse, fleshy, or
coriaceous.
Inflorescence.—The inflorescence consists of terminal or axillary
cymes of varying density. The flowers occur on short pedicels
at the tip of a stem or branch in a compact cluster or head, as in
H. brevicornis, or they may be borne loosely on long pedicels at
the tip of a branch or at a node, as in H. Schiedeana. In the
Central American and the more primitive of the South American
species, the inflorescence is usually a spicate, racemose, or more
or less umbellate cyme. A pair of small, linear, foliaceous bracts
is usually present at the base of the inflorescence, often bearing
tiny, undeveloped buds in their axils. Frequently the bracts
approximate the leaves in size and structure, and in H. involucrata
they form an involucre almost entirely enveloping the inflores-
cence.
Calyx.—The calyx is persistent, foliaceous, 4-parted, and the
segments united only at the extreme base and arranged in pairs;
the inner pair represents an inner cycle, the second an outer cycle.
The segments may be caudate as in the type species, H. corni-
culata, or spatulate, obovate, elliptic, lanceolate-linear, or linear,
with intergradations, with obtuse, acute, acuminate, mucronate
or apiculate tips. The tips very rarely are reflexed, as in H.
Schiedeana. The length of the calyx varies from one-third to
nearly equal that of the corolla. The calyx-segments are from
1- to 3-nerved, often reticulately veined at the tip, and both
surfaces are often papillate. Squamellae are usually found on
the inner surface of the calyx-segments at their base (see fig. 1).
They vary in size, shape, number, and position on the lobes, and
[Vor. 20
128 ANNALS OF THE MISSOURI BOTANICAL GARDEN
are usually distinguishable only under the dissecting lens after
the herbarium specimens have been boiled. Squamellae occur
in many of the Contorti both on the calyx and the corolla-lobes.
Engler and Prantl? have referred to similar structures in several
genera of the Gentianaceae as discs.
util
Fig. 1. Types of squamellae found on the calyx in the genus Halenia.
Corolla.—The corolla is marcescent, campanulate, 4-lobed,
white, yellow, yellow-green, green, or purple, ranging in length
from 5 mm. to nearly 3 cm. The lobes are triangularly ovate or
obovate, acute, acuminate, apiculate, mucronate or obtuse, often
auriculate, with an entire, erose, or crisped margin, and fre-
quently papillate on both surfaces. The veining of the dex-
trorsely convolute lobes may or may not be reticulate. The
corolla-tube varies from one-fourth to three-fourths the length
of the entire corolla. At or near the base of the tube, opposite
each lobe, is a tubercle which may be merely a slight swelling in
the lower portion of the tube, as in H. brevicornis, or a definite
*? Gilg in Engler & Prantl, Nat. Pflanzenfam. 47: 89. 1895.
1933]
ALLEN—THE GENUS HALENIA 129
spur longer than the corolla, as in H. guatemalensis (pls. 8-11).
It frequently happens that the corolla lacks spurs entirely in the
axillary flowers or in those occurring late in the season, as, for
example, in H. deflexa and other species. Not only in Halenia
is this situation apparent, but it is recalled that in other normally
spurred plants, for example, Linaria canadensis, spurs are fre-
quently absent. "Therefore it has seemed advisable to discon-
tinue heterantha as a variety of H. deflexa, since the occurrence of
these spurless forms is more or less frequent within the genus.
Gilg noted that in certain species of Halenia, for example, H.
brevicornis, etc., the flowers on the main stalk are normally large
and wide open, while below, on the same stem, they are definitely
smaller, apparently not opening at all or else very slightly. The
structure is similar, with the exception of the absence of spurs.
The ovary in these abnormal flowers contains fewer seeds, and
the capsule is much reduced in size. This would indicate,
according to Gilg, a gradual reduction of chasmogamous flowers
which depend on insect pollination, to more or less cleistogamous
flowers. The shape of the spurs varies exceedingly, from slender
to very thick and broad, or from spreading and ascending to
pendulous and incurved. Intergrading forms are frequent. The
spurs are frequently conspicuously veined and apparently
glandular.
Stamens.—The stamens are equal in number to, and are borne
alternate with, the lobes of the corolla at or near the summit of
the tube. The filaments are adnate to the corolla, and the
anthers are often enfolded in the bud by the margins of the
corolla-lobe. The anthers are versatile, deltoid, ovate or oblong,
often mucronate and papillate. The filaments are usually linear,
but they may become more or less dilated and, rarely, papillate.
The pollen grains are constant for the genus, being more or less
tetrahedrally spherical, and having three pores.
Pistil.—The pistil is sessile. The stigma is usually sessile and
cleft to expose the two inner stigmatic surfaces; the lobes may be
truncate or ovate. The ovary is bicarpellate, the margins of each
carpel being infolded and becoming the region of attachment for
the numerous ovules.
Fruit.—The fruit is a flattened, lanceolate to lanceolate-
[Vor. 20
130 ANNALS OF THE MISSOURI BOTANICAL GARDEN
obovate capsule, frequently subfalcate, and usually exserted. It
is unilocular at maturity, though in the young state it appears to
be more or less two-celled. The fruit dehisces septicidally along
the inner surface of each locule tip.
Seed.—' The seeds vary in shape, being globose to ovoid or
elliptic, often flattened. The surface is reticulate in the majority
of the South American species and in the brevicornis complex,
except for var. latifolia. The remaining species show the surface
of the seeds to be minutely granular instead of reticulate. They
vary in size from 0.5 to 1 mm. in diameter, and in color from dull
greenish-brown and yellow-brown to dark, shiny brown, the latter
usually being typical of those with reticulate surface. The age of
the plant and the conditions attending its collection no doubt
have their influence on the color, size, and, to some extent, the
texture of the seed-coat.
FLoRAL ANATOMY
The major portion of the anatomical investigation of the
Gentianaceae, particularly the Menyanthoideae, has been con-
cerned with the stem and leaf structure. The most complete
anatomical data is found in Gilg’s treatment of the Gentianaceae
in ‘Die Natürlichen Pflanzenfamilien.^? Solereder’s ‘Systematic
Anatomy of the Dicotyledons'* gives very little additional in-
formation. Since that time more attention has been given to
floral morphology as a separate study. Stolt,? in 1921, made an
exhaustive cytological survey of the flowers of several genera,
among which was included Halenia elliptica, an Asiatic species
having affinities with Halenia deflexa of North America. Inci-
dentally, Stolt inserted a diagram of the transverse section
through the ovary, which indicates clearly the vascular system
of that portion of the flower.
In order to make the present monograph as complete as possible,
anatomical study was undertaken. Fresh flowers of Halenia
deflexa, a species with both spurred and spurless forms, were
obtained by the author in Vermont and preserved in 70 per cent
40 Gilg in Engler & Prantl, Nat. Pflanzenfam. 4?: 50. 1895.
“| Solereder, Systematic Anatomy of the Dicotyledons 1: 548-550. 1908.
* Stolt in K. Svensk. Vet.-Akad. Handl. 61"; 1-56. 1921.
1933]
ALLEN—THE GENUS HALENIA 131
alcohol. The pickled material was dehydrated and embedded
in paraffin following the butyl alcohol method outlined by
Zirkle, sectioned at 10 u, and stained with crystal violet and
erythrosin. The accompanying drawings (pl. 12) were made with
the aid of a “Promi” microscopic drawing and projecting appara-
tus. The xylem has been cross-hatched in order to differentiate
it from the other vascular elements.
Transverse sections of the spurred form of Halenia deflexa
show that the vascular system of the pedicel is an amphiphloic
siphonostele (pl. 12, fig. 1). Approaching the receptacle, the
stele enlarges and assumes a rhombic form (fig. 2). The de-
currence of the outer lobes of the calyx is apparent. Shortly
thereafter (fig. 3), the midribs (a) of the two outer calyx-lobes
leave the receptacular stele (r). They migrate outward, and
from either, two lateral traces (a’) are given off (fig. 4). At this
point, four lacunae (z) appear, prior to the severing of the calyx
from the receptacle. In the succeeding illustration (fig. 5) these
lacunae (z) merge into two crescent-shaped fissures, and the
midribs of the two inner calycine lobes (b) leave the stele.
In fig. 6 the calyx-tube is entirely free from the receptacle, and
in the axil of each potential lobe are visible 4-8 minute emergences
or squamellae (e). The stele has again assumed a more or less
rhombic shape, but from the 4 angles, traces (c) depart centri-
fugally, soon resolving into the midrib (n) and two laterals (c’)
destined to supply each of the 4 corolla-lobes (fig. 7).
The disruption of the residual vascular cylinder is continued,
and the 4 staminal traces (f) are fully differentiated. At this in-
terval the corolla-tube is virtually free from the receptacle.
In the succeeding illustration (fig. 8) lacunae appear at m, the
ovary at this point appearing bilocular. "The residual stele con-
sists of two roughly semi-circular masses with a concentration of
lignified elements (l) at either end. The calyx-lobes are free,
and sections near the tip of the reflexed spurs are found, the
origin of which is to be described.
In the following illustration (fig. 9) the origin of the spurs is
apparent, and a cleft (y) is visible, indicating the sinus. Simul-
taneously, 4 protrusions occur on the inner surface of the corolla,
55 Zirkle in Science, N. S. 71: 103. 1930.
[Vor. 20
132 ANNALS OF THE MISSOURI BOTANICAL GARDEN
below the sinus, preparatory to the severing of the staminal
filaments (f). The ventral traces (1) and dorsal (k) are dis-
cernible. The ovules (0) are evident, showing their position in
the axils of the placentae (p).
In the final stage, taken from a section through the tip of the
flower (fig. 10), the differentiation of the corolla-lobes and the
isolation of the staminal filaments (f) are complete. The pla-
centae (p) have diminished in size.
The spurless form shows a similar vascular system except for
the absence of spurs, the presence of fewer ovules, and finally,
a more pronounced dorsal trace (k), after the cessation of ovular
production.
GEOGRAPHICAL DISTRIBUTION
The accompanying maps show three centers of distribution of
the species of Halenia in America. The first (figs. 2, 4) extends
from Labrador and Newfoundland, south to New York and west
to British Columbia and Montana. The one species and its
variety found in this area grow in moist or dry situations in
calcareous, slaty, or alluvial soil, in open woods or fields, on
stream banks or along the sea-shore, usually in the shade. The
habit varies with the habitat. This distribution follows closely
the northern region of glaciation, and coincides with the usual
distribution areas of herbaceous species common in that territory.
This same species has been collected three times in the State of
Mexico, but has not been reported from the intervening region,
a fact that might suggest a previously more continuous distribu-
tion of the species from the northern Rockies along the mountain
ranges to the Mexican Sierras.
The second area comprises a region extending from the Chiri-
cahua Mountains of New Mexico and Arizona, southward to
Costa Rica in Central America. Here are a few wide-spread
mountain species; but for the most part, they are endemics
occurring in volcanic areas (fig. 3).
The third large center of distribution is the northern part of
South America, where the genus is represented by a relatively
large number of endemic species (fig. 5).
All of the North and Central American species, with the
Ld ee d l3
Fig. 2. Map of southern Canada and northern United States, showing the geographical distribution of the species of Halenia: 1 = H. defleza; 2 = H. deflexa var. Brentoniana.
14. H. brevicornis
Fig. 3. Map of southern United States, Mexico, and Central America showing the geographical distribution of Halenia.
16. . var. multiflora 4. H. crassiuscula 5. H. Palmeri. 3. H. recurva 24. H. Shannonit
20. var. ovata 13. H. decumbens 8. H. plantaginea 26. H. rhyacophila 23. f. compacta
21. var. Tuerckheimii 25. H. guatemalensis 9. f. grandiflora 28. var. procumbens
29. H. caleoides 30. var. latifolia 22. H. platyphylla 27. var. macropoda
6. H. Conzaitii 12. H. nudicaulis 10. H. Pringlei 11. H. Schiedeana
1933]
ALLEN—THE GENUS HALENIA
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Maps of North and South America, showing the principal areas of distribution of Halenia.
by permission of the George F. Cram Company).
exception of Halenia alata and H. brevicornis and its varieties, are
spurred forms which show a definite relationship to one another.
Those from South America are of two types: spurred type,
similar to the North American species (excluding brevicornis and
[Vor. 20
134 ANNALS OF THE MISSOURI BOTANICAL GARDEN
alata) and showing affinities with them; non-spurred type, with
tubercles instead of spurs and quite different in habit from the
brevicornis and alata. Halenia brevicornis and its varieties form
a connecting link between the second and third areas, as well as
a morphological link, so to speak, between the primitive Swertiella
and the more advanced Haleniastrum. The varieties of brevi-
cornis do not extend further south than Guatemala, though the
species proper is found in South America. Here again, as in
Mexico and Central America, exist endemies which show a
development almost parallel with that which has occurred north
of the equator. An example of this is clearly shown by Halenia
decumbens, from the mountains of Mexico, which bears a striking
resemblance to H. Weddelliana, a species from Ecuador, Colombia,
and Peru.
SYSTEMATIC POSITION
The Gentianaceae, as pointed out by Gilg,“ shows closer
affinity to the Loganiaceae than to any other family in the
Contorti, yet the differences are so well marked that the two
families are never confused.
Grisebach separates the Gentianaceae into two subfamilies, the
Gentianoideae and the Menyanthoideae. In the first subfamily
the leaves are always simple, entire, sessile, and never al-
ternate. The aestivation of the corolla is never valvate. The
second subfamily, the Menyanthoideae, has alternate and mostly
petiolate, sometimes trifoliate, leaves, and the aestivation is
induplicate-valvate. Halenia belongs to the first division. The
structures called squamellae, described early in the text, are
never found in the Menyanthoideae but may occur in other
genera of the Gentianaceae, at the base of the corolla or calyx-
lobes.
The nearest relative of Halenia is Swertia. The most primitive
forms of Halenia, particularly the South American species, are
often confused with Swertia. The primitive members of Halenia,
instead of the definitely spurred corolla typical of the majority
of the species of the genus, possess small, knob-like protuberances
or nectaries which upon casual examination might pass unnoticed
“ Gilg in Engler & Prantl, Nat. Pflanzenfam. 42: 50. 1895.
1933]
ALLEN—THE GENUS HALENIA 135
or be taken for the nectaries which are structurally distinct and
characteristic of Swertia. More detailed study reveals the fact
that the depressions or spurs of Halenia are without the marginal
fringe which is always conspicuous in Swertia (fig. 6). Halenia
never has a corona, but in Swertia it is frequently present. The
corolla-lobes of Halenia are dextrorsely convolute in the bud,
whereas the reverse is the case in Swertia. Anatomical study
discloses other generic differences.
a b C d
Fig. 6. Swertia perennis L.: a, bud; b, interior surface of petal. Halenia brevicornis
(HBK.) Don: c, bud; d, interior surface of petal.
Within the genus itself two rather distinct sections are ap-
parent. The more primitive, which was mentioned above as
being near Swertia and possibly originating from it, may be called
tentatively Swertiella. It contains those species from South
America which were considered by Gilg as being most primitive.
Halenia alata of Mexico also belongs to this section. The Swer-
tiellae may be characterized by the absence of spurs or the pres-
ence of small tubercles or prominences at the base of the corolla,
which are not visible usually from the outside. They are coarse,
fleshy, foliose plants, and for the most part with dense inflores-
cences. The second section, Haleniastrum, is more advanced
and is readily recognized by the presence of spurs of varying
[Vor. 20
136 ANNALS OF THE MISSOURI BOTANICAL GARDEN
length and shape. Both herbaceous and more or less woody
types comprise this section.
An interesting discussion of the phylogeny of the Gentianaceae
was presented by T. H. Huxley* before the Linnaean Society of
London, April 7, 1887. Confining his study to the structure of
the corolla, he separated the family into two groups; the first
division Permelitae was segregated mainly on the presence of à
series of nectarial cells on the inner surface of the cup. The
Permelitae were again divided into four groups with various
BREVIGORNIS
— DASYANTRA xe du HÉRZOGII mig WERERBAUERI
mapa — T anarTLA ELATA TERRIEN.. Tien €—
umir M 98 EA ps LE
CURANDA sso KILLIPII
HYPOTHETICAL ANCESTOR
Fig. 7. Chart showing the probable phylogenetic relationship of species of Halenia.
modifications of floral structure. The first, Actinanthe, which he
considered the most primitive and the least differentiated, con-
tained among other elements, Exadenus. The latter was in-
cluded as a transitional stage on account of its rudimentary spurs,
leading to the second division, the Keratanthe, containing only
one genus Halenia, which represents an extreme modification of
the Exadenus type. The third related division, the Lophanthe,
5 Huxley, T. H. Jour. Linn. Soc. Bot. 24: 101. 1887.
1933]
ALLEN—THE GENUS HALENIA 137
depicts another outgrowth of the Actinanthe type, but with
filamentous appendages or fimbriae. Swertia and Frasera fall
into this category.
Text-figure 7 shows the probable relationship of the various
species of Halenia, according to the author's interpretation.
For the sake of convenience, in the taxonomic treatment of
the genus, the North and South American species are taken up
separately in each section.
ABBREVIATIONS
In the citations of specimens examined the following abbrevia-
tions have been used to denote the various herbaria from which
specimens were used for study.
ANSP = Herbarium of the Academy of Natural Sciences of Philadelphia.
B n.
G = Botanical Garden, Berlin
BB = Brooklyn Botanical Garden.
BM - British Museum of Natural History, London.
B = Herbarium of the Botanical Garden, B ls.
C — Herbarium of the En of Chicago, deposited in the Field Museum
of Natural Histo
CAS = Herbarium of the California Academy of Sciences.
D = Dudley Herbarium of ^r burns Jr. University.
DH = Delessert Herbarium of Gen
F = Herbarium of the Field Musei of Natural History.
G = Gray Herbarium of Harvard University.
HP = Herbarium of H. Pittier, Director de Museo Comercial, Caracas, Venezuela.
HJP - Herbarium of the Jardin des Plantes, Paris
IAC = Herbarium of the Iowa Agricultural Collage.
K = Herbarium of the Royal Botanic Gardens, Kew
L = Linnaean Herbarium, Linnaean Society of cm London.
M = Herbarium of the Missouri Botanical Mera
MU = Herbarium of the University of Missour
NY = Herbarium of the New York Botanical rte
P — Parry Herbarium deposited at the Iowa Rosipulturel College.
S = Herbarium of the Botanical Museum, Stockholm.
SM = Herbarium of the State Museum, Albany, New York.
UC = Herbarium of the Botanical Mri of the University of Copenhagen.
US — United States National Herbariu
i = Herbarium of the Natural History ‘Museum, Vienna.
TAXONOMY
Halenia Bórckh. in Roemer, Arkiv für Botanik 1!: 25. 1796;
Ruiz & Pavon, Fl. Peruv. 3: 21, pl. 242, fig. 1. 1802; Endl. Gen.
Pl. 601. 1836-40; Grisebach, Obs. Gent. 36. 1836; G. Don, Gen.
[Vor. 20
138 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Hist. 4: 177. 1838; Grisebach, Gen. & Sp. Gent. 322. 1839;
Dietrich, Syn. Pl. 2: 918. 1840; Hooker, Fl. Bor.-Am. 2: 67,
pl. 155-6. 1840; Grisebach in DC. Prodr. 9: 128. 1845;
Grisebach in Linnaea 22: 45. 1849; Weddell, Chlor. And. 2:
74. 1859; Benth. & Hooker, Gen. Pl. 2: 817. 1876; Hemsl.
Biol. Cent.-Am. Bot. 2:351. 1882; Baillon, Hist. Pl. 10:142. 1891;
Gilg in Engler & Prantl, Nat. Pflanzenfam. 42: 89. 1895;
Conzatti, Fl. Syn. Mexico, 174. 1897; Rouy, Ill. Pl. Eur. 17: pl.
412. 1902; Gilg in Fedde, Rep. Spec. Nov. 2:52. 1906; Robinson
in Gray's Manual, ed. 7. 659. 1908; Britton & Brown, Ill. Fl. 3:
15, fig. 3365. 1913; Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 93.
1916; Johnson, Tax. Fl. Pl. 488. fig. 340. 1931; Briquet in
Candollea 4: 317. 1931; Rydberg, Fl. Cent. N. Am. 636. 1982.
Tetragonanthus Gmelin, Fl. Sib. 4: 114, pl. 53. 1769; Kuntze,
Rev. Gen. Pl. 2: 431. 1891; Britton, Manual, 734. 1901;
Small, Fl. Southeastern U. S. 931. 1913; Rydberg, Fl. Rocky
Mts. 666. 1922.
Swertia Linnaeus, Amoen. Acad. 2: 344. 1751; Jussieu, Gen.
Pl. 158. 1791; Michaux, Fl. Bor.-Am. 1: 97. 1803; Humboldt,
Bonpland & Kunth, Nov. Gen. & Sp. Pl. 3: 174. 1818; Kunth,
Syn. Pl. 2: 266. 1823; Schlectendal & Chamisso in Linnaea 5:
122. 1830.
Ceratia Persoon, Syn. Pl. 1: 287. 1805; Hedw. Gen. 181.
1806.
Exadenus Grisebach, Obs. Gent. 36. 1836; Gen. & Sp. Gent.
322. 1839; in DC. Prodr. 9: 128. 1845.
Glabrous caulescent annual, biennial, or perennial herbs.
Root fibrous, varying from typically herbaceous to extremely
ligneous. Leaves membranaceous or fleshy, opposite, entire,
sessile or petiolate, usually 3-5-veined, veins frequently sub-
merged in fleshy type. Inflorescence a terminal or axillary,
subumbellate, or rarely racemose or spicate cyme. Calyx 4-
parted, foliaceous, linear, lanceolate, ovate or spatulate, usually
papillate, often bearing small squamellae at the base of each lobe.
Corolla 4-parted, white, yellow, green, or purple, marcescent,
campanulate, the tube of varying length; the lobes dextrorsely
convolute, elliptic to ovate, obtuse to acute or apiculate, entire,
crisped or erose, frequently auriculate and papillate. Stamens
1933]
ALLEN—THE GENUS HALENIA 139
4, included, adnate to the corolla-tube at varying heights, and
alternate with the corolla-lobes; filaments linear, occasionally
somewhat dilated; anthers 2-celled, ovate, oblong or subtriangu-
lar, versatile. Carpels 2, sessile, the edges being infolded to
form a parietal placenta bearing many ovules; stigma sessile,
composed of two oblong or ovate lobes, the inner surfaces of
which are stigmatic. Fruit a compressed capsule, lanceolate to
ovate, often subfalcate, septicidially dehiscent from the tip.
Seeds globose or slightly flattened, brown or greenish tan,
granular or reticulate.
Type species: H. corniculata (L.) Druce in Rept. Bot. Exch.
Cl. Brit. Isles 3: 419. 1914.
SYNOPSIS OF THE SECTIONS OF THE GENUS
Plants usually coarse with fleshy leaves, rarely slender with thin, herbaceous
leaves; stem usually leafy; spurs absent, or present as very small inconspicu-
ous ida frequently obscured by calyx; distribution chiefly Sout
esie eer errr eT CORE D 1. Swertiella
Planta usually slender, with thin, herbaceous leaves; om leafy or unio
spurs present; distribution North and South America.......... 2. Haleniastrum
SECTION 1. SWERTIELLA
KEY TO NORTH AMERICAN SPECIES AND VARIETIES
1. Leaves mostly radical; stem more or less BOBRDOSÓE ID sis d ns seruo OS 1. H. alata
1. Leaves mostly cauline; stem not scapose.
2. Leaves ovate, less than 1.2 cm. long......... 2f. H. brevicornis var. ovata
2. Leaves not ovate, longer than 1.2 cm.
3. Pedicels filiform, slender, elongate; habit decidedly spreading.......
VE OP RO Bik "ke win fe RA n GE g. H. brevicornis var. Tuerckheimii
3. Pedicels stouter than filiform, ——
4. Corolla without distinct spur
5. Inflorescence strict................ 2c. H. brevicornis var. micranthella
5. Inflorescence compact.
6. Leaves linear, slender... 60 Lese cecil 2. H. brevicornis
6. Leaves ovate to lanceolate, coarse....2a. H. brevicornis var. latifolia
4. Corolla with small, but distinct spurs.
5. Spurs thick, conical, more or less pendulous...................
n a BS Rs. 6 96% 4 so X o oc 2b. H. brevicornis var. multiflora
5. Spurs blunt, spreading, hog .2e. H. brevicornis var. chihuahuensis
5. Spurs slender, divergent. ............. 9d. H. brevicornis var. divergens
1. H. alata (Mart. & Gal.) Hemsl. Biol. Cent.-Am. Bot. 2:
351. 1882.
Exadenus alatus Mart. & Gal. Bull. Acad. Brux. 11!: 372.
1844; Walper's Rep. Bot. Syst. 6: 508. 1846-7.
[Vor. 20
140 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Tetragonanthus alatus Kuntze, Rev. Gen. Pl. 2: 431. 1891.
Small perennial with 1-2 simple erect slightly winged stems,
about 0.5-1 dm. high; numerous radical leaves, up to 3.5 em.
long and .4 em. broad, crowded, attenuate into long petioles,
oblanceolate, obtuse, 3-nerved; lower cauline leaves almost twice
the length of the radical, subsessile; upper linear, obtuse, sessile;
inflorescence consisting of a few (4-6) terminal flowers on 4-
winged pedicels; calyx foliaceous, slightly shorter than corolla,
segments oblong, 3-nerved, papillate, acute; corolla yellow,
subrotate, up to .6 em. long, tube about one-half as long as the
entire corolla; corolla-lobes ovate, obtuse; filaments linear;
anthers ovate; capsule broadly ovate; seeds yellow-brown,
globose, granular.
Distribution: rocky forests of southern Mexico.
Specimens examined:
Vera Cruz: in forests and on trachytic rocks on Mt. Orizaba, alt. 2250-3000 m.,
June—Oct. 1840, Galeotti 7221 (BG, B TYPE, DH, K, V); Mt. Orizaba, alt. 3000-3125
m., Aug. 1838, Linden 934 (DH, K).
2. H. brevicornis (HBK.) G. Don, Gen. Hist. 4: 177. 1838;
Wedd. Chlor. And. 2: 77. 1859; Hemsl. Biol. Cent.-Am. Bot.
2: 352. 1882; Gilg in Engler & Prantl, Nat. Pflanzenfam. 4?:
89. 1895.
Swertia brevicornis HBK. Nov. Gen. & Sp. Pl. 3: 174. 1818.
S. parviflora HBK. Nov. Gen. & Sp. Pl. 3: 174. 1818.
S. parviflora var. « angustifolia Sch. & Cham. in Linnaea 5: 122.
1830.
S. cucullata Sessé & Mociño, Fl. Mex. 79. 1894.
Halenia parviflora G. Don, Gen. Hist. 4: 177. 1838; Wedd
Chlor. And. 2: 77. 1859; Hemsl. Biol. Cent.-Am. Bot. 2: 352.
1882.
H. paucifolia Hemsl. Biol. Cent.-Am. Bot. 2: 352. 1882.
H. erythraeoides Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p.
105. 1916.
Exadenus parviflorus Griseb. Gen. & Sp. Gent. 322. 1839.
E. brevicornis Griseb. l. c. 323.
E. paucifolius Mart. & Gal. Bull. Acad. Brux. 11!: 372. 1844.
Tetragonanthus paucifolius Kuntze, Rev. Gen. Pl. 2: 431. 1891.
T. parviflorus Kuntze, l. c.
T. brevicornis Kuntze, l. c.
1933]
ALLEN—THE GENUS HALENIA 141
Annual, 1.5-4.5 dm. high; root slender; stems erect, slender,
usually simple below and branched above, sometimes branched
from the base, slightly angled, and striate; leaves sessile, sub-
connate, lower linear to lanceolate, up to 3 cm. long, .4 cm.
broad, usually prominently uninerviate, upper shorter, narrower,
up to 2 em. long, linear; inflorescence compact cymose clusters,
pedicels erect, 1.8 em. or less in length; calyx-segments lanceolate,
approximately one-half to two-thirds the length of the corolla,
usually 3-nerved; corolla .4—.8 cm. long, tube one-half or slightly
more the length of the entire corolla; corolla-lobes broadly to
narrowly ovate, acute, more or less auriculate; spurs merely
small depressions near the base of the corolla-tube, usually not
visible to the naked eye; stamens about .2 cm. long, attached
usually near the middle of the tube; filaments linear, anthers
usually deltoid; ovary lanceolate, about .5 cm. long; seeds
minute, ovoid to subglobose, brownish, finely reticulate.
Distribution: isa in mountains, from Mexico, southward to Peru.
Specimens exami
Mexico:
San Luis Porosi: Oct. 1879, Schaffner 421 (BG, BM, F, NY, UC, US); region of
San Luis Potosi, alt. 1500-2000 m., 1878, Parry & Palmer 600 (ANSP, BM, F, G,
IAC, K, M, NY, US)
VERA Cruz: Mt. Orizaba, alt. 1875 m., 1841-43, Liebmann 10775 (UC); same
locality and date, alt. 2500 m., Liebmann 10778 (UC); same locality, 1840, Galeotti
7219 (B TYPE of Exadenus paucifolius, DH); “in dumetis prope Jalapam," 1830,
Schiede & Deppe 247a (BG TYPE of Swertia parviflora var. « angustifolia, BM, M)
GUANAJUATO: near city Guanajuato, alt. 1650 m., Humboldt & Bonpland (BG
TYPE not seen, HJP, M photo.).
PUEBLA: vicinity of Puebla, ravines, Hacienda Alamos, route Vera Cruz, alt. 2170
m., Nov. 5, 1907, Arsène 2098 (G, M, US); mountains, Esperanza, Aug. 1907, Purpus
2697 (BG, BM, CAS, F, M, US); Chinantha, alt. 1750-2000 m., May 1841, Liebmann
10776 NY, UC).
Mexico: La Cima, alt. 2500 m., | alt. 1500 m., Aug. 1904, Kuntze 23783
(NY); Volean de Toluca, Heller 401
tee 1864-70, Ghiesbreght 137 He K, NY); Ghiesbreght 618 (G, K, M).
XICO WITHOUT LOCALITY: Tate (DH); Oct. 8, 1897, Berlandier 1207 ex 13 (DH);
Berlandier 1207 (V); Pavon (DH); Wawra 425 (V).
CENTRAL AMERICA:
CosrA Rica: *in monte Reventado,” alt. 2250 m., coll. of 1847, Oersted 10779
(UC)
NicanAGUA: El Viejo, Oersted (M).
2a. H. brevicornis var. latifolia (Sch. & Cham.) Allen, n.
comb.
[Vo 20
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Swertia parviflora var. 8 latifolia Sch. & Cham. in Linnaea 5:
122.
Ezadenus parviflorus var. 6 latifolius Griseb. Gen. & Sp. Gent.
322. 1839; in DC. Prodr. 9: 128. 1845
Halenia parsiftera var. latifolia Herrsl. Biol. Cent.-Am. Bot.
2:351. 1882.
A more sturdy plant than the species, attaining a height of 6
dm.; leaves larger than species, 1.5-4 em. long, .3-1.5 cm. broad,
acute or obtuse, lowermost smaller, ovate, rotund, and with
longer petioles; inflorescence frequently less compact than that
of species and with longer pedicels; calyx-segments lanceolate,
foliaceous, one-half to two-thirds the length of the corolla,
papillate, 3-nerved; corolla .6-.8 em. long, tube about equalling
the ovate, frequently papillate, auriculate lobes; spurs short,
consisting of very slight angular protrusions froi the extreme
base of the corolla-tube, giving the corolla a square appearance.
Distribution: mountains of Mexico.
Vera Cruz: “in dumetis prope Jalapam,” 1830, Schiede & Deppe 247b (BG TYPE
of Swertia parviflora var. 8 latifolia Sch. & Cham., BM, M, S, V); same locality, coll.
of 1833, Beyrich (S); region of Orizaba, Oct. 20, 1866, Powpesü 3126 (BG, B, K, 8,
UC, US).
PuxBLA: Manzanilla, alt. 2250 m., Nov. 24, 1908, — E 703 (B, US); Barrancas,
Hacienda Alamos, route Vera Oris, alt. 2170 m., Dec. 10, 1907, Arsène (M, US);
Boca del Monte, alt. 2300 m., Nov. 16-19, 1907, Anim fa
More tos: Lecima, Sierra de Ajusco, Aug. 18, 1896, Harehberger 13? (ANSP, G,
US
).
TLaxcaLa: Chiautempan, alt. 2250 m., Nov. 10, 1908, Arsène 1711 (B, US).
Mexico: Ajusco Mountains, 1905, Luton & Lemmon (CAS); Valley of Mexico,
ate Le Ixtaccihuatl, Jan. 1903, Purpus (CAS); due de Agua, Nov. 1
Purpus 1762 (CAS, F, G, M, US); Sierra de las Cruces, alt. 2375 m. , Sept. 12, 1904,
Pringle 13120 (BG, F, G, K, 8, UC, US); Eslava, Salazar (US); near Salazar, Sept.
14, 1903, Rose & fee 7026 (US); Désierto Viejo, near Mexico, Sept. 6, 1865,
Bourgeau 799 (BG, B, DH, G, K, NY, 8, UC, US).
MICHOACAN: vicinity of Morelia, alt. 2500 m., Oct. 26, 1911, Arsène 5610 (M, US).
Mexico witHout Locauity: Berlandier 1207 (DH); 1832, Alaman (DH).
2b. H. brevicornis var. multiflora (Benth.) Allen, n. comb.
Halenia multiflora Benth. Pl. Hartw. 24. 1839
Plant more sturdy than the species, frequently branched
above, rarely branched at the base, 4.5 dm. or less high; leaves up
to 1.3-2.5 em. long, .2-.9 em. broad, obtuse or acute, ovate to
1933]
ALLEN—THE GENUS HALENIA 143
narrowly lanceolate, frequently faintly 3-nerved, midvein always
prominent; inflorescence usually densely flowered, though com-
pactness of arrangement varies; mature corolla with small thick
rounded pendant spurs at its base.
Distribution: in «oi ERN of Mexico.
Specimens examin
San Luis Porosi: in posta baie of San Miquelito, Aug. 1877, Schaffner 88 (G);
Alvarez, Sept. 28-Oct. 3, 1902, Palmer 160 (CAS, F, G, M, NY, UC, US); Parry &
Palmer 600b (IAC, M).
ZACATECAS: on the Sierra de los Morones, near Plateado, Sept. 1, 1897, Rose 2732
US).
Janisco: ‘‘in pinetis Bolaños,” 1839, Hartweg 210 (BM, DH, G, K, NY, V);
Sierra minis: west of Bolafios, Sept. 15-17, 1897, Rose 2962 (US); banks of ravines
near Guadalajara, alt. 1250 m., Oct. 21, 1903, Pringle 11636 (BG, F, G, K, US);
hills near Guadalajara, Oct. 14, 1889, Pringle 2735 (C, IAC); Rio Blanco, Oct. 1886,
Palmer 680 (G, NY, US); same locality and date, Palmer 683 (G).
GUERRERO: between Ajusinapa and Petatlan, alt. 1250-1750 m., Dec. 14, 1894,
Nelson 2126 (US).
Oaxaca: Sierra de San Felipe, alt. 2000 m., Oct. 13, 1894, Smith 665a (M, NY, US);
same locality, alt. 1800 m., Aug. 15, 1898, Conzatti & Gonzalez 878 (G); San Pedro
Nolasco, alt. 1875 m., Oct. 1840, Galeotti 1490 (B, DH)
2c. H. brevicornis var. micranthella (Briq.) Allen, n. comb.
Halenia micranthella Briq. in Candollea 4: 320. 1931.
Plant 1.5-6 dm. high; leaves linear-lanceolate, often obtuse,
3-nerved, the lower long-petiolate, the upper linear, sessile;
inflorescence usually less compact, but more strict, than in the
species, and the stem and pedicels more erect; the nodes of the
inflorescence approximately equidistant, giving the appearance
of a narrow raceme; corolla campanulate, but more narrowed at
the base than in be species; calyx-segments usually one-half the
length of the corolla; corolla-lobes broadly ovate, acuminate,
auriculate; spurs reduced to minute depressions, frequently not
visible to the naked eye; anthers usually broadly ovate; filaments
varying, usually linear, rarely dilated.
Distribution: mountains of Mexico.
Specimens examined:
HipaLGo: El Chico, near Pachuca, Sept. 1905, Purpus 1761 (CAS, F, G, M, NY,
US); wet meadows, Sierra de Pachuca, alt. 2450 m., Aug. 13, 1898, Pringle 6964
(ANSP, BG, B, CAS, DH, F, G, IAC, K, M, NY, S, US, V); same locality, Sept. 8,
1899, Pringle 7943 (BG, F, G, K, M, NY).
Mexico: Sierra de a o Nov. 9, 1903, Pringle 11842 (BG, F, G, K, S, UC, US);
near Ozumba, alt. 2000 m., Nov. 3, 1902, Pringle 11329 (G, UC, US).
Souta Mexico wiht Locauity: July 1841, Liebmann 10777 (UC).
[Vor. 20
144 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Briquet based his new species on Pringle 6964. This is cited
in the original publication as 1964, but this is merely a typo-
graphical error. The Pringle specimens are about 12 cm. high.
The additional material cited above undoubtedly belongs to the
same species based on Pringle 6964, but for the most part it
consists of plants over 15 cm. in height.
It is possible that there are two distinct plants under Purpus
1761, but since the variation in the brevicornis complex, as a
whole, is so pronounced, these differences have been considered
as variations typical of the variety. Hence, all sheets of Purpus
1761 have been determined as var. micranthella.
2d. H. brevicornis var. divergens“ Allen, n. var.
Similar to var. multiflora but with more slender spurs which
diverge, making the corolla broader at the base than at the tip.
Distribution: central Mexico.
Specimens ex t
Mıcnoacan: Loma Sta. Maria, vicinity of Morelia, alt. 2000 m., Sept. 4, 1910,
Arsène 55 (F); vicinity of Morelia, near La Huerta, alt. 1950 m., Sept. 1, 1910,
Arsène (M TvPrE, US); Loma Sta. Maria, alt. 2050 m., Sept. 19, 1910, Arsène (M,
US); same locality, alt. 1950 m., Sept. 4, 1910, Arsène 5957 (K, M, S, US); same
locality, Oct. 28, 1910, Arsène 5864 (K, M, US)
Vera Cruz: Orizaba, 1853, Müller (NY).
Mexico wrrnovT Locauity: 1858, Sumichrast (DH).
2e. H. brevicornis var. chihuahuensis‘’ Allen, n. var
Similar to var. multiflora but with lower leaves always elliptic-
ovate, about 1 cm. long, increasing in length and acuteness and
decreasing in width as they approach the summit of the stem;
inflorescence more loosely arranged, and spurs more blunt but
not quite so pronounced as in multiflora; calyx-segments three-
fourths the length of the corolla, and usually narrower.
** H. brevicornis var. divergens Allen, var. sse intei a var. multiflora calcari-
bus tenuioribus divergentibus latioribus basi quam summo.—Mi1cHoACAN: vicinity of
Morelia, near La Huerta, alt. 1950 m., Sept. 1, 1910, Arsène (M TYPE,
*' H. brevicornis var. chihuahuensis Allen, var. nov.—Differt a var. multifor
folis inferioribus semper elliptico-ovatis ca. 1 cm. longis, prope summum caulis
longitudine augmentibus et latitudine deminuentibus; inflorescentia laxiora; Ld
bus obtusis; calycis segmentibus 34 corollae longitudini adaequantibus, plerumque
angustioribus.—CmrHuAHUA: pine plains, base of the Sierra Madre, Sept. 26, 1888,
Pringle 1664 (BG, BM, B, CAS, DH, M yer, S, V).
1933]
THE GENUS HALENIA 145
ALLEN
Distribution: mountains of Chihuahua, Mexico.
Specimens examine
CHIHUAHUA: pine plains, base of the Sierra Madre, Oct. 4, 1887, Pringle 1830
(ANSP, CAS, F, G, K, NY, US); same locality, Sept. 26, 1888, Pringle 1664 (BG,
BM, B, CAS, DH, M ryps, S, V); mesa, west of Hop Valley, Sierra Madre Mountains,
alt. 1750 m., Sept. 17, 1903, Jones (BM, M, US); southwestern Chihuahua, Aug.-
Nov., 1885, Palmer 408 (ANSP, G, IAC, K, NY, US)
2f. H. brevicornis var. ovata‘? Allen, n. var.
Plant 5-6 dm. high; stems erect, simple below, bearing short
floriferous branches only at tip; leaves about 12 pairs, shortly
petiolate, frequently bearing buds in the axis; lower leaves
broadly ovate, usually with a prominent midvein, reticulate,
less than .5 cm. long, .3—4 em. broad, abruptly acuminate;
middle cauline leaves ovate, 1-1.2 cm. long, .7-.8 cm. broad,
acuminate; upper leaves up to 1.5 em. long, lanceolate; inflores-
cence and flowers similar to multiflora type, but spurs more slender,
slightly incurved, and divaricate.
Distribution: known only from type locality.
Specimens examined:
Mexico: Tepic, Jan. 5-Feb. 6, 1892 Palmer (US TYPE).
2g. H. brevicornis var. Tuerckheimii (Briq.) Allen, n. comb.
Halenia Tuerkheimii Briq. in Candollea 4: 317. 1931 (dedi-
cated to H. von Tuerckheim).
Slender graceful stem, loosely branching, up to 6 dm. high;
leaves lanceolate to lincar-lanedelata or elliptic, about 1 cm. long,
.5 em. broad, lower cauline leaves becoming more acute, faintly
3-nerved, midvein prominent; middle cauline leaves 3 cm. or
less long, .5 em. broad; inflorescence in loose, terminal or axillary,
few-flowered cymes, each flower borne on a long slender attenuate
pedicel; first flowers with very small, though definitely formed,
spurs, at the base of the corolla-tube; later and usually axillary
owers of more slender habit and without spurs.
48H. brevicornis var. ovata Allen, var. nov.—Planta 5-6 dm. "T caulibus
erectis, infra simplicibus modo summo ramos breves floriferos gerentibus; foliis ca.
12 geminis, breve petiolatis, saepe in axibus gemmas gerentibus; foliis inferioribus
late ovatis, medio-nervo plerumque prominenti, reticulatis, minusquam .5 em. longis,
3—.4 em. latis, abrupte Ead foliis mediis caulinis 1-1.2 cm. longis, .7—.8 em.
latis, ovatis, acuminatis; foliis superioribus usque ad 1.5 cm. longis, lanceolatis;
inflorescentia et floribus multiflorae similibus sed calcaribus tenuioribus, parvulum
incurvatis divaricatisque.—MExico: Tepic, Jan. 5-Feb. 6, 1892, Palmer (US TYPE).
[Vor. 20
146 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: known only from type locality.
Specimens examine
GUATEMALA: “Alta Verapes, Fichtenwülder bei San Joaquin," alt. 1000 m., Dec.
1907, von T'uerckheim 2041 (F, DH, G, NY, US, V).
From an examination of the types of Halenia brevicornis, H.
parviflora, H. multiflora, and Exadenus paucifolius, etc., it ap-
pears that this group presents a complex, all members of which
show variation in habit and spurs, which can not be considered
specific differences but differences of degree. Careful perusal
of the specimens available discloses the fact that nearly every
locality produces some variation in the species. Thus, Chihuahua
gives rise to a definitely spurred form, while Guatemalan material
presents an entirely different aspect habitally. Ecological ex-
periments, as well as field work, would doubtless prove highly
valuable as a supplement to the taxonomic treatment of this
complex. At present there is no distinction which warrants the
retention of these as distinct entities. To draw a line between
these forms mentioned and treat them specifically would com-
plicate further an already confused situation. Therefore it has
seemed advisable to draw attention to these differences, or
variations from the specific form, by relegating them to the
status of variety of form, with the note that this is merely an
arbitrary disposal, that intergradation is existent and that en-
vironment is in a large measure responsible for the variation
found within the species.
KEY TO THE SOUTH AMERICAN SPECIES
1. Leaves always thin, herbaceous; stem and branches more or less slender.
2. Plant slender; flowers in enthenls less than 1 em. long
3. Stem ueuadly branched above, sparingly leafy; baal rosette wee
boxed Spy ea a vidi Verc D XO Eier RrrDDCR Ce - breskoornte
3. Stem not branched above, leafy; leaves adpressed; a. ern
DUSenb.s o ioo direc Ra der yon DC rs aub DAS 3. H. adpressa
2. Plant coarse; flowers in anthesis usually more than 1 em. long.
3. Nodes 8 or less; leaves not apiculate...................... . H. macrantha
3. Nodes 12 and more; leaves apiculate..................... 5. H. Karstenii
1. Leaves more or less fleshy; stem and branches coars
2. Upper leaves of inflorescence subinvolucrate and buds rs —
H
2. Upper leaves of inflorescence not subinvolucrate.
3. Rosette usually present; stem-leaves linear-lanceolate.
4. Plant usually less than 30 em. high; leaves not more than 5 em. long.
5. Calyx-lobes minutely hirtellous.
1933]
ALLEN—THE GENUS HALENIA 147
6. Plant usually less than 20 cm. high, or if more than 20 em. high,
H.
owers in axils of all caulineleaves................ 7. H. inaequalis
6. Plant 10-30 em. high; flowers in axils of upper leaves only..
GREG RS 6b UE POTETE sys 5 CRORE TREE 8. H. . viridis
5. Calyx-lobes not hirtellous.
6. Stem-leaves less than 3 pairs, linear................ 9. H. Schultzei
6. Stem-leaves more than 3 ud lanceolate to HART ERR
7. Cauline leaves less than 2 em. long............ 10. H. gentianoides
7. Cauline leaves more than 2. Ca, ONE. ees 11. H. stellarioides
4. Plant usually more than 30 cm. high; leaves more than 5 cm. long.
5. Inflorescence verticillate; leaves ternate............ 12. H. verticillata
5. Inflorescence not verticillate; leaves not ternate.
Nodes more gn Dialed ate i NOE ME 18. H. foliosa
6. Nodes less than 1
7. Root-stalk aia conspicuously larger than stem.
8. Flowers more than 1.5 cm. long, 8-10 mm. broad........
BR ss datdxpe we Pas uA V aaa eT ae 14. H. dasyantha
8. Flowers less than 1.3 cm. long, 5-7 mm. broad...... 16. H. elata
7. Root-stalk not swollen, scarcely larger than stem
8. Plant less than 30 em. high.................... 16. H. Tolimae
8. Plant more than 30 cm. high
9. porcine elongate, spicate; niim > con-
em ER TE RET TEE E ee . H. hygrophila
9. Inforseec spreading; calyx-venation Be bua
eT D 18. H. parallela
3. Rosette absent; pron es ovate to ovate-lanceolate........ 19. H. major
2. Halenia brevicornis (HBK.) G. Don, Gen. Hist. 4: 177.
1838
For synonymy and description, see North American species,
page 140 of this work.
Specimens inge.
SOUTH AME
VIRA Trujillo & Mérida, alt. 1300-4900 m., 1842, Linden 456 (DH).
CoroMnzia: Bogota, Dec. 19, 1853, Holton 19 (464) (NY); Dept. Cundinamarca, -
moist grassy loam, southwest of Sibate, alt. 2600-2800 m., Oct. 13-15, 1917, Pennell
2434 (M, NY, US); Dept. Antioquia, La Sierra, Medellin, alt. 2000 m., Jan. 4, 1931,
Archer 1348 (M); Dept. El Cauca, open banks near Rio Cauca, Coconuco to Popayan,
alt. 2000-2500 m., June 18, 1922, Killip 6889 (US); field, near Rio San Andreas,
Calaguala, Coconuco, alt. 2500-2800 m., June 18, 1922, Pennell 7154 (ANSP, G,
NY, US); field, north of Coconuco, alt. 2300-2500 m., June 11, 1922, Pennell &
Killip 6480 (ANSP).
Ecvapon: near Quito, coll. of 1859, Jameson (DH, G, NY, UC, V).
Peru: steep grassy slopes, Mito, alt. 3000 m., April 8-18, 1923, Macbride 3431
(F, M); 1840, Mathews 3133 (DH
Attention should be Salted to the fact that the Swertia brevi-
cornis of Humboldt, Bonpland & Kunth, which Gilg has relegated
[Vor. 20
148 ANNALS OF THE MISSOURI BOTANICAL GARDEN
to synonymy, is a very short much-branched specimen, differing
markedly in habit from the S. parviflora HBK. A close examina-
tion of the type of the former reveals the fact that the terminal
branch has been broken off. Subsequent lateral offshoots give
an entirely different appearance habitally. ‘This situation is
apparent also in the specimens collected by Oersted in Central
America, which Gilg in 1915 labeled H. parviflora.
3. H. adpressa*? Allen, n. sp.
Perennial, .5-2.5 dm. high; root coarse; stem usually solitary,
erect, slender; basal leaves in dense rosette, oblanceolate, 1-1.5
em. long, .3 em. broad; cauline leaves 5-6 pairs, sessile, lanceolate,
1-1.5 em. long, .2-.3 em. broad, acute; inflorescence a several-
flowered cyme, pedicels 2 cm. long; calyx-lobes lanceolate, .4—.6
em. long, .15 em. broad, obsoletely 3-nerved, midvein prominent;
corolla .7—.9 em. long, yellow, tube slightly less than one-half
the length of the entire corolla; lobes ovate, auriculate, papillate,
apiculate; spurs thick or slender, pendulous, more or less diver-
gent, giving the flower a triangular appearance just preceding
anthesis; stamens approximately .4 em. long; filaments linear,
anthers oval; capsule lanceolate; seeds ovoid.
Distribution: known only from Colombia.
Specimens examine
CoroMBia: Dept. Bentesder, Páramo de las Vegas, alt. 3700-3800 m., Dec. 20—21,
1926, Killip & Smith 15679 (M cian NY, US).
Species very similar to H. brevicorn
4. H. macrantha Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p.
105. 1916
Perennial herb, 5-6 cm. high, with root curved-erect, ligneous,
49H. adpressa Allen, sp. nov.—Herba perennis, usque ad 2.5 dm. alta; radice
crassa; caule plerumque solitario, erecto, tenui; foliis basalibus in rosula densa,
oblanceolatis, 1.-1.5 cm. longis, .3 cm. latis; foliis caulinis 5-6 geminis, sessilibus,
lanceolatis, 1-1.5 cm. longis, .2-.3 cm. latis, acutis; inflorescentia multiflorifera cyma,
pedicellis 2 cm. longis; calycis lobis lanceolatis, .4—.6 cm. longis, .15 cm. latis, ob-
solete 3-nerviis, medio-nervo prominenti; corolla .7—.9 em. longa, flava, tubo 14 totae
corollae longitudini parum subaequanti; corollae lobis ovatis, auriculatis, papil-
latis, apiculatis; calcaribus crassis vel tenuibus, gibbis, pendulis, plus minusve
divaricatis; staminibus ca. .4 cm. longis; filamentis linearibus, antheris ovalibus;
capsula lanceolata; seminibus ovoideis.—CoroMnBiA: Dept. Santander, Páramo de
las Vegas, alt. 3700-3800 m., Dec. 20-21, 1926, Killip & Smith 15679 (M TYPE, NY,
US).
1933]
ALLEN—THE GENUS HALENIA 149
rather elongate, covered with the blackish remains of marcescent
leaves; flowering stem single, erect, internodes 4-5 cm. long, with
no rosette leaves before anthesis; leaves herbaceous, acute,
3-nerved veins sunken above, prominent below; lower cauline
leaves oblanceolate, gradually narrowed into a long but broad
petiole, 6-7 cm. long, .5-1.5 cm. broad; upper leaves lanceolate
to ovate-lanceolate, broadly sessile, about 3 cm. long; inflores-
cence an apical 3-flowered cyme, solitary in axils of the upper-
most euphylloid leaves; pedicels of apical flowers 3 cm. long,
those of laterals 2 cm.; Locos oblanceolate, 1-1.3 cm. long,
3-nerved, acute; Boro about 1.5 em. long, green, tube about
one-third the length of the entire corolla; lobes ovate to broadly
ovate, very acute, somewhat apiculate; spurs large, globose,
conical, calluses about .3 cm. long at the base of the tube.
Distribution: Colombia
No specimens exicoloed, but description compiled from original publication.
(TYPE, Kalbreyer 702, BG).
5. H. Karstenii Gilg in Engl. Bot. Jahrb. 54: Beibl. 118,
105. 1916.
Biennial or perennial herb up to 5 dm. high; root branched;
stems simple, covered with leaf-bases for 7-8 cm. at intervals of
less than .4 cm., internodes up to 4—5 cm. long; leaves in pairs at
the nodes, sessile, lanceolate, narrowed at base, 2-5.5 cm. long,
up to .9 em. broad, 3-nerved, veins excurrent, forming a mucro;
inflorescence 4—12 flowers in terminal and axillary cymes, pedicels
erect, up to 4 cm. long; calyx-lobes lanceolate, .6—.8 cm. long,
.2-.25 em. broad, acuminate, 3-nerved, papillate; corolla about
1 em. long, tube not quite one-half the length of the entire
corolla; lobes ovate, subrotund, erose at apex; spurs midway up
the corolla tube, small, scarcely noticeable, glandular convex
depressions in the corolla; stamens about .3 cm. long, at the ori-
fice of the tube; filaments linear, anthers oval, acute; capsule
15-18 cm. long, ovate; seeds ovoid, light brown, wrinkled.
Distribution: páramos of Bogota.
ned:
Specimens examine
CoLoMBia: Páramo de Bogota, Karsten (V TYPE).
6. H. subinvolucrata Gilg in Engl. Bot. Jahrb. 54: Beibl. 118,
p. 99. 1916.
[Vor. 20
150 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Perennial herb, about 2 dm. high; root multifibrous; subter-
ranean stem vertical, short, thick, flowering stem erect, loosely
leafy; leaves thickly herbaceous or somewhat fleshy, 3-nerved,
veins sub-parallel, deeply sunken above, prominent below;
basal leaves linear-lanceolate, gradually narrowed at the base
into narrow petiole, 4-5 cm. long, .4-.5 em. broad; lower cauline
leaves similar to basal leaves; upper cauline leaves broadly
sessile, lanceolate or lanceolate-ovate to ovate, 2-4 cm. long,
.6-1 em. broad; inflorescence terminal or axillary, almost sessile
3-flowered cymes, peduncles .3—.6 em. long; floral leaves sheathing
inflorescence and longer than flowers; calyx-lobes oblong-ovate,
about 1 em. long, up to .3 em. broad, 5-nerved, minutely hirtellous,
acute; corolla about 1 em. long, tube about one-third or less the
length of the entire corolla; lobes oblong-ovate, .7 cm. long,
.3—.35 em. broad, subrotund, erose at apex; spurs scarcely prom-
inent blackish maculations at the base of the tube; stamens
about .2 cm. long; anthers ovate-oblong; filaments linear.
Distribution: Venezuela.
Specimens examined
VENEZUELA: high mountains of Trujillo and Mérida, alt. 1300-4900 m., 1842,
Linden 437 (DH TYPE).
7. H. inaequalis Wedd. Chlor. And. 2: 78. pl. 53 C. 1859;
Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 99. 1916.
Herb 1.5-2(-3) dm. high; stem aditafy. erect, loosely leafy,
with internodes about 4-5 cm. long, and inflorescence comprising
the upper third of the stem, or stem short, leafy, with internodes
equidistant, 1-2 em. long, arising in axil of every leaf; radical
leaves petiolate, lanceolate to oblanceolate, 2-3 em. long; upper
cauline leaves sessile, lanceolate to elliptic, minutely hirtellous;
flowers disposed in small loose pedunculate cymules; peduncles
erect or slightly nodding, 1-2 em. long; calyx-lobes lanceolate
or oblong-lanceolate, acute, margin minutely hirtellous; corolla
scarcely 1 cm. long, exceeding the calyx by one-fourth its length;
lobes ovate, acute; spurs inconspicuous.
Distribution: known nir from Venezuela.
Specimens examine
VENEZUELA: "PN de Timotes, Mérida, alt. 3800 m., Sept. 4, 1921, Jahn 558
(HP, US); same locality, alt. 3600 m., Jan. 21, 1922, Jue 839 (HP, US); Mérida,
alt. 3000 m., 1846, Funck & Schlim 901 (DH TIN).
1933]
ALLEN—THE GENUS HALENIA 151
Funck & Schlim 1148 is cited by Weddell in the original
publication of the species, but Gilg has placed this number under
H. viridis.
8. H. viridis E Gilg in Engl. Bot. Jahrb. 54: Beibl.
118, p. 100. 1916.
erba viridis Griseb. in Linnaea 22: 43. 1849; Wedd.
Chlor. And. 2:62. 1859.
H. inaequalis Wedd. Chlor. And. 2: 78. 1859; Gilg, l. c.
Perennial herb, 1-3 dm. high; stem solitary, erect, thick,
simple, internodes 4—5 cm. long; basal leaves in rosette, more or
less coriaceous, sessile, lanceolate, subequal, up to 6 cm. long, .4
cm. broad, minutely hirtellous, 3-nerved; cauline leaves 3-4
pairs, approximately 5 cm. long; inflorescence a narrow racemi-
form cyme arising in the middle of the stem, with simple 3-5-
flowered cymules; pedicels erect before anthesis, cernuous after,
about 2.5 cm. or less long; calyx-lobes oblong-lanceolate, minutely
hirtellous, about .4-.5 cm. long, nerved; corolla about 1 cm.
long, greenish, tube about one-third the length of the entire co-
rolla; lobes elliptic-lanceolate, rather obtuse, somewhat erose at
apex; spurs inconspicuous; stamens .5 cm. long; anthers ovate;
capsule oblong-lanceolate.
Distribution: known wy from Venezuela.
Specimens examine
VENEZUELA: Sierra Nevada: Mérida, alt. 3300 m., 1846, Funck & Schlim 1148 (DH
TYPE); Laguna Mucuy, Cabeceras del Saisay, Mérida, 4200-4300 m., April 19, 1930,
Gehriger 92a (HP).
9. H. Schultzei* Gilg, n. sp.
Caespitose mat-like perennial, about 3 dm. high; root tough,
fibrous; stems one or more, erect, simple, slender, internodes 5—7
50H. Schultzei Gilg, sp. nov.—Herba perennis, caespitosa, ca. 3 dm. alta; radice
lenta, fibrata; caulibus 1—multis, simplicibus, tenuibus, internodiis 5-7 cm.
folis basalibus in rosula densa, in petiolis longis tenuibus attenuatis, ad basin
dilatatis, lineari-lanceolatis, ca. 5 cm. longis, .5 cm. latis, nerviis, acutis; foliis caulinis
plerumque 2 geminis, sessilibus, linearibus, plerumque ca. 1.5 cm. longis, nerviis,
acutis; inflorescentia pauco-florifera cymis terminalibus lateralibusque, pedicellis
tenuibus, 1-1.5 cm. longis, ad apicem recurvatis; calycis lobis lanceolatis, ca
longis, acutis; corolla latissime campanulata, ca. 1.3 cm. longa, flava; corollae lobis
ovatis, obtusis marginibus erosis vel crispis; calcaribus inconspicuis.—(TYPE,
Schultze 1304, BG).
[Vor. 20
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN
em. long; basal leaves in dense rosette, narrowed into long
slender petioles, dilated at base, linear-lanceolate, about 5 cm.
long, .5 em. broad, nerved, acute; stem-leaves usually 2 pairs,
sessile, linear, usually about 1.5 em. long, nerved, acute; inflores-
cence few-flowered in terminal and lateral cymes, pedicels slender,
1-1.5 em. long, recurved at tip; calyx-lobes lanceolate, about .6
em. long, acute; corolla very broadly campanulate, about 1.3 cm.
long, yellow; corolla-lobes ovate, rather obtuse, erose or crisped
margins; spurs not discernible.
Distribution: Colombia
ecimens examined, but description compiled from photograph. (TYPE,
Schultze 1304, BG).
10. H. gentianoides Wedd. Chlor. And. 2: 78. pl. 53B. 1859.
Perennial herb, up to 3 dm. high; root more or less thick,
ligneous; stems sterile or flowering, long, loosely leafy; basal
leaves in a dense rosette, petiolate; cauline leaves 2-6 pairs,
subsessile, lanceolate, acute, 3-nerved; inflorescence lateral or
terminal, small racemiform cymes, peduncles more or less resu-
pinate; calyx-lobes oblong, up to .7 cm. long, papillate, acuminate,
prominently 3-nerved; corolla about 1 cm. long, tube approxi-
mately one-third the length of the entire corolla; lobes ovate,
rotund, crenulate-erose; spurs reduced to inconspicuous glandular
areas not visible to the naked eye; stamens approximately .4 cm.
long; filaments linear, anthers ovate, acuminate; capsule lanceo-
late.
Distribution: páramos of Colombia.
p mens examined:
Corownia: Páramo de Bogota, Karsten (V); same locality, Goudot (BG TYPE,
M photo, V); same locality, T'riana 1964 (DH, V); same locality, Jan. 17, 1854,
Holton 467 (DH, G); Guasca, 1919, Ariste-Joseph A423 (US).
11. H. stellarioides Gilg in Engl. Bot. Jahrb. 54: Beibl. 118,
p. 100. 1916.
Perennial? herb, up to 3 dm. high; flowering stem erect, loosely
leafy, internodes 4-7 cm. long; basal leaves lacking; cauline
leaves sessile, lanceolate to linear-lanceolate, 2.5-3.5 cm. long,
gradually decreasing toward the summit, .4—.5 cm. broad, acute,
3-nerved; inflorescence lateral or terminal, 5-7-flowered cymes,
pedicels more or less erect, up to 1.5 em. long; calyx-lobes lance-
1933]
ALLEN—THE GENUS HALENIA 153
olate or oblong-lanceolate, about .8 em. long, .25 cm. broad,
acute, 3-nerved, or nerves obsolete; corolla about 1 cm. long,
tube about one-third the length of the entire corolla; lobes ovate-
oblong, upper margin subcrenulate, acute; spurs semi-globose
callous prominences at the base of the tube.
Distribution: páramos of Colombia.
No specimens examined, but description compiled from original publication and
photograph. (TYPE, Lehmann 3080, BG).
12. H. verticillata Gilg in Engl. Bot. Jahrb. 54: Beibl. 118,
p.103. 1916.
Annual herb, slender, up to 1 m. high; root fibrous; stem .5 cm.
thick, becoming black, covered with the remains of early leaves,
densely leafy at the middle or just below the middle of the stem, in-
ternodes up to 8 em. long; basal leaves in dense whorls for a varying
distance up the stem, connate, sessile, linear-lanceolate, 10 cm. or
less long, .8 cm. broad, acuminate, 5-nerved fleshy; cauline leaves in
7-8 whorls of 3, lanceolate (extreme upper subt
ovate-lanceolate), 5-nerved, 5-6 cm. long, gradually decreasing
in length toward the summit, with corresponding increase in
width; flowers numerous, disposed in 3-5 verticillate, axillary
and terminal cymose clusters; pedicels erect, up to 3.5 cm. long;
calyx-lobes oblong-lanceolate, papillate on under surface of veins
and entire upper tip, up to 1.0 em. long, .2-.35 cm. broad, acu-
minate to acute, 5-nerved, veins parallel, becoming confluent at
tip; corolla apple-green or yellowish-green, up to 1.7 cm. long,
tube about one-third the length of the entire corolla; lobes
broadly ovate, subrotund, erose and papillate at tip; spurs sub-
globose glandular prominences at the base of the corolla.
Distribution: wet rS about voleanos, Colombia.
Specimens examine
CoroMnia: ‘“‘Cauca am Vulkan," Sotará, 3500 m., Lehmann 6190 (BG TYPE, F);
Dept. of El Cauca, Mt. Pan de Azucar, alt. 3500-3700 m., June 16, 1922, Pennell
7052 (ANSP, G, NY, S, US).
13. H. foliosa Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p. 101.
1916.
Biennial herb (?), up to 8 dm. high; root stout (?); flowering
stems l-many, erect, simple, few or no leaves at base, lower
internodes 1-1.5 cm. long, upper 5-7 cm. long; numerous pairs
[Vor. 20
154 ANNALS OF THE MISSOURI BOTANICAL GARDEN
of lower cauline leaves, thickly herbaceous, gradually narrowed
toward the base but dilated again at base, lanceolate, 3-5 cm.
long, about 1 em. broad, acute, 3-5 nerved; upper cauline leaves
herbaceous, gradually narrowed toward the base and broadly
sessile, ovate-elliptie, 3-5 em. long, about 1 em. broad, acute to
very acute, 5-nerved, veins parallel, sunken above, prominent
below; terminal and axillary, loosely arranged, many-flowered
elongate cymes, comprising a thyrsoid inflorescence 10 cm. long,
pedicels about 2.5 em. long; small upper leaves shorter than the
inflorescence; calyx-lobes ovate-oblong, .7—.8 cm. long, .3 em.
broad, acute or very acute, 3-nerved; corolla about 1.3 cm. long,
tube about one-fourth the length of the entire corolla; spurs
semi-globose protuberances at the base of the tube.
Distribution: plrunos of Colómbia.
Specimens examine
CoroMnia: Dept. of ‘Bolivar-Antioquia, Páramo de Chaquiro, alt. 3000-3200 m.,
Feb. 23, 1918, Pennell 4268 (NY, US). (TYPE not seen, Stübel 276, BG, M photo).
14. H. dasyantha Gilg in Engl. Bot. Jahrb. 54: Beibl. 118,
p.103. 1916.
Perennial herb, (1.5—-)3—7 dm. high; root thick ;stem erect, simple,
more than .5 em. thick, fleshy, brown in dried specimens, faintly
striate, covered with remains of marcescent leaves, 2—5 internodes
up to 10 em. long; basal leaves in a dense rosette, broadly elliptic,
up to 10 em. long, nearly 2 cm. broad, acute, 5-nerved; 3-5 pairs
of cauline leaves, sessile, dilated at in| oblong-lanceolate, 4—10
em. long, their length gradually decreasing toward the summit,
1-2 em. broad, acute, 5-nerved; inflorescence axillary and ter-
minal, many-flowered racemose cymes, usually 10 em. long;
pedicels erect, for most part up to 3.5 cm. long; calyx-lobes ovate
to ovate-lanceolate, papillate, up to 1 cm. long, .4-.5 cm. broad,
acute to abruptly acuminate, 3-nerved; squamellae frequently
scale-like lobed bodies; corolla 1.5-2 cm. long, pale greenish-
yellow, tube nearly equal in length to the entire corolla; lobes
ovate, subrotund, erose at apex, somewhat papillate; spurs small
subglobose prominences at the base of the corolla-lobes, almost
obscured by the calyx; stamens approximately .4 cm. long,
attached at the summit of the tube; filaments linear, anthers
ovate-oval; stigma reflexed; capsule up to 2 cm. long; seeds
elliptical, reticulate.
1933]
ALLEN—THE GENUS HALENIA 155
Distribution: moist grassy páramos, or dry open woods, Colombia.
Specimens examined:
CoroMBia: Dept. of Caldas, Páramo del Quindio, alt. 3700-4200 m., Aug. 15-20,
1922, Pennell & Hazen 9997 (ANSP, NY, US); Dept. of Tolima, Páramo de Ruiz,
alt. 3500-3800 m., Dec. 16-17, 1917, Pennell 3001 (NY, M, US); Dept. of Cauca,
Páramo de Buena Vista, Huila group, Central Cordillera, alt. 3000-3600 m., Jan
1906, Pittier 1111 (US). (TYPE not seen, Lehmann 2065, BG, M photo.).
This species is very similar to H. elata but is, on the whole, a
larger and coarser plant. Since the material is scanty and the
geographical location different, H. dasyantha has for the present
been maintained as a distinct species.
15. H. elata Wedd. Chlor. And. 2:78. 1859.
Perennial, up to 5 dm. high; stem thick, coarse, erect, loosely
leafy; numerous basal leaves in dense rosette, broadly lanceolate,
attenuate into petiole, dilated at base, 7-10 cm. long, approxi-
mately 1 cm. broad, 5-nerved, acuminate; cauline leaves more or
less sessile, shorter, lanceolate, acute; inflorescence numerous
loose terminal and lateral cymes, peduncles up to 3 cm. long;
calyx-lobes oblong-lanceolate to ovate-lanceolate, .7—.8 cm. long,
3-nerved, papillate, acute; corolla hardly one-fourth more than
the length of the calyx, tube slightly more than one-half the
length of the entire corolla; lobes ovate, obtuse; spurs inconspicu-
ous tubercules at the extreme base of the corolla; stamens .6
em. long; anthers narrowly ovate, filaments linear.
Distribution: in ie vas Nevada of Santa Marta, Venezuela.
Specimens exami
VENEZUELA: MN Nevada de Santa Marta, Caracas, 1844, Funck 4156 (DH
TYPE).
16. H. Tolimae Gilg in Engl. Bot. Jahrb. 54: Beibl. 118, p.
101. 1916.
Perennial herb, 2.5-3 dm. high; root thick; root-stalk thick,
densely covered with remains of marcescent leaves; one to few
flowering stems, thick, erect, simple for the most part, very
narrowly winged, internodes 3-6 cm. long; basal leaves arranged
in rosette, slightly narrowed into long broad petioles, dilated at
the base, lanceolate, 5-8 cm. long, up to .5-.6 em. broad, 3-5-
nerved, acute; stem-leaves 2-3 at a node, sessile, broadly lanceo-
late, 4-6 cm. long, .6—7 cm. broad, their size gradually de-
creasing toward the summit, 3-5 subparallel veins, sunken above,
[Vor. 20
156 ANNALS OF THE MISSOURI BOTANICAL GARDEN
prominent below, reticulate; inflorescence few-flowered (5-7)
cymes, axillary and terminal; pedicels more or less resupinate, up
to 3 em. long, the central one longer; calyx-lobes ovate to ovate-
oblong, papillate, .7—.9 cm. long, .25—.35 em. broad, subacuminate,
3-nerved; corolla up to 1.3 em. long, light greenish-yellow, tube
slightly less than one-third the length of the entire corolla; lobes
ovate, erose and papillate at the tip; spurs glandular subglobose
protuberances at the base of the tube, obscured by the calyx;
stamens approximately .4 cm. long; filaments linear, anthers
ovate; capsule lanceolate, apiculate, 2 cm. long; seed ovoid-
elliptical, very minutely reticulate, pale tan.
Distribution: grassy páramos of Colombia.
Specimens examined:
CoroMnBia: Dept. of Caldas, Páramo del Quindio, alt. 4100-4400 m., Aug. 15-20,
1922, Pennell & Hazen 9841 (ANSP, G, NY, US); (typ not seen, Stübel 228, BG,
M photo.); bare loam slopes below snow, same locality and date, alt. 4300-4500 m.,
Pennell & Hazen 9894 (ANSP, NY, S, US).
This last-cited specimen has leaves broader than those of the type, but is similar
otherwise. H. Tolimae appears to be closely related to H. elata.
17. H. hygrophila Gilg in Engl. Bot. Jahrb. 54: Beibl. 118,
p.102. 1916.
Biennial herb, about 6 dm. high; root ligneous; subterranean
root-stalk covered with darkened leaf-bases; flowering stem
fleshy, erect, simple, 1 cm. thick, faintly winged, internodes up
to 8 em. long; 30-40 basal leaves in a dense rosette, linear, up to
12 em. long, .5-.9 em. broad, acute, 5-nerved; cauline leaves
yellow-green, 3-4 pairs, the lower lanceolate, the upper sub-
tending the inflorescences, ovate-lanceolate, sessile, 5-7 «cm.
long, gradually decreasing in size toward the summit, approxi-
mately .8-1.2 em. broad, 7-nerved, parallel veins confluent at
tip; inflorescence terminal and axillary in upper stem-leaves, in
dense many-flowered cymes, giving spicate appearance, pedicels
erect, up to 3 cm. long; calyx-lobes ovate-oblong to ovate,
papillate, .9-1.0 cm. long, narrowly acute, 5-nerved, reticulate;
corolla up to 1.5 em. long, yellowish-green, tube approximately
one-third the length of the entire corolla; lobes ovate, erose at
apex; spurs angular sac-like prominences at the extreme base of
the corolla lobes, obscured by calyx; stamens approximately
.7 em. long, attached at the orifice of the tube; filaments linear,
1933]
ALLEN—THE GENUS HALENIA 157
anthers broadly oblong, apiculate; capsule lanceolate; seeds
elliptic, pale tan, reticulate.
etnias páramos of Andes, Colombia.
Speci d:
diens : Páramo of Guanacas, Central Andes of Popayan, 3000-3600 m.,
Lehmann 7990 (F, à. BG TYPE, M photo, US)
18. H. parallela* Allen, n. sp.
Perennial herb, up to 6 dm. high; root-stalk thick; flowering
stem single, erect, simple, or branched above, striate, lower inter-
nodes 1-1.5 em. long, upper 4-6 cm. long; numerous basal leaves
in rosette, almost sheathing, lanceolate, 3-5-nerved, about 9
em. long, 1 em. broad; cauline leaves about twelve pairs, more
remote toward the apex, sessile, lanceolate to elliptic, 4-6 cm.
long, gradually decreasing toward the summit, approximately 1
cm. broad, 3-5 subparallel veins sunken above, prominent below;
inflorescence 1 or more many-flowered axillary and terminal
cymes, pedicels more or less erect, up to 3.5 cm. long, the lateral
somewhat shorter than the terminal; calyx-lobes ovate, papillate,
.7-.9 cm. long, .4 em. broad, attenuately acute, prominently
5—7-nerved, veins subparallel; corolla about 1.3 em. long, probably
greenish?, tube about one-fourth the length of the entire corolla;
lobes broadly ovate, slightly papillate, crisped toward the tip,
abruptly acuminate; spurs large globose protuberances at the base
of the tube, obscured by calyx; stamens nearly .5 cm. long,
attached at the orifice of the tube; filaments linear, anthers
oblong; capsule lanceolate.
5 H. parallela Allen, sp. nov.—Herba perennis, usque ad 6 dm. alta; radice crassa;
aule florifero solitario, erecto, simplice vel supra ramoso, striato; internodiis
enn 1-1.5 em. longis, superioribus yet: m. longis; foliis iuo multis,
in rosula densa, fere Serin bur, lanceolatis, P vi iis, ca. 9 em. longis, 1 cm.
latis; foliis caulinis ca. duodecem geminis, ad apicem remotioribus, seii. lanceo-
latis vel ellipticis, 4-6 cm. longis, sensim sursum decrescentibus, ca. 1 cm. latis,
3-5-nerviis, nervis subparallelis, supra immersis, infra prae infor-
escentia cymis 1- I apri axillaribus et terminalibus; pedicellis plus
minusve erectis, usque ad 3.5 cm. longis, lateralibus terminalibus brevioribus;
calycis lobis ovatis, popillats, 4-. 9 cm. longis, .4 cm. latis, attenuate acutis, pro-
minente 5-7-nerviis, nervis subparallelis; corolla ca. 1.3 cm. longa, forte viridula,
tubo ca. 14 totae corollae longitudini adaequanti; lobis late ovatis, parum papillatis,
ad apicem crispis abrupte acuminatis; calcaribus magnis globosis gibbis ad basin
tubi calyce obscuratis; staminibus ca. .5 cm. longis, tubi summo adjunctis;
filamentis linearibus, antheris oblongis; capsula lanceolata.—V ENEZUELA: Páramo de
La Negra, Mérida, Dec. 1927, Gutzwiller 32 (HP TYPE, US)
[Vor. 20
158 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: Venezuela.
Specimens examin
VENEZUELA: Páramo de La Negra, Mérida, Dec. 1927, Gutzwiller 32 (HP TYPE,
US).
Species near H. foliosa and H. hygrophila.
19. H. major Wedd. Chlor. And. 2: 79. 1859.
Annual probably, up to 6 (?) dm. high; (root not seen); stem
erect, simple below, branched above, loosely leafy, very narrowly
winged and striate; leaves sessile, subconnate, broadly elliptic,
2-7 em. long, .5-2 em. broad, somewhat abruptly acuminate,
3-5-nerved; inflorescence terminal and axillary 3-6-flowered sub-
umbelliform cymes, pedicels 4 cm. long, usually erect, frequently
nodding at apex; calyx-lobes oblanceolate to subspatulate, up
to .8 em. long, .25 em. wide, 3-nerved, midvein very prominent ;
corolla 1.0-1.3 em. long, tube almost one-half the length of the
corolla; lobes ovate, erose at apex; spurs small upcurved conical
protrusions almost at the orifice of the tube; stamens approxi-
mately .5 em. long, at the orifice of the tube; filaments linear;
anthers not seen; capsule ovate, 1.3-1.6 cm. long; seeds oval,
brown-black, wrinkled.
Distribution: shrub zone, mountain bases, Colombia.
d:
BIA: Dept. of El Cauca, Mt. Pan de Azucar, alt. 3300-3600 m., June 16,
1922, Pennell 7084 (ANSP, NY, US); Pag of Cundinamarca, Sibate, alt. 2700-2800
m., Oct. 13-15, 1917, Pennell 2438 (M, US); Dept. of Caldas, Cerro Tatama, alt.
3400-3700 m., Sept. 8-10, 1922, slg 10575 (US). (TYPE not seen, Goudot, HJP).
SECTION 2. HALENIASTRUM
KEY TO NORTH AMERICAN SPECIES AND VARIETIES
l. m spreading to ascending.
ant less than 2.5 dm. high.
3. Biennial; flowers white; distribution Mexico............ 20. H. crassiuscula
3. Annual; flowers purple; distribution Canada. 23a. H. deflexa var. Brentoniana
2. Plant more than 2.5 dm. high.
3. Spurs, if present, about !4 the length of the corolla......... 21. H. Pringlet
3. Spurs less than 1% the length of the corolla.
4. Annuals; leaves mostly cauline.
5. Stem-leaves linear. ............ ccc cece cc ccc eeeecees 22. H. recurva
5. Stem-leaves lanceolate to ovate..........00... 0000 cece 23. H. deflexa
4. Perennials; leaves mostly radical.
. Strict, many-flowered, spike-like inflorescence.
D DEM RE Lieber eres q9EXECODCKERK ER" wens 24. H. rhyacophila
6. Stem procumbent.............. 24a. H. rhyacophila var. procumbens
1933]
ALLEN—THE GENUS HALENIA 159
5. Loose, broad, few-flowered inflorescence............... lesus.
YogeAet "PTS REPE A 24b. H. rhyacophila var. macropoda
1. he pendulous to incurved.
. Leaves not apiculate or very rarely so; ender not mucronulate.
3. Basal rosette absent; leaves mostly cau
4. Habit erect; leaves less than 3 cm. (he
D.C NE I oo ees ses ec tee ee CES 25. H. Palmeri
5. Leaves lanceolate or ovate.
6. Calyx-lobes mostly obtuse, appressed........... 26. H. Conzattit
6. Calyx-lobes acute, reflexed.............. suus. 27. H. Schiedeana
4. Habit prostrate; leaves 5-12 cm. long.................. 28. H. caleoides
3. Basal leaves present, cauline few o
4. Spurs incurved, 14 or less than 4 the linen of the corolla.
5. Sterile branches present; leaves numerous. ......... 29. H. platyphylla
5. Sterile branches absent; leaves less than 15.
6. Flowers 1.5 cm. or less lon
7. Flowers less than 1 em. long; spurs rudimentary. .30. H. nudicaulis
7. Flowers more than 1 cm. long; spurs 14-14 the length of the
oo ee ik os oe haa a dod ae ee 31. H. plantaginea
6. Flowers more than 1.5 cm. long...31a. H. plantaginea f. grandiflora
4. Spurs spreading, about l4 the length of the corolla.
5. Stems always erect.
Gea ves lineaments coos Se ie lene oU 32. H. Shannonii
0. Leaves elipüeal...........Leeees. 82a. H. Shannonii f. compacta
5. Stems more or less deceumbent.................ss. 33. H. decumbens
2. Leaves conspicuously apiculate; calyx-segments mucronulate.
3. Plant more than 2 dm. hi 5 Ou Bul ee Oe eed OE . H. guatemalensis
3. Plant less than 2 dm. high........... 34a. H. guatemalensis var. latifolia
20. H. crassiuscula Robinson & Seaton in Proc. Am. Acad.
28:113. 1899.
Small caespitose biennial of dense habit, slightly fleshy; stems
erect, 0.4-1.0 dm. high, narrowly winged, much branched;
radical leaves broadly oblanceolate to elliptic, 2 cm. long, obtuse,
attenuate into long petiole nearly equalling the blade, 3-nerved;
eauline leaves 1-3 pairs, narrowly oblanceolate to oblong, nar-
rowed at the base; inflorescence dense compact umbellate cyme;
flowers terminal or axillary, pedicellate, after anthesis slightly
nodding, not at all resupinate; calyx-segments lanceolate to
oblanceolate, .45-.6 cm. long, obtuse, papillate, 3-nerved; corolla
white, up to 1.5 cm. long, about .5 em. broad at base, tube .35-.4
em. long; lobes oblong-elliptic, acute; spurs .4 cm. long, arising
slightly below the midpoint of the tube, slender, spreading, and
curved upwards; stamens .25 cm. long, anthers ovate; filaments
linear; capsule lanceolate, frequently subfalcate, acute, 1.4 cm.
long; seeds globose, light yellow-brown, granular.
[Vor. 20
160 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: bare alpine summits, Mexico.
Specimens examine
Mexico: Nevado de Toluca, bare alpine summits, alt. 3500 m., Sept. 2, 1892,
Pringle 4229 (ANSP, BG, BM, B, CAS, C, DH, F, G TYm, IAC, K, M, NY, S, US,
V); Ixtaccihuatl, wet meadows, alt. 3000-3250 m., March-July 1903, Purpus 318
(CAS, M, US); Popocatepetl, Aug. 7-8, 1901, Rose & Hay 5999 (US).
21. H. Pringlei Robinson & Seaton in Proc. Am. Acad. 28:
113. 1893.
Halenia candida Ramirez in Inform. Secret. Foment. Mexico
(Excurs. Mont. Ajusco), 34. 1895; Estud. Hist. Nat. 102. 1904.
Biennial, of glaucous aspect; root thick, tough, ligneous; stem
usually solitary, occasionally caespitose, simple or nearly so,
scape-like, slender, erect, 1-2.5 dm. high; leaves less than 3 cm.
long, radical leaves elliptic to narrowly oblanceolate, faintly 3-
nerved, attenuate below into slender petioles, usually as long as
the leaf-blade and persistent; cauline leaves 1-2 pairs, sessile,
short, sublinear, 1.5-3 cm. long, about .3 em. broad; flowers
disposed in terminal, or occasionally lateral, few-flowered um-
belliform cymes, pedicels less than 2 cm. long, usually about .8
em.; calyx-lobes oblong-spatulate, .3—.5 cm. long, acuminate, 3-
nerved; corolla white, .8-1.5 cm. long, spurless in the majority
of cases; when spurs present, slender, spreading, and curved-
assending, 1.5-1.8 em. from tip to tip, with prominent veins and
glandular tips; corolla-tube up to .2 cm. long; corolla-lobes
elliptic, acuminate; stamens .2-.35 em. long; anthers narrowly
ovate, minutely papillate, filaments linear, slightly uncinate;
capsule lanceolate, acute, exserted; seeds subglobose, yellow-
brown, granular.
Distribution: springy meadows of central and south Mexico.
Specimens examine
Mexico: springy alpina meadows, Sierra de las Cruces, alt. 2450 m., Aug. 28,
1904, Pringle 13121 (BG, C, UC, G, K, US); same locality, Aug. 23, 1892, Pringle
4209 (ANSP, C, CAS, DH, F, G TYm, IAC, K, M, NY, 8, US, V); same locality,
June 1895, €—— 908 (US).
X1CO: without locality, 1920-21, Reiche 36 (BG).
The habit of this species 1s very similar to that of H. nudicaulis.
The Pringle specimens cited above were collected in August, and
the plants are smaller and grow less luxuriantly than the single
specimen collected in June by Altamirano. The specimens
collected later in the season very rarely possess spurs, while the
1933]
ALLEN—THE GENUS HALENIA 161
earlier plants show a distinctly spurred corolla. This condition
is shown in other species to a somewhat less extent, and is in all
probability traceable to variation in environmental conditions.
22. H. recurva (Sm.) Allen, n. comb.
Swertia recurva Smith in Rees, Cyclopedia 34: sub Swertia.
1819.
Halenia Rothrockit Gray in Proc. Am. Acad. 11: 84. 1876;
Rothrock, Rept. Wheeler Exped. 195, pl. 21. 1878; Hemsl.
Biol. Cent.-Am. Bot. 2: 353. 1882.
Tetragonanthus Rothrockii Heller, Cat. N. Am. Pl. 6. 1898,
and ed. 2, 16. :
Annual, 2.5-5 dm. high; stem simple, often branched above;
basal leaves less than 3.5 cm. long, .6 cm. broad, elliptic-lanceolate
to spatulate; cauline leaves remote, lance-linear, 1.5—4 cm. long,
about .35 em. broad, obscurely 3-nerved, midrib prominent
below; inflorescence a loosely flowered subumbellate cyme;
flowers on slender pedicels, .5-3 cm. long, often in sevens; calyx
lobes lanceolate, elongate-acute, up to .6 cm. long, uninerviate,
papillate; corolla bright-yellow, about 1-1.2 cm. long, tube less
than one-half the length of the entire corolla; corolla-lobes ovate,
subacuminate, delicately veined, papillate; spurs curved, hori-
zontal or ascending, up to 1.6 cm. from tip to tip; anthers broadly
oblong, mucronate, papillate; filaments slightly obovate; capsule
ovate-lanceolate; seeds yellow-brown, subglobose-ovoid, granular.
Distribution: southern United States and Mexico.
Specimens examined:
UNITED STATES:
Arizona: Mt. Graham, alt. 2250 m., Sept. 1874, Rothrock 733 (ANSP, F, IAC, M,
US); same locality, Aug. 1874, Rothrock (G, NY); Chiricahua Mts., Sept. 22. 1931,
Jones 28603 (M); Barfoot Park, Chiricahua Mts., alt. 2000-2050 m., rolling andesitic,
recently lumbered pine land, Sept. 8, 1906, Blumer 1859 (BG, D, G, K, ^
US, V); same locality, alt. 2480 m., Sept. 22-23, 1914, Eggleston 10774 (US); Apache
Pass, Chiricahua Mts., Sept. 1881, Lemmen & Lain mon (CAS); Hermitage, Chiricahua
Mts., Sept. 1881, Lemmon & Lemmon (CAS); Rucker Valley, Chiricahua Mts.,
B 1881, collector unknown 1874 (CAS, DH, F, SM); White Mts., Aug. 1878, fw
(F); summit of White Mts. (Springerville-Fort Apache Road), Apache Reservation,
alt. 2270-2880 m., Aug. 29, 1919, Eggleston 15781 (F); Riverside Ranger Station,
Greer, Apache Forest, alt. 2700 m., Aug. 24, 1920, Eggleston 17137 (NY, US);
grassy flats near Brinkley’s Ranch, White Mts., Aug. 5, 1915, Ellis 20 (US); Riggs
Flat, Pinaleno Mts., alt. 2000 m., Sept. 23, 1917, Shreve 5373 (G); Columbia Trail,
Pinaleno Mts., alt. 2500 m., Sept. 13, 1914, Shreve 4312 (CAS, US).
[Vor. 20
162 ANNALS OF THE MISSOURI BOTANICAL GARDEN
New Mexico: Mogollon Mts., on or near the west fork of the Gila River, Socorro
Co., alt. 2125 m., Aug. 14, 1903, Metcalfe 501 (M, NY); swampy ground, divide
of Mogollon Mts., Sept. 7, 1881, Rusby 264 (ANSP, BM, CAS, F, K, M, NY, US);
same locality, Aug. 1881, Rusby (IAC, NY).
M .
EXICO:
Cuinuanvua: Mt. Mohinora, Sept. 1, 1898, Nelson 4868 (G, US); cool slopes,
Sierra Madre, alt. 1750-2375 m., Sept. 27, 1888, Pringle 1663 (BG, BM, B, CAS, DH,
M, NY, 8, V); same locality, Sept. 24, 1887, Pringle 1329 (ANSP, BG, C, G, K,
NY, US); Meadow Valley, Sierra Madre Mts., alt. 1750 m., Sept. 17, 1903, Jones
(S); Sierra Madres, near Colonia Garcia, alt. 2000 m., ent, 6, 1899, Townsend &
Barber 309 (BB, F, DH, G, M, NY, US); Escalon, Mutis (L).
CoanviLa: Sierra Madre, 40 m. south of Saltillo, July 1880, Palmer 839 (ANSP, G,
K, US).
DunaNao: Barranca, below Sandia Station, alt. 1625 m., Oct. 13, 1905, Pringle
13588 (G, 8, UC, US); Sierra de Candela, alt. 3000 m., Aug. 27, 1903, Endlich 53
).
JALISCO: Sierra de Tequila, alt. 2000 m., July 5, 1893, Pringle 5465 (G).
The name Swertia recurva Smith was given to the specimen
collected by Mutis and sent to Linnaeus, now preserved in the
herbarium of the Linnaean Society of London. The description
published in Rees’ ‘Cyclopedia’ was inadequate, and later the
specific name recurva was placed under deflexa, the well-known
northern species. Since few of the succeeding monographers
ever saw the original Swertia recurva Sm., it is not strange that
the error persisted. Over fifty years later, Gray described
Halenia Rothrockii as a new species.
si
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ALLEN—THE GENUS HALENIA
THE SENSITIVITY OF ORCHID SEEDLINGS
TO NUTRITIONAL IONS
F. L. WYND
Assistant in the Henry Shaw School of Botany of Washington University
I. INTRODUCTION
The germination and growth of orchid seedlings on artificial
media have received considerable attention from plant physiolo-
gists. Almost invariably, however, their investigations have
been concerned with the carbohydrate metabolism to the ex-
clusion of the inorganic nutritional relationships. The purpose
of the present paper is to report the results of a study on the sensi-
tivity of germinating orchid seeds and young seedlings to different
nutritional ions. The few available data on inorganic nutrition
are scattered in the papers dealing primarily with other phases
of orchid development on artificial media.
Burgeff (09) pointed out that orchid seeds germinated more
favorably when nitrogen was available as ammonium sulphate
than as a nitrate salt. This fact led Knudson (22) to modify
Pfeffer's solution, and this is the solution frequently mentioned
in orchid literature as ‘‘Knudson’s Solution B." Clement (724,
’24a) has said that the nutrition of different species of Odonto-
glossum varies widely, but he did not elucidate the matter further.
Ballion and Ballion (’24) made a similar statement, that in their
work with Cattleya, Miltonia, and Odontoglossum the nutritional
conditions varied according to the species, but here again no
specific information was given. La Garde (29) found that his
solution L gave about 10 per cent better growth than did Knud-
son's solution B, and attributed this superiority to the slight
increase of nitrogen and the large increase of potassium and
phosphorus in his solution. Smith (32) used Knudson's solution
B, but found that doubling or trebling the amount of ammonium
sulphate gave better growth and a deeper color to his seedlings.
In so far as we have been able to ascertain, the above brief
review is complete as to published data concerning the inorganic
nutrition of orchid seedlings.
ANN. Mo. Bor. GARD., Vor. 20, 1933. (223)
[Vor. 20
224 ANNALS OF THE MISSOURI BOTANICAL GARDEN
II. METHODS
Livingston ('19) has clearly defined four possible criteria for
the comparison of different nutrient solutions, as follows:—
(1) The volume-molecular proportion of any one salt.
(2) The osmotic proportion of any one salt.
(3) The total volume-molecular concentration.
(4) The total osmotic concentration.
As it is impossible to vary all the features of the composition of
a series of solutions regularly at the same time, it is necessary to
select arbitrarily one characteristic as the basis for comparison.
Since it is probable that the solution must act upon plants pri-
marily in an osmotic way, we have followed Livingston and many
others in using the total osmotie concentration as the basis for
comparative studies.
The triangular representation of a series of solutions of three
components varying in any of the above ways was first introduced
in physical chemistry by Schreinemakers (093) and again by
Bancroft (02). Schreiner and Skinner (710) first applied this
method of investigating nutrition in their work on fertilizers.
Since 1910 it has been used by various investigators to study
the nutrition of several agriculturally important plants: wheat,
by Shive (15), McCall (16), Livingston and Tottingham ('18),
Van Alstine (19), Meier and Halstead (21), and by Tottingham
and Rankin (22); buckwheat, by Shive (715, '17) and by Shive
and Martin (718, ’18a); potato, by Johnston ('24) and by Martin
and Shive (720); celery, by Poole and Fant (22); cranberry, by
Addoms and Mounce (731); peach seedlings, by Davidson (728).
After investigating the work of the above authors, it appeared to
us that the triangular system of arranging a varying series of
nutrient solutions provided the most satisfactory approach to
the problem of the reaction of orchid seedlings to their inorganic
substrate.
Seeds for the investigation were furnished from the green-
houses of the Missouri Botanical Garden through the courtesy of
Dr. George T. Moore, the Director. Flowers of Cattleya Trianae
Linden and Rchb. f. were pollinated November 17, 1930, and the
largest and apparently the best pod was picked May 3, 1932,
after a development period of almost eighteen months. It has
1933]
WYND—ORCHID SEEDLINGS AND NUTRITIONAL IONS 225
been the author’s experience that in general the capsules first to
mature furnish seeds of lower vitality than those maturing later;
hence the one requiring the longest time to mature was used to
supply seeds for the present work.
The nutrient solutions investigated were those of Type I and
Type IV, as described by Livingston (19). By the use of only
three salts it is possible to arrange solutions containing the six
major nutritional ions in six different combinations. Of these
six possible combinations, only two will have all their ions added
in unlike combinations. The compositions of the solutions used
are given in tables 1 and rr. In both types the total osmotic
TABLE I
TYPE I—KH:;PO:, Ca(NO??, MgSO:
Molecular proportion Volume-molecular concentration
Culture
No. KH;PO, Ca(NOs)2 MgSO* KH;PO* Ca(NO3); MgSO*
R1 S1 1 1 6 0027 .0027 0161
S2 1 2 5 0025 .0049 0123
S3 1 3 4 0024 0071 0094
S4 1 4 3 0022 0089 0067
S5 1 5 2 0022 0108 0043
S6 1 6 1 0020 0122 0020
R2 S1 2 1 5 .0053 .0027 0132
S2 2 2 4 .0049 .0049 0099
S3 2 3 3 .0047 .0071 0071
S4 2 4 2 .0045 .0090 0045
S5 2 5 1 .0041 0104 0021
R3 Sl 3 1 4 0076 .0025 0101
S2 3 2 3 0072 . 0048 0072
S3 3 3 2 0068 . 0068 0045
S4 3 4 1 0065 .0086 0021
R4 S1 4 1 3 0099 0025 0074
S2 4 2 0094 .0047 0047
S3 1 90 .0068 0022
R5 $81 5 1 2 0123 .0024 0049
S2 5 2 1 0118 .0047 0023
R6 S1 6 1 1 0145 0024 0024
[Vor. 20
226 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE II
TYPE IV—K380,, Ca(HiPOQ9s Mg(NO3):*
Molecular proportions Volume-molecular proportions
Culture
xs K;SO, |Ca(H;POj,*|Mg(NOss|| K;SO, | Ca(H;PO,.| Mg(NOs)2
Ri S1 1 1 6 .0018 .0018 .0108
82 1 2 5 .0018 .0036 .0092
S3 1 3 4 .0019 .0056 .0075
R2 S1 2 1 5 0037 0018 0091
82 2 2 4 0037 0037 0074
S3 2 3 3 0037 0056 0056
R3 S1 3 1 4 0056 .0019 0075
82 3 2 3 0056 0037 0056
83 3 3 2 0056 0056 0038
R4 S1 4 1 3 0076 0019 0057
S2 4 2 2 0077 0039 0039
83 4 3 1 0078 0059 0019
R5 Sl 5 1 2 0097 0019 0039
82 5 2 1 0098 0039 0019
R6 $1 6 1 1 .0116 .0019 .0019
* The solutions containing the highest proportion of Ca(H;PO,); could not be pre-
pared, because sterilization at the temperature and pH used caused precipitation.
concentration was one atmosphere. Mallinckrodt salts of high-
est reagent quality were used. One-hundred-ce. portions of the
nutrient solutions were placed in 200-cc. Erlenmeyer flasks, 1.75
per cent of Merck's powdered “Reagent” agar added, and steri-
lized by autoclaving at 20 Ibs. pressure for 20 minutes. Follow-
ing the work of La Garde (’29), who found that maltose was the
best source of carbohydrate, a 2 per cent concentration of this
sugar was used in all cultures. A study of the hydrolysis of this
sugar under our conditions of sterilization showed that about
10 per cent was hydrolyzed to glucose.
In addition to the six major nutrient ions there were added
one-half part per million of manganese as manganese sulphate,
and one-half part per million of boron as sodium borate. A
ferric phosphate suspension was prepared as described by Living-
1933]
WYND— ORCHID SEEDLINGS AND NUTRITIONAL IONS 227
ston (’19) and 1 cc. added to each liter of nutrient solution.
This gave ferric phosphate in the concentration of about 3 mgms.
per liter.
Owing to the sensitivity of orchids to the acidity of the medium,
each solution was so adjusted that the pH after sterilization was
4.8 to 5.0. Tables mr and rv indicate the pH relationships be-
fore and after sterilization. The seeds were sterilized by shaking
TABLE III
THE ACIDITY RELATIONSHIPS OF TYPE I SOLUTIONS*
A B C
Sol. eps Fe, a Mn, ee pH after 1.75% agar
and 2% maltose | py as made up | PH adjusted before | a aged and auto-
autoclaving claved
R1 Sl 5.74 3.93 4.9
82 5.10 3.91 5.0
S3 4.93 3.90 5.0
S4 4.90 3.90 5.0
S5 4.83 3.90 5.0
S6 5.08 3.90 5.1
R2 S1 4.86 3.93 4.8
82 4.75 3.88 5.0
S3 4.73 3.93 4.9
S4 4.73 4.10 5.0
S5 83 4.10 5.0
R3 S1 5.26 4.22 5.0
82 4.86 4.24 5.0
53 4.83 4.20 5.0
S4 76 4.20 5.0
R4 81 4 4.20 5.0
S2 4.81 4.22 5.0
53 4.24 4.9
R5 S1 4.86 4 4.9
S2 4.83 4.22 4.9
R6 S1 4.80 4.29 4.9
*'The pH determinations indicated in columns A and B were obtained by the
quinhydrone electrode, while the results in Column C were obtained colorimetrically
by the use of brom-cresol-green as an indicator.
[Vor. 20
228 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE IV
ACIDITY RELATIONSHIPS OF TYPE IV SOLUTIONS
DETERMINATIONS MADE AS INDICATED IN TABLE III
A B C
Sol. plus Fe, B, Mn
y, 2n, . pH after 1.75% agar
and 2% maltose pH as made up pH adjusted. before |" | (ded and auto-
autoclaving claved
R1 S1 4.3 4.1 5.0
82 3.71 4.18 4.9
S3 3.5 4 4.9
R2 S1 3.89 5.0
S2 3.67 4.20 4.9
53 3.59 4.29 4.9
R3 S1 4.05 3.90 4.9
S2 3.73 4.22 5.0
S3 3.64 4.30 4.8
R4 S1 3.91 3.91 5.0
S2 3.69 4.25 4.9
S3 .54 32 4.9
R5 Sl 4.34 3.81 4.9
82 3.78 4.15 5.0
R6 S1 4.27 3.90 5.0
20 minutes in à small vial of calcium hypochlorite solution pre-
pared as described by Wilson (15). They were then inoculated
directly into the flasks of nutrient agar by a platinum wire in-
oculating needle. The flasks were incubated at 20-25? C. in a
specially shaded greenhouse compartment. During the warmer
summer days the greenhouse temperature could not be held
below 30°, and on rare occasions 35°, but the short duration of
these high temperatures apparently was without any harmful
effect.
III. RESULTS
Type I.—The growth period of these cultures extended from
April 15 until November 12, 1932. At the end of this time, the
1933]
WYND— ORCHID SEEDLINGS AND NUTRITIONAL IONS 229
diameter of the seedlings was measured by a low-power microscope
equipped with a calibrated eye-piece micrometer according to
the method of Quednow (730). The flasks had been very lightly
inoculated so as to eliminate crowding of the growing seedlings.
The figures in table v indicate the diameter in microns, each
TABLE V
GROWTH OF SEEDLINGS IN TYPE I SOLUTION. THE FIGURES
EPRESENT DIAMETERS IN MICRONS
Solution Series 1 Series 2
R1 S1 915 910
82 1100 1130
S3 970 1020
S4 1015 1010
S5 925 940
S6 810 830
R2 S1 (Contaminated) (Contaminated)
S2 925
S3 855 845
S4 600 740
S5 740 690
R3 S1 805 715
82 660 740
S3 690 725
S4 585 445
R4 S1 860 690
S2 620 620
S3 620 670
R5 S1 635 640
S2 475 585
R6 Sl 680 615
based on an average of 25 individuals. Figure 1 shows graphi-
cally the comparative development on the different media used.
Figures 3 and 4 indicate the areas of the triangles corresponding
to greatest, medium, and least growth. The circles are drawn to
scale and represent comparatively the magnitude of the seedlings
of the respective cultures.
[Vor. 20
230 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Type IV.—The growth period of these cultures lasted from
April 28 until November 20, 1932. Table vr presents the data
for the growth on these solutions. As before, the diameter is
indicated in microns and is the average of 25 determinations.
Figure 2 shows graphically the comparative development.
ha Ri
Rive
Ri Ss + e amaaan ene e ee A a
R 5s —————— n fare aan m atas s ttai B MM n t
Ra S2 LTD
Rid ——————— BTE
Ra Si ———————— MM —Ó—M Mo M ne esti ars a RR eae
Re 53 Ri $e RE A RR RR t s e Se A anae
f: $6 LM
5$1 np —»—————— xd
15$ A oem an Dd
355 CES OR urnam ars ce aa
$i LS Ty
$ Ry 9*1
Ed 3 LE
RA $2 Re 3!
Re $5 w3:
F 2
Res Wis
a b
Fig. 1. a: diagrammatic representation of the relative diameters of seedlings
germinated on Type I solution. Series 1; b: relative diameters of seedlings germinated
on Type I solution. Series 2.
SI $55!
$551 Ris
RISI Ysi
Rasi PE rs nena ee ae ae a ee A
WSI MSI
LIETI M $2
$3$2 RIS
Rass 1252
2$2 f4$32
$52 RSS
Rode Red
RES! $353
R233 R233
RAS3 T $3
Riss Riss
a
Fig. a: diagrammatic — of the relative diameters of seedlings
germinated on Type IV solution. Series 1; b: relative diameters of seedlings germi-
nated on Type IV solution. Series 2.
Figures 5 and 6 show the areas of the triangles producing greatest,
medium, and least growth, the circles being drawn as before to
indicate the average diameters of the seedlings of the respective
cultures.
1933]
WYND—ORCHID SEEDLINGS AND NUTRITIONAL IONS 231
TABLE VI
GROWTH OF SEEDLINGS IN TYPE IV SOLUTION. THE FIGURES
REPRESENT DIAMETERS IN MICRONS
Solution Series 1 Series 2
R1 S1 920 990
S 825 825
S3 550 595
R2 Sl 955 950
S2 740 740
S3 635 665
R3 S1 965 965
S2 765 770
S3 750 690
R4 S1 1010 950
S2 730 730
S3 570 630
R5 S1 975 1035
S2 715 715
R6 S1 675 705
IV. Discussion
Examination of figs. 3 and 4 shows perfect agreement between
the duplicate series of Type I in the areas of greatest growth.
These areas are characterized by having very low concentrations
of KH;PO, The five best cultures in row 1 have only one-eighth
of the total osmotic concentration due to this salt. The ratios
of calcium nitrate to magnesium sulphate are seen to vary be-
tween wide limits, .1667 to 2.500, with no significant effect on
growth. The areas of medium and least growth do not show such
perfect agreement in the duplicate series, but a comparison shows
that the areas of medium growth correspond in general to those
having the medium KH;PO, concentration, and the areas of least
growth correspond to those having the highest proportion of this
salt. This relation is particularly clearly shown in fig. 4. From
these data one may be led to the conclusion that germinating
orchid seeds and young seedlings are comparatively sensitive to
[Vor. 20
232 ANNALS OF THE MISSOURI BOTANICAL GARDEN
varying concentrations of KH;PO,, but relatively insensitive to
wide variations in Ca(NO;); and MgSQ,.
Since the ions were present as salts, it is impossible to determine
which ion is responsible for the result. It is possible to add the
six ions in but one other combination of three salts so that each
anion will be linked with a different cation. This combination
is represented by the Type-IV triangle. It is unfortunately
impossible to prepare the complete Type-IV series because at
the pH used (5.0) the solutions having the higher Ca(H;PO;),
concentrations precipitated on sterilization due to the decom-
position of this compound.
(EP v.
JA QU
K Ha Po,
Fig. 3. Type I, series 1, showing Fig. 4. Type I, series 2, showing
areas of high, medium, and low yield. areas of high, medium, and low yield.
Culture R2 S1 omitted because of Culture R2 $1 omitted because of
Aspergillus contamination. Aspergillus contamination.
Growth on the possible solutions of this type shows a remark-
able result. The areas of greatest growth perfectly agree and
are characterized by solutions having the minimal proportion of
the total osmotic concentration due to Ca(H;PO). Again, the
ratios of the other salts, KSO, to Mg (NOs). vary between wide
limits, .1667 to 2.500, with very little effect on growth. The
areas of medium growth correspond to those of medium concen-
tration of this salt, and those of least growth correspond to the
areas of high concentration of this salt.
This comparison of the growth on solutions of Type I with
that on Type IV indicates that in both cases greatest growth was
1933]
WYND—ORCHID SEEDLINGS AND NUTRITIONAL IONS 233
associated with the lowest proportions of the phosphate salt,
irrespective of which ion it accompanied, and that the relative
proportions of all other ions were of comparatively little effect.
The fact that growth was inversely proportional to the concen-
tration of the H:PO, ion seems to be related significantly to the
results of Gregory (28). This author studied the effect of the
different ions on growth by an ingenious mathematical treatment
of the results published previously by Johnston (’24) concerning
the growth of potato plants in three-salt solutions. In studying
duplicate series, Gregory found that in each the fraction of the
total growth produced by the H;PO, ion was a negative quantity.
Fig. 5. Type IV, series 1, showing Fig. 6. Type IV, series 2, showing
areas of high, medium, and low yield. areas of high, medium, and low yield.
The cultures of high concentration The cultures of high concentration
Ca(H;PO,) omitted (see text). Ca(H2PO,)2 omitted.
In general, he also found that the growth produced by other
negative ions was much less than that produced by the positive
ions.
If we indicate the concentration of each ion in a “molar”
sense and compare the composition of the best solution of Type I
and Type IV in table vi, an interesting analogy is apparent.
Despite the great apparent variation in composition of these two
solutions, we see that in both the concentrations of the negative
ions are of the same order of magnitude, while those of the positive
ions vary greatly. It seems therefore probable that growth was
[Vor. 20
234 ANNALS OF THE MISSOURI BOTANICAL GARDEN
being conditioned in both cases by the negative rather than the
positive ions.
TABLE VII
CONCENTRATION OF IONS IN “MOLS" OF THE BEST SOLUTION
OF TYPE I AND TYPE IV
Type I Type IV
H3PO,7 .0025 .0038
NO; . 0098 .0078
SO, - 0123 0097
K* 0025 0194
Ca* * 0049 0019
Mg* * 0123 0039
These results need not be interpreted as contradictory to those
of Smith (732), who found that better growth was obtained in
solutions having larger amounts of ammonium sulphate added to
Knudson's solution B. Smith added nitrogen in the ammonium
ion, while we used the nitrate ion. It is entirely possible that
orchid seedlings might react favorably to an increase of nitrogen
as the ammonium ion, while growth would not improve with
increases in the form of nitrate. This is indeed made very prob-
able by the work of Burgeff ('09), who found better growth with
ammonium than with nitrate salts.
We are at a loss to relate these results to those of La Garde
(29) who states, in regard to superiority of his solution L over
that of Knudson’s solution B, as follows:—'' The seedlings ap-
peared darker in color and had progressed relatively further in
development. This effect might be ascribed to the larger dose
of phosphorus and potassium.” Experiments now in progress,
which will be reported later, show that La Garde’s solution L is
indeed a very favorable orchid medium, but it seems probable that
we may not ascribe this quality to the increase of phosphorus
and potassium per se.
This study further emphasizes the suggestion made by some
authors that the Ca:Mg ratio is not of such importance as it
was once thought. At least the limits of variation used in our
experiments were not sufficiently great to produce any significant
effect.
1933]
WYND—ORCHID SEEDLINGS AND NUTRITIONAL IONS 235
V. SUMMARY
Seeds of Cattleya Trianae Linden and Rchb. f. were germinated
on three-salt solutions of Type I and Type IV, with the following
results :—
(1) The seedlings showed best development on media having
low proportions of the total osmotic concentration due to the
phosphate salt, irrespective of whether it was present as the
potassium of the calcium compound.
(2) The proportion of all other ions within the limits used
appeared to have relatively little effect on germination and growth.
(3) The negative ions appeared of greater importance in con-
ditioning orchid seedling development than the positive ions.
(4) The best growth in Type I solutions was found in culture
R1 S2 having the composition :—
OT ee aurea ad 0025 M
à po eM 0049 M
Mg... oe sro xr Pe ee rae 0123 M
(b) The best growth in Type IV solutions occurred in culture
R5 S1 containing :—
BBG. cen sons 0s Xx cs chee .0097 M
Ca(H;PO4s. . eee cea lior ey .0019 M
Mg(NO3s. ...... eere rni rt rrr 0039 M
VI. ACKNOWLEDGEMENTS
The author wishes to thank Dr. George T. Moore, Director of
the Missouri Botanical Garden, for furnishing seeds for this
investigation; and also to express his appreciation to Elinor Al-
berts Linder, formerly Orchidologist to the Missouri Botanical
Garden, for helpful suggestions while employed at this institu-
tion; and also to Dr. E. S. Reynolds, under whose supervision the
work was carried out.
[Vor. 20
236 ANNALS OF THE MISSOURI BOTANICAL GARDEN
BIBLIOGRAPHY
Schreinemakers, F. A. H. ('93). Konzentrierung oder Verdünnung einer Lösung bei
nter Temperatur. Zeit. Phys. Chem, 11: 81-85. 1893.
Bancroft, A. D. (02). Synthetic analysis of solid phases. Jour. Phys. Chem. 6:
178- 1902
Burgeff, H. (09). Die Wurzelpilze der Orchideen; ihre Kultur und ihr Leben in
flanze. Jena, 1909.
Schreiner, O., and Skinner, J. J. (10). Ratio of DA nitrate, and potassium
in absorption and growth. Bot. Gaz. 50: 1-30. 1910.
Shive, J. W. (15). A three salt nutrient solution *, plants. Am. Jour. Bot. 2:
157-160. 1915.
Wilson, J. K. (15). Calcium hypochlorite as a seed sterilizer. Ibid. 420-427. 1915.
McCall, A. G. (16). Physiological balance of nutrient solution for plants in sand
cultures. Soil Sci. 2: 207-253. 1916.
Shive, J. W. (17). A study of physiological er 2 buckwheat in three-salt
solution. N. J. Agr. Exp. Sta. Bull. 319: 917.
— — ——, and Martin, W. H. (18). A decre Ped of salt requirements for
pe: “and for mature buckwheat plants in solution cultures. Jour. Agr. Res.
: 151-175. 1918.
— — ——, (18a). A comparison of salt requirements for young and for
mature buckwheat plants in water culture and sand culture. Am. Jour. Bot. 5:
186-191. 1918.
Livingston, B. E., and Tottingham, W. E. (18). A new three-salt nutrient solution
for plant cultures. Ibid. 337-346. 1918.
, (19). A plan for comparative research on the salt requirements of repre-
sentative agricultural plants. Baltimore, 1919.
Van Alstine, E. (19). The relation of salt proportion to the growth of wheat in sand
cultures. N. J. Agr. Exp. Sta. 40: 366-374. 1919.
Martin, W. H., and Shive, J. W. (’20). A study of the salt requirements of the
potato. N. J. Agr. Exp. Sta. Rept. 41: 409-412. 1920
Meier, H, F. A., and Halstead, C. E. (21). Hydrogen-ion eiticntitwiibt relations
in a ihreoquit solution. Soil Sci. 11: 325-352. 1921.
Tottingham, W. E., and Rankin, E. J. (22). Nutrient solutions for wheat. Am.
Jour. Bot. 9: 270-276. 1922.
Knudson, L. (22). Non-symbiotie germination of orchid seeds, Bot. Gaz. 73:
1-25. 1922.
Poole, R. F., and Fant, G. W. (22). Further study of the relation of various
fertilisers mixtures to the growth of celery in muck soil. N. J. Agr. Exp. Sta.
Rept. 43: 395-399.
Clement, E. (24). Germination : E TU and other seed without fungal
aid. Orchid Rev. 32: 233-2
— ———, (248). The apa odia of orchid seeds. Ibid. 359-
366. 4.
Ballion, M., and Ballion, G. (24). The non-symbiotic germination of orchid seeds
in Belgium. Jbid. 305-309. 1924.
Johnston, E. S. (24). Growth of potato plants in sand cultures treated with the
“six types" of nutrient solutions. Md. Agr. Exp. Sta. Bull. 270: 53-86. 1924.
Davidson, O. W. ('28). An application of the triangle system in determining a
nutrient solution suitable for research with the peach in sand culture. Am.
Soc. Hort. Sci., Proc. 25: 354-358. 1928.
1933]
WYND—ORCHID SEEDLINGS AND NUTRITIONAL IONS 237
Gregory, F. G. (’28). The differential effect of the ions of three-salt solutions on the
growth of potato plants in sand cultures. Roy. Soc. Lond., Proc. B. 102: 311
1928.
La Garda; R . 29). Non-symbiotic germination of orchids. Ann. Mo. Bot. Gard.
Quednow, K. G. (30). Beiträge zur Frage der Aufnahme gelóster Kohlenstoffver-
bindungen durch Orchideen und andere Pflanzen. Bot. Archiv. 30:51-108. 1930.
Addoms, Ruth M., and Mounee, F. C. (31). Notes on the nutrient requirement
and the histology of the cranberry (Vaccinium macrocarpon Ait.) with special
reference to mycorrhiza. Plant Physiol. 6: 653-669. 1931.
Smith, F. E. V. (32). Raising orchid seedlings asymbiotically under tropical
conditions. Gard. Chron. 91: 9-11.
Annals
of the
Missouri Botanical Garden
Vor. 20 APRIL, 1933 No. 2
A REVISION OF THE NORTH AMERICAN SPECIES
OF VERBENA!
LILY M. PERRY
Instructor in Botany, University of Georgia, Athens, Georgia
Formerly Rufus J. Lackland Research Fellow in Botany
in the Henry Shaw School of Botany of Washington University
INTRODUCTION
Verbena, the type-genus of the family Verbenaceae, is pre-
eminently American. Owing to inherent variability and varia-
tions caused by adaptation to local conditions of growth, the group
is arather complex one. Moreover, since the last comprehensive
monograph of the genus was published so long ago that it contains
only some twenty of the hundred or more described North
American species and varieties, considerable material has been
incorrectly named. It appeared, therefore, that a critical
examination of the group might be helpful in untangling the
existing confusion of identities. The following paper is an at-
tempt to present the results of a study of the North American
species, together with a brief review of the historical, morpho-
logical, and phylogenetic aspects of the species under considera-
tion.
The materials studied are indicated in the text by abbreviations
for the herbaria to which they belong: the Academy of Natural
Sciences of Philadelphia (ANSP); Field Museum of Natural
History (F); Gray Herbarium of Harvard University (G); the
Royal Botanic Gardens, Kew (K); the Missouri Botanical
1 An investigation carried out at the Missouri Botanical Garden in the Graduate
Laboratory of the Henry Shaw School of Botany of Washington University, and
submitted as a thesis in partial fulfillment of the requirements for the degree of
doctor of philosophy in the Henry Shaw School of Botany of Washington University.
Issued July 10, 1933.
Ann. Mo. Bor. Garb., Vor. 20, 1933. (239)
[Vor. 20
240 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Garden (MBG); New York Botanical Garden (NY); Pomona
College (P); United States National Herbarium (US); photo-
graphs of types and authentic specimens have been received from
the Museum of Natural History, Paris (Par.); the Herbarium of
the Botanical Garden and Museum, Berlin-Dahlem (Bot. Mus.
Berl.-Dahl.), and the Herbarium of the Botanical Garden, Madrid
(Herb. Bot. Gard. Madrid).
For the photographs and the use of the loans, as well as for the
many courtesies received while visiting different herbaria, the
writer is truly grateful and hereby acknowledges her indebtedness
to all who have given assistance in this way. She expresses her
hearty thanks and cordial appreciation to Dr. George T. Moore,
Director of the Missouri Botanical Garden, for the use of the
extensive privileges of the excellent library and the herbarium.
She is also indebted to Miss Nell Horner, Librarian, for any
suggestions in bibliographical matters. To Dr. J. M. Greenman,
who has directed and supervised the work, giving ready aid and
helpful suggestions at all times, the writer expresses her particular
gratitude for the encouragement and personal interest shown
throughout this study.
Taxonomic HISTORY
The derivation of the word Verbena is obscure. The name has
been handed down to us as representing certain herbs used in
sacred rites by the ancients. Although apparently one of the
well-known plants in botanical history, Verbena was described
comprehensively for the first time by Tournefort,! whose descrip-
tion formed the basis of the synopsis of the genus as accepted by
Linnaeus.’
In the ‘Species Plantarum,’ Linnaeus? records fourteen species
of which eight have been removed since to other genera. The
latter fact is sufficient to indicate that the Linnaean concept was
too inclusive. It was held, however, until 1806, when Jussieu,‘
in an article on ''Observationes sur la famille des plantes
1 Tournefort, Institutiones Rei Herbariae, 200, pl. 94. 1700.
2 Linnaeus, Gen. Pl. 12. 1754.
3 Linnaeus, Sp. Pl. 18-21. 1753.
* Jussieu, Ann. Mus. Paris 7: 71-73. 1806.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 241
Verbénacées," pointed out that the genus was a composite one
containing several units. Amongst these he showed that Verbena
differed from the others in having four stamens and four nutlets.
This definition was accepted by Endlicher* under the heading
'" VERBENAE VERAE," and it has continued to the present time.
Although the genus is predominantly American, the earlier
species were described largely from plants grown in various
European gardens. ‘This was unfortunate, since such types are
often extinct or inaccessible or sometimes difficult to interpret
on account of probable fusion or other modifications appearing
in the species after some few years of cultivation. The first
treatment of the North American species, as such, is that of
Michaux.® He recognized eight species, all from the region of
the middle and southeastern United States. From time to time
new species were collected by pioneer explorers. Kunth’ de-
scribed eight from the Mexican region, Bentham: added three
more, and Martens and Galeotti® reported eleven, of which six
were new. ‘The ablest and most inclusive work on the genus as
a whole is that of Schauer.!° In this he accepted seventy-one
species and listed fourteen more as insufficiently known; of these,
twenty-seven are indigenous to North America. Appearing
about the same time as Schauer’s work is that of Walpers,1
whose review might be considered as an assembling of the litera-
ture of the genus rather than the results of a special monographie
study; nevertheless, it is comprehensive and contains a few new
species. Subsequently no publication of the North American
species ?n toto has appeared, although the works of Gray” and
Small! are very helpful in so far as regional floras are concerned.
The last two publications contain the only keys to the group
under consideration.
The type-species of the genus is V. officinalis L.
5 Endlicher, Gen. Pl. 633 m 3085). 1838.
6 Michaux, Fl. Bor.-Am. 2: 18-15. 1803.
1 HBK. Nov. Gen. et n 2: 272-271. 1818.
8 Bentham, Pl. Hartw. 21. 39.
? Martens & Galeotti, Bull. Acad. Brux. 11?: 320-324. 1844.
10 Schauer in DC. Prodr. 11: 535-556. 1847.
1! Walpers, Rep. 4: 13-33. 1844-48; 6: 686-687. 1846-47.
? Gray, Syn. Fl. N. Am. 2!: 335-338. 1878.
13 Small, Fl. Southeast. U. S. ed. 1, 1008-1011. 1903.
[Vor. 20
242 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Gross MORPHOLOGY
Roots.—Some species are annual with a fibrous root system
branching from a slender primary root; others are perennial with
a larger tap-root. ‘They maintain themselves from year to year
by crowns from which clustered new shoots rise near the base of
the old ones. A few species show a tendency to root at the nodes.
Probably, for the most part, the main purpose of these secondary
roots is to aid in obtaining nourishment for the plant, as instance,
V. canadensis, a hardy plant with stems rooting only at the lower
nodes. Nevertheless, in V. teucriifolia, they may be a possible
means of propagation, as definite roots are found at more or less
remote nodes; occasionally, both roots and fruiting spike are
developed from the same node. In many cases, it is difficult to
tell whether a species is an annual or only a short-lived perennial.
The question of duration depends in some measure on climatic
factors.
Stems.—' l he stems are slender, terete, or, more commonly,
quadrangular in cross-section with sharp or obtuse angles, and
chiefly pubescent. Apparently all are herbaceous, the perennials
dying down to the ground in winter and reappearing again in the
spring. ‘The shorter forms branch freely with decumbent-
ascending or prostrate branches; the taller are usually erect with
branches loosely ascending-spreading. All the lower branches
are opposite, but the upper may be somewhat irregularly placed.
Leaves.—In the majority of the species of the section Glandu-
laria the leaves are deeply cleft, whereas those of the Verbenaca
are in large part merely dentate or incised. As a whole they are
membranaceous, varying greatly in size and somewhat in thick-
ness. In a few species they are strictly sessile; in most, however,
they are either subsessile or short-petioled and tend to be strongly
veined beneath. All have more or less short, sheathing, and
connate bases. In general the leaves are opposite, but at times
they are subalternate or irregularly disposed on the upper part
of the stem; they are rarely ternate, and this arrangement appears
to be an individual expression rather than a specific or even a
varietal character.
Pubescence.—The trichomes are simple and unicellular, varying
in quantity, length, rigidity, coarseness, and direction. None
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 243
of the pubescence is truly hispid, but in some species the hairs
are slightly stiffer than in others; this indument, as exemplified
in the calyx of V. bipinnatifida, is designated as ‘‘hispid-hirsute.”’
In V. prostrata and V. Gooddingii the pubescence is character-
istically soft-villous, but in the majority of species the general
tendency is to approach a hispidulous-hirsute to strigillose
condition. In the former instance, the trichomes, especially
along the stem and on the inflorescence, are of different lengths
and more or less spreading; whereas in the latter, as in V. tam-
pensis, they are short, approximately equal, and appressed.
Often the hairs of the upper surface of the leaf spring from
minute bulbous bases; these, according to Solereder," are calcified
or silicified hairs, and the trichomes with somewhat pustulate
bases, in V. carolina and occasionally in other species, are cystolith-
hairs. In any case, Solereder did not find cystoliths dissociated
from the hairs. A viscid condition is rarely present, but glandular
hairs are widely distributed throughout the genus.
Inflorescence.—The inflorescence is terminal at the apices of
stems or branches. The flowers are subtended by bracts and are
borne in solitary or somewhat panicled or cymosely arranged
spikes. Although these vary considerably, it is fairly easy to
determine the section to which a plant belongs by the type of
its spike.
In the majority of the indigenous North American species of
the Verbenaca, the spikes are elongated and tend to be irregularly
placed, as in Verbena carolina, so that they appear more or less
panicled. In V. sphaerocarpa and V. litoralis, however, they are
commonly arranged in simple or compound cymes. A spike
may be either very slender with the flowers somewhat remote, or
dense with imbricated flowers. In general, the elongated and
comparatively narrow spike with small and inconspicuous flowers
is a distinctive feature of the Verbenaca native to our continent.
On the other hand, the spikes characteristic of the section
Glandularia are shorter and relatively broad with larger and more
showy flowers. In the earlier stages of anthesis these are in
fascicle-like clusters; later, when the rhachis elongates, the in-
florescence becomes spicate.
14 Solereder, Syst. Anat. Dicot. ed. 2, 1: 631-633. 1908.
[Vor. 20
244 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Calyx.—The calyx is tubular, 5-lobed or subtruncate, 5-
nerved and persistent. The nerves protrude somewhat beyond
the margin of the lobes, forming calyx-teeth; these vary more or
less specifically and occasionally intraspecifically. The nerves
are unequal in length and the posterior one, along which the
calyx ruptures to release the schizocarp, is always the shortest.
In the Glandularia, the calyx is at least twice as long as the
schizocarp and at maturity the teeth and lobes are more or less
contorted. In contrast, the calyx of the Verbenaca is generally
shorter and may be either connivent, concealing the schizocarp
as in V. scabra, or, as in such species as V. urticifolia and V.
officinalis, open, disclosing the apices of the nutlets. With the
developing ovary, the calyx enlarges a little at the base; usually
it is somewhat angled, but in V. tumidula it is round at the base
and tends to be inflated. Although only relative in value, the
pubescence of the calyx is probably of more worth in specific
determination than the other previously mentioned characters.
Corolla.—' The corolla is salverform and various in color. The
tube is either straight or incurved, as long as or much longer than
the calyx, and uniform or slightly enlarged at the throat. The
degree of the exsertion of the corolla-tube is apparently a conveni-
ent and obvious character, but one to be handled with care;
first, it is somewhat variable, and second, when the corolla falls
after anthesis, it does not drop at once but may be some little
time working its way out of the calyx-tube; hence, the length of
the exserted portion appears much greater than it really is. The
limb is lobed and may be either conspicuous (6-12 mm. broad) or
inconspicuous (2 mm. broad). The shape and the direction of
the lobes may be of value in field identification but are not at all
satisfactory in work on herbarium material. The throat inside
is bearded with straight minutely roughened retrorse hairs and
at the orifice with moniliform ones; outside and beyond the calyx
it is either glabrous or finely pubescent. The inner hairiness
is more prominent in the section Glandularia.
Stamens.—The stamens are didynamous, often very short-
stalked, and in the North American species, inserted on the upper
middle half of the corolla-tube, but not exserted. In V. bipin-
natifida and all the related species of its section, the anterior pair
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 245
is very near the orifice of the throat and usually bears a gland-
like appendage on the connective. The stamens are too much
alike to be useful in specific differentiation.
Pistil.—The ovary is superior, bicarpellary, and entire or very
shallowly lobed at the apex. The style varies in length with the
length of the corolla-tube, bearing the mature stigma approxi-
mately in the region of the lower anthers. The typical form is
slender and bilobed, the posterior lobe being sterile and laterally
compressed, the anterior stigmatic and broad. The relation of
these lobes to each other is reasonably constant throughout each
section of the genus. In V. scabra, however, the broad stigmatic
surface appears to be subtended on either side by a sterile style-
lobe. Another interesting variation of the style is produced by
V. quadrangulata; here, the base is enlarged or thickened into a
somewhat hemispherical-angulate stylopodium and persists as
an integral portion of the nutlets.
Fruit.—The fruit of Verbena is a schizocarp enveloped by the
persistent calyx. As previously stated, the calyx splits along the
posterior nerve to release the schizocarp attached to a more or
less round-quadrangular gynobase. For the most part, four
nutlets or pyrenae develop, two from each biovulate carpel.
Generally the nutlets are either subcylindrie or subtrigonous,
but in V. quadrangulata they are beaked cylindric, and in V
tumidula subovoid. The dorsal surface is more or less convex
with the lateral surfaces scarcely definite, rather appearing as
continuations of the dorsal portion and adjoining the commis-
sural face or faces. This is not always the case, since in V.
tumidula the lower part of the nutlet seems inflated and the
lateral is somewhat ventricose and smooth. The surface of the
nutlet varies from essentially smooth, as in V. carolina and its
allies, to striate or sulcate, and is oftenest raised-reticulate or
reticulate-scrobiculate on the upper part. Occasionally, the
character or scope of these depressions is of specific value, as in
V. gracilis; but usually a certain type of nutlet is characteristic
of a group of species rather than a single one.
The commissure may be more or less muricate or practically
smooth. In the subcylindric pyrenae of the section Glandularia,
the commissural face is commonly convex and does not extend
[Vor. 20
246 ANNALS OF THE MISSOURI BOTANICAL GARDEN
to the tip of the nutlet; whereas in the trigonous nutlets of the
Verbenaca, the commissure ordinarily has two faces meeting at
right angles, and is as long as the nutlet.
The gynobase varies with the size and shape of the pyrenae
which it bears. In V. carolina and V. Ehrenbergiana, it is nearly
flat and almost orbicular, but in species with larger nutlets, as in
V. simplex and V. neomexicana, it is slightly deeper and in large
part somewhat quadrangular. It shows its greatest develop-
ment in the section Glandularia where the pyrenae are largest
and often somewhat broadened at the base. Perhaps the gyno-
base should be used as a character complementary to that of the
nutlets, but its definite value is rather elusive.
The more important morphological characters, which have
been used in the delineation of the genus and its sections, are
illustrated with a legend (pl. 13).
Hybrids.—It is a well-accepted fact that the cultivated races
of Verbena have arisen through the more or less fusing of species
attractive for horticultural purposes. Granting this, without
field observation and knowledge gained through experimental
work, the writer does not propose to discuss the question of
hybridization in Verbena. Nevertheless, it is necessary to point
out that there is a considerable number of specimens amongst the
collections of this genus from the Middle States, which show
variation in several directions or a combination of the characters
of two or more species; it is the prevailing opinion that these
plants are hybrids.
RELATIONSHIPS AND DISTRIBUTION
The genus Verbena is a member of the tribe Verbeneae accord-
ing to Engler and Gilg,5 or of the Huverbeneae according to
Briquet in Engler and Prantl.^ It is distinguished from its
immediate relatives by the four dry and hardened nutlets. In
this paper fifty-one species are recognized, four naturalized and
forty-seven indigenous. With the exception of a few border-line
species, it is relatively easy to separate the genus into two fairly
distinct sections in which the majority of the members are so
5 Engler & Gilg, Syllab. der Pflanzenf. eds. ; AJ 10, 339. 1924.
Engler & Prantl, Nat. Pflanzenf. 4?*: 146.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 247
different that it would seem as if they had followed divergent
trends for a long period of time. In attempting to present any
genus in à natural order, it is highly desirable to know what
species are primitive. However, without an intimate knowledge
of the more diversified South American species, it seems prefer-
able to state the evidence derived from the study of the North
American group and leave the conclusions vague rather than more
or less arbitrary.
The section Verbenaca is probably the older of the two. It
has the wider distribution, ranging from Canada to Panama as
well as from the east to the west coast of United States and Mex-
ico. Also, it is larger, containing thirty-two species. A number
of these are annual, the remainder are perennial, but it is often
hard to distinguish the members by this criterion. Although
admittedly varying and apparently intergrading, the species as
a whole are more stable and more readily recognized than those
of the section Glandularia. The flowers are much smaller and
for the most part are produced in greater abundance; the nutlets
also are abundant and in some cases minute. The combination
of these characters leads one to conclude that this section is the
more primitive one.
Within the section, some species appear to be more closely
related than others. V. officinalis, V. menthaefolia, V. Halei,
and V. riparia form a natural group possessing the same general
habit, type of pubescence and of nutlets. These are species of
more or less adjacent regions except V. officinalis, which has
been introduced from Europe. Again, V. Ehrenbergiana, V.
carolina, V. longifolia, and V. recta are very closely related, with
V. urticifolia and V. scabra only a little farther removed. The
last five species differ from the first mentioned in their more or
less upright habit and less divided leaves, also in their smaller
and somewhat smoother nutlets. It would appear as if all had
arisen very close to the ancestral form and diverged only a little
in the process of adaptation to environment.
V. hastata and V. simplex are quite distinctive in habit, but the
abundance of intermediate forms occurring between these and
the two species, V. stricta and V. bracteata, would seem to indicate
a close relationship amongst these units. V. Orcuttiana, although
[Vor. 20
248 ANNALS OF THE MISSOURI BOTANICAL GARDEN
somewhat isolated, appears to have its nearest affinity in V.
simplex.
A third closely related group contains V. canescens, V. neo-
mexicana, V. plicata, and V. gracilis; the common characters
here are those of leaf-incision, pubescence, and general features
of inflorescence. V. perennis resembles this group in all respects
except in its quite aberrant pubescence of short antrorse and
hispidulous trichomes; in this character it simulates more closely
the first-mentioned group.
The following are more or less isolated species of limited
distribution and with fairly static characters: V. xutha, V.
prostrata, and V. robusta. V. carnea is probably a relic of some
ancient form; the distinctive character of the schizocarp gives
no clue to its affinities, but rather emphasizes the anomaly of the
species.
The section Glandularia seems to be the more nascent. The
species are perennial, developing from a crown, and have a more
or less sprawling to upright habit. The characters are highly
variable and some species are difficult to distinguish, apparently
passing into each other. Particularly is this true of V. ambrosi-
folia, V. Wrightii, and V. ciliata. Some of the specimens from
northern Mexico have been most perplexing; while it is perfectly
evident that they belong to this group, none of them is typical.
V. bipinnatifida is a closely related and wide-ranging species;
by comparison, its characters are fairly clear-cut. The remaining
species of the section are easily associated in groups or are
anomalous. V. canadensis, V. maritima, and V. tampensis are
very much alike either in habit or in floral structure. V. tam-
pensis, however, is one of the species of this group where the
glands may or may not be present on the anthers, and may
represent the development of some transitional stage between
the two sections. V. elegans of Mexico is very closely parallel to
V. canadensis of the southeastern and central United States. V.
delticola seems to belong in this association also; yet it is interest-
ing to note that its fruit shows a tendency to develop a beak,
and this may be significant of some little affinity with V. quad-
rangulata, a species so aberrant that it has been treated as a
separate genus. V. Gooddingi V. tumidula, and V. pumila
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 249
might very well be related to any of the previously mentioned
species of the section. V. setacea resembles V. Gooddingw var.
nepetifolia in habit, but it has such long calyx-teeth that one is a
little hesitant about expressing any ideas of its relationship. V.
teucriifolia and V. ciliata appear to intergrade in Mexico. The
former, as well as V. pumila, shows a tendency toward reduced
corollas and perhaps parallel development. V. lilacina and V.
amoena, are without close relatives.
Considering the genus as a whole, some species are in a state
of flux showing many atypical forms and covering fairly large
geographie areas; others are just as widespread and yet are
characterized by more static characters; and again, others are
endemie showing a very limited distribution and appearing as
outlying members of the group. The species probably vary
from youth to age in the order mentioned.
As previously stated, the genus Verbena is fairly widely
distributed in North America; its range extends from southern
Canada to Panama and from the east coast to the west both in
United States and in Mexico. It is found also in the West
Indies and on islands in the Pacific off the coasts of California
and Mexico. It occurs in great abundance in Texas, New Mexico,
and Arizona, south along the Sierra Madre and throughout the
eastern part of Mexico. It is also fairly well distributed in
southern Mexico, but only a few species are reported from
Central America. The region of greatest specific concentration
for North America is in the state of Texas, where twenty out of
the forty-seven indigenous North American species are rep-
resented. Amongst these species are members of both sections,
some with very definitely clear-cut characters and more or less
limited distribution, others with fluctuating characters and more
widely distributed.
By way of contrast, let us consider the outlying members of
the genus which have stable characters and are isolated or in-
habit very limited areas—V. lilacina, V. sphaerocarpa, V. Or-
cuttiana, V. setacea, V. macrodonta, V. amoena, and V. subuligera;
or again species such as V. carnea and V. quadrangulata, which
are singular but have a larger geographic range. All these
appear to be relies of a more ancient distribution. Undoubtedly
[Vor. 20
250 ANNALS OF THE MISSOURI BOTANICAL GARDEN
V. lilacina and V. sphaerocarpa, endemics of Cedros and Socorro
Islands respectively, developed as a result of geographic isolation.
The same may be true of the Lower Californian entities, V.
macrodonta, V. setacea, and V. Orcuttiana. Perhaps V. amoena
from Lecheria, a locality some little distance north of the city of
Mexico, and V. subuligera, from the Sierra Madre west of Durango,
owe both their restriction and their differentiation to the presence
of mountainous barriers. However, the same conclusion is
hardly valid for either V. carnea, ranging from North Carolina
to Florida and west to Texas, or V. quadrangulata, of southern
Texas and Tamaulipas; yet they are probably relic endemics
which have survived in favoring areas. If this be true, here is
another bit of interesting evidence supporting the idea that the
Verbenaca is the older of the two sections. Of the nine species
enumerated above as being of more ancient derivation, four, V.
setacea, V. lilacina, V. amoena, and V. quadrangulata, have been
placed in the section Glandularia; yet, in the flowers examined,
the anthers are definitely unappendaged. To be sure, this
disposition of the species is not in accord with the previous
classifications and may not be acceptable; nevertheless, it seems
logical to believe there must be transitional forms in the develop-
ment and evolution of species; and in this case, the major portion
of the characters of the species in question are those of the
section Glandularia. Two other species which have developed
a little farther along this same line are V. tampensis and V. pumila;
gland-like appendages may or may not be present on the anthers;
if present, they are usually very small.
It may be of interest to note that, in agreement with Jordan’s
law of distribution, the species associated in groups usually
occupy adjacent regions.
Now as to origin.—It is possible that the progenitors of Verbena
were very widespread at the beginning of the Pleistocene and
with the glacial advance migrated southward; later, when more
favorable climatic conditions developed they spread northward,
establishing themselves in the present regions of distribution.
Another possibility is that the centre of distribution is in South
America and migration has been northward. However this
may be, V. litoralis is the only known species indigenous to both
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 251
countries and does not seem to be very closely related to the
remaining North American species. Without a detailed in-
vestigation of the South American group, it seems preferable to
reserve assumptions in the matter.
Economic VALUE
Verbena is of very little value economically. It was introduced
into cultivation in European gardens relatively early in the his-
tory of botanical explorations. Although not so popular as it
was once, on account of its vigor and hardiness, it is still used in
many gardens to impart bright colors to the landscape.
TAXONOMY
Verbena [Tourn.] Linn. Gen. Pl. 12. 1754.
Obletia Rozier, Jour. Phys. 1: 367, pl. 2. 1773.
Glandularia Gmel. Syst. Veg. 920. 1791.
Billardiera Moench, Meth. 369. 1794.
Shuttleworthia Meisn. Pl. Vasc. Gen. 290. 1839, and Comm.
198. 1839.
Uwarowia Bunge, Acad. St. Petersb. Bull. Sci. 7: 278. 1840.
Calyx tubular, 5-toothed, 5-nerved, at maturity often slightly
enlarged at the base, persistent. Corolla minute to conspicuous,
hypocraterimorphous, subbilabiate, 5-lobed; tube erect or in-
eurved, uniform or slightly enlarged above, equalling the calyx-
lobes or surpassing them; lobes imbricate in the bud, broadly
oblong, obtuse or retuse, unequal. Stamens 4, didynamous,
included; anthers ovate, unappendaged or the connective of the
upper pair expanding extrorsely into a clavate and gland-like
appendage; filaments usually short, attached above the base of
the corolla-tube. Ovary superior, entire at apex or very shortly
4-lobed, bicarpellary, 4-loculed; ovules in pairs, one to each
locule, anatropous; style terminal, mostly bilobed, the anterior
lobe stigmatose and the posterior usually sterile. Schizocarp
included in the persistent calyx, dry, hardened or brittle, separat-
ing into 4 homomorphous nutlets. Seed erect; embryo straight;
endosperm none.—Chiefly American. Herbs or shrubs, erect or
decumbent to prostrate, pubescent or occasionally glabrous.
Leaves opposite, rarely ternate or verticilate, or the upper
[Vor. 20
252 ANNALS OF THE MISSOURI BOTANICAL GARDEN
alternate (irregular), variously dentate, incised or dissected, not
often entire. Spikes terminal, rarely axillary, peduncled or
sessile, densely flowered or elongate and slender with flowers
remote, solitary, cymose or paniculate. Flowers hermaphroditic,
zygomorphic, sessile and bracteate.
SECTION 1. VERBENACA Schauer
1. VERBENACA Schauer in DC. Prodr. 11: 536. 1847.
Sterile style-lobe adjacent to the stigmatie surface or sub-
lateral but usually not protruding beyond it; ovary entire at the
apex with the style attached at the distal end, or, if at all lobed,
style inserted in the indefinite depression between the lobes;
schizocarp commonly not constricted along the lines of cleavage;
anthers unappendaged. Annuals or herbaceous perennials with
prostrate, decumbent-ascending or erect stems. Flowers mostly
not showy. Calyx rarely more than twice as long as the schizo-
carp and not contorted beyond it. Species 1-32.
Ser. I. PACHYSTACHYAE Schauer. Heads or spikes, at least in
anthesis, crowded and short, not at any time greatly elongated
or open, generally disposed in compound cymes.—Chiefly
introduced South American species.
KEY TO THE SPECIES
A. Leaves semiamplexicaul and subcordat
B. Inflorescence glandular; bracts ditioni longer than the —
corolla-tube 2-3 times as long as the calyx................0... rigida
B. Inflorescence not glandular; bracts barely equalling or only dd. ex-
ceeding the calyx; corolla-tube scarcely twice as long as the calyx. .
Vuarasrumzrag. T S T E S RI eL EROR EVUY. bonarlóits
A. Leaves not semiamplexicaul nor subcordate, tapering into a cuneate-
attenuate subsessile or petiolar base.
C. Spikes short (3-5 mm. long), very dense and appressed-pubescent.
D. Fruiting-calyx ascending; schizocarp longer than broad, raised-
reticulate above, striate toward the base............ 3. V. brasiliensis
D. Fruiting-calyx spreading; schizocarp as broad as long, practically
feat: ad: PIE ESPE E ke oe T RD E 4. V. sphaerocarpa
C. Spikes 5-10 mm. long, dense at the apex, somewhat open bulón & nd
ROAS Pe ii cf ie A te ese o PENNE UN 5. V. litoralis
1. V. rigida Spreng. in L. Syst., Cur. Post. 42: 230. 1827.
V. venosa Gill. & Hook. Bot. Misc. 1: 167. 1830.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 253
V. bonariensis var. rigida O. Kuntze, Rev. Gen. 3?: 255. 1898;
Briq. Ann. Conserv. & Jard. Bot. Genéve 7-8: 291. 1904.
Stems 2-6 dm. tall, sharply 4-angled in cross-section, scabrous-
pubescent; leaves oblong to oblong-lanceolate or narrowly
obovate, sessile, more or less semiamplexicaul, subcordate,
sharply and coarsely serrate, midrib and veins impressed above
but prominently reticulated beneath, scabrous and more or less
hispidulous on both surfaces, margins of older leaves somewhat
revolute; spikes usually short, dense and cylindrical, disposed in
subternate cymes with the laterals pedunculate; bracts lanceolate,
acuminate-subulate, usually closely imbricated and longer than
the calyx, glandular-pubescent, ciliate; calyx 4 mm. long, glan-
dular-pubescent, lobes acute with short mucronate-subulate
tips; corolla-tube twice (-thrice) as long as the calyx, pubescent
without; corolla-limb 5-7 mm. broad; stamens inserted on the
lower middle of the corolla; nutlets trigonous, slightly broader
at the base than at the apex, scarcely 2 mm. long, raised-reticulate
on the upper half, striate toward the base; commissural faces
muricately scabrous.
Distribution: indigenous to South America; introduced in the West Indies and
southern United States.
Specimens examined:
Norts Carona: roadside, Williamston, Martin Co., 4 July 1922, Randolph 688
(G).
Grorata: uncultivated ground, in perennial spreading patches, 26 May 1928,
Reade (NY); vicinity of Thomson, McDuffie Co., 1 Sept. 1907, Bartlett 1122 (P).
ALABAMA: Mobile, Mohr (US); Mobile, June 1919, Graves 525 (MBG, US).
LourisiANA: Mandeville, 15 Aug. 1912, Pennell 4204 (NY); Covington, Aug. 1919,
Arsène (US); Catalpa, 24 Aug. 1912, Pennell 4332 (NY); Baton Rouge, 28 March
1904, Billings 49 (G); roadsides, Plaquemines Co., July 1883, Langlois (NY).
Texas: Orange, 1914, Young 662 (P, US); Huntsville, 24 May 1917, E. J. Palmer
12038 (MBG); Houston, 1872, Hall 433 (G, MBG, NY, P, US); Houston, 1913,
Fisher 460, 625 (US); Houston, 25 Sept. 1915, Thurow (US); Houston, 22 May 1917,
E. J. Palmer 12001 (MBG); 16 km. southeast of Houston, 12 April 1925, Small &
Wherry 11813 (NY).
Mexico: Vera Cruz: Zacuapan, May 1913, Purpus 6413 (F, G, MBG, NY, US).
West INDIES:
BERMUDA: St. George's, 13 July 1905, Moore 2984 (G); near Devil's Hole, 13 June
1905, Harshberger (ANSP, G, MBG); near Hamilton, 31 Aug.-21 Sept. 1905, Brown
& Britton 153 (ANSP); Tucker’s Town, 3 May 1912, Collins 45 (G); Tucker's Town,
8 Aug. 1913, Collins 269 (G, NY); fields, Montrose, Sept. 1913, Brown, Britton &
Wortley 1645 (ANSP, NY); St. David's Island, 22 May-2 June 1919, Brown 693
(ANSP, NY).
[Vor. 20
254 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Cuna: near Farallones, Oriente, 5 Aug. 1913, Leon 3910 (NY).
Jamaica: road to Salt Hill, St. Andrew, 6 May 1915, Harris 11969 (G, MBG, NY);
Blue Mountains, St. Andrew, 20 March 1916, Perkins 1024 (G); near Fairfield, 3-7
Sept. 1908, Britton 3176 (NY).
GuADELOvPE: without locality, 1893, Duss 3470 (NY).
MARTINIQUE: without locality, Sept. 1901, Duss 4697 (NY).
This South American species is commonly cultivated. It has
escaped and established itself in various places. Kuntze and
Briquet have regarded it as a variety of V. bonariensis, but
lacking a considerable amount of material for examination, the
writer prefers, at present, to maintain it as a specific entity.
2. V. bonariensis L. Sp. Pl. 20. 1753.
V. elongata Salisb. Prodr. 71. 1796.
? V. quadrangularis Vellozo, Fl. Flum. 16. 1825; 1: pl. 39. 1827.
Stems 1 m. more or less in height, 4-angled in cross-section,
somewhat scabrous-pubescent; leaves decussately opposite, lance-
olate, subauriculate, semiamplexicaul, acutely serrate, entire
toward the base, rugose and hirtellous above, spreading-pubescent
beneath, particularly along the prominently reticulated veins;
spikes compact, mostly short, commonly sessile and crowded in
dense fasciculate cymes; bracts lanceolate-acuminate, barely
equalling or slightly exceeding the calyx, pubescent; calyx 3 mm.
long, pubescent, lobes acute with short subulate tips; corolla-tube
scarcely twice the length of the calyx, pubescent without; corolla-
limb inconspicuous; nutlets 2 mm. long, trigonous, chiefly striate,
slightly raised-reticulate at apex; commissural faces scarcely
reaching the tip of the nutlet, muricate-scabrous.
Distribution: native of South America; introduced into southern United States
and the West Indies.
Specimens examined:
Sourn CanoLINA: roadsides near Charleston, Curtiss 1963 in part (F, G, MBG,
NY); Charleston Neck, 28 Sept. 1853, Gibbes (NY, US); Charleston, 30 April 1912,
Robinson 127 (G); about 6 km. south of Charleston, 9 Nov. 1929, Moldenke 148
(MBG, NY).
Georata: along canal bank, Augusta, Cuthbert 267, 358 (NY).
LovistaNa: vicinity of Covington, 1920, Arsène 11859, 12534 (F, US).
ARKANSAS: roadsides near Forrest City, 17 Oct. 1925, E. J. Palner 29303 (G, MBG,
NY).
TExas: Huntington, 15 Sept. 1923, Tharp 2658 (US).
West INDIES
BERMUDA: waste ground, 29 March-3 May 1909, Marble 787 (ANSP, NY); Pem-
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 255
broke Swamp, 21 June 1905, pica (ANSP, G, MBG, NY); roadside, Pem-
broke, 31 Aug.-20 Sept. 1905, Brown & Britton 98 (ANSP, G, NY, P); Hamilton,
12 July 1905, Moore 2946 (G); E: Hamilton 16 Jan. 1912, Robinson 94 (G); Tucker's
Town, Collins 58 (G), 266 (G, N
JAMAICA: Blue Mountains, 14 ie 1890, Hitchcock (MBG); Blue Mountains, Mt.
Hybla, 5 Apr. 1916, Perkins 102 (G); St. Helen's Gap, 7 April 1909, Taylor 4244 (NY);
St. Helen's Gap, St. Andrew, 4 March 1920, Mazon & Killip 569 (G, NY); Cinchona,
26 July 1903, Nichols 168 (G, MBG, NY); Cinchona, 22 Dec. 1905, Harris 9132
(NY); Arntully, 25 Aug. 1927, Orcutt 2736 (G, MBG).
This is another introduced South American species evidently
related to V. rigida, but it is a larger and coarser plant with
smaller flowers as well as less harsh and somewhat viscid pubes-
cence.
3. V. brasiliensis Vellozo, Fl. Flum. 17. 1825; 1:pl. 40. 1827.
V. litoralis var. brasiliensis Briq. Ann. Conserv. & Jard. Bot.
Genéve 7-8: 202. 19
Stems 1 m. more or less in height, 4-angled in cross-section,
practically glabrous below, somewhat scabrous-pubescent above,
slightly contracted at the nodes; leaves decussate, lanceolate,
tapering into a POE iterate: subsessile or petiolar base,
sharply or incised serrate, strigillose and somewhat pustulate
above with veins impressed, sparsely pubescent beneath; spikes
compact, mostly short and strict, usually sessile in open cymes;
bracts scarcely as long as the calyx, lanceolate, subulate at apex,
ciliate; calyx approximately 3 mm. long, somewhat appressed-
pubescent, lobes acute with short subulate tips; corolla-tube a
little longer than the calyx, pubescent without; corolla-limb
inconspicuous; nutlets trigonous, about 2 mm. long, strongly
striate, raised-reticulate at apex; commissural faces scarcely
reaching the tip of the nutlet, muricate-scabrous.
Distribution: Glide America; introduced into southern United States.
Specimens examined:
2 ea ballast dumps at Wilmington, 2 July 1897, Small 5707 (G,
MBG, NY); dry sandy soil, open woods about 3 km. south of Wilmington, 25 July
1922, Biidolph 1004 (G).
Groraia: savanna, 27 May 1927, Korthof (US); Brunswick, 12 May 1930, Mol-
denke 1184 (MBG, NY).
FLoripa: waste place, Chipley, 24 May 1930, Blanton 6598 (MBG, US); waste
ground, Pensacola, 24 July 1899, Curtiss 6490 (G, MBG, NY, US); ballast ground,
Pensacola, 30 April 1903, Tracy 8706 (F, G, MBG, NY, US).
ALABAMA: ballast ground, estuary of Mobile River, 4 July 1893, Mohr (US); Port
[Vor. 20
256 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Eads, 22 Aug. 1900, Lloyd & Tracy 20 (F, G, MBG, NY, US); north bank, Bayou
Terrebonne, Houma, 29 May 1914, Wurzlow (NY).
A species very similar in habit to V. litoralis but readily
distinguished by the difference in inflorescence. The spikes of
V. brasiliensis are short, compact, sessile, and regularly arranged
in open cymes; whereas those of V. litoralis are longer, compact
or somewhat elongate, peduncled, and arranged in more or less
paniculate cymes.
4. V. sphaerocarpa Perry," n. sp.
Stems erect, square in cross-section, branched above, glabrous
or sparsely scabrous-pubescent; leaves oblong, tapering at the
base into a short petiole, 3-6(-10) em. long, acute, sharply
serrate, the upper often entire, scabrous-pubescent above with
trichomes minutely bulbous at the bases, more or less short-
strigillose on both surfaces, prominently veined beneath; spikes
short and dense, subsessile or short-pedunculate, cymosely
arranged; bracts ovate-lanceolate, shorter than the calyx, acute,
pubescent; calyx scarcely 2 mm. long, connivent over the schizo-
carp, appressed-pubescent, teeth very short, acutish; corolla-
tube protruding slightly beyond the calyx; corolla-limb et
1.5-2 mm. broad; schizocarp 1 mm. long, approximately 1
in diameter; nutlets very faintly striate or essentially ans
commissural faces muriculately scabrous.
Distribution: known only from Socorro Island.
= ek tage examined:
x1co: CoLIMA: Socorro Island: March-June 1897, Anthony 380 (G, MBG
But US); 27 May-3 July 1903, Barkelew 231 (G, MBG, P, US); 9 May 1925,
Solis 70 (US).
This endemic from Socorro Island is most nearly related to V.
1 V. sphaerocarpa Perry, sp. nov., annua vel perennis (basi ignota); caule erecto
quadrangulari glabro vel sparse scabro-pubescente; foliis oblongis basi in brevem
petiolum attenuatis 3-6(-10) em. longis acutis, infimis argute serratis, superioribus
saepe integris, omnibus scabro-pubescentibus supra subtusque plus minusve breviter
strigosis et reticulatis; spicis brevibus et compactis subsessilibus vel breviter pe-
dunculatis in cymas dispositis; bracteis ovato-lanceolatis acutis pubescentibus
calyce brevioribus; calyce vix 2 mm. longo connivente adpresso-pubescente; calycis
dentibus brevissimis acutiusculis; corollae tubo paulo exserto; corollae limbo circiter
1.5-2 mm. lato; schizocarpio 1 mm. alto circiter 1 mm. lato; coccis obsolete striato-
reticulatis.—Collected on Socorro Island, Mexico, March-June 1897, Anthony 380
(MBG), TYPE
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 257
litoralis and has a similar habit, but is easily separated on the
distinctive characters of the inflorescence. The spikes are
shorter and denser, the flowers are smaller and so crowded that
the lower ones appear to be inserted at right angles to the rhachis
of the spike; moreover, the schizocarp is fully as broad as or even
broader than long, an unusual trait not found elsewhere in the
North American species of Verbena.
5. V. litoralis HBK. Nov. Gen. et Sp. 2: 276, pl. 137. 1818.
V. caracasana HBK. I. c. 275.
V. bonariensis var. littoralis Hook. Bot. Misc. 1: 166. 1830.
V. affinis Mart. & Gal. Bull. Acad. Brux. 11?: 322. 1844.
V. littoralis « pycnostachya Schauer in DC. Prodr. 11: 542.
1847, excluding V. brasiliensis Vell. Fl. Flum. 1: pl. 40.
V. littoralis 8 leptostachya Schauer in DC. Prodr. l. c.
V. Hanseni Greene, Pittonia 3: 308. 1898.
V. litoralis var. caracasana Briq. Ann. Conserv. & Jard. Bot.
Genéve 7-8: 292. 1904
Stems approaching 1 m. in height, square in cross-section,
somewhat fastigiately branched above, glabrous or sparsely
strigillose, slightly contracted at the nodes; leaves lanceolate or
oblong, tapering into a very short petiole or a subsessile base,
3-10 cm. long, decussate, more or less sharply and coarsely
serrate, sparsely strigillose on both surfaces, scabrous and some-
what rugose above, prominently veined beneath; spikes terminal,
fairly dense, pedunculate, cymosely arranged or tending to be
panicled, often elongate; bracts ovate-lanceolate, acuminate, sub-
equalling or somewhat shorter than the calyx, glabrate; calyx
2-2.5 mm. long, finely strigillose, subtruncate, teeth minute,
subulate; corolla-tube variable in length, always somewhat longer
than the calyx; corolla-limb inconspicuous, 2.5-3 mm. broad;
nutlets trigonous, hardly 2 mm. long, striate, somewhat reticulate
at the apex; commissural faces about as long as the nutlet,
muricate-scabrous.
Distribution: Mexico, Central and South America; introduced into the United
tates.
Specimens examined:
Lourstana: north bank, Bayou Black, Houma, 5 and 9 May 1914, Wurzlow (NY).
CALIFORNIA: Clinton, Amador Co., June 1896, Hansen 2025 (MBG).
[Vor. 20
258 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Mexico:
San Luis Porosr: on mountains around the city, San Luis Potosi, 1876, Schaffner
718 (G); Alvarez, 28 Sept.—3 Oct. 1902, Palmer 141 in part (G).
Nayarit: Tepic, 5 Jan.-6 Feb. 1882, Palmer 2019 (US), 2014a (G, NY).
Jasco: fields, Guadalajara, 11 Aug. 1902, Pringle 11093 (F, G, MBG, NY, US).
HipALGOo: near Ixmiquilpan, 1905, Rose, Painter & Rose 9077, 9151 (US)
Vera Cruz: Orizaba, 27 July 1891, Seaton 27 (F, G, NY, US); Huatusco, April
1857, Mohr (US); Fortin, Feb. 1883, Kerber 311 (US); Santa Rosa, 13 Aug. 1926,
Fisher 168 (US).
TABASCO: Mayito, 10 April 1889, Rovirosa (ANSP, NY, US).
PuEBLA: Pahuatlan, 14 June 1913, Salazar (US); near Puebla, 20 Oct. 1908,
Arsène (US).
MicHoacan: low valley, Zinapecuaro, 2 May 1849, Gregg 764 (MBG); Quinceo,
11 Nov. 1909, Arsène (US); Morelia, Aug.-April 1840, Galeotti 781 (K), TYPE col-
lection of V. affinis; Morelia, Coronilla, 8 Aug. 1909, Arséne (US).
Oaxaca: wet meadows, Sierra de San Felipe, 11 Sept. 1894, Pringle 4877 (ANSP,
G, MBG, NY, P, US); Sierra de San Felipe, 6 Oct. 1894, C. L. Smith 222 (MBG, US);
San Jacinto, 25 Sept. 1895, L. C. Smith 794 (G).
CENTRAL AMERICA:
GUATEMALA: uneultivated places, Oct. 1885, Bernoulli 128 (NY); without locality,
1892, Heyde 610 (US); waste places near railway bridge, 5 June 1909, Deam 6180
(G, US); vicinity of Los Amates, Dept. Izabal, 24 May 1922, Standley 24404 (US);
Coban, Dept. Alta Verapaz, May 1886, von T'uerckheim 904 (ANSP, F, G, NY, US);
near the Finca Sepacuite, Dept. Alta Verapaz, 25 March 1902, Cook & Griggs 148
(US); Canjutz, Dept. San Marcos, 1 Sept. 1922, Salas 32 (US); Guatemala, 1923,
Ruano 332, 403 (US); Guatemala City, 1912, Aguirre 4 (US); near Guatemala, July
1921, T'onduz 627 (US); Chiapas, Dept. Santa Rosa, Dec. 1892, Heyde & Lux 4370
(G, NY, US); Santa Rosa, May 1892, Heyde & Lux 3019 in part (G).
Honvuras: vicinity of Siguatepeque, Dept. Comayagua, Standley 56082 (F, US).
SAN BALVADOR: vicinity of Ateos, Dept. La Libertad, 17 April 1922, Standley
23326 (G, US); Volean de San Vicente, Dept. San Vicente, 7-8 March 1922, Standley
21486 (G, NY, US); Cerro de San Jacinto, near San Salvador, 8 Feb. 1922, Standley
20617 (G, US); San Jacinto, 5 April 1905, Velasco 8848 (G, US); Volean de San
Salvador, 7 April 1922, Standley 22842 (US); vicinity of San Salvador, 1905, Renson
291 (NY, US); San Salvador, 1922, Calderon 729 (G, NY, US), 925 (G, US)
Nicaraaua: Casa Colorada and vicinity, south of Managua, 27 June 1923, Mazon,
Harvey & Valentine 73856 (NY, US); Las Nubes and vicinity, south of Managua, 28
June 1923, Maxon, Harvey & Valentine 7478 (US).
Costa Rica: without locality, 19 June 1874, Kuntze 2109 (NY, US); without
locality, April 1910, Worthen (MBG); Los Ayotes, near Tilaran, Prov. Guanacaste,
21 Jan. 1926, Standley & Valerio 45430 (US); Aguacate, Nov. 1846, Oersted 11522
(US); vicinity of San Jose, Feb. 1924, Standley 33293 (US); vicinity of La Palma, on
the road to La Hondura, 17-18 July 1923, Maron & Harvey 7951 (US); Cerro de
Piedra Blanca, above Escasu, 31 Jan. 1924, Standley 32652 (US); near Cartago,
1845-8, Oersted 11324 (US); Cartago, Oct. 1887, Cooper 5890 (F, G, MBG, NY, US).
PANAMA: fields along Rio Caldera above El Boquete, 5 Feb. 1918, Killip 3510 (US).
This is a widely distributed species with very distinctive habit
and somewhat variable inflorescence. In some specimens the
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 259
spikes appear to remain compact, in others they tend to elongate.
Schauer used this difference to separate the forms pycnostachya
and leptostachya, although he frankly admits the difficulty of
distinguishing the two owing to the intermediate phases. V.
affinis is characterized by à somewhat coarser floral pubescence;
this, however, seems to be a variable feature and, as such, does
not appear to merit more than passing mention.
Ser. II. LEPTOSTACHYAE Schauer. Spikes slender and open
or compact at anthesis, greatly elongating in fruit, solitary or in
simple cymes or panicled.—North America.
KEY TO THE SPECIES
A. Spikes panicled at the apices of stems and LE subtended chiefly by
inconspicuous bracts; floral bracts not prom
B. Spikes very slender, elongated and graceful, ined with remote fruits.
Leaves 1-2-pinnatifid or 3—5-cleft or deeply incised.
D. Corolla-limb not less than 3 mm. broad; nutlets 1.5 mm. or more
E. diccre a little longer than broad.
F. eise) usually densely glandular and somewhat viscid-
WAIDOMOORNG ss.. ederin ke 6 oe ROB rre ene . V. officinalis
E. Schizocarp about twice as long as broad.
G. Leaf-blades diverse in outline, the basal incised-dentate, the
middle stem-leaves 1-2-pinnatifid, the upper sparingly
hot pd or entire; bracts one-half as long as the calyces at
ts n NPEREETE SEPIUS Pon a ae hoe os 8. V. Halei
G. Leaf-blades jud in outline; bracts about as long as the
onkpoes at snthesia. «205 RR esa rore 9. V. riparia
D. Corolla-limb eldest more than 1 mm. broad; nutlets 1.5 mm. or
len Joni. ia cap cesses e Eee eee 10. V. Ehrenbergiana
C. Leaves serrate or shallowly incised.
. Leaves subsessile or rider with attenuate base
I. Stems mum leaves mostly crenate-serrate; Nee
ulou
hispidulous... 7... 2.22. ov EO OS LUE . V. carolina
I. Stems inis leaves acutely serrate; calyx pe glabrous
TOME. ES E E N . longifolia
H. ly petioled with round MG Mon
x aid et im spreading; calyx-lob gmatic surf
subtended by two sterile ike 1-3- MP MROCDNENUT Ve 14. V. scabra
J. Fruiting-calyx ascending; calyx-lobes not connivent; stigmatic
e subtended by one sterile style-lobe........ 18. V. urticifolia
B. Spikes thicker or densely flowered, usually with contiguous fruits.
[Vor. 20
260 ANNALS OF THE MISSOURI BOTANICAL GARDEN
K. Spikes subsessile or very short-peduncled, forming a "— in-
florescence, not strict; corolla-limb 2 mm. broad.............. 2. V. recta
K. Spikes peduncled, usually not indictum strict; corolla-limb à 5
M e ere rer yee PES eT Se TT TRY TIT eT Ors Free 16. V. hastata
A. Spikes solitary or in 3's at the apices of stem and branches, or res
and subtended by leafy bracts at the base; floral bracts prominent or no
L. Schizocarp readily separating into 4 nutlets at maturity.
. Leaves serrate-dentate or shallowly incised, or predominantly entire.
N. Leaves serrate or edic or shallowly incised; spikes with
more or less contiguous fr
O. Plants coarse, more or less d hirsute-pubescent or hirsute-
villous; leaves —— to ovate-orbicular; spikes
stout at anthesis, 7-10 m
P. Leaves aai + to ovate-acuminate, short-petiolate;
corolla-limb 5-6 mm s
Q. Plants em A ineei spikes flexuous, elongated and
open in fruit; bracts not exceeding the fruiting calyx;
nutlets 2 mm. dis shallowly scrobiculate above, sulcate
toward the base... isses ese 21. V. macrodonta
Q. Plants lucis oO spike strict, dense in fruit; bracts
surpassing the fruiting calyx; nutlets 2.5 mm. long, retic-
ulate-scrobiculate above, striate toward the base........
NEUE SERGE ENG T ECRAPRERRUREVERPELEAXCPDUUR 20. V. MacDougalii
P. Leaves ovate-orbicular, sessile; corolla-limb 8-9 mm. broad..
MEET ToT RET Te Pe eee eae TIT ee NT 19. V. stricta
O. Plants more slender, hirtellous or — strigillose with short
trichomes, canescent or not; leaves linear to narrowly elliptical
or spathulate; spikes at an nibus 5-6 mm. broad.
R. Leaves strigillose; inflorescence not p——
kia seit eave ta ani wh Wa Oke prie. 7. V. engin
R. — riiin inflorescence glandular-pubescent.
8. ; bracts lanceo-
nione: sinh corolla-limb 3—4 mm. broad. . .18. V. Orcuttiana
8. Leaves hirtellous and canescent; rhachis — ie
bracts ovate-acuminate; eoeolia-limb 6-8 mm
V Ed MERE E sw acea les 26b. V. neomexicana var. ‘Meridia
N. Leaves predominantly entire (mostly ^ie the lower ones w
few salient teeth); spikes with more or less remote fruits. . 27. y. perennis
M. Leaves deeply incised-dentate or pinnatifid or 3-cleft.
T. Spikes not essentially bracteose; flowers readily seen; corolla-limb
U. Leaf-blades, at least the lower ones, oblong-ovate or obtusely
elliptie-ovate, not narrowly elongated, usually 3-cleft with the
segments incised-dentate.
V. Leaves petioled; pubescence various, but not rns hirsute;
spikes usually stout, if slender not greatly elong
W. Leaves not plicate, venation not noticeably int near
the margin; spikes stout at anthesis, elongating or not in
ruit.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 261
X. Plants pilose to hirsute-villous; leaves not scabrous above;
spikes elongating in fruit; bracts not WE the
s i f MEER TE m . V. prostrata
X. Plants sparsely hirsute; leaves scabrous E spikes
occasionally elongating in fruit; bracts slightly sur-
passing the calyx. 36 cos cc chases Eas oS 23. V. robusta
W. Leaves more or less strongly plicate; venation a aed d
whitish near the margin; spikes not stout at anthesis. .
jis dint ae Capes as late a see eee ee 25. y. aplioala
V. Leaves subsessile or at most very short-petioled; pubescence
coarsely hirsute; spikes slender and greatly elongated. . 24. V. zutha
U. Leaf-blades, at least the lower ones, oblong-lanceolate to spathu-
late, narrowly elongated, usually incised-pinnatifid or incised-
dentate (excl. V. plicata).
Y. Plants coarse with a low somewhat compact habit, canescent-
hirsute; leaves subpinnatifid, contracted into a broadly
margined semiamplexicaul or petiola
Z. Inflorescence somewhat glandular-hirsute; bracts lanceolate.
ba dos dbi db das o id vb Mea pd ED Sie . canescens
Z. Inflorescence sparsely, if at all, glandular and densely hermes
bracts ovate, abruptly acuminate......................
eer ee ee Per re err a. V. canescens var. Roemeriana
Y. Plants more slender with a taller E open habit, hirsute to
canescent-hirtellous; leaves .pinnately cleft to incised-
entate, with a narrowly margined petiolar base.
AA. Corolla-limb 4 mm. broad; commissural faces extending to the tip of the
nulle... ovo err cas eer RERO ICE ET Ced 26. V. neomezicana
AA. Corolla-limb 6-10 mm. broad; commissural faces scarcely extending to
the tip of the nutlet.
BB. Plant more or less hirsute; leaves MUS cleft; bracts lanceolate. .
"Tr 6a. V. neomexicana var. zylopoda
BB. Plants canescent-hirtellous; leaves Rena or somewhat m eeply
incised; bracts ovate................. 26b. V. neomexicana var. hirtella
T. Spikes usually bracteose with somewhat foliaceous bracts at th
ani of the spike; flowers inconspicuous; corolla-limb 2.5-3 m
road (larger in V. plicata).
CC. md delicate; leaves incised-pinnatifid to pinnately cleft, not pli-
cate; inflorescence glandular-hirtellous; nutlets 1.5-2 mm. long,
scrobiculate practically to the base...................4. 28. V. gracilis
CC. Plants coarse; leaves incised-dentate to subpinnatifid or 3-cleft, pli-
cate or an inflorescence hirsute or pun nutlets
2-2.5 mm. long, not scrobiculate to the bas
DD. Leaves more or less strongly plicate, UN whitish-
veined near the margin... .. r or ZU Pre LEVE D 25. V. plicata
DD. oie ue plicate nor conspicuously whitish-veined near the
EE. Plants j EA leaves spreading-hirsute; nutlets raised-
reticulate above, striate below
FF. Leaves with a subpetiolar or scialinxplixicadl base; bracts
[Vor. 20
262 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ovate (a little longer than the oe abruptly acumin-
ate, ascending.............. 29a. anescens var. Roemeriana
FF. Leaves narrowed into a margined Seide: bracts linear-
lanceolate (much longer than the flowers), usually re-
81.
8 8. EE E E TE V. bracteata
EE. Plants hirsute-pubescent; leaves appressed-hirsute; nutlets
only faintly reticulate or essentially smooth. ..... 80. V. subuligera
L. Schizocarp tardily separating into 4 nutlets at maturity.......... 32. V. carnea
6. V. officinalis L. Sp. Pl. 20. 1753.
V. spuria L. l.c.
V. domingensis Urb. Symb. Ant. 5: 484. 1908.
Stems ascending or erect, branched, glabrous or nearly so;
leaves 2-7 cm. long, strigillose on both surfaces; basal and lower
stem-leaves more or less ovate, narrowed below into a petiole,
1-2-pinnatifid or 3—5-cleft with parts incised; upper leaves similar
but smaller and less divided; spikes paniculately disposed or in
3's or solitary, slender and elongate; bracts usually about half as
long as the calyx; calyx 2-2.5 mm. long, glandular-pubescent,
subtruneate, teeth minute; corolla-tube a little longer than the
calyx; corolla-limb about 4 mm. broad, segments more or less
rounded; nutlets trigonous, barely 2 mm. long, strongly striate,
slightly reticulate above; commissural faces muricate.
Distribution: Europe; introduced in waste places in eastern North America and
West Indies.
Specimens examined:
MassaAcuusETTS: Rowley, Oakes (G).
RuopkE IsnAND: Warwick Neck, 1848, Thurber (G).
New York: Brooklyn, 13 Sept. 1879, von Schrenk (MBG).
New Jersey: ballast, Camden, 30 Aug. 1874, Parker (G); roadside, Longacoming,
Camden Co., 23 July 1867, Parker (G); Cold Spring, Cape May Co., 30 Aug. 1917,
Gershoy 583 (G).
PENNSYLVANIA: Philadelphia, 1844, Lea (MBG); Lancaster, 21 Aug. 1861, Porter
(G); banks of Susquehanna, Harrisburg, Oct. 1852, Porter (G); York Furnace, York
Co., 28 June 1899, MacElwee 873 (MBG).
DEgrLAwARE: Wilmington, 1845, T'atnall (G).
District OF CoruMsBia: vicinity of Washington, 23 June 1878, Ward (MBG).
VinaiNIA: Parksley, 11 Sept. 1902, Norton (MBG); Bedford Co., 30 June 1870,
Curtiss (MBG); Marion, 1892, Small (MBG).
ORTH CAROLINA: without data, Curtis (MBG); waste grounds, Moyock, 1 July
1922, Randolph 587 (G); dry sandy bank about 3 km. west of Plymouth, 4 July 1922,
Randolph 645 (G); Clarkton, 23 June 1897, Biltmore Herbarium 4762 (G, NY);
Cranberry Forge, July 1895, F. Wislizenus 1214 (MBG).
Sourn CAROLINA: streets in Graniteville, 22 May 1899, Eggert (MBG).
Georata: Rome, Chapman (MBG).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 263
Fronipa: without data, Rugel 121 (F, MBG).
ALABAMA: Collinsville, 29 July 1897, Eggert (MBG); Attalla, 9 July 1898, Eggert
(MBG).
LourstANA: Port Eads, 22 Aug. 1900, Tracy & Lloyd (G, MBG, NY).
TENNESSEE: along the banks of Doe River, Carter Co., 16-17 July 1891, Small &
Heller 484 (G, MBG); Knoxville, July 1893, Ruth (MBG); Hollow Rock, Carroll Co.,
5 Aug. 1897, m (F, MBG).
West INDIE
BERMUDA: Hamilton Parish, 11 July 1905, Moore 2939a (G); Flatts, 31 Aug.-20
Sept. 1905, Brown & Britton 28 (ANSP, G, NY); Flatts, 3 Aug. 1915, Collins 267
(G, NY); Bailey's Bay, 10 Feb.-9 March 1908, Brown 492 (ANSP, G, NY); St.
George’s, 18 Jan. 1912, Robinson 113 (G).
Santo Dominao: Angostura del Rio Yaqui, 8 May 1887, Eggers 1828 (NY), TYPE
collection of V. domingensis; Culo de Maco, Prov. Azua, Aug. 1912, Fuertes 1856
(NY); Loma Rosilla, Prov. de la Vega, July 1912, Fuertes 1771 (NY); vicinity of
Mission, Fonds Varettes, 12 April-4 May 1920, Leonard 3939 (G, NY).
Cusa: without data, Wright 3658 (G); Vecindad de Vento, May 1906, Baker 2591
(NY, P); Playa de Marianao, 29 March 1911, Britton & Cowell 103826 (NY); near
Playa de Marianao, Leon (NY), Leon & Edmunds (NY); cultivated field, Campo
Florido, Prov. Havana, 13 March 1905, Curtiss 677 (ANSP, G, MBG, NY).
On the whole, the specimens from Santo Domingo and Cuba
differ from the typical V. officinalis in their slender and more
elongate habit; the inflorescence is scarcely as glandular, the
flowers are smaller, and the nutlets often do not exceed 1.5 mm.
in length. Nevertheless, the Cuban specimens vary greatly in
size, and Curtiss 677 is hardis separable from typical V. officinalis.
Since many of the specimens are rather poor, it appears probable
that they may very well represent an impoverished condition.
Urban himself was somewhat uncertain of the status of his
species as he appended the following note in a later publication:
“ An re vera a formis V. officinalis L. separanda?'
7. V. menthaefolia Benth. Pl. Hartw. 21. 1839.
V. setosa Mart. & Gal. Bull. Acad. Brux. 11?: 321. 1844.
Stems decumbent or ascending, branched, sparsely and
minutely hispidulous; leaves ovate, tapering at base into a
margined petiole, 3-6 cm. long, deeply cleft or subincised with
divisions remotely serrate-dentate, strigillose on both surfaces,
somewhat pustulate above; spikes panicled, slender, elongate,
compact only at the apex; bracts ovate-lanceolate, acuminate,
variable in length, usually shorter than the calyx, ciliate and
sparsely strigillose; calyx 2.5-3 mm. long, strigillose, sparsely (if
at all) glandular, teeth minute; corolla-tube only slightly longer
[Vor. 20
264 ANNALS OF THE MISSOURI BOTANICAL GARDEN
than the calyx; corolla-limb about 6 mm. broad, segments more
or less truncate; nutlets trigonous with convex back, 2-2.5 mm.
long, striate, raised-reticulate above; commissural faces muricate.
Distribution: Arizona to southern California, northwest Mexico to Oaxaca.
Specimens examined:
ARIZONA: north of Yuma, 26 April 1906, Jones (P).
CaLrFOonNIA: San Diego, May 1852, Thurber 555 (G), June 1875, Palmer 308
(MBG), 5 May 1903, Abrams 3406 (G, MBG, NY, P), 6 July 1915, Macbride &
Payson 781 (G),17 April 1918, Carlson (G), in canyons, Aug. 1918, Spencer 971 (G, P),
waysides, 31 Oct. 1919, Spencer 1414 (G, P); Otay Creek, San Diego Co., 27 March
1923, Peirin 8379 (P).
Mexico
LOWER CALIFORNIA: Salton River Crossing, 27 April 1894, Schoenfeldt 2915 (G,
NY, US); sandy roadside, Tia Juana, 1 Feb. 1920, Bartram (ANSP); ranch, 46 km.
southwest of Tia Juana, 13 April 1925, Jones (P); base of Cucopa Mountains, 6
April 1905, MacDougal 168 (NY).
Sonora: vicinity of Alamos, 16 March 1910, Rose, Standley & Russell 12934 (US);
vicinity of Hermosillo, 6 March 1910, Rose, Standley & Russell 12451 (US); vicinity
of Navojoa, 21 March 1910, Rose, Standley & Russell 13130 (US).
SINALOA: collection of 1921, Ortega 4215 (US); vicinity of Fuerte, 25 March 1910,
Rose, Standley & Russell 13447 (US); vicinity of San Blas, 24 March 1910, Rose,
Standley & Russell 13422 (NY, US); Topolobampo, 15-25 Sept. 1897, Palmer 268
(US); near Plomosas, 18 July 1897, Rose 1763 (US).
Curnuanva: valley of the San Pedro, Ortiz, 11 April 1887, Pringle 1599 (MBG).
CoaHUILA: near Saltillo, 7 May 1848, Gregg 11 (MBG); near Saltillo, 14 July 1848,
Gregg 265 (G, MBG); vicinity of Saltillo, May 1898, Palmer 191 (G, MBG, NY, US);
vicinity of Buena Vista, 24 July 1848, Gregg 276 (MBG); valley of Parras, 11 April
1847, Gregg 406 (MBG, NY); San Lorenzo de Laguna, about 120 km. southwest of
Parras, 1-10 May 1880, Palmer 1042 (ANSP, G, US).
Duranco: near El Salto, 12 July 1898, Nelson 4577 (MBG, US); bottom-lands
and ravines, Durango, April-Nov. 1896, Palmer 153 (F, G, MBG, NY, US), 356
(G, MBG, NY, US).
San Luis Porosi: Alvarez, 28 Feb.-3 Oct. 1902, Palmer 141 in part (F, G, MBG,
NY, US); San Luis Potosi, 1878, Parry & Palmer 717 (G).
AGUASCALIENTES: near Aguas Calientes, 10 Oct. 1903, Rose & Painter 7799
US).
GUANAJUATO: margin of stream, Sirena Mountain, 1894, Duges (G); Leon, 1839,
Hartweg 175 (K Tyre, NY
QUERETARO: vicinity of Queretaro, 1912, Basile 99 (US); Queretaro, 1910-13,
Arsène & Agniel 10242 (F, G, MBG, US); Queretaro, July 1914, Arsène 9998 (US).
Vera Cruz: Maltrata, 20 Aug. 1891, Seaton 7 (F, G, US); Huatusco, 1857, Mohr
S).
HipaLao: Sierra de Pachuca, 20-24 July 1905, Rose, Painter & Rose 8753 (G, NY,
US); Nopala, Aug. 1913, Salazar (US); Moran, 1840, Galeotti 778 (K), TYPE collec-
tion of V. setosa.
Mexico: near Tlalnepantla, 6 July 1905, ps Painter & Rose 8382 (G, US);
Lomas de Santa Fe, July 1928, Lyonnet (US); V ley of Mexico, 1865-66, Bourgeau
860 (US), 547 (G); Valley of Mexico, 27 June Ag Pringle 8534 (ANSP, F, G, MBG,
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 265
NY, P, US); mountains between Toluca and Mexico City, 28 June 1910, Rusby 181
(NY).
More tos: La Cascada, 29 May 1901, Pringle 9529 in part (F, US).
MicHoACcAN: Morelia-Rincon, Arsène 2798 (G); east of Maravatio, 30 April 1844,
Gregg 823 (MBG).
Oaxaca: mountains, San Juan del Estado, 18 June 1894, L. C. Smith 27 (G);
Sierra de San Felipe, 20 Aug. 1894, Pringle 5715 (G); between Coixtlahuaca and
Tamazulapam, 12 Nov. 1894, Nelson 1943 (US); between Las Sedas and Salome, 30
Aug. 1921, Conzatti 4207 (US).
In the specimens cited from California south including Sinaloa,
the inflorescence is more densely strigillose than in the collections
from the southern part of Mexico, the calyces are about 1 mm.
longer, with teeth strongly unequal and the subtending bracts
often as long as the calyces. Although this apparently indigenous
species has been known generally as V. officinalis, it has somewhat
harsher pubescence and is scarcely, if at all, glandular. The
fruiting calyx tends to be connivent, concealing the apex of the
schizocarp rather than open and disclosing it. Perhaps these
are differences only of degree and may be merely variations of V.
officinalis; nevertheless, for the present it seems preferable to
retain the name V. menthaefolia for the American representative.
8. V. Halei Small, Bull. Torr. Bot. Club 25: 617. 1898.
V. leucanthemifolia Greene, Pittonia 5: 135. 1903.
Stems usually several from a woody base, erect, ascendingly
branched, glabrous or strigillose above; leaves 3-10 cm. long,
strigillose on both surfaces, diverse in outline—the basal and
lower stem-leaves oblong to ovate, tapering into a petiole ap-
proximately as long as the blade, irregularly dentate or incised;
the middle stem-leaves 1—2-pinnatifid with shorter petioles; the
upper sparingly dentate or entire; spikes paniculately disposed,
slender and elongate; bracts about one-half as long as the calyx,
appressed, ciliate; fruiting calyx 3-3.5 mm. long, strigillose,
subtruncate, the broad nerves terminating in unequal subulate
teeth; corolla-tube scarcely longer than the calyx; corolla-limb
6—7 mm. broad, segments retuse; nutlets trigonous, approximately
2.5 mm. long, usually prominently striate, raised-reticulate at
apex; commissural faces muricate.
Distribution: Alabama to Texas, Mexico
ALABAMA: without data, Buckley (MBG); p 11 May 1904, Deweys (G).
[Vor. 20
266 ANNALS OF THE MISSOURI BOTANICAL GARDEN
MississIPPI: cultivated grounds near coast, May 1859, Hilgard (MBG); Ocean
Springs, 14 Sept. 1891, Seymour 50 (G); Ocean Springs, June 1892, Skehan 109 (G,
MBG), 46 (G); Biloxi, 9 June 1900, Tracy (NY); Natchez, June 1898, Shimek
(MBG).
LourstaNna: without data, Riddell 1268 (NY); without data, Hale (G), 245 (NY);
sandy open ground, Natchitoches, 24 April 1915, E. J. Palmer 7369 (MBG); Alex-
andria, 20 Aug. 1847, Gregg (MBG); vicinity of Covington, 1920, Arsène 11831,
12242 (US); Gretna, opposite New Orleans, 5 May 1899, Ball 344 (G, MBG, NY);
Madisonville, 14 May 1888, Joor (MBG); along Calcasieu River near Lake Charles,
11 April 1925, Small & Wherry 11774 (NY); Cameron, 4 July 1903, Tracy 8709
(F, G, MBG, NY); vicinity of Cameron, 4 Dec. 1910, McAtee 1953 (US).
OKLAHOMA: south of Dougherty, Murray Co., 1 May 1926, Stratton 62 (MBG).
Texas: Texarkana, Letterman (MBG); railroad near Polk, 13 June 1898, Eggert
(MBG); between Iowa Park and Electra, Wichita Co., 20 Aug. 1921, Ferris & Duncan
3337 (MBG); near Longview, 19 April 1899, Biyori (MBG); Dallas, Reverchon 42
(G), 732 (MBG); Dallas, Jones (P); North Dallas, 27 June 1899, Eggert (MBG);
Brazos, July 1843, Lindheimer 155 (77) (MBG); Tarrant Co., Ruth 108 (G, MBG,
NY, US); Van Zandt, April 1929, Ezell 5699 (US); prairies near Granbury, Hood Co.,
4 May 1900, Eggert (MBG); Abilene, 19 May 1902, Tracy 7996 (F, G, MBG, NY);
near Comanche, 8 May 1900, Eggert (MBG); Waco, 1904, Pace 22 (MBG); Gurley,
20 April 1907, Howell 362 (US); San Augustine, 19 April 1916, E. J. Palmer 9485
(MBG); College Station, Brazos Co., 28 April 1917, E. J. Palmer 11715 (MBG);
Mill Creek, April 1839, Lindheimer (MBG); near Houston, March 1842, Lindheimer
155 (106) (MBG); prairies, Houston, 1872, Hall 432 (MBG, P); Houston, 21 April
1899, Eggert (MBG); Houston, 17 April 1900, Bush 28 (MBG, US); Galveston
Island, May 1843, Lindheimer (MBG); Galveston Island, 22 Sept. 1901, T'racy 7533
(F, G, MBG, NY); Alvin, 22 April 1918, Young (P); Columbia, 9 April 1899, Bush
84 (G, MBG); Columbia, 20 April 1900, Bush 77 (G, MBG, NY, US); sandy prairies,
Columbia, 29 March 1914, E. J. Palmer 5044 (MBG); Austin, 1 May 1915, Young 77
(MBG); Austin, 7 April 1922, Tharp 2818 (US); flood plains of Colorado River, near
Austin, 2 April 1929, Armer 5385 (US); Crab Apple Creek, Gillespie Co., Jermy 203
(MBG); Kerrville, 4 June 1916, E. J. Palmer 10037 (MBG); New Braunfels, April
1851, Lindheimer 1076 (537) (G, MBG, NY); near Bracken, 13 July 1903, Groth 75
(G, NY); Bexar to Austin, April 1828, Berlandier 322, 1592 (G); San Antonio, 1900,
Wilkinson (MBG); San Antonio, 1911, Clemens & Clemens 969, 970 (P); San An-
tonio, April 1922, Schulz 766 (US); Sutherland Springs, Wilson Co., Aug. 1879,
Palmer 1043 (G); near Sabinal, 21 April 1925, Small & Wherry 11998 (NY); Del Rio,
20 April 1930, Jones 26229 in part (P); Uvalde, 11 May 1918, E. J. Palmer 13564
(MBG); Uvalde, 28 April 1928, E. J. Palmer 33604 (MBG, NY); Millett, 11 May
1897, Trelease (MBG); Cuero, 22 March 1907, Howell 313 (US); prairies near Goliad,
8 April 1900, Eggert (G, MBG); Refugio, 8 March 1916, E. J. Palmer 9111 (MBG);
Corpus Christi, 5-12 March 1894, Heller 1419 (G, MBG, NY, US); Corpus Christi,
1 May 1913, Orcutt 5867 (MBG); Kingsville, 25 March 1920, High 52 (MBG); near
Laredo, Aug. 1899, Mackenzie 96 (MBG); near Laredo, 6 April 1901, Eggert (MBG);
Rio Hondo, Cameron Co., Sept. 1913, Chandler (G, MBG); Sierra Blanca, 11 April
1930, Jones 26229 in part (MBG).
Mexico: TAMAULIPAS: Matamoros, Berlandier 1511, 3016 (G, MBG); vicinity of
Tampico, 1-31 Jan. 1910, Palmer 78 (F, G, MBG, NY, US).
A species closely related to V. menthaefolia and V. officinalis,
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 267
but readily distinguished by the diverse outline of the leaves
and the somewhat more slender achenes.
9. V. riparia Raf. ex Small & Heller, Mem. Torr. Bot. Club
3:12. 1892.
? V. hastata var. B oblongifolia Nutt. Gen. 2: 40. 1818.
V. urticifolia var. riparia Britton, Mem. Torr. Bot. Club 5:
276. 1894.
Stems 6-15 dm. tall, erect, sparsely pubescent or glabrate,
widely branched; leaves oblong to ovate, 4-12 em. long, petioled,
pinnatifid or nearly tripartite toward the base, sparsely strigillose
on both surfaces, venation prominent beneath; spikes paniculately
disposed, slender, elongate; bracts lanceolate-ovate, as long as the
calyx at anthesis, acuminate; calyx 3 mm. long, minutely glandular-
pubescent, subtruncate, teeth minute; corolla-tube only slightly
longer than the calyx, puberulent without; corolla-limb 3.5 mm.
broad; segments more or less rounded, the middle posterior one
emarginate; nutlets oblong, 2-2.5 mm. long.
Distribution: New Jersey and Virginia to North Carolina (acc. to Small).
Specimens examined:
VirGINIA: Marion, peden Co., 1 July 1892, Small (P); Marion, Smyth Co., 6
July 1892, Small (G, M
NoRTH CAROLINA: near E Caldwell Co., 3 July 1891, Small & Heller (F); near
falls of Yadkin, Stanley Co., 18 Aug. 1891, Small & Heller (F).
The material at hand is too scanty and too immature to give
many clues to the probable relationship of this species. It
would seem as nearly related to V. officinalis as to either V.
hastata or V. urticifolia. It is characterized by pinnatifid or
tripartite leaves, minutely glandular-pubescent inflorescence,
and fruit about twice as long as thick.
10. V. Ehrenbergiana Schauer in DC. Prodr. 11: 548. 1847.
Stem erect, branched, hirsute; leaves trifid with lateral lobes
small, somewhat ovate with cuneate base narrowed into a mar-
gined petiole, attenuate at apex, 4-8(-10) cm. long, coarsely
serrate-dentate, strigillose above, hirtellous below, also somewhat
paler and prominently veined; spikes paniculately disposed,
very slender, remotely flowered; bracts ovate, about one-half as
long as the calyx or shorter, subulate, ciliate; fruiting calyx
[Vor. 20
268 ANNALS OF THE MISSOURI BOTANICAL GARDEN
about 1.5 mm. long, strigillose, lobes very short, obtuse, mucro-
nate; corolla inconspieuous, limb about 1 mm. broad; nutlets
tigonone with convex back, 1 mm. long; commissural faces
meeting sharply at right angles, almost smooth.
Distribution: Mexico.
Specimens examined:
Mkxico:
Coanuita: Saltillo, July 1880, Palmer 2037 (G).
NvuEvo — moist places near Monterey, July 1888, Pringle 1948 (ANSP, F, G,
MBG, NY,
AN Luis ix oTOSI: Tancanhuitz, Feb. 1888, Seler 722 (G, US); Rio Verde, 17 Nov.
1910, Orcutt 5423 (MBG); Bagre, Minas de San Rafael, July 1911, Purpus 5451 (F,
G, MBG, NY, US).
Hipauco: Cazadero, April 1841, Liebmann 11335 (US).
Vera Cruz: Wartenberg, near Tantoyuca, 1858, Ervendberg 153 (ANSP, G).
PuEBrA: Pahuatlan, 12 July 1913, Salazar (US).
Mexico: near Los Reyes, Ehrenberg 713 (Bot. Mus. Berl.-Dahl. TYPE, MBG phot.).
In general habit this species somewhat approaches V. carolina.
It differs, however, in its tripartite leaves, smaller flowers, and
fruits. The eollsotion from Saltillo is much more densely
hirsute on all parts, but apparently is conspecific.
11. V. carolina L. Syst. ed. 10, 852. 1759; Sp. Pl. ed. 2,
29. 1762.
caroliniana Willd. Sp. Pl. 1: 119. 1798.
polystachya HBK. Nov. Gen. et Sp. 2: 274. 1818.
biserrata HBK. Ll. c. 275.
veronicaefolia HBK. L. c.
hirsuta Mart. & Gal. in Bull. Acad. Brux. 11?: 321. 1844.
. mollis Mart. & Gal. l. c. 323.
. paucifolia Mart. & Gal. l. c. 324, as V. pauciflora M. & G.
in Walp. Rep. 6: 687. 1846-47.
V. caroliniana forma or var. polystachya (Kunth) Loes. in
Fedde, Rep. Sp. Nov. Veg. 9: 362. 1911.
Stems erect or ascending, usually solitary, branched, hirsute-
hispid; leaves lanceolate-oblong or somewhat elliptical, 3-8(-12)
em. long, obtusish or acute, subsessile or narrowed into a short
petiole, coarsely crenate-serrate, strigose and somewhat pustulate
above, hirsute-strigose especially along the midrib and veins
beneath; spikes paniculately disposed, slender, open in fruit;
bracts ovate, varying in length, usually about one-half as long as
Must Bis i Big Bia Me
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 269
the calyx or shorter, acuminate-subulate, somewhat spreading,
ciliate; fruiting calyx approximately 2 mm. long, with the obtuse
ROAS lobes connivent, hispidulous; corolla-tube scarcely
exserted; corolla-limb inconspicuous, about 2 mm. broad; mature
schizocarp inclosed by calyx; nutlets trigonous with convex
back, hardly 1.5 mm. long, faintly striate; commissural faces
meeting at right angles, almost smooth.
Distribution: Arizona and Mexico to Salvador, Central America.
Specimens examined:
Arizona: Wilgus Ranch, Chiricahua Mountains, 12 Oct. 1907, Blumer 1783 (G,
MBG, NY, US); Cave Creek, Chiricahua Mountains, 2 Sept. 1929, Harrison & `
Kearney 6144 (US); Santa Rita Mountains, 15 July 1881, Pringle (G); Fort Huachuca,
July 1893, Wilcox (NY); Huachuca Mountains, 4 Sept. 1903, J pes E US); Ramsay
Canyon, Huachuca Mountains, 30 Sept. 1929, Jones 25006 (G, N
Nevapa: Carson Valley, Aug. 1872, Lemmon
Mexico: without data, Graham (G); collection of 1844, ‘Galeotti 795A (G, US).
Lower CALIFORNIA: Sierra de San Francisquito, 1 Oct. 1899, Brandegee (NY).
Sonora: Santa Cruz, 22 Oct. 1893, Mearns 2627 (US); Cochuto, 1 Oct. 1890,
Hartman 94 (G, NY, US); La Cruz de los Canadas, 3 Nov. 1890, Lloyd 448 (G).
SiNALOA: near Colomas, foothills of Sierra Madre, 15 July 1897, Rose 1677 (US);
Santa Lucia, 1919, Dehesa 1551 (US).
Curavanva: Norogachi, Aug.-Nov. 1885, Palmer 364 (ANSP, G, US); San Diego
Canyon, Sierra Madre Mountains, 16 Sept. 1903, Jones (P); Guayanopa Canyon,
Sierra Madre Mountains, 23 Sept. 1903, Jones (P).
DvnaNGo: rich bottom-lands, vicinity of Durango, Apr.-Nov. 1896, Palmer 339
(F, G, MBG, NY, US).
Nayarit: west of Tepic, 31 May 1849, Gregg 1001 (MBG); Tepic, 5 Jan.-6 Feb.
1892, Palmer 2057 (US).
Jauisco: Cuidad Guzman, 14 Oct. 1921, Kempton & Collins (US).
GvANAJUATO: Guanajuato, 1880, Duges (G).
Hipataco: Real del Monte, 15 Sept. 1910, Clokey 1865 n d ; near Moran,
Humboldt & Bonpland (Par. TYPE of V. veronicaefolia, MBG p
Vera Cruz: Sierra Templada in general, June-Oct. 1840, ze | 735 (K), TYPE
PI of V. hirsuta; Orizaba, 1853, Mueller (NY); Orizaba, 1855, Mueller 766,
87 (NY); Orizaba, 1855-7, Botteri 180 (F, G, MBG, US); Orizaba, 1857, Mohr (US);
ata Schiede 88 (NY); Jalapa, Jan. 1894, C. L. Smith 1737 (G).
PuEBLA: region of Atoyac, near Puebla, 15 July 1909, Nicholas (US); Tezuitlan,
9 June 1910, Orcutt 4045 (F).
Mexico: near city of Mexico, Humboldt & Bonpland (Par. TYPE of V. biserrata,
MBG phot.); valley of Mexico to Tizapan, 19 Aug. 1865-66, Bourgeau 119 (G, US);
Chalco region, 4 Oct. 1921, Kempton & Collins (US); near Tlalpam, 1905, Rose,
Painter & Rose 8496 (US); Tlalpam, 9 Aug. 1910, Orcutt 3488 (F, US); Eslava, 15
June 1901, Pringle 9312 (F, G, MBG, NY, US); near Toluca, 18 Sept. 1889, Pringle
2813 (F, G
More os: fields, La Cascada, 29 May 1901, Pringle 9529 in part (G, MBG, NY);
Toro, 5 Aug. 1924, Fisher (F, MBG); Tepoztlan, autumn 1926, R. Redfield 4 (US).
Micuoacan: Morelia, Coronilla, 8 Aug. 1909, Arsène 3000 (US); Quinceo, near
[Vor. 20
270 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Morelia, 11 Nov. 1909, Arsène (US); on slope of voleanie mountain, Xorullo, Hum-
boldt & Bonpland (Par. TYPE of V. polystachya, MBG phot.).
Corma: Colima, 9 Jan.-6 Feb. 1891, Palmer 1156 (G, NY, US).
Oaxaca: mountains near Oaxaca, June-Oet. 1840, Galeotti 737 (K), TYPE collec-
tion of V. mollis; Sierra de San Felipe, 12 Sept. 1894, Pringle 4892 (ANSP, G, MBG,
NY, US); Sierra de San Felipe, 2 Oct. 1894, C. L. Smith 224 (MBG, NY, US);
vicinity of La Parada, 19 Aug. 1894, Nelson 1021 (US); Cuyamecalco, 5 Sept. 1895,
L. C. Smith 634 (G).
CENTRAL AMERICA:
GUATEMALA: uncultivated places, Oct. 1865, Bernoulli 127 (N Y); without locality,
Sept. 1927, Morales 786 (US); without locality, 1892, Heyde 120, 530, 477 (US); San
Siguan, Dept. Quiche, April 1892, Heyde & Lua 3018 (ANSP, MBG, US); Coban,
Dept. Alta Verapaz, Nov. 1902, von Tuerckheim 8442 (F, G, NY, US); Coban, Dec.
1906, von Tuerckheim IT 651 (F, MBG); Tactic, Dept. Alta Verapaz, May 1886, von
Tuerckheim 913 (G, US); vicinity of Los Amates, Dept. Izabal, 24 May 1922, Standley
24443 (MBG, US); Proderos, Dept. Guatemala, July 1921, Tonduz 628 (US);
Chilloui, Dept. Guatemala, 20 June 1921, Rojos 44 (US); Volean Pacaya, Dept.
Amatitlan, July 1892, Shannon 3638 (US); San Vincente, Pacaya, Dept. Amatitlan,
5 May 1921, Tonduz 488 (US); San Lucas, Dept. Solola, 16 Feb. 1906, Kellerman
5825 (US); San Pedro, 1913, Tejada 72 (US); Tejutla, 1913, Tejada 5? (US); Santa
Rosa, Dept. Santa Rosa, May 1892, Heyde & Lux 3019 (G in part, NY, US); vicinity
of Siguatepeque, Dept. Comayagua, 14-27 Feb. 1928, Standley 55944 (F, US).
SALVADOR: vicinity of Apastepeque, Dept. San Vincente, 4 March 1922, Standley
21331 (G, MBG, US); vicinity of San Salvador, Renson 175 (NY, US); Plazuela de
Aculhaca, San Salvador, Dec. 1906, Velasco 8999 (US); Amatepeque Hill, near San
Salvador, 2 Feb. 1907, Pittier 1909 (US); San Salvador, 1922, Calderon 794 (G, US);
vicinity of San Salvador, 30 March-24 April 1922, Standley 22414 (G, NY, US);
vicinity of Santa Tecla, Dept. La Libertad, 10 April 1922, Standley 23083 (G, US)
Finca San Nicolas, 1923, Choussy 36 (US).
This species is probably most nearly related to V. urticifolia
and V. scabra, but is easily set apart from them by its subsessile
or very short-petiolate leaves; moreover, their areas of distribu-
tion are practically distinct. The collections cited appear to be
conspecific, notwithstanding the fact that they vary greatly in
size, serration of leaves, and length of floral bracts. The speci-
mens from the northern part of the range are slightly coarser,
with leaves more remotely crenate-serrate, yet they match very
closely Pringle 4892, from the Sierra de San Felipe, Oaxaca. In
the latter, however, the leaves are more densely pubescent on
the under surface. The specimens from Guatemala and Sal-
vador have somewhat crowded spikes and are, as a whole,
smaller in every way and more densely hispidulous. However,
since most of the collections were made from December to Febru-
ary, it seems as if this more compact habit might be a seasonal
rather than a geographic variation.
.
,
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 271
A specimen labelled *'Inter Tallahassee et St. Marks, Florida,
legit Rugel, April-June 1843" appears in the herbarium of the
Missouri Botanical Garden. As this is the only representative
of V. carolina seen from Florida, and as it is so far from the
natural range of the species, it would appear to be an error in
labelling. Another collection worthy of mention is Nelson 752.
It is almost glabrous, and the lobes of the corolla are emarginate,
a rather unusual feature in this series. The lack of pubescence
suggests V. longifolia, but unfortunately none of the available
material of the species is in sufficiently good condition to reveal
the character of the corolla. Although the general habit of the
collection is that of V. carolina, it 1s rather doubtfully con-
specific.
12. V. recta HBK. Nov. Gen. et Sp. 2: 277. 1818.
V. caroliniana forma or var. recta (Kunth) Loes. in Fedde, Rep.
Sp. Nov. Veg. 9: 362. 1911.
Stem erect, branched, hispidulous, reddish; leaves lanceolate-
oblong to ovate with cuneate base narrowed into a short margined
petiole, 4-8 em. long, acute, serrate-dentate, reticulately veined,
strigillose on both salare, minutely pustulate above; spikes
numerous, congested at the apices of stems and ee short-
cylindrical, usually dense but not strict; bracts a little shorter
than the calyx, ovate-lanceolate, acuminate, hispidulous; fruiting
calyx about 2 mm. long, hispidulous, lobes short, obtuse, mu-
cronate, commonly connivent; corolla-tube scarcely exserted;
corolla-limb inconspicuous; nutlets trigonous with convex back,
1.5-2 mm. long, faintly striate; commissural faces meeting at
right angles, practically smooth.
Distribution: Mexico.
Specimens examined:
MExico:
Hipauao: Real del Monte, 12 July 1913, Salazar (US); between Pachuca and Cerro
Ventoso, Humboldt & Bonpland 4066 (PAR. TYPE, MBG phot.).
Pursta: Huitzuilzilapam, 9 June 1910, Orcutt 3950 (F).
Mexico: Cima Station, 30 Aug. 1905, Pringle 13597 (G, US); Toluca, Berlandier
1222 (US).
ORELOS: Tres Marias, 4 July 1901, Rose & Hay 5310 (US); Toro, 5 Aug. 1924,
Fisher 320 (MBG, US).
Oaxaca: vicinity of Cerro San Felipe, 1894, Nelson 1105 (US); Sierra de San
Felipe, 9 Aug. 1894, Pringle 4769 (ANSP, F, G, MBG, NY, US).
[Vor. 20
272 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Verbena recta, generally determined in the material at hand as
“aff. V. polystachya HBK.," is easily distinguished from this
closely related species by the short hispidulous pubescence, the
commonly broader leaves, and the compact cylindrical congested
spikes. The floral characters of both are similar, although
generally the fruiting calyx of V. carolina is more acutely ovoid
and the subtending bract is much sharper at the apex.
13. V. longifolia Mart. & Gal. Bull. Acad. Brux. 11?: 323.
1844
Stems erect, tall, glabrous, obtusely 4-angled; leaves lanceolate
to elongate-elliptie, 10-12 cm. long, the upper somewhat smaller,
decussate, short-petiolate, acutely serrate from below the middle
to the apex, appressed-pubescent or very short-strigillose on both
surfaces, venation prominent beneath; spikes paniculately
disposed, elongate, slender, open in fruit, glabrous; bracts ovate,
about one-half as long as the calyx, acute-acuminate, minutely
ciliate; calyx 2 mm. long, with the obtuse lobes connivent over
the schizocarp, practically glabrous; corolla-tube scarcely pro-
truding beyond the calyx; corolla-limb inconspicuous; nutlets
trigonous, hardly 1.5 mm. long, smooth or faintly striate; com-
missural faces smooth.
Distribution: southern Mexico.
Cami ens examined:
Mex!
VERA ‘Cavs: Colipa, March 1841, Liebmann 11318 (US); near Chila, April 1888,
Seler 724 (G, US).
PukBLA: in vicinity of San Luis Tultitlanapa, July 1908, Purpus 3406 (F, G,
MBG, NY, US).
Morevos: San Anton, near Cuernavaca, 14 Oct. 1904, Seler 4194 (G, US); Cuer-
navaca, 28 Oct. 1905, Seler 4347 (G, US).
Oaxaca: Tehuantepec, 19 April 1910, Orcutt 3321 (F, MBG, US).
A rather singular species combining the foliar characters of V.
litoralis with the inflorescence characters of V. carolina.
14. V. scabra Vahl, Eclog. Am. 2: 2. 1798, not V. scabra
Marnock, Floricult. Mag. 5: 87, pl. 54, fig. 2. 1840-41.
Stem 1 m. more or less tall, erect, solitary, simple or branched,
hispidulous; leaves ovate to elongate-ovate, 3-10(-13) em. long,
2.5-5 em. broad, petiolate, serrate-dentate, acute or obtusish,
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 273
scabrous and commonly strigillose above, less scabrous and some-
what paler beneath, also hispidulous along the veins; spikes
paniculately disposed, slender, pedunculate, copiously and closely
flowered; bracts ovate-acuminate, about half as long as the calyx,
hispidulous; fruiting calyx 2.5-3 mm. long, hispidulous, ovoid,
with the somewhat unequal lobes acutely connivent, diverging
from the rhachis of the spike 45° or more; corolla-tube scarcely,
if at all, longer than the calyx; corolla-limb about 2 mm. wide,
lobes obtuse; anthers glandless; stigmatic surface midway
between two almost equal obtusish sterile lobes; nutlets trigonous,
1-1.3 mm. long, faintly striate, reticulate above; commissural
faces extending to the top of the schizocarp, meeting sharply
at right angles, muriculate.
Distribution: North Carolina, Florida west to California, south into northwestern
Mexico; West Indies.
Specimens examined:
NonTH CAROLINA: moist rich soil, open swamp land, Wilmington, 25 July 1922,
Randolph 1012 (G).
LORIDA: without locality, 1842-49, Rugel 156 (F, MBG); common in everglades,
27 Aug. 1925, O'Neill (MBG); vicinity of Mayport and Jacksonville, 1870—76,
Keeler (NY); near Jacksonville, 25 June-13 July 1894, Curtiss 5111 (MBG, NY);
Lake City, 21 July 1893, Quaintance (MBG); vicinity of Eustis, Lake Co., 1-15 July
1894, Nash 1248 (G, MBG, NY); low ground, Orange Co., 22 July 1902, Fredholm
5416 (G, MBG, P, US); open low ground, near Lake Okeechobee, 20 May 1925,
E. J. Palmer 27462 (MBG); east shore of Lake Okeechobee, 11-25 Nov. 1913,
Small & Small 4337 (NY); east shore of Observation Island, 11-25 Nov. 1913,
Small & Small 4403 (NY); hammocks along Taylor Creek, 11-25 Nov. 1913, Small
& Small 4341 (NY); Fort Lauderdale, 19-25 Nov. 1903, Small & Carter 1072 (F,
NY); moist sandy soil, Hollywood, Broward Co., 10 Feb. 1930, Moldenke 591,
599a (MBG, NY); everglades west of Miami, 3 Nov.-7 Dec. 1912, Small 4020 (NY);
hammocks near Miami River, 26 Nov.-20 Dec. 1913, Small & Small 4520 (NY);
Biscayne Bay-Indian River, Palmer 397 (G, MBG); Myers, Lee Co., July-Aug. 1900,
Hitchcock 269 (MBG); sandy ditch, south of Estero, Lee Co., 14 April 1930, Moldenke
972 (MBG, NY); Sanibel Island, 18 May 1901, Tracy (NY); Manatee, 8 May 1900,
Tracy 6652 (G, MBG, NY); Hernando Co., June-July 1898, Hitchcock (MBG); near
Tallahassee, Berg (NY); shore of Dog Island Sound, near Lanark, 26 July 1920,
Harper 242 (NY, US); streets of Apa[lachicola], Chapman (MBG).
Mississipri: Scranton, Jackson Co., 5-6 Aug. 1896, Pollard 1191 (G, MBG, NY);
Ocean Springs, 13 Aug. 1895, Skehan (MBG).
LouistANA: South Pass, 20 Aug. 1900, Tracy & Lloyd 22 (G, MBG); New Orleans,
Riddell (G); New Orleans, Drummond (NY); New Orleans, 21 Oct. 1885, Joor
(MBG)
Texas: without locality, July 1851, Lindheimer 618 (MBG); Beaumont, Jefferson
Co., 11 Sept. 1916, E. J. Palmer 10692 (MBG); Beaumont, Sept. 1904, Kirn 2139
(P); Houston, July 1842, Lindheimer (MBG); Comanche Springs, New Braunfels,
[Vor 20
274 ANNALS OF THE MISSOURI BOTANICAL GARDEN
etc., Aug. 1851, Lindheimer 1077 (G, MBG, NY); Leon River, Oct. 1850, Bigelow
(G); San I and vicinity, 1898, Stanfield (NY); New Braunfels, 1847, Lind-
heimer (MBG).
Arizona: north of Rice, Apache Reservation, 4 Oct. 1927, Harrison 4897 (US);
Santa Cruz River, Tucson, 11 May 1881, Pringle (G); banks of Santa Cruz River,
near Tucson, 18 July 1884, Pringle (ANSP, F, G, NY, P, US)
CaLrFORNIA: without locality, Wallace (G); Cienega, on Santa Monica R. R.,
yon 6 (G); Los Angeles, 1892, Davidson (G); near Pasadena, 17 Dec. 1892, Mc-
Clatchie (N Y); San Bernardino, Oct. 1891, Parish (MBG), 5338 (NY); San Bernar-
dino Valley, Parish & Parish 1043 (F, MBG); San Bernardino Valley, 25 July 1909,
Parish 7149 (P); 3 km. south of San Bernardino, 2 Sept. 1924, Johnston (P); Upland,
July 1917, Johnston (P); 8 km. southeast of Chino, Munz & Johnston 11289 (P);
Cocomozo Mountains, Aug. 1881, Parish & Parish 11143 (G).
Mexico:
Lower CALIFORNIA: San Tomas, 15 July 1885, Orcutt 1802 (MBG).
CoanuiLA: Soledad, a section of low mountains with few oaks, 40 km. southwest
from Monoclova, 9-19 Sept. 1880, Palmer 1040 (ANSP, US); San Lorenzo de Laguna,
120 km. southwest of Parras, May 1880, Palmer 1040 (G)
West INDIES:
Cuna: without locality, 1865, Wright 3659 (G); Santa Rosalia, 2 Aug. 1895,
Combs 389 (G, MBG, NY); valley of San Juan, vicinity of Matanzas, 14 March 1903,
Britton, Britton & Shafer 292 (NY); Havana, 15 Oct. 1908, Leon 685 (NY); Havana,
21 April 1919, Leon & Edmunds 8719 (NY); Rio Arimao, 22 March 1910, Britton &
Wilson 5771 (NY); Isle of Pines, 8 March 1916, Britton, Wilson & Leon 15256 (NY);
Pinar del Rio, 7 April 1924, Krig 3187 (NY).
Jamaica: Port Antonio, Dec. 1890, Hitchcock (MBG); Port Antonio, Aug. 1910,
Lang 102 (ANSP); a near Black River, 12 Sept. 1907, Harris 9987 (NY); road-
side, Hopeton, 13-22 Sept. 1907, Britton 1527 (NY); Och Rios and vicinity, 4 April
1908, Britton & Hollick E (NY); banks of Cabaritta River, Meylersfield, 16 Dec.
1914, Harris 11808 (G, MBG, NY).
Harti: Prov. Barahona, Aug. 1910, Fuertes 391 (NY); Prov. La Vega, July 1912,
Fuertes 1758 (NY); Miragoane and vicinity, July 1927, Eyerdam 201, 432 (G).
Porto Rico: without locality, Read (ANSP); Bayamon, 22 March 1885, Sintenis
1074 (MBG, NY); Aybonito, 3 Nov. 1885, Sintenis 2010 (ANSP, P); Vega Baja, 5
Nov. 1913, Stevens & Hess 4260 (NY); El Teudal, Coamo River, 13 Feb. 1922,
Britton, Britton & Brown 6019 (NY); border of Mary Lake, near Luquillo, 3 April
1922, Britton, Britton & Brown 70387 (NY).
Brrmupa: Smith's Parish, 12 July 1905, Moore 2947 (G, NY); Hamilton Parish,
9 July 1905, Moore 2874 (G, NY); cultivated land north of Hamilton Parish, 31
ug.-20 Sept. 1905, Brown & Britton 878 (ANSP, G, NY); Washington Sound,
Sept. 1913, Brown & Britton 1681 (ANSP, NY); roadside, 3 Aug. 1913, Collins 268
(G, NY
Verbena scabra, often confused with V. urticifolia, is easily
separable by the very scabrous upper surface of the leaf and the
rather conspicuous divergence of the fruit from the rhachis of
the spike. A quite distinctive feature is the position of the
stigmatie surface apparently between two almost equal sterile
1933]
PERRY— NORTH AMERICAN SPECIES OF VERBENA 275
style-lobes. This condition is somewhat approached in V.
carolina, but in the other species examined the second sterile
lobe has never been so definitely developed.
15. V. urticifolia L. Sp. Pl. 20. 1753.
V. diffusa Poir. in Lam. Encyc. 8: 550. 1808.
V. diffusa Desf. ex Spreng. in L. Syst. 2: 748. 1825.
V. urticifolia var. simplex Farwell, Papers Mich. Acad. Sci.
3: 108. 4.
Erect herb 5-15 dm. tall; stem solitary, simple or more often
branching from near the base, hirtellous to almost glabrous;
leaves 8-20 cm. long, petiolate, blades broadly lanceolate to
oblong-ovate, with rounded base decurrent on the petiole, short-
acuminate or acute, coarsely and somewhat doubly crenate-
serrate, hirtellous or glabrate on both surfaces, often minutely
pustulate above; spikes paniculately disposed, slender, peduncu-
late, more or less sparsely flowered; bracts ovate-acuminate,
very short, ciliate; calyx 2 mm. long, pubescent particularly
along the nerves, lobes obtuse, not connivent, teeth short,
subulate, subequal; corolla-tube scarcely exserted; corolla-limb
about 2 mm. wide, lobes obtuse; mature schizocarp exposed at
the distal end; nutlets trigonous with a convex back, about 2
mm. long, faintly striate; commissural faces meeting at right
angles, almost smooth.
Distribution: Quebec and Ontario, Maine to Nebraska and southward.
pecimens examined:
UEBEC: vicinity of Longueuil, Aug. 1916, Victorin 3124 (MBG); Cape Tour-
mente, 40 km. below Quebec, 12 Aug. 1922, Victorin 15750 (G).
Ontario: Casselman, 24 Aug. 1891, Scott (G); Niagara Falls, 1830, ? Torrey 154
(NY).
Maine: North Berwick, 25 Sept. 1897, Fernald & Parlin 928 (G).
New HampsuireE: Walpole, 28 July 1901, Williams (G); Ashuelot, 1 Aug. 1898,
Robinson 559 (G).
Vermont: East Middlebury, 14 July 1908, Williams (G); Rutland, 31 Aug. 1899,
Eggleston 1531 (G); Manchester, 21 July 1898, Day 147 (G); Brattleboro, 2 Aug.
898, Robinson 134 (G).
ASSACHUSETTS: Lowell, 28 July 1927, Beattie (P); slopes above Connecticut
River, Holyoke, 7 Sept. 1926, Seymour 601 (MBG, NY); Waverley, 10 Aug. 1878,
Lane (G); Belmont, 27 Sept. 1891, Deane (G); Milton, 3 Aug. 1887, Kennedy (G);
South Framingham, 25 July 1890, Sturtevant (MBG); Dedham, 22 Aug. 1897,
Williams (G); Adams, 24 Aug. 1901, Day ?9 (G); Manchester, 18 Aug. 1895, Williams
(G); Monson, Aug. 1897, Morris (MBG); Nonquit, 1888, Sturtevant (G).
[Vor. 20
276 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Ruope IsrnAND: dry roadside banks and thickets northeast of Great Salt Pond,
Block Island, 20 Aug. 1913, Fernald, Hunnewell & Long 10262 (G); Providence,
July 1844, Thurber (G); Providence, 20 July 1892, J. F. Collins (G).
Connecticut: in damp woods, Wethersfield, Wright (G); Hartford meadow, 3
Aug. 1882, Wright (G); New Haven, 1858, Eaton (MBG).
New York: open pasture, Canton, 18 July 1914, Phelps 800 (G); Bolton Landing,
Lake George, 11 Aug. 1906, Seler 4560 (G); Vaughns, north of Hudson Falls, 8 Aug.
1912, Burnham (G); near station, Ithaca, 4 Aug. 1915, Hames & McDaniels 4870 (G);
Fall Creek, Ithaca, 12 July 1916, Munz 616 (P); Six Mile Creek, Ithaca, 8 July 1878,
Trelease (MBG); Fleischmanns, Delaware Co., 30 July 1892, von Schrenk (MBG);
Port Chester, 3 Sept. 1886, Stabler (G).
New Jersey: Green Pond, Morris Co., 21 Sept. 1886, Britton (NY); Denmark
Pond, 7 Aug. 1910, Mackenzie 4746 (G, MBG); Watchung, Somerset Co., 7 July
1930, Moldenke 1330 (MBG, NY); South Amboy, July 1892, Halsted 175 (G, NY).
PENNSYLVANIA: about Tunkhannock, 18 July 1929, Osterhout (P); Meadville, Aug.
1893, Curtis (P); Harrisburg, 20 July 1888, Small (G); Lancaster, 29 Aug. 1900,
Heller (G); woodlands, Angora, 9 July 1899, MacElwee 882 (MBG); near Philadel-
phia, 29 July 1871, Redfield (MBG); near Philadelphia, 1890, Greenman 1376 (G,
MBG).
ELAWARE: marsh along Delaware River, south of Newcastle, 2 Aug. 1923,
Tidestrom 11572 (G).
ManvLAND: College Park, Symons (MBG); Chesapeake City, 11 July 1923,
Tidestrom 11413 (G)
District or COLUMBIA: gravelly and cindery artificial soil, northeast Washington,
24 July 1893, Boettcher 226 (G, MBG); Anacostia, 19 July 1893, Boettcher 176 (G,
MBG); Brookland, 19 July 1912, Holm (MBG); near Long Bridge, 11 July 1891,
Blanchard (MBG)
Vrana: Chisels Run, west of Williamsburg, 22 June 1921, Grimes 3794 (NY);
Reed Creek, Wythe Co., 23 July 1892, Small (MBG, NY); Marion, Smythe Co., 20
July 1892, Small (MBG
Nortu CAROLINA: Onobuim Station, 13 Sept. 1885, Britton (NY).
Sourn CAROLINA: near Anderson, 8 July 1919, Davis 9117, 8458 (MBG); Santee
Canal, Ravenel (G); Charleston, 19 Aug. 1859, Gibbes (NY).
FrLomipa: Apalachicola, Chapman (MBG); near Tallahassee, May 1843, Rugel
(MBG).
ALABAMA: Tensaw, 22 Aug. 1904, Tracy 8037 (G, MBG, NY).
Mississipri: Woodville, 7 Sept. 1887, Joor (MBG); Taylorville, 2 Aug. 1903,
Tracy (NY).
LourstaNa: without data, Hale (G); Natchitoches, 14 June 1915, E. J. Palmer
8000 (MBG); waste pps Alexandria, 5 June 1899, Ball 556 (G, MBG); Lockport,
23 Oct. 1919, Guidroz 3
Ouro: Shaker Pond, near RE 8 July 1896, Greenman 1380 (MBG); Oberlin,
June 1895, Hicks (MBG); Oxford, 6 June 1910, Overholis (MBG); Cincinnati, 18
Oct. 1930, Stephenson (MBG)
West Virama: Tygart River valley, above Huttonsville, 23 Sept. 1904, Green-
man 244 (G); banks of Blackwater River, Hendricks, 10 Sept. 1904, Greenman 246
(G); low ground west of Salt Pond, 28 June 1922, laa c 474 (G).
Inp1ANA: East Chicago, 10 Aug. 1910, Lansing 28
Kentucky: Stamping Ground, 1 July 1930, vw 308 (MBG); Rockdale, July
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 277
1928, Runyon 1260 (US); Nicholasville, 18 July 1923, McFarland 92 (MBG); Bowling
Green, Aug. 1896, Price (MBG); Calvert City, 16-19 June 1909, Eggleston 4841 (NY).
TENNESSEE: borders of woods, Knoxville, July 1898, Ruth 833 (MBG), 788 (NY).
Wisconsin: St. Croix Falls, July 1899, Baker (P); Green Bay, 7 July 1878, Schuette
(NY
InumNors: Bowmanville, Cook Co., 7 July 1896, Chase (MBG); Chicago, 19 Aug.
1907, Greenman 1877 (MBG); Starved Rock, June-Sept. 1921, Thone 50 (MBG);
Urbana, 6 July 1899, Gleason 718 (G); Olney, 17 July 1927, Ridgway 2831 (MBG);
near Alton, 24 July 1927, Bucholz (MBG); Freeburg, Aug. 1917, Hertel (MBG).
MINNESOTA: open grounds, Collegeville, Stearns Co., 29 July 1912, Chandonnet
(MBG); near Minneapolis, July 1891, Aiton 8467 (P); meadow, Northfield, 1912,
Goldsmith 119 (NY); Houston Co., July 1912, Freiberg (MBG).
Iowa: Black Hawk Co., 22 July 1929, Burk 591 (MBG); Ames, Hitchcock (MBG);
Ames, Sept. 1873, Bessey (G); Ames, Sept. 1909, Campbell 67 (G, MBG); Bentons-
port, July 1920, Graves 1691 (MBG); Decatur Co., 13 July 1903, Anderson (MBG).
Missounr: Hannibal, Marion Co., Davis 73, 127 4a, 3458, 8591, 4450, 4565 (MBG);
Ilaseo, Ralls Co., 13 Aug. 1915, Davis 3852 (MBG); Aberdeen, 11 Sept. 1911, Davis
950 (MBG, NY); St. Louis, Aug. 1838, Riehl 135 (MBG, NY); St. Louis, 11 Aug.
1894, Glatfelter (MBG); St. Louis, Sept. 1899, Baker (P); Creve Coeur Lake, 24 Aug.
1930, Kellogg 16277 (MBG); Fern Glen, St. Louis Co., 14 July 1906, Johnson (MBG);
Allenton, Letterman (MBG); Allenton, 27 July 1884, Kellogg (MBG); Kimmswick,
15 July 1885, F. Wislizenus 282 (MBG); Victoria, 6 July 1890, Hitchcock (MBG);
Shepard Mountain, near Arcadia, 22 July 1915, Greenman 3766 (MBG); Poplar
Bluff, 14 Aug. 1892, Dewart 15 (MBG); Jefferson City, July 1866, Knause (MBG);
Brumley to Bagnell, 21 Sept. 1897, T'release 719 (MBG); Springfield, 31 July 1892,
Dewart 18 (MBG); Willard, 9 July 1919, Blankinship (P); Swan, 24 Sept. 1899, Bush
452 (MBG); Jackson Co., 7 Aug. 1893, Bush (MBG); bottoms, Cass Co., 24 July
1864, Broadhead (MBG); Webb City, 4 July 1902, E. J. Palmer 198 (MBG); gravel
bars, Butler Creek, Noel, 9 Sept. 1913, E. J. Palmer 4229 (MBG
ARKANSAS: waste places, near Corning, 21 Aug. 1896, Eggert (MBG); near McNab,
5 Oct. 1923, Greenman 4412 (MBG); Fulton, 18 Sept. 1900, Bush 884 (MBG).
H Dakora: Brookings Co., 1903, Hia (MBG); Mitchell, July 1903,
Hoffstetter (MBG); sand hills near La Creek P. O., Bennett Co., 14 Aug. 1911, Visher
2263 (NY); Vermilion, 11 Sept. 1911, Visher (MBG P
NEBRASKA: Dickson’s Bluffs, on Missouri River, 12 July 1853-4, Hayden (MBG);
near Plummer Ford, Dismal River, 8 Aug. 1893, Rydberg 1716 (NY); Lincoln, Aug.
1899, Hedgcock (MBG); Franklin, 1893, Laybourne (MBG).
Kansas: Riley Co., 16 Aug. 1892, Waugh (MBG); low woods, Riley Co., 12 July
1895, Norton 389 (G, MBG, NY); Manhattan, Sept. 1893, Norton (MBG); vicinity
of Arkansas City, south of Arkansas River, 2 July 1929, Rydberg & Imler 482 (NY).
OKLAHOMA: near Alva, 11 July 1913, Stevens 1673 (G, MBG, NY); near Tonkawa,
4 Aug. 1913, Stevens 1821 (G); 16 km. south of Stillwater, 14 July 1927, Stratton 158
MBG); near Cleo, 19 July 1913, Stevens 1742 (G); vicinity of Fort Sill, 3 July 1916,
Clemens 11749 (MBG); Davis, 10 July 1916, Emig 716 (MBG); near Grant, 4 June
1916, Houghton 4015 (G); “Arkansas,” 23 July 1894, Bush 432 (MBG).
Texas: Tarrant Co., 10 Aug. 1926, Killiam 6933 (US); Dallas, Aug. 1876, Rever-
chon (G), ?34 (MBG); Houston, May 1842, Lindheimer 195 (MBG); Columbia, 13
Oet. 1900, Bush 1482 (MBG)
Verbena urticifolia is readily distinguished by petiolate and
[Vor. 20
278 ANNALS OF THE MISSOURI BOTANICAL GARDEN
rarely scabrous leaves, remotely flowered slender spikes, and
unconnivent fruiting calyces. On account of its close resemblance
to V. scabra the two were confused, and V. urticifolia formerly
was believed to be a native of tropieal America. It would now
appear to be indigenous in the Middle Eastern States, establishing
itself as à weed in various places.
16. V. hastata L. Sp. Pl. 20. 1753.
V. pinnatifida Lam. Tab. Encyc. 1: 57. 1791, as V. hastata
y Poir in Lam. Encyc. 8: 546. 1808.
V. paniculata Lam. Tab. Encyc. 1:57. 1791.
V. paniculata var. pinnatifida Schauer in DC. Prodr. 11: 546.
1847.
V. hastata var. pinnatifida Gray, Syn. Fl. N. Am. 2!: 330.
1878; Britton, Mem. Torr. Bot. Club 5: 276. 1894.
V. hastala var. paniculata Farwell, Ann. Rept. Comm. Parks
and Boul. Detroit 11: 82. 1900.
V. hastata f. rosea Cheney, Rhodora 4: 245. 1902.
V. hastata var. paniculata f. rosea Farwell, Papers Mich. Acad.
Sci. 2:37. 1923.
Stems 4-15 dm. tall, branched above, rough-pubescent with
short antrorse hairs; leaves lanceolate, oblong-lanceolate or
occasionally ovate-lanceolate, 5-15(-18) em. long, gradually
acuminate, petiolate, coarsely or incised serrate, often hastately
3-lobed at base, veins impressed above, rough-pubescent on both
surfaces; spikes strict, usually numerous in a panicle, pedunculate,
compact; bracts lanceolate-subulate, commonly a little shorter
than the calyx; calyx 2.5-3 mm. long, pubescent, lobes acute with
short subulate tips, more or less connivent; corolla-tube somewhat
longer than the calyx, pubescent without; corolla-limb 3-4.5
mm. broad; nutlets about 2 mm. long, nearly smooth or very
faintly striate; commissural faces muriculate or almost smooth.
LN Nova Seotia to British Columbia, south throughout the United
Sta
sail Scotta: bank of Five-mile River, Hants Co., 19 July 1920, Pease & Long
22350 (G).
New Brunswick: Nepisiguit, 30 July 1873, Fowler (MBG); alluvium of Nash-
waak River, Nashwaak, 31 July 1922, Fernald & Pease 25247 (G
Quesec: La Trappe, 5 Aug. 1926, Louis-Marie 141 (G); swamp, ‘Ascot, Sherbrooke
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 279
Co., 20 July 1923, koh (G); sand-plains, northwest of Three Rivers, 1 Aug. 1923,
Chenin & Knowlton (G).
Ontario: Ashdod, ss July ee Fowler (MBG); Kingston, Sept. 1897, Fowler (G);
Aylmer, 5 Aug. 1912, Fisher (MBG).
Marne: gravelly shore of St. John River, St. Francis, 5 Aug. 1893, Fernald (G);
Foxcroft, 18 July 1895, Fernald 296 (G, MBG); North Berwick, 28 Aug. 1891, Parlin
(G); Eliot, 27 Aug. 1895, Williams (G).
New Hampsuire: bank of “Connt” [Connecticut] River, 1894, Jesup (G); Suc-
cess, Coos Co., 24 Aug. 1908, Moore 4316 (G); roadside, Walpole, 28 July 1901,
Williams (G); roadside, East Jaffrey, 10 July 1897, Robinson 196 (G).
Vermont: Weybridge, 8 July 1908, Williams (G); fields, Brandon, 7 July 1921,
Dutton (MBG); Mechanicsville, 6 Aug. 1906, J. W. Anderson (G); Manchester, 6
July 1898, Day 149 (G); Brattleboro, 2 Aug. 1898, Robinson 153 (G).
MassACHUSETTS: by Merrimac River, Lowell, 8 Aug. 1927, Beattie (P); Melrose,
17 July 1876, Morong (MBG); Waverly, 10 Aug. 1878, Lane (G); South Framingham,
11 Aug. 1888, Sturtevant (MBG); Jamaica Plain, Faxon (G); Wigwam Pond, Dedham,
6 Oct. 1901, Williams (G): Nine Mile Pond, Cape Cod, 4 Sept. 1898, Greenman 398
(G, MBG); shore of Wequawket Pond, Centreville, 6 Sept. 1896, Williams (G);
Centreville, 20 Aug. 1902, Cheney (G); Nonquit, 29 Aug. 1888, Sturtevant (MBG);
Fisher’s Pond, West Tisbury, Martha’s Vineyard, 4 Sept. 1917, Seymour 1324 (G);
Monson, July 1897, Morris (MBG); near Water Shop Pond, Springfield, 5 Aug. 1924,
Seymour 511 (MBG); South Worthington, Hampshire Co., 17 Aug. 1912, Robinson
RHODE IsrAND: oec Olney (G); Providence, 16 July 1892, Collins (G);
dense wet thickets at borders of sphagnous swamps southwest of Harbor Pond,
Block Island, 19 Aug. 1913, Fernald & Long 10263 (G).
Connecticut: waste ground near river, Stafford, 9 July 1922, Weatherby 5076
(MBG); Southington, 10 Aug. 1897, Bissell (MBG); Southington, 31 July 1898,
Andrews (G); Oxford, 11 Aug. 1888, Harger (G); New Haven, 1858, Eaton (MBG).
New York: Granville, 22 Aug. 1924, Drushel 6351 (MBG); Port Chester, 3 Sept.
1886, Stabler (G); Willets Neck, Long Island, 8 Aug. 1853, Hexamer & Maier (G);
Peconic, 15 Aug. 1895, von Schrenk (MBG); Fleischmanns, Delaware Co., 15 Jul
1892, von Schrenk (MBG); Dryden, 3 Sept. 1915, Dean & Thomas 4874 (G); low
flats north of the PRÉ op 16 July 1915, MacDaniels 4873 (G); near Bool’s
Brook, Ithaca, Munz 617 (P); Keene Valley, Essex Co., 1 Aug. 1891, von Schrenk
(MBG); Canton, 17 E 1914, Phelps 799 (G).
New JEnsEY: Fort Lee, 30 July 1921, Rydberg (P); Plainfield, 9 Aug. 1877, T're-
lease (MBG); Watehühg: 11 July 1930, Moldenke 1339 (MBG); New Market, July
1892, Kelsey 173 (G).
PENNSYLVANIA: banks of Susquehanna River at Perdix, 4 Aug. 1926, Heller 14222
(MBG); near Harrisburg, 13 July 1888, Small (G); near Philadelphia, 1889, Greenman
1273 (G); shores of Schuylkill River, Fairmount Park, 22 July 1871, Redfield (MBG).
DELAWARE: along Delaware River, south of Newcastle, 2 Aug. 1923, Tidestrom
11551 (G).
Maryan: below Havre de iex Park, 2 Aug. 1902, Shull 156 (MBG); Ellicott
City, 3 Aug. 1916, Arséne 672 (M
District or COLUMBIA: near ed Bridge, Washington, 11 July 1891, Blanchard
(MBG).
Virainia: Langley, 10 Aug. 1901, W. Palmer (US); South Fork, Holston River, at
St. Clair’s Bottom, Smyth Co., 30 July 1892, Small (G, MBG).
[Vor. 20
280 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fromipa: Apalachicola, Chapman (MBG).
Onto: dry places, July, Riehl (MBG); alluvial soil, Windham Township, Portage
Co., 12 Aug. 1924, Webb 5441 (G); Shaker Pond near Cleveland, 8 July 1896, Green-
man 1379 (G, MBG); near Cleveland, Greenman 1377, 1882 (MBG).
West Virainia: near Tygart Junction, 24 Sept. 1914, Moore 2569 (G, P); Marlin-
ton, 29 July 1930, Berkley 1804 (MBG); Woods Co., 7 Aug. 1897, Pollock (MBG).
MicniaAN: meadow, Turin, 9 July 1901, Barlow (G); thickets along West Maple
River, west of Pellston, 7 Aug. 1917, Gates 10708 (MBG); near Trenton, 13 Aug.
ES Chandler (US); — College, 1890, Baker (P); South Haven, 4 Sept.
1911, Lansing 3340 (G); B n Harbor, 4 July 1911, Lansing 3241 (G); New
Buffalo, 20 July 1911, rani (G
DIANA: open woods, East lines: 10 Aug. 1910, Lansing 2805 (G).
TENNESSEE: Hollow Rock, 5 Aug. 1897, Eggert (MBG); along railroads, MeNairy
Co., July 1893, Bain 444 (G).
Wisconsin: along Wolf River, near Keshena, 13 Sept. 1925, E. J. Palmer 28738
(MBG); St. Croix Falls, Polk Co., July 1899, Baker (P); near Beef Slough, Buffalo
Co., 13 Aug. 1926, Fassett & Hotchkiss 3322 (G); near Mirror Lake, Sauk Co., July
1903, Eggert (MBG).
ors: meadow, State Street, Chicago, 4 Aug. 1907, Greenman 1953 (MBG);
old beaches, Lake Chicago, Chicago, 5 Sept. 1909, Steele 106 (MBG); waste ground,
39th and Stoney Island, Chicago, 12 July 1913, H. H. Smith 5654 (G); west of Kan-
kakee, 3 Aug. 1912, Sherff 1649 (MBG); waste land, Champaign, 31 July 1900,
Gleason 1945 (G); Beardstown, July 1842, Geyer (MBG); Mississippi bottoms,
Shepherd, Pike Co., 29 Aug. 1914, Davis 3565 (MBG); banks of the Mississippi,
opposite St. Louis, July 1842, Engelmann (M BG).
Minnesora: shore, De Soto Lake, Itaska Park, Becker Co., 13 Aug. 1929, Grant
3073 (G, MBG); fields, Collegeville, 29 July 1912, Chandonnet (MBG); Rockville,
July 1896, Campbell (P); Swan Lake, Minneapolis, July 1892, Ballard (G).
Iowa: Ames, Sept. 1873, Bessey (G); Ames, Sept. 1909, Pammel 78 (MBG);
ledges, Boone Co., 1898, Pammel 1806 (MBG); Grinnell, Aug. 1875, Jones (P);
Grinnell, Aug. 1877, Jones (G, MBG); Decatur Co., 13 July 1903, Anderson (MBG).
Missoumi: Medill, 24 Aug. 1920, Bush 9165 (MBG, P); Palmyra, 5 Oct. "h
Davis 8578, 8590 (MBG); Oakwood, Davis 3192, 3817 (MBG); St. Louis Co.,
Aug. 1877, Eggert (MBG, P); Creve Coeur Lake, 14 Aug. 1927, Kellogg 1131 dios
St. PM July 1839, Lindheimer (MBG); banks of Chouteau's Pond, St. Louis, Aug.
1, Engelmann (MBG); south banks of Chouteau's Pond, St. Louis, July 1842,
Races 337 (MBG); banks of River des Peres, near St. Louis, Sept. 1841, Engel-
mann (MBG); St. Louis, Aug. 1861, Engelmann (G); Allenton, 18 June 1896, Kellogg
(MBG); Courtney, 31 Aug. 1904, Bush 2214 (MBG); Webb City, E. J. Palmer 197,
198, 1085 (MBG); Neck City, 4 Aug. 1916, E. J. Palmer 10568 (MBG); Joplin, 10
July 1897, T'release 716 (MBG).
Manirosa: low ground, Muskeg Island, Lake Winnipeg, 11 Aug. 1884, Macoun
ie
TH Daxkora: peninsula of Lake Ibsen, 1 Aug. 1899, Lunell (G); Grand Forks,
10 pes 1893, Brannon 128 (MBG); edge of slough, Fairmount, 22 July 1912, Berg-
man 2335 (MBG).
Sours Dakora: Brookings, 1 Aug. 1893, Thornber (MBG); Oakwood Lakes,
Brookings Co., 27 July 1903, Johnson (MBG); Iroquois, 9 Aug. 1894, Thornber (G);
Forestburg, 18 July 1910, Visher 4439 (MBG).
NEBRASKA: Dickson’s Bluffs on the Missouri, 12 July 1853, Hayden (MBG);
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 281
prairies, Middle Loup River, near Thedford, 8 Sept. 1893, Rydberg 1515 (G); St. Paul,
24 July 1909, Bates (G); Lincoln, 9 July 1885, Webber (MBG); waste places, Nuckolls
Co., Aug. 1899, Hedgcock (MBG).
KANSAS: Olathe, Aug. 1892, Hitchcock (MBG); wet places, Riley Co., 16 July
1895, Norton 390 (G, MBG); banks of Joy Creek, about 8 km. from Osborne City, 11
July 1894, Shear 202 (G); Reno Co., 24 July 1899, White 174 (MBG).
OKLAHOMA: near Lamont, 2 Aug. 1913, Stevens 1791 (G, MBG); Doby Springs,
Harper Co., 20 Aug. 1927, Stratton ved (MBG); elose to edge of Lake Ivanhoe, near
M 11 Oct. 1913, Stevens 2901
Texas: 1.6 km. north of Canadian = on Amarillo-Dalhart Road, Oldham Co.,
27 Aug. 1921, Ferris & Duncan 8513 (MBG)
SASKATCHEWAN: without locality, 1857-8, Bourgeau (G).
Montana: Hot Springs, Flathead Lake, 8 Sept. 1908, Jones 8963 (P).
WvoniNa: Goose Creek, 10 < 1896, A. Nelson 2258 (G, MBG); Sheridan, 24
July a A. Nelson 8439 (MBG
OLORADO: Wray, 11-13 Aug. 1919, Eggleston 15551 (P); Timnath, 22 July 1901,
or (P); Fort Collins, 7 July 1897, Crandall 2014 (G); along Platte River,
Denver, 16 Aug. 1910, Eastwood 41 (G, MBG); foothills near Golden, 1 Aug. 1878,
Jones 523 (P); Brighton, Sept.-Oct. 1908, E. L. Johnston 507 (MBG); Georgetown,
24 July 1878, Jones (P); Canon City, 1872, Brandegee (MBG).
New Mexico: valley of the Mimbres, 1851, Wright 1498 (ANSP, G).
Arizona: Flagstaff, 1891, MacDougal 566 (NY); Hassayampa Creek, Fort Whipple,
6 June 1865, Cowes & Palmer 279 (MBG).
Ivano: Boise, 18 Aug. 1911, Clark 253 (G, MBG); Falk’s Store, Canyon Co., 28
June 1910, Macbride 304 (MBG).
Uran, Salt Lake City, 18 June 1878, Jones 487 (P); near Salt Lake City, 14 July
1902, Pammel & Blackwood 3638 (G); American Fork Canyon, 3 Aug. 1880, Jones
1487 (P).
Nevapa: Panaca, 5 Sept. 1912, Jones (P).
British CoLUMBIA: Alberni, Vancouver Island, July 1915, Carter (G).
WasniNGTON: Meyers Falls, 20 Aug. 1902, Kreager 469 (G); Alma, Okanogan Co.,
July 1897, Elmer 537 (MBG, P); foot of Priest Rapids, 17 July 1903, Cotton 1396
(G); Yakima region, 1882, Brandegee (MBG); Walla Walla, July 1898, Savage,
Cameron & Lenocker (MBG); Prosser, 14 July 1929, Gotfredson 90 (P).
OnEGOoN: near Pendleton, 11 Sept. 1896, Leiberg 2630 (G); Multnomah Co., July
1877, T. Howell 292 (G); Mount Scott, Multnomah Co., 15 Aug. 1902, Sheldon
11167 (G, MBG, P); low ground, Salem, 30 July 1917, J. C. Nelson 1804 (G).
CALIFORNIA: near San Joaquin River, Louis Park, Stockton, 26 Sept. 1927, Stan-
ford ?21 (P).
This rather distinct species shows a tendency to blend with
several members of the Verbenaca in the region of the Middle
States. Some botanists have regarded the variability in the
compactness and the profusion of the inflorescence, as well as
the tendency of the leaves to develop hastate lobes near the base,
as being of nomenclatorial value. Unfortunately in the rather
complete series of specimens at hand, the writer has not found
any combination of characters varying greatly in one direction;
[Vor. 20
282 ANNALS OF THE MISSOURI BOTANICAL GARDEN
and, at the present time, sees no reason to maintain other than
the specific category.
17. V. simplex Lehm. Ind. Sem. Hort. Hamb. 17. 1825; in
Nova Acta K. Acad. Leop. (Pugill. Pl. 1: 37) 14: 824. 1828;
Linnaea 3: Litt.-Ber. 10. 1828.
V. angustifolia Michx. Fl. Bor.-Am. 2: 14. 1803, not V.
angustifolia Mill. Gard. Dict. no. 15. 1768.
V. rugosa Muhl. ex Willd. Enum. Hort. Berol. 633. 1809, not
V. rugosa Mill. Gard. Dict. no. 18. 1768, nor V. rugosa D. Don
in Sweet, Brit. Fl. Gard. II. 4: pl. 318. 1838.
Stems chiefly erect, branched above, branches ascending,
sparsely pubescent or strigillose; leaves lanceolate or spathulate,
tapering into a subsessile base, 3-10 em. long, subacute or ob-
tusish, serrate or serrulate, reticulately rugose above and oc-
casionally scabrous, somewhat prominently veined beneath,
glabrate or sparsely strigillose on both surfaces; spikes solitary
at the apices of stems and branches, short- pedunoled, strict,
usually somewhat crowded; bracts lanosolateanbulate, commonly
shorter than the calyx, glabrate or glabrous; fruiting calyx 4
(-5) mm. long, sparsely pubescent, lobes acuminate; corolla-tube
scarcely longer than the calyx, with scattering hairs outside the
throat; corolla-limb about 6 mm. broad; nutlets trigonous, 2.5
mm. long, raised-reticulate above, striate toward base; com-
missural faces fully as long as the nutlets, muriculate-scabrous.
Distribution: Ontario and Vermont, south to Florida, west to Oklahoma and
Nebraska.
Specimens examined:
ONTARIO: rocky ground, Belleville, 20 June 1876, Macoun 1305 (G).
Vermonr: without locality, 5 July 1903, Blanchard 26, 60 (G), 161 (NY).
Massacuuserts: dry hillside, Sheffield, 27 Aug. 1902, Hoffman (G).
Connecticut: railroad cut, Fair Haven, 18 June 1886, Harger (G); New Haven,
Eaton (G); New Haven, 26 June 1884, Safford 216 (US).
New York: sandy field north of Tripoli, 25 July 1920, Burnham (G); Pine Plains,
1875, Hoyeradt (G).
Jersey: Scotch Plains, 13 Aug. 1877, Trelease (MBG); Camden Co., 22
June 1871, Parker (G); Atco, 22 June 1871, Redfield 6430 (MBG); Sicklerville, 21
June 1894, Brinton (G); Oakland, 25 July 1909, Mackenzie 4213 (MBG)
PENNSYLVANIA: without data, Muehlenberg (Bot. Mus. Berl.-Dahl. TYPE of V.
rugosa, MBG phot.); Philadelphia, Nuttall (G); Whiteland, Chester Co., 26 June
1910, Bartram 1021 (G); vicinity of Conewago, 28 May 1889, Small (US); York
Furnace, 28 June 1899, MacElwee 760 (MBG, NY); Mercersburg, Green (G)
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 283
Maryann: Great Falls, 6 Aug. 1905, House 1863 (MBG); Hurlock, 29 May 1919,
C. P. Smith 3186 (G).
District OF CoLuMB1A: Washington, 26 May 1889, Sudworth (G); Joy City, 19
June 1891, Blanchard (MBG).
Vrana: Bluemont, 25 May 1905, House 875 (US); open woods near Churchview,
5 June 1921, Leonard & Killip 534 (US); north of Williamsburg, 17 June 1921,
Grimes 8788 (NY); Bedford Co., June 1873, Curtiss (MBG); Chilhowie, 4 Aug. 1892,
Small (F, NY); about Cumberland Gap, Lee Co., 27 July 1892, Small (F, G, MBG,
US); Blue Ridge, 22 July 1891, Seymour 48 (G), 49 (MBG).
NonTH CAROLINA: near Chapel Hill, Ashe (US); Morgantown, 1872, Ruger (US);
near Biltmore, 17 June 1897, Biltmore Herbarium 4759b (G, NY, US); Madison Co.,
27 July 1880, J. D. Smith (US).
Sours CAROLINA: Abbeville District, July 1855, Hexamer & Maier (G).
Grorata: Dalton, 9 Aug. 1900, Harper 385 (NY, US); Chickamauga Creek, near
Ringgold, 6-12 Aug. 1895, Small (F, NY); between Taylor Ridge and Lafayette, 3
July 1900, P. Wilson 161 (NY, US).
Fronipa: without locality, 1842-49, Rugel 127 (F, MBG, US).
AMA: Cullman, 29 May 1892, Mohr (US); Blount Springs, 5 May 1898, Baker
(P); Blountsville, 6 May 1898, Baker (US); Choctaw Co., 13-15 Oct. 1896, Schuchert
(US).
Mississippi: Agricultural College, Oktibbeha Co., 11-17 Aug. 1896, Pollard 1312
(F, G, MBG, NY, P, US).
Onro: on limestone soil, Castalia, 27 July 1894, Moseley Er Sandusky, 28 July
1894, Moseley (F, US); Oxford, 16 June 1910, Overholts (MBG
West VIRGINIA: Shenandoah Junction, June 1891, Millepaugh 865 (NY); along
Roanoke River, south of Roanoke, 2 June 1891, Small & Heller 482 (G); roadside,
south of Williamsburg, 13 June 1921, Grimes 3716 (G); Sweet Springs, 14 Sept. 1903,
Steele 314 (G, MBG, NY, US).
InpIANA: near boundary line to Galien, St. Joseph Co., 15 June 1911, Nieuwland
2685 (MBG, US); about 3 km. west of Goldsmith, 9 July 1913, ae 13619 (MBG);
about 1.6 km. west of Palmyra, 22 June 1916, Deam 2035? (G, U
Kentucky: without data, Short (NY); Stamping Ground, 29 ae 1930, Singer
178 (US); Smithfield, 30 May 1909, Eggleston (NY); Nicholasville, 8 June 1923,
McFarland 109 (MBG); Mammoth Cave, Edmonson Co., May 1899, Palmer (NY);
Bowling Green, 21 May 1899, Price (MBG); near Franklin, 22 May 1926, Anderson
& Woodson 47 (MBG); Kuttawa, 27 Sept.-9 Oct. 1909, Eggleston 5237 (MBG, NY).
ENNESSEE: Carter Co., 8 July 1880, J. D. Smith (US); Knoxville, Ruth (F, P),
731 (US), 740 (NY), 765 (MBG); roadside at base of Chilhowee Mountains, Curtiss
1955 (F, G, MBG, NY); Chattanooga, 25 Aug. 1876, Engelmann (MBG); West Nash-
ville, 26-27 May 1909, Eggleston 4430 (MBG, NY, US); Kingston Spring, 5 Aug.
1897, Eggert (MBG); Hollow Rock, Carroll Co., 14 Aug. 1897, Bilimore Herbarium
4759 (G, MBG, NY, US).
ue Baraboo, 1861, Hale (G, MBG).
Inuinors: Stony Island, June 1911, Greenman 3640 (MBG); Stony Island, 25 June
1914, H. H. Smith 5923 (G, MBG); near 91st Street, Chicago, 28 July 1907, Greenman
1981 (MBG); South Chisage, 18 July 1913, H. H. Smith 5670 (G); Rock Island, 17
Aug. 1866, Engelmann (MBG); East St. Louis, 9 July 1898, Norton (MBG); bluffs
near Prairie du Pont, 20 June 1876, Eggert (MBG); Red Bud, 3 June 1888, Pammel
(MBG); Shawneetown, 26 May 1919, E. J. Palmer 15262 (MBG); Mound City,
June 1859, Vasey (G).
[Vor. 20
284 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Iowa: without locality, 17 July 1875, Arthur 24 (MBG); Cedar Falls, 12 July 1926,
Pammel 272 ys Iowa City, Hitchcock (MBG); Columbus Junction, 27 Aug. 1899,
Pammel 1699 (MBG).
Missounr: Hannibal, 27 June 1917, Davis 3588 (MBG); Ethel, 11 June 1915,
Bush 7586a (MBG); Silex, 29 May 1915, Davis 4552 (MBG); prairie, St. Louis, May
1833, Engelmann 336 (MBG); banks of Mississippi above St. Louis, Oct. 1841,
—— (M BG); St. Louis, Sept. 1841, Engelmann (G, US); mineral region south-
of St. Louis, June 1845, King (MBG); Pacific, 13 June 1897, Trelease 713
(MBG); Pacific, 3 June 1918, Greenman 4125 (MBG); Gray Summit, 16 June 1927,
Kellogg 1180 (MBG); Allenton, 6 July 1911, Letterman (MBG); near Sulphur Springs,
May 1927, Steyermark 292 (MBG); Crystal City, 14 May 1887, Trelease (MBG);
Washington, 25 June 1888, Pammel (MBG); Victoria, 8 July 1890, Hitchcock (MBG);
Jefferson Co., 20 June 1876, Eggert (MBG); near Hillsboro, June, Riehl 456 (MBG);
Hillsboro, 24 May 1885, F. Wislizenus 280 (MBG); near St. Genevieve, 27 Ma
1928, Greenman 4578 (MBG); Jerome, 24 May 1914, Kellogg 496 (MBG); Sheffield,
17 June 1915, Bush 7647 (G, MBG, US); Westport, 17 May 1896, Bush 914 (US);
Independence, 1 June 1895, Tindall (MBG); south of Cedar Gap, 22 May-3 June
1911, Lansing 2976 (F, G); about 2 km. west of Mansfield, 5-12 June 1911, Lansing
135 (F, G); Springfield, 31 July 1892, Dewart 42 (MBG); Willard, 24 July 1919,
Blankinship (P); limestone barrens, Ash Grove, 24 Aug. 1912, Standley 9336 (US);
Carterville, 8 July 1910, E. J. Palmer 2981 (MBG); Webb City, E. J. Palmer 200,
3319 (MBG); Swan, 24 Sept. 1899, Bush 450 (MBG); Galena, Stone Co., 27 Ma
1914, E. J. Palmer 5769 (MBG); Gainesville, 26 June 1928, E. J. Palmer 34766
(MBG); Eagle Rock, 24 June nv Bush 206 (MBG, US); Noel, McDonald Co., 9
Sept. 1913, E. J. Palmer 4232 (MBG
ARKANSAS: Beaver Station, Eureka Springs, Glatfeller (MBG).
NEBRASKA: Wahoo, June 1890, Rydberg 154 (NY).
ANSAS: Lawrence, Stevens (US); between Sereni var Hopes 20 June 1929,
Rydberg & Imler 120 (MBG); sterile soil, Anderson Co. , Hitchcock 791 (G,
MBG, NY, US); Mound City, 18 July 1887, Kellerman pe
OKLAHOMA: near Miami, 26 Aug. 1913, Stevens 2299 (G, ved near Copan, 15 Aug.
1913, Stevens 2080 (G); Foyil, 5 Aug. 1894, Bush 433 (MBG); on ereek bank, near
Pawhuska, 9 Aug. 1913, Stevens 1980 (G); Caddo, 20 range 1891, Sheldon 49 (US);
near Idabel, 18 May 1916, Houghton 3622, 3646 (G, MBG).
This species appears to be very closely related to V. hastata.
In its typical form, it is readily distinguished by its elongate
spikes and narrow leaves. Often, however, intermediate forms
occur between the two, making it somewhat difficult to find clear
lines of demarcation between them.
18. V. Orcuttiana Perry,'? n. sp.
5 V. Orcuttiana Perry, spec. nov., herbacea verisimiliter perennis; caulibus
erectis quadrangularibus sparse trábesbentibus ramosis; foliis lanceolato-ellipticis
spathulatisve in petiolum alatum attenuatis 4-6 em. longis grosse serratis lineato-
rugosis adpresso-pubescentibus supra subtusque reticulatis dense patento-hirtellis;
spicis pedunculatis compactis elongatisque; bracteis lanceolato-acuminatis sparse
ciliatis nervo medio et margine decurrentibus calyce brevioribus; calyce 4 mm. longo
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 285
Stems several from a common base, 4-angled in cross-section,
glabrous or very sparsely hirtellous, branching; leaves lanceolate-
elliptical to spathulate, tapering into a margined petiole (1-2
em. long), 4-6 em. long, decussate, coarsely serrate, rugose and
appressed-pubescent above, more densely spreading-pubescent
or hirtellous beneath and prominently veined; spikes pedunculate,
solitary or somewhat panicled, strict, mostly dense, elongate;
rhachis more or less angulate; bracts lanceolate-acuminate,
shorter than the calyx, midrib and margins more or less decurrent
along the rhachis, sparsely ciliate; calyx 4 mm. long, appressed-
pubescent and finely glandular, teeth short-subulate or acuminate,
more or less connivent above the schizocarp; corolla-tube about
as long as the calyx; corolla-limb 3-4 mm. broad; nutlets tri-
gonous, 2 mm. long, raised-reticulate at the apex, striate toward
the base; commissural faces extending to the tip of the nutlet,
muriculate-scabrous.
Distribution: Lower California.
Specimens examined:
Mexico:
Lower CALIFORNIA: Pinery, 27 July 1883, Orcutt (US); Hanson's Ranch, 29 July
1883, Orcutt (G, NY, US); table-lands, Hanson’s Ranch, 30 July 1883, Orcutt 909
G TYPE); mountains, northern Lower California, 8 July 1885, Orcutt (US); Sierra
Juarez, 12 July 1924, Gallegos 2342 (US)
This species, which has been passing as V. litoralis, is much
like V. neomexicana var. xylopoda in the finely glandular spike
and the angle of insertion of the flowers. It differs, however,
in the shorter nutlets, the smaller corollas, and the type of the
pubescence. In V. Orcuttiana, the trichomes are short and
somewhat hirtellous and the pubescence of the inflorescence is
closely appressed. In gross habit, it is scarcely to be distin-
guished from V. simplex, but the latter has somewhat harsher
pubescence and larger non-glandular flowers.
19. V. stricta Vent. Hort. Cels. 53, pl. 53. 1800.
V. Alopecurus Cav. Descr. 68. 1802.
adpresso-pubescente tenuiter glanduloso; calycis dentibus breviter subulatis vel
acuminatis; corollae tubo vix exserto; corollae limbo 3-4 mm. lato; coccis subtrigonis
2 mm. longis dorso striatis apice elevato-reticulatis; commissura muriculata.—Col-
lected on table-lands, Hanson's Ranch, Lower California, 30 July 1883, Orcutt 909
(G), TYPE.
[Vor. 20
286 ANNALS OF THE MISSOURI BOTANICAL GARDEN
V. rigens Michx. Fl. Bor.-Am. 2:14. 1803.
cuneifolia Raf. Med. Repos. N. Y. II. 5: 360. 1808.
stricta B ? mollis Torr. Ann. Lyc. N. Y. 2:234. 1827.
mollis Raf. Atl. Jour. 146. 1832.
stricta f. roseiflora Benke, Rhodora 34: 10. 1932.
stricta f. albiflora Wadmond, Rhodora 34: 19. 1932.
Stems 3-12 dm. tall, subterete, simple or branched above,
rather densely hirsute; leaves ovate or suborbicular, 6-10 cm.
long, sessile or nearly so, sharply and mostly biserrate, thickish,
hirsute and rugose above, densely hirsute-villous and prominently
veined beneath; spikes solitary or several, short-pedunculate,
thick, usually quite dense both in flower and in fruit; bracts
lanceolate-subulate, approximately as long as the calyx, hirsute,
ciliate; calyx 4-5 mm. long, densely hirsute, lobes acuminate;
corolla protruding slightly beyond the calyx, pubescent without;
corolla-limb 8-9 mm. broad; nutlets trigonous, 2.5 mm. long,
raised-reticulate above, strongly striate below ; commissural faces
reaching tip of nutlet, muriculate or almost smooth.
Distribution: eastern and central United States, from Pennsylvania westward
through the Rocky Mountains. Probably introduced into other localities.
Specimens examined:
(Herb. Bot. Gard. Madrid TYPE of V. Alopecurus, MBG phot.).
Massacuuserts: Fall River, 2 Sept. 1908, Sanford (G
Connecticut: waste ground, Nangatuck, 19 July 1908, Blewitt 14 (G); Bridgeport,
27 Aug. 1892, Hames (G).
New York: pasture south of Pulpit Rock, Ithaca, 2 Aug. 1919, A. J. Hames
12797 (Q).
New Jersey: ballast, New Durham, 15 July 1893, Van Sickle (US).
PENNSYLVANIA: along Lincoln Highway at Gap, Lancaster Co., Urban (G);
vicinity of Conewago, Sept. 1892, Small (NY); Conewago, Sept. 1892, Heller 638
(US).
hin ec Da
Onto: Dayton, Short (MBG).
MicniGAN: Constantine, 2 July 1923, Fisher 19 (MBG); Pine Lake, Charlevoix Co.,
18 Aug. 1917, Ehlers 642 (P).
InpIANA: East Chicago, 10 Aug. 1910, Lansing 2810 (G); Lake Maximkuckee, 9
Aug. 1889, Evermann 970 (US); sand pit northeast of Winona Lake, 2 Aug. 1897,
Deam (MBG); west of Palmyra, 22 June 1916, Deam 20357a (G
KENTUCKY: waste places, Hickman, 14 Aug. 1897, Biltmore Herbarium 3653a
(NY); Calvert City, Marshall Co., 16-19 aee 1909, Eggleston 4837 (NY); Wickliffe,
16 Aug. 1923, McFarland & Anderson 253 (MBG).
TENNESSEE: Hickmann, Gattinger (US); Henderson, June 1892, Bain 328 (NY).
Wisconsin: Mirror Lake, 15 July 1903, Eggert (MBG); Waupaca, 1907, Garesche
(MBG); Trempeleau, 1861, Hale (G); Oregon road, Madison, 29 July 1889, T'release
(MBG).
1933]
PERRY— NORTH AMERICAN SPECIES OF VERBENA 287
InnrNors: Stony Island, Chicago, H. H. Smith 5645, 6028 (G, MBG), 5952 (G);
Romeo, 26 July 1897, Umbach (MBG); Starved Rock, La Salle Co., June-Sept. 1921,
Thone 88 (MBG); Oquawka, 1872, Patterson (G, NY); Bloomington, July 1886,
Robinson (G); Carthage Township, 2 Sept. 1916, Gates 9994 (MBG); Decatur, 19
June 1896, Gleason 377 (G); Athens, Aug. 1863, Hall (MBG); East Hannibal, 13
June 1913, Davis 16 (MBG); Shepherd, 25 June 1915, Davis 6370 (MBG); Mississippi
River, bluffs north of Alton, 4 Aug. 1910, Sherff (G); Bonpas Township, 15 July 1925,
Ridgway 2431 (MBG); E. St. Louis, 28 July 1900, Eggert (MBG); Grand Tower, 22
Aug. 1900, Gleason (G).
Minnesota: Perham, Ottertail Co., 8 Aug. 1912, Chandonnet (MBG); St. Anthony,
7 July 1888, Schuette (G, NY); Willmar, July 1892, Frost (G); Morton, July 1890,
MacMillan (P); Lake City, 28 July 1883, Manning (G); Winona Co., Aug. 1901,
Holzinger (NY); Houston Co., July 1912, Freiberg (MBG).
owa: Fayette Co., 12 July-4 Sept. 1904, Fink 251 (US); Hawkeye, Aug. 1896,
Gardner 588 (NY); Cerro Gordo Co., 14 Aug. 1899, J'ones petal: Ames, 29 Aug. 1896,
Pammel 85 (G, MBG, NY); Grinnell, Aug. 1877, Jones (NY, P); Mount Pleasant,
7 July 1898, Ball 1585 (MBG); rn ig Aug. 1920, a 1947 (MBG); Ben-
tonsport, July 1920, Graves 1994 (M
Missouri: suburbs of Hannibal, pies 1202, 1613, 2791, 2960, 2962, 3227, 3589,
4461, 4487 (all MBG); Chain of Rocks, Aug. 1915, Beckwith 48 (MBG); St. Louis Co.,
15 July 1872, Redfield 522 (MBG); St. Louis, Aug. 1838, Riehl 196 (MBG, NY); St.
Louis, Aug. 1841, Engelmann (G); Allenton, Letterman (MBG); Williamsville,
Wayne Co., 27 June 1914, E. J. Palmer 6107 (MBG); Jerome, 16 June 1914, Kellogg
498 (MBG); Warsaw to Linn Creek, Camden Co., July-Aug. 1913, Emig 107 (MBG);
Cole Camp, Benton Co., 12 July 1897, Trelease 717 (MBG); railroad embankments,
Mansfield, 5-12 June 1911, Lansing 3164 (G); Randolph, 17 July 1898, Mackenzie
261 (MBG); Independence, 26 June 1895, Bush 475 (MBG, NY); Willard, 9 July
1919, Blankinship (P); Carterville, 8 July 1910, E. J. Palmer 2980 (MBG); Webb
City, 3 Aug. 1902, E. J. Palmer 199 (MBG); Turkey Creek, Joplin, 10 July 1897,
Trelease 718 (MBG); Swan, 25 Sept. 1899, Bush 670 (MBG).
ARKANSAS: waste places, Harvey 1958 (G, NY); Big Lake, 20-22 June 1911,
McAtee 2053 (NY, US); Jonesboro, Craighead Co., 4 July 1927, Demaree 3553
(MBG); Fayetteville, Harvey 61 (MBG); Fort Smith, 1853-4, Bigelow (US).
Sovru Daxora: Big Stone Lake, Roberts Co., July 1922, Over 14386 (US); Wind-
sor Township, Brookings Co., 27 July 1903, Johnson (MBG); Forestburg, 3 July 1910,
Visher 4450 (MBG); 16 km. south of Interior, 29 June 1929, E. J. Palmer 37627 (G,
MBG); near Fort Meade, Black Hills, 19 June 1887, Forwood 299 (US); canyons,
Lead, 9 Aug. 1913, Carr 118 (G, MBG, NY, US); Rapid City, 22 July 1912, Visher
1507 = Hot Springs, 3 Aug. 1892, Rydberg 982 (US).
EBR : Lincoln, 27 June 1885, Webber (MBG); E bel City, 26 Aug.
1926, Heller 14290 (MBG); South Fork of Platte, July 1 H. Engelmann (G,
MBG); Alma, 21 June 1897, Pammel (MBG); Anselmo, 8 m 1889, Webber (US);
Broken Bow, 7 July 1897, Pammel (MBG); Callaway, 27 June 1901, Bates (G);
Dismal River, south of Thedford, 27 June 1893, Rydberg 1422 (G, NY, US); North
Platte, July 1896, Plank (NY); Ogallala, 16 June 1925, Jones (P).
Kansas: Pottawatomie Co., 1895, Hitchcock 972 (US); prairie, Riley Co., 26 July
1895, Norton 391 (G, MBG, NY, US); Concordia, 24 July 1929, Benke 5164 (G, NY),
forma roseiflora; hill 8 km. from Osborne, 11 July 1894, Shear 191 (G); Florence,
28-30 July 1903, Griffiths 5050 (US); vicinity of Caney, 29 June 1929, Rydberg & Imler
433 (MBG, NY); Syracuse, 28 July 1893, C. H. Thompson 154 (G, MBG, NY, US).
[Vor. 20
288 ANNALS OF THE MISSOURI BOTANICAL GARDEN
OKLAHOMA: Verdigris, 2 Aug. 1894, Bush 435 (MBG); 12 km. west of Pawnee, 27
July 1927, Stratton 249 (MBG); near Tonkawa, 5 Aug. 1913, Stevens 1865 (G);
Doby Springs, 20 Aug. 1927, Stratton 408 (MBG); near Shattuck, 11 Oct. 1913,
Stevens 2930 (G); Norman, 10 Oct. 1914, Emig 365 (MBG); near Granite, 17 June
um Stevens 1084 (G); PEC Mountains near Davis, 23 June 1917, Emig 787
MBG); vicinity of Fort Sill, 16 Aug. 1916, Clemens 11748 (MBG); near Cache, 25
eg 1913, Stevens 135414 (G, MBG); near Grant, 2 June 1916, Houghton 4000 (G,
MBG, NY).
Texas: Lipscomb, 1 July 1903, Howell 64 (US); prairies, near Canadian, 11 Aug.
1900, Eggert (MBG); Dallas, Reverchon (G, NY), 735 (MBG).
ONTANA: bottoms, Crow Agency, 14 July 1901, Blankinship (G).
Wyomina: Whalen Canyon, 16 July 1894, A. Nelson 538 (G); Hartville, 20 July
1894, A. Nelson 505 (G, MBG).
Cotorapo: Wray, 1-4 July 1919, Eggleston 15224 (MBG); Denver, 2 Sept. 1910,
Eastwood 90 (G, MBG, US).
New Mexico: creek bottom, Santa Fe, 1847, Fendler 597 in a (ANSP, G).
WasurNGTON: Meyers Falls, 20 Aug. 1902, Kreager 475 (G, N Y, US).
A very distinct species somewhat incapable of sharp delimita-
tion on account of the tendency to hybridize with neighboring
species. Normally it is recognized by its stout compact spike,
imbricated flowers, and ovate-orbicular sessile leaves.
20. V. MacDougalii Heller, Bull. Torr. Bot. Club 26: 588.
1899.
V. MacDougalii mut. rosella Cockerell, Am. Nat. 36: 809.
1902
Stems 3-8 dm. tall, stout, obtusely four-angled, simple or
occasionally branched, cinereous-green, hirsute-pubescent ; leaves
oblong-elliptical or elongate-ovate, 6-10 em. long, short-petiolate
or narrowed into a subpetiolar base, coarsely and irregularly
serrate-dentate, hirtellous, rugose and minutely pustulate above,
densely pilose-pubescent and prominently veined beneath;
spikes solitary or sometimes several, short-pedunculate, thick,
comparatively dense both in flower and in fruit; bracts lanceolate-
subulate, for the most part noticeably longer than the calyx,
pubescent, ciliate; calyx 4-5 mm. long, rather densely pubescent,
lobes very obtuse, terminating in short subulate teeth; corolla-
tube scarcely protruding beyond the calyx; corolla- limb 6 mm.
broad; nutlets trigonous with convex back, 2.5 mm. long, raised-
tetioulate toward the distal end, strongly or faintly striate below;
commissural faces reaching tip of nutlet, muriculate or almost
smooth.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 289
Distribution: ae Wyoming to New Mexico and Arizona.
Specimens examined:
WYOMING: Platte! Canyon, Laramie Co., 2 July 1901, A. Nelson 8354 (MBG).
Cotoravo: Palmer Lake, 22 July 1895, Osterhout (NY); La Veta, 14 July 1896,
Shear 3577 (NY); Cucharas Valley, near La Veta, 20 July 1900, Vreeland 636 (NY);
Stonewall, July 1912, Beckwith 188 (NY); Arboles, 10 July 1899, Baker 565 (F, G,
MBG, NY, P, US).
New Mexico: without locality, 1847, Fendler 597 in part (MBG); Vermejo Park,
Colfax Co., 31 Aug. 1913, Wooton (NY); vicinity of Ute Park, Colfax Co., 2 Sept.
1916, Standley 14224 (NY); foot of Baldy Mountain, near Elizabethtown, Oct. 1898,
St. John (US), mut. rosella; near Sierra Grande Union, 18 June 1911, Standley 6065
(US); Jicarilla Apache Reservation, near Dulce, 20 Aug. 1911, Standley 8239 (US);
southeast of Cuba, 22 Aug. 1915, Read 19 (US); Sandia Mountains, 15 July 1914,
Ellis 258 (MBG, US); Santa Fe Co., 10 July 1847, Edwards (NY); Santa Fe Co.,
1889, Brandegee (MBG); Santa Fe Co., 20 July 1898, Greene 77 (NY); Santa Fe
Canyon, 3 Oct. 1913, Rose, Fitch & Parkhurst 17717 (US); Winsar’s Ranch, Pecos
River National Forest, 6 July 1908, Standley 4223 (F, G, MBG, NY, US); near
Pecos, 15 Aug. 1908, Standley 4927 (F, G, MBG, NY, US); west of Las Vegas, St.
John (P); Las Vegas, 26 June 1893, Mulford 39 (MBG); vicinity of Las Vegas,
Anect 57 (G), 211 (NY); mountains west of Grant’s Station, 2 Aug. 1892, Wooton
(NY, US); White Mountains, Lincoln Co., 26 July 1897, Wooton 208 (MBG, NY,
P, US); Cloudcroft, 1912, Stearns 342, 358 (US); Cloudcroft, Aug. 1920, Schulz 255
(NY); Mescalero Reservation, Sacramento Mountains, 21 July 1905, Wooton (US).
Arizona: Clark’s Valley, 1 Aug. 1883, Rusby (NY, US); South Fork, Little
Colorado River, Apache Forest, 23 Aug. 1920, Eggleston 17106 (NY); 16 km. south
of Rowe’s Point on the Grand Canyon, 5 Nov. 1899, Ward 11 (US); Flagstaff, 29
Aug. 1884, Jones (P); vicinity of Flagstaff, 8 July 1898, aire a 249 (ANSP, F,
G, NY TYPE); im pines, Flagstaff, 16 Aug. 1922, Hanson A148 (F, MBG).
Uram: Mammoth Creek, near head of Sevier River, 10 Sept. 1894, Jones 6026
(MBG, NY, P.
This southwestern representative of V. stricta is readily distin-
guished by the short-petiolate elongated leaves, the compact
spikes, and the floral bracts surpassing the calyx.
21. V. macrodonta Perry,!? n. sp. PI. 14.
19 V. macrodonta Perry, spec. nov., herbacea (basi ignota) verisimiliter perennis;
caule 1-1.5 m. alto erecto hirsuto-hispidulo ramoso; foliis elongato-ovatis basi
cuneata in brevem petiolum alatum attenuatis 10-14 em. longis biserratis vel mu-
cronulato-denticulatis supra scabro-hirsutis et inconspicue pustulatis subtus reticu-
latis hirsutis; spicis paniculatis glanduloso-hirsutis basi foliolatis compactis deinde
elongatis basi laxisque; bracteis lineari-lanceolatis subulatis ciliatis calyci subae-
quantibus; calyce 5 mm. longo glanduloso aliquantulum viscido-pubescente; calycis
dentibus 1.5 mm. longis subulatis; corollae tubo paulo exserto extus glabro vel
uberulo; corollae limbo 5-6 mm. lato; coccis subtrigonis 2 mm. longis dorso sulcatis
superiore parte tenuiter Birrae commissura muriculata.—Collected on the
road from Miraflores to San Bernardo Ranch in Sierra La Laguna, Lower California,
about 750 m. alt., 20 Jan. 1906, Nelson & Goldman 7425 (MBG), TYPE.
[Vor. 20
290 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Stem 1-1.5 m. tall, erect, branching, hirsute-hispidulous;
leaves elongate-ovate with cuneate base narrowed into a short
(1-2 em. long) margined petiole, 10-14 cm. long, coarsely and
sharply biserrate-dentate with mucronate teeth, rugose with
veins impressed and scabrous-hirsute above with minutely
pustulate hairs, prominently veined and hirsute beneath; spikes
panicled, subtended by leafy bracts, dense before anthesis,
becoming elongated and open in fruit, glandular-hirsute; floral
bracts lanceolate-linear, approximately equalling the fruiting
calyx, subulate, ciliate; calyx 5 mm. long, glandular, somewhat
viscid-pubescent, teeth 1.5 mm. long, subulate; corolla-tube
protruding a little beyond the calyx, glabrous or puberulent
without; corolla-limb 5-6 mm. broad; nutlets trigonous, 2 mm.
long, shallowly scrobiculate on the upper half, tending to be
sulcate toward the base; commissural faces muriculate.
Distribution: Lower California.
Specimens examined:
Mexico: Lower CALIFORNIA: road from Miraflores to San Bernardo Ranch in
Sierra La Laguna, about 750 m. alt., 20 Jan. 1906, Nelson & Goldman 7425 (MBG
TYPE, US).
Verbena macrodonta is a coarse plant with large thickish leaves
and open inflorescence, in a measure similar to V. MacDougalit,
but differing in its less strict habit, somewhat remote fruits, more
glandular calyces, and shorter plumper nutlets.
22. V. prostrata R. Br. in Ait. Hort. Kew. ed. 2,4:41. 1812.
V. lasiostachys Link, Enum. Hort. Berol. 2: 122. 1822.
Stems at first erect or ascending, at length spreading and
diffusely branched, sparsely villous; leaves ovate or oblong-ovate
with cuneate base tapering into a margined petiole, 5-10 cm.
long, commonly 3-cleft, coarsely and irregularly serrate-dentate
with apiculate teeth, or incised, veins impressed above, more or
less conspicuous below, both surfaces pilose to sparsely villous;
spikes solitary or more often loosely paniculate, dense before
anthesis, becoming elongated and open in fruit; bracts lanceolate-
subulate, not longer than the calyx, villous and at times finely
glandular; calyx 4-5 mm. long, villous or glandular-hirsute,
subtruncate, the distal end connivent above the schizocarp,
teeth subulate; corolla-tube a little longer than the calyx, very
1933]
PERRY—NORTH AMERICAN SPECIES OF YERBENA 291
sparsely pubescent outside or apparently glabrous; corolla-limb
3-5 mm. broad; nutlets oblong-trigonous with convex back,
2-2.5 mm. long, slightly thicker at the distal end, raised-reticulate
above, striate below, often with striae fading out toward the base;
commissural faces more or less muricate, approximately reaching
the tip of the nutlet.
Distribution: ae and California.
Specimens examin
OnEGON: dry um along railroad, about 1.5 km. north of Comstock, 18 June 1919,
J. C. Nelson 2669 (G); banks of Umpqua at Roseberg, 2 June 1928, Thompson 4412
(MBG); Glendale, 19 June 1902, Jones (P); Pleasant Creek, near Wimer, 26 July
1892, Hammond 322 (MBG, NY, US); Grant’s Pass, 24 June 1884, T. Howell 174
(US); Grant’s Pass, 3 July 1887, T. Howell (F), 1249 (MBG); gravel bar about 5 km.
above mouth of Rogue River, 8 July 1919, Peck 8702 (G, MBG, NY); in clearing
about hotel, Agness, 22 June 1917, J. C. Nelson 1429 (G); Dryden, 16 June 1904,
Piper 6160 (US); near Medford, Aug. 1922, Epling 5445 (MBG); south of Ashland,
19 May 1898, Applegate 2228 (US).
CALIFORNIA: near Yreka, Siskiyou, 19 June 1876, Greene 860 (F, G, MBG, NY);
Yreka, 2 July 1910, Butler 1621 (MBG, P, US); Mount Shasta, 13-27 July 1892,
Palmer 2529a (US); Soda Springs, Nevada Co., 30 July 1881, Jones 2598 (P); Round
Valley, Mendocino Co., 20 May-20 June 1898, Chestnut (US); near Clear Lake,
1865, Torrey 417 (G, NY); Clear Lake, Lake Co., 12 July 1929, Blankinship (MBG);
southern slope of Mount Sanhedrin, Lake Co., 19 July 1902, Heller 5919 (G, MBG,
NY, P, US); west of Windsor, near Russian River, Sonoma Co., 27 June 1902, Heller
5785 (F, G, MBG, NY, P, US); Oak Knoll, 5 May 1901, Braunton 378 (US); Irish-
town, Amador Co., June 1893, Hansen 964 (MBG); West Point Bridge, Calaveras
Co., 7 July 1896, Hansen 1823 (US); Stanford University, Santa Clara Co., April
1898, Abrams (P); San Jose, 14 July 1899, Pammel 187 (MBG); Santa Cruz, 21 June
1881, Jones (G, P); in pine woods, Pacific Grove, 25 May 1903, Heller 6778 (F, G,
MBG, NY, P, US); Carmel Bay, Sept. 1902, Elmer 4045 (G, P); Monterey, Hartweg
1924 (G, NY); Monterey, Aug. 1917, Parish 11590 (P); Santa Lucia Mountains,
May 1898, Plaskett 142 (NY, US); San Antonio River, 1880, Vasey (US); Wood’s
Creek, Fresno Co., 19 June 1910, Clemens (P); San Luis Obispo, 26 June 1876, Palmer
342 (MBG, NY, US); San Luis Obispo Co., 19 June 1887, Summers (MBG, P); Santa
Barbara Co., 1865, Torrey 416 (G, NY); Santa Barbara, May 1902, Elmer 3846
(MBG, P, US); seep, base of ocean bluffs, 32 km. northwest of Santa Barbara, 26
March 1925, Munz 9295 (P); Sulphur Mountain Spring, Ventura Co., 1-2 June 1908,
Abrams & McGregor 6 (G, NY, US); Ventura, 13 April 1923, Kendall (P); San
Buenaventura, March 1861, Brewer 229 (US); Los Angeles, 1860-62, Brewer 31 (G,
US); Los Angeles, 29 May 1891, Fritchey 29 (MBG); near Mesmer, 10 June 1917,
Johnston 1820 (P, US); near Pasadena, June 1893, Haynes (P); San Antonio Canyon,
28 June 1917, Johnston 1608 (P); edge of woods, Palomar Mountain, 11 Aug. 1918,
Spencer 994 (G, P); wayside, Mesa Grande, 2 June 1919, Spencer 1164 (G); Spencer
Valley, near Julian, San Diego Co., 20 June 1903, Abrams 3787 (F, G, MBG, NY, P);
bottom of canyon south of “Lemon Tank," west coast, San Clemente Island, 10
April 1923, Munz 6734 (P).
Although, from the original publication, one would naturally
[Vor. 20
202 ANNALS OF THE MISSOURI BOTANICAL GARDEN
infer that V. prostrata is Aiton’s species, it is in all probability
Robert Brown's. Aiton, in his acknowledgments (postscript to
the fifth volume of Hortus Kewensis), mentions the new matter
added by his friend Robert Brown, some without reference to
his name; more tangible evidence is furnished by Schauer, who,
in his monograph, indicates the specimen at Kew as V. prostrata
R. Br
The species is comparatively easy to distinguish by its de-
cumbent habit, soft villous pubescence, and elongated spikes.
It closely resembles V. robusta, which is much harsher and of
limited distribution.
The following specimens differ from the species in having
scarcely glandular inflorescence and fruiting calyx only 2.5-
3 mm. long shortly conniving beyond the fruit:
Catirornia: Three Rivers, Tulare Co., 9 July 1904, Culbertson 4210 (MBG, P);
San Bernardino, Sept. 1886, Parish & Parish 969 (F, US); near San Bernardino,
May 1894, Parish 2819 (MBG, P); Inglewood, Los Angeles Co., 31 May 1902,
Abrams 2481 (P); San Diego Co., 1875, Palmer 309 (G, MBG); southern California,
1876, Parry & Lemmon 842 (NY).
23. V. robusta Greene, Pittonia 3: 309. 1898.
Stems 6-9 dm. high, erect, paniculately branched above,
glabrate or sparsely hirsute; leaves ovate or oblong-ovate with
cuneate base tapering into a margined petiole, 4—7(-10) cm.
long, usually 3-cleft, irregularly serrate-dentate with apiculate
teeth, or incised, rugose and scabrous-pubescent above, less
harshly pubescent beneath with veins prominently reticulate;
spikes often crowded, subsessile, usually dense but occasionally
elongated; bracts lanceolate-subulate, more or less exceeding
the calyx in length, densely glandular-hirtellous; calyx 4 mm.
long, densely glandular-hirsute, lobes obtusish, terminating in
very unequal acuminate-subulate teeth; corolla-tube a little
longer than the calyx, puberulent without; corolla-limb 3-4 mm.
broad; nutlets oblong-trigonous, 2-2.5 mm. long, raised-reticulate
above with striae fading out toward the base; commissural
faces more or less muricate, approximately extending to the tip
of the nutlet.
Distribution: California and Lower California.
Specimens examined:
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 203
CALIFORNIA: vicinity of Ione, Dec. 1904, Braunton 1268 (MBG, NY); Tiburon,
Marin Co., 26 July 1900, Eastwood (G); valley back of Berkeley, 23 Oct. 1880,
Engelmann (MBG); West Berkeley, May 1887, Greene (US); salt marshes, Oakland,
Sept. 1886, Congdon (G); Temescal, Alameda Co., Aug. 1891, Michener & Bioletti 123
(G); Crystal Springs Lake, San Mateo Co., July 1903, Elmer 4950 (MBG, P, US);
San Luis Obispo, 1876, Palmer 34114 (F, G, NY, US); west Sherman Canyon, 18
June 1901, Braunton 99 (US); Valdez Bay, Santa Cruz Island, 5 Sept. 1927, Jones
(P); sandy soil near coast, Santa Catalina Island, 19 July 1915, Macbride & Payson
850 (G); Santa Catalina Island, 20-25 July 1917, Eastwood 6500 (US); dry stream-
bed, pebbly beach canyon, Santa Catalina Island, 13 May 1928, Dunkle 1955 (P);
Avalon, Santa Catalina Island, Sept. 1896, Trask (MBG); Avalon, May 1897,
Trask (F, US); Avalon, Aug. 1901, Trask (NY); Avalon, 28 April 1914, Carlson
(US); Avalon, 13 June 1915, Carlson (G, MBG); Jamuel Valley, San Diego Co., 26
June 1875, Palmer (G), 310 (F, MBG, NY)
Mexico: Lower CALIFORNIA: near Rancho Salina, foot of Guatay Grade, about
6 km. south of Rio Guadalupe, 11 Sept. 1929, Wiggins & Gillespie 3977 (MBG);
Santa Tomas, 15 July 1885, Orcutt 1301 (MBG, US).
Verbena robusta has been much confused with the nearly
related V. prostrata, but is quite readily distinguished from the
latter by the brighter green color of the herbage, the scabrous
upper surface of the leaves, and the usually dense spikes. More-
over, the mature calyx lacks the marked tendency toward sub-
connivent lobes, a characteristic of V. prostrata. The schizo-
carps of the two are very much alike. The collection Orcutt 1301
has a greatly elongated inflorescence and the leaves are not
particularly scabrous. The specimens Braunton 99 and 1263
show unusually long floral bracts. These are probably atypical
phases of the species or possibly hybrids.
24. V. xutha Lehm. Del. Sem. Hort. Hamb. 7, 8. 1834;
Linnaea 10: Litt.-Ber. 115. 1835-6.
V. strigosa Hook. Comp. Bot. Mag. 1: 176. 1836, not V.
strigosa Cham. Linnaea 7: 256. 1832.
V. Lucaeana Walpers, Rep. 4: 23. 1844-48.
Stems upright, 6-10 dm. tall, branched, hirsute-hispid; leaves
oblong or broadly ovate in outline, 5-8(-12) cm. long, incised-
pinnatifid or more often trifid with segments coarsely dentate,
the lateral much smaller than the middle segment and close to
the sessile base of the blade, strigose above, canescent and spread-
ing-hirsute below, trichomes particularly prominent along the
somewhat paler midrib and veins; spikes elongate, somewhat
strict, not compact except at anthek: short-pedunculate; bracts
[Vor. 20
204 ANNALS OF THE MISSOURI BOTANICAL GARDEN
lanceolate-subulate, subequalling or commonly a little shorter
than the calyx, strigillose, ciliate; calyx 3-4 mm. long, strigose-
hirsute, lobes acuminate-subulate; corolla-tube approximately
as long as the calyx, the throat pubescent without; corolla-limb
5-8 mm. broad; nutlets 2 mm. long, raised-reticulate on the upper
half, faintly striate below, commissural faces reaching the apex
of the nutlet, muricate or muriculately scabrous.
Distribution: Alabama to Texas.
Specimens examined:
ALABAMA: Navy Cove, Aug. 1889, Mohr (US).
Louisiana: without data, Hale (G, MBG), Drummond (G); vicinity of Alexandria,
9 June 1899, Ball 605 (MBG, US); Chopin, 6 May 1915, E. J. Palmer 7556 (MBG);
St. Martinville, 30 May 1893, Langlois (MBG, US); Pointe a la Hache, 25 June
1884, Langlois 123 (US); New Orleans, 1832, Drummond 253 bis (K TYPE of V.
wider yrs Schriever, 8 June 1917, Munz 1607 (P); Cameron, 4 July 1903, Tracy
8708 ( , MBG, NY, US).
RKANBAS: near Homan, 10 June 1898, Eggert (MBG); Texarkana, Aug. 1881,
Letterman (MBG, US).
Texas: Gladewater, 18 June 1901, Reverchon 2582 (MBG, NY); near Longview,
7 June 1899, Eggert (MBG); Corsicana, 3 Oct. 1900, Reverchon 2118 (MBG, US);
Huntsville, 1920, Warner 43 (US); sandy land, Brazos Co., 1899, Ness (G); Brazos,
July 1899, Lindheimer (MBG); dry banks of Brazos River, 23 June 1917, Munz 1470
(P); College Station, 24 July 1899, Reverchon (MBG); black land prairie, Montgomery
Co., 18-21 July 1900, Dixon 473 (F, G, NY); Burton, 26 May 1872, Hall 434 (F, G,
; , P, US); Austin, 12 July 1920, Tharp 667, 668 (US); Austin, Aug. 1921,
Schulz 679, 701 (US); San Marcos and vicinity, 1898, Stanfield (N Y); Industry, 1844,
Lindheimer 145 (MBG); Cypress City, May 1877, J. Ball (MBG); near Houston,
April 1842, Lindheimer 154 (G, MBG); swamps, Houston, Aug. 1904, Kuntze 23811
(NY); Lotus, about 16 km. west of Houston, 8 Aug. 1921, Ferris & Duncan 8268
(MBG); Strand, Jefferson Co., 9 April 1924, Tharp 3166 (MBG); Galveston, 6 June
1920, Fisher 212 (US); Columbis, 5 Oct. 1900, Bush 1275 (MBG); Columbia, 23
Sept. 1901, Bush 899 (MBG); Hallettsville, 9 Aug. 1912, Fisher 122 (US); Ottine,
30 Aug. 1926, Bogusch 1235 in part (P); Sequin, 17 June 1903, Groth 18? (F, G, NY,
US); Cave Lake, Jackson Co., 30 June 1915, Drushel 2842 (MBG)
This species is easily distinguished by its erect habit, coarse
pubescence, elongated open spike, and sessile commonly trifid
leaves. It has often been confused with V. canescens var.
Roemeriana, but the latter is a smaller and more compact plant.
25. V. plicata Greene, Pittonia 5: 135. 1903.
Coarse herb with stems decumbent to ascending, branched,
hirtellous; lower leaf-blades elliptic-ovate, narrowed into a
margined petiole of approximately the same length, 1-3(-4) em.
long, broadly obtuse, plicate, coarsely incised-dentate, often
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 205
3-lobed, more or less canescent, rugose and somewhat appressed-
hirsute above, hirsute beneath and prominently marked (particu-
larly near the margin) with whitish veins; upper leaves similar
but smaller, often appearing spathulate; spikes terminal on
stems and branches, ordinarily not compact; bracts ovate-
lanceolate, usually exceeding the calyx, at times barely equalling
it, acute, hirsute, midrib often noticeable; calyx 3.5-4 mm. long,
more or less glandular-hirsute, lobes very obtuse or subtruncate,
terminating abruptly in subulate teeth; corolla-tube scarcely
longer than the calyx; corolla-limb 4-6 mm. broad, anterior lobe
retuse; nutlets cylindric-trigonous, 2-2.5 mm. long, shallowly
scrobiculate above, changing to indefinitely striate toward the
base; commissural faces finely muricate-scabrous to practically
smooth, not reaching the tip of the nutlet.
Distribution: Texas, New Mexico, Arizona, and northern Mexico.
Specimens examined:
Texas: without data, Berlandier 2506 in part, 644 (MBG); without locality, Sept.
1881, Havard (US); southwestern Texas, Reverchon 118 (G); western Texas, 1851-2,
Wright 1496 (G, MBG, NY); sandy roadside, Chillicothe, 28 Sept.-3 Oct. 1906, Ball
1171 (US); sands, Estelline, 25 May 1904, Reverchon 4314 (MBG); sandy waste,
Garza Co., 6 June 1925, Ruth 1289 (US); Post, 22 May 1925, Wooton (US); Lubbock,
24 April 1930, Demaree 7539 (US); moist open ground along creeks, Sweetwater,
Nolan Co., 27 May 1918, E. J. Palmer 18730 (MBG); north of Colorado, Mitchell
Co., June 1900, Eggert (G, MBG); prairie north of Stanton, Martin Co., 13 June
1900, Eggert (MBG); rocky and sandy soils, Comanche, 8 Aug. 1877, Reverchon 834
in part (MBG); near Comanche, 10 May 1900, Eggert (G, MBG); rocky prairies,
rown Co., 10 Aug. 1877, Reverchon 737 in part (MBG); Brown Co., April 1882,
Reverchon 737 (US); San Angelo, 19 May 1908, Reverchon 1953 (MBG); Barstow, 14
April 1902, Tracy & Earle 30 (F, G, MBG, NY), Tyre collection; Barstow, 15 April-
3 May 1902, Tracy & Earle 41 (NY); Oxona, 13 April 1930, Jones 26221 (P); Davis
Mountains, 5 Aug. 1918, Young (P); cliffs back of Fort Davis, Davis Mountains,
9-12 July 1921, Ferris & Duncan 2726 (MBG, NY, P); about 5 km. east of Study
Butte, Brewster Co., 30 June 1931, Moore & Steyermark 5299 (MBG); Alpine, 7
June 1926, E. J. Palmer 30523 (MBG); near Boquillas, 17 April 1919, Hanson 619
(MBG, US); Austin, 12 May 1872, Hall 429 (F, G, NY); gravel bars of Blanco
River, Blanco, 5 April 1918, E. J. Palmer 13282 (MBG); near Feodora, Terrell Co.,
26 April 1928, E. J. Palmer 33537 (MBG, NY); Devils River, Valverde Co., May
1913, Orcutt 6235 (MBG); Corpus Christi Bay, Dec. 1879, Palmer 2038 (G); Laredo,
Aug. 1879, Palmer 2040 (G); Laredo, Berlandier 1485 (= 225) in part (G); Laredo, 21
March 1903, Reverchon 3904 (G, MBG, US); Laredo, 1913, Orcutt 5555, 5717 (MBG);
near Pharr, Hidalgo Co., 6 April 1931, McKelvey 1756 (G).
New Mexico: plains, Carrizoza, 8-19 May 1902, Earle 606 (NY); in valley near
Gray, 26 July 1900, Earle 427 (NY, US); neighborhood of San Miguel, 12 Aug. 1847,
Fendler 594 (G, MBG).
ARIZONA: desert prairie, north of Tucson, 24 April 1913, Greenman & Greenman 28
[Vor. 20
296 ANNALS OF THE MISSOURI BOTANICAL GARDEN
(MBG); Tucson, 2 May 1892, T'oumey 806 (US); slopes west of Tucson, 30 Dec. 1919,
Bartram 256 (ANSP); Cienega, near Pantano, 14 June 1881, Pringle (F, G, MBG);
valley near Camp Lowell, 8 June 1882, Pringle (ANSP, F, NY); near Fort Lowell,
15 Sept. 1900, Griffiths 1595 (NY).
Mexico:
Crimuanva: Colonia Juarez, Sierra Madre Mountains, 11 Sept. 1903, Jones (P).
Coanuita: Sierra Mojada Mountains, 19 April 1892, Jones 372 (P, US).
In habit V. plicata is somewhat similar to V. neomexicana, but
in inflorescence it strongly resembles V. canescens var. Roemeriana.
Its distinetive character is foliar and is most easily seen in the
basal and the lower stem-leaves. The leaf is obviously petioled,
often 3-lobed, and very shallowly incised-dentate; moreover, the
veins beneath are whitish and particularly prominent near the
margin owing to the plication of the leaf and the apparent
broadening of the veins in this region.
26. V. neomexicana (Gray) Small, Fl. Southeast. U. S. ed. 1,
1010. 1903, and ed. 2, 1913.
V. canescens var. neomexicana Gray, Syn. Fl. N. Am. 2!: 337.
1878.
V. officinalis var. hirsuta Torr. Bot. Mex. Bound. 2:128. 1859.
Plant slender; stems upright, branched, hirsute; leaves 1-5 cm.
long, pinnately cleft or almost parted, segments incised or coarsely
toothed, rugose, somewhat scabrous and finely pustulate above,
the veins more or less prominent beneath, hirsute on both sur-
faces; spikes solitary or tending to be panicled, usually short-
peduncled, hirsute; bracts lanceolate-acuminate, commonly not
longer than the calyx; calyx about 3 mm. long, hirsute-pubescent
and very slightly glandular, teeth short and subulate; corolla-
tube scarcely longer than the calyx; corolla-limb approximately
4 mm. broad; nutlets trigonous with convex back, 2 mm. long,
very shallowly reticulate-scrobiculate on the upper half, longi-
tudinally striate below; commissural faces extending to the tip
of the nutlets.
Distribution: ie and New Mexico.
Specimens examin
Texas: Fort Davis, 13 Sept. 1918, Young 1703 (US).
New Mexico: White Mountains, Lincoln Co., 12 Aug. 1897, Wooton 646 (NY);
Ruidoso Creek, Lincoln Co., 3 July 1895, Wooton (NY, US); Kingston, Sierra Co., 6
July 1904, Metcalfe 955 (US); borders of thickets, near Coppermines, 1851, Wright
149? (G tyre, MBG, NY, US); Pinos Altos Mountains, 1880, Greene (F, MBG);
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 207
bottom of Tierra Blanca Canyon, Gila Forest, 18 Aug. 1916, Chapline 609 (NY);
G S. Ranch, vicinity of Silver City, 27 Aug.-12 Sept. 1911, Holzinger (US);
Mogollon Mountains, on or near West Fork of Gila River, Socorro Co., 28 Au
1903, Metcalfe 612 (G, MBG, NY, P, US
The collections of Young 1708, Wooton 646, and Wooton
(collection of July 3, 1895) are by no means typical of the species,
but perhaps are better placed here than elsewhere. This species
appears to be very closely related to V. canescens and V. gracilis.
It differs from both, however, in the upright habit and the nutlets.
In both V. canescens and V. gracilis the commissural face does
not extend to the tip of the nutlet and the striae on the dorsal
surface are less conspicuous.
26a. Var. xylopoda Perry,” n. var.
Stem somewhat coarser; pubescence shorter, denser and more
glandular; calyx 4 mm. long, glandular-hirsute, teeth acuminate;
corolla-limb 6-10 mm. broad.
Distribution: ia eT California, and northern Mexico.
ecimens ex d:
ARIZONA: Clarkdale, 17 Sept. 1921, W. W. Jones 344, 346 (G); Rio Verde, 8 Sept.
1865, Cowes & Palmer 571 (MBG); Skull Valley, 28 April 1903, Jones (MBG, P, US);
near Oracle, 20 April 1930, Harrison & Kearney 6689 (NY, P); Chiricahua Moun-
tains, Cochise Co., 1 May 1894, W. W. Price (P); Paradise, Chiricahua Mountains,
16 Sept. 1907, Blumer 2170 (F, US); Chiricahua Mine, 21 Oct. 1907, Blumer 1804
(F, G, MBG, NY, US); Warren, Cochise Co., 20 May 1915, Carlson (US); Bisbee,
3 Oct. 1892, Mearns 1013 (US); near Fort Huachuca, 1882, Lemmon 2857 (G); Fort
Huachuca, Aug. 1892, Wilcor (NY); Huachuca Mountains, 3 Sept. 1903, Jones (P);
Ash Canyon, Huachuca Mountains, 6 Aug. 1909, Goodding 334 (G, NY); Huachuca
Mountains, 3 Sept. 1928, Harrison & Kearney 5796 (US); Santa Catalina Mountains,
27 July 1917, Munz 1149 (P); Sabino Canyon, Santa Catalina Mountains, 21 April
1922, Hanson A1130 (MBG TYPE); about 13 km. south of Vail, 31 Aug. 1903, Jones
(P); foothills of the Santa Rita Mountains, 11 May 1884, Pringle (ANSP, F, G, US);
Santa Rita Forest Reserve, 8-13 Sept. 1902, Griffiths 3431 (US); Santa Rita Range
Reserve, 12 May 1912, Wooton (US); slopes about Calabasas, 21 April 1908, Tides-
trom 872 (US); Baboquivari Mountains, 28 March 1927, Peebles, Harrison & Kearne
8790 (US); Baboquivari Mountains, 26 Sept. 1927, Harrison 4778 (US); Baboquivari
Mountains, 12 April 1928, Gilman 1120 (P).
CALIFORNIA: without locality, 1876, Palmer (G), 33914 (US).
Mexico:
Lower CALIFORNIA: Big Canyon of Tantillas Mountains, 10 Sept. 1875, Palmer
(G), sis (F, MBG).
20 Var. xylopoda Perry, var. nov., typicam simulans sed indumentum brevius
densiusque; calyce 4 mm. longo glanduloso-hirsuto; calycis dentibus acuminatis;
corollae limbo 6-10 mm. lato.—Collected on rocky slopes, Sabino Canyon, Santa
Catalina Mountains, Arizona, 21 April 1922, Hanson A1130 (MBG), TYPE.
or. 20
298 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Sonora: Guadalupe Canyon, 27 Aug. 1893, Merton 2042 (US); '"Niggerhead
Mountains, near monument no. 82," Aug. 1893, Mearns 1887, 1918 (US); Fronteras,
June 1851, Thurber 446 (G, NY).
CurIHuAnvA: Saint Eulalia Hills, 30 July 1885, Wilkinson (US); San Diego, 26
April 1891, Hartman 608 (G); rocky hills, near Chihuahua, 16 April 1885, Pringle 270
(ANSP, F, G, US); vicinity of Chihuahua, 8-27 April 1908, Palmer 52 (F, G, MBG,
NY, US); “between Sacramento and Chihuahua," 24 Aug. 1846, Wislizenus 150
(MBG); in the Sierra Madre, 21 June-29 July 1899, Nelson 6161 (G, US); near
Colonia Garcia, 29 July 1899, T'ownsend & Barber 192 (F, G, MBG, NY, P, US);
between Colonia Garcia and Pratt's Ranch, below Pacheco, 22-24 Aug. 1899,
Nelson 6271 (G, US).
This variety differs from the species in the denser and more
glandular pubescence and the larger corolla. The nutlets also
are slightly longer, with the reticulations somewhat deeper than
in the species, and the commissural faces hardly extend to the
tip of the nutlets. The specimens Nelson 6161, 6271 and Town-
send & Barber 192 closely resemble the above variety in in-
florescence, but are more like V. neomexicana in the long and
somewhat sparsely hirsute indument on the lower part of the
stem; Pringle 270 and Wilkinson approach variety hirtella.
26b. Var. hirtella Perry," n. var.
Plants densely canescent-hirtellous; leaves more or less shal-
lowly incised; bracts usually broadly ovate-acuminate; corolla-
limb about 8 mm. broad.
Distribution: Texas, New Mexico, and Coahuila, Mexico.
Specimens examine
Texas: rocky ii between Van Horn Wells and one 2 July 1852, Parry,
Bigelow, Wright & Schott (NY, US); Sivermore Peak, Davis Mountains, 9-12 July
1921, Ferris & Duncan 2607 (MBG, NY); sand bars of Sieb Davis Mountains, 11
July 1926, E. J. Palmer 30791 (MBG); Valentine, 28 April 1930, Jones 2622/ in part
(P); near Shafter, Presidio Co., 26 April 1931, McKelvey 2046 (G); Pinto Canyon,
near Ruidosa, 13 April 1919, Hanson 645 (G, NY, US); Chisos Mountains, 12 Aug.
1915, Young 112 (MBG); foothills of Chisos Mountains, 22 May 1928, E. J. Palmer
84065 (MBG "tyre, NY); gravelly mu north side of Chisos Mountains, 27 June
1931, Moore & tavernas 8277 (MBG).
New Mexico: Socorro Mountains, 11 July 1897, Herrick 715 (US); low moun-
tains west of San Antonio, 14 April 1908, Wooton 3852 (US).
Mexico: CoanuiLa: Sierra de Parras, Purpus 1094 (F, G, MBG, NY, P).
The pubescence of this variety is much finer and shorter than
?! Var. hirtella Perry, var. nov., planta dense hirtello-canescens; foliis plus minusve
breviter incisis; bracteis fere late ovato-acuminatis; corollae limbo circiter 8 m
lato.—Collected on ns foothills of the Chisos Mountains, Texas, 22 May 1928,
E. J. Palmer 84065 (MBG), TYPE.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 209
in the above. The leaves are not so deeply incised and often
tend to be elongated. The leaves of Purpus 1094 are so narrow
and shallowly incised that it appears superficially like V. perennis;
nevertheless, the character of the pubescence allies it with this
variety.
27. V. perennis Wooton, Bull. Torr. Bot. Club 25: 262. 1898.
Stems several from a woody base, divaricately ascending-
erect, more or less strictly branched, glabrate or often finely
glandular and slightly hispidulous with short stiff antrorse
hairs; leaves predominantly linear, 1-4 cm. long, entire or
pinnately few-lobed, erect-ascending, sparsely hispidulous, mar-
gins revolute; spikes terminal, pedunculate, slender-filiform,
elongate; T ovate, 1.5-3 mm. long, acute, hispidulous,
ciliate; calyx 4-5 mm. long, pubescence more abundant along the
nerves, lobes subequal, short, acute; corolla-tube slightly longer
than the calyx, pubescent; corolla-limb 5-7 mm. broad, lobes
repand; fruit more or less remote; schizocarp 3 mm. long, strongly
constricted along the lines of cleavage; nutlets subcylindric,
reticulate-scrobiculate except at base; commissural faces smooth
or slightly scabrous and not extending to the tip of the nutlet.
Distribution: Texas and New Mexico.
Specimens examined:
Texas: Glass Mountains, 25 Aug. 1925, Tharp 3682 (US); Loyola, 5 Nov. 1902,
Seler (G); Guadalupe Mountains, 1882, Havard 197 (G); Guadalupe Mountains, 13
Aug. 1916, Young (MBG); rough grassy slopes, McKittrick Canyon, Guadalupe
Mountains, 23 July 1931, Moore & Steyermark 3611 (MBG); Guadalupe Mountains,
8 Aug. 1931, Clarke 4250 (MBG).
New Mexico: Guadalupe, Oct. 1881, Havard (US); Queen, Aug. 1909, Wooton
(MBG, US); Queen, 12-20 Aug. 1924, Standley 40686 (US); Otero Co., 17 Aug. 1899,
Standley (US); White Mountains, Lincoln Co., 21 July 1897, Wooton 187 (G, MBG,
NY, P), type collection; Ruidoso Creek, White Mountains, 5 Aug. 1901, Wooton
(MBG); north of El Capitan Mountains, Lincoln Co., 31 Aug. 1900, Earle 387
(MBG, NY, P, US); Capitan, 8-19 May 1902, Earle 619 (NY); Gray, Lincoln Co.,
6 June 1898, Skehan 20 (F, G, MBG, NY, P, US); Berendo Creek, 13 May 1905,
Metcalfe 1568 (F, G, MBG, NY, US).
The relationship of this species is somewhat anomalous. The
lobing of the leaves and the character of the nutlets seem to ally
it with V. canescens and its relatives; whereas the pubescence
and the predominance of practically entire linear-oblong leaves
recall V. simplex. It could scarcely be confused with either,
[Vor. 20
300 ANNALS OF THE MISSOURI BOTANICAL GARDEN
since the character combination of an open spike, very narrow
leaves, and sparsely short-strigillose hairs is not found elsewhere
in the group under consideration.
28. V. gracilis Desf. Cat. Hort. Paris, ed. 3, 393. 1829.
V. remota Benth. Pl. Hartw. 21. 1839.
V. arizonica Gray, Proc. Am. Acad. 19: 95. 1883.
Low diffusely branched herb; branches decumbent to ascending,
more or less glandular, canescent, hirsute; leaves ovate, cuneate
at base and narrowed into a margined petiole, 1-3 cm. long,
incised-pinnatifid to pinnately cleft, segments oblong, subincised,
acutish, midrib and veins impressed, sometimes glandular,
hirsute particularly on the lower surface; spike terminal, slender,
more or less sparsely flowered, sessile, the lowermost flowers
often appearing in the axils of the upper leaves; bracts linear,
attenuate, gradually reduced in length toward the tip of the spike,
usually much longer than the calyx; calyx 3 mm. long, more or
less glandular, hirtellous, subtruncate, teeth minute and subulate;
corolla inconspicuous, the tube somewhat longer than the calyx,
the limb about 2-3 mm. broad; nutlets 1.5-2 mm. long, finely
scrobiculate from the apex practically to the base; commissural
faces muricately scabrous.
Distribution: Arizona, Utah, and Mexico.
Specimens examined:
(Par. Tyre, MBG phot.). Described from specimen cultivated in the Botanical
Garden of Paris.
Arizona: Gardiner's Spring, 25 June 1882, Pringle (ANSP, F, G, NY); Gulching
Ground, Pine Canyon, Chiricahua Mountains, 5 July 1907, Blumer 1612 (F, G, MBG,
NY, US); San Pedro River, Mexican Boundary Line, 12 Oct. 1892, Mearns 1116
(US); Tanner’s Cafion, near Fort Huachuca, 1882, Lemmon (G); Fort Huachuca,
1890, Patzky (US); Fort Huachuca, Wilcox (NY, US), 29 (US).
Uran: mesa east of Monticello, 25 July 1911, Rydberg & Garrett 9201 (NY).
Mexico:
Cuinuanua: waste places, Chihuahua, 26 May 1885, Pringle 54 (G); near Chihua-
hua, 7 May 1887, Pringle (F, MBG); vicinity of Chihuahua, 1-21 May 1908, Palmer
200 (US).
DvnaNao: city of Durango and vicinity, April-Nov. 1896, Palmer 135 (F, G,
MBG, NY, US), 911 (US); city of Durango, 1 Aug. 1898, Nelson 4593 (MBG, US);
Otinapa, 25 July-5 Aug. 1906, Palmer 456 (G, US).
AGUASCALIENTES: (Zacatecas according to the Kew specimen): Hartweg 174 (G,
K tyre of V. remota, NY).
San Luis Porost: region of San Luis Potosi, 1878, Parry & Palmer 722 (G, MBG,
US); valley of San Luis Potosi, 1876, Schaffner 720 (ANSP, G, MBG).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 301
QUERETARO: Queretaro, 2100 m. alt., July 1904, Kuntze 23444 (NY).
Hipauco: Nopala, 1 Aug. 1913, Salazar (US); bare hills above Pachuca, about
2500 m. alt., 23 July 1898, Pringle 7590 (F, MBG, P); dry calcareous mesas, near
Metepec Station, about 2500 m. alt., 22 June 1904, Pringle 13159 (F, G, US).
Mexico: vicinity of Mexico, ? Berlandier 578 (MBG, US); valley of Mexico,
Pedrigal, about 2200 m. alt., 1 Sept. 1896, Pringle 6539 (ANSP, F, G, MBG, NY,
US); Santa Fe, 26 June 1865-6, Bourgeau 361 (G); near Tlalpam, 15 July 1901,
Rose & Hay 5488 (US); near Tlalpam, July 1905, Rose, Painter & Rose 8495 (G, US);
lava beds, Tizapan, about 2250 m. alt., 24 Aug. 1900, Pringle 9135 (G, MBG,
NY).
Oaxaca: near Mitla, June 1888, Seler 22 (G).
V. gracilis is probably a relative of V. canescens and V. neo-
mexicana. It is easily recognized by its slender habit, usually
long-attenuate bracts, tiny flowers, and essentially scrobiculate
nutlets. The specimen from Utah is atypical, but for practical
purposes seems better referred here.
29. V. canescens HBK. Nov. Gen. et Sp. 2: 274, pl. 136.
1818.
Low coarse herb; stems several and branched, decumbent to
ascending, canescent, hirsute; leaves oblong-lanceolate, 1-5 em.
long, acute, remotely incised-dentate or subpinnatifid, contracted
into a margined entire semiamplexicaul or petiolar base, rugose
and hirtellous above, the trichomes often with minute bulbous
bases, canescent-hirsute and somewhat conspicuously veined
beneath, margins revolute; spikes sessile or short-peduncled,
mostly solitary, loose-flowered or compact, glandular-hirsute;
bracts lanceolate, acuminate, variable in length, often exceeding
the calyx, hirsute, ciliate; calyx about 3 mm. long, subtruncate,
with very short inconspicuous teeth; corolla-tube slightly longer
than the calyx; corolla-limb 4-6 mm. broad, lobes retuse; nutlets
subcylindrical, 2 mm. long, raised-reticulate above, longitudinally
striate at least half way from base to apex; commissural faces
muricate-scabrous, not reaching the tip of the nutlets.
Distribution: Nevada and Mexico.
Specimens examined:
Nevapa: Caliente, 9 May 1892, Jones 554 (P).
Mexico:
CoanuirLa: Saltillo and vicinity, 15-30 April 1898, Palmer 25 (F, G, MBG, NY,
US); Saltillo, 1200 m. alt., 22 Aug. 1913, Adole 22 [Arsène 10626] (F, G, MBG, NY,
US); valley near Saltillo, 1848-49, Gregg 99, 246 (MBG); canyon and elevated portion
of Sierra Madre, south of Saltillo, 25 July-1 Aug. 1880, Palmer 1047 (ANSP, G, US);
[Vor. 20
302 ANNALS OF THE MISSOURI BOTANICAL GARDEN
near Buenavista, 22 ry "e^ Gregg (NY), 740 (MBG); near San Pablo, 29 April
1847, Gregg (NY), 645 (MBG).
ZACATECAS: near Conception del Oro, 11-14 Aug. 1904, phe 803 (F, G, MBG,
NY, US); Zacatecas, 1800-2100 m. alt., Aug. 1903, Purpus 467 (US).
AGUASCALIENTES: stony places, rim Calientes, Hartweg 177 (G, NY); near
Aguas Calientes, 20 Aug. 1901, Rose & Hay 6212 (NY, US); near Ape Calientes, 9
Oct. 1903, Rose & Painter 7741 (G, NY, US).
San Luis Porosi: Angostura, June 1911, Purpus 6515 (F, G, NY, US); in sands
around San Luis Potosi, 1876, Schaffner 719 (ANSP, G); San Luis Potosi, 1878,
Rose, Painter & Rose 8971, 9162 (US).
EBLA: Cerro Guadalupe, 20 June 1909, Nicolas (US); margin of Atoyac, 20
June 1910, Nicolas (US); barren hills about Esperanza, 2660 m. alt., 17 Aug. 1905,
Pittier bed (US
VER UZ: Ceiba, M Mueller 1215 (NY); Esperanza, 2400 m. alt., 14 Aug.
1891, Boson 822 (F, G, U
: Cerro de r Ae 16 Oct. 1921, Conzatti 4288 (US); dry calcareous
hills, Las Sedas, 1800 m. alt., 11 Aug. 1894, Pringle 4784 (ANSP, G, MBG, NY, US);
Las Sedas, about 1900 m. alt., 27 June 1895, L. C. Smith 412 (G); Las Sedas, 1950 m.
alt., 30 Aug. 1921, Conzatti 4194 (US).
This is a low foliose species closely related to V. neomexicana
and V. plicata, but distinguished from them by the semiamplexi-
caul elongate leaves and rather characteristic compact habit.
The specimens cited from Coahuila are excellent examples of
phases intermediate between the species and the variety. The
collection from Nevada seems somewhat out of range. Possibly
it was carried in as a weed or escaped from cultivation.
29a. Var. Roemeriana (Scheele) Perry, new comb.
V. Roemeriana Scheele, Linnaea 21: 755. 1848.
Plants more densely hirsute, sparsely (if at all) glandular;
floral bracts broadly ovate at base, abruptly acuminate, concave
at anthesis, later slightly recurved, for the most part overtopping
the calyx; dorsal surface of the nutlets reticulate-scrobiculate
approximately two-thirds of their length, striate toward the
base, nutlets usually more slender than in the species.
Distribution: Texas and Mexico.
)ecimens examine
Texas: without ioonlity:, 1846, Lindheimer 500 (G, MBG, US); without locality,
1885, Reverchon 737 in part (MBG); Red River, Wichita Co., 10 April 1922, Tharp
1361 (US); sandy soils, Brown Co., Reverchon 1961 (G, NY); near Brownwood,
Brown Co., 1 Nov. 1925, E. J. Palmer 29535 (MBG); Tom Greene Co., 1879, Tweedy
(NY); Dove Creek, Tom Greene Co., May 1880, Tweedy 118, 246 (US); San Angelo,
13 May 1903, Reverchon 3903 (MBG); Legion Creek, Gillespie Co., Jermy 182 (MBG);
gravel bars of Blanco River, Blanco, 5 April 1918, E. J. Palmer 13283 (MBG);
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 303
Austin, 16 May 1872, Hall 430 (F, G, MBG, NY, P, US); Austin, 1 April 1918,
Young 112 (MBG); San Antonio to Austin, 18 April 1925, Small & Wherry 11969
(NY); Dryden, 22 April 1930, Jones 26226 (MBG, P); Langtry, Valverde Co., May
1913, Orcutt 6111 (MBG); rocky hills at Devils River, 10 Sept. 1900, Eggert (MBG);
Lacey's Ranch, Kerr Co., 1 June 1916, E. J. Palmer 10002 (MBG); Kerrville, 7-14
May 1894, Heller 1782 (G, MBG, NY, US); San Marcos, 6 Nov. 1897, T'release
(MBG); New Braunfels, April 1850, Lindheimer 294 (MBG), 1074 (F, G, MBG, NY,
US); Ozona, 30 April 1930, Jones 26222 (P); Rock Springs, 17 April 1930, Jones
26223 (P); Del Rio, 20 April 1930, Jones 26225 (P); Fort Clark, Kinney Co., 27 Feb.
1893, Mearns 1237 (US); Fort Clark, 10 May 1893, Mearns 1456 (US); gravel and
sand deposits along stream, Uvalde, 11 May 1918, E. J. Palmer 13563 (MBG); dry
rocky ground along small stream, Uvalde, 24 April 1928, E. J. Palmer 33605 (MBG);
near Uvalde, 30 April 1928, E. J. Palmer 33646 (NY); 16 km. west of Uvalde, 8
June 1931, Moore & Steyermark 3005 (MBG); near Verde Creek, near Hondo, San
Antonio to Del Rio, 21 April 1925, Small & Wherry 11980 (NY); San Antonio, April
1911, Clemens & Clemens 9?4 (MBG, P), 971, 972, 973 (P); San Antonio, 16 March
1916, E. J. Palmer 9188 (MBG); on limestone hill, near Bracken, 27 June 1903, Groth
16 (F, G, NY); in fields near Rio Frio, July 1829, Berlandier 2054, 644 in part (G);
El Jardin, 10 March 1924, Runyon 629 (US); woods near Colorado River, near
Wharton, 12 April 1925, Small & Wherry 11826 (NY); stony hills near Goliad, 9
April 1900, Eggert (G, MBG); near Mathis, San Patricio Co., 5 April 1931, McKelvey
1710 (G); west San Diego, 31 July 1931, Clark 4041 (MBG).
Mexico:
NveEvo Leon: Walnut Grove, 27 May 1847, Gregg 791 (MBG, NY); Monterey, 26
May 1847, Wislizenus 825 (MBG); Monterey, 17-26 Feb. 1880, Palmer 1044 (ANSP,
G, US); Pico Chico, near Monterey, 19 March 1900, Canby 194 (US); Monterey, 26
July 1926, Fisher 235 (US).
TAMAULIPAS: vicinity of Victoria, 320 m. alt., 1 Feb.-9 April 1907, Palmer 82
(F, G, MBG, NY as no. 8, US); from Santander to Victoria, Nov. 1830, Berlandier
827 (G, MBG, NY); Jaumave, 1931, von Rozynski 13, 17, 185 (F).
Differing from the species chiefly in its coarser less glandular
pubescence and the broadly ovate abruptly acuminate bracts.
A few of the above-cited specimens show transitional phases,
but, as a whole, are closer to the variety.
30. V. subuligera Greene, Pittonia 1: 156. 1888.
Stems probably several from a common base, procumbent,
hirsute-pubescent; leaves ovate, gradually narrowed into a sub-
petiolar base, 1.5 to 4 cm. long, pinnately cleft, larger segments
incised or sharply dentate, appressed-hirsute on both surfaces
but especially beneath on the somewhat prominently reticulated
veins; spikes short-petiolate, elongate, conspicuously bracted,
in fruit interrupted; bracts lanceolate-subulate, twice as long as
the calyx, ascending-spreading, reflexed in age, hirsute; fruiting
calyx about 3 mm. long, pubescent, hirsute along the nerves,
[Vor. 20
304 ANNALS OF THE MISSOURI BOTANICAL GARDEN
lobes very short, strongly connivent over the schizocarp; corolla-
tube scarcely longer than the calyx; nutlets subcylindric-trigonous,
2 mm. long, only faintly striate; commissure muriculate or
smoothish, not reaching the top of the nutlet.
Distribution: Mexico.
Specimens examined:
Mexico: DunaNGo (?): "from the Sierra Madre, west of Durango," about 2400
m. alt., Sept. and Oct. 1881, Forrer (ANSP, F, G, NY, US), Tyre collection.
Although the general habit of this species is much like that of
V. bracteata, the latter differs in having obtusish leaf-segments
and sharply reticulate-striate nutlets with commissural faces
fully as long as the nutlet.
31. V. bracteata Lag. & Rodr. in Anal. Cienc. Nat. 4: 260.
1801.
V. bracteosa Michx. Fl. Bor.-Am. 2: 13. 1803.
V. squarrosa Roth, Catalect. Bot. 3: 3. 1800.
V. canescens Chapman, Fl. Southeast. U. S. 307. 1860.
V. bracteosa var. brevibracteata Gray, Syn. Fl. N. Am. 2!: 336.
1878.
V. rudis Greene, Pittonia 4: 152. 1900.
V. confinis Greene, Pittonia, l. c.
V. bracteosa var. albiflora Cockerell in Daniels, Fl. Boulder,
Colo. 204. 1911.
V. imbricata Wooton & Standley, Contr. U. S. Nat. Herb. 16:
166. 1913.
Zapania bracteosa Poir. in Lam. Encyc. 8: 843. 1808.
Stems usually several from a common base, diffusely branched,
decumbent or ascending, rarely erect, coarsely hirsute; leaves
1-4(-6) em. long, pinnately incised or usually 3-lobed (lateral
lobes narrow and divaricate, middle lobe large, cuneate-obovate,
incisely toothed or cleft), narrowed into a short margined petiole,
hirsute on both surfaces, midrib and veins slightly prominent
beneath; spikes terminal, sessile, comparatively thick, conspicu-
ously bracted; bracts much longer than the calyx, spreading-
ascending, recurved in age, coarsely hirsute, the lowermost often
incised and leaf-like, the upper linear-lanceolate, acute-acuminate,
entire; calyx 3-4 mm. long, hirsute particularly along the nerves,
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 305
lobes very short, connivent over the schizocarp; corolla-tube
protruding slightly beyond the calyx, very finely pubescent
outside the throat; corolla-imb 2.5-3 mm. broad; nutlets tri-
gonous, approximately 2 mm. long, sharply raised-reticulate
above, striate below; commissural faces reaching to the distal
end of the nutlet, muricately scabrous.
Distribution: waste places, Maine to Florida, west to California, and northern
exico.
Specimens examined:
(Herb. Bot. Gard. Madrid Tyre, MBG phot.). Described from specimen culti-
vated in the Botanical Garden of Madrid.
Ontario: Point Edward, Lake Huron, 19 July 1901, Macoun (NY).
Maine: Cumberland, 12 July 1902, Chamberlain 418 (G).
MassAcHUSETTS: Dedham, 4 Aug. 1900, Rich (G); Worcester, 27 June 1904,
Dewhurst (G).
Connecticut: Winsted, Winchester, 5 Sept. 1909, Fernald (G); Bridgeport, 4
Sept. 1908, Hames 8137 (G
New York: Montauk Paint. 23 July 1895, von Schrenk (MBG
New JEnsEY: Weehawken, Aug. 1894, Van Sickle (US); Kaisa Point, Camden,
20 July 1866, Parker (G, MBG).
PENNSYLVANIA: "Cementon," valley of Lehigh River, 20 Aug. 1923, Churchill
(G); Lancaster, 29 Aug. 1900, Heller (US).
Nortu Carona: mountains of North Carolina, June 1872, Leroy & Ruger (NY);
Statesville, June 1872, Ruger (US
GEORGIA: Cartersville, Ravenel (NY); Thompsons Mills and vicinity, Gwinnett
Co., 14 May 1908, Allard 207 (MBG, US); Stone Mountain, 24 May 1897, Eggert
MBG); Ocmulgee River, swamp palen Macon Island, 8-9 July 1895, Small (NY);
Smithville, 18 Aug. 1885, J. D. Smith (MBG).
LORIDA: without data, Buckley (MBG), Chapman (MBG); Apalachicola, 5 Aug.
1889, Biltmore Herbarium 1082b (NY, US).
ALABAMA: roads and uncultivated fields, Sept. 1843, Rugel (MBG); Stevenson,
5 Sept. 1877, Ward (US); Valley Head, July 1898, Ruth 495 (NY), 518 (MBG);
Wilcox Co., 18 May 1840, Buckley (NY); sandy ground, near Atalla, Etowah Co.,
9 July 1898, Eggert (MBG, NY); Auburn, 13 May 1898, Earle & Baker (P, US);
Mobile, 22 May 1884, Mohr (US).
LouisiANA: vicinity of Alexandria, 5 June 1899, Ball 553 (G, MBG, NY, US).
B Sandusky, 14 Aug. 1920, Moseley (G).
Micuiaan: Vincent Lake, Cheboygan Co., 15 Aug. 1917, Ehlers 627 (G); Manistee,
8 Aug. 1882, Morong 1O D
INDIANA: of Bass Lake, Steuben Co., 25 July 1906, Deam 1272 (MBG,
NY, US); Michigan City, 6 July 1903, Mell & Ea (MBG).
Kentucky: without data, Rafinesque (G); southern Louisville, 19 Aug. 1892,
—€— vog
ENNE : Kin ngston Spring, 19 Aug. 1897, Eggert (MBG); Sherwood, 9 June
1897, oed (MBC): Memphis, 22 June 1851, Fendler (G).
Wisconsin: Fort Howard, 22 July 1887, Schuette (G, US); Madison, 27 Aug. 1893,
Churchill (G); near Mirror Lake, Sauk Co., 13 July 1903, Eggert (MBG).
[Vor. 20
306 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Inurors: Fountaindale, 1887, Bebb (G); Elgin, 27 Aug. 1912, Sherf 1797 (MBG);
Havana, 18 Aug. 1904, Gleason (G); Monticello, 17 June 1886, Seymour & Waite
(G); Athens, Hall (US); East St. Louis, 11 June 1879, Eggert (P).
Minnesota: Princeton, July 1892, Sheldon (G, US); Collegeville, 29 July 1912,
Chandonnet (MBG); hills between Minnehaha and Fort Snelling, 5 July 1888,
Schuette (G); Fort Snelling, 26 July 1899, Mearns 524 (US); Lake City, 25 July 1883,
Manning (G).
Iowa: Lawler, 1890, Rolfs (G); Fayette Co., 30 July 1894, Fink (US); Black Hawk
Co., 15 July 1929, Burk 542a (MBG); Ames, 5 Aug. 1896, Ball (G, MBG, NY, US);
‘lene, Aug. 1904, Fawcett 12 (G, MBG, NY, US); Council Bluffs, Vasey (G).
Missourtr: Hannibal, Davis 2691, 3587, 9089 (MBG); Eolia, 26 May 1914, Davis
2414 (MBG); St. Louis, Aug.-Sept. 1838, Riehl 9 (MBG); St. Louis, Sept. 1841,
Engelmann (G, US); Jefferson Ave., St. Louis, 2 July 1875, Eggert (MBG); Kimms-
wick, 15 July 1885, F. Wislizenus 284 (MBG); Williamsville, Wayne Co., 27 June
1914, E. J. Palmer 6106 (MBG); Poplar Bluff, 12 July 1930, Kellogg 16276 (MBG);
Sibley, 30 June 1906, Bush 4010 (G, MBG); Courtney, Bush 2109, 8537 (MBG);
bluffs of Pomme de Terre River, Polk Co., 29 July 1891, Trelease 715 (MBG);
vicinity of Gates, 26 Aug. 1912, Standley 9362 (US); Joplin, Jasper Co., 10 July
1897, Trelease 494 (MBG); Webb City, 1 July 1903, E. J. Palmer 454 (MBG);
Swan, 28 Sept. 1899, Bush 601 (MBG).
ANSAS: Monette, 14 June 1927, Demaree 3353 (MBG); Jonesboro, 15 June
1927, Demaree 8371 (G, MBG); Batesville, 1 Sept. 1897, T'release (MBG); Fayette-
ville, Harvey 62 (MBG); Fort Smith, 1853-4, Bigelow (US); Fulton, 22 Sept. 1900,
Bush 1034 (MBG).
RTH DAKOTA: Leeds, 10 Sept. 1896, Lunell (G, US); Fairmount, 22 July 1912,
Bergman 2370 (MBG); Mandan, 1915, Sarvis 122 (US); Dickinson, 14 Aug. 1908,
Holgate (G, NY).
SovrHu Dakora: Swan Creek, 20 July 1911, Visher 3859 (MBG); White River
Valley, 8 July 1911, Visher 2028 (NY); Deadwood, 23 July 1913, Carr 179 (G, MBG,
US); Lead City, 7 July 1892, Rydberg 934 (NY, US); Black Hills, near Fort Meade,
19 June 1887, Forwood sd (US).
NEBRASKA: "very common on sandy banks of Yellowstone River," 1853-4,
Hayden 10 (MBG); gt. James, 24 June 1893, Clements 2612 (G, US); Lincoln, 20
June 1888, Webber (US); near Dix, 28 Aug. 1926, Heller 14303 (MBG).
Kansas: prairie, Riley Co., 18 June 1895, Norton 392 (G, MBG, NY, US); Kearney
Co., 27 Aug. 1897, Hitchcock 1129 (MBG); bottom lands of Arkansas River, south
of Kendall, 15 June 1929, Rydberg & Imler 1008 (MBG); Ulysses, 27 June 1893,
Thompson 42 (MBG, NY, US); vicinity of Richfield, 20 Sept. 1912, Rose & Fitch
17106 (US).
KLAHOMA: Sapulpa, 22 July 1894, Bush 436 (MBG); near Page, 20 June 1914,
Blakeley 1462 (G); near Alva, 1913, Stevens 68014, 2889 (G); Woods Co., 21 June
1899, White 206 (MBG); vicinity of Fort Sill, 22 May 1916, Clemens 11750a (MBG);
near Snyder, 23 June 1913, Stevens 1192 (G, NY); near Camp, 11 May 1913, Stevens
891 (MBG).
Texas: Dallas, Reverchon 736, 2116 (MBG); Tarrant Co., Ruth 109 (G, NY);
Fort Worth, Ruth 109 (US); Polytechnic, Ruth 109 (MBG); Abilene, 22 May 1902,
Tracy 8001 (G, MBG, NY, US); west of Big Spring, Sept. 1881, Havard (US);
Amarillo, 2 Sept. 1910, Ball 1676 (US); banks of Red River, Randall Co., 13 Aug.
1900, Eggert (MBG); Canyon, 2 June 1918, E. J. Palmer 18861 (MBG); Lubbock,
10 June 1917, E. J. Palmer 12496a (MBG); El Paso, 7 June 1917, Clemens (P); below
1933]
PERRY— NORTH AMERICAN SPECIES OF VERBENA 307
Dofiana, Parry, Bigelow, Wright & Schott (US); valley of Rio Grande, 64-80 km.
below El Paso, ? 1851, Wright 567 (G), pu (G, MBG, NY); valley of Rio Grande,
St. Elizani, May-Oet. 1849, Wright 454 (G, US).
ALBERTA: Crow's Nest Pass, Aug. 1897, Macoun 24268 (N
Montana: about 20 km. above Glendive, 17 July 1883, bue (US); near Glendive,
19-21 Aug. 1884, J. Ball (G); Thompson's Falls, 27 Aug. 1892, Sandberg, MacDougal
& Heller 972 (G, P, US).
Wyomina: Yellowstone National Park, 1 Aug. 1902, Mearns 2698 (US); Mammoth
Hot Springs, 22 Oct. 1902, Mearns 5009 (NY, US); Casper, 6 July 1901, Goodding 207
(G, MBG, NY, P, US); Blue Grass Hills, 8 July 1894, A. Nelson 320 (G, MBG, NY,
US); Laramie Peak, 8 Aug. 1895, A. Nelson 1652 (P); Laramie, 24 July 1900, A.
Nelson 7671 (G, MBG, NY, P, US).
Coronapo: Fort Collins, 12 July 1884, Sheldon 38 (G); Denver, 10 Aug. 1910,
Eastwood 25 (G, MBG, US); Manitou, 12 Aug. 1901, Clements 9 (G, MBG, NY, US);
Cheyenne Canyon, Colorado Springs, 16 Aug. 1915, Drushel 4887 (MBG); vicinity
of La Junta, 26 Sept. 1913, Rose & Fitch 17503 (MBG, US); Deer Run, 25 Aug. 1901,
Baker 920 (G, MBG, NY, P, US); Cimarron, 29 July 1901, Baker 288 (G, MBG, NY,
P); Arboles, 18 June 1899, Baker 564 (F, G, MBG, P), Tyre collection of V. rudis.
EW Mexico: without locality, 1847, Fendler 587 (ANSP, G, MBG, US), 592
(MBG); Raton, 21-22 June 1911, Standley 6268 (US); Cimarron Canyon, 21-24
Aug. 1903, Griffiths 5560 (MBG); Farmington, 8 Aug. 1904, Wooton 2831 (US TYPE
of V. imbricata); Las Vegas, 1927, Arsène 18543, 18593, 18634, 18885, 18961, 18964
(all P); vicinity of Las Vegas, 1926, Arsène 18399 (ANSP), 18407 (MBG); near Pecos,
San Miguel Co., 20 Aug. 1908, Standley 5136 (MBG, NY, US); mouth of Indian
Creek, Pecos River National Forest, 25 July 1908, Standley 4549 (G, MBG, NY, US);
Upper Rio Tesuque, 24 July 1908, Standley 4458 (MBG, US); Santa Fe, 20 July
1926, Arsène & Benedict 15734 (MBG); vicinity of Bernalillo, 11 July 1926, Arsène &
Benedict 16790 (ANSP); Sandia Mountains, 19 Aug. 1926, Arsène & Benedict 16600
(MBG); Sandia Mountains, July-Aug. 1914, Ellis 221 (MBG, US); Capelin Canyon,
14 Aug. 1914, Ellis 221 (NY); near Albuquerque, 20 June 1926, E. J. Palmer 31117
MBG); Roswell, 24 Aug. 1900, Earle 351 (MBG, P, US); Queen, 2 Aug. 1909,
Wooton (MBG, US); near Fort Craig, 1 Aug. 1880, Rusby 336 (MBG, NY); Organ
Mountains, Dona Ana Co., 30 Aug. 1897, Wooton 409 (G, MBG, NY, US), TYPE
collection of V. confinis; Mesilla Valley, Dona Ana Co., 6 Aug. 1907, Wooton &
Standley 3330 (F, MBG); Kingston, 18 June 1904, Metcalfe 1008 (G, MBG, NY, P,
US); Berendo Creek, Sierra Co., 20 May 1904, Metcalfe 897 (MBG); Gila River
bottom, near Cliff, Grant Co., 13 June 1903, Metcalfe 137 (G, MBG, NY, P, US).
Arizona: Tuba Oasis, 15-31 July 1920, Clute 93 (G, MBG, NY, US); vicinity of
Flagstaff, 13 July 1898, MacDougal 286 (ANSP, G, NY, US); Flagstaff, 13 Aug.
1922, Hanson A.147 (MBG); near Prescott, 26 July 1927, Peebles, Harrison &
Kearney 4243 (US); 16 km. west of MeNary, 23 June 1930, Goodman & Hitchcock
1817 (MBG, NY).
Ipano: valley of Clearwater River, Nez Perces Co., 30 May 1892, Sandberg, Mac-
Dougal & Heller 264 (G, MBG, NY, P, US); Falk's Store, Canyon Co., 7 July 1910,
Macbride 200 (MBG, US); alkaline flats, sink of Big Lost River, 16 Aug. 1895,
Henderson 4070 (US); near St. Anthony, 4 July 1901, Merrill & Wilcox 776 (G, NY,
US); Boise, July 1892, Mulford (G, MBG); Boise, 1911, J. A. Clark 55 (G, MBG,
P, US), 279 (G, MBG, NY, P, US).
Uran: Oquirrh Mountains, Salt Lake City, 9-16 July 1902, Pammel & Blackwood
8589 (G, MBG); Salt Lake City, June 1869, Watson 823 (G, NY, US); Gunnison, 25
[Vor. 20
308 ANNALS OF THE MISSOURI BOTANICAL GARDEN
June 1875, Ward 256 (G, US); Greenriver, 22 June 1894, Jones 5478 (MBG, NY, P,
US); along Bullion Creek, above Marysvale, 21 July 1905, Rydberg & Carlton 7048
(G, ule US); Milford, 4 June 1902, Goodding 1022 (MBG).
NEvADA: Winnemucca, 1 Sept. 1897, Hillman (P); saline flats, Las Vegas, 6 May
1905, Goodding 2312 (G, MBG, NY); Colorado River bottoms, 24 km. east of Search-
light, 6 June 1915, Parish 10288 (G, MBG).
RITISH COLUMBIA: Kamloops, 13 June 1889, Macoun (NY, US).
ASHINGTON: without locality, 1889, Vasey 468 (G, NY, US); Cascade Mountains
to Fort Colville, about lat. 49°, 1860, Lyall (G); Oroville, 26 June 1911, Jones (P);
Meyers Falls, 21 Aug. 1902, Kreager 4?4 (G, NY, US); near Egbert Spring, Douglas
Co., 1 July 1893, Sandberg & Leiberg 340 (G, MBG, NY, US); Sentinel Bluffs, 15
July 1903, Cotton 1363 (G, US); Pullman, 21 July 1896, Elmer 324 (MBG, NY);
Wawawai, June 1896, Elmer (P); near river, Prosser, 10 July 1902, Cotton 621 (G,
MBG, US
OREGON: without locality, 1871, Hall 398 (G, MBG, NY); margin of shallow lake,
Eastern Oregon, 25 June 1898, Cusick 1967 (G, MBG); Snake River at mouth of
Cache Creek, Wallowa Co., 28 May 1897, Sheldon 8202 (MBG, NY); about 5 km.
above mouth of Clark’s Creek, 9 Sept. 1897, Sheldon 8856 (MBG, NY, US); Pilot
Rock, 2 Sept. 1896, Brown 58 (MBG, US); Pilot Rock, July 1902, Grifiths & Hunter
18 (NY, US); Butter Creek, Umatilla Co., 14 Sept. 1894, Leiberg 902 (G, MBG, NY,
US); about 16 km. west of Boardman, 17 July 1928, Thompson 4880 (G, MBG, US);
between Mosier and Rowena, 28-30 July 1922, = 9487 (MBG, P); The Dalles,
on the Columbia, 2 Aug. 1880, Englemann (MBG).
CaLrFORNIA: near Monterey, Bolander 426 (G); Jolon, 1880, Vasey 512 (US);
between Tulare and Tulare Lake, 25-30 Aug. 1892, Palmer 2699 (US); near Lone
Pine, Inyo Co., 14 June 1891, Coville & Funston 953 (US); near San Bernardino, 3
June 1891, Parish & Parish 2171 (NY, US); Soldiers Home, 20 June 1902, Abrams
2574 (G, MBG, NY, P); near ponds, Laguna, Munz 2207, 6349, 6598 (P); Laguna
Canyon, 26 July 1916, Crawford (MBG, P); Blue Lake, Imperial Valley, 1 April
1903, Abrams (MBG, NY).
Mexico:
Lower Cauirornia: Tia Juana, Orcutt (F, x 1228 (MBG); Ensenada, 31 Aug.
1886, Ming 1549 (MBG); Mancadero, 31 Aug. 1889, Orcutt 1546 (MBG).
Sonora: Alamo, 20 May 1925, Kennedy 7088 2 (US).
sunu tg 80 km. south of Juarez, 1911, Stearns 7 (F, US); Bolsom de Mapini,
13 April 1847, Gregg 483 (MBG
Pio ILA: near San Juan, valley of Nazas, 11 May 1847, Gregg 686 (MBG); —
of Nazas, near San Lorenzo, 11 May 1847, Gregg 623 (MBG); El Toro, near Movan
July 1910, godes 4524 in part (US); San Lorenzo de Laguna and vicinity, inre
west of Parras, 1-10 May 1880, Palmer 1048 (ANSP, G, US).
A readily recognized and widely distributed species varying
greatly in habit and pubescence as well as in length and position
of floral bracts. The typical form is a sprawling hirsute plant
with laxly ascending branches and spreading bracts. In New
Mexico it appears as a nearly erect herb (V. imbricata Woot. &
Standl.) with shorter and ascending or appressed bracts (var.
brevibracteata Gray). Some phases of this variation occur
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 309
throughout the northwest range of the species, but since these
tendencies to vary are apparently lacking in correlation, they do
not seem to be worthy of nomenclatorial attention.
32. V. carnea Medic. Bot. Beobacht. 1783: 131. 1784.
V. caroliniana Michx. Fl. Bor.-Am. 2: 14.
V. carolinensis Small, Fl. Southeast. U. S. ed. 1, 1009. 1903,
and ed. 2, 1913.
Phryma caroliniensis Walt. Fl. Car. 166. 1788.
Styleurodon caroliniamum Raf. Fl. Tellur. 2: 104. 1836.
Cinereous perennial; stems simple or sparingly branched above,
ascending, puberulent; leaves spathulate to oblong or the upper
oblong-hastate, sessile, shallowly serrate-dentate, scabrous above,
less harsh and spreading-pubescent along the prominently
reticulated veins of the lower surface; spikes terminal, peduncu-
late, chiefly solitary but occasionally in 3’s, slender with crowded
tips, more open in fruit; bracts lanceolate, Mou one-half shorter
than the calyx, glandular-pubetcetitl calyx about 5 mm. long,
glandular-pubescent, lobes acute, unequal; corolla-tube slightly
longer than the calyx, pubescent without; corolla-limb about 5
mm. broad, segments somewhat truncate; anthers glandless;
schizocarp 3 mm. long, not readily separating into 4 nutlets,
longitudinally sulcate and commonly scrobiculate on the upper
part.
Distribution: North Carolina to Florida, and west to Texas.
Specimens examined
NonTH Carouina: Pinehurst, 3 Sept. 1897, Katzenstein (G); open pine woods 3
km. south of James City, 11 July 1922, Randolph (G); Burgaw, Aug. 1878, Hyams
(US), June 1879, Hyams (MBG)
Soutu CAROLINA: Santee Canal, Ravenel (G); sandy open pine woods near Navy
Yard, Charleston, 27 April 1912, Robinson 26 (G); Summerville, June 1891, Taylor
(US); Bluffton, 1881, Mellichamp (US); sand bank near Orangeburg, 8 May 1907,
House 8276 (NY); Aiken, July 1870, Ravenel (NY, US); sandy ground north of
Graniteville, 22 May 1899, Eggert (MBG).
Greonara: sandy hills north of Augusta, 22 May 1899, Eggert (MBG); Augusta, 14
May 1900, Cuthbert pe: A E Alexander, Ellis (P); Flint River at Albany, 24-28
May 1895, Small (F,
FLORIDA: siht nS Buckley (G, MBG); without locality, Chapman (NY);
Lake City, 27 Feb. 1893, Rolfs 191 (MBG); pine barren, Duval Co., 29 April 1902,
Fredholm 5136 (G, MBG, US); Oakwood, Duval Co., 1 June 1893, Fredholm 110
(P); dry pine barrens, near Jacksonville, May, Curtiss 1959 (G, MBG, NY, US),
8 May 1884, Curtiss 4765 (G, NY, US), 29 April 1893, Curtiss 4386 (MBG, US);
[Vor. 20
310 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Hibernia, March 1869, Canby (G, NY, US); St. Augustine, 1877, Reynolds (NY
Gainesville, 11 May 1925, O'Neill 986 (US); dry sandy woods, Irvine, 29 April 1930,
Moldenke 1091 (MBG, NY); Ocala, Marion Co., 2 April 1879, J. D. Smith (US);
high pine land, vicinity of Eustis, 1-15 May 1894, Nash 601 (G, MBG, US); pine
lands at Lake Helen, 28 April 1906, Deam 1799 (US); Fort Meade, March 1880,
J. D. Smith (US); between Tallahassee and St. Marks, April 1843, Rugel (NY); sandy
open dep near Tallahassee, 9 April 1929, E. J. Palmer 35235 (G, MBG); Aspalaga,
Chapman (MBG); y T barrens, near Apalachicola, 11 Aug. 1889, Biltmore
Miler: peri (G, NY, U
LABAMA: without locality, 1859, aag (G); Auburn, Lee Co., 10 May 1896,
Earle & Underwood (NY); Auburn, 10 Feb. 1897, Earle & Baker (NY); Pinewoods,
Buckley (NY); woods, Spring Hill, 6 Aug. 1897, Bush 312 (MBG, NY, US); Spring
Hill, 25 June 1915, Drushel (MBG); open barrens, Spring Hill, June-July 1919,
Graves 621 (MBG, US); Mobile, May 1875, Curtiss (MBG); Mobile, June 1879,
Mohr (US).
Mississippi; sandy soil, southern Mississippi, 1859, Hilgard (MBG); Ocean
Springs, May 1892, Seymour & Earle 118 (G, MBG); Ocean Springs, 8 May 1895,
Skehan (MBG); Biloxi, 21 April 1898, Tracy 4981 (G, MBG, NY, US); Long Beach,
6 Aug. 1891, Joor (MBG); Harrison Co., 19 April 1927, Woodson & Anderson 1577
(MBG).
UISIANA: vicinity of Covington, 30 April 1920, Arséne 12117 (US), 8 May 1920,
Arsène 11820 (US).
Texas: sandy open ground, Fletcher, Hardin Co., 25 April 1916, E. J. Palmer
9559 (MBG).
One of the most distinct species of the genus in North America.
It is readily recognized by the long slender graceful spike, the
sessile elongate leaves, and the tardy splitting of the schizocarp.
The last character is probably Walter's reason for placing this
species in the genus Phryma.
SECTION 2. GLANDULARIA Schauer
2. GLANDULARIA Schauer in DC. Prodr. 11: 550. 1847.
Sterile style-lobe adjacent to stigmatic surface and protruding
well beyond it; ovary in later stages definitely but shallowly
lobed at the distal end, with the style apparently inserted in a
depression between the lobes; schizocarp constricted along the
lines of cleavage; connective of the upper anthers chiefly appen-
daged. Herbaceous perennials with prostrate, decumbent, as-
cending, and sometimes erect stems. Flowers usually doo: at
first fascicled or somewhat corymbose, later spicate. Calyx
usually more than twice as long as the schizocarp and constricted
or contorted beyond it. Species 33-51.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA dll
KEY TO THE SPECIES
A. Leaves tapering into a margined petiole or subsessile or sessile, at least not
subauriculate and semiamplexicaul at the base.
B. Nutlets with a very definite beak parallel to the axis of the schizocarp..
bee CEM noa Nen coe ce ie bios ve nS wo See 33. V. quadrangulata
B. Nutlets with an indefinite beak or none.
C. Nutlets suggesting a tendency toward a beak (a slight protrusion
horizontal to the axis of the schizocarp); commissural face practi-
cally reaching the tip of the nutlet................... 34. V. delticola
C. Nutlets without a beak; commissural face not reaching the tip of the
nutle
D. Nutlets subovoid, lateral surfaces ventricose and smooth, not at all
similar to the dorsal surface. ..................... 39. V. tumidula
D. Nutlets Oe a E surfaces not ventricose and usually
scrobiculate, similar e dorsal surfac
E. Leaves shallowly Gee mi or oe
F. Corolla-tube slightly longer than the calyx.
G. Calyx villous-hirsute, somewhat glandular................
DUM NN Cr CAPE 47a. V. Gooddingii var. bise
G. Calyx short-strigillose, not glandular........... 38. V. tampensis
F. Corolla-tube at least one-half longer than the calyx.
H. Plants prostrate-decumbent; leaves varying from cuneate to
orbieular-ovate; calyx-teeth short (1-1.5 mm. long)......
37.
H: ies ascending, decumbent only at base; leaves ovate to
gate-ovate; calyx-teeth long (2-3 mm.)....35. V. canadensis
E. Leaves 3-dleft, inclsod-plumdd or bipinnatifid.
I. Corolla-tube protruding well beyond the calyx.
J. Corolla-tube approximately twice as long as the calyx.
K. Plants somewhat hirsute; both surfaces of the leaves simi-
lar in color; spikes elongating in age; calyx-teeth 2-3
mm. long. iusso EBEN MER d 0 S4 ex ns 85. V. canadensis
K. Plants premio < lower surface of leaves lighter
than the upper; spikes compact at maturity; calyx-
teeth ie surpassing 2 mm. in len
L. Plants prostrate-decumbent; spikes few-flowered..36. V. elegans
L. edo Fine g decumbent only at base; spikes
y-Bowered. ii. cenas en 36a. V. elegans var. asperata
J. Cardia bibi kope to one-half longer than the cal
M. Floral bracts equalling or exceeding the calyx; esiyx not
glandular, hispid-hirsute
N. Leaves bipinnatifid, ultimate segments linear-oblong..
vvv ENERO eor 40. V. bipinnatifida
N. Leaves 3-cleft, with segments remotely incised or AE
ultimate segments much broader than in the
DTE ot 40a. V. Watana var. boil
M. Floral braets shorter than the calyx; calyx for the most
part somewhat glandular, hispid-hirsute to villous-
pubescent.
[Vor. 20
312 ANNALS OF THE MISSOURI BOTANICAL GARDEN
O. Leaves sessile, strigose-hispid............. 46. V. Andrieuxit
O. Leaves subsessile to short-petiolate, hispidulous-hirsute
to hirtellous
P. Plants coarse; leaves usually 2.5-4 em. long, hispid-
ulous-hirsute; corolla-liimb 6-12 mm. broad;
nutlets 2.5-3 mm. long
Q. Calyx-teeth long (2-3 mm.).
R. Plants usually tall (2-4 dm.); leaves bipinnatifid
ith ultimate segments lanceolate; calyx-teeth
not especially conspicuous in mature fruit.
S. Calyx glandular................ 41. V. ambrosifolia
Calyx not glandular.41a. V. ambrosifolia f. eglandulosa
R. — usually low (1-2 dm.); leaves trifid with
egments more or less incised; calyx-teeth
mesas long in mature fruit...........
UXWEREEREISATUKMERS 49a. V. ciliata var. longidentata
Q. Calyx-teeth short (usually less than 2 mm.).
T. Plants decumbent-ascending or prostrate
densely hirsute; calyx somewhat glandular,
hispidulous-hirsute, scarcely viscid
U. Plants with decumbent-ascending loose habit;
leaf-margin merid revolute........ 48. V. ciliata
U. Plants with prostrate compact habit; le
margin strongly sinus te. .43b. V. ciliata var. pubera
T. Plants ascending-erect, more or less hirsut
T aene plandular-hirsute and oi
cid-pubescent.................0. 42. V. Wrightà
P. Plants Neid esed smaller, 1-2.5 cm. long,
"page sor tu 5-6 mm. broad; nutlets
Aerated ads TAA EIE OER 44. V. racemosa
I. Corolla-tube slightly ue than the c
V. Calyx-teeth acute-subulate, short dg than half as long as
the calyx-tube).
W. Plants repent, slender; spikes few-flowered, scarcely pro-
truding beyond the subtending leaves.
X. Corolla-limb 5-7 mm. broad............ 46. V. teucriifolia
X. Corolla-limb 3 mm. broad: 46a V. teucriifolia var. corollulata
W. Plants decumbent-ascending, stouter; spikes many-flow-
ered, subsessile or short-pedunculate.
Y. Plants more or less hirsute, decumbent; corolla incon
spicuous, limb 3-5 mm. broad............ 48. V. pue
Y. T ke ap ig ascending; corolla con-
, limb 8-9 mm. broad........ 47. V. Goodding&
V. Clysleih m a elongate (approximately half
s long as the calyx-tube
Z. Plante decumbent-ascending, Kpaka leaves
ovate, trifid with the segments coarsely dentate. 49. V. setacea
Z. Plants erect, sparsely hirsute; leaves narrow ae oe
_Mipianatisd, with the segments linear.......... 50. V. lilacina
A. I I i at the ay er ees 51. V. amoena
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 313
3. V. quadrangulata Heller, Contr. Herb. Franklin &
Heat College (Bot. Expl. S. Texas) 1: 84, pl. 6. 1895.
V. pumila f. albiflora Standl. Field Mus. Publ. Bot. 4: 256.
1929
H ain PF. deg eae Small, Fl. Southeast. U. S. ed.
1, 1011. 1908, and ed. 2, 1
Low plant more or PH m branched from the base;
stems prostrate-ascending, rooting at the lower nodes, hirsute;
leaves 1-3 cm. long, broadly ovate, with cuneate base contracted
into a very short margined petiole, incised-pinnatifid or 3-cleft,
with lobes incised, strigose-hirsute on both surfaces; spikes
compact, sessile or nearly so, terminal; bracts somewhat shorter
than the calyx, narrowly lanceolate, hirsute, ciliate; fruiting
calyx 5(-6) mm. long, hirsute, particularly along the ribs, lobes
short, subulate-tipped; corolla-tube slightly longer than the
calyx, practically glabrous without; corolla-limb 2.5-3 mm.
broad, lobes emarginate; anthers unappendaged; ovary sur-
mounted by subhemispheric-angulate stylopodium; schizocarp
constricted along the lines of cleavage; nutlets 4 mm. long,
crowned with a smooth obtusish beak, shallowly reticulate-
scrobiculate above, longitudinally striate and somewhat abruptly
enlarged at base.
Distribution: Texas and northern Mexico.
Specimens examined:
Texas: sandy beach and open flats, Rio Bravo del Norte, Schott (NY); Devils
River, Val Verde Co., 15 May 1913, Orcutt (MBG); creek between Del Rio and
Comstock, 22 April 1925, Small & Wherry 12010 (NY); “Brackett” (? Brackettville),
22 Aug. 1900, Trelease 101 (MBG); Spofford Junction, 22 March 1900, Canby 193
(US); Uvalde, 28 April 1928, E. J. Palmer 33592 (G, MBG, NY); San Antonio,
April 1922, Schulz 767 (US); 24 km. south of San Antonio, 28 April 1921, Schulz
475 (US); woods near Colorado River, near Wharton, 12 April 1925, Small & Wherry
11824 (NY); railroad north of Moore Station, Frio Co., 5 April 1901, Eggert (MBG);
sandy open ground, Pleasanton, 16 May 1916, E. J. Palmer 1747 (MBG); El Jardin,
10 March 1924, Runyon 628 (US); Eagle Pass, April 1883, Havard (US); Kennedy-
Beeville, 15 March 1929, Tharp 5538 (US); Kennedy-Portland, 17 March 1929,
Tharp 5610 (US); Corpus Christi Bay, Dec. 1879, Palmer 1046 (G); Corpus Christi,
10 March 1894, Heller 1888 (G, MBG, NY), Type collection; San Diego, 1885-6,
Croft 78 (NY); Kingsville, 27 March 1920, High 75 (MBG); between Laredo and
Bejar, Feb. 1828, Berlandier 1485 =225 (G); Laredo, Feb. 1891, Dodge 154 (US);
sandy ground, Laredo, 6 April 1901, Eggert (G, MBG); Laredo, 21 March 1903,
Reverchon 3902 (G, MBG, NY, P, US); Laredo, 1913, Orcutt 5542, 5780 (MBG);
San Antonio, 1926, Pagel 2208 (F); Guadalupe, 168 km. southwest of San Antonio,
[Vor. 20
314 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Sept. 1879, Palmer 2039 (G); chaparral near cun ge 6 April 1925, Small &
Wherry 11903 (NY); Brazos Santiago, 1889, Nealley (F, U
Mexico: TaMAULIPAS: east of Matamoros, May on Berhondior 3018 —1518
(MBG).
Verbena quadrangulata is an anomalous species often confused
with V. pumila on account of the strong similarity in habit, but
is perhaps more closely related to V. delticola, the only other
known North American species with a tendency toward develop-
ing a nutlet with a beak. The style is enlarged at the base into a
persistent subhemispherical body; hence, when the nutlets split
apart they appear as if elongated at the apex into a beak-like
appendage. Although the anthers are not glandular and the
flowers not showy, the species seems to belong to the section
Glandularia.
34. V. delticola Small, n. sp.
Stems decumbent to ascending, branched, more or less hirsute;
leaves ovate-deltoid, 3-7 cm. long, with truncate-cuneate base
narrowed into a margined petiole, obtusish or acutish, coarsely
serrate-dentate, often trilobed, usually thin, sparsely appressed-
hirsute on both surfaces, trichomes above often with minute
bulbous bases; spikes peduncled, fascicle-like in anthesis, be-
coming elongated in fruit; bracts linear-attenuate, shorter than
or equalling the calyx; calyx 7-8 mm. long, sparsely glandular,
hirsute, lobes slender, subulate, unequal; corolla-tube protruding
well beyond the calyx, pubescent or glabrate without; corolla-
limb probably 10 mm. broad, segments emarginate; nutlets
3 mm. long, subcylindrical, reticulate from the apex to the defi-
nitely broadened base; commissural face practically reaching
the tip of the nutlet, muricately scabrous.
2 V, delticola Small, spec. nov., herbacea verisimiliter perennis; caulibus decum-
bentibus vel ascendentibus ramosis plus minusve hirsutis; foliis ovato-deltoideis basi
cuneata in petiolum alatum attenuatis 3-7 cm. longis obtusiusculis vel acutis grosse
serrato-dentatis saepe trilobis utrinque sparse adpresso-hirsutis; spicis pedunculatis;
bracteis lineari-attenuatis calyce brevioribus vel subaequant tibus; calyce 7-8 mm.
longo sparse glanduloso hirsuto; calycis dentibus subulatis inaequalibus; corollae
tubo exserto extus pubescente vel glabrato; corollae limbo circiter 10 mm. lato
segmentis emarginatis; coccis subcylindricis 3 mm. longis superne reticulatis.—
Collected at Las Palmas Ranch, vicinity of Brownsville, Texas, 1-5 Aug. 1921
Ferris & Duncan 3161 (NY), TYPE.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 315
Distribution: Texas and Mexico.
Specimens examined:
Texas: Edinburg, Hooker 5999 (US); Samfordyce, 1927, me 7224 (US);
chaparral near Harlingen, 16 April 1925, Small & Wherry 11901 (NY); Brazos
Santiago, 1889, Nealley 117, 118 (US); Reynoldsville, Cameron Co., 11 April 1905,
Lewton 141 (US); Las Palmas Ranch, vicinity of Brownsville, 1-5 aay 1921, Ferris
& conic 3161 (MBG, NY Type).
MeExic
NO Leon: Rinconada, 24 April 1847, Gregg 752 (MBG); Guajuco, March
1880, Palmer 1051 (G, US); Walnut Grove, near Monterey, 7 Feb. 1847, Gregg 202
(MBG); river gravel, near Monterey, 11 July 1888, Pringle 2228 (G); Sierra Madre,
near Monterey, 25 Aug. 1903, Pringle 11843 (G, US); Loma del Obispodo, Monterey,
1 Feb. 1907, Safford 1221 (US); Monterey, Guadalupe, May 1911, Arsène 6129 (G,
MBG, NY, US); Monterey, 10-11 March 1923, Tharp 1826 (US).
TAMAULIPAS: Victoria, 23 May 1898, Nelson 4424 (G, US); vicinity of Victoria,
1 Feb.-9 April 1907, Palmer 39 (F, G, MBG, NY, US); San Fernando to Jimeney,
26-27 Feb. 1902, Nelson 6628 (? 6028) (G, MBG, NY, US); vicinity of Tampico,
1-31 Jan. 1910, Palmer 90 (F, G, MBG, NY, US
San Luis Potost: La Hoya, Liebmann 11314 (US).
PoEBLA: near Metlaltoyuca, 31 Jan. 1918, Goldman 42 (US).
V Cruz: Wartenburg, near Tantoyuca, April 1858, Ervendberg 236 (G);
Colipa, March 1841, Liebmann 11313 (US).
Superficially, this species bears a strong resemblance to V.
canadensis, but usually the leaves are not so deeply incised. Its
distinctive character is found in the nutlet. Commonly, in the
section Glandularia, the schizocarp is shallowly lobed at the apex;
hence, the style appears to be attached in a very definite de-
pression and ordinarily the commissural face does not reach the
tip of the nutlet. In V. delticola, however, the depression is
indefinite, the commissural face practically reaches the tip, and
the separate nutlets viewed from the lateral or ventral surface
suggest a tendency toward developing a beak. In Safford 1221
a small beak is present.
ih V. canadensis (L.) Britton, Mem. Torr. Bot. Club 5: 276.
Aiia Jacq. Hort. Vind. 2: 82, pl. 176. 1772; Linn. f.
Suppl. 86. 1781; Bot. Mag. pl. 308. 1795; Bot. Reg. pl. 294.
V: O TETA Retz. Svenska Vet. Akad. Nya Stockh. Handl. 34:
143, pl. 5. 1778.
V. Obletia Medic. Act. Acad. Theod.-Palat. 3: 194, pl. 7. 1775.
V. longiflora Lam. Tab. Encye. 1: 57. 9
[Vor. 20
316 ANNALS OF THE MISSOURI BOTANICAL GARDEN
V. rubra Salisb., Prodr. 71. 1796.
V. Lamberti Sims in Bot. Mag. pl. 2200. 1821.
V. Aubletia var. Drummondi Lindl. Bot. Reg. pl. 1925. 1837.
V. Lamberti var. rosea Sweet, Brit. Fl. Gard. II. 4: 363. 1838.
V. Drummondii Hort. ex G. Don in Loud. Hort. Brit. Suppl.
2:080. 1839.
V. Drummond (Lindl. Baxt. ex Small, Fl. Southeast. U. S.
ed. 1, 1011. 1903, and ed. 2, 1913.
V. canadensis var. Lamberti 'Thell. Fl. Advent. de Montpellier,
428. 1912.
Buchnera canadensis L. Mant. 88. 1767.
Billardiera explanata Moench, Method. 369. 1794.
Anonymos caroliniensis Walt. Fl. Carol. 164. 1788.
Glandularia caroliniensis J. F. Gmel. Syst. Veg. 22: 920. 1791.
G. Aubletia Nutt. Trans. Am. Phil. Soc. 5: 184. 1837.
Stems decumbent to ascending, rooting at the lower nodes,
more or less branched, spreading-hirsute or glabrate; leaves
ovate to elongate-ovate, 3-9 em. long, 1.5-4 em. broad, with
truncate or cuneate base narrowed into a margined petiole, in-
cised or incised-pinnatifid or 3-cleft, appressed-hirsute or glabrate
on both surfaces; spikes pedunculate, fascicle-like in anthesis,
becoming elongated in fruit; bracts shorter than (or occasionally
as long as) the calyx, linear-attenuate, hirsute, usually ciliate;
calyx glandular-hirsute, in fruit 10-13 mm. long, lobes very
slender, subulate-setaceous, unequal, the posterior lobe much
shorter; corolla-tube about twice as long as the calyx, glabrous
without or finely pubescent or glandular; corolla-limb 11-15 mm.
broad, segments emarginate; anther-glands not minute; schizo-
carp at maturity constricted along the lines of cleavage; nutlets
3(-3.5) mm. long, subcylindrical with slightly broadened base,
reticulate-scrobiculate; commissural face muricate-scabrous.
Distribution: North Carolina to Florida, and west to Colorado and Texas.
ed:
NonTH CanoriNA: roadsides, Kittrell, 11 April 1889, Sturtevant (MBG).
SourTH Carona: Abbeville District, Hexamer & Maier (G).
GEoRGIA: by roads and margins of fields through lower Georgia and middle
Florida, March 1843, Rugel (MBG).
Fiorma: Fort King, Alden (NY); St. Nicholas, Duval Co., July 1898, Lighthipe
599 (NY); Mous Creek, St. John’s Co., 15 April 1879, J. D. Smith (US); rich woods
in Daytona, 30 March 1906, Deam 1831 (US); Ocala, 4 April 1879, J. D. Smith (US).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 817
ALABAMA: above Tuscaloosa, 8 May 1875, E. A. Smith (US); above Tuscaloosa,
11 April 1892, Ward (US).
LovuisiANA: without locality, Hale (G, US), Short (NY); Minden, 15 April 1901,
Trelease (MBG); Winnfield, 13 April 1912, Petersen (NY); Natchitoches, 16 March
, E. J. Palmer 7001 (MBG); Chopin, Natchitoches Parish, 23 March 1915,
E. J. Palmer 7071 (MBG); vicinity of Covington, 20 March 1920, Arsène 11982 (US);
Jacksonville, Drummond (G); Cameron, 4 July 1903, Tracy 8707 (G, MBG, NY,
US).
Kentucky: Shelbyville, Flint (G).
TENNESSEE: Cedar Glades, north of Lavergne, 4 May 1898, Eggert (MBG).
Iowa: Council Bluffs, Vasey (G).
Missourt: St. Louis, May 1833, Engelmann 334 (MBG); St. Louis, 8 July 1910,
Sherff 333 (G); Windsor Springs, 30 May 1890, Hitchcock (MBG); St. Louis Co., 16
May 1879, Eggert (MBG, US); Allenton, 27 April 1887, Eggert (MBG); west of
Kimmswick, 13 April 1918, Drushel 3659 (MBG); Kimmswick, 30 April 1905, Johnson
(MBG); Hillsboro, 24 May 1885, F. Wislizenus 285 (MBG); Crystal City, 19 Aug.
1886, Eggert (MBG); sandy ground north of Crystal City, 6 May 1891, Eggert (MBG);
Siliea, 18 April 1896, Eggert (MBG); Shot Hill, near Selma, 30 May 1923, Greenman
4231 (MBG); Victoria, 10 May 1890, Hitchcock (MBG); Old Mines, Washington Co.,
19 Aug. 1928, Kellogg 1958 (MBG); Blackwell, St. Francois Co., 18 April 1897, T'release
714 (MBG); Central, 28 Aug. 1898, T'release 1161 (MBG); banks of Pilot Knob Creek,
9 Sept. 1859, Engelmann (MBG); Pilot Knob, July 1867, Engelmann (MBG); Granite
Mountain, Iron Co., 24 May 1918, Greenman 4076 (MBG); Ironton, 4 May 1923,
Epling 6134 (MBG); Shepard Mountain, Iron Co., 26 May 1918, Greenman 3870
(MBG); Shut In, Stout Creek, Iron Co., 28 May 1916, Drushel 2762 (MBG); near
Silvermine, Madison Co., 20 May 1927, Greenman (MBG); Mine La Motte, 19 May
1927, Greenman (MBG); Grandin, 5 May 1901, Bush 318 (MBG); dry rocky banks,
Shannon Co., 13 April 1889, Bush 1168 (MBG); Jerome, 5 April 1914, Kellogg (MBG);
Jackson Co., 28 March 1864, Broadhead (MBG); Jackson Co., 28 May 1893, Bush
283 (G, MBG); Independence, 1 June 1895, Tindall (MBG); Dodson, 27 April 1904,
Bush 1933 (MBG); rocky woods, Vale, 13 April 1908, Bush 4922 (MBG, US); barrens,
Greenwood, 28 Oct. 1915, Bush 2901 (G, MBG, NY); Greenwood, 25 April 1911,
Bush 6439 (MBG); Jasper Co., 16 Aug. 1893, Bush (MBG); Oronogo, 10 July 1910,
E. J. Palmer 2996 (MBG); open banks, Webb City, 19 April 1903, E. J. Palmer 559
(MBG); Webb City, 4 May 1902, E. J. Palmer 304 (MBG); prairie, vicinity of Pearl,
22 Aug. 1912, Standley 9171 (US); Willard, 20 July 1919, Blankinship (P); Springfield,
31 July 1892, Dewart 74 (MBG); vicinity of Springfield, 29 Aug. 1911, Standley 8367
(US); barrens, Swan, 25 Sept. 1899, Bush 569 (MBG); Swan, 7-9 Oct. 1915, Eggles-
ton 12240 (NY, US); Galena, Stone Co., 27 May 1914, E. J. Palmer 5682 (MBG);
limestone slopes, ‘‘ Bald Joe," Stone Co., 30 April 1924, E. J. Palmer 24616 (MBG);
near Seligman, Barry Co., 4 April 1926, E. J. Palmer 29804 (MBG).
ARKANSAS: Eureka Springs, 19 April 1899, T'release (MBG); top of dolomite hill
along White River, near Beaver, Carroll Co., 23 Oct. 1925, E. J. Palmer 29333 (G,
MBG, NY); bluffs, Van Buren, 5 April 1929, Demaree 6411 (US); 14 km. west of
Fort Smith, 1853-4, Bigelow (US); Fort Smith, 1 April 1928, Benke 4566 (G); Hot
Springs, 31 Oct. 1899, T'release (MBG); Malvern, Letterman (MBG, US); Prescott,
8 April 1900, Bush 545 (MBG); clay barrens, Fulton, Hompetibd Co., 28 April
1914, E. J. Palmer 5407 (MBG); Red River, Pitcher
Kansas: Princeton, 19 May 1919, Street (P); between Olathe and Pleasanton,
Miami Co., 18 June 1929, Rydberg & Imler 22 (MBG, NY); open woods, Cherokee
[Vor. 20
318 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Co., 1896, Hitchcock 790 (G, MBG, NY, US); vicinity of Edna, 28 June 1929, Ryd-
berg & Imler 379 (NY).
OKLAHOMA: near Miami, Ottawa Co., 26 Aug. 1913, Stevens 2840 (G, MBG, NY,
US); Sapulpa, 22 July 1894, Bush 437 (G, MBG, NY, US); Perkins, 14 June 1893,
Waugh 168 (MBG); onland woods, Muskogee, 30 Sone 1918, E. J. Palmer 14283
MBG); Norman, 17 April 1915, Emig 415 (MBG, US); near Page, 20 May 1914,
Blakeley 1471 (G); near Paul's Valley, Garvin Co., 19 April 1913, Stevens 109 (G,
MBG); Colbert Station, 19 June 1891, Sheldon 26 (US); near Idabel, 18 May 1916,
Houghton 3643 (G, MBG).
Texas: Denison, 22 April 1904, Reverchon (MBG); Denison, 15 March 1904,
Reverchon (MBG); wet places, T. & C. Junction, Bowie Co., 5 Sept. 1900, Eggert
G, MBG); sandy ground north of Longview, Gregg Co., 19 April 1899, Eggert
(MBG); open ground, Longview, 21 April 1915, E. J. Palmer 7124 (MBG); sands,
Dallas, Reverchon (G), 740 (MBG); Newland, near Dallas, 26 March 1901, Rever-
chon 2533 (MBG, NY); Grapeland, Houston Co., 28 May 1917, E. J. Palmer 12056
(MBG); Grapeland, 8 June 1920, Tharp 838 (G, NY, US); Livingston, Polk Co., 10
April 1914, E. J. Palmer 5185 (MBG); Sabine River, 25 km. north of Orange, 18
April 1899, Bray 65 (US); Dayton, Liberty Co., 21 May 1917, E. J. Palmer 11979
(MBG); sandy open ground, near Conroe, Montgomery Co., 15 April 1929, E. J.
Palmer 83319 (G, MBG, NY); Houston, Feb. 1842, Lindheimer (G); woods, Houston,
6 April 1872, Hall 436 (MBG, abi P, US); Houston, Fisher (MBG), 33, 40, 47, 707
(US); Galveston, Van Huff (M
CoronaApo: Rocky neri pe 40-41’, 1868, Vasey (G, MBG).
Verbena canadensis is one of the most easily recognized of the
North American species belonging to the section Glandularia.
It has a large corolla with tube about twice as long as the calyx
and, at maturity, an elongated spike with somewhat remote
truita and conspicuous calyx-teeth. The leaves are variously
lobed or cleft. Although these variations have been used at
sundry times to segregate species, they do not seem to be constant
nor are they supported by definite geographic ranges, hence
scarcely merit recognition.
36. V. elegans HBK. Nov. Gen. et Sp. 2: 273. 1818.
V. moranensis Willd. ex Spreng. Syst. 2:750. 1825.
V. canadensis subsp. elegans Thell. Fl. Advent. de Montpellier,
428. 1912
V. canadensis var. Ehrenbergii Thell. l. c.
Stems prostrate-decumbent, tending to root at the older nodes,
more or less branched, sparsely hispidulous-hirsute; leaves
lanceolate-ovate with cuneate base narrowed into a margined
petiole, 2-5 cm. long, incised-pinnatifid or trifid with the divisions
less deeply cleft, somewhat appressed-hirsute on both surfaces,
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 319
dark green above, lighter beneath, often with branches or fascicles
of smaller leaves in the axils; spikes few-flowered, pedunculate,
terminal, fascicle-like; bracts mostly one-half to two-thirds as
long as the calyx, lanceolate, attenuate, glandular-hirsute,
ciliate; calyx 8-10 mm. long, glandular-hirsute, teeth short,
subulate and unequal; corolla-tube 15-18 mm. long, finely but
sparsely pubescent without, throat gibbous; corolla-limb 10-12
mm. wide; nutlets subcylindrical, 3.5-4 mm. long, reticulate-
scrobieulate chiefly on the upper half, lower part striate; com-
missural face muriculate.
Distribution: vocum Mexico.
Specimens examined
Mexico:
Hipauco: Sierra de Pachuca, 2700 m. alt., 22 July 1898, Pringle 6908 (ANSP, F,
G, MBG, NY, P, US); Sierra de Pachuca, 21 and 22 July 1901, Rose & Hay 6556
(US); mountains, Pachuca, July 1905, Purpus 1433 (MBG, P); between Pachuca
and Real del Monte, 19 July 1905, Rose, Painter & Rose 8668 (NY, US); Real del
Monte, 9 May 1910, Clokey (MBG); Moran, Humboldt & Bonpland 4063 (Bot. Mus.
Berl.-Dahl. type of V. moranensis, MBG phot.); under oaks and firs, Sierra de
Ajusco, ca. 2800 m. alt., 26 Aug. 1902, Pringle 11092 (F, G, MBG, NY, US)
Mexico: lomas de Santa Fe, Aug. 1927, Lyonnet 178 (US).
Vera Cruz: near Chila, Distr. Ozuluama, April 1888, Seler 723 (G).
Oaxaca: Tehuantepec, 1906, Gandoger (MBG).
The above species is related to V. canadensis but is easily
separable by its slender more or less prostrate habit, few-flowered
compact spikes, and shorter calyx-teeth. The nutlets of the two
are about the same size, but the reticulations in V. elegans are a
little coarser than those in the more northern species.
36a. Var. asperata Perry,” n. var.
Stems decumbent-ascending, hirsute-hispid; leaves variously
cleft or lobed; spikes dense, many-flowered; calyx glandular,
hispidulous-hirsute, teeth a little longer than in the species.
Distribution: Sonora ane Chihuahua to San Luis Potosi.
DECORUS examine
XICO
Sonora: ice Madre, 15 Dec. 1890, Lumholtz 445 (G); Badehuachi, 2 Dec. 1890,
Ya 446 (G); Hermosillo, 1888, M. A. Crawford (G).
23 Var. asperata Perry, var. nov., caules ascendentes hirsuto-hispidi; foliis bi-
pinnatifidis vel multifidis; spicis densis multifloris; calyce glanduloso hispidulo-
hirsuto.—Colleeted at San Antonio, Coahuila, Mexico, 31 Aug. 1848, Gregg 356
(MBG), TYPE.
[Vor. 20
320 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Curnuanva: southwestern Chihuahua, Aug.-Nov. 1885, Palmer 295 (G, US).
DunaANco: San Ramon, 21 Apr.-18 May 1906, Palmer 191 (G, MBG, NY, US).
Coanuita: Sierra de Parras, Oct. 1910, Purpus 4974 (F, G, MBG, US); 72 km.
east of Saltillo, July 1880, Palmer 1052 (G, US); San Antonio, 31 Aug. 1848, Gregg
3856 (MBG type); Sierra Encarnaciore, 28 July 1896, Nelson 3896 (US).
San Luis Porosi: near Morales, 1876, Schaffner 716 (G); near San Luis Potosi,
1878, Parry & Palmer 720 (ANSP, F, MBG, US); Alvarez, 5-12 Sept. 1902, Palmer
51 (F, G, MBG, NY, US).
This variety, as compared with the species, is stouter and more
erect with coarser pubescence and larger spike. Palmer
differs from the other collections in having very shallowly lobed
ovate leaves. Parry & Palmer ?20 shows much variation in
pubescence, some plants being scabrous, others scarcely so at all.
Purpus 4974 is lacking the coarse pubescence but appears more
closely allied here than elsewhere.
37. V. maritima Small, Bull. N. Y. Bot. Gard. 3: 436. 1905.
Stems branched at base, decumbent or prostrate, sparingly
pubescent or glabrate; leaves cuneate to orbicular-ovate, 1-4(-6)
em. long, tapering into a margined petiole, incised-dentate or
somewhat lobed, sparsely pubescent or glabrate on both surfaces;
spikes terminal, pedunculate, fascicle-like in anthesis, becoming
elongate in fruit; bracts linear-lanceolate, about one-half as long
as the calyx, acuminate, pubescent, ciliate; fruiting calyx 10-13
mm. long, appressed-pubescent, often glandular, teeth short,
slender, subulate; corolla-tube at least one-half longer than the
calyx, pubescent without; corolla-limb 10-15 mm. broad; anthers
with or without glands; nutlets subcylindric with broadened
base, 4 mm. long, scrobiculate; commissure narrow, muricately
scabrous.
Distribution: Florida.
Specimens examined:
FLoripa: sandy ridges bordering the ocean, eastern Florida, Curtiss 1963 in part
(G, MBG, NY); sandy soil, Merritt’s Island, 9 Dec. 1929, Moldenke 219a (NY);
near Cape Canaveral, 15 July 1896, Curtiss 5706 (G, MBG, NY, P, US); Cape
Canaveral, 2-5 April 1904, Burgess 638 (NY); Fort Pierce, 8-9 April 1904, Burgess
713 (NY); sand dunes, Hobe Sound, 19 March 1921, Randolph 52 (G); West Jupiter,
10 April 1904, Burgess 783 (NY); near beach, Palm Beach, 19 Nov. 1914, Small 2124
(NY); dry hammock south of Palm Beach, 1 May 1918, Small 8509 (NY); beach
opposite Miami, Nov. 1904, Small 2100 (NY); opposite Miami, Feb. 1911, Small,
Carter & Small 3311 (NY); Cape Florida, 29 March 1904, Britton 296 (NY) ; pine
lands, Small Island, northwest of Perrine, 16 Jan. 1909, Small & Carter 2994 (NY);
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 321
sandy soil, Redlands, 2 Feb. 1930, Moldenke 549 (MBG, NY); sandy soil, Golden
Beach, 10 Feb. 1930, Moldenke 586 (MBG, NY); Camp Jackson, 25 March 1904,
Britton 220 (F, NY); Everglades, west of Camp Jackson, Dade Co., 6-9 May 1904,
Small & Wilson 1961 (NY); Royal Palm Hammock, 20 Feb. 1915, Small & Small
5422 (NY); Paradise Key and vicinity, 21-29 Sept. 1917, Safford & Mosier 210 (US);
Hammer Key, Everglades, 12 May 1918, Small 8594, 8599 (NY); in sand near
Cocoa Beach, 16 March 1930, O'Neill 6309 (MBG, US); between Cutler and Long-
view Camp, 9-12 Nov. 1903, Small & Carter 1077 (NY).
This native of the sand dunes and the hammocks of Florida
resembles V. canadensis in inflorescence, but is readily distin-
guished by the creeping habit and the cuneate or orbicular-ovate
leaves.
38. V. tampensis Nash, Bull. Torr. Bot. Club 23: 104. 1896.
Stems erect from a decumbent or creeping base, pubescent or
glabrate; leaves ovate to oblong-ovate, 2-7 cm. long, 1.5-4 cm.
broad, truncate at base, tapering into a margined petiole (2 or
more cm. long), coarsely serrate-dentate or shallowly incised,
strigillose or glabrate on both surfaces, paler beneath; spikes
pedunculate, fascicle-like in anthesis, becoming elongated in
fruit; bracts linear-subulate, about one-half as long as the calyx,
strigillose, ciliate; fruiting calyx 12-13 mm. long, strigillose,
lobes very slender, subulate, unequal; corolla-tube somewhat
longer (about 2-4 mm.) than the calyx, pubescent without;
corolla-limb 10-12 mm. broad; anthers glandless or with a minute
gland on the connective of each of the upper pair; schizocarp at
maturity slightly constricted along the lines of cleavage; nutlets
subcylindrical with broadened base, 4 mm. long, reticulate-
scrobiculate above, striate toward base; commissure minutely
scabrous.
Distribution: Florida.
Specimens examined:
FLoriwa: without locality, 1842-49, Rugel 305 (F, MBG, NY, US); Daytona, 24
Jan. 1907, Mell (MBG); hammock south of Daytona, 8 May 1918, Small 8686 (NY);
pine lands east of Eustis, 7 May 1918, Small 8469 (NY); Titusville, 31 March 1914,
Mattern (US); dry sandy soil along roadside, 3 km. east of Fort Christmas, Orange
Co., 9 Dec. 1929, Moldenke 212 (MBG, NY); hammock along Indian River, Cocoa,
9 May 1918, Small 8732 (NY); low open woods, Indian River, June, Curtiss 1963 in
part (G, MBG, NY, US); Okeechobee region, Brevard Co., 18 April 1903, Fredholm
6804 (G); above Fort Lauderdale, 12 March 1920, J. P. Young 737 (US); Indian
River at Biscayne Bay, 1874, Palmer 6433 (MBG); Tampa, May 1876, Garber (US);
Tampa, 24 Aug. 1895, Nash 2470 (G, NY), TYPE collection; Tampa, 7 March 1898,
[Vor. 20
322 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Pollard (US); St. Petersburg, April 1921, Beckwith 772 (US); Palmetto, 8 May 1900,
Tracy 6650 (G, MBG, NY, US); Braden River, 13 km. southeast of Manatee, 13
June 1918, Barrett 13 (US); dry sandy soil north of Venice, 23 April 1930, Moldenke
1039 (MBG, NY); Owanita, Lee Co., 18 March 1907, W. Kellogg (G); thicket along
river, Pondilla, Lee Co., 8 March 1927, Standley (US).
Verbena tampensis, apparently endemic in peninsular Florida,
is very similar in habit to V. canadensis. It differs in having
more shallowly incised or dentate leaves, a strigillose calyx,
slightly longer nutlets, and essentially glandless anthers. The
nutlets are enlarged at the base somewhat more than those of
closely related species.
39. V. tumidula Perry,” n. sp.
Stems 15-20 cm. long, decumbent-ascending, branching,
pilose-hirsute; leaves broadly ovate with cuneate base narrowed
into a margined petiole, 2-4 cm. long, trifid with segments incised
or coarsely crenate-dentate, appressed-pubescent or strigillose
above, hirtellous beneath; spikes short-pedunculate, protruding
slightly beyond the uppermost leaves, compact; bracts ovate-
lanceolate, acuminate, not exceeding the calyx in length, ciliate;
mature calyx 8-9 mm. long, inflated at base, hirsute, finely
glandular, teeth subulate, 1.5 mm. long; corolla-tube about 11
mm. long, puberulent without; corolla-limb 8-10 mm. broad;
anthers not glandular; nutlets 3 mm. long, dorsal surface retic-
ulate-scrobiculate, lateral surfaces ventricose and smooth;
commissural face almost smooth.
Distribution: Texas and New Mexico.
— ger examine
“from W. Texas to El Paso, New Mexico," May-Oct. 1849, Wright 456
m US); Barkedale, 7 May 1918, E. J. Palmer 18512 (MBG TYPE).
New Mexico: collection of 1851-2, Wright 1503 in part (G, NY).
V. tumidula Perry, spec. nov., herbacea; caulibus A vel ascen-
dentibus ramosis piloso-hirsutis; foliis late ovatis basi cuneata in petiolum alatum
contractis trifidis segmentis inciso- vel grosse crenato-dentatis Sei hirtellis supra
adpresso-pubescentibus vel strigosis; spicis breviter pedunculatis paulo exsertis;
bracteis ovato-lanceolatis acuminatis ciliatis calyce fere paulo brevioribus; calyce
m. longo basim inflato nd et iini glanduloso; calycis dentibus 1.5
mm. longis; corollae tubo circiter 11 mm. longo extus puberulo; corollae limbo 8-10
m. lato; eonneetivo antherarum SUpAHanm inappendiculato; coccis subovoideis
3 mm. longis dorso reticulato-scrobiculatis lateribus ventricosis et glabris; diae
subtiliter muriculata. Collected at Barksdale, Texas, 7 May 1918, E. J. Palmer
18512 (MBG), TYPE.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 323
This species undoubtedly belongs to the section Glandularia,
although apparently the anthers are glandless. Superficially,
it resembles V. Gooddingii var. nepetifolia, but the spikes are not
so large nor so showy. Its distinctive characters are the ventri-
cose nutlets and the inflated fruiting calyx. The bracts, too,
are broader and shorter than those of nearly related species of
this group.
40. V. bipinnatifida Nutt. Jour. Acad. Nat. Sci. Phila. 2: 123.
1821; Schauer in DC. Prodr. 11: 553. 1847.
Glandularia bipinnatifida Nutt. Trans. Am. Phil. Soc. N.S.
5:184. 1837.
More or less diffusely branched from the base; stems loosely
ascending, occasionally rooting at the lower nodes or from sub-
terranean branches, hispid-hirsute; leaves petiolate, blades
2-6 cm. long, bipinnately parted or tripartite with divisions more
or less bipinnatifid, lobes linear or oblong, appressed-hirsute on
both surfaces, margin at times revolute; spikes pedunculate,
fascicle-like in anthesis, becoming elongate in fruit; bracts mostly
longer than the calyx, linear-subulate, hispid-hirsute, ciliate;
fruiting calyx 8.5-10 mm. long, pubescent, hispid-hirsute along
the nerves, lobes very slender, subulate-setaceous from a broader
base, unequal; corolla-tube about one-half longer than the calyx,
pubescent without; corolla-limb 8-10(-12) mm. broad, lobes
emarginate; nutlets cylindrie with slightly broadened base,
mostly 3 mm. long, reticulate-scrobiculate above, striate toward
the base; commissure muricately scabrous.
Distribution: South Dakota to Alabama, westward to Arizona and northern
exico
Specimens examined:
a : between Cahaba and Beloit, Dallas Co., 26 April 1927, Harper 15
(G, N US): on limestone outcrops along small creeks, near Demopolis, Marengo
Co., | May 1925, E. J. Palmer 27209 (MBG); Spring Hill, 1918, Graves 536, 1946
(MBG).
Mississippi: Ocean Springs, 1892, Skehan 47 (G).
UISIANA: vicinity of Alexandria, 19 May 1899, Ball 401 MBG, NY, US);
Cotes S xS St. Mary Co., 17 July 1893, Langlois (MBG, US).
Miss : prairies, Upper Misscurt 21 June 1839, Geyer (US); Courtney, 6 June
1894, Bush p^ (G, MBG); Courtney, 13 June 1906, Bush 4029 (G, MBG, NY, US).
ARKANSAS: “Red River,” Nuttall (G, NY), TYPE collection; dry gravelly hills,
Fulton, Hempstead Co., 17 June 1915, E. J. Palmer 8045 (MBG); near Homan, 10
[Vor. 20
324 ANNALS OF THE MISSOURI BOTANICAL GARDEN
June 1898, Eggert (MBG); along railroads, Miller Co., 19 June 1908, Eggert (NY);
Texarkana, Aug. 1880, Letterman (MBG, NY).
Soutn Daxora: Fort Pierre, Nicollet’s Northwest Expedition (US); Pierre, 2 Sept.
1891, T. A. Williams (MBG, NY); near McClure, June 1910, O. E. White (MBG);
Kennebec, Lyman Co., 1 July 1914, Over 3177 (US); Pine Ridge Reservation,
Washabaugh Co., 22 Aug. 1911, Visher 21382 (F, NY); White River, 16 July 1896,
T. A. Williams (NY); White River flood plain, 4 Aug. 1914, Over 2103 (US); White
River Valley, Shannon Co., 8 July 1911, Visher 2177 (NY); Fall River Falls, 18
June 1892, Rydberg 985 (G, NY, US).
NEBRASKA: Spencer, Boyd Co., 25 July 1893, Clements 2776 (G, US); Callaway,
4 July 1901, Bates (G).
Kansas: Cloud Co., 19 May 1888, Carleton (MBG); near Osborne City, 19 May
1894, Shear 38 (G, NY, US); Manhattan, 24 May 1901, A. Nelson 8245 (G, MBG,
NY, US); plains, Ellis Co., 16 July 1895, Hitchcock 893 (G, MBG, NY, US); Smoky
Hill, June 1867, Parry 15? (G, MBG); vicinity of Hays, 20 July 1929, Rydberg &
Inler 1212 (NY), 1253 (MBG, NY); prairies, Medicine Lodge, 12 Sept. 1890, Smyth
306 (US); Morton Co., July 1891, Carleton 177 (US); vicinity of Richfield, 20 Sept.
1912, Rose & Fitch 17105 (US).
OkLAnoMa: Vinita, 3 Aug. 1877, Gurney & Monell (MBG); Grant Co., 20 June
1899, White 19? (MBG); Woods Co., 29 June ?1899, White 165 (MBG); near Alva,
Woods Co., 28 Sept. 1913, Stevens 2860 (G); near Whitehorse, 31 May 1913, Stevens
699 (G, MBG, NY, US); about 17 km. north of Boise City, Cimarron Co., 22 Aug.
1927, Stratton 449 (MBG); near Shattuck, Ellis Co., 20 May 1914, Clifton 3085 (G,
P); Hollis, Harmon Co., 21 June 1913, Stevens 1108 (G); 10 km. southwest of Hollis,
6 Aug. 1927, Stratton 817 (MBG); vicinity of Fort Sill, 27 April 1916, Clemens 11751
(MBG); on false Washita between Fort Cobb and Fort Arbuckle, 1868, Palmer 4?
(NY); Arbuckle Mountains, Davis, 29 Oct. 1914, Emig $95 (MBG); near Davis, 23
June 1917, Emig 788 (MBG); Mannsville, Johnston Co., April 1916, Griffith 3456
(G); Price’s Falls, Murray Co., 30 April 1926, Stratton 10 (MBG); near Crusher
Spur, Murray Co., 11 April 1913, Stevens 14 (G, MBG, NY, US); near Camp, Texas
Co., 12 May 1913, Stevens 427 (G); Caddo, 22 June 1891, Sheldon (MBG), 48 (US).
Texas: without data, Lindheimer 282 (MBG), 289 (F, G, MBG, NY, US), 307
(G); Canadian, Hemphill Co., 10 Aug. 1900, Eggert (MBG); Canadian, July 1903,
Howell 111 (US); vicinity of Terrell, Kaufman Co., 4 May 1904, T'yler (US); Dallas,
Reverchon ?89 (MBG, US), 1962 (F, G, MBG, NY, US); Dallas, Reverchon (G, NY,
US); Dallas, 22 June 1899, Eggert (MBG); dry ground, near Garland, 24 June 1899,
Eggert (MBG); Fort Worth, Ruth 107 (F, G, MBG, NY, US); Weatherford, 26 May
1902, Tracy 7999 (F, G, MBG, NY, US); Corsicana, Navarro Co., 1 June 1915,
E. J. Palmer ?821 (MBG); near Granbury, Hood Co., 4 May 1900, Eggert (MBG);
Granbury, 4 Sept. 1914, E. J. Palmer 6510 (MBG); near Abilene, 7 June 1900,
Eggert (MBG); Abilene, 19 May 1902, T'racy 8000 (F, MBG); Killeen, 1 Oct. 1891,
Ward (US); Walker Co., 6-12 May 1910, Dixon 561 (F, P); near Austin, Travis Co.,
18 April 1929, E. J. Palmer 33389 (G, MBG); Crab Apple Creek, Gillespie Co.,
Jermy 183 (MBG); about 25 km. west of San Felipe, March 1844, Lindheimer 146
(MBG); San Marcos, 11 June 1897, Trelease (MBG); Comanche Spring, New
Braunfels, Lindheimer 1072, 1073 (F, G, MBG, NY, US); northwest of New Braun-
fels, 14 Sept. 1913, Pennell 5444 (NY); limestone soil mixed with sand, near Bracken,
Groth 80 (F, G, NY, US); Rock Springs, 17 April 1930, Jones 26228 in part (MBG);
Del Rio, 18 April 1930, Jones 26229 in part (P); Fort Clark, Kinney Co., Mearns
1252, 1274, 1894 (US); Eagle Pass, Rio Bravo del Norte, Feb.-March 1852, Schott
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 325
(NY); San Antonio de Bexar, Feb. 1828, Berlandier 1449189 (G); between
Trinity River and Bexar, June 1828, Berlandier 429 —?1749 (G); San Antonio, 4 April
1901, Eggert (G); San Antonio, 1897, Wilkinson 9 (MBG); San Antonio, 15 Aug. 1906,
Ball 909 (US); vicinity of San Antonio, April 1919, von Schrenk (MBG); San Antonio
to Austin, 18 April 1925, Small & Wherry 11968 (NY); 16 km. west of San Antonio,
8 June 1931, Moore & Steyermark 8001 (MBG); along Trinity River, near Liberty, 11
April 1925, Small & Wherry 11781 (NY); Columbia, 27 March 1900, Bush 449 (MBG,
NY, US); Columbia, 3 May 1900, Bush 205 (MBG, US); Cuero, 22 March 1907,
Howell 308 (US); prairies, near Victoria, 10 April 1900, Eggert (MBG); Victoria, 27
April 1905, Mazon $808 (US); Victoria, 7 March 1916, E. "i Palmer 9098 (MBG);
from Goliad to Bexar, May 1834, Berlandier 2428 2998 (G, MBG).
?MoNTANA: sandy soil along "Vellawatons and upper Missouri, 1853-4, Hayden
(MBG).
New Mexico: without data, 1850-2, Wright 1502 (G, NY, US), 1604 in part
(ANSP); below Dofiana, Parry, Bigelow, Wright, Schott (NY); Santa Fe, 20 July
1898, Earle 84 (MBG, NY); Roswell, Aug. 1900, Earle & Earle 526 (MBG, NY).
Arizona: Bill Williams Mountain, 22 July 1898, MacDougal 317 (ANSP, F, G,
NY); by streams of Santa Catalina Mountains, 13 April 1881, Pringle (G); Santa
Rita Mountains, 31 July 1927, Peebles, Harrison & Kearney 4549 (US); Miller’s
Canyon, Huachuca Mountains, July 1909, Goodding 251, 495 (G, NY); road to
Rustler’s Park, Chiricahua Mountains, 18-19 June 1930, Goodman & Hitchcock 1211
(MBG); Barfoot Park, Chiricahua Mountains, 31 Aug. 1906, Blumer 1345 (F, G,
MBG, NY).
Mexico: Cuinvanua: near Colonia Garcia, 21 July 1899, Townsend & Barber 139
(F, G, MBG, P, US); near Colonia Garcia, 1899, Nelson 6117 (G, US); Round
Valley, Sierra Madre, 17 Sept. 1903, Jones (P).
In the specimens from Arizona and Mexico, it should be noted
that the pubescence is hispidulous and the spikes at maturity
are, for the most part, dense rather than elongated.
The typical form of the species is very easily recognized by the
hispid-hirsute pubescence, particularly of the inflorescence, the
lack of glands, the long floral bracts, the elongating spike, the
deeply cut leaves, and a rather coarse habit. Apparently it
intergrades with V. ciliata in northern Mexico, with V. ambrosi-
folia in New Mexico, and at times with V. canadensis in the south-
western part of its range. Nevertheless, in spite of the many
intermediate phases, it seems preferable to maintain each group
as specific entities.
40a. Var. latilobata Perry,” n. var.
Leaves trifid with the segments remotely incised or lobed,
* Var. latilobata Perry, var. nov., folia trifida; segmentis grosse incisis lobisve supra
strigosis subtusque praesertim in nervis et venis hirtellis hirsutisve; spicis maturitate
densis vel elongatis.—Collected between San Pedro and Fronteras, Sonora, Mexico,
20-24 Sept. 1890, Hartman 906 (G), TYPE.
[Vor. 20
326 ANNALS OF THE MISSOURI BOTANICAL GARDEN
strigillose above, hirtellous-hirsute, especially along the veins
beneath; spike compact or somewhat elongated in age; bracts
variable in length, usually equalling or exceeding the calyx;
floral and nutlet characters as in the species.
Distribution: southern Arizona and northwestern Mexico.
Specimens examined:
ONA: Galiuro Mountains, 9 July 1894, — (NY); Bisbee, Oct. 1908,
Qoodding 37 (NY); Flagstaff, 17 June 1887, Mearns (NY); Fort Huachuca, Aug.
1893, Wilcox (NY); Fort Huachuca, Sept. 1894, Wilcox (US); Huachuca Mountains,
29 June-5 July 1903, Griffiths 4833 (US); Huachuca Mountains, 3 Sept. 1903,
Jones (P); Miller's Canyon, Huachuca Mountains, 26 Aug. 1910, Goodding 883
(G, NY); Santa Rita Mountains, 20 Sept.-4 Oct. 1902, Griffiths & Thornber 159 (US);
Santa Rita Mountains, 24 Aug. 1903, Jones (P, US).
Mexico:
CHIHUAHUA: at base of Mt. Mohinora, about 13 km. from Guadalupe y Calvo,
23-31 Aug. 1898, Nelson 4856 (G, US); Colonia Juarez, Sierra Madre Mountains,
11 Sept. 1903, Jones (P); San Diego Canyon, Sierra Madre Mountains, 16 Sept.
1903, Jones (P); without data, Wright 1503 in part (G, NY).
Sonora: between San Pedro and Fronteras, 20-24 Sept. 1890, Hartman 906 (G
TYPE); San Pedro, 14 Sept. 1890, Hartman 880 (G); Cananea, 27 Sept. 1908, Donnelly
(P); Cananea, 20 Aug.-1 Sept. 1914, Murdoch (F).
DunaNao: without data, Garcia 331 (US); Sierra Madre, west of Durango, Sept.—
Oct. 1881, Forrer (F).
The variety differs from the species chiefly in foliar characters.
41. V. ambrosifolia Rydb. in Small, Fl. Southeast. U. S. ed.
1, 1011. 1337. 1908, and ed. 2, 1913.
More or less diffusely branched from the base; stems loosely
decumbent-ascending, somewhat hirsute; leaves short-petiolate,
blades 2-6 em. long, bipinnatifid with the ultimate segments
lanceolate, appressed-hirsute on both surfaces, margin slightly
revolute; spikes pedunculate or subsessile, fascicle-like in anthesis,
somewhat elongating in fruit; bracts a little shorter than the
calyx, lance-subulate, hirsute, ciliate; fruiting calyx 8-9 mm.
long, glandular, pubescent, hirsute particularly along the nerves;
teeth subulate; corolla-tube protruding one-third to one-half
beyond the calyx, pubescent without; corolla-limb 6—8(-10)
mm. wide; nutlets subeylindrie often with slightly broadened
base, 2.5-3 mm. long, reticulate-scrobiculate above, striate
below; commissure muricately scabrous.
Distribution: Oklahoma and Texas to Arizona and northern Mexico.
ecimens examine
OKLAHOMA: near onion, Cimarron Co., 15 May 1913, Stevens 484 (G, MBG).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 327
Texas: 16 km. east of Comstock, 9 June 1931, Moore & Steyermark 3009 (MBG);
Alpine, 30 May 1928, E. J. Palmer 34240 (NY); prairies near Marfa, 14 May 1901,
Eggert (MBG); foothills below McKittrick Canyon, 23 July 1931, Moore & Steyer-
mark 3622 (MBG); Sierra Blanea, May 1913, Orcutt 6096, 619? (MBG); northeast
base of Quitman Mountains, near Sierra Blanca, 4 July 1921, Ferris & Duncan 2474
(MBG).
oLonADO: New Windsor, 18 June 1905, Osterhout 3165 (G); Boulder, July 1891,
Penard 348 (NY); Denver, July 1892, Eastwood (NY); South Denver, 8 June 1891,
E. C. Smith (MBG); foothills, below Colorado Springs, 29 May 1878, Jones 122 (P);
Cafion City, May 1871, Brandegee (MBG); Pueblo, 10 July 1873, Greene 45 (G);
road to Walsenburg, Pueblo, 13 June 1917, E. L. Johnston 986 (MBG); mesas near
Pueblo, 14 May 1900, Rydberg & Vreeland 5676 (NY); Rocky Ford, Otero, 8 June
1900, Osterhout (G fragm., NY Type); vicinity of La Junta, 26 Sept. 1913, Rose &
Fitch 17504 (MBG, US).
New Mexico: collection of 1847, Fendler = in part (ANSP, F, G, MBG, US);
south of Raton on the road to Taos, 30 June 1929, Mathias 545 (MBG); Tierra
Amarilla, Rio Arriba Co., 18 April-25 May 1911. Eggleston 6644 (US); Ojo Caliente,
ear Fairview, 18 July 1904, Wooton (US); in canyon near Santa Fe, 29 June 1846,
Wislizenus 525 (MBG); about Santa Fe, 8-9 Sept. 1881, Engelmann (MBG); near
Pecos, San Miguel Co., 15 Aug. 1908, Standley 4951 (F, G, MBG, NY, US); Pecos,
18 June 1927, Arsène 18615 (P); vicinity of Las Vegas, Anect 100, 143 (G), 179 (NY),
Arsène 18793 (P); Nara Visa, 21 April 1911, Fisher (MBG); vicinity of Bernalillo,
12 July 1926, Arsène & Benedict 16306 (MBG); vicinity of Albuquerque, 6 Oct
1913, Rose & Fitch 17801 (MBG, US); near Albuquerque, 21 June 1926, E. J.
Palmer 31192 (MBG); between Carrizozo and Socorro, 21 July 1928, Wolf 2890 (P);
Socorro, May 1881, Vasey (F, MBG, US); White Mountains, Lincoln Co., Aug.
1897, Wooton 368 (G, MBG, NY, P); Capitan, 8-19 May 1902, Earle 688 (NY);
Lincoln, 31 July 1900, Earle & Earle 286 (MBG, P); South Spring, 2-4 May 1903,
Griffiths 4244 (US); Mimbres River, Grant Co., 20 Aug. 1904, Metcalfe 1231 (MBG);
plains south of White Sands, Dofia Ana Co., 28 Aug. 1897, Wooton 642 (MBG,
NY, US); Queen, 2 Aug. 1909, Wooton (MBG, US); Organ Mountains, Dofia Ana
Co., 16 Aug. 1893, Wooton (MBG
rentem Warsaw Mill, Pima Eo. 3 Dec. 1893, Mearns 2677 (MBG, NY, US);
about 3 km. south of Holbrook, 17 June 1901, Ward (NY, US); Little Colorado,
1869, nisi (US).
Mex
See Candelaria Mountains, 1911, Stearns 104 (US).
Nurvo Leon: Monterey, 17-26 Feb. 1880, Palmer 1050 (MBG, US).
CoanvuILA: about 9 km. east of Saltillo, April 1880, Palmer 1050 (G); Sierra Mojada,
20 April 1892, Jones 634 (P); Sabinas, 21 May 1902, Nelson 6789 (NY, US); Carneros
Station, 31 May 1890, Pringle 3467 (F); fields near Carneros Station, 10 Nov. 1904,
Pringle 13157 (F, G, US
An examination of the type specimens of V. ambrosifolia and
V. Wrightii reveals differences probably sufficient to suggest
distinct but closely related species. Typically the former is
characterized by long calyx-teeth and coarser pubescence;
whereas the latter has short calyx-teeth, finer and somewhat
viscid pubescence. On the other hand, a study of the aggregate
[Vor. 20
328 ANNALS OF THE MISSOURI BOTANICAL GARDEN
specimens of the two shows a high variability in the combination
of these characters; moreover, their geographic ranges practically
coincide. For (hose reasons, V. ambrosifolia is maintained as a
species with very doubtful status.
41a. Forma eglandulosa Perry,? new form.
Differing from the species only in the more hirsute-hispid
pubescence of the flower and the lack of glands.
Distribution: Texas to Arizona, and Sonora, Mexico.
Specimens examined:
Texas: “El Paso to Monument no 53," Sept. 1892, Wagner 974 (US).
New Mexico: Albuquerque, 21 June 1926, E. J. Palmer 31158 (MBG); 8 km.
west of Magdalena, 20 July 1904, Wooton 2835 (US); Organ Mountains, 14 July
1897, Wooton (US); mesa just west of Organ Mountains, 13 June 1906, Standley
(US); Santa Rita, 1 Aug. 1911, Holzinger (MBG rypx, US); Deming, 13 July 1917,
Munz 1231 (P); Los Playas, 1854-5, Antisell 186 (NY); near White Water, 11 Sept.
1893, Mearns 2278 (US); Animas Valley, 2 Oct. 1893, Mearns 2486 (US).
Arizona: Douglas, 13 May 1915, Carlson (G, US); Santa Catalina Mountains, 15
Sept. 1896, T'oumey (NY, US); Dragoon, 18 July 1920, W. W. Jones 185 (G); Cochise,
12 Oct. 1900, Griffiths 1919 (NY).
Mexico: Sonora: San Pedro River, 15 Oct. 1892, Mearns 1138 (US).
This is a phase of the species lacking glands and only separable
from V. bipinnatifida by the short bracts.
42. V. Wrightii Gray, Syn. Fl. N. Am. 2!:: 337. 1878.
Stems usually several from a common base, decumbent-
ascending to erect, branched, sparsely hispid-hirsute; leaves sub-
sessile or contracted into a short margined petiole, blades 2-3
(-4) em. long, bipinnatifid or trifid, with the divisions more or
less deeply incised, ultimate segments narrowly lanceolate,
hirtellous to hirsute on both surfaces; spikes short-pedunculate,
in fruit more or less compact; bracts shorter than the calyx,
lanceolate, acute or acuminate, hirsute-ciliate; fruiting calyx
-9 mm. long, densely glandular, somewhat viscid-pubescent,
nerves hirsute, lobes short, subulate, unequal; corolla-tube
11-12 mm. long, pubescent around the throat; corolla-limb 6-8
mm. broad, lobes retuse; nutlets 2.5-3 mm. long, reticulate-
Borabieulate above, siate toward the base; commissure muri-
cately scabrous.
** Forma eglandulosa Perry, forma nova; calyce hirsutiori-hispido oe —
Colleeted at Santa Rita, New Mexico, 1 Aug. 1911, Holzinger (MBG), T
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 329
Distribution: bee Colorado, New Mexico, and Arizona.
Specimens exa ;
Texas: San picti Havard (US); Barstow, 16 April 1902, Tracy & Earle 61
(F, G, MBG); northeast of Grand Falls, 17 May 1917, Clawson 13923 (US); near
Feodora, Terrell Co., 26 May 1928, E. J. Palmer 33583 (MBG); Upper Blue Creek
Canyon, Chisos i cope 1 July 1931, Moore & iibi $324 (MBG); Fort
Davis, 9-12 July 1921, is & Duncan 2647 (MBG, P); Davis Mountains, 23
April 1902, T'racy & EL 168i in part (F, G, MBG, La foothills, Davis Mountains,
15 April-3 May 1902, Tracy & Earle 162a (NY); canyon of Upper Limpia Creek,
10 June 1926, E. J. Palmer 30672 (MBG); Guadalupe Mountains, 9 Sept. 1916,
Young (MBG); flats near Van Horn, 13 May 1901, Eggert (G, MBG); Vieja Moun-
tain, Oct. 1883, Havard 97 (G, US); Canutillo, Franklin Mountains, 12 July 1911,
Barlow (US); mountains near Fronteras, 22 March 1852, Wright (MBG, NY), 1504
in part (G Type); Hueco Mountains, March 1851, Thurber 143 (G); about 8 km.
north of Shafter, 10 April 1919, Hanson 551 (MBG ).
CoronRApo: Durango, 20 July 1898, Baker, Earle & Tracy 531 (G, MBG, P).
New Mexico: without locality, 1851-2, Wright 1603 in part (MBG); Copper
Mines, 28 July 1851, Bigelow (NY); Rabbit Ear Creek, 28 June 1846, Wislizenus 483
(MBG); Las Vegas, 26 June 1895, Mulford 87 (MBG, NY); hills, Santa Fe, 15 May
1897, Heller 3536 (G, MBG, P); Sandia Mountains, 6 Sept. 1884, Jones 457 (P);
Sandia Mountains, 28 April 1914, Ellis 17 (MBG); about 3 km. east of Albuquerque,
1915, Kammerer 83 (MBG); Albuquerque, Sept. 1884, Jones (NY, P); Albuquerque,
5 Sept. 1909, Rusby (NY); mountains west of Grant’s Station, 2 Aug. 1892, Wooton
(US); South Camp, Magdalena Mountains, July 1897, Herrick 704 (US); White
Mountains, Lincoln Co., 20 Aug. 1897, Wooton 864 (G, MBG, P); near Lincoln
National Forest, 1903, Plummer (US); Vaughn, 10 Sept. 1921, Harwood (P); plains,
White Sands, Otero Co., 25 Aug. 1899, Wooton (US); High Rolls and vicinity, 21-28
May 1902, Viereck (ANSP); Alamogordo, 7 April-24 May 1902, Rehn & Viereck
(ANSP, MBG, P); Kingston, Sierra Co., 9 July 1904, Metcalfe 1090 (G, MBG, NY,
P, US); Gallina’s Canyon, Black Range, Grant Co., 17 Aug. 1915, Pilsbry (ANSP);
Mangas Springs, about 29 km. northwest of Silver City, 11 June 1903, Metcalfe 126
(G, MBG, P); near Silver City, June 1880, Greene (MBG, P); Silver City, 8 May
1919, Eastwood 8468 (G); Rincon, 16 May 1890, Jones (P); Big Burros Ranger
Station, Gila Forest, Grant Co., 11 Sept. 1920, Eggleston 17231 (MBG, NY); Organ
Mountains, 8 May 1895, Wooton (P); Organ Mountains, 11 June 1906, Standley
(MBG, US); Filmore Canyon, Organ Mountains, Dofia Ana Co., 25 May 1905,
Wooton (US); Van Pattens, Dofia Ana Co., 29 Aug. 1894, Wooton (US); Slaughter
Canyon, 12-20 Aug. 1924, Standley 40625 (US).
RIZONA: Long H. Ranch to St. John’s, 6-15 Aug. 1913, Griffiths 5190 (US);
between Springerville and St. John’s, Apache Co., 15 Sept. 1917, Marsh 14224 (MBG).
On account of its strong variability, this species has been
exceedingly difficult to define. It was first set apart as an erect
annual, with anther-glands as high and almost as large as the
anther-cells. In the series of specimens at hand, the anther-
glands are neither constant in size nor in height relative to the
anther-sacs. The main characters appear to be the glandular
somewhat viscid pubescence of the calyx, the very short acute-
subulate calyx-teeth, and the somewhat compact spikes.
[Vor. 20
330 ANNALS OF THE MISSOURI BOTANICAL GARDEN
43. V. ciliata Denth. Pl. Hartw. 21. 1839.
Stems several from a common base, decumbent, branched,
hirsute-hispidulous; leaves short-petiolate or subsessile, subbi-
pinnatifid or commonly trifid, with the divisions more or less
deeply incised, ultimate segments linear-oblong with slightly
revolute margin, hirsute-strigillose on both surfaces; spikes
subsessile or short-pedunculate, in fruit more or less compact;
bracts a little shorter than the calyx, lanceolate, subulate or
acuminate, hirsute-ciliate; calyx 7-8 mm. long, somewhat
glandular, hirsute-hispidulous, lobes short, subulate, unequal;
corolla-tube 10-12 mm. long, pubescent without; corolla-limb
6-8 mm. broad, lobes retuse; nutlets 2.5-3 mm. long, reticulate-
scrobiculate above, striate toward base; commissural faces
muricately scabrous.
Distribution: New Mexico, Arizona to southern Mexico.
Specimens examined:
Ne Ew Mexico: “Upper Corner Monument, parallel 31° 47," 3 May 1892, Mearns
108 (G, NY).
ARIZONA: without locality, 1892, Towmey 305% (US); Ash Fork, 18 June 1901,
Barber 106 (US); Ash Fork, 20 June 1903, Griffiths 4757 (US); Aguila, 3 April 1930,
Jones 26231 (MBG, P); Snowflake, 30 July 1897, Zuck (US); Pinedale, 1-15 Aug.
1897, Hough 109 (US); Fort Grant, Bonita, 18 July 1917, Munz 1249 (P); between
Tombstone and Bisbee, 23 May 1928, Peebles 5353 (US).
Mexico: without data, Coulter (ANSP); without locality, 26 July 1885, Schu-
mann 1071 (G, US).
Sonora: San Bernardino, Aug. 1852, T'hurber 769 (G).
CnurinvAnua: Saint Eulalia plains, 26 Sept. 1885, Wilkinson (US); highlands near
Chihuahua, 28 April 1847, Gregg (G, MBG, NY); valley near Chihuahua, 13 Sept.
1846, Pringle 1117 (F, US); vicinity of Chihuahua, 8-27 April 1908, Palmer ?9 (F,
G, MBG, NY, US); vicinity of Madera, 27 May-3 June 1908, Palmer 281 (G, NY,
US).
DvunaNao: Valley of Nazas, Bolson de Mapimi, 11 May 1847, Gregg 632 (MBG);
near El Salto, 12 July 1898, Nelson 4559 (MBG, US); Durango and vicinity, April-
Nov. 1896, Palmer 345 (F, G, MBG, NY, US); Durango, 1 Aug. 1898, Nelson 4604
(MBG, US); Otinapa, 25 July-5 Aug. 1906, Palmer 401 (F, G, MBG, NY, US).
OAHUILA: near Saltillo, 1847, Gregg 57, 257 (MBG); Saltillo, 15-30 April 1898,
Palmer 74 (F, G, MBG, NY, US); Saltillo, 24 Aug. 1926, Fisher 223 (US); Oro, 18
Aug. 1903, Rose & Painter 6431 (US); La Ventura, 2-5 Aug. 1896, Nelson 3920 (US);
Parras, March 1905, Purpus 1095 (NY); Valley of Nazas, ius 49, Gregg 26 (MBG);
El Toro, near Movano, July 1910, Purpus 4524 in part (F, MBG).
SAN Lala Porosr: near San Luis Potosi, 1876, Vis Md 717 (ANSP, G); near San
Luis Potosi, 1878, Parry & Palmer 719 (ANSP, ;
ZACATECAS: Zacatecas, July 1904, Kuntze 424 jeg Hacienda de Cedros, 1908,
Lloyd 159 (US); near Plateado, 3 Sept. 1897, Rose 2769 (US); near San Juan Capis-
trano, 19 Aug. 1897, Rose 2435 (US).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA Sol
AGUASCALIENTES: near Aguas Calientes, 9 Oct. 1903, Rose & Painter 7742 (US).
Nayarit: Tepic, 5 Jan.-6 Feb. 1892, Palmer 2055 (US); Ixtlan, 19 Feb. 1927,
Jones 23244 (MBG, NY, P).
Jauisco: near Guadalajara, 11 Aug. 1902, Pringle 11091 (F, MBG, NY, US);
near Guadalajara, 23 Feb. 1907, Safford 1398 (US); Guadalajara, Aug. 1909, Furness
(F, US); Rio Blanco, June-Oct. 1886, Palmer 35 (G, NY, US).
GUANAJUATO: Leon, 1839, Hartweg 176 (G, NY), TYPE collection; Guanajuato,
1894, Duges 496, 496a (G); Obregon, 28 July 1926, Fisher 207 (US).
QUERETARO: near San Juan del Rio, 17 Aug. 1905, Rose, Painter & Rose 9503 (US);
Queretaro, 1910-13, Arsène & Agniel 10252 (F, G, MBG, yt US).
Hripauao: above Pachuca, 23 July 1898, Pringle 7691 (F,
VERA Cruz: Orizaba, 1853, Mueller 325, 1209, 1319 pals Oraibi 4 Aug. 1891,
Seaton 150 (F, G, US); Maltrata, Jan. 1883, Kerber 255 (US); Maltrata, 16 Aug.
1891, Seaton 391 (F, G, NY, US).
PuEBLA: Cholula, 1 Jan. 1899, Deam (F), 86 (G); Tehuacan, Galeotti 736 (NY, US);
near Tehuacan, 8 Sept. 1906, Rose & Rose hee (G, NY, US); vicinity of San Luis
sare umen Se July 1908, Purpus 3407 (G,
Mexico: meadows near Lecheria, 12 d nx Pringle 13158 (F, G, US); Valley
of Mexico, 1865-6, Bourgeau 120 (G); Eslava, 15 June 1901, Pringle 9313 (G, MBG,
NY, US); Amecameca, 24 July 1924, Fisher (F), 319 (MBG, US); near San Angel, 9
April 1849, Gregg 648 (MBG).
MicnoacaN: low valley, Zinapecuaro, 2 May 1849, Gregg 756 (MBG); Querendaro,
28 Oet. 1895, Seler 1174 (G); Los Reyes, 8-12 Feb. 1903, Nelson 6858 (US).
DERE voleano of Colima, 14 July 1892, Jones 686 (P).
OAXACA: Cerro de las Soledad, 20 Nov. 1895, Seler 1379 (G); Las Sedas, 8 Sept.
1894, c. L. Smith 221 in part (US).
This is a variable and wide-ranging species closely related to
the bipinnatifida-Wrightii-ambrosifolia complex and scarcely
capable of sharp delimitation. Its best characters are the
hirsute-hispid pubescence, the usually sessile compact spike, the
medium-short somewhat glandular calyx subtended by a slightly
shorter bract. The specimens, Seaton 150, Pringle 9318, and
Gregg 648, have typical spikes, but the leaves are small and the
stems tend to be repent, suggesting a possible intergradation
with V. teucritfolia.
43a. Var. longidentata Perry,?’ n. var.
Hispidulous-hirsute plant of variable size and more or less
open habit; leaves bipinnatifid; calyx-teeth 2-2.5 mm. long.
Distribution: BÉ Texas, New Mexico, and Tamaulipas.
Specimens examined:
Texas: Post, 22 ie 1925, Wooton (US); Lubbock, 14 Máy 1930, Demaree 7685
27 Var. longidentata Perry, var. nov., planta hispidulo-hirsuta; foliis bipinnatifidis;
calycis dentibus 2-2.5 mm. longis.—Collected at Matamoros, Mexico, April 1836,
Berlandier 3020 (MBG), TYPE.
[Vor. 20
332 ANNALS OF THE MISSOURI BOTANICAL GARDEN
(US); Vista, May 1892, T'release (MBG); woods near Colorado River, near Wharton,
12 April 1925, Small & Wherry 11884 (NY); hills near San Antonio, 4 April 1901,
Eggert (MBG); El Jardin, 10 March 1924, Runyon 630 (US); Laredo, Feb.-March
1913, Orcutt 5657 (MBG); Corpus Christi, 11 May 1900, Bailey 258 (US); Corpus
Christi, May 1913, Orcutt 5792 (MBG); Aransas Pass, 1922, Schulz 845 (US);
Kingsville, 25 March 1920, High 59 (MBG); Brownsville, 4 May 1900, Bailey 221
(US).
New Mexico: Hagerman, 26 April 1929, Benke 5046 (F, G, MBG).
Mexico: TAMAULIPAS: Matamoros, April 1836, Berlandier 1520 (G), 3020 (G,
MBG rypr, NY).
Scarcely differing from the species except in the relatively very
long calyx-teeth and the often somewhat less deeply cleft leaves.
43b. Var. pubera (Greene) Perry, n. comb.
V. pubera Greene, Pittonia 5: 136. 1903.
Plants with more or less prostrate compact habit; ultimate
segments of the leaves linear-oblong, margins strongly revolute.
Distribution: rop to Arizona.
Specimens exam t
EXAS: Davis om 23 April 1902, Tracy & Earle 162 in part (MBG), TYPE
collection of V. pubera; Sierra Blanca, El Paso Co., 22 May 1913, Orcutt 6172 (MBG).
New Mexico: low mountains west of San Antonio, 14 April 1908, Wooton 3849
(US); near Silver City, May 1880, Greene (MBG, P); Deming, 29 April 1884, Jones
3801 (NY, P, US); Deming, 9 April 1930, Jones 26232 (MBG, P); Lordsburg, 3
July 1891, Evans (MBG); east of Lordsburg, 5 — 1930, Jones 26227 (MBG, P);
Hudson Hot Springs, 2 —— Ag diee 804 (N
ARIZONA: Cameron's, 16 June 1929, Jones (P); pM of Flagstaff, 8 Aug.
1922, Hanson A146 (F. MBG, fo Peach Springs, 15 June 1930, Jones 25480
(MBG, P); Dewey, 16 July 1922, W. W. Jones 180 (G); Taylor, 1 Aug. 1897, Zuck
(MBG, NY); Chiricahua Mountains, 5 April 1897, T'oumey (US).
This variety is a low plant with several stems from a common
base, short internodes, and a tendency to branch freely. The
leaves are more finely dissected than in the species and the margins
are more revolute.
44. V. racemosa Eggert, Torreya 2: 123. 1902.
V. pulchella Greene, Pittonia 5:136. 1903.
Stems several from a common base, ascending-erect, branched,
pubescent; leaves bipinnatifid or trifid with segments deeply
cleft, ultimate segments linear, somewhat pubescent or hirtellous
on both surfaces; spikes subsessile, elongate but dense at maturity ;
braets scarcely so long as the calyx, lanceolate, acuminate,
ciliate; calyx 5-6 mm. long, sparsely glandular-hirtellous, teeth
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 333
short, acute-subulate; corolla-tube 7-9 mm. long, puberulent or
glabrous without; corolla-limb 5-6 mm. broad; nutlets 2 mm.
long, reticulate-scrobiculate almost to base; commissural faces
muricately scabrous.
Distribution: perg New Mexico?
DT exam -
Exas: Upton Co., 2 May 1929, Cory 664 (G); Stockton, June 1881, Havard (US);
Beier of Davis Mauniatus, 21 April 1902, Tracy & Earle 106a (F, G); low valleys
near Sierra Blanca, El Paso Co., 15 May 1901, Eggert (G, MBG); Sierra Blanca, 22
May 1913, Orcutt 6184 (MBG); valley of the Rio Grande below Dofiana, Parry
jaa Wright & Schott in part (US).
w Mexico: without locality, 1851-2, Wright ?1498 (NY), 1501 (G).
Fine pubescence, short calyx-teeth, small leaves, and elongated
profusely floriferous spikes are the distinctive characters of this
species. It is probably a close relative of V. ciliata.
45. V. Andrieuxii Schauer in DC. Prodr. 11: 553. 1847.
Stems diffuse; branches procumbent, sub4-angled, hispid;
leaves 4 cm. long, cuneate at base, sessile, deeply trifid, rugose
with veins impressed above, strigose-hispid on both sides, middle
lobe pinnatifid, lateral lobes spreading, lanceolate, acutish,
margin incised or entire, subrevolute; spikes terminal and axillary,
short-pedunculate, oblong, crowded, hispid, subglandular; bracts
one-third shorter than the calyx, subulate-lanceolate; calyx 8-9
mm. long, teeth subulate; corolla-tube one-half longer than the
calyx, puberulent, corolla-limb medium; schizocarp one-half the
length of the calyx.
isi pe southern Mexico.
Specimens examined:
duo
?Mexico: between Mexico and Oaxaca, Andrieux 138 (DeCandolle Herb. TYPE, F
phot., K, MBG phot.).
Oaxaca: mountain ridge, west side of valley of Cuicatlan, 10 Nov. 1894, Nelson
1895 (US); Tieneguilla, 5 April 1895, L. C. Smith 348 (G); valley of Oaxaca, 2 Oct.
1894, Nelson 1613 (G, US)
The above description is a close translation of the original.
Even with the photograph of the type, the species is difficult to
interpret without floral dissections. For the present, it seems
preferable to maintain its status, referring here the above some-
what similar collections.
[Vor. 20
334 ANNALS OF THE MISSOURI BOTANICAL GARDEN
46. V. teucriifolia Mart. & Gal. Bull. Acad. Brux. 11°: 322.
1844.
V. exilis Schauer in DC. Prodr. 11: 553. 1847.
Stems several from a common base, prostrate or repent, more
or less diffusely branched, 4-angled, glabrate or hirtellous;
leaves 1.5-2.5 em. long, cuneate at base, narrowed into a margined
petiole, pinnatifid-incised, segments linear-oblong, obtusish,
often glabrous or sparsely strigillose on both surfaces, nerves
impressed above, subglaucescent beneath; margin slightly rev-
olute; spikes terminal, sessile or short-peduncled, scarcely
emerging from the highest leaves, few-flowered; bracts approxi-
mately one-half as long as the calyx, lanceolate, acuminate-acute,
ciliate; calyx 5-7 mm. long, sparsely hirtellous or occasionally
hirsute, lobes short, acute; corolla-tube a little longer than the
calyx; corolla-limb 5-7 mm. broad; anther-glands minute;
nutlets 2.5-3 mm. long, chiefly reticulate-scrobiculate; commissu-
ral face muriculate or almost smooth.
Distribution: Nexioe and Guatemala.
Specimens examine
Mkxico:
San Luis Porosi: Huaxalote, Ehrenberg 131 (Bot. Mus. Berl.-Dahl. TYPE of V.
exilis; MBG phot.).
doce ye Leon, June-July 1841, Liebmann 11329 (US).
Hiparco: near Real del Monte, 2 June 1899, Rose & Hough 4484 (US).
Vera Cruz: sand hills, Perote, Halsted (NY); Perote, near Jalapa, 1894, C. L. Smith
1474 (G); peak of Orizaba, June-Oct. 1840, Galeotti 777 (K), TYPE collection; Vaqueria
del Jacal and Orizaba, Oct. 1841, Liebmann 11328 (US); “Talvare,” Feb. 1843,
Liebmann 11330 (US); Boca del Monte, 13 March 1894, Nelson 196, 223 (US).
: common in dry volcanic sand, Chalchicomula, 8 April 1890, Stone
(ANSP); Chalchicomula, 20 Feb. 1892, J. G. Smith 464 (MBG); near Chalchicomula,
15 March 1894, Nelson 242 (US); Popocatepetl, 7-8 Aug. 1901, Rose & Hay 6049
(US).
Mexico: La Cima, Jalapa, Aug. 1904, Kuntze 23738 (NY); Sierra de las Cruces,
Salazar, 13 Aug. 1896, Harshberger 41 (ANSP, G, US); lava beds, near Eslava, 19
July 1910, Rusby 124 (NY); near Toluca, 23 June 1889, Pringle 2927 (F, G); valley of
Toluca, 15 Aug. 1892, Pringle 4180 (ANSP, F, G, MBG, NY, US); San Angel, 12
Aug. 1910, Orcutt 3551 (F, MBG, US).
More tos: Toro, 5 Aug. 1924, Fisher (F, MBG), 337 (US); Tres Marias, 11 Aug.
1906, Pringle 13792 (G, US).
Mıcnoacan: Morelia, 9 July 1910, Arsène (F).
OAXACA: : Chinantle, May 1841, Liebmann 11308, 11316 (US).
ENTRAL AMERICA: GUATEMALA: near Quezaltenango, 23 June 1882, Lehmann
1603 (US); Quezaltenango, Oct. 1886, von T'uerckheim 1068 (ANSP, G, US).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 335
The specimens cited vary greatly in degree of pubescence.
The larger number are sparsely hirtellous or strigillose. The
following are densely hirtellous-hirsute: Liebmann 11329, Nelson
196, 223, Smith 1474, Halsted, Smith 464, Nelson 242
The Guatemalan collections appear to be conspecific, although
both the cilia of the bracts and the trichomes along the nerves
of the calyx are somewhat coarser and stiffer than those of the
Mexican material. V. teucriifolia is a slender plant easily
separated from its nearest relatives by its prostrate or repent
habit and few-flowered spikes.
46a. Var. corollulata Perry;?? n. var.
Usually finely and densely pubescent; calyx 5 mm. long,
spreading-pubescent, lobes very short; corolla inconspicuous,
tube barely protruding beyond the calyx, limb 3 mm. broad;
nutlets 2.5 mm. long.
Distribution: Mexico.
Specimens examined:
XICO:
San Lurs Porosi: region of San Luis Potosi, 1878, Parry & Palmer 718 (G, MBG
TYPE, US), 7/9 in part (ANSP, F, MBG, US).
ZACATECAS: plains, LaHonda, 18 Aug. 1890, Pringle 8651 (G); Zacatecas, 1903,
Purpus 187 (US).
Hipauao: Hacienda Palmar, near Pachuca, 21 July 1905, Rose, Painter & Rose
8823 (G, NY, US); Telles, Sept. 1910, Orcutt 4146 (F, MBG).
The variety differs from the species chiefly in its smaller
corollas; perhaps it is only a dimorphic form.
47. V. Gooddingii Briq. Ann. Conserv. & Jard. Bot. Genève
10: 103. 1907.
V. verna var. fissa Nelson, Am. Jour. Bot. 18: 437. 1931.
Stems usually several from a common base, erect or decumbent-
ascending, cinereous-green, branched, densely pilose or some-
what villous, often glandular; leaves 3-5 cm. long, tapering at
the base into a short margined petiole, 3-cleft, divisions coarsely
toothed or incised, cinereous-green, more or less villous-hirsute
28 Var. corollulata Perry, var. nov., planta dense pubescens; calyce 5 mm. longo
patenti-pubescente; corolla inconspicua, tubo vix exserto, limbo 3 mm. lato; coccis
2.5 mm. longis.—Collected in the region of San Luis Potosi, Mexico, 1878, Parry &
Palmer 718 (MBG), TYPE.
[Vor. 20
336 ANNALS OF THE MISSOURI BOTANICAL GARDEN
on both surfaces, midrib and veins slightly prominent beneath,
margins scarcely (if at all) revolute; spikes pedunculate, fascicle-
like in anthesis, somewhat elongated in fruit; bracts usually a
little shorter than the calyx, lanceolate, acuminate, villous-
hirsute, long-ciliate; calyx 8.5-11 mm. long, villous-hirsute, more
or less glandular, lobes slender, subulate, unequal; corolla-tube
very little longer than the calyx, pubescent without; corolla-
limb 8-9(-12) mm. wide, segments retuse; nutlets 3(-3.5) mm.
long, subcylindrie, reticulate-scrobiculate except at the striate
base.
Distribution: Utah, Arizona to California, and Lower California.
Specimens examined:
Uram: southern Utah, 1876, Johnson (US).
Nevapa: sandy soil, Charleston Mountains, about 1800 m. alt., May-Oct. 1898,
Purpus 6061 (P, US); Trout Creek, Charleston Mountains, about 1900 m. alt.,
1926, Jaeger (P); Calientes, about 1400 m. alt., 29 April 1914, Jones (P); Kernan,
Valley Meadow Wash, 28 April 1902, Goodding 645 (F, G, MBG, P), Tyre collection.
Arizona: without locality, 1891, MacDougal 603 (NY); Chloride, about 1350 m
alt., 15 April 1903, Jones (P); Hackberry, 24 May 1884, Jones (F, P, US); Peach
Springs, June 1884, Lemmon (US); Peach Springs, April 1893, Wilson 99 (US);
Kingman, 11 March 1912, Wooton (US); Skull Valley, about 1300 m. alt., 28 April
1903, Jones (P); Hillside, about 1100 m. alt., 1 May 1903, Jones (P); near Williams,
11 April 1905, Wilcor (US); Jerome Divide, 25 July 1921, W. W. Jones (MBG);
east of Jerome Junction, 1 May 1908, Tidestrom 898 (US); Rio Verde, Fort Whipple,
30 Aug. 1865, Cowes & Palmer 498 (MBG); Fort Verde, 1888, Mearns (NY); Natural
Bridge, 23 April 1904, F. M. Chamberlain 56 (US); Bradshaw Mountains, 22 June
1892, Toumey 305a (US); 16 km. south of Black River, White Mountains, about
1800 m. alt., 22 June 1930, Goodman & Hitchcock 1297 (MBG, NY); Galiuro Moun-
tains, 29 July 1894, Toumey (US); Tucson, 16 May 1896, Zuck (NY, US); washes,
near Baboquivari Mountains, 24 Feb. 1923, Hanson A1021 (F, MBG); Baboquivari
Mountains, 10 March 1926, T'hackery & Leding 1100 (US); south of Warren, Sulphur
Springs Valley, 3 June 1915, Carlson (US); Lowell, May 1884, W. F. Parish 196, 197
(G); wash near Agua Caliente, 20 Jan. 1920, Bartram 255 (ANSP).
CALIFORNIA: east side, summit of Providence Mountains, 29 May 1861, Cooper
(US); vicinity of Bonanza King Mine, east slope of Providence Mountains, about 1200
alt., 21-24 May 1920, Munz, Johnston & Harwood 4252 (P); Rock Spring, San
Bernardino Co., May 1876, Palmer (G), 539 (F, MBG, NY, US); southern part of
San Diego Co., 1875, Palmer (G).
Mexico: Lower Cauirornias: Pifione Forest, 25 July 1883, Orcutt (F).
In habit V. Gooddingii is much like V. ciliata, but differs in the
villous-hirsute pubescence and the larger flowers. It is readily
recognized in its typical form by the long-ciliate character of
the bracts, the short dense villous spikes, and the shaggy pubes-
cence.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 337
47a. Var. nepetifolia Tidestrom, Proc. Biol. Soc. Wash. 38:
15. 1925.
V. arizonica Briq. Ann. Conserv. & Jard. Bot. Genéve 10: 102.
1907, not V. arizonica Gray, Proc. Am. Acad. 19: 95. 1883.
V. verna Nelson, Am. Jour. Bot. 18: 436. 1931.
Leaves broadly ovate, coarsely and unevenly dentate, at
times lobed, abruptly narrowed into a cuneate base; pubescence
varying from strongly pilose to densely villous.
Distribution: Colorado, Nevada, Arizona, and northwest
Mexico.
Specimens examined:
eae southern Colorado, 1867, Parry (US).
Nevapa: El Dorado Canyon, Lincoln Co., 1880, T. W. Davis (MBG); El Dorado
at Nelson, 30 April 1907, Jones (P).
ARIZONA: Oatman, 4 May 1928, Thackery 348 (US); Hackberry, 7 March 1912,
Wooton (US); Camp Verde to Prescott, Aug. 1896, Fernow (US); by streams of
Santa Catalina Mountains, 13 April 1881, Pringle in part (F, G, MBG); Tucson,
1871-5, Rothrock (US); Tucson, 1881, Vasey (US); Tucson Mountain, 7 March 1901,
Griffiths 2421 (NY); Tucson Mountain, 13 March-23 April 1903, Griffiths 3489 (MBG,
US); Laboratory Hill, vicinity of Tucson, April 1908, Rose 11968 (US); Tucson,
1909, Parish (P); vicinity of Tucson, 1910, Rose, Standley & Russell 15176, 15181
(NY, US); Tucson, 1911, Beard (MBG); Tucson Mountain, oe 11, 1913, Greenman
& Greenman 56 (MBG); Tumanoc Hill, Tucson, 25 July 1916, J. A. Harris C16375
(NY); rocky slopes west of Tucson, 26 Dec. 1919, Bartram b (ANSP, US); near
Tucson, 17 March 1927, Harrison & Kearney 3646 (US); Sells, Papago Reservation,
24 Feb. 1926, Loomis & Thackery 910 (US); Huachuca Mountains, Aug. 1882,
Lemmon (US); south Huachuca Mountains, 31 May 1930, Peebles 6755 (NY);
Yucca, 14 May 1884, Jones 3901 (F, P, US), TYPE collection of V. arizonica Briq.,
36 (G)
Mexico:
Lower CALIFORNIA: Pifione Forest, 6 Oct. 1882, Orcutt (US); San Jacinto, 18 July
1885, Orcutt 1303 (F, MBG, NY); Rosario, 3 May 1886, Orcutt 5141 (F, MBG, NY);
Agua Dulce, 48 km. southeast of San Fernando, 9 Sept. 1905, Nelson & Goldman
7123a (MBG, US)
ONORA: ‘‘Niggerhead Mountains, near Monument no. 82," Aug. 1893, Mearns
1904, 1914 (US); Alamos, 26 March-18 April 1890, Palmer 307 (G, US); Alamos, 27
Jan. 1899, Goldman 292 (G, NY, US); Sierra de Alamos, 14 March 1910, Rose,
Standley & Russell 12840 (NY, US); Alamo, West Magdalena, 17 May 1925, Kennedy
7033 (US); Pinocate Mountains, 21 Nov. 1907, MacDougal 73 (US); La Cienega, 18
July 1911, Goodding 952 (US).
SiNALOA: without locality, 1922, Ortega 4541 (US); sandy soil along river, vicinity
of Fuerte, 25 March 1910, Rose, Standley & Russell 13449 (NY, US).
The variety differs from the species in its much less indented
and broader leaves. The pubescence is much more variable and
at times harsh, suggesting an intergradation with V. bipinnatifida.
[Vor. 20
338 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The leaf-forms recall V. canadensis, but the spike is scarcely
distinguishable from V. Gooddingit.
48. V. pumila Rydb. in Small, Fl. Southeast. U. S. ed. 1, 1010.
1903, and ed. 2, 1913.
V. inconspicua Greene, Pittonia 5: 137. 1908.
V. brevibracteata Eggert, Torreya 2:124. 1902, not V. bracteosa
var. brevibracteata Gray, Syn. Fl. N. Am. 2!: 336. 1878
Stems usually several from a common base, branched, de-
eumbent-ascending, hirsute, often finely glandular; leaves 1.5-3
em. long, obtusely triangular with cuneate or truncate base
contracted into a short narrowly margined petiole, trifid, occa-
sionally lobed, divisions variously incised, appressed-hirsute on
both surfaces; spikes short-peduncled to sessile, fairly compact;
bracts almost as long as the calyx, linear-lanceolate, hispid-
hirsute; fruiting calyx about 6 mm. long, pubescent, hispidulous
along the nerves, at times finely glandular, lobes subulate, short;
corolla-tube a little longer than the calyx, slightly, if at all,
pubescent without; corolla-limb 3-5 mm. broad; anther-glands
minute or wanting; nutlets 2.5(-3) mm. long, reticulate-scrobic-
ulate except at base; commissural faces muriculate.
Distribution: erect Texas, New Mexico; and Sonora and Sinaloa, Mexico.
Specimens examine
OKLAHOMA: Caton, 8 May 1895, Bush 1275 (MBG, NY mvrE); Oklahoma City,
11 May 1891, Carleton 134 (US); vicinity of Fort Sill, 6 May 1916, Clemens 11752
(MBG); Arbuckle Mountains, Davis, 1 April 1916, Emig 401 (MBG); Price’s
Falls, Murray Co., 30 April 1926, Stratton 11 (MBG); near Crusher Spur, Murray
Co., 12 April 1913, Stevens 28 (G, MBG, NY, US); near Tishomingo, 15 April 1916,
Houghton 2570 (G).
Texas: indefinite data, Lindheimer 501 (F, G, MBG, US); Dallas, 17 March
1876, Reverchon 788 (MBG, US), 1963 (F, G, MBG, NY), 2117 (MBG); Poly-
technic, 10 April 1913, Ruth 110 (MBG); Tarrant Co., 10 June 1913, Ruth 110 (G);
Haskell, 1898, Morton (MBG); sandy ground, near Granbury, Hood Co., 6 May
1900, Eggert (MBG); sandy ground, Big Spring, Howard Co., 11 June 1900, Eggert
(MBG); Round Top Mountain, Comanche Co., 9 May 1900, Eggert (MBG); near
Comanche, 10 May 1900, Eggert (G, MBG); San Saba River, Brady, 16 April 1926,
Studhalter 1106 (US); San Saba, 8 May 1917, E. J. Palmer 11843 (MBG); plains
west of Pecos, 20 April 1902, Tracy & Earle 106 (F, G, MBG, US); foothills, Davis
Mountains, 23 April 1902, Tracy & Earle 178 (F, G “MBG, NY, US); sandy ground,
near Bastrop, 17 April 1929, E. J. Palmer 33367 (MBG, NY); its hills, Austin, May
1872, Hall 428 (G), 431 (MBG, NY, P); Austin, Bray 98 (NY), Young (G,
MBG), Young 114 (P), E. J. Palmer 9389 (MBG), Tharp 1862, 1364 (US), Armer
6881 (US); gravel bars of Blanco River, Blanco, 5 April 1918, E. J. Palmer 13281
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 339
(MBG); Big Branch, Gillespie Co., Jermy 184 (MBG); New Braunfels, March 1850,
Lindheimer 434 (MBG), 1075 (F, G, MBG, NY, US); Lacey’s Ranch, Kerr Co., 1
June 1916, E. J. Palmer 10008 (MBG); Kerrville, 28 March 1916, E. J. Palmer 9278
(MBG); Hungerford, 4 March 1914, E. J. Palmer 4840 (MBG); Wharton, 18 March
1914, E. J. Palmer 4977 (MBG); Bexar Co., Jermy (MBG), 84, 85, (US); San Antonio,
March 1882, Havard (US); San Antonio, Jermy (NY), Jermy 209 (G), Wilkinson 8,
108 (MBG), Trelease 99 (MBG), Eggert (MBG), Bush 1185 (MBG), Clemens & Clem-
ens 976 (MBG, P); Rock Springs, 17 April 1930, Jones 26220, 26228 (P); gravel and
sand bars, small streams, Uvalde, 11 May 1918, E. J. Palmer 13562 (MBG); near Vic-
toria, 11 April 1900, Eggert (G, MBG); Riverside, Walker Co., 24 March 1918, E. J
Palmer 13175 (MBG); Brownsville, 23 Jan. 1919, Hanson (MBG); Brownsville, 30
Jan. 1919, Hanson 322 (G, NY, US); Brownsville, 14-15 March 1923, Tharp 1871
(US); southwest Texas, 1851-2, Wright 1500 (ANSP, F, G, MBG, NY).
New Mexico: without data, 1851-2, Wright 1501 (ANSP); Carlsbad Cavern, 5
May se pum 109 (US)
MExic
xod: Alamos, 26 March-8 April 1890, Palmer 326 (G); along an arroyo,
vicinity of Alamos, 13 March 1910, Rose, Standley & Russell 12745 in part (NY, US).
SiNALOA: San Blas, 2 Feb. 1927, Jones 23243 (MBG).
Verbena pumila has often been confused with V. quadrangulata
and, superficially, closely resembles it, but is readily separated
on the beakless nutlets.
49. V. setacea Perry,’ n. sp. Pl. 15.
Stems decumbent-ascending, soft-pubescent; leaves 3-5 cm.
long, with cuneate-truncate base narrowed into a margined
petiole, trifid with segments coarsely dentate, lateral lobes small,
veins somewhat prominent beneath, soft-villous-pubescent on
both surfaces; spikes short-peduncled; bracts linear-lanceolate,
about three-fourths as long as the calyx, subulate-setaceous,
soft-pubescent, ciliate; calyx 6 mm. long, villous-pubescent,
teeth (2 mm.) about half as long as the tube, subulate-setaceous;
corolla-tube 7 mm. long, pubescent without; corolla-limb about
6—7 mm. broad; anthers not glandular; mature nutlet not seen.
29 V, setacea Perry, spec. nov., herbacea vel basi suffruticosa; caulibus decum-
bentibus ramosis ramis spoon iode piloso-pubescentibus; iie is basi cuneata in
petiolum alatum attenuatis 3-5 cm. longis tri tatis lateralibus
parvis subtus venoso-reticulatis utrinque molliter villoso-pubescentibus; spicis
breviter pedunculatis; bracteis lineari-lanceolatis subulato-setaceis piloso-pubescent-
ibus ciliatis calyce paulo brevioribus; calyce 6 mm. longo villoso-pubescente, calycis
dentibus subulato-setaceis 2 mm. longis; corollae tubo 7 mm. longo extus pubescente;
limbo fere 6-7 mm. lato; connectivo antherarum superiorum inappendiculato;
fructu immaturo.—Collected at Calmalli, Lower California, Jan.-March 1898,
Purpus 195 (P), TYPE.
[Vor. 20
340 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Distribution: Lower California.
Specimens examined:
Mexico: Lower CALIFORNIA: rocks, Calmalli, Jan.-March 1898, Purpus 195
(P TYPE).
The gross habit of this plant is very like that of V. Gooddingii
var. nepetifolia. Unfortunately the inflorescence is immature and
only a few flowers are in anthesis, hence it is rather difficult to
say what are the characters of the spike or of the mature nutlets.
Moreover, the corolla may be larger than appears in this speci-
men. ‘The flower itself is similar to that of V. lilacina, but the
two plants are so different in habit it would seem as if this were
perhaps only a superficial resemblance. The species is readily
distinguished by its general habit and the long setaceous calyx-
teeth
50. V. lilacina Greene, Bull. Calif. Acad. Sci. 1: 210. 1885.
Stems erect, much branched, 0.5-1 m. high, very sparsely
hirsute or glabrous except just below the spike; leaves 3-5 cm.
long, contracted at base into a margined petiole, bipinnatifid
(upper pinnatifid), divisions remote, ultimate lobes chiefly
linear and acute, somewhat scabrous and strigillose on both
surfaces, rugose above, midrib prominent beneath; spikes fascicle-
like, cymosely arranged, long-pedunculate, bracts somewhat
shorter than the calyx, lanceolate-setaceous, pubescent, ciliate;
calyx about 7 mm. long, appressed-pubescent, short-hirsute
along the nerves, lobes unusually long (2.5-3 mm.), attenuate
into subulate-setaceous teeth; corolla-tube protruding very little
beyond the calyx; corolla-limb 10 mm. broad, segments emargin-
ate; anthers not glandular; nutlets almost smooth, slightly
enlarged at the base; commissural face muricate-scabrous, not
reaching the tip of the nutlet.
Distribution: Known only from Cedros Island.
Specimens examined:
XICO:
Lower CALIFORNIA: Cedros Island: 29 April 1885, Greene (G); 1889, Palmer (G),
677 (F, NY); March-June 1897, Anthony 288 (F, G, MBG, US); 12 March 1911,
Rose 16155 (NY, US).
This is an anomalous species of uncertain relationship. It has
an erect open habit with more or less glabrous stem and long
internodes somewhat suggesting V. neomexicana (§ Verbenaca);
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 341
nevertheless, the short stout dense spike does not point to an
affinity with this section. Although the anthers are not glandu-
lar, the sterile style-lobe protrudes well beyond the stigmatic
surface, and, in mature fruit, the style appears to have been
inserted in a deep depression at the apex of the schizocarp;
hence the species is regarded provisionally as a member of the
section Glandularia.
51. V. amoena Paxton, Mag. Bot. 7: 3, pl. 1840.
V. grandiflora Sesse & Mocino, Pl. N. Hispan. ed. 1, 6. 1887-90
[La Naturaleza, II. 1. App.].
Stem about 5 dm. tall, decumbent or ascending, retrorsely
hispidulous; leaves 5-8 cm. long, bipinnatifid, with the lower
part entire, forming a broadly margined subauriculate and semi-
amplexicaul base, divisions remote, linear-oblong, sparsely
incised, appressed-pubescent above, hispidulous beneath with
midrib and veins prominent; internodes 4-6 cm. long; spike
terminal, fascicle-like in anthesis, pedunculate; bracts linear-
lanceolate, subulate, pubescent; calyx about 10 mm. long,
densely pubescent, particularly along the nerves, glandular,
teeth slender, subulate; corolla-tube slightly longer than the
calyx, pubescent without, particularly around the throat; corolla-
limb about 9 mm. broad; anthers unappendaged; mature fruit
not seen.
Distribution: Mexico.
Specimens examined:
Mexico:
MicHoacan: Puruandiro, Sesse & Mocino 99 (Herb. Bot. Gard. Madrid TYPE,
MBG phot.).
Mexico: hills, Lecheria, 5 July 1904, Pringle 13434 (US).
This unique species is readily recognized by its coarse habit,
together with its bipinnatifid, subauriculate and semiamplexicaul
leaves. Although Paxton’s description is not accurate, the
specimen agrees well with his plate and with the photograph of
V. grandiflora.
DOUBTFUL OR LITTLE-KNOWN SPECIES
‘í VERBENA ERINOIDES Lam." This species has established itself
in several places. It belongs to a South American species-
[Vor. 20
342 ANNALS OF THE MISSOURI BOTANICAL GARDEN
complex needing critical study to determine accurately its real
identity.
V. GRANDIFLORA Ortega, Hort. Matr. Dec. 2. 1797. Although
the description is inadequate for positive identification, the
phrase ‘‘Semina saepius duo" would seem to indicate that the
species does not belong to the genus Verbena.
V. nEPENS Spreng. Erst. Nachtr. d. Beschrieb. d. Bot. Gart.
Univ. Halle, 40, no. 51. 1801. The literature on this species is
very vague. The plant described is said to be a native of Santo
Domingo. According to Lamarck,? the vervain of Santo
Domingo is a species of heliotrope.
V. TRIFIDA HBK. Nov. Gen. et Sp. 2: 273, pl. 134. 1818.
This is a true Verbena, but none of the specimens at hand agrees
with the description; and the floral drawings are insufficient to
reveal its affinities.
V. BARBATA Grah. Edinb. New Phil. Jour. 176. 1827. This
is undoubtedly a member of the section Verbenaca, but the de-
scription is too meagre to identify it.
V. INCARNATA Raf. Atl. Jour. 154. 1832. Here again the
description is too inadequate for specific identification.
V. DELICATULA Mart. & Zucc. in Otto & Dietr. Allg. Garten-
zeit. 2: 245. 1834. Unfortunately the essential characters,
which would separate this species from its allies, are not clearly
defined. The description is so much like that of V. barbata that
the writer suspects that these two species are identical.
V. MamruEsrn Turez. Bull. Soc. Nat. Mosc. 36?: 196. 1863.
In the group to which this species evidently belongs, the units
are very closely related; hence without more specifie definition
the species is obscure.
V. PAUCIFOLIA Turez. Bull. Soc. Nat. Mosc. 36?: 196. 1863.
This may be V. carolina, although the leaves of the latter are
somewhat broader; but until such time as the type may be
studied it seems preferable to withhold any decision regarding
it.
V. INTEGRIFOLIA Sesse & Mocino, Pl. Nov. Hispan. 6. 1887
[La Naturaleza, II. 1. App.]. No known Mexican species of
Verbena has entire leaves.
30 Lamarck, Encyc. 1, Suppl. 5: 469. 1817.
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA
V. SCABRELLA Sesse & Mocino, l. c.
343
Apparently this species
does not belong to the genus, but anything further regarding
its identity is unknown to the writer.
V.
12: 12
synonymy under V. canadensis.
AUBLETIA var. LAMBERTI M. E. Jones, Contr. West. Bot.
1908. This combination, nomenclatorially, belongs in
Its entity is clearly with the
section Glandularia, but with which species it should be associated
is unknown.
LIST OF EXSICCATAE
The collectors’ numbers are printed in italics.
Unnumbered collections are in-
dicated by a dash. The number in parenthesis is the species number used in this
revision.
Abrams, L. R. — (22, 31); 2481 (22);
2574 (31); 3406 (7); 3787 (22); 9487
(31).
Abrams, L. R. & McGregor, E. A. &
(22
Adole, Bro. 22 (29).
Allard, H. A. 20? (31).
Anderson, I. W. — (16).
Anderson, J. P. — (15, 16).
Anderson E. S. & Woodson, R. E., Jr.
4? (17); 1577? (32).
Andrews, L. — (16).
Anect, Bro. 5? (20); 100, 143, 179 (41);
211 (20).
Anthony, A. W. 288 (50); 380 (4).
Antisell, T. 186 (41a).
Applegate, E. I. 2228 (22).
Armer, * A. 5381 (48); 5385 (8).
Arséne — (1, 5, 11, 46); 672 (16);
2798 b ; 8000 (11); 6129 (34); 9998
(7); 10626 (29); 11820 (32); 11831
(8); 11859 (2); 11982 (35); 12117 (32);
12242 (8); 12534 (2); 18399, 18407,
18543, 18593 (31); 18615 (41); 18684
(31); 18793 (41); 18885, 18961, 18964
(31).
Arsène & Agniel, 10242 (7); 10252 (43).
Arséne & Benedict, 15734 (31); 16306
(41); 16600, 16790 (31)
Arthur, J. C. 24 (17).
Ashe, W. W. — (17).
Bailey, V. 221, 258 (43a).
Bain, S. M. 328 P 444 (16).
Baker, C. F. , 16, 17); 288, 564
(31); 565 (20): (31); 2591 (6).
Baker, C. F., Earle, F. S. & Tracy, S. M.
531 (42 `
Ball, C. ; $44 (8); 401 (40);
553 n. 556 Eo. 605 (24); 909 (40);
1171 (25); 1585 (19); 1675 (31).
Ball, J. — (31).
Ballard, C. A. — (16).
Barber, H. S. 106 (43).
Barkelew, F. E. 231 (4).
Bartlett, A. H. 1122 (1).
artram, E. B. — (7); 255 (47); 256
(25); 257 (478); 1 1021 (17).
Basile, b 99 (7).
Bates, — "o 19, 40).
Beard,
Beattie, F br — (us 16).
Beaumont —
Beckwith, F. 48 (19); 138 (20); 772 (38).
Benke, H. C. 4566 (35); 5046 (43a); 5164
19).
Berg, H. K. — (14).
Bergmann, L. S. — (31); 2385 (16);
2370 (31).
Berkley, E. E. 1304 (16).
Berlandier, J. 322 ES 429 —?1749 (40);
344
578 (28); 644 (25, 29a); 827 (29a);
1222 (12); 1449=189 (40); 1485 (=
225) in part (25, 33); 1511 (8); 1520
(43a); 1592 (8); 2054 (29a); 2428 =
998 (40); 2506 in part (25); 3016 (8);
$018 — 1518 (33); 3020 (43a).
Bernoulli, G. 127 v as (5).
Bessey, C. E. — (15
Bigelow, J. M. Mer 2 31, 35, 42).
Billings, F. H. 49 (1).
Biltmore Herbarium, /082b (31); 8653a
(19); 4759, 4759b (17); 4761a (32);
4762 (6).
Bissell, C. H. — (16).
Blakeley, O. W. 1462 (30; 1471 (35).
Blanchard, F. — (15, 17).
Blanchard, W. H. 26, ^ 161 (17).
Blankinship, J. W. — (15, 17, 19, 22,
35).
Blanton, F. S. 6598 (3).
Blewitt, A. E. 14 (19).
Blumer, J. C. 1345 (40); 1612 (28);
1783 (11); 1804, 2170 (26a).
Boettcher, F. L. J. 176, 226 (15).
Bogusch, E. R. 1236 (24).
Bolander, H. N. ee (31).
Botteri, M. 180 (11).
Bourgeau, E. — (16); 119 (11); 120 (43);
860 (7); 861 (28); 547 (7).
Boyce, H. T. E. — (1
— ki 8. — (11,
B. 466
Brannon, i 0.1
Braunton, E. 99 ain 328 (22); 1263
(23).
16, 20, 41);
Bray, W. L. 65 (35); 98 ay ET (292).
Brewer, W. H. 31 a 229 (22).
Brinton, J. B. — (17
Britton, N. L. — LM 220, 296 (37);
1627 (14); 3176 (1).
Britton, N. L. & Brown, M. S. 28 (6);
98 (2); 158 (1); 878, 1681 (14).
Britton, N. L. & Cowell, 10326 (6).
i . & Hollick, 2704 (14).
. & Wilson, P. 5771 (14).
Britton, N. L., Britton, E. G. & Brown,
M. S. 6019, 7037 (14).
Britton, N. L., Britton, E. G. & Shafer,
J. A. 292 (14)
[Vor. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Britton, N. L., Brown, M. S. & Wortley,
1645 (1).
Britton, N. L., Wilson, P. & Leon, Bro.
15256 (14).
Broadhead, G. C. — (15, 35).
Brown, H. E. 58 (3 E
Brown, 8. 492 (6); 693 (1)
Brown & Brit qiio Britton & Brown.
Bryant, M. L. 88 (22)
Bucholz, A. F. — (15).
Buckley, S. B. — (8, 31, 32).
Burgess, A. B. 638, 713, 783 (37).
Burk, M. 542a (31); 591 (15).
Burnham, 8. H. — (15, 17).
Bush, B. F. — (15, 17, 35); 28, 77, 84 (8);
205 (40); 206 (17); 283 (35); 312 (32);
318 (35); 851 (40); 432 (15); 433 (17);
435 (19); 436 (31); 437 (35); 449 (40);
450 (17); 452 (15); 475 (19); 545 (35);
569 (35); 570 (19); 601 (31); 884 (15);
$99 (24); 914 (17); 1034 (31); 1168
(35); 1185 (48); 1275 (coll. of 1900)
(24); 1275 (coll. of 1895) (48); 1482
(15); 1933 (35); 2109 (31); 2214 (16);
2901 (35); 4010 (31); 4029 (40); 4922,
6439 (35); 7586a, 7647 (17); 8537 (31);
9165 (16).
Butler, G. D. 1621 (22).
Calderon, S. 729 D HM (11); 925 (5).
Campbell, J. E. — (16).
Campbell, J. R. 67 TA
Canby, W. M. — (32); 193 (33); 194
(29a).
Carleton, M. A. —, 177 (40).
Carleton, M. L. 134 (48).
Carlson, J. I. — (7, 23, 26a, 41a, 47).
Carr, W. P. 118 wt 179 (81).
Carter, W. R. — (16).
Olubealt. E. : 418 (31).
oe E. B. & Knowlton, F. H.
16).
Chamberlain, F. M. 56 A
Chandler, H. P. — (8, 16).
Chandonnet, Z. L. — (15, 16, 19, 31).
Chapline, W. R. 609 (26
Chapman, A. W. — (6, 14, 15, 16, 31,
Chass, A — (15).
Cheney, C. I. — (16).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA
Chestnut, V. K. — (22
Choussy, F. 36 (11).
Churchill, J. R. — (31).
Clark, J. A. 55 (31); 253 (16); vila (31).
Clark, O. M. 4041 (29a); 4250 (27).
Clawson, A. B. 13923 (42
Clemens, Mrs. J. — (22, 31); 11748 (19);
11749 (15); 11750a (31); 11761 (40);
11752 (48).
Clemens, Mr. & Mrs. J. 969, 970 (8);
971, 972, 978, 974 (29a); 976 (48).
Clements, F. 2612 (31); 2776 (40).
Clements, F. E. & E. S. 9 (31
Clifton, R. L. 3085 (40).
Clute, W. N. 93 (31).
Clokey, I. W. — (36); 1 11).
Collins, F. S. 45 (1); 53, pel EE 267 (6);
268 (14); 269 (1).
Congdon, J. W. — (23).
Conzatti, C. 4194 (29); 4207 (7); 4288
9
).
O. F. & Griggs, 148 (5).
Cooper, J. G. — (47).
Cooper, J. J. 5890 (5).
Cory, V. L. 664 (44).
Cotton, J. s. 621, 1363 (31); 1896 (16).
Coues, E. & Palmer, Ed. 279 (16); 498
(47); 671 (26a).
Coulter, — (43).
Coville, F. V. & BE F. 968 (31).
wen, J. H.
Crandall, C. S. 25 (16).
Croft, M. B. 78 (33
Culbertson, n W. 4810 (22)
Curtis, C. D. — (15
Curtis, M. A. — (6).
Curtiss, A — (6, 17, 32); 677 (6);
i (17); 1959 (32); 1963 in part (2,
38); 4386, 4765 (32); 6111 (14);
Bn (37); 649 E A
Cusick, W. C. 1 31).
Cuthbert, A. i. i 268 (32); 858 (2).
Davidson, A. — (1
Davis, J. 16 (19); 73, 950 (15); 1202 (19);
127 4a (15); 1613 (19); 2414, 2691 (31);
345
$791, 2960, 2962 (19); 3192 (16);
$227 (19); 3453 (15); 3565, 8578 (16);
$587 (31); 3588 (17); 9589 (19); 3590
(16); 3591 (15); 3817 (16); 3852, 4450
(15); 4562 (17); 4461, 4487 (19); 4565
(15); 6870 (19); 8458 (15); 9039 (31);
Day, M. A. 79, 147 (15); 149 (16).
Deam, C. C. — (19); 86 (43); 1272 (31);
6180 (5); 13619, 20857 (17); 20857a
(19).
Deam, Mrs. C. C. 1799 (32); 1831 (35).
Dean & Thomas, 4874 (16).
Deane, W. — (15).
Dehesa, M. P. 1561 (11).
Demaree, D. 3353, 3371 S vae (19);
6411 (35); 7589 (25); 7685 (48a).
Dewart, F. W. 15, 18 Wen d (17); ?4
(35).
Deweys, Mrs. F. E. — (8).
Dewhurst, B. L. — (31).
Dixon, R. A. 473 (24); 561 (40).
Dodge, C. K. 154 (33).
Donnelly, L. L. — (40a).
Drummond, T. — (14, 24, 35); 253 bis
(24).
Drushel, J. A. — (32); 2762 (35); 2842
(24); 3659 (35; hi D; 6351 (16).
6, 496a (43)
E (id.
H. — (19); 8137 (31).
Eames, E. H. & McDaniels, L. H. 4870
(15).
Earle, F. S. 84 (40); 387 (27); 427, 606,
(25); 619 (27); 638 (41).
Earle, F. S. & Baker, C. F. — (31, 32).
Earle, F. S. & E. S. 236 (41); 361 (31);
387 (27); 526 (40).
Earle, F. S. & paar — (32).
Eastw : 3, 41); 25 (31); 41
(16); ps (19); s (23); 8468 (42).
Eaton, D. C. — (15, 16, 17)
Edwards, — (20).
Eggers, H. F. A. von, 1828 (6).
Eggert, H. — (6, 8, 15, 16, 17, 19, 24,
346
25, 29a, 31, 32, 33, 35, 40, 41, 42, 43a,
44, 48).
Eggleston, W. W. — (17); 1581 (15);
4430 (17); 4837 (19); 4841 (15); 5237
(17); 6644 (41); 12240 (35); 15224
(19); 15551 (16); 17106 (20); 1?231
(42).
Ehlers, J. H. 627 (31); 642 (19).
Ehrenberg, C. 131 (46); 713 (10).
Ellis, C. C. 17 (42); 221 (31); 258 (20).
Elmer, A. D. E. —, 324 (31); 537 (16);
3846, 4045 (22); 4950 (23).
Emig, W. H. 107, 365 (19); 395 (40); 401
(48); 415 (35); 716 (15); 787 (19); 788
(40
Greeters, G. — (16, 17, 19, 23, 31, 35,
41); 334 (35); wer an; 337 (16).
Engelmann, H. —
Epling, C. C. 5445 - 6134 (35).
Ervendberg, L. C. 153 (10); 236 (34).
Evans, W. H. — (43b).
Evermann, B. W. 970 (19).
Eyerdam, W. J. 201, 432 (14).
Ezell, A. 5699 (8).
Fassett, N. C. & Hotchkiss, N. 3322 (16).
Fawcett, H. S. /2 (31).
Faxon, C. E. — (16
Fendler, A. — (31); 586 iu 587 (31);
594 (25); 597 in part (19, 20).
Fernald, M. L. — (16, 31); 296 (16).
Fernald, M. L., Hunnewell, F. W.
Long, B. 10262 (15).
Fernald, M. L. & Long, B. 10268 (16).
Fernald, M. L. & Parlin, J. E. 928 (15).
Fernald, M. L. & Pease, A. 8. 25247 (16).
Fernow, B. E. — (47a).
Ferris, R. S. & Duncan, C. D. 2474 (41);
2607 (20b); 2647 (42); 2786 (25);
3161 (34); 3268 (24); 3337 (8); 8518
(16)
Fink, B. — (31); 251 (19).
Fisher, G. L. — (11, 16, 35, 41, 46);
19 (19); 33, 40, 47 (35); 122 (24); 168
(5); 207 (43); 212 (24); 223 (43);
235 (29a); 282 (25); 319 (43); 820 (19);
33 7 (46); 460, 625 (1); 707 (35).
— (35).
Forrer, A. — (30, 40a).
[Vor. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Forwood, W. H. 298 (31); 299 (19).
Fowler, J. — (16).
Fredholm, A. 110, 5136 (32); 6416 (14);
5804 (38).
Freiburg, G. W. — (15, 19).
Fritchey, J. A. 29 (22).
Frost, W. D. — (19)
Fuertes, M. 391, 1758 (14); 1771, 1856
(6).
Furness, D. R. —
Galeotti, H. 736 (11); 736 (43); 737 (11);
777 (46); 778 (T); 781 (6); 795A (11).
Gallegos, J. M. 2342 (18
Gandoger, M. — (36)
Garber, A. P. — (38).
Garcia, P. I. 331 (40a).
Gardner, J. R. 553 (19).
Garesche, E. E. — (19).
Gates, F. C. 9994 (19).
Gates, F. C. & M. T. 10708 (16).
Gattinger, A. — (19).
Gershoy, A. 583 (6).
Geyer, C. A. — (16, 40).
Gibbes, L. R. — (2, 15).
Gilman, M. F. 1120 (26a).
Glatfelter, N. M. — (8, 15, 17).
Gleason, H. A. — (19, 31); 377 (19);
718 (15); 1945 (16).
Goldman, E. A. 42 (34); 292 (47a).
Goldsmith, G. ^d 9 (15
Goodding, L. N. 37? (402); ; 207 (81);
251 (40); 334 (26a); 495 (40); 645 (47);
883 egy 952 (47a); oi 2312 (31).
Goodm oJ. iteheock, C. L
1211 (40); 1297 w^ Pe (31).
phe ee LT A. 90 (16).
Graham
Grant, M. L. pa 73 (16).
Graves, E. W. 525 (1); 536 (40); 621 (82);
1691 (15); 1946 (40); 1947, 1994 (19).
Green, T. — (17).
Greene, E. L. — (23, 26, 42, 43b, 50);
45 (41); 77 (20); e e `
Greenman, 44, 245 (15);
398, 1273 (16); 1376 y 1877, 1379
(16); 1380 (15); 1382 (16); 1877 (15);
1953 (16); 1981, 3640 (17); 3766 (15);
3870, 4076 (35); 4125 (17); 4231 (35);
4412 (15); 4578 (17
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 347
Greenman, J. M. & Greenman, M. T.
28 (25); 56 (47a).
Gregg, J. — (8, 29, 43); 11 (7); 26, 57
(43); 99 (29); 202 (34); 246 (29); 257
(43); 265, 276 (7); 355 (86a); 406
(7); 483 (31); 545 (29); 623 (31); 632
(43); 686 (31); 648 (43); 740 (29); 752
(34); 756 (43); 764 (5); 791 (29a);
823 (7); 1001 (11)
Griffith, F. 3456 (40).
Griffiths, D. 1595 (25); 1919 (41a); 2421
(47a); 8431 (26a); 3489 (47a); 4244
(41); 4757 (43) 4833 (40a); 5050 (19);
6190 (42); 5560 (31
Griffiths & Hunter, 18 (31).
Griffiths, D. & Thornber, J. J. 159
(40a).
Grimes, E. M. 3716, 8788 (17); 3794
(15).
Groth, B. H. A. 16 (29a); 80 (40); 75
(8); 187 (24).
Guidroz, H. 3 (15).
Gurney, J. & Monell, J. T. — (40).
Hale, T. J. — (8, 15, 17, 19, 24, 35);
245 (8)
Hall, E. — (19, 31); 393 (31); 428 (48);
429 (25); 430 (29a); 431 (48); 432 (8);
438 (1); 484 (24); 435 (35).
Halsted, G. 176 (15).
Hammond, E. W. 322 (22).
Hansen, G. 964, 1823 (22); 2026 (5).
Hanson, H. C. — (48); A146 (43b);
A147 (31); A148 (20); 322 (48); 551
(42); 619 v pud (26b).
Hanson, H. C . E. A1021 (47);
A1130 (26a).
Harger, E. B. — (16, 17).
Harper, R. M. 16 (40); 242 (14); 385
(17).
Harris, J. A. C16375 (47a).
Harris, W. 9132 (2); 9937, 11808 (14);
Harrison, G. J. 4778 (26a); 4897 (14).
Harrison, G. J. & Kearney, T. H. 3646
(47a); 5796 (26a); 6144 (11); 6689
a).
Harshberger, J. W. — (1, 2); 41 (46).
Hartman, C. V. 94 (11); 608 (26a); 880,
906 (40a).
Hartweg, T. 174 = 175 (7); 176 (43);
177 (29); 1924 (22).
Harvey, F. L. " n 62 (31); 1958 (19).
Harwood, R. D. — (
Havard, V. — jp 27, 31, 33, 42, 44, 48);
97 (42); 197 (27).
Hayden, F. V. — (15, 16, 40); 10 (31).
Haynes, D. — (22).
Hedgcock, G. G. — (15, 16).
Heller, A. A. — (15, 31); 638 (19); 1388
(33); 1419 (8); 1732 (29a); 5785, 5919,
6778 (22); 14222 (16); 14290 (19);
14303 (31
Heller, A. A. & E. G. 3536 (42).
Henderson, L. F. 4070 (31).
Herrick, C. L. 704 (42); 715 (26b).
Hertel, H. — (15).
Hexamer, A. C. & Maier, F. W. — (16,
17, 35).
Heyde, E. T. 120, 477, 580 (11); 610 (5).
Heyde, E. T. & Lux, E. 3018 (11); 3019
in part (5, 11); 4370 (5).
Hicks — (15).
High, M. M. 52 vn 59 (48a); 76 (33).
Hilgard, E. — (8, 32).
Hillman, F. H. — (31
Hitchcock, A. S. — (2, 14, 15, 16, 17,
32, 35); 269 (14); 393 (40); 790 (35);
791 (17); 972 E 1129 (31).
Hoffstetter, G. — (15).
Holgate, W. R. — ie
Holm, T. — (15).
Holzinger, I. M. — (19, 26, 41a).
Hooker, L. H. 5999 (34).
Hough, W. 109 (43).
Houghton, H. W. 2570 (48); 3622 (17);
3643 (35); 3646 (17); 4000 (19); 4015
(15
Hosen, H. D. 875, 1363 (17); 8276 (32).
Howell, A. H. 64 (19); 111, 308 (40);
313, 362 (8).
Howell, T. 174 4 (22); = (16); 1249 (22).
Hoyeradt,
Humboldt, A. p Bonpland, A. — (11);
4063 (36); 4066 (12
Hyams, M. E. — (32
Jaeger, E. C. — (47).
Jermy, G. — (48); 84, 85 (48); 182 (29a);
183 (40); 184 (48); 203 (8); 209 (48).
348
Jesup, H. G. — (16).
Johnson, A. G. — (15, 16, 19, 35).
Johnston, E. L. 507 (16); 986 (41)
Johnston, I. M. — (14); 1320, 1608 (22).
Jones, M. E. — (7, 8, 11, 16, 19, 20, 22,
23, 25, 20a, 40, 40a, 42, 43b, 47, 47a);
36 (47a); 122 (41); 372 (25); 457 (42);
487, 523 (16); 554 (29); 634 (41);
686 (43); 1487 (16); 2598 (22); 3801
(43b); 3901 (47a); 5478 (31); 6026 (20);
8963 (16); 23243 (48); 23244 (43);
25006 (11); 25480 (43b); 26220 (48);
26224, in part (26b); 26225, 26226
(29a); 26227 (43b); 26228 in part (40,
48); 26229 in part (8, 40); 26231 (43);
26232 (43
Jones, b Pd esu 180 (43b); 185
6a).
Katzenstein, O. — (32).
Keeler, H. D. — (14).
Kellerman, W. A. — (17); 5825 (11).
Kellogg, J. H. — (15, 16, 35); 496 (17);
498 (19); 1130 (17); 1131 (10); 1958
(35); 15276 (31); 15277 (15).
Kellogg, W. — (38
Kelsey, J. A. 173 (10).
Kempton & Collins — (11).
Kendall, M. L. — (22).
a G. G. — (15
nedy, P. B. 7032 ( on; 7033 (47a).
Kerber, E. 255 (43); 311 (5).
Killiam, O. L. 6933 (15).
Killip, E. P. 3510 (5).
z
=
S
=
Ez
E
A
w
Knowlton, C. H. — - (16).
Korthoff, I. — (3).
Kreager, F. O. 469 (16); 474 (31); 475
19).
(
Krig, J. T. 3187 (14).
Kuntze, O. 424 (43); 2109 (5); 23444
(28); 23738 sey 23811 (24).
Lane, W. C. — 6).
Lang, H. A. x 4).
Langlois, A. ^i — (1, 24, 40); 123 (24).
[Vor. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Lansing, O. E. — (16); 2805 (16); 2806
(15); 2810 (19); 2976, 3135 (17); 3164,
(19); 3241, 3340 (16)
Laybourne, W. A. — (15).
Lea, M. C. — (6).
Lee, D. W. 109 (48).
Lehmann, F. C. 1603 (46).
ond J. B. 902 (31); 2630 (16).
, J. G. — (28, 47, 47a); 2857
(26a); 3075 (11).
Leon, Frére — (6); 685 (14); 3910 (1).
Leon, Frére & Edmunds, Frére — (6);
8719 (14).
Leonard, E. C. 3939 (6).
Leonard, E. C. & Killip, E. P. 534 (17).
Leroy & Ruger — (31).
Letterman, G. W. — (8, 15, 17, 19, 24,
, 40).
Lewton, F. L. 141 (34).
Liebmann, F. M. 1/1308 (46); 11318,
11314 (34); 11316 (46); 11318 (13);
11328, 11329, 11330 (46); 11335 (10).
Lighthipe, L. H. 599 (35).
Lindheimer, F. — (8, 14, 16, 24, 35);
145 (24); 146 (40); 154 (24); 155 (8);
195 (15); 232, 289 (40); 294 (29a);
307 (40); 434 (48); 500 (292); 501
(48); 618 (14); 1072, 1073 (40); 1074
(29a); 1075 (48); 1076 (8); 1077 (14).
Lloyd, C. E. 448 (11).
Lloyd, F. E. 159 (43).
Lloyd, F. E. & Tracy, S. M. — (6); 20
(3); 22 (14).
Loomis, H. F. & Thackery, F. A. 910
(47a).
Louis-Marie, Pére, 141 (16).
Lumholtz, C. 445, 446 (36a).
Lunell, J. — (16, 31).
Lyall, D. — (31)
Lyon, 6 (1
Lyonnet, E. — (7); 173 (30).
Macbride, J. F. 200 (31); 304 (16).
Macbride, J. F. & Payson, E. B. 781
(7); 850 (23).
MacDaniels, L. H. 4873 (16).
MacDougal, D. T. 73 (47a); 153 (7);
249 (20); 286 (31); 317 (40); 566 (16);
608
MacElwee, A. 760 (17); 873 (6); 882 (15).
1933]
PERRY— NORTH AMERICAN SPECIES OF VERBENA
M T K. K. 96 (8); 261 (19); 4218
(17); 4746 (15).
del C. — (19).
Macoun, J. — (16, 31); 1305 (17);
24268 (31
McAtee, W. L. 1953 (8); 2053 (19).
McClatchie, A. J. — (14).
McFarland, F. T. 92 (15); 109 (17).
McFarland, F. T. & Anderson, W. 253
(19).
McKelvey, S. D. 1710 (29a); 1756 (25);
2046 (26b).
Manning, W. H. — (19, 31).
Marble, D. W. 737 (2).
Marsh, C. D. 14224 (42).
Mathias, M. E. 545 (41)
Mattern, E. S. & W. I. — (38).
Maxon, W. R. 3808 (40).
Maxon, W. R. & Harvey, F. L. 7951 (5).
Maxon, W. R., Harvey, & Valen-
nh 7356, 7478 (5).
.R.& me E. P. 569 (2).
Sera, E. A. — (40a, 47); 108 (43);
524 (31); 1013 (26a); 1116 (28); 1138
(41a); 1237 (29a); 1252, 1274, 1894
(40); 1456 (29a); 1887 (26a); 1904,
1914 (47a); 1918 (26a); 2278, 2486
o>) 2627 (11); 2677 (41); 2698,
9 (31).
Fa C. D. — (38).
Mell, C. D. & Knopf, E. C. — (81).
Mellichamp, K. — (32).
Meredith, H. B. — (6).
Merrill, E. D. phy ox E. N. 776 (31).
Merton, E. C. 2042 (26:
Metcalfe, O. B. 126 42); 187 (81);
612 (26); 897 (31); 955 pud 1008 (31);
1090 (42); 1231 (41); 1 (27).
Michener, C. A. & iden i5 T. 123
(23).
Millspaugh, C. F. 866 (17).
Mohr, C. — B 17, 24, 31,
32).
Molby, E. 7224 (34).
Moldenke, H. N. 148 (2); 212 (38);
219a, 549, 586 (37); 591, 599a, 972
(14); 1039 (38); 1091 (32); 1184 (3);
1330 (15); 1339 (16).
Moore, A. H. 2569 (16); 2874 (14);
349
2939a (6); 2946 (2); 2947 (14); 2984
; 4816 (16).
Moore, J. A. & a J. A. 3001
(40); 3005 (29a); 3009 (41); 3277
(26b); 3299 (25); 3324 (42); 3611
(27); 3622 (41).
Morales, R. 786 (11).
Moron
E+ a 7).
Mueller, F. — (11); 325 (43); 766, 887
(11); 1209 (43); 1215 (29); 1319 (43).
Mulford, A. I. — (31); 37 (42); 39 (20).
Munz, P. A. 616 (15); 617 (16); 1149
(26a); 1231 (41a); 1249 (43); 1470,
607 (24); 2207, 6349, 6598 (31);
m 9295 (22).
z, P. A. rodar. I. M. 11289 (14).
"unm P. A., Johnston, I. M. & Harwood,
R. D. 4252 (47).
Murdoch, J. — (40a).
cos G. V. 601 (32); 1248 (14); 2470
Nea I G. C. — (33); 117, 118 (84).
Nelson, A. 320 (31); 505, 538 (19); 1652
(31); 2258 (16); 7671 (31); 8245 (40);
8354 (20); 8489 (16).
Nelson, E. . 196, 223, 242 (46);
1021 (11); 1105 (12); 1518, 1895 (45);
1943 (7); 3896 (36a); 3920 (43); 4424
(34); 4559 (48); 4577 (7); 4593 (28);
4604 (43); 4856 (40a); 6117 (40);
6161, 6271 (26a); 6628 (34); 6789
(41); 6858 (43).
Nelson, E. W. & Goldman, E. A. 7123a
(47a); 7425 (21).
Nelson, J. C. 1429 (22); 1804 (16); 2669
(2 zn
ess, H. — (24).
dee cw — (11).
Nichols, G. E. 163 (2).
Nicolas, — (29).
Nicollet's Northwest Expedition, —
(40
idrico à A. 2685 (17).
Nor — (6, 15, 17); 389 (15);
pe Ne: E (19); 592 (31).
350)
Nuttall, T. — (17, 40).
Oakes, W. —
Oersted, A. 8. 11084, 11324 (5).
Olney, S. T. —
O'Neill, H. — "s 986 ha 6309 (37).
Orcutt, C. R. — (18, 31, 33, 47a); 909
(18); 1228 (31); - de 1302 (14);
1303 (472); 1546, 1549 (31); 2736 (2);
3321 (13); 3488 (11); 3551 (46); 3950
(12); 4045 (11); 4146 (46a); 5141
(47a); 5423 (10); 5542 (33); 5556 (25);
5657 (43a); 5717 (25); 5780 (33);
5792 (43a); 5867 (8); 6096 (41); 6111
(292); en (43b); 6184 (44); 6197
(41); 6235 (25).
Ortega, J. G. p M; 4541 (47a).
Osterhout, G. E. (15, 16, 20, 41);
$8166 (40).
Over, W. H. 2103 (40); 8177 (40); 14386
(19).
eo L. O. — (15, 17).
Pace, L. 22 (8).
Pagel, L. E. 2208 (33).
Palmer, Edward — (17, 26a, 41, 47, 50);
25 (29); 85 (43); 39 (34); 47 (40); 51
(36a); 62 (26a); 74 (43); 78 (8); 79
(43); 82 (29a); 90 (34); 135 (28);
141 in part (5); 141 in part, 158, 191
(coll. of 1898) (7); 191 (coll. of 1906)
(36a); 200 (28); 268 (7); 281 (43);
295 (36a); S03 (29); 307 (47a); 308
(7); 809 (22); 310 (23); 312 (26a); 326
(48); 339 (coll. of 1876) (47); 339
(coll. of 1896) (11); 33914 (26a);
1g (23); 342 (22); 345 (43); 356
(7); 864 (11); 397 (14); 401 (43);
456 (28); 6?7 (50); 911 (28); 1040
(14); 1042 (7); 1043 (8); 1044 (29a);
1046 (33); 1047 (29); 1048 (31); 1050
(41); 1051 (34); 1052 (36a); 1156 (11);
2014a, 2019 (5); 2037 (10); 2038 (25);
2039 (33); 2040 (25); 2055 (43); 2057
(11); 2529a (22); 2699 (31); 6433 (38).
Palmer, E. J. 197 (16); 198 (15, 16);
199 (19); 200 (17); 304 (35); 454 (31);
559 (35); 1085 (16); 1747 (33); 2980
(19); 2981 (17); 2996 (35); 8319 (17);
4229 (15); 4232 (17); 4840, 4977 (48);
6044 (8); 5185, 5407, 5682 (35);
[Vor. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
5769 (17); 6106 (31); 610? (19); 6510
(40); 7001, 7071, 7124 (35); 7369 (8);
7566 (24); 7821 (40); 8000 (15); 8045,
9098 (40); 9111 (8); 9183 (29a); 9278,
9339 (48); 9485 (8); 9559 (32); 10002
(29a); 10003 (48); 10037 (8); 10568
(16); 10692 (14); 11715 (8); 11843
(48); 11979 (35); 12001, 12038 (1);
12056 (35); 12496a (31); 13176,
18281 (48); 13282 (25); 18283 (29a);
18512 (39); 13562 (48); 13563 (29a);
18664 (8); 13730 (25); 13861 (31);
14988 (35); 15262 (17); 24616 (35);
27209 (40); 27462 (14); 28738 (16);
29303 (2); 29333 (35); 29535 (29a);
29804 (35); 30523 (25); 30672 (42);
30791 (26b); 31117 (31); 81158 (41a);
31192 (41); 33319 (35); 33367 (48);
33389 (40); 33537 (25); 33583 (42);
33592 (33); 33604 (8); 33605, 33646
(29a); 34065 (26b); 34240 (41); 34766
(17); 355235 (32); 37627 (19).
Palmer, W. — (16
Pammel, L. H. — (17, 19); 78 (16); 88
(19); 187 (22); 272, 1699 (17); 1806
(16
Pam nai L. H. & Blackwood, R. E.
3589 (31); 3638 (16).
Parish, S. B. — (14, 47a); 2819 (22);
pend 7149 (14); 10288 (31); 11590
ors W. F. 196, 197 (47).
Parish, S. B. & W. F. 969 (22); 1043 (14);
2171 (31); 11148 (14).
Parker, C. F. — (6, 17, 31).
Parlin, J. E. — (16).
Parry, C. C. — (47a); 167 (40).
Parry, C. C., Bigelow, J. ia Wright, C.
& Schott, A. — (26b, 31, 40, 44).
Parry, C. C, & AMA J^ e p^
(22)
Parry, C. C. & Palmer, E. 717 (7); a
719 in part (46a); 719 in part (43);
720 (36a); 722 (28).
Patterson, H. N. — (19).
Patzky, — (28).
Pease, A. S. & boy B. 22350 (16).
Peck, M. E. 8
Peebles, R. H. po a ; 6756 (47a).
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA
Peebles, R. H., Harrison, G. J. &
Kearney, T. H. 3790 (26a); 4243 (31);
4549 (40).
Peirson, F. W. pi (7).
Penard, E. 348 (41).
Pennell, F. W. 4204, 4332 (1); 5444 (40).
Perkins, J. R. 1024 (1); 102 (2).
Peterson, N. F. — (
Phelps, O. P. 799 (18); 800 (15).
Pilsbry, H. A. —
Piper, C. V. 6160 d
Pitcher, — (35).
Pittier, H. 414 (29); 1909 (11).
Plank, E. N. — (19).
Plaskett, R. A. 142 (22).
Plummer, F. G. — (42
Pollard, C. L. — (38); 1191 (14); 1312
17).
Pollock, W. M. — (16).
( ;
Pringle, C. G. — (11, 14, 25, 26a, 28, 40,
47a); 54 (28); 270 (26a); 1117 (43);
1599 (7); 1948 (10); 2228 (34); 2813
(11); 2927 (46); 3467 (41); 3551
(46a); 4180 (46); 4769 (12); 4784 (29);
4877 (5); 4892 (11); 5715 (7); 6539
(28); 6908 (36); 7590 (28); 7591 (43);
8534 (7); 9135 (28); 9312 (11); 9318
(43); 9529 in part (7, 11); 11091 (43);
11092 (36); 11093 (5); 11843 (34);
13157 (41); 13158 (43); 13159 (28);
13434 (51); 18597 (12); 13792 (46).
Purpus, C. A. 13? (46a); 195 (49); 467
(29); 1094 (26b); 1095 (43); 1433 (36);
3406 (13); 3407 (43); 4524 in part
(31, 43); 4974 (36a); 5451 (10); 5515
(29); 6061 (47); ia m
Quaintance, A. L. —
Rafinesque, C. S. — ap.
Randolph, L. F. & F. — (32); 52
(37); 474 (15); 587, ib (6); 688 (1);
1004 (3); 1012
Ravenel, H. W. — (15, 31, 32).
Read, A. D. — (14); 19 (20).
Reade, J. M. — (1).
Redfield, J. H. — (15, 16); 522 (19);
6430 (17).
351
Redfield, R. 4 (11).
Rehn, J. A. & Viereck, H. L. — (42).
Renson, C. 175 (11); 291 (5).
Reverchon, J. — (15, 19, 35, 40); 42 (8);
118 (25); 782 (8); 734 (15); 735 (19);
736 (31); 737 in part (25, 29a); 738
(48); 739 (40); 740 (35); 834 in part,
1953 (25); 1961 (29a); 1962 (40); 1963
(48); 2116 (31); 2117 (48); 2118, 2532
(24); 2533 (35); 8902 (33); 3903 (29a);
4814 (25).
Riddell, J. L. 1268 (8).
Ridgway, R. 2431 (19); 2831 (15).
Riehl, N. — (16); 9 (31); 185 (15); 196
(19); 456 (17).
Robinson, B. L. — (19); 26 (32); 94 (2);
118 (6); 127 (2); 134 (15); 153, 196
(16); 559 (15); 672 (16).
11).
; 191 (32).
Rose, J. N. 1677? Hew 1763 (7); 2485,
2769 (43); 11968 (47a); 16155 (50).
Rose, J. N. & Fitch, W. R. 17105 (40);
17106, 17503 (31); 17504, 17801 (41).
Rose, J. N., Fitch, W. R. & Parkhurst,
17717 (20).
ose, J. N. & Hay, R. 5310 (12); 5488
(28); 5556 (36); 6049 (46); 6212 (29).
Rose & Hough, = : 6).
se, J. N. & Pai , 4. H, 6481 (43);
7741 (29); 7742 io 7799 (1).
Rose, J. N., Painter, J. H. & Rose, J. S.
8382 (7); 8495 (28); 8496 (11); 8668
(36); 8753 (7); 8823 (46a); 8971 (29);
9077, 9151 (5); 9162 (29); 9503 (43).
Rose, J. N. & Rose, J. S. 11398 (43).
Rose, J. N., Standley, P. C. & Russell,
P. G. 12451, 12934 (7); 12745 in part
(48); 12840 (47a); 13130, 13422,
13447 (7); 18449, 15176, 15181 (47a).
Rothrock, J. F. — (47a).
Rovirosa, J. N. — (5).
von Rozynski, H. W. 13, 17, 135 (29a).
Ruano, J. M. 332, 03 (5).
Rugel, F. — (15, 31, 32, - 121 (6);
127 (17); 156 a 305 (38
Ruger, M. — (17, 31).
352
Runyon, R. 628 (33); 629 (29a); 630
und E (15 ak
H. — (20, 42); 124 (46);
"T Ay 336 (31).
Ruth, A. — (6); 107 (40); 108 (8); 109
(31); 110 (48); 495, 518 (31); ?31, ?40,
765 (17); 788, 833 (15); 1289 (25).
Rydberg, P. A. — ; 154 (17); 932
(19); 934 (31); 935 (40); 1422 (19);
1515 (10); 1716 (15).
Rydberg, P. A. & Carlton, E. C. 7043
(31).
Rydberg, P. A. & Garrett, A. O. 9201
28).
Rydberg, P. A. & Imler, R. H. 22 (35);
120 (17); 379 (35); 433 (19); 482 (15);
1008 (31); 1212, 1253 (40).
Rydberg, P. A. & Vreeland, F.
5676 (41).
St. John, Mrs. O. — (20).
Safford, W. E. 216 fs 1221 (34); 1398
R. —
(43).
Safford, W. E. & Mosier, 210 (37).
Salas, G. 32 (5).
Salazar, F. — (5, 7, 10, 12, 28).
Sandberg, J. H. & Leiberg, J. B. 340
(31).
Sandberg, J. H., MacDougal,
E. XT.
& Heller, A. A. 264, 972 (31).
Sanford, S. N. — (19).
Sarvis, J. T. 122 (31).
Savage, E., Cameron, F. E. &
Lenocker, J. E. —
Schaffner, J. G. 716 (362); 717 (43);
718 (5); 719 d 720 (28).
Schiede, C. J. W. 8
Schoenfeldt, L. Pie (T.
Schott, A. — (33, 40).
von ^n H. — (6, 15, 16, 31, 40,
48).
Schuchert, C. — (17).
Schuette, J. H. — (15, 19, 3
Schulz, E. D. 255 (20); je (33); 679,
701 (24); 766 (8); 767 (33); 845 (43a).
Scott, W. — (15).
Schumann, W. 1071 (43).
Seaton, H. E. 7 (7); 27 (5); 150 (43);
3822 (29); 391 (43).
Seler, C. & E. — (27); 22 (28); 722 (10);
ANNALS OF THE MISSOURI BOTANICAL GARDEN
[Vor. 20
783 (36); 724 (13); atb 1379 (43);
4194, 4347 (13); 4560 (15).
Sesse, M. & Mocino, J. M. 99 (51).
Seymour, A. B. —, 48, 49 (17); 50 (8).
Seymour, F. C. 511 (16); 601 (15); 1324
(16).
Seymour, A. B. & Earle, F. S. 118 (32).
Seymour, A. B. & obe — (81
Shannon, W. C. 3638 (11).
Shear, C. L. 38 nit 191 (19); 202 (16);
3577? (20).
Sheldon, E. P. — (31, 40); 26 (35); 38
(31); 48 (40); 49 (17); 8202, 8856
(31); 11167 (16).
Sherff, E. E. — (19); 333 (35); 1649 (16);
Short, C. W. — (17, 19, 35).
Shull, G. H. 156 (16).
Singer, J. W. 178 (17); 308 (15).
Sintenis, P. 1074, 2010 (14).
Skehan, J. — (14, 32); 20 (27); 46 (8);
4r (40); 109 (8).
Small, J. K. — (6, 9, 15, 16, 17, 19, 31,
32); 2100, 2124 (37); 4020 (14);
5707 (3); 8469 ve 8509, 8594, 8599
(37); 8686, 8732 (38).
Small, J. K. & Qe. J. J. 1072 (14);
1077, 2994 (37).
Small, J. K., Carter, J. J. & Small, G. K.
38311 (3
Small, J. K. 2 Heller, A. A. — (9); 432
"e 484
T E p Small, G. K. 4337, 4341,
^ 4520 (14); 6422 (37).
Small, J. K. & Wherry, E. T. 11774 (8);
11781 (40); 11818 (1); 11824 (33);
11826 (29a); 11834 (48a); 11901 (34);
11903 (33); 11968 (40); 11969, 11980
(29a); 11998 (8); 12010 (33
ie A K. & Age P, 1961 (37).
Smith, s n part (43); 222 (5);
"ig S e A (48): 173? (11).
Smith, C. P. 3186
e
Smith, E. A. — (35. -
Smith, E. C. — (41).
Smith, H. H. 5645 (19); 5654 (16);
5670, 5923 (17); 5952, 6028 (19).
Smith, J. D. — (17, 31, 32, 35).
1933]
PERRY— NORTH AMERICAN SPECIES OF VERBENA
Smith, J. G. — 464 (46).
Smith, L. C. 27 (7); 348 (45); 412 (29);
Spencer, M. F. 971 (7); 994, 1164 (22);
1414 (7).
Stabler, L. M. — (15, 16).
Standley, P. C. — (27, 38, 41a, 42);
4223 (20); 4458, 4549 (31); 4927 (20);
4951 (41); 5136 (31); 6065 (20);
6268 (31); 8239 (20); 8367, 9171 (35);
9336 (17); 9362 (31); 14224 (20);
20617 (5); 21831 (11); 21486 (5);
22414 (11); 22842 (5); 23083 (11
23326, 24404 (5); 24443 (11); 32652,
33293 (5); 40625 (42); 40686 (27);
55944 (11); 56082 (5).
Standley & Valerio, 45430 (5).
Stanfield, S. W. — 2 24.)
Stanford, E. E. ?21 (16).
Stearns, E. S. ? PN 104 (41); 542, 358
“we
(20
Steele, E. S. 106 (16); 314 (17).
Stephenson, B. C. — (15
Stevens, G. W. 1 4 (40); 28 (48); 109
(85); 391 (31); ys (40); 484 (41);
680%, (31); 699 (40); 1034 (19); 1108
(40); 1192 (31); 135414 (19); 1673,
1742 (15); 1791 (16); 1821 (15); 1865
(19); 1980, 2080, 2299 (17); 2340 (35);
2850 (40); 2889 (31); 2901 (16); 2930
Stevens, W. C. — (17).
Stevens, W. H. 28 (48).
Stevens & Hess, 4260 (14).
Steyermark, J. A. 292 (17).
Stone, W. — (46).
Stratton, R. 10 (40); 11 (48); 62 (8);
158 (15); 249 (19); 317 (40); 400 (16);
403 (19); 449 (40).
Street, M. H. — (35).
Studhalter, R. A. 1106 (48).
Sturtevant, E. L. — n 16, 35).
Taylor, A. 4244 (2 (2).
353
Taylor, K. A. — (32).
Tejada, R. 57, 72 (11).
Thackery, F. A. 348 (47a).
Thackery & Leding, 1100 (47).
Tharp, B. C. 667, 668 (24); 838 (35);
1361 (29a); 1362, 1364 (48); 1826 (34);
1871 (48); 2558 (2); 2818 (8); 3166
(24); 3682 (27); 5538, 5610 (33).
Thompson, C. H. 154 (19).
Thompson, H. 42 (31); 4412 (22); 4880
(31
Thone, F. H. 50 (15); 88 (19).
Thornber, J. J. — (16).
Thurber, G. — (6, 15); 143 (42); 446
(26a); 555 (7); 769 (48).
Thurow, — (1).
Tidestrom, I. 872 (26a); 898 (47);
11418 (15); 11551 (16); 11572 (15).
Tindall, C. W. — (17, 3
Tonduz, A. 488 (11); 627 (5); 628 (11).
Torrey, J. 154 (15); 416, 417 (22).
Toumey, J. W. — (40a, 41a, 43b, 47);
805a (47); 305% T 306 (25).
Townsend, C. & Barber, C. N.
ed TU 192 do.
Tra M. — (8, 14, 15); 4981 (32);
0050 jm 6652 ad 7588, 7996 (8);
7999, 8000 (40); 8001 (31); 8037 (15);
8706 (3); 8707 (35); 8708 (24); 8709 (8).
F. S. 30, 41 (25);
art (42, 43b); 162a (42); 178 (48).
dA & mm Lloyd & Tracy.
Trask, B. — ;
Trelease, W. — (8, 15, 16, 17, 19, 29a,
31, 35, 40, 43a); pe (48); 101 (33);
494 (31); 718 (17); 714 (35); 715 (31);
716 (16); 717, 718 (19); 719 (15); 1161
)
(35).
von Tuerckheim, H. 77 651 (11); 904 (5);
913 (11); 1068 (46); 8442 (11).
Tweedy, F. —, 113, 246 (29a).
Tyler, F. J. — (40 i?
Umbach, L. M. — (19).
Van Sickle, W. M. — (19, 31).
Vasey, G. — (17, 22, 31, 35, 41, 47a);
468, 512 (31).
[Vor. 20
354 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Velasco, L. V. 8848 ( Eu hen (11). Wislizenus, A. 150 (26a); 325 (29a);
Victorin, M. 3/24, 1 15). 483 (42); 525 (A1).
Viereck, H. L. — (4 E Wislizenus, F. 280 (17); 282 (15); 284
Visher, S. S. — (15); 1507 (19); 2028 (31); 285 pee 1214 (6).
(31); 2132, 2177 (40); 2263 (15); Wolf, C. B. 2
8359 (31); 4439 (16); 4450 (19). Woodson, R. E 198 (35).
Vreeland, F. K. 636 (20); 804 (43b). Wooton, E. O. — (20, 25, 26, 26a, 27,
ui n F. 974 (41a). 31, 41, 41a, 42, 43a, 47, 47a); 187 (27);
Wallac — (14). 208 (20); 363 (41); 364 (42); 409
Ward, L F. — (6, 24, 31, 35, 40, 41); (31); 642 (41); 646 (26); 2831 (31);
11 (20); 256 (31). 2835 (41a); 2836 (31); 3849 (43b);
Warner, S. R. 43 (24). 8852 (26b).
Watson, S. 823 (31). Wooton, E. O. & Standley, P. C. 3330
augh, F. A. — (15); 168 (35). (31).
Weatherby, C. A. 5076 (16). Worthen, G. C. — (5).
Webb, R. J. 5441 (16). Wright, C. — (15, 42); 454 (31); 4
Webber, H. J. — (16, 19). (39); 567 (31); 1496 (25); 1497 m
—_ M. 165 (40); 174 (16); 197 (40); 1498 (16); 21498 (44); 1499 (31);
6 (31). 1500, 1501 in part (48); 1501 in part
Sn O. E. — (40). (44); 1502 (40); 1508 in part (39, 40a,
Wiggins, I. W. & Gillespie, D. K. 3977 42); 1504 in part (40, 42); 3658 (6);
(23). 3659 (14)
Wilcox, T. E. — (11, 26a, 28, 40a, 47). Wurzlow, E. C. — (3, 5).
Wilkinson, E. H. — (8, 26a, 43); 8 (489); Young, H. A. — (17).
9 (40); 108 (48). Young, J. P. 737 (38).
Williams, E. F. — (15, 16). Young, M. 8. — (8, 25, 27, 42, 48); ??
Williams, T. A. — (40). (8); 112 E 29a); 114 (48); 662
Wilson, N. C. 99 (47). (1); 1708 (26).
Wilson, P. 161 (17). Zuck, M. — y 43b, 47).
INDEX TO SPECIES
New species, varieties, and combinations are printed in bold face type; synonyms,
in italics; and previously published names, in ordinary type.
Page Page
Anonymos caroliniensis........... 316 UDerOWÍa. coord xen 251
Billardiera................ ees 251 Verbena.......... uses 251
€ "RERE EEE A 36 («oS AED 316 JC MTM 257
Buchnera canadensis,............ 316 Alopecurus.... 0.0.00. e cece 285
Glandularia. ...... eer 251 ambrosifolia.................. 326
BINE in bs nh revu» STARS 316 forma eglandulosa .......... 328
bipinnatifida... a...an ee 323 amoena. r E EE AA 341
caroliniensis...... lessen 316 Andrieuxii..................05 333
Helleranthus quadrangulatus....... 313 angustifolia PIXWRAG PAGG EAT ERE 282
ee ee TT PETIT 251 NONE ur EFVEKEXE: 300, 337
Phryma ici i EG d USYE T 309 Aubletia....... e elles
Pep PRETT IEEE 251 var. Drumm a, ss. 316
1933]
PERRY—NORTH AMERICAN SPECIES OF VERBENA 300
Page Page
PAAR 0S eee ee ees 342 gracilis, 6.0. <5 2% ee ed 300
bipinnatifida.................. 323 randi loras cisco ee ores me 341
var AMODA 60 dtd as 325 imr LER ie oS see CDM ER 342
USER NOLES fee tia eee oe ate ee 208: 3HB8l601. oa aera ee ee 265
DOUATIGNAIS sere Ae coches otra 254 Ha mae papaya Soak eee RE 257
bonariensis hastata: «2. sau cee E TA 278
VACS OT OUE renes a 257 hastata
aa A E E 253 var. oblongifolia.............. 267
DIACOR OU E E N 304 VAT PONCO a 640 Gas 278
eos UR ENG box 304 TINS 0800. o eua 278
var 12) Poner EE 304 var. pinnatifida. oss s 278
var. ERAS No ATA AN 304 Mreta S E A E A ee 268
DFASHIODSIBS e a aA eso outs eph a 255 lax oer ONOCN. MI
OOTOLO 0e o 338 s MUROS ACTEUR 342
ONE se sin aii ss RR 315 ACU. ooo NET te ee 338
fe gatos DD et ee 342
var. Ehrenbergii. ............ 318 UOTE eve CUM ERAN 316
BUDEP: INO, oo oi ees 318 VAT. T0860... eene 316
VET LOAIOCT ech no 316 LASTOGLOCIYS = as Ec 290
CATIOBDOIH seers Soe eee 301 leucanthemifolia...........0055 265
ar. Roemeriana............ 302 BUA, E ee eee 340
CUREHNCOH S Eder vd trs MER RR. 304 ey renee TT T 257
VAT: n60m6éxtcana., .... s ec eae 296 litoralis
COP GC 0 EAE oe DN a 257 VAL Draschensis | Sd REEL 255
Den ecu me oo Sere e 309 VAT CO Gcasqna.. ee ck ere 257
ORTUDDUSR NS io conet dta 268 lit is
COME. eee i 268, 309 B leptostachya. .............. 257
orma or var. pu "ee 2 a Dycnostachyja ..«.. b. eee 257
forma or var. recía. .......... 271 DUM. eis a A 315
COE derer verre ya 309 D ADMIRE TRITT 272
alibi p MAR PT 330 HOG Loser ee eme esta 293
var. longidentata...... ...... 331 Mae DONA. acosa does ris 288
ee a e ee 332 acDougalii mut. rosella. ...... 288
DAR E E ME 304 BEDA 6.43. epo EX ran 289
cuneiolua. v 286 IBS nek, PEPEE TE ea E E 320
delicatula.. aena a aaa 342 AE ar ai i AREA A ESTO 342
delticola 22 n 9 E E 314 menthaefolia. .... -iie e arrenak 263
aunn Ea POE E N A NE 275 CD RENE r 268, 286
dOTmATgensis- oos Se ee eee 262 SROVONCRAS « voveo bas a 31
DONON s ons oe ee re 316 TOOMGENIGANG . ov Fe os nests 296
Ehrenbergiana................ 267 var. Wirtella. : ....: 2.9 985 298
[MISC EDD EOS LS 318 warszylopoda....4.. 2.0 297
var. asperata................ 319 UDIIPNg. or ee eee 315
CHO HORRORE EET EET 254 DNENGE Lees RUE e 315
PBELIGRIGN ae erue sd se 341 OIDODAIDU o ore que EP eT 262
Aldi M M E 334 officinalis var. hirsuta. ......... 296
GOOG DEEP Eee ees oso orn o 335 DOFCULHanA - cuve. ud o rs 284
var. eile Ng 337 DREN, @ os 5:- - se vas STU 278
356
Page
var. pinnatifida.......... LL. 278
DEUM ev rk a xao Ys 268
BINNEN. coxrkrrsa kv d Cao 268
paucifolia..............Luesue. 342
WOON PRIMETIME 299
pinnatifida chee awed sad eee ees 278
ee LE T ee 294
polystachya . . 6... ce cee ee 268
prostrata... 0... lessen 290
C CP PEERS POPE EPOR 332
aiino ee ee AARON 332
WER ONNE ERES EEE T TE E 338
ai forma albiflora.......... 313
quadrangularis... .......ssc.... 254
soci ia ETELE EP TENER 313
^j líí MÀ 332
"dO: AUOD PEPCPRRATTPPMSE RM 271
TOMOA, onc xr er ea 300
jn. PEE 342
Ju S TERRE TETTE ITILTI 286
HN LL cocco XE eens 252
TAR Loo keh x4 xxv RERI 267
| a ry rrr eda 292
Roemeriana.........i sees. 302
PIPILL o cep TREE YEAR 316
xo ^ PROFERRE PLE TET 304
i rr 282
d EERTE TEET ET EEE E 272
Se ee ee ee T 272
penis TERE end ceed bred eaten 343
[Vor. 20, 1933]
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Page
BMRONA 5. 6 44 4s Pi ER RE 339
NND. ioo rr ne SAE ate rT 263
s; or^ AREE USC 282
ono MD 1730942351 ATEEN TT 256
EEEREN AT ET 262
ee ee ET 304
sion 93-8 32 E89 EARN XR REX BAS 285
gio e UIE S 12g E Eu 286
Pt Ms EEEE 286
forma arene in | him | 2 e m| a S516] 8 51A |
e p = > 2 e^ [» e = z z 7 "d uonnpog
?|l$|2$|5|"*|*;$|"]|s|H|** |] BIE
UALIT Hud SNVUSD NI SNOILOTOS SQO0IHVA
I FIAVL
JHL AO NOLLISOdIXOO
1933]
WYND—NUTRIENT SOLUTIONS FOR ORCHIDS 365
per million, in table r1; and the growth data, in table rm. All
solutions received 1 cc. of a M/200 suspension of ferric phosphate
per liter prepared as described by Livingston (719).
TABLE II
COMPOSITION OF THE SOLUTIONS IN PARTS PER MILLION
OF THE NUTRIENT COMPONENTS
Ca | Mg K PO, | SO, | NO; | NH, quud Cl
La Garde 360 | 98 287 | 698 | 390] 388 | 283 | 308 658
Crone 213 49 387 153 474 623 140
Knudson 169 25 112 137 462 525 137 225
Sachs 310 49 387 307 474 623 140 302
Pfeffer 135 20 186 140 78 545 123
Shive A 144 | 360 703 | 1710 | 1440 448 101
Schimper 291 | 42 260 | 235 | 168 | 1071 241 260
Shive B 208 | 480 563 | 1368 | 1930 | 646 145
Knop 169 | 25 134 | 175 98 | 525 118 58
Tottingham 399 | 347 700 | 1235 | 1390 | 1547 348
Hansteen-
Cranner 202 | 60 129 | 314 | 1098 322 | 250 458
Zinzadze 49 | 17 62 47 | 112 51 15 23 58
TABLE III
GROWTH DATA, BASED UPON THE AVERAGE OF 25 SEEDLINGS
Solution Height in Diameter in| % Total pH at ps E ~
microns microns salt planting experiment
La Garde 4243 1590 . 400 4.9 3.8
Crone 3760 1213 225 5.5 4.5
Knudson 3360 1145 . 200 5.0 4.5
Sachs 3258 1300 .300 D. 4.5
Pfeffer 2953 1135 .150 5.0 4.6
Shive A 2908 1250 .700 4.8 4.5
Schimper 2835 1223 .945 5.1 4.5
Shive B 2793 1213 .812 4.8 4.4
Knop 2360 1145 .162 5.0 4.5
Tottingham 1868 900 .820 4.9 4.5
Hansteen-
Cranner No growth | No growth .295 4.9 3.9
Zinzadze No growth | No growth .052 5.1 3.7
The fact that no growth occurred on the solution of Hansteen-
Cranner and of Zinzadze was probably due to instability of the
or, 20
366 ANNALS OF THE MISSOURI BOTANICAL GARDEN
pH of these solutions as shown by their high acidity at the end of
the experiment. In this connection it is interesting to note that
7 § § 8 8 8 g 8$
F The relation of growth to caleium content. Zinzadze, Pfeffer, Shive A,
Knop, Knudson, Hansteen-Cranner, Shive B, Crone, Schimper, Sachs, La Garde,
Tottingham.
| |
8 8 5 8 8 $ 8 8 § 8
Fig. 2. The relation of growth to magnesium content. Zinzadze, Pfeffer, Knop,
Knudson, Schimper, Sachs, Crone, Hansteen-Cranner, La Garde, Tottingham,
Shive A, Shive B.
1933]
WYND— NUTRIENT SOLUTIONS FOR ORCHIDS 367
3500 4
2800 4
700 4
8 Š 8 8 S 8 8 8
Fig. 3. The relation of growth to potassium. Zinzadze, Knudson, Hansteen-
Cranner, Knop, Pfeffer, Schimper, La Garde, Sachs, Crone, Shive B, Tottingham,
Shive A
$500
2.5 9 3$ 2 $134 73
Fig. 4. The relation of growth to phosphate. Zinzadze, Knudson, Pfeffer,
Crone, Knop, Schimper, Sachs, Hansteen-Cranner, La Garde, Tottingham, Shive B,
Shive A.
[Vor. 20
368 ANNALS OF THE MISSOURI BOTANICAL GARDEN
= > Y T - de cones xS aaa ver
3 5 3 8 3 P E § Po]
Fig. 5. The relation of growth to sulphate. Pfeffer, Knop, Zinzadze, Schimper,
La Garde, Knudson, Crone, Sachs, Hansteen-Cranner, Tottingham, Shive A,
ive B.
Shive
4200 |
3600
2800 |
2100 |
5
SO: @ E § P RP b
Fig. 6. The relation between growth and nitrate. Zinzadze, La Garde, Shive A,
Knop, Knudson, Pffefer, Crone, Sachs, Shive B, Schimper, Tottingham.
La Garde's solution also was strongly acid at the end of the
experiment, probably because of the greater growth of the
numerous seedlings upon it. The use of ammonium carbonate by
1933]
WYND—NUTRIENT SOLUTIONS FOR ORCHIDS 369
La Garde was probably for its buffer action against this unde-
sirable change of acidity with growth. The effectiveness of the
concentration of carbonate used as a buffering agent was tested
by preparing La Garde's solution with and without carbonate
and then comparing the titration curves obtained with N/100
hydrochloric acid. In both cases, the titration curves were
$500
2800
2100}
1400[
i 5 : 5 ;_—,
8 8 2 3 3 3 3
Fig. 7. Relation between growth and ammonia. Zinzadze, Knudson, La Garde,
Hansteen-Cranner.
identical. The solutions were also analyzed for carbonate after
autoclaving, but no positive test could be obtained. The auto-
claving at 20 pounds pressure for 20 minutes at the initial pH
necessary (4.25) undoubtedly destroyed the small amount of
carbonate present.
Examination of table 1 shows that the best growth occurred
on the solution of La Garde, followed by that on Crone's and
[Vor. 20
370 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Knudson’s. While the seedlings on La Garde’s solution were
conspicuously larger than those on Knudson’s solution, the Knud-
son seedlings showed a definitely superior root development.
The Crone seedlings were actually larger than those on Knudson’s
solution, but they did not appear to be so green. Of all the
solutions tested, we would regard those of La Garde and Knudson
to be the most satisfactory.
4200 |
3600 |
2100 }
n Lond [2d to to CA CA
o o 3 9 S 8 a
Fig. 8. Relation between growth and total nitrogen. Zinzadze, Shive A, Knop,
Pfeffer, Crone, Sachs, Shive B, Knudson, Schimper, Hansteen-Cranner, La Garde,
Tottingham.
The graphs indicate the comparative growth in relation to the
concentrations in parts per million of each ion. It is apparent
that the quality of the solutions is not related to a specific amount
of any one ion. The author regards the superiority of La Garde’s
solution to be due to the particular complex of nutritional factors,
and not to the specific effect of any particular ion. The nature
of its superiority might well be a favorable condition of permea-
bility of the cells produced by chemical means not yet understood.
: 933]
WYND—NUTRIENT SOLUTIONS FOR ORCHIDS 371
4200
3500
2800
2100
1400
700
a g B 3} à 85 & 1n TEM
Fig. 9. Relation of dia to chlorine. Zinzadze, Knop, Schimper, Sachs,
Hansteen-Cranner, La Gar
| |
à E P o
o
2
£
ost’ f
Fig. 10. Relation of growth to per cent total salt. Zinzadze, Pfeffer, Knop,
Knudson, Crone, Hansteen-Cranner, Sachs, Schimper, La Garde, Shive A, Totting-
ham, Shive B.
In all figures, the vertical distance represents the height of the
seedlings in microns, and the horizontal distance represents the
[Vor. 20, 1933]
312 ANNALS OF THE MISSOURI BOTANICAL GARDEN
concentrations in parts per million of the ions indicated. The
vertieal lines representing the growth on the various media are
listed from left to right.
CONCLUSIONS
1. Seeds of Cattleya Trianae Linden & Rchb. f., were germinated
and grown on a number of published nutrient solutions. The
best growth was obtained on La Garde's solution, which was
closely followed by that obtained on the media of Crone and of
Knudson.
2. No growth was obtained on the solutions of Hansteen-
Cranner and Zinzadze. This was probably due to the unstable
pH of these solutions.
3. The solutions studied differed in so many factors that the
effects of any given species of ion were modified and obscured.
Hence the nutritional value of the various solutions may not be
interpreted as the effects of the concentrations of specific ions.
This is clearly illustrated by the graphs.
The author wishes to thank Dr. E. S. Reynolds, Plant Physi-
ologist, Missouri Botanical Garden, for his coóperation, and also
Dr. G. T. Moore, Director of the Missouri Botanical Garden, for
a generous supply of orchid seeds.
BIBLIOGRAPHY
Benecke, W., and Jost, L. (24). Pflanzenphysiologie 1: 135-137. Jena, 1924.
Duggar, B. M. (24). Plant physiology, p. 145. New York, 1924.
Knudson, L. (22). Non-symbiotie germination of orchid seeds. Bot. Gaz. 73: 1-25.
1922.
La Garde, R. V. (29). Non-symbiotic germination of orchids. Ann. Mo. Bot.
Gard. 16: 499-514. 1929.
Livingston, B. E, (19). A plan for coóperative research on the salt requirements of
epresentative agricultural plants. Baltimore, 1919.
maine D. T. (01). Plant physiology, p. 224. New York, 1901.
Shive, J. W. ('15a8). A study of physiological balance in Aus media. Physiol.
Res. 1: 327-397. 1915
, (15b). A three salt nutrient solution for plants. Am. Jour. Bot. 2:
157-160. 1915.
Tottingham, W. E. (14). A quantitative chemical and physiological study of
nutrient solutions for plant cultures. Physiol. Res. 1: 133-245. 1914.
Wynd, F. L. (33). The sensitivity of orchid seedlings to nutritional ions. Ann.
Mo. Bot. Gard. 20: 223-237. .
Zinzadze, S. *. (26). Eine neue Náührlósung. Ber. deut. bot. Ges. 44: 461-470.
Annals
of the
Missouri Botanical Garden
Vor. 20 SEPTEMBER, 1933 No. 3
THE FOLIOSE AND FRUTICOSE LICHENS
OF COSTA RICA. I
CARROLL WILLIAM DODGE
M ycologist to the Missouri Botanical Garden
Professor in the Henry Shaw School of Botany of Washington University
While there exists much literature on various groups of lichens
from various parts of tropical America, at present I know of no
recent work covering the whole area or that of any constituent
country whereby the traveler may identify the lichens he meets,
or the student in an herbarium of the temperate zone may readily
determine any considerable portion of the miscellaneous material
sent in by collectors. It is hoped that this work may furnish
such a manual, although no one realizes better than I the huge
amount of monographie work necessary before a book approach-
ing completeness and accuracy can be published.
I first came under the spell of the rich flora of Costa Rica
during a two-weeks visit in the summer of 1925 and at that time
formed the resolution to spend my first sabbatical year in that
country in the study of its fungus and lichen flora. During the
subsequent years much material in addition to my own col-
lections came to me for identification. Plans were matured
and in early September, 1929, my family and I landed in Costa
Rica and made our headquarters in San José, the capital of the
country. In the latter part of September I was joined by Mr.
W. Stephen Thomas, a student at Harvard, who accompanied me
on most of the collecting trips until the middle of March. Mrs.
Dodge also spent some time in the field and much is owed to her
keen observation and helpful care of the collections. Even the
small daughter, at the age of four, added a number of specimens
Issued October 6, 1933.
ANN. Mo. Bor. Garb., Vor. 20, 1933 (373)
[Vor. 20
374 ANNALS OF THE MISSOURI BOTANICAL GARDEN
not otherwise easily obtained, through her skill in climbing
guayabos (Psidiwm guajave L.) which were too small and too brittle
to be climbed by an adult. We left Costa Rica in June, 1930,
and after a brief stay at the Farlow Herbarium to sort over
material and give tentative determinations I took the more
puzzling specimens to Europe where the months from August
to December, 1930, were spent in the larger herbaria.
Very many persons and institutions have contributed to the
success of this undertaking both at home and abroad, and to all
] tender my most hearty thanks in grateful acknowledgment of
their many kindnesses. For financial assistance, I am grateful
to the John Simon Guggenheim Memorial Foundation which
appointed me as a Fellow to Costa Rica for the year 1929-30
and to Europe from October to December, 1930; to the President
and Fellows of Harvard College and to the Farlow Herbarium,
for equipment, incidental expenses, and for an assistant in the
field; to my colleague, Prof. Greenman, and former colleagues,
Professors Ames, Barbour, and Thaxter, and to Mr. Paul C.
Standley of the Field Museum, for excellent advice in planning
my trip and for helpful letters of introduction. Members of
the staff of the United Fruit Company were also very helpful
both in planning the trip and placing the facilities of that great
company at my disposal in Costa Rica, especially Dr. J. R.
Johnston of the Boston office, and Messrs. Kress, Fuller, Craw-
ford, Stübbe, and George Catt, the director of its Botanical
Garden and Experiment Station at Siquirres, who accompanied
me on several trips in that vicinity.
Among Costa Ricans, I am deeply indebted to the following
for advice and hospitality, without which I should have been
unable to cover so much territory so thoroughly in so little time:
Bernado Yglesias Rodriguez, the Director of the Escuela Na-
cional de Agricultura, who placed the facilities of the botanical
laboratory at my disposal and enabled me to spend three very
profitable weeks at the ancestral Yglesias finca, Guayabillos, on
the upper slopes of the western face of Irazü; to Anastasio Alfaro,
formerly director of the Museo Nacional (and later to his suc-
cessor, the late J. Fidel Tristan), who placed the facilities of the
library and herbarium (Wercklé) of the institution at my dis-
1933]
DODGE—LICHENS OF COSTA RICA. I 375
posal and suggested the plan of my trip to Guanacaste, as well as
contributing several specimens; to Dr. Alberto M. Brenes, the
botanist at the Museo Nacional, who collected extensively for
me in the vicinity of San Ramón, an interesting region which
I did not have an opportunity to visit personally; to Ricardo
Chavarría Flores, formerly city engineer of Cartago, who en-
abled me to collect extensively at his finca in the vicinity of
Santiago de Cartago and the Rio Birrís; Juvenal Valerio Rodri-
guez, of the Instituto de Alajuela, who planned trips in the
vicinity of Alajuela and to the Cerros de Zurquí; Otón Jiménez
Luthmer, of the Botica Oriental, a well-known amateur botanist,
who gave generously of his time in suggesting trips and making
plans, although his business duties prevented his accompanying
me in the field except for a short time on our trip to Guanacaste;
to Ferdinand Nevermann, the coleopterist, for numerous pleasant
collecting trips, including a three-weeks stay at fincas of which
he is manager in Limón Province, and for much helpful information
in connection with my large insect collections; to Ruben Torres
Rojas, of Cartago, for specimens, although I did not have the
pleasure of a personal contact with him during my short visits to
Cartago; and to Charles H. Lankester, for helpful suggestions.
Besides these who are more or less professionally interested
in natural history, the following were generous in their hos-
pitality and enabled me to visit regions which would have
been otherwise quite inaccessible to the traveler: Gonzalo Volio
and his son Carlos, Antonio Sobrado and his brothers, Carlos
Collado, Fernando Castro, José Castro Araya, Antonio Gutiérrez,
José Luis Sancho, Carlos Piedra and son, Alexander Ross and
sons, Robert Hanckel, and H. J. Marks.
While in Europe I was indebted to the curators, acting curators,
and staffs of the following institutions for permission to study
the herbaria under their care: the Art Galleries of the Glasgow
Corporation (Stirton Herbarium); British Museum of Natural
History; Royal Botanic Gardens at Kew; Universitetets Bot-
aniske Museum, Oslo; Botaniske Trädgård, Upsala; Riksmuseet,
Stockholm; Botaniske Museum, København; K. Botanische
Garten, Berlin-Dahlem; K. Botanische Museum, München;
Conservatoire Botanique and Herbier Boissier of the Université
[Vor. 20
376 ANNALS OF THE MISSOURI BOTANICAL GARDEN
at Genéve; and the Laboratoire de Cryptogamie at Paris, as well
as the private herbarium of M. le docteur M. Bouly de Lesdain
of Dunkerque. I am also indebted to the following lichenolo-
gists, whom I met during my trip, for helpful suggestions and
criticism: Miss A. L. Smith, Bernt Lynge, Einar DuRietz, the
late Karl Schulz-Korth, J. Motyka, and Ove Hgeg. Since my
return to America I have had access to or seen specimens from
the following herbaria, for which I gratefully acknowledge my
indebtedness to the curators: Farlow Herbarium of Harvard
University, New York Botanical Garden, National Herbarium
of the Smithsonian Institution, the Field Museum of Natural
History, Chicago, the herbaria of the University of Michigan
and of the Missouri Botanical Garden. I am also grateful to
the Director of the Missouri Botanical Garden and to the Chan-
cellor of Washington University for a leave of absence after the
appointment to the staffs of those institutions, in order that I
might complete certain studies in the Farlow Herbarium of
Harvard University before I left that institution. Finally I
wish to thank any others through whose hospitality and assistance
this work has been made possible.
GEOGRAPHY
Topographically Costa Rica may be divided into four main
phytogeographic regions: the Atlantic Coastal Plain, the Meseta
Central, the Subalpine Region, and the Pacific Coastal Region,
the latter less well known and less homogeneous.
The Atlantic Coastal Plain is characterized by its gentle slope
from the sandy beaches up to about 100 m. or even higher in
the poorly known Llanos de Santa Clara and in the Sarapiquí,
San Carlos and Rio Frío valleys. The beaches are interrupted
only by shallow harbors at Cahuita, Limón, and Moin, the first
being protected by a coral reef, the two latter being partially
protected by the small island Uvita, now used as a quarantine
station for Puerto Limón. Several large rivers rising in the
highlands cross this plain and during their flood states overflow,
forming spectacular anastomosing channels which leave small
pools in the dry season. Most of the river mouths are closed by
shifting bars which hinder or prevent the use of the rivers for
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1933]
DODGE—LICHENS OF COSTA RICA. I 377
shipping. Just back of the beach there is an almost continuous
series of lagoons, which may be dredged and may be used for
transport of the farm produce to the railroads and ports. These
lagoons extend most of the way from San Juan del Norte to
Limón.
The soil is quite fertile where drainage is sufficient and much
of the forest has been cleared for agricultural purposes, in the
Llanos de Sta. Clara and Limón. The rainfall is heavy and quite
evenly distributed, although somewhat less during January,
February, and March, the so-called dry season (verano). Dur-
ing most of the year the sun shines until about noon, then showers
may be expected for either a short time or for the rest of the
afternoon. The nights are usually clear and cool.
During the colonial era there was very little permanent settle-
ment of this region although many attempts were made. Some
cacao plantations were developed at Matina, and their product
taken overland via Cartago to Honduras and Guatemala or
quietly sold to the English pirates and smugglers who frequently
entered the mouth of the Matina River for this purpose. With
the advent of the Northern Railroad in 1870-1890 the banana
industry spread rapidly over most of the available land from the
Parismina to the Estrella River valleys and up the Sixaola
from Bocas del Toro. When the banana wilt or Panama disease
(Fusarium cubense E. F. Smith) and increasing soil exhaustion
began to threaten this industry, coconuts, pineapples, and
cacao were extensively planted in suitable localities. Coconuts
were à prompt and complete failure owing to bud rot and other
factors, although they still furnish enough for local consumption.
Pineapple cultivation has recently been abandoned for economic
reasons, although the product was much superior to that of
Hawaii. Cacao cultivation has been only partially successful
owing to high labor costs. Recently much land has been practi-
cally abandoned and is gradually returning to second-growth
forests. While small plantations of forest trees, as Hevea,
Ochroma, etc., are still in an experimental stage, they seem to be
successful botanically but owing to the high cost of labor they
are still doubtful economically. Maize is sometimes grown on
the Llanos de Santa Clara.
[Vor. 20
378 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The Atlantic Coastal Plain penetrates far inland in the Tortu-
guero, Sarapiquí, San Carlos and Río Frío systems, but since
I have not had an opportunity to examine them personally and
have seen very few specimens from them it is impossible to
discuss them further. Before the railroad was built the Río
Sarapiquí was the main route from San José to the Atlantie, and
a few villages still remain. ‘There are a few scattered settlements
on the Río San Carlos and Río Frío, as may be seen from the
relations of Thiel and Cespedes, but except for a few specimens
by Pittier from the upper Sarapiquí Valley and from the vicinity
of San Rafael on the R. San Carlos, I know of no lichen collections
from this region.
'The Atlantie Transition Zone from 100 to 900 m. is evident
mostly in narrow river valleys, the only botanically well-known
portions being the southeastern portion of the Llanos de Santa
Clara around Guápiles, formerly an important farm of the United
Fruit Company used for their earlier experimental plantings
of forest trees, and the valley of the Reventazón.
In the valley of the Reventazón, tertiary sedimentary rocks
are evident in the cafions and railway cuts. The slopes are very
steep, due to the great erosion caused by the heavy rainfall
which is about the same as that of the Coastal Plain. Bananas
and coffee form the principal crops in the broader side valleys,
the banana being comparatively short-lived but much used as a
shade for the coffee. Since this development is comparatively
recent, it still remains to be seen whether this is à permanent
stage or a temporary phase in the agricultural development of
the region. Formerly sugar cane was extensively cultivated
in this region and considerable areas remain in Juan Vifias and
Santiago de Cartago. Pineapples are grown for the domestic
markets near Turrialba and to a less extent in some other places.
Owing to the rapid erosion it seems doubtful whether much of
the land which has been cleared on such steep slopes can remain
permanently in cultivation. "There are available a few collections
from Angostura near Turrialba by Polakowsky and my own from
the beautiful valley of the Río Pejivalle near the United Fruit
Company's farm of that name, as well as the even more extensive
ones from Santiago de Cartago and occasional specimens from
1933]
DODGE—LICHENS OF COSTA RICA. I 379
Juan Viñas (Naranjo) by Ørsted and scattered numbers by
Pittier.
On the practically inaccessible slopes and in the valleys where
transportation is too costly to make farms economically profitable
yet, some forests are still to be found. These still resemble
those of the Atlantic Coastal Plain with their abundant epiphytes,
bromeliads being conspicuous, especially Tillandsia usneoides.
There are many small areas among the hills where fogs keep
the humidity much higher than in neighboring areas. These
are quite evident by the abundance of the long pendulous species
of Usnea which replace Tillandsia usneoides as a prominent
feature of the landscape. Species of Leptogium, Parmelia,
Anaptychia, and mosses are also very abundant. Cladonias
begin to appear and replace the similarly colored moss of the
lower elevations. In the top of the forest trees and in the isolated
trees of the potrero (pasture), crustose species are abundant. In
the lower levels of the forest, lichens are rarer and usually sterile,
perhaps due to the small amount of light. Trees whose bark
scales off rapidly, such as the guayabo (Psidium guajave L.),
are rarely found to have lichens, except for a few fruticose
species of Ramalina and Usnea. Nephroma was found in one of
these Usnea-Tillandsia zones on an exposed ‘‘knife-edge’”’ pro-
truding into the cañon of the Reventazón near Santiago de
Cartago.
The Meseta Central, occupying the huge central valley between
the Cordillera Central and the Cordillera de Talamanca, consists
of long gentle slopes between 1000 and 1700 m. cut by deep
river gorges. The pass of Ochomogo and the Cerro de Carpintera
divide this meseta into two portions, the eastern valley of Guarco,
comprising the upper portion of the valley of the Reventaz6n
and its tributaries, and the western valley of San José, comprising
the upper part of the valley of the Rio Grande de Tarcoles and
its tributaries. The valley of Guareo is bounded on the north
by the upper slopes of the volcanoes Irazü and Turrialba and on
the south by the continental divide in the Cordillera de Talamanca.
Most of the large tributaries enter from the south and are not
explored fully from a geographical standpoint. I have seen
lichen collections from the valley of the Río Navarro and the
[Vor. 20
380 ANNALS OF THE MISSOURI BOTANICAL GARDEN
lower part of the Orosi. The Río Grande is by far the largest
and longest of the tributaries but its flora is wholly unknown to
me. Coffee is the principal crop though considerable amounts of
garden vegetables, especially chayote and yuca, are raised to
supply not only the local markets but also the demands of the
Atlantic Coastal Plain. Cartago, the colonial capital, is the
only city but there are numerous small towns. The collections
of C. Wercklé, ‘‘near Cartago 1200-1500 m.," in San José and
Berlin, seem to be from the region about Aguacaliente and the
lower slopes of the Cerro de Carpintera, judging from species
represented.
The valley of San José is bounded on the north by the continental
divide of the western portion of the Cordillera Central, the volcano
Pods, and Barba with its lesser peaks, the Cerros de las Caricias,
Las Lajas, and Zurquí, on the east by Irazú and the Cerro de
Carpintera, on the south by the Cerros de Escasi and Piedra
Blanca, on the west by the Cerros de Aguacate. In the valley
of San José the slopes descend gently to about 650 m. This
valley is much drier than the Guarco with a longer dry season
from December to April and less rain in the other months. The
winds are strong, especially in the dry season. Practically the
whole region has been denuded of forests, partly in the interests
of agriculture and partly owing to the fiendish glee of the in-
habitants who deliberately set fires to the dry grass on windy
days in order to watch it burn. By the end of the dry season
practically the whole valley outside the cafetales and cane fields
is black from the charred remains of grass fires.
The underlying rock of both valleys is largely volcanic, while
the soil above is largely of aeolian origin from the volcanic dust
which is still being regularly emitted from Irazü and occasionally
from Pods. In some places fields of voleanic boulders exist in
such profusion that they make agriculture as difficult as do the
glacial boulders in parts of New England. The greater portion
of both valleys is devoted to coffee growing, with sugar cane and
upland rice in the lower levels. Formerly considerable excellent
tobaeco was grown but this crop has practically disappeared.
The cities of Alajuela, Heredia, and San José, and many important
towns are located in this valley.
1933]
DODGE—LICHENS OF COSTA RICA. I 381
Crustose lichens are abundant in the cafetales both on the
coffee and its shade, but practically no attempt was made to
collect them since the thalli are mostly young and sterile and
we hesitated to injure the trees by removing portions of bark.
The living fence posts, mostly species of Erythrina (poró), were
utilized, however, since they are regularly and heavily pruned
back. These yielded most of the crustose species from this area.
Another fruitful source was the roadside bank. These banks are
caused by erosion and by cutting down with spades to secure
material to fill in holes in the road. The depth of a road in
most places is proportional to the slope and to its age, a recently
laid-out road having only slight banks while an old road often
has vertical sides rising 15-30 feet above the wheel tracks.
These banks have a wealth of lichens, bryophytes, and fungi
during the rainy season (invierno) but usually dry out, except
in occasional moist shady spots, during the dry season. Con-
sequently most of the species are annuals and disappear quite
completely during the dry season. These banks are even more
conspicuous and important in the Subalpine Region.
The Subalpine Region probably includes most of the land in
Costa Rica above 1700 m., but is known botanically only on the
Cordillera Central and the northernmost mountains of the Cor-
dillera de Talamanca as far as the Cerro de las Vueltas. Pods,
Irazú, and Turrialba are largely cleared on their southern slopes
and used for dairy farms. Barba is still largely wooded and
inaccessible, but Standley and Maxon have explored the slopes
of the lower peaks of the Cerros de las Caricias, las Lajas, and
Zurqui. I also had the opportunity to spend two days on the
lower slopes of Zurquí. Standley and J. Valerio secured an
extensive series of lichens from Cerro de las Vueltas and neigh-
boring peaks within reach of Santa Maria de Dota, in 1925-6.
Various collectors have brought back small series of specimens
from Pods. Standley in 1924 and I in 1929 collected on the
south slope of Turrialba for a short period, and I spent about
three weeks at Guayabillos on the west slope of Irazü, as well
as climbing to the crater twice from the south side. Ever since
the time of Ørsted many collectors have brought in a few speci-
mens from lrazá. The alpine and subalpine areas of Irazü
[Vor. 20
382 ANNALS OF THE MISSOURI BOTANICAL GARDEN
are much less rich than the other mountains probably because
of the constant dust. On my last ascent I did not find even a
sterile thallus above 3000 m., while Ørsted reports lichens from
the summit. Botanically this Subalpine Region is the most
interesting, as it has a high percentage of Colombian and other
South American species, as well as most of the endemic species.
The Pacific Coastal flora is too poorly known to classify into
geographical divisions. I have been told by lumbermen that
there are many interesting local distributions of the forest trees
but these observations are too fragmentary for generalizations.
For present purposes we may subdivide into the province of
Guanacaste the peninsula of Nicoya, the great river valleys of
the R. Naranjo, R. General, and R. Grande de Térraba or
Diquís, and the Peninsula of Osa. Of these regions I have
personally visited only the first and the last. The peninsula
of Nicoya has not been explored at all as far as concerns lichens,
while our knowledge of the great river valleys is due solely to
two journeys by Pittier, Biolley, and Tonduz, in January and
February, 1891, and by Tonduz from October 18, 1891, to April
17, 1892 (see p. 388).
The province of Guanacaste consists of a low coast range
penetrated by the Bahías de Culebra and de Salinas, a broad
valley of the R. Tempisque about 40 km. wide and 60 km. long,
rising gently from sea level to about 200 m. and sloping again
gently to the Lago de Nicaragua, with Liberia, the provincial
capital, at its center, and the west slope of the Cordillera de
Tilarán (or de Guanacaste) on its east. Ørsted crossed the plain
from the Tempisque to the Bahía de Salinas and explored the
region of R. Sapoa and the coast, but I have seen no lichens from
this region. Pittier collected along the shore of the Bahía de
Salinas and may have penetrated the low coast range. Other-
wise it is unknown as far as lichens are concerned. "The plain
of the Tempisque is used for grazing with some sugar cane grown
in the southwest part in the vicinity of El Paso. The underlying
rock and soil in Liberia is white cascajo, a soft pumice laid down
in water and hence somewhat stratified. Further south there
are clay deposits. Rainfall is high during the rainy season but
the dry season is hot and dusty, unofficial temperature in the
1933]
DODGE—LICHENS OF COSTA RICA. I 383
shade rising as high as 108°F. in the hottest part of the day and
correspondingly cool at night. Wind velocities are high where
the trade winds sweep through the passes between the volcanoes
in the first half of the dry season, so that most of the perennial
vegetation is that of the arid regions and may account for the
similarity of the flora with that of Nicaragua and the north.
Here grass fires are even more frequent in proportion to the
population and more destructive, hence the region is seldom
visited by botanists; in fact I have failed to find any collections
from this region or any mention of it in botanical works. In
1930 I was able to spend about two weeks in the vicinity of
Liberia.
The western slopes of the Cordillera de Tilarán are used for
grazing, with small cafetales located in sheltered, moister areas.
'The vegetation does not appear much different from the plain of
the Tempisque, especially on the exposed slopes, but in the shel-
tered ravines more delicate types survive. This region has not
been visited by botanists to any great extent, although Standley
and J. Valerio collected in the vicinity of Tilarán in 1926, and
Valerio probably made studies of the local flora of Tilarán
while he was stationed there as a teacher some years ago. Be-
sides a week in Tilarán I spent about a week at 700 m. on the
slopes of Santamaría, the next peak south of La Vieja, and a
week at La Granadilla on the plateau between the R. de las
Cafias and the R. San José which culminates at 960 m. in the
Cerro de San José.
In Santamaría and Tilarán, even during the dry season, much
of the time the weather was characterized by mists and high
winds and much rain, while a short distance lower and to the
west there was neither mist norrain. However, the brief intervals
of hot sun are sufficient to keep the xerophytic vegetation on
the wind-swept prairie, while the sheltered ravines and shaded
hillsides have mesophytie types. At Tilarán we crossed the
continental divide into the headwaters of the Río Frío system
with a change in vegetation even more striking than in the pass
at Ochomogo in the Meseta Central.
The Cerro de San José rises sharply above the surrounding
peaks to 960 m., somewhat higher than the near-by peaks of the
[Vor. 20
384 ANNALS OF THE MISSOURI BOTANICAL GARDEN
continental divide. It seems to be composed of stratified
sedimentary rock sloping 40? S, 70° W. To the west rises the
lower exploded crater of Cerro Pelado, while to the east lie the
lower peaks which form the junction of the Cordillera de Tilarán
and the Cordillera Central. Though the region is considerably
drier than the same elevations on the Cordillera de Tilarán,
yet it maintains a luxuriant forest vegetation where it has not
been cleared for potreros. Being an isolated wind-swept and
fire-swept peak, the vegetation of the summit has a stunted,
subalpine appearance. The Hacienda Granadilla on the plateau
between the Río de las Cañas and the Río de San José at about
900-600 m. furnished much interesting material, especially as
we were fortunate in finding two huge pochotes (? Bombaz elliptica
HBK.) which had blown over so recently that the crustose
lichens were still fresh and the clusters of orchids and bromeliads
were still in bloom. These furnished about the only collections
to show the flora of the tree tops in Guanacaste.
The region between the Cerro de San José and Puntarenas has
not been visited by botanists to my knowledge, although it is
quite accessible on account of the Abangares and other gold
mines which have been exploited more or less continually since
colonial times. Since the region is of a different geological
formation it would probably well repay investigation, as would
also the calcareous Santa Catalina range between the R. Tempis-
que and R. Bebedero at the head of the Gulf of Nicoya.
The peninsula of Osa between Golfo Dulce and the Pacific
consists of low mountain ranges, probably of sedimentary rock;
at least such were the outcrops observed along the shore and in
the gorge of the Quebrada de la Laja, a tributary of the Río Nuevo.
G. Cufodontis, of the Austrian expedition of 1930, visited the
region in the immediate vicinity of Puerto Jiménez (Santo
Domingo, on many old maps and charts, on the small bay just
above Puntarenitas). Two weeks later I visited the same region,
also the Hacienda Sándalo between the Tigre and Terrones
Rivers (marked Necki on the charts). The coastal plain about
a mile wide slopes gradually from the beach to about 30 m. and
then more steeply up the spurs of the range which follows the
west coast of the Golfo Dulce. The forest is still virgin in most
1933]
DODGE—LICHENS OF COSTA RICA. I 385
places and consists of a rather open palm and hardwood forest
characteristic of well-drained areas. The coast between the tide
levels has a characteristic mangrove swamp. Mr. Dunlap,
of the United Fruit Company, visited this region a few years ago
but I have not learned of any specimens taken. Mr. H. J.
Marks, the proprietor of the Hacienda Sándalo, has sent a few
specimens of wood and herbarium specimens of forest trees to
Prof. 8. J. Record of Yale University.
Practically nothing is known of the flora of the islands off
the west coast. The only one which has been visited by bota-
nists, Isla Coco, has so far yielded very few foliose or fruticose
species. Dr. H. K. Svenson, botanist of the Astor expedition, one
of the latest to visit the island, describes it as follows: ‘‘ The lichen
flora must be very inconspicuous; I did not go out of my way to
look for lichens but if any large forms had been in my path, I
believe I would have picked them up. Cocos Island is entirely
covered by a rank vegetation which the sunlight rarely pene-
trates.” So far the only species reported are one Leptogium and
two Parmelias, all wide-ranging species of the tropical lowlands.
BoTANICAL EXPLORATION
Of early botanical exploration in Costa Rica the little that is
known apparently indicates that few or no cryptogams were
collected. Martin Sessé and José Mariano Mociño probably
visited the Pacific Coast between 1795 and 1804. The Sulfur,"
with George Barclay as botanist, entered the Golfo de Nicoya
about 1840 and about the same time Emmanuel Friedrichstahl
visited a portion of Nicaragua and Costa Rica though his collec-
tions were not carefully labeled as to locality, all the sheets being
marked Guatemala.
Between 1845 and 1851, the “Herald” probably visited
Pedregal, the harbor for Davíd, Panamá, as Seemann the botanist
brought back from Boquete and the neighboring voleano of
Chiriquí, lichens which were studied by Churchill Babington and
published by Seemann (1852-1857). These specimens are in
Babington's herbarium at Cambridge University and in the
Royal Botanic Gardens at Kew.
The first to make extensive collections was Anders Sandge
[Vor. 20
386 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Ørsted (1816-1872), to whom we owe the first elaborate accounts
of the natural history of Costa Rica. For details of his travels
and lists of plants collected at various places, see Ørsted (1848,
1851, 1863), Hemsley (1878-1888), and Durand & Pittier (1891).
He entered the country at Puntarenas, passing over the old
carretera which ascended the Cerros de Aguacate thence via
Alajuela to San José, where he made his headquarters for some
time. From this point he visited C. Escazú and Pacaca, pene-
trating to the mountains of Jarís to the south. He also visited
Candelaria, probably in the vicinity of Tablazo, judging from
the list of plants he collected in that locality. In May, 1847,
he explored Poás and Barba. Earlier in that year he had spent
some time studying the volcano Irazü, making his headquarters
in Cartago. From this point he visited Aguacaliente and Ujarrás
(probably the valley below the modern Paraíso), then made the
very difficult trip to Moin over the old trail now largely aban-
doned, passing Cervantes, Río Birrís, and Quebrada Honda to
Naranjo (the modern Juan Viñas), thence to Turrialba, keeping
along the shelf of land above the lake near Tunnel Camp, Bonilla,
ete. to the eustoms house on the R. Reventazón (probably below
the present La Junta) where he crossed the Reventaz6n in
canoes and proceeded along the coastal plain to Moin.
Another long trip was his visit to Guanacaste and adjacent
Nicaragua, the details of which were published in 1848 from a
letter he had sent home. He does not state in what year this
trip was undertaken, but since the mention that Puntarenas was
then a free port (this having been decreed in 1847) and since he
was visiting voleanoes in February and May, 1847, it is probable
that he made the journey in 1848. He was persuaded by govern-
ment officials to explore a possible route for an inter-oceanic
canal between the Lago de Nicaragua and the Pacific. The
party, headed by Francisco Gutierrez, left San José on February
17 for Puntarenas, where they took a huge canoe up the Gulf
of Nicoya and thence up the Río Tempisque. On March 8 they
set out with a pack train from Santa Rosa for the Hacienda
Sapoa just above the junction with R. Bolaños. Thence they
visited H. Las Animas above the junction with the R. Gua-
chepelin, ascending that valley and descending the R. Tortuga,
1933]
DODGE—LICHENS OF COSTA RICA. I 387
the next river east of the R. Sapoa. @rsted may have visited
some of the islands in the Lago de Nicaragua before he ascended
the Río Sapoa to the junction of the Sansapote and overland to
the H. Sapoa. Another trip was taken to the Golfo de Bolafios,
thence over the height of land and down the Sansapote to its
junction with the Sapoa.
Ørsted also visited San Juan del Norte (Greytown), crossing
the pass between Barba and Irazü at La Palma and descending
along the Río Sucio and the R. Sarapiquí. It is not known
whether he sailed from San Juan del Norte or from Puntarenas,
which was frequently visited by Danish vessels at that period,
thirteen having called there in 1853.
Warscewiez, coming from Guatemala via El Salvador and
Nicaragua, is said to have met Ørsted in Nicaragua before
reaching Costa Rica via the R. Sarapiquí. He evidently visited
Irazá and Turrialba, but I have seen no lichens from his collec-
tions.
Orsted's huge collections are to be found in Kjgbenhavn
where their study has only recently been completed. "They were
distributed about 1931. The fungi were sent to Elias Magnus
Fries at Upsala and were published promptly by him in 1851,
portions of the collections being found in both Kjøbenhavn and
Upsala. The lichens were sent to Th. M. Fries but only a single
number, Stereocaulon obesum 'Th. M. Fr., was published. Some
have since been named and inserted in the herbarium at Upsala.
Through the kindness of Professor N. Svedelius, the remaining
unnamed specimens were turned over to me for study and are
reported in this work. At present none of Orsted's lichens are
at Kjøbenhavn but it is probable that a complete set of duplicates
will eventually be deposited in that herbarium.
Ørsted was soon followed in Costa Rica by a number of natu-
ralists who have left us long accounts of geology and natural
history. Many resided in Costa Rica for several years, but few
appear to have collected lichens.
During his visit, Polakowsky collected a few specimens studied
by Nylander (1876) and apparently distributed quite widely,
as I have seen a complete set in Berlin and scattered numbers in
other herbaria.
[Vor. 20
388 ANNALS OF THE MISSOURI BOTANICAL GARDEN
A new era for Costa Rican natural history began with the
establishment in 1887 of the Instituto Fisico-geografico Nacional
de Costa Rica, with Henri Pittier as its first director. Due to
his enthusiastic explorations and those of his fellow botanist,
Tonduz, a large herbarium was assembled in San José and
materials were sent to many foreign collaborators for study,
resulting in the publication of Durand and Pittier (1891-1901).
The lichens which were studied by Müller Argau were largely
from two extensive expeditions to the southwestern portion of
the country and scattered numbers collected by others in various
parts of the country. Pittier (1891) described these expeditions
in detail. As these collections will be frequently mentioned I
have thought it wise to include an abstract of their itinerary.
On January 15, the first expedition reached San Marcos de
Dota, the capital of the canton of Tarrazü, the next night Santa
María de Dota, and the C. del Angelo, small valley of El Copey
1790 m., and El Roble 2670 m., on the following day. On
January 18 they passed Alto de la Baraja 2933 m., Rancho de las
Vueltas, C. de las Vueltas 3019 m., and Ojo de Agua 2760 m.;
on January 19 the Paramo de las Vueltas and the C. de Buena
Vista 3299 m., and La Muerte at 3130 m.; on the 20th, southward
along the slopes of Buena Vista to Lagunilla 1857 m., the Alto del
Palmital at 1211 m., reaching El Géneral on the evening of
January 21 where they stayed until January 28, 1891, and whence
Biolley returned to San José. Pittier and Tonduz continued to
Buenos Aires, Térraba, and Boruca, whence Pittier returned by
a somewhat different route to San Marcos and San José, arriving
on February 23, while Tonduz stayed in the field until March 8,
1891.
The following season Tonduz left San José on October 18,
1892, more or less retracing the journey of the previous season to
Buenos Aires, thence down the R. Grande de Térraba or Diquís,
stopping some time at Boruca, finally down the river to the coast
and back by sea to Puntarenas, stopping at several points,
arriving on April 17, 1893. The material from these two trips
furnished the bulk of material upon which Müller Argau (1891,
1893) based his account of the lichens.
'These collections are to be found in the Herbier Müller Argau,
1933]
DODGE—LICHENS OF COSTA RICA. I 389
which was purchased by the Herbier Boissier and is now at the
Université de Genéve. I have been told that a duplicate set
exists in Brussels, but apparently none was returned to the Museo
Nacional in San José.
C. Wercklé, an engineer who resided in Costa Rica for many
years, collected much in the vicinity of Cartago. His specimens
were determined by Lindau and may be seen in Berlin and in
Wercklé’s herbarium at the Museo Nacional de Costa Rica.
Wercklé was not at all discriminating in selecting samples for
Lindau, so that most of the material in San José needed revision.
The Calverts (1917) spent a year, 1909-1910, in Costa Rica
and have given us a good account of its natural history. They
brought back some plants which are in the herbarium of the
University of Pennsylvania, but I have seen only one lichen from
them (at the Farlow Herbarium). Maxon and assistants have
visited Costa Rica for ferns and brought back some lichens
which were sent to G. K. Merrill for study and are now to be
found in the Farlow Herbarium (which acquired the Merrill
herbarium) and in the National Herbarium of the Smithsonian
Institution at Washington.
Paul C. Standley, in 1924 and again in 1925-6, in company
with Juvenal Valerio Rodriguez and Ruben Torres Rojas, made
very extensive collections of lichens although his primary interest
was angiosperms. These lichens were likewise sent to G. K.
Merrill who was studying them at the time of his death. As
curator of the Farlow Herbarium I continued this study in
preparation for my trip to Costa Rica and am here reporting
these collections.
In 1925, during a two-weeks’ visit, I secured a large amount of
material from Cerro Carpintera and again in 1929-30 I secured
several thousand specimens of cryptogams which are still being
studied. The most complete set of these plants is in my personal
herbarium and the next set, lacking only a few numbers of the
commoner species, is at the Farlow Herbarium.
Karl Danielson, an assistant to H. E. Stork, in 1928 collected
a few numbers of lichens which are now at the University of
Michigan. These are also reported here for the first time,
duplicates being found in the Farlow Herbarium and in my own
herbarium.
[Vor. 20
390 ANNALS OF THE MISSOURI BOTANICAL GARDEN
In 1920 and 1930 the Costa Rican government botanist,
Alberto M. Brenes, collected many lichens in the vicinity of
San Ramón and a few from other parts of the country. These
were also turned over to me for study and are to be found in the
Museo Nacional, with duplicates in my own herbarium and in the
Farlow Herbarium.
The Oesterreichische Costa Rica Expedition, under the leader-
ship of Otto Porsch (1932) with G. Cufodontis as assistant
botanist, spent the late verano and early invierno in Costa Rica,
visiting much the same territory as I did. If lichens were
collected they were probably studied by Zahlbruckner but I
have seen no publication on this group, although many of the
other groups are already published.
FLonisTICS
It is too early to draw any very definite conclusions regarding
the distribution of plants and the affinities of the flora, but
certain typical distributions are evident.
Families and genera which are abundant in the northern
United States and are often widespread in the North Temperate
Zone are usually found at the higher altitudes and reach their
southern limits in the mountains of Colombia, except for oc-
casional isolated species farther south. Such an example is
the Peltigeraceae, which are widespread and abundant in the
northern part of the United States. Only two genera, Nephroma
and Peltigera, reach the American tropics. Nephroma is rep-
resented by a single species seen in only one restricted locality
in Costa Rica. Peltigera is represented by about eight species,
mostly strictly tropical with only one undescribed species reaching
Magellanes, Chile.
Another typical example is a northern family, which is char-
acteristic of the Atlantic Coastal Plain, the Collemaceae. Here
genera are mostly common to the eastern and western hemispheres
but subgenera and species are much more restricted in distri-
bution. These are mostly found on the coastal plains of Costa
Rica, being characteristic on the plains of Guanacaste and
occasional on the Atlantic Coastal Plain but not yet reported
from Panama. The species belonging in these genera from the
1933]
DODGE—LICHENS OF COSTA RICA. I 391
higher elevations have relations with those of the mountains
of Colombia and not with those north of the Isthmus of Nicaragua.
A third type of distribution is found in those families which
are predominantly southern, as in the Stictaceae. Here we find
them highly developed and characteristic in the highlands,
relatively rare in the lowlands. The genera and subgenera are
world-wide in distribution and the species also have wide ranges,
becoming less frequent in numbers and in species in the North
Temperate Zone. In this group, however, species of the moun-
tains of Costa Rica may extend into North America along the
Atlantic Coastal Plain, even as far as the Avalon peninsula of
Newfoundland, while the species of the Coastal Plain in Costa
Rica are southern in their ranges.
Finally some cosmopolitan families in which it is difficult
to determine a center of distribution have genera or subgenera
which are strictly limited to tropical America.
Further consideration of problems of distribution may be
deferred until the systematic enumeration is complete. In
general the affinities of the flora seem to be with eastern Brasil
but since we have very few collections available from the inter-
mediate mountain ranges of eastern Colombia and southern
Venezuela and British Guiana, it is likely that stations of many
of these species will eventually be discovered in these regions.
Relatively few species are common to Mexico and Costa Rica
or to Perú and Costa Rica, and practically no Costa Rican species
extend as far south as Chile.
In the following systematic enumeration I have included in
the keys to species all those which have been described from
tropical America, whether they have been found in Costa Rica
or not. I have not included species which were not originally
described from this area, although they may have been reported
from it, since often these reports have been based on misdeter-
mined specimens. While it is evident that political boundaries
do not always coincide with natural floristic divisions, for the
purposes of this study tropical America may be defined as
Mexico and the West Indies southward to Perú, Bolivia, Para-
guay, Brasil, and Uruguay. These are quite natural in the pres-
ent state of our knowledge, as the lichen flora of northern
[Vor. 20
392 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Mexico which might be expected to show affinities with that of
the southwestern United States is practically unknown. Simi-
larly, that of northern Argentina, which might be expected to be
similar to that of adjacent Paraguay, has been little explored
for cryptogams. Northern Chile, largely desert, has also been
little studied, while the flora south of the desert has relatively
few species which have been reported from tropical America as
here defined. The greatest difficulty occurs in the exclusion of
southern Florida where occur many species which are tropical
rather than northern in their affinities. Fortunately for our
purposes comparatively few species have been described from
this region.
KEY TO TROPICAL AMERICAN FAMILIES
O
FOLIOSE AND FRUTICOSE LICHENS
Apothecia more or less exposed, paraphyses growing above the asci to =
a tissue, asci early evanescent, leaving an enclosed — spore mas
Gidea um).. ° CONIOCARPINEAE
Thallus foliose or -frutiooss, corticate, apothects enel. —
Apothecia linear, elongate-elliptic or angular, not forming à a ‘masadic
cu da dad Cannas celcy i040 kine od ede E EE A NE GRAPHIDINEAE
Thallus fruticose, erect or dependent, with a basal layer attached to the
substrate, corticate [not yet reported between Lower California and
Perú]; See rr ee er en eee ree ree RoccELLACEAE
Apothecia poten not forming a mazediu ..CYCLOCARPINEAE
Thallus strictly crustose, but aang piede appearing fruticose
owing p the proliferation of the apothecia from the margins, easily
mistaken for a hepatic on cursory examination; Guiana........ Polystroma
Thallus loosely byssine from filaments of Cladophora or Trentepohlia, apo-
thecia with light-colored parathecium.................. COENOGONIACEAE
Thallus filiform or dwarf fruticose, or squamose, very — iam foliose
[deferred for further treatment with the crustose speci
gr symbiont Scytonema or Stigonema, apothecia more or m sunk
llus, small and easily overlooked.................. EPHEBACEAE
Algal avoddon ot DIR RP ENERO ECRES PYRENOPSIDACEAE
Algal symbiont Révularía......... sese LICHINACEAE
Thallus gelatinous, swelling greatly when moistened, dwarf fruticose,
squamose or large foliose, algal symbiont usually Nostoc........ OLLEMACEAE
Thallus definitely foliose or fruticose, or if small, the form of the thallus
not due to the algal symbiont.
Hypothallus and rhisoids highly — thallus squamose to small
foliose, upper gal symbiont usually Nostoc. eae
Wypothallus evanescent.
Thallus large foliose, algal symbionts Nostocaceae or Palmellaceae,
spores fusiform to acicular, 2 or more cellec
1933]
DODGE—LICHENS OF COSTA RICA. I 393
Both surfaces corticate, lower surface interrupted by more or less
highly developed breathing pores, apothecia with well-de-
veloped parathecia or amphithecia, stipitate or sessile... . .STICTACEAE
Only upper cortex developed, the lower surface tomentose with
more or less highly developed network of veins and tufted
rhizoids, apothecia attached to the thallus over their whole
under surface without true parathecium or amphithecium
aad surrounded fe the torn remains of the tissue cover-
ing the young apothecium...................00005 PELTIGERACEAE
Thallus oni or small foliose, only upper surface corticate, algal
symbiont Pleurococcus, ups ecia is well-developed light-
colored parathecium, spores va
No secondary thallus or podetia pu apothecia sessile. ......
t ope op OST HYLLOPSORACEAE
Secondary thallus or podetia present, varying from simple sta
apothecium without algae to highly developed branched
or infundibuliform structures, primary thallus squamose or
small foliose, hie evanescent or degenerate, perhaps crus-
tose in qome species s on Taal eds sero oe LADONIACEAE
Thallus large foliose or Ee. algal symbiont Pleurococcus, spores
ne- or two-celled or muriform, never elongate-fusiform to
acicular.
Apothecia with black parathecium, spores one-celled [in Gyrophora
aplocarpa from Pert, the only species of the family so far re-
Orted|. c anaa e EDD RUNE a ae GYROPHORACEAE
ieee with bright-colored MESS ds
Spores not placodiomorphou
Thallus PEON ai piga Rae PRAE PARMELIACEAE
Thalis Irulioo8b. ..... zx e E TUE. USNEACEAE
Spores dimora
Spores hyaline, thallus usually bright yellow, at least the
epi THE
CIUM OTaANgE « eoim Se G ELOSCHISTACEAE
Spores brown, balie usually more or less glaucous, epithe-
Mum brown OF DrUInoso. o NEL e HYSCIACEAE
SPHAEROPHORACEAE
Thallus foliose or fruticose, corticate on both surfaces or the
lower surface incompletely corticate, with Protococcus. Apo-
thecia sessile on the margin, or on the lower side of the thallus,
open at first or enclosed by an amphithecium.
Only two genera have been reported from tropical America,
both fruticose with solid axis and terminal apothecium. Acro-
scyphus sphaerophoroides Lév. without an amphithecium has
been reported from Mexico and Perú but has not been found in
Costa Rica. Sphaerophorus is typically Subarctic-Antarctic
in distribution, coming southward on the higher mountains with
[Vor. 20
394 ANNALS OF THE MISSOURI BOTANICAL GARDEN
a single species on the higher mountains of tropical America.
The whole family is greatly in need of revision.
SPHAEROPHORUS MELANOCARPUS (Sw.) DC. apud Lam. & DC.,
Fl. Franç. ed. 2, 6: 178. 1805, excl. syn.
Lichen melanocarpus Swartz, Nova Gen. Sp. Pl. Prodr. 147. 1788.
Type: Jamaica, Swartz.
Thallus ashy glaucescent or pale above, pale below, partly terete,
partly compressed, smooth, not rugose, highly branched, 1-6 cm.
high, primary branches 1-4 mm. thick, KOH yellow above,
KOH - below, CaOCL, — ; cortex 20 y thick, subpellucid, of
thick-walled conglutinate irregularly woven hyphae; medulla
I —, composed of hyphae 4-8 u thick; mazedium oblique in
the ends of the thicker terete branches, becoming disciform,
exciple lacerate, ascospores 7-11 y in diameter. Spermatogonia
immersed in the tips or in the lower surface of the branches,
opening by a blackened verruciform mouth; spermatia oblong,
1 x 3 y [description from Nylander and Vainio].
The Costa Rican material does not altogether agree with the
above description, but there is so much variation in macroscopic
appearance that only a careful monograph can settle the number
of species and their relationships.
Cartago: F. V. Turrialba, 2000-2400 m., Standley 35313.
San José: L. la Chonta, n. e. Sta. María de Dota, 2000-2100 m., Standley 42290;
C. de las Vueltas, 2700-3000 m., Standley & J. Valerio 43832.
Heredia: C. Zurquí, 2000-2400 m., Standley & J. Valerio 50436.
ROCCELLACEAE
This strictly maritime family has not yet been reported between
Lower California and Perú, probably because no suitable mari-
time habitats have been visited by botanists, for the group is
abundant in such strictly tropical habitats as the Galápagos
Islands. I did not have an opportunity to visit such myself, but
had I been able to visit the cliffs along the various headlands
between Puntarenas and Golfo Dulce, it is possible that I might
have secured some representatives of this family.
COENOGONIACEAE
Thallus spongy byssoid, either adnate or forming dimidiate
shelving masses, homoeomerous with Trentepohlia or Cladophora,
1933]
DODGE—LICHENS OF COSTA RICA. I 395
whose filaments are partially surrounded by hyphae. Apothecia
with pseudoparenchymatous parathecium; asci 8-spored; spores
hyaline, one- or two-celled; spermatia exobasidial.
Thallus with Trenlepohlia ee ee JE MI Coenogonium
Thallus with Cladophora, apothecia unknown ............. sess. Racodium
COENOGONIUM Ehrenb.
CorNocontum Ehrenberg apud Nees ab Esenbeck, Horae
Physicae Berol. 120. 1820
Thallus loosely spongy byssoid, either adnate or forming
dimidiate shelving masses (suggesting a thin species of Poly-
stictus), homoeomerous with Trentepohlia which is partially
surrounded by a network of hyphae. Apothecia scattered on
the upper surface, scutiform, usually with a short stipe, with a
parathecium of thin-walled pseudoparenchyma without a me-
dulla; paraphyses unbranched, often with swollen tips; asci 8-
spored; spores hyaline, fusiform, ellipsoidal to elongate, one- or
two-celled; spermagonia spherical, spermatia exobasidial, fusi-
form, straight; spermatiophores mixed with anaphyses.
Clements segregated the species of Coenogonium with uni-
cellular spores as Holocoenis. However, his proposal has not been
followed by other authors, although Vainio and Zahlbruckner
have proposed sections on this basis, Vainio using Coenobiatora
and C torina, while Zahlbruckner incorrectly adopted
Holocoenis and Coenobiatorina.
This family is very aberrant among lichens in several respects
and perhaps would be better dropped and the species distributed
among the fungi and algae. It seems more logical to regard the
group as a case of parasitism of fungi on algae, as neither fungus
nor alga seems to influence the development of the other, and
there has been no further evolution resulting from the acquisition
of a photosynthetic unit by the fungus. Taking this view, one
would recognize about four or perhaps five species of fungi from
tropical America, one species with unicellular spores about
6-10 x 2.5—4 y. belonging in Patinella; and three or four species
with 2-celled spores belonging in Orbilia or a segregate from that
genus. Among the algae one would recognize a dozen or more
species of Trentepohlia which are often subject to the attacks
[Vor. 20
396 ANNALS OF THE MISSOURI BOTANICAL GARDEN
of Patinella and Orbilia. In the following key to the tropical
American species of Coenogonium, as it is commonly understood
by lichenologists, it will be noted that out of nineteen names
proposed, four were based on specimens in which no apothecia
were found. Four other species originally described in Coeno-
gonium have already been transferred to Trentepohlia. I have
earefully investigated the ascocarps of the fertile Costa Rican
specimens and can find few characters which would separate
species from each other, except spore size and septation. For
lack of time to investigate this question thoroughly, I have here
followed the traditional arrangement, as to do otherwise would
involve serious nomenclatorial difficulties. To adopt the view
that this group should be excluded from the lichens would
invalidate Coenogonium, as based on a combination of characters
of parasite and host and involve redescription of its species in
Patinella, Orbilia, and Trentepohlia.
KEY TO THE TROPICAL AMERICAN SPECIES OF COENOGONIUM
Apothecial dise livid, cell walls aliis spores unicellular; algae 11-16 u
in diameter; French Guiana. ............. 0.0 ccc eee eee eee ee ees C. Leprieurit
Apothecia carneous, yellowish or waxy white, never livid.
Thallus dimidiate, not adnate to the substrate
Thallus thick, rigid, with white subsilky villum, zonate, sterile; Brasil. .
"EE eee eee Te eT rer ey Terre re . Echinus
Thallus thinner.
Filaments 7-9 & in diam., cells not very distinct, greenish, sub-
iffuse, WAROROIE So oi dics bie oy ERE PRAE REaMESA PERF subvirescens
Filaments 12 u in diam., rigid: BINE 5554044400944 REPE AU RR C. Linkit
Filaments thicker, 16-20 » in diam., cen m rigid.
Cells 4-5 times as long as broad; Mexico................. C. confervoides
Cells about twice as long as broad; spores T. 8.5 x 2-3 u; Brasil. .
————————————— eee C. aorocephalum
Filaments and paraphyses thick, imperfectly known............. C. andinum
Thallus adnate to effuse
Filaments moniliform, 18 4 in diam.; thallus glaucous green to tawny
brown, margins paler; spores 2-celled, 3-4 times as long as
broad: CUDA: 1422453 09 X493 EEEIEE SOREN ANNETTA C. rica ais
Filaments partially moniliform, partly cylindrical, cells 20-
m.; € M Luo oa ERUDEERCOIERR TE SASL ES C. Missis
Filaments uniformly ia
Filaments 23-36 u in diam.; Sion apothecia 0.6-0.7 mm. in
diam.; spores u- 15 x 25-35 u OBOE okt ies Se yn C. disjunctum
filis 20-28 uw in diam.; ijon niae A 6-10 x3 à;
or-
bonia or ionii —— —— PIT E . interpositum
1933]
DODGE—LICHENS OF COSTA RICA. I 397
Filaments 17-20 y in diam.; cells 30-50 » long; spores 8 x 2.5 u, 2-
edad: Costa Bi .....eoeoe eorr err ERAS P E C. interponendum
Filaments b m in Pc thallus ashy green, pulvinate, n^ to 2
eu DONE TIG. eie ores rre S vL iiio
Filaments 12-16 n in pon thallus yellowish; spores 8-10 x 3 u
Meek Kaan 85:5 0k os os ee 1. i C. interplerum
hace 12-14 yw in diam.; thallus yellowish to green; St. Vince
DCN SY es errr ere ee ee eprieurii v. panne
Filaments 8-11 » in diam., conglutinate in fancidion 70-90 u
diam.; e 7-10 3.5-4.5 y; Bolivia. -cees vats ss ade a C. paie ans
Filaments 4-8
Filaments age aps spermatia 7-9 x 15 u; paraphyses cla-
o PAR, oon ee OS eee C. dialeptizum
Filaments articulate; spermatia not reported.
Cells 2.5-4 times as long as broad, paraphyses obovoid;
C.
aiT AEE E esa EE ee ae ee rn pannosum
ells 1.5-3 times as long as broad, sterile; Brasil......... C. depressum
Filament size not given, rugulose, articulate-branched, coalesced
into fascicles; Venesuels..... ... <0 610 s00 eto RR VH C. Tuckermani
Cornocontum LEPRIEURII Nyl., Ann. Sci. Nat. Bot. IV, 16:
89. 1862.
C. Linkii var. Leprieurii Mont., Ann. Sci. Nat. Bot. III, 16:
47. 1851
Holocoenis Leprieurii Clements, Genera of fungi, 174. 1909.
Type: Guayane Française, Leprieur, sine no.
Algae loosely woven, yellowish, 11-16 u in diameter. Apo-
thecia plane or convex, excluding the white margin, dise livid
from the first; asci about 20 u in diameter; spores oblong to
fusiform or ellipsoidal, unicellular, 6-10 x 2.5-4 u; paraphyses
slender, apices clavate.
I have not seen this species from Costa Rica but Dodge &
Nevermann 7399, from jungle at Castilla farm, 20 m., Limón
Province, seems to be a variety of this species. The fungus is
not well developed but has a livid disc, the margin is still thick
and white, and the alga is of much smaller diameter. The
exciple is composed of isodiametric cells with colored walls. It
appears to be the same fungus attacking a much smaller alga,
and should be referred to Patinella.
COENOGONIUM SUBVIRESCENS Nyl., Flora 57: 72. 1874.
C. Leprieurii var. subvirescens Nyl., Ann. Sci. Nat. Bot. IV,
16:89. 1862.
Type: Brasil, Amazonas, Rio Negro, Spruce 28, is type of the
[Vor. 20
398 ANNALS OF THE MISSOURI BOTANICAL GARDEN
species, while C. Leprieurii var. subvirescens was described from
rench Guiana without collector.
Algae somewhat interwoven but more or less parallel, 7-9 y,
forming a loose dimidiate to subcircular thallus depending upon
position and shape of substrate, not or obscurely zonate, pale
to dark olive buff, cells about 21-24 y long, septa not very evident,
other cell walls thick, chloroplasts appearing somewhat in the
shape of a dumb-bell with the pigment more or less massed at
the ends of the cells.
Apothecia about 800 y. in diameter, tapering below to a short
stipe about 250 y. in diameter, composed of thick-walled pseudo-
parenchyma with the outer 40 u composed of a palisade of elongate
cells, perpendieular to the surface; parathecium about 80 y
thick, homogeneous with the rest of the apothecium, the hypo-
am very thin, filamentous, of cells with thinner walls. The
hymenium is about 80 y tall and is composed of filiform para-
physes with clavate tips and slender clavate asci. The asco-
spores are not mature in my specimens but seem to be slender,
2-celled, probably about the size of most of the other spores in
the genus except C. disjunctum.
Limén: Cadiz, Dodge 7452; Castilla farm, 20 m., Dodge & Nevermann 7451; Monte
Verde, K. Danielson 73; Siquirres, I m., Dedin Catt & Thomas 5574; La
bier 70-80 m., Standley 368
Car : R. Pejivalle, 650-800 m., “re & Thomas 4408; El Muñeco, on R.
aan 1400- 1500 m., Standley & J. Valerio 61055; C. Carpintera, 1500-1850 m.,
Standley 35596.
Puntarenas: Osa, Golfo Dulce, Dodge 7450.
Guanacaste: Q. Serena near Tilarán, 700 m., Standley & J. Valerio 46242; Santa-
maría, 720-850 m., Dodge 7015.
CoENoGONIUM Linxir Ehrenberg ap. Nees ab Esenbeck,
Horae Phys. Berol. 120. 1820.
C. controversum Pers. ap. Gaudich. in Freycinet, Voy. Uranie,
Bot. 214. 1826 (nom. nud.).
Type: Brasil, Sta. Catharina, Chamisso. The type of C.
controversum came from the walls of the aqueduct of Corcovado,
Rio de Janeiro, Gaudichaud.
Algae soriewliag interwoven but more or less parallel, about
12 v in diameter, forming a somewhat rigid dimidiate to sub-
orbicular thallus, not or obscurely zonate, pale to dark olive-
buff, cells 40-60 y. long, septa very difficult to observe, chloro-
1933]
DODGE—LICHENS OF COSTA RICA. I 399
plasts small and scattered, outer cell walls thick, partially covered
with anastomosing fungal hyphae.
Apothecia warm buff, about 850 y in diameter, tapering
sharply to a short stipe about 240 y. in diameter and up to 300 y.
long, pseudoparenchymatous, without palisade layer at the sur-
face; parathecium about 100 u thick, homogeneous with the rest
of the apothecium, the hypothecium very thin and soon evanes-
cent; the hymenium about 80 u tall, composed of filiform para-
physes with swollen tips and cylindrical asci. The ascospores
are not fully mature in my specimens but probably the common
size for the genus.
Cartago: C. Carpintera, 1800 m., K. Danielson 9.
Alajuela: La Palma de S. Ram in. 1100 m., Brenes 175.
Guanacaste: El Silencio, near Tilarán, 750 m., Standley & J. Valerio 44619.
COoENOGONIUM CONFERVOIDES Nyl., Flora 41:380. 1858.
? C. andinum Karsten ap. Nyl., Bot. Zeit. 20: 178. 1862.
Type: Mexico, Orizaba, Fr. M üller (Schimper Herb.). Type of
C. andinum from Colombia, 2000 m., Lindig 2560, also Perú,
Weddell, and southern Brasil, Guillemin. Since I have been
unable to see the types, the interpretation of this species is
difficult. In Ann. Sci. Nat. Bot. IV, 11: 242. 1859, Nylander
amplified his very inadequate description, basing his emendations
on a specimen from Tahiti, Lépine 14. When he monographed
the genus in Ann. Sci. Nat. Bot. IV, 16: 91. 1862, he cited
Lepine 14 from Tahiti first, adding Guadeloupe Island, Duchas-
saing, Brasil, Gaudichaud, and Weddell, and Chile, Gay, not
citing the original Mexican type. Since none of these descriptions
was based on specimens having apothecia its application is
further confused. Since the descriptions only state how it
differs from C. Linkii, I have assumed that the thallus is more or
less dimidiate with a larger alga than in that species. Nylander
also notes that a specimen from Colombia, Bogota, 2700 m.,
Lindig 887, has algal filaments 12-18 » in diam. Unfortunately
I have been unable to see any of the above-mentioned specimens.
The following description is based on Costa Rican specimens.
Algae somewhat interwoven but more or less parallel, 16—20
(-28) u in diameter, forming a soft dimidiate or suborbicular
thallus, not zonate, pale to deep olive buff, cells about 60-100 y.
[Vor. 20
400 ANNALS OF THE MISSOURI BOTANICAL GARDEN
long, septa plainly visible, chloroplasts rather indistinct in these
specimens, often alternate cells more or less collapsing, giving
a somewhat moniliform appearance to old filaments, outer walls
thick, not densely covered with hyphae. The collapsed cells
sometimes have the coloring matter collected at the ends, giving
the dumb-bell appearance to the chloroplast as in C. subvirescens.
Apothecia warm buff, 560 x in diameter, patelliform, with
stipe about 160 u in diameter and 300 ų long, pseudoparenchymat-
ous, of very thick-walled cells which tend to arrange their long
diameters perpendicular to the surface of the apothecium, but
not forming a definite palisade as in C. subvirescens; the para-
thecium 40-50 x thick, homogeneous with the rest of the apo-
thecium, the hypothecium 20-30 y. thick of interwoven hyphae,
the hymenium about 60 u tall, composed of filiform paraphyses
with spherical tips and slender cylindrical asci about 5 y in di-
ameter; ascospores distichous, 4-celled, 10-12 x 3 u. [The asco-
spores are immature and so closely packed in the ascus that I
am not certain of size and septation, as I have been unable to
free spores from the ascus.]
Limón: Waldeck, Dodge 7453.
Cartago?: Morpho Valley, 1420 m., K. Danielson 51 [This locality is uncertain and
Danielson's barometrie readings are apt to be a little high, but it probably in the
vieinity of the upper Reventazón below Cartago, where Lepidoptera of the genus
Morpho are not uncommon.]
COENOGONIUM HETEROTRICHUM Müll. Arg., Bull. Soc. R.
Bot. Belg. 32: 162. 1893.
Type: Costa Rica, San Marcos de Dota, Tonduz 6115.
Thallus pulvinate, cespitose-effused, deep olive buff to dark
olive, algae somewhat interwoven, dimorphic, larger filaments
20-25 y. in diameter, cells about 40 u long, chloroplasts disciform,
scattered; smaller filaments submoniliform, the cells of smaller
diameter being about 7-8 y. and 12 y. long, those of larger diameter
being about 10-12 y. and of about the same length.
Fungus unknown. In Brenes 29a the fungal hyphae are
brown instead of hyaline.
San José: S. Marcos de Dota, 1200 m., Tonduz 6115, type.
Alajuela: La Palma de S. Ramón, 1250 m., Brenes 29a.
COENOGONIUM INTERPONENDUM Nyl. ap. Polakowsky, Jour.
Bot. Brit. & For. 15: 225. 1877.
1933]
DODGE—LICHENS OF COSTA RICA. I 401
Type: Costa Rica, Cartago, Angostura, H. Polakowsky 496.
Algae loosely interwoven, forming an adnate irregular thallus,
deep olive buff to citrine drab, cells about 17-20 y. in diameter,
30-50 u long, pigment quite uniformly dispersed throughout
the cell, yellowish green. All my Costa Rican material is sterile.
Nylander states that the spores are fusiform, 2-celled, 8 x 2.5 y.
Limón: Carmen, Dodge 7449.
Cartago: Angostura, H. Polakowsky 496, TYPE.
Alajuela: La Palma de S. Ramón, 1250 m., Brenes, 53a, 116, 3
Guanacaste: H. Q. Azul on lower slopes of v. Tenorio, 400- she m. l Dédye & Thomas
042.
CoENOGONIUM INTERPOSITUM Nyl, Ann. Sci. Nat. Bot. IV,
16:91. 1862.
Type: Borbonia, Lepervanche, Meziéres; Louisiana, Hale.
Thallus loosely tomentose intertangled, more or less pulvinate,
algal cells 20-28 u in diameter and 60-88 y long, chloroplasts
appearing the shape of a dumb-bell with the pigment massing
at the ends of the cells.
[Apothecia pale carneo-luteous; spores oblong, simple, 6-10
x 3 u; paraphyses slender or medium.]
As all my Costa Rican material is sterile the reference of it
to this species is doubtful. It quite possibly belongs with C.
disjunctum Nyl., based on specimens from Martinique and Cuba,
Wright 170, which reaches a slightly larger diameter of algal
filament and much larger 2-celled spores. Both species are said
to resemble C. confervoides and perhaps should be placed in the
C. confervoides group.
Cartago: C. Carpintera, 1560-1700 m., Dodge & Thomas 7920.
Alajuela: La Palma de S. Ramón, 1100-1250 m., Brenes 53, 115.
COENOGONIUM INTERPLEXUM Nyl., Ann. Sci. Nat. Bot. IV,
16: 92, pl. 12, f. 20, 21. 1862.
Type: Colombia, 2200 m., Lindig 2561.
Thallus loosely interwoven, intricate, algal cells 12-16 » in
diameter and 30 y long, chloroplasts more or less disciform,
scattered.
[Apothecia fleshy-yellow, plane, 1 mm. in diameter, margin
waxy, fleshy to white; spores short-fusiform, 2-celled, 8-10 x
3 u; paraphyses medium, apex clavate.]
Alajuela: Piedades de S. Ramón, 900 m., Brenes 408.
Puntarenas: Boruea, 560 m., T'onduz 6114.
[Vor. 20
402 ANNALS OF THE MISSOURI BOTANICAL GARDEN
COENOGONIUM PANNOSUM Müll. Arg., Flora 64: 234. 1881.
Type: Brasil, São Paulo, Apiahy, Puiggari 1026.
Thallus loosely interwoven, dark olive buff to citrine drab,
more or less adnate to substratum, algal cells 6-8 » in diameter,
28-32 y long, pigment migrating toward the ends of the cells.
[Apothecia 350-500 y. in diameter, pale or orange white, plane,
with paler and thinner margin, finally immarginate and slightly
convex, becoming more flesh color; hymenium hyaline, para-
physes very slender with obovoid head about three times as
thick as the stalk; asci slender, cylindrical, 8-spored; spores 2-
locular, 6-7 x 2 y, fusiform with acute ends.]
Since my Costa Rican material is sterile its reference here is
uncertain and it may be only very young algae of C. subvirescens
Nyl., with which it agrees in general appearance, but there is no
trace of the formation of a dimidiate thallus.
Guanacaste: H. Q. Azul, on slope of V. Tenorio, 500-600 m., Dodge & Thomas 6654.
COENOGONIUM DEPRESSUM Müll. Arg., Flora 64:525. 1881.
Type: Brasil, São Paulo, Apiahy, Puiggari 1034.
Thallus adnate, pale olive buff, filaments short, flexuous,
depressed, subintricate; cells 4-8(-13) y in diameter, 16-20 y
long, somewhat inflated in the middle but scarcely enough to
give a moniliform appearance, chloroplasts no longer distinct.
As I have not compared the Costa Rican material with the
type I cannot be certain of the identity.
Cartago: alpine region of Irazüá, Ørsted.
COENOGONIUM IMPLEXUM Nyl., Ann. Sci. Nat. Bot. IV, 16:
92. 1802.
Type: Australia, Victoria, Jarvin, Ferd. M üller.
[Similar to C. interplexum but algal cells a little smaller, 10-13 y
in diameter, spores a little larger, 8-11 x 3.5-4.5 u, paraphyses
thicker.]
The following collection has been referred here by Müller
Argau, but I have not studied it.
Puntarenas: Boruca, 560 m., T'onduz 6118.
EPHEBACEAE
Thallus dwarf fruticose, branched, more or less filiform,
without rhizinae, crustose or small squamose, with Scytonema
1933]
DODGE—LICHENS OF COSTA RICA. I 403
or Stigonema. Apothecia small, often with very small puncti-
form disc, scarcely visible; paraphyses well developed or absent;
asci 8-spored; spores hyaline, 1—2-celled.
The individuals of this family are so small that they are easily
overlooked both in collecting and in sorting material, and so the
few species here reported are probably not representative of this
group in Costa Rica.
KEY TO TROPICAL AMERICAN GENERA OF EPHEBACEAE
Thallus crustose to small squamose, homoeomeric; Brasil........ Pterigyopsis atra
Thallus dwarf fruticose, dark, thickly branched.
Apothecia sunken in swellings of the thallus, single or gregarious; spores
unicellular; paraphyses present; Brasil.....................lsuu. Ephebeia
Apothecia sessile on the thallus, either lateral or terminal.
Thallus without pseudoparenchymatous cortex or medulla; paraphyses
filiform, simple; asci 8-spored; spores ovoid or spherical, unicellular
Fu M cnc De TIME ce. UE Ade Thermutis
Thallus with pseudoparenchymatous cortex and medulla.
PENNE hv e or REP MISSAE EPA REND Leptogidium byssoides
Loc 2 bono. C oe 585s 0 OS Rh a Oe ea 0 ee Polychidium
THERMUTIS E. Fr.
THERMUTIS E. Fr., Syst. Orb. Veg. 1: 392. 1825.
Gonionema Nyl., Mem. Soc. Sci. Nat. Cherbourg 3: 163. 1855.
Thallus dwarf fruticose, thickly branched and filiform, without
rhizinae, Scytonema present with the hyphae running in the
gelified sheath. Apothecia small, lateral, saucer-shaped to
almost spherical; parathecium often highly developed and thick;
hypothecium light-colored; paraphyses unbranched, filiform,
tips not swollen; asci clavate, thin-walled, 8-spored; spores
hyaline, ellipsoidal, elongate, unicellular, thin-walled. Sperma-
gonia lateral or terminal, sessile, more or less spherical; spermatia
small, ovoid, or elongate.
THERMUTIS VELUTINA (Ach.) Fw., Linnaea 23: 170. 1850.
Lichen velutinus Ach., Lichenog. Supe: Prodr. 218. 1798.
Gonionema velutina Nyl., Act. Soc. Linn. Bordeaux 21: 262.
1856.
This species has been reported by Müller Argau from the
following localities in Costa Rica. The specimens were not
studied while I was in Genève.
San José: San Marcos de Dota, 1200 m., Tonduz 5378.
Guanacaste: Boruca, 560 m., T'onduz 5371; Térraba, Tonduz 5372.
[Vor. 20
404 ANNALS OF THE MISSOURI BOTANICAL GARDEN
A small sterile specimen from Guanacaste: near Tilarán, 500-
690 m., Dodge & Thomas 8081, may belong in Polychidium but
the specimen is not definitely determinable as to genus.
COLLEMACEAE
Thallus gelified, crustose to foliose or dwarf fruticose, with or
without rhizoids, sometimes umbilicate, homoeomerous with
Nostoc. Apothecia from sunken almost perithecia to sessile
apothecia usually with an amphithecium, occasionally with
parathecium; paraphyses simple; asci 8-spored; spores hyaline,
spherical to acicular, straight or twisted, 1-celled to muriform,
usually with a thin wall (except in Physma and section Lem-
phospora of Lempholemma).
This family is widely distributed in the temperate and tropical
zones, the more highly developed members occurring in the
latter. It is possible that some of the smaller and less conspicuous
genera have been overlooked or have been included in the crustose
material in the preliminary sorting. So far only four genera
have been found in Costa Rica.
KEY TO TROPICAL AMERICAN GENERA OF COLLEMACEAE
Spores unicellular.
Thallus crustose, not gelified, parathecium present; spores ellipsoidal,
thin-walled.
Brasi
FORD REITI TERI ER x Wa REA FEX EXXTAMESVEUSURHS C cats L. Finkii
Thallus squamulose, gelified; spores soberiol, thin-walled but surrounded
by a thick vire sheath.
Thallus not corticate.......... 0. ccc ccc cece cece cc eceaseues Lempholemma
Thallus corticate, wholly pseudoparenchymatous; Juan Fernandez
ECER TEE EE IE a on EE eee Lemmopsis PER TETT E
Thallus foliose, gelified; spores ellipsoidal to fusiform, thick-walled,
h
surrounded by a gelified sheath.............. 0.000. essere Physma
Spores phragmospores or dictyospores.
Cortex not developed; apothecia with parathecia only................ Collema
Cortex of pseudoparenchyma; apothecia with amphithecia (parathecia
often also present)............ 0c ccc ccc c cece cece hr eptogium
LEMPHOLEMMA Körb.
LEMPHOLEMMA Körber, Syst. Lich. Germ. 400. 1855.
Type species: Lempholemma compactum Körber.
Thallus from verrucose, squamulose, dwarf fruticose to foliose
in our species, gelified when moist, clothed with rhizinae below,
1933]
DODGE—LICHENS OF COSTA RICA. I 405
homoeomerie, without cortex, with Nostoc. Apothecia superfi-
cial or terminal, mostly sunken in the thallus (not in our species) ;
amphithecium either with or without cortex; parathecium color-
less, urceolate; hypothecium colorless; paraphyses filiform,
simple; asci 8-spored, clavate, often twisted below; spores
hyaline, fusiform, ellipsoidal-ovoid or spherical, smooth.
The genus has been divided into eight sections of which only
one section Lemphospora is tropical. This section is characterized
as having apothecia without cortex, spores more or less spherical
with a thick gelified sheath. Lempholemma Dussii (Vainio)
Zahlbr., Cat. Lich. Univ. 3: 23. 1925 [Collema Dussii Vainio,
Ann. Acad. Sci. Fenn. A6': 114. 1915] has been described
from Guadeloupe in the Antilles.
LEMPHOLEMMA (LEMPHOSPORA) oblique-peltatum (Eschw. ap.
Martius) Dodge, comb. nov.
Collema oblique-peltatum Eschw. ap. Martius, Icon. Pl. Cryptog.
Fasc. 2: 27, pl. 11, f. 2 [between 1828 and 1833].
Type: Brasil, near Pará, Martius.
Thallus foliose, up to 5 cm. in diameter, drying grayish olive
to dark olive gray, margin thick, broadly subdichotomously
lobed, lobes up to 3 mm. broad and 2-3 mm. long, smooth,
surface minutely pitted although appearing practically smooth
to the naked eye, without soredia or isidia; thallus 200—400 y.
thick, homoeomerous, without cortex, but provided with a thin
layer of tangled rhizinae below. Apothecia subsessile, often
appearing obliquely attached to the thallus in section, up to
2.5 mm. in diameter, margin of the same color as the thallus,
with periclinal folds and wrinkles, disc tawny to mars brown;
amphithecium appearing lobulate in section (due to the periclinal
wrinkles), homogeneous with the thallus, 200-300 y thick but
extending beyond the parathecium as much as 800 u; para-
thecium pseudoparenchymatous, highly developed, about 100 yu
thick below, thinning out above to about 50 y; hypothecium
about 40 y. thick, of highly gelified hyphae; thecium about 100 y.
1 I have been unable to locate the date of the second fascicle of this work. The
cover of the copy in the Missouri Botanical Garden gives the pages of text and plates
in each fascicle but only the dates 1828-1834. Martius in his ‘Flora Brasiliensis’ 1:
233-234. 1833, quotes from the ‘Icones’ citing page, hence it must have been issued
before that date. In this work he placed it in the subgenus Enchylium.
[Vor. 20
406 ANNALS OF THE MISSOURI BOTANICAL GARDEN
tall; paraphyses about 1-2 y. in diameter, clavate above, septate,
cells 4-6 u long, with walls highly gelified so that they appear
as slender rows of cells imbedded in a gel; asci clavate-cylindrical,
about 80 x 16 u, containing 8 monostichous spores; ascospores
broadly ellipsoidal, 12 x 8 y. while still in the ascus, with a thick
gelified wall.
While I have not studied the type specimen of this species, a
careful study of the description and the figures shows that it
belongs in Lempholemma section Lemphospora rather than in
Collema. Judging from determinations observed in various
herbaria, this species seems to have been confused with Lepto-
gium vesiculosum (L. bullatum), L. tremelloides, and L. foveolatum,
although the abundant development of rhizoids below should
have separated it. While it is minutely scrobiculate-foveolate
(visible under hand-lens), it is not conspicuously so to the naked
eye as in L. foveolatum. Besides my Costa Rican material I
have also seen it from Eustis, Lake Co., Florida, Nash 2024,
determined by Eckfeldt as L. faveolatum (sic) Nyl.
Limón: Waldeck, Dodge 7404; near Siquirres, 70-170 m., Dodge, Catt & Thomas
5588, 8024; Hamburg, 55 m., Standley & J. Valerio 48767.
Guanacaste: H. Santamaría, = m., Dodge & Thomas 6898; near Tilarán, 340-
670 m., Dodge & Thomas 6644, 80
Puntarenas: near Corozál, dd Dodge 8023; Puerto Jiménez, Brenes 839.
LEMPHOLEMMA (LEMPHOSPORA) dichotomum Dodge, sp. nov.
Type: Costa Rica, Guanacaste, H. Granadilla, Dodge &
Thomas 6736.
Thallus foliosus, ad 5 em. diametro, 400-650 y crassitudine
metiens, griseo-olivaceus, margine crassiuscula, integra, lobatus,
dichotomus, lobi 1-1.5 mm. lati, divergentes, superne longi-
tudinaliter rugosus, sine sorediis isidiisque, inferne rhizinis nigris
densissimis intertextis obsitus, homoeomerus, nostocaceus, sine
strato corticali. Apothecium peltatum, basi constrictum, ad
2.5 mm. diametro metiens, margine concolori, rugosa, disco
fulvo castaneove; amphithecium sectione lobatum, homogeneum,
150-300 y crassitudine inferne, sed ad 500 y superne et ultra
marginem parathecii; parathecium pseudoparenchymatieum,
ad 160 uw crassitudine inferne, attenuatum ad 80 y superne;
hypothecium filamentosum, ad 80 y crassitudine, hyphis geli-
factis; thecium ad 120 y altum; paraphyses filiformes, apicibus
1933]
DODGE—LICHENS OF COSTA RICA. I 407
decompositis brunneis, in materia gelata fixi, cellulis cylindricis,
l a diametro, 4-6 y longitudine metientibus; asci 60 x 10 y,
cylindriei; ascosporae octonae, monostichae, late ellipsoideae,
8 x 12 y. (dum in ascis sunt), membrana incrassata, gelifacta.
Thallus foliose, up to 5 cm. in diameter, 400-650 u thick,
grayish olive, margin thick, smooth, dichotomously lobed, lobes
1-1.5 mm. broad, divergent, upper surface longitudinally wrinkled,
without isidia or soredia, lower surface covered with a dense
black nap of rhizinae, homoeomerous, with Nostoc, without
cortical layer either above or below. Apothecium constricted
at the base, up to 2.5 mm. in diameter, margin concolorous,
irregularly wrinkled, dise tawny to chestnut; amphithecium in
sections appearing lobed, homogeneous with the thallus, 150-
300 u thick below but extending up to 500 u above and beyond
the margin of the parathecium which is pseudoparenchymatous,
about 160 » thick below, thinning out to 80 y at the margin;
hypothecium filamentous, 80 » thick, of gelified hyphae; thecium
about 120 u tall, paraphyses filiform, the apices decomposing
brown, imbedded in a gel, cells cylindric, about 1 y in diameter,
4—6 y. long; ascospores 8 per ascus, monostichous, broadly ellip-
soidal, about 8 x 12 y. (while still in the asci), with a thick gelified
wall. This species has also been seen from Barro Colorado
Island, Gatun Lake, Panamá.
Guanacaste: H. Granadilla, 540 m., Dodge & Thomas 6737.
PHYSMA Mass.
PuysMA Massalongo, Neag. Lich. 6. 1854.
Dichodium Nyl., Bull. Soc. Linn. Normandie II, 2: 43. 1868.
Type: Physma Boryanum Massalongo.
Thallus foliose with rhizinae beneath, corticate with several
layers of pseudoparenchymatous cells; algae Nostoc. Apothecia
superficial, lecanorine, with broad dise, thick margin; hypo-
thecium light-colored; paraphyses filiform; asci 8-spored; spores
colorless, ellipsoidal or fusiform, unicellular with thick almost
warty wall or with a gelified sheath. Spermagonia sunk in the
thallus, showing above by the dark swelling, surrounded by
pseudoparenchyma; spermatiophores simple or forked, septate,
cells short; spermatia short, straight.
[Vor. 20
408 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Only two species are known from tropical America, P. chilensis
Hue from Chile and P. pruinosum Vainio, from the Antilles.
Material reported from America as belonging to P. byrsinum
(Ach.) Müll. Arg. (P. byrsea (Ach.) Tuck.) is probably mis-
determined. Apparently the genus is most highly developed
and wide-spread in Oceania and adjacent Asia.
PHYSMA PRUINOSUM Vainio, Ann. Acad. Sci. Fenn. A67:112. 1915.
A single sterile specimen collected near Cartago, 1500 m.,
C. Wercklé, May 1900 (in Mus. Nac. 17233, and in Herb. Bot.
Gard. Berlin), was referred to Physma byrsinum by Lindau. It
is possible that this species belongs to Physma pruinosum Vainio
or it may possibly be a very young thallus of Leptogiwm marginel-
lum (Sw.) S. F. Gray. Its spermagonia are marginal and its
spermatia are ellipsoidal and small.
COLLEMA Wigg.
CornLEMA Wiggers, Primit. Fl. Holsat. 89. 1780.
? Gabura Adanson, Fam. Pl. 2:6. 1763?
Scylenium S. F. Gray, Nat. Arrang. Brit. Pl. 1: 398. 1821.
Type species: Collema Lactuca (Web.) Wiggers [Lichen crispus L.].
Thallus foliose or squamulose to almost crustose, gelified
when moist, lying on the substrate without rhizinae, homoe-
omerous, not corticate, hyphal system loose; algae Nostoc.
Apothecia at first sunken, erumpent, sessile or scutellate and
constricted below, with amphithecium; parathecium either
present or absent, both parathecium and hypothecium either
of interwoven hyphae or pseudoparenchymatous; paraphyses
simple, adherent, mostly septate; asci 8-spored; ascospores
colorless, cylindric, acicular, fusiform, long-ellipsoidal or ovoidal to
almost cubical, ends obtuse or acute, sometimes becoming
muriform, thin-walled, without gelified sheath. Spermagonia
sunken in the thallus or in thalline warts, with light-colored wall;
spermatiophores simple or branched, without sterigmata, septate
with short cells; spermatia short, oblong to ellipsoidal, straight.
KEY TO TROPICAL AMERICAN SPECIES OF COLLEMA
Amphithecium corticate; spores acicular........... isses COLLEMODIOPSIS
[C. lherminieri Hue, from Guadeloupe, is the only species of this typically
northern group so far reported. C. Granadillae is here reported from
Costa Rica.]
1933]
DODGE—LICHENS OF COSTA RICA. I 409
Amphithecium not cort
Spores oblong to amet more or less muriform; thallus laciniate,
Tore Or NM EEUE. iL soos Sivan eR ern nen sete BLENNOTHALLIA
Lobes n than 0. 5 mm. broad; apothecia unknown; Chile....... C. millegranum
Lobes more than 0.5 mm. broad; apothecia well developed; Mexico.
M rr NEMESIS C. masioandes
TE clavate, ins esae to fusiform, siguen rarely up to 3-septate,
thallus narrowly laciniate or incised...... DICOLLEMA
nd clavate, 14-21 x 4.5-5.5 u; Brasil, soon , and Argentina....
mH m TIN. C. corynesporum
Spores ellipsoidal or fusiform.
Margins of laciniae nodose-granular................00e00 00s C. pycnocarpum
po. PLI p; Brasil. orii oh oe bk na ees o rr v. Minarum
Sporos 10-44 x 6-4 u; Brasil oi ces se. Ace ei ee pas One v. Malmei
Spores 10-15.5 x 4-5.5 u; Brasil and Colombia.......... v. crassiusculum
Spores 14-17 x 4-5 u; New York and New Jersey............ v. typicum
Spores 16-25 x 3-7 u, 3-septate; Virginia southward.......... C. cyrtaspis
Spores 16-30 x 4-5 yu, 1-3-septate; S Pr be EM C. pycnocarpoides
Margins of laciniae not nodose-granula
Laciniae 1-3 mm. broad, spores 12- 14 x 5-6 u; Paraguay...... C. crenatum
Laciniae under 1 mm. broad, more or less canaliculate below; Cuba.
pe gt a ee en erm n ar ROSE . stellatum
Laciniae thin, flat; spores 11-17 x 4-5 w................ C. solenarium
Spores fusiform to acicular, more than 3-septate, over 30 » long; thallus
broadly laciniate, often fenestrate...............00. SYNECHOBLASTUS
Thallus isidiose; spores over 100 x in length; Brasil
Parathecium present, spores 125-175 x 8-4 u........... sun. d rade
Parathecium absent, spores somewhat over 100 nu inlength........ C.
Thallus not isidiose; spores not over 95 y in length.
Apothecia white-pruinose.
Apothecia small, crowded on short bullate prominences; spores
0-80 x 3-5 wu; thallus pustulate, not fenestrate; southern
United States and southward along coastal plains. ...... E OS
ae sessile or nearly so, larger; thallus more or less fen
6; tropical highlands. ELDER ese C. Howophtilmsum
Piat 77-92 x 624 iy MOXIOO s a a a a eese typicum
Spores 55-74 x 5-7 m, 7-1l-septate; Colombia............ v. pease
Spores 42-58 x 4-5 u, 6-9-septate; Brasil................ v. brasiliense
Spores 36-50 x 7 p, only 5-septate. ............... sss. C. leucocarpum
Apothecia not white-pruinose.
Spores 48-66 x 6-8 u; Colombia............... eee eee C. implicatum
Spores 38-50 x 6-8 p; Chile... REEL ahs ss hee tases C. chilenum
E 40-45 x 5-6.5 u; apothecia sessile, margins turgid; Mex-
E vier s UE O Gea lk tol Rae Sos o eB C. turgidulum
bas, 30-37 x 2.5-3 pu; apothecial margins thin, blackening;
DIOR o eonun ere aa ee . baculiferum
CoLLEMODIOPSIS Vainio, Étude Lich. Brésil 1: 234. 1890.
Type species: Collema nigrescens (Huds.
[Vor. 20
410 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Spores slender, several-celled phragmospores, never muriform,
apothecia with a pseudoparenchymatous cortex.
This subgenus is typically northern in its distribution. Only
C. lherminieri Hue from Guadeloupe has been described previ-
ously from the American tropics.
CoLLEMA (CoLLEMopIopsis) Granadillae Dodge, sp. nov.
Type: Costa Rica, Guanacaste, H. Granadilla, Dodge & Thomas
6576.
Thallus irregulariter subpinnatifide lobatus, lobis tenuibus,
ad 1 mm. latis, viridi-nigricans, superne elevato-rugosus verruco-
susque ad subisidiosus, inferne reticulatim elevato-rugosus,
cinerascens, ad 400 y crassitudine, decorticatus, rhizinis desti-
tutus. Apothecium planum, peltatum, basi constrictum, 0.5-
1.0 mm. diametro metiens, omnino nigrum, margine verrucosa;
amphithecium 150-250 yu crassitudine, pseudoparenchymatice
corticatum, intus algis nostocaceis subrectis; parathecium grosse
pseudoparenchymaticum, bene evolutum, 100—120 y crassitudine
inferne, attenuatum ad 25-35 y superne; hypothecium hyphis
tenuibus dense contextum, 25-30 y crassitudine; thecium 150-
170 y altitudine, paraphyses arcte cohaerentes septati, cylindrici,
filiformes, ad 1 u diametro metientes, clavato-capitati; ascosporae
octonae, polystichae, rectae, fusiformes vel aciculares, 5-septatae,
97-42 x 4-5 y.
Thallus irregularly subpinnately lobed, lobes slender, about
1 mm. broad, dark greenish black, with elevated folds and
wrinkles above with masses of small subisidiose warts, below
reticulately deeply scrobiculate wrinkled, more or less ashy,
about 400 y thick, without cortex or rhizoids. Apothecium
peltate with constricted base, plane, about 0.5-1 mm. in diameter,
wholly black, margin verrucose; amphithecium 150-250 y thick,
corticate with several layers of pseudoparenchyma below, within
the filaments of Nostoc more or less straight and loosely inter-
woven; parathecium of large-celled pseudoparenchyma, about
100-120 y thick below, thinning above at the margin to 25-35 y;
hypothecium of densely woven slender hyphae, 25-30 y thick;
thecium 150-170 u tall; paraphyses closely adherent, septate,
cylindrical, filiform, about 1 u in diameter, clavate, capitate at
the apex; ascospores 8 per ascus, polystichous, straight, fusiform
or acicular, 6-celled, 37—42 x 4-5 u.
1933]
DODGE—LICHENS OF COSTA RICA. I 411
Guanacaste: H. Granadilla, Dodge & Thomas 6576; near Tilarán, Standley & J.
Valerio 44529, 640—660 m., Dodge & Thomas 6560.
CoLLEMA LEUCOPEPLA (Tuck.) Schneider, Guide Study Lich.,
181. 1898.
Collema nigrescens var. leucopepla Tuck., Syn. N. Am. Lich.
1:148. 1882.
Type: not stated in original description, based on material
from South Carolina, Georgia, Florida, Alabama, and Louisiana.
It is probable that material collected in Puntarenas, Boruca,
560 m., Tonduz 5373, and determined by Müller Argau as C.
nigrescens var. caesium Ach., belongs here but I did not have
time to study it critically while I was in Genéve. This species
of the lower Atlantic and Gulf Coastal Plain in the United States
has also been reported by Vainio from Mexico.
DICOLLEMA (Clements) Dodge, n. m ^
Dicollema Clements, Gen. Fung. 74.
Type species: Collema pycnocarpum Ny i
Thallus narrowly laciniate or incised, spores clavate to fusiform,
typically 2-celled, rarely 4-celled.
The material referred to the type species of this genus by
different authors is quite variable and the whole group needs a
thorough revision. Pending such a study, I am proposing
several varieties of C. pycnocarpum Nyl. to cover the more
conspicuous variations which seem to have separate geographical
ranges, those with the smaller spore sizes being rather more
southern in their distribution.
COLLEMA PYCNOCARPUM Nyl., Syn. Meth. Lich. 1: 115. 1858.
Type: United States [on journey from New York to Philadel-
phia], Moré.
Thallus dark or pale green, medium in size, granulate-nodose.
Apothecia rufous, crowded, almost contiguous, at first plane,
then somewhat convex, quite small, 0.5 mm. or a little larger;
spores 8 per ascus, oblong or oblong-ellipsoid, simple or uniseptate,
14-17 x 4-5 y.
Northern Atlantic Coastal Plain.
Var. Minarum Dodge, var. nov.
Collema pycnocarpum Vainio, Etude Lich. Brésil 1: 238. 1890.
Type: Brasil, Minas Geraes, Sitio, 1000 m., Vainio 734.
[Vor. 20
412 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Sporae oblongae, ellipsoideae vel fusiformes, 1-septatae,
6-0 x 3 y.
Var. Malmei Dodge, var. nov.
C. pycnocarpum Malme, Ark. f. Bot. 198: 7. 1924.
Type: Brasil, Rio Grande do Sul, Canóas near Porto Alegre,
Malme 536.
Sporae oblongo-ellipsoideae vel ellipsoideae, uniseptatae, 10—
14 x 3-4 y.
Var. crassiusculum (Malme) Dodge, comb. nov.
Forma crassiusculum Malme, Ark. f. Bot. 198: 7. 1924.
Collema pycnocarpum Nyl., Acta Soc. Sci. Fenn. 7:428. 1863.
Type: Brasil, Matto Grosso, Corumba, Malme.
Sporae ellipsoideae, utroque apice obtusae, raro acutae vel
altero obtusae, altero acutae, 10-15.5 x 4-5.5 y.
The specimens cited by Nylander from Colombia, Bogotá,
2600 m., Lindig 2872, have spores 10-14 x 4.5-5.5 u.
COLLEMA CYRTASPIS Tuck., Proc. Amer. Acad. Arts & Sci.
5:387. 1802.
Type: no type mentioned in the original description.
Thallus deeply and irregularly lobed, verrucose, deep blue
green, darkening on drying, verrucae much larger than in C.
pycnocarpum, about 400 y thick; algal layer about 100 y thick,
composed of coiled and tangled filaments of Nostoc imbedded in
a gel, cells spherical, about 4 u in diameter; medulla gelified,
traversed by very loosely tangled hyphae about 2-3 y in diameter
and occasional straight filaments of Nostoc. Apothecia immersed
in the verrucae when young, becoming plane or even convex,
up to 2 mm. in diameter, margin slightly verrucose when young,
practically disappearing as the apothecium becomes increasingly
complex; amphithecium about 160 » broad, of the same texture
as the thallus; parathecium lacking; hypothecium filamentous,
quite highly developed, about 40 y thick, not conspicuously
thinner toward the margins, hyphae 2-3 » in diameter, very
loosely woven; thecium about 80 y tall; paraphyses filamentous,
clavate, expanded above, imbedded in a gel; asci clavate, about
40 x 8 u, 8-spored; ascospores subfusiform, 2-4-locular, 16-25 x
3-7 u [16-20 x 4-4.5 u in Costa Rican specimens].
This species is widely distributed in the southern Atlantic
1933]
DODGE—LICHENS OF COSTA RICA. I 413
and Gulf Coastal Plain in the United States, extending up the
Mississippi Valley to Iowa.
Guanacaste: Liberia, 100 m., Dodge & Thomas 8016, 8027.
SYNECHOBLASTUS (Trevis.) Vainio, Étude Lich. Brésil 1:
234. 1890.
Lathargium S. F. Gray, Nat. Arrang. Brit. Pl. 1: 399. 1821,
p. p.
Synechoblastus 'Trevis., Caratt. Tre Nuov. Gen. Coll. 3. 1853,
p. p.
Type species: as subgenus based on C. glaucophthalmum N yl.
Apothecia without pseudoparenchymatous cortex, spores fusi-
form to more or less acicular, many-celled, mostly over 30 y long.
This subgenus is typically northern. Only two species are
known from Costa Rica, where they are temperate species at
elevations between 1000 and 1800 m., and are also found in the
highlands of Colombia at somewhat greater altitudes.
COLLEMA GLAUCOPHTHALMUM Nyl., Syn. Meth. Lich. 1: 114,
115. 1858.
Type: Mexico, Orizaba, Fr. M üller.
Thallus olivaceous-fuscous, medium size, expanded, more or
less fenestrate and dissected, scrobiculate, and often granuliferous.
Apothecia glauco-lilac colored, plane and somewhat concave,
margin prominent, thin; ascospores 77-92 x 6-7 y.
Var. GRANATENSE Hue, Jour. de Bot. [Morot] 20:12. 1906.
Collema glaucophthalmum Nyl., Acta Soc. Sci. Fenn. 7: 428.
1863, non loco alio.
Type: Colombia, Choachí, 2600 m., Lindig 813.
Thallus dusky yellowish green to almost black, irregularly
lobed and fenestrate, smooth or somewhat scrobiculate, coarsely
wrinkled and subverrucose, especially toward the tips, margins
thick and rounded, 800-1000 u thick, homoeomerous, of loosely
tangled filaments of Nostoc, cells 5-6 x 2.5-3 y, ellipsoidal.
Apothecia peltate, constricted at the base, up to 3 mm. in di-
ameter, margin at first prominent, smooth, becoming thinner,
less prominent and verrucose at maturity; disc chalky white to
vinaceous-russet; amphithecium about 160 y. thick, homogeneous
with the thallus; parathecium pseudoparenchymatous, 40 u
thick, thinning to 20 u or even disappearing above at the margin;
[Vor. 20
414 ANNALS OF THE MISSOURI BOTANICAL GARDEN
hypothecium filamentous, of large very densely interwoven
hyphae about 40 y thick; thecium about 120 y tall; paraphyses
filiform, about 1 u in diameter, not swollen above; asci clavate,
about 20 y in diameter, 8-spored; ascospores polystichous,
fusiform to acicular, 55-74 x 5-7 u.
The whiteness of the apothecial dise is very variable on the
same thallus, in general being more pronounced on young and
rapidly growing apothecia and gradually disappearing on older
and more exposed ones. It seems to be a semi-crystalline
deposit which slowly dissolves away, disclosing the vinaceous-
russet disc formed by the discoloration of the upper portion of
the gel surrounding the paraphyses. This variety has previously
been reported only from Colombia.
Cartago: near R. Birrfs above Santiago, 920-1340 m., Dodge 8011, Dodge & Thomas
8015; Carpintera, 1700 m., K. Danielson 102.
San José: Sta. María de Dota, 1500-1800 m., Standley & J. Valerio 44150.
CoLLEMA IMPLICATUM Nyl., Acta Soc. Sci. Fenn. 7: 428. 1863.
Type: Colombia, Villeta, 1100 m., Lindig 749; Cundinamarca,
Bogotá, 2400-2600 m., Lindig.
Thallus dark greenish black, broadly lobed and fenestrate,
smooth near margins, becoming deeply reticulate-scrobiculate in
older portions of the thallus, verrucae scattered, more elevated,
often subisidioid, margins thin and semipellucid, homoeomerous,
of loosely tangled filaments of Nostoc, cells 4 x 5 u, heterocysts
5x8 y, much more closely tangled in an outer zone about 150 y
thick. Apothecia peltate, constricted at the base, up to 4 mm.
in diameter, margin at first prominent, verrucose, becoming
thinner, less prominent, scarcely visible as the apothecium
becomes expanded and convex, dise cameo-brown, without
pruina in young apothecia; amphithecium about 100 p thick,
homogeneous with the thallus; parathecium pseudoparenchy-
matous, about 80 à thick below, thinning out to a single layer
of thick-walled cells 6 & wide above at the margin; hypothecium
of densely woven hyphae about 20 u thick; thecium about 100 y
tall; paraphyses about 2 y in diameter, with clavate brown tips;
asci clavate, 8-spored, 12-16 y in diameter; ascospores poly-
stichous, fusiform to acicular, 48—66 x 6-8 u, 7—9-septate.
This species is very closely related to C. glaucophthalmum
1933]
DODGE—LICHENS OF COSTA RICA. I 415
and has been reduced to synonymy by some authors. It differs
from the previous species in several minor details of proportion
of measurements and in the lack of a well-developed pruina.
It is to be hoped that some one will be able to study the two
forms in the field to determine the constancy of this character.
In a series of specimens one can find considerable variation in
the amount of pruina in C. glaucophthalmum. It should be
noted that the two species occupy the same areas in Costa Rica.
Cartago: above R. Birrís at Santiago, 1140-1180 m., Dodge 8017; Carpintera,
1700 m., K. Danielson 101c.
The following species has not been found in Costa Rica but
it is closely related to this group.
COLLEMA (SYNECHOBLASTUS) Ramboi Dodge, sp. nov.
Type: Brasil, Rio Grande do Sul, Porto Alegre, B. Rambo 74.
Thallus parvus, adscendens, obscure viridi-nigricans, angusti-
lobatus, fenestratus clathratusque, verrucis elevatis isidioideis,
marginibus plus minusve integris, tenuibus, pellucidis, 130-140 y
crassitudine, homoeomerus, filamentis nostocaceis laxe implexis,
in zona exteriori 40 y crassitudine dense contextis, cellulis ad
3x6 yu. Apothecium peltatum, basi constrictum, planum,
0.5-0.6 (— 1.0) mm. diametro, margine tenui, verrucosa sub-
crenulatave, disco obscure castaneo nigroque; amphithecium 20 p
crassitudine, cum thallo homogeneum; parathecium deest; hypo-
thecium 20 u crassitudine, hyphis tenuibus dense contextum;
thecium 160 y altitudine; paraphyses filiformes, ad 1 y diametro,
apicibus inflatis; asci clavati, 120 x 8-10 »; ascosporae octonae,
polystichae, aciculares, immaturae, plus quam 100 y. longitudine,
multiloculares.
Thallus small, elevated, dark greenish black, narrowly lobed
and fenestrate, verrucae elevated and isidioid, margins more or
less smooth, thin, pellucid, 130-140 u thick, homoeomerous, of
loosely tangled filaments of Nostoc, cells about 3x 6 y, much
more closely tangled in an outer zone about 40 y thick. Apothecia
peltate, constricted at the base, 0.5-0.6 (- 1.0) mm. in diameter,
plane, margin very thin, verrucose, subcrenulate, finally almost
disappearing while the expanded apothecium remains plane,
dise dark chestnut to black; amphithecium 20 y broad, homo-
geneous with the thallus; parathecium absent; hypothecium 20 y.
[Vor. 20
416 ANNALS OF THE MISSOURI BOTANICAL GARDEN
thick, of slender densely woven hyphae; thecium about 160 u
tall; paraphyses filiform, about 1 y in diameter with greatly
nidi tips, imbedded in à hymenial gel; asci clavate, 8-spored,
120 u long, 8-10 u wide; ascospores polystichous, 8 per ascus,
acicular, immature and very difficult to measure but somewhat
more than 100 u long, many-celled.
LEPTOGIUM 5$. F. Gray
Leproacium S. F. Gray, Nat. Arrang. Brit. Pl. 1: 400. 1821.
Type species: Leptogiwm tremelloides S. F. Gray, excl. syn.
Thallus foliose in the tropical species, gelified, below naked
(covered with rhizinae in section Mallotium), with pseudo-
parenchymatous cortex above and below, algae Nostoc. Apo-
thecia sunken in the thallus at first, emerging and often short-
stipitate; amphithecium often corticate with several layers of
pseudoparenchyma below, usually of a single layer above;
parathecium usually present and pseudoparenchymatous, often
thinning out above at the margin, sometimes of large, more or
less parallel hyphae; paraphyses simple and filiform, often with
apex variously thickened; asci clavate to cylindrical, 8-spored;
ascospores usually imbricately monostichous, occasionally distich-
ous, hyaline, usually broadly fusiform with acute to acuminate
ends (acicular in section Leptogiopsis), usually muriform (except
in Leptogiopsis), with thin walls.
Four of the seven sections into which Leptogium is divided are
found in tropical America, all four of them in Costa Rica. Lep-
togiopsis differs from all the other sections in its acicular, never
muriform spores, and perhaps should be treated as a separate
genus, more or less related to section Synechoblastus of Collema.
It is endemie in tropical America, with a single unidentifiable
fragment yet found in Costa Rica. Mallotiwm, to which may
also be referred the section Leptolobaria of Vainio based on L.
callithamnium from Chile and the Antilles, has highly developed
rhizinae below, and often more or less tomentum above, especially
in the vicinity of the apothecia. It is widespread in its distribu-
tion, being found in the temperate zones and evidently preferring
the colder regions of the tropies, not occurring below about
1300 m., in Costa Rica not seen below 3000 m. Diplothallus, with
1933]
DODGE—LICHENS OF COSTA RICA. I 417
its two separate layers of thallus, each with its own cortex,
connected by pillars of pseudoparenchyma which on drying
cause punctate depressions in the thallus, is endemic in tropical
America and confined to the temperate regions, in Costa Rica
occurring between 1200 and 1700 m., coming down to lower
levels in the mountain near H. Santamaria of the Cordillera de
Tilarán in Guanacaste. Euleptogium, which contains the most
species, is found from sea level to about 1800 m. in Costa Rica,
although most of the species of this section have a narrower
altitudinal range. The lowland species ascend the river valleys
but mostly drop out at about 1000 m. These are usually wide-
spread at low elevations, extending from the Atlantic and Gulf
Coastal plains of the United States to southern Brasil and
Paraguay. The species of the temperate region of Costa Rica
are mostly confined to the mountains from southern Mexico to
the northern Andes and the mountains of eastern Brasil.
The morphology of the apothecium is not altogether clear and
considerable confusion exists in the nomenclature of the various
parts. In the following discussion, I have treated the tissue,
usually well differentiated, underlying the hypothecium, as the
parathecium, whether it extends to the surface of the thecium
or not and whether or not it is pseudoparenchymatous. In a
very few species the parathecium as thus defined thins out and
disappears near the edge of the thecium, and the hyphae of the
hypothecium extend up to the surface of the thecium. It is
possible that some would prefer to regard the parathecium as
formed of two layers, the lower of which is pseudoparenchymatous
and the upper filamentous, with only the upper extending to
the surface of the thecium. The amphithecium (thalline margin
of earlier lichenologists) is essentially homogeneous and continuous
with the thallus, although the algae are sometimes more densely
tangled and in many species the cortical cells on the under side
of the amphithecium (morphologically continuous with the upper
surface of the thallus) or even the under side of the thallus
below the apothecium, may proliferate, forming a pseudoparen-
chyma of several layers of cells sometimes even thicker than
the algal zone.
[Vor. 20
418 ANNALS OF THE MISSOURI BOTANICAL GARDEN
KEY TO THE TROPICAL AMERICAN SPECIES OF LEPTOGIUM
Spores acicular, never mu
Thallus efiulateseroieulate; French Guiana. [Spores 6-celled, 35 x 10 y,
teste Leighton, in specimen from Amazonas, Brasil].......... L. reticulatum
Thallus far reticulate-scrobiculate.
Spores 48 u long, 8-10-celled; Florida.......... L. fusisporum (Tuck.) Dodge
Spores 50-60 x 7-8 u, 10-12-celled; Mexico................... pressum
Spores IT x 3.5-5 yu, 10-12-celled; Brasil.............. L. iégapols vou"
Spores murifor
Thallus of im lene each with its own cortex, attached to each other
by columns of tissue... nnne DIPLOTHALLUS
AB ME TI oi 9555055 9-5 0 0 ee teens L. diaphanum
Thallus of a single lamella, not as above.
Thallus with a thick tomentum of rhizinae below............ MALLOTIUM
Apothecia on lower — lobes of thallus broad, imbricate. . . . L. resupinans
Apothecia on upper su
Upper surface of thallus tomentose, at least near apotheci
Margins of sinuses “iy lobes medium, sinuate; spec
margins microphylline................. 0.00000 e eee L. inflecum
Margins of thallus dios dissected.
Upper v of th and more g
MEN A ERATE E E E T L. papillosum
-— n of the thallus arachnoid-tomentose, lobes
nutely dissected, not otherwise isidiose; Chile. L. callithamnium
Upper parma of thallus glabrous except in the vicinity of the
apothecia; apothecial margin isidiose, scattered groups
of isidia sometimes on the upper ica of the thallus.
POC TET CETELE TT ee TETE ET inflecum v. isidiosulum
Thallus corticate with a single layer of cells on both yi without
ae Po. me ee Tee Seen ee bree Te EULEPTOGIUM
Thallus scrobiculate.
Margin of thallus papillate-denticulate, olivaceous, thallus deeply
reticulate-scrobiculate; parathecium filamentous; spores 22-
MINE. rire a4 5895 05 659004959440858 050 EXER L. olivaceum
Margin of thallus smooth, thallus plumbeo
Thallus deeply reticulate-scrobiculate; nab 30-40 x 12-16 y.
— rre L. foveolatum
Thallus shallowly scrobiculate; spores 20-30 x 8-10 y. . .L. microstictum
Thallus variously wrinkled on drying, but not scrobiculate, usually
h when moist
Apothecia on tips of long swollen processes of the thallus.
Thallus lobes narrow, dichotomously branched, wrinkles ir-
TOA ry ia sn E he ae doa aoe ak eaten R RITE ES . stipitatum
Thallus lobes broader, not conspicuously dichotomously
branched, wrinkles predominantly go ied or radial,
secondary wrinkles mostly periclinal to m
— radially wrinkled to node spores 23-32 x
MIB TCI" . vesiculosum
"mE. with periclinal wrinkles, foliolate or isidiose.
'Thallus and apothecia not isidiose.
1933]
DODGE—LICHENS OF COSTA RICA. I 419
Apothecia small, spores 22-30 x 10-12 u, margin wrinkled.
Thallus lobes broad c.s oou rre) L. depuis
Thallus lobes narrower, margins crisped........... ampestre
Apothecia rather large with more or a foliolate margins.
PATE i Nie hn Gre anki ob ps anne RS See v. macrocarpum
Thallus and apothecia isidiose, often densely so; spores
BO-40 (65) x 12-17 poss coo L. coralloideum
Apothecia marginal, sessile or on short solid stalks, very minute.
Thallus dark green to black, exciple isidiose [spores 24-30
ads e u, transverse septa 5, fide Malme];
CTC ALC: ERO IINE D I IE L. chloromelan
Thallus weil to plumbeous.
Exciple isidiose, parathecium 150 y thick; thecium 150-
y. CD LE. . marginellum
Exciple verruculose, iiio 100 & thick; thecium
120-140 y tall; Paraguay......... eee L. pilcomayense
Exciple crenulate, esie ink 30 u thick; thecium 110-
0 a tall; Paragüsy. o n e eR . microcarpum
ay
Apothecia scattered on upper surface of the thallus.
Thallus thick, 250 4 or more, roughened bi by more or
less conical outgrowths but rhizinae absen
d Ach or slightly wrinkled, lobes E A obo-
eet 2 oscó i Eia eg noe UR EE xi L. hypotrachynum
EI wrinkled above, lobes appressed, short, wide
margins sometimes crisped; Brasil.......... . mattogrossense
Smooth or only slightly wrinkled below................. L. sessile
Thallus thinner, seldom reaching
Thallus more or less isidiose.
Isidia blackening, cylindrie; parathecium well developed.
Spores 18-23 x 8-10 u, transverse septa 3, rare
Brasil and Paraguay........ pichneum Malme excl. syn.
Spores 22-30 x 11-13 y, transverse Aaa Donde et:
MERE EE simplicius v. pichneoides
Isidia concolorous, parathecium not well developed. .
Nr ege E E ERI RIT L. doninta Nyl.
Thallus 100 „ thick; apothecia not isidiose; spores
u, transverse septa usually 5; isidia
coralloid branched or somewhat flattened; Brasil.
k^ woe Fuse For AAA L. austroamericanum (Malme) Dodge
Thallus about 30 u thick; apothecial margins verrucose,
spores 16-23 x 8-9 m, transverse septa 3; isidia
often flattened and microphylline..............
eres ee L. denticulatum Malme non Nyl.
Isidia of hemispheric concolorous granules, thallus rigid,
eranl undulate, wrinkled toward the margin;
Paraguay: sse o. as NARRA e i HEAR A Y ap L. granulare
Thallus not isidioe
Parathecium Pee er or, if pseudoparenchymatous,
very thin and inconspicuous.
[Vor. 20
420 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Cortex of amphithecium a single layer i — thallus
50-75 u thick; apothecia rarely 1 in diam.;
spores 15-19 x 6-8 p with 3 federa septa;
DM ariista $e aes POR S ed L. Puiggarit
Cortex of amphithecium a single layer of cells above,
becoming thick, pseudoparenchymatous below;
apothecia 1-3 mm. in diam.
Thallus greenish black, 80-100 u thick; spores 15-22
X 6-8 u, with 3-5 transverse septa; Brasil. . . L. brasiliense
Thallus ashy, Mic or bluis
Thallus papulose and wrinkled above, opaque,
130-160 » thick; spores 20-30 x 7-12 u.. L. pulchellum
Thallus minutely wrinkled above, opaque, 150-
170 u thick; spores 24-28 x 12 w........ L. Standleyi
Thallus minutely wrinkled above, margins in-
rolled; spores 38-46 x 16-18 w.......... L. dimorphum
Thallus smooth or minutely wrinkled on drying,
pellucid, less than 100 x thic
E 22-28 x 8-10 4; thecium 120-150 u tall;
thallus 80-100 p thick... L. azureum
Spores 18-22 x 10-12 „, with 3-5 transverse
— LM 90-120 ,& tall; thallus 35-
OO atbiok. oor boa 6s 40a RS L. Tuckermani
Spores 18-24 x 8-9 u with 3-5 transverse septa;
thecium 140-160 u tall; thallus 35-50 pu
ge | es ee LER "A Schiffneri
Parathecium peeudoperenchy matous.
Spores 35-40 x 16-20 4; thallus plumbeous to — amphi
thecium smooth; Bolivia.................. L. laevius (Nyl ) Dodge
Spores 30-40 x 12-17 u; thallus rather dark plumbeous to bla
Laciniae 1-1.5 mm. broad, apothecia not over 1.5 mm. in Pep
thecium 170 4 tall; paraphyses 1.5-2 u thick; Brasil....
ETT Tee ee AMOUR EATER KORR EA E SAAREEN TS L. Lafayetteanum
Laciniae up to 8 mm. broad, apothecia 1-2.5 mm. in diam.;
thecium 190-220 , tall; paraphyses 2.5-3 u thick; Brasil.
DES ARRXVECECNRELERE SUE X RO EY ESO En eke L. pachycheilum
Spores 25-35 x 12-15 4; apothecia 1-2.5 mm. in diam.; thecium
150-180 4 tall; paraphyses 2-2.5 4 thick; Brasil..........
Sins bak Ev OURS s 400440 EO RUE E VR L. ulvaceum perius non Pers.
Spores 22-30 x 11-13 u; apothecia 1.5-2.2 mm. in diam.;thec
150-170. 5 Illo sisse ERE Wr Eo tn Eee bs Ls simplicius
Spores 18-34 x 9-15 u; thallus dark green to plumbeous; amphi-
thecium granulate to wrinkled.
Thalline lobes ascendant, gyrose-plicate. . L. conchatum (Tuck.) Dodge
oromelan auct. non Sw.)
Thalline lobes narrower, edges erect, crisped. L. stellans (Tuck.) Dodge
DrirLorHALLUS Vainio, Étude Lich. Brésil 1: 222. 1890.
Thallus of two lamellae connected by pillars. Each lamella
1933]
DODGE—LICHENS OF COSTA RICA. I 421
has a cortex of a single layer of isodiametric cells above and
below, drying impressed-punctate. Rhizinae absent, spores
muriform.
LEPTOGIUM DIAPHANUM (Sw.) Mont., Ann. Sci. Nat. Bot. III,
10: 134. 1848.
Lichen diaphanum Sw., Nov. Gen. Sp. Pl. Prodr. 147. 1788.
Parmelia diaphana Ach., Meth. Lich. 223. 1803.
Collema diaphanum Ach., Lichenog. Univ. 654. 1810.
Leptogium punctulatum Nyl. ap. Fournier, Mexic. Pl. 1: 1.
1872.
Leptogium tremelloides var. impressopunctatum Tuck. ap.
Williams, Amer. Nat. 29: 482. 1895.
Type: Jamaica, Swartz, in Riksmuseet, Stockholm, carefully
described by Malme, Ark. f. Bot. 198: 27. 1924.
Thallus mineral gray, lobes rounded, ascending, crowded,
impressed-punctate, otherwise quite smooth, consisting of two
layers, each about 40 u thick, composed of a row of pseudo-
parenchymatous cells about 8 u in diameter on each surface
and an algal layer of Nostoc between, the layers connected by
pseudoparenchymatous pillars whose contraction in drying form
the depressions. Apothecia borne on the upper layer, peltate,
constricted at the base, 1-1.5 mm. in diameter, margin light buff,
smooth, dise chestnut; amphithecium corticate with thick
pseudoparenchyma below, about 80 y thick, thinning to two
layers of cells above next the disc, algal layer about 30 y thick
below the thecium; parathecium absent; hypothecium of densely
woven, large, thick-walled hyphae about 50 y thick; thecium
120-130 x tall; paraphyses filiform, 1-2 y thick, with clavate
tips, forming a brown epithecium imbedded in a gel; asci clavate
to cylindric, about 12-16 y in diameter, thin-walled tips thickened
and staining deep blue with iodine; spores 8 per ascus, imbricately
monostichous, fusiform, muriform, 16-25 x 7-8 y, with 3-5
transverse septa.
This species is found in the mountains from Mexico and the
West Indies, Dominican Republic, and Jamaica (900-1200 m.)
to Minas Geraes (1400-1500) m., Bolivia (1900 m.), and Perü
(1700 m). In Costa Rica it is found from 1200 to 1700 m. and
descends to 800 m. on the mountain back of the farmhouse at
[Vor. 20
422 ANNALS OF THE MISSOURI BOTANICAL GARDEN
H. Santamaría where the peculiar weather conditions enable
many species to flourish below their normal altitudes. Ap-
parently it needs high humidities as it is found mostly in localities
of frequent and long-continued fogs (see p. 379).
Cartago: R. Birrís above Santiago, 1220-1340 m., Dodge & Thomas 7938; Cartago,
R. Torres R. 143; Carpintera, 1700 m., K. Danielson 103.
San José: Sta. María de Dota, 1500-1800 m., Standley & J. Valerio 43208.
Heredia: C. Central de Zurquí, 1600-1700 m., Dodge, J. Valerio & Thomas 4624.
Guanacaste: H. Santamaría, 720-850 m., Dodge, Jiménez & Thomas 7017.
MarroTiUM Ach., Lich. Univ. 644. 1810.
Mallotium Gray, Nat. Arrang. Brit. Pl. 1:399. 1821.
Type: Collema saturninum (Dicks.) Ach.
Thallus foliose, cortex of pseudoparenchyma above, below
tomentose with rhizinae; spores muriform.
This subgenus, considered as a separate genus by many
authors, is cosmopolitan in distribution, mostly along cold
foggy coasts or in mountains, being abundant in species in
northern Europe and in Patagonia, Tierra del Fuego, and Ant-
arctic Islands. Our tropical species seem confined to much
higher elevations (above 1800 m. in Costa Rica) than the other
members of Leptogium. It is quite distinct in appearance, often
tomentose above, so that sterile specimens might be taken for
Erioderma or Umbilicaria on macroscopic examination.
Our two species, L. inflerum and L. papillosum, are both
Mexican, the former extending to Pert, the latter not known
south of Costa Rica.
Leprocium papillosum (Bouly de Lesdain) Dodge, comb. nov.
Leptogium Hildenbrandii var. papillosum Bouly de Lesdain,
Lich. Mexique, 30. 1914.
Type: Mexico, Michoacan, Puebla, H. Batan, G. Arséne
Brouard 4212; Morelia, C. Azul, G. Arséne Brouard 3999.
Thallus mineral gray above, dark olive buff below, lobes
subpinnatifid, with small, lacerate, almost isidioid margins,
papillate-granulose, sometimes isidiose above, densely tomentose
below, algal layer about 60 y thick, corticate on each surface,
with cells 5-6 w in diameter, the cortex of the lower surface
giving rise to a dense covering of rhizinae. Apothecia sub-
marginal but too immature in our specimens to show details of
structure well.
1933]
DODGE—LICHENS OF COSTA RICA. I 423
San José: L. de la Chonta, n. e. Sta. María de Dota, 2000-2100 m., Standley 42285;
near Sta. María de Dota, 1500-1800 m., Standley 41643.
LEPTOGIUM INFLEXUM Nyl., Flora 41: 377. 1858; Syn. Meth.
Lich. 1: 132. 1858.
Type: Mexico, Orizaba, Fr. M üller.
Thallus deep glaucous gray, dark olive buff below, lobes ir-
regular, rounded, margins crisped and subascending, smooth
above except in the vicinity of the apothecia, densely tomentose
below, algal layer about 60 y. thick, corticate on both sides with
two layers of cells about 8-10 y in diameter, the lower surface
giving rise to rhizinae, as also the outer cortical cells in the vicinity
of the apothecia. Apothecia large, up to 4 mm. in diameter,
margin more or less tomentose below, phyllophorous, concolorous
with the thallus, disc orange rufous to auburn, edge of para-
thecium lighter; amphithecium with pseudoparenchymatous
cortex about 100 u thick, thinning to about one or two cells
thick above, algal layer of Nostoc about 80 y thick; parathecium
pseudoparenchymatous, about 20 y. thick below the hypothecium,
expanding upward to thickness of 100 u above; hypothecium
30 u thick, of densely woven hyphae; thecium 240 y tall; para-
physes filiform, 1-2 u in diameter, with swollen clavate tips
forming a brown epithecium; asci 20-25 y in diameter, cylindrical;
ascospores monostichous, 8 per ascus, broad, fusiform, muriform,
with 7 transverse septa, about 35-40 x 13-17 y.
Superficially the thallus suggests the texture of L. tremelloides,
while the apothecia might easily be mistaken for L. phyllocarpum,
although the rhizinae below should be easily observed.
Cartago: F. Voleán de Turrialba, 2000—2400 m., Standley 34966, 35174.
San José: Quebradillas, 7 km. n. Sta. María de Dota, Standley 42899; C. de las
Vueltas, 2700-3000 m., Standley & J. Valerio 43823a.
Var. iSiDIOSULUM Nyl., Acta Soc. Sci. Fenn. 7: 429. 1863;
Ann. Sci. Nat. Bot. IV, 19: 289. 1863.
Type: Colombia, Paramo Choachí, 3600 m., Lindig.
This variety differs in the thallus being partly isidio-furfu-
raceous, especially the margins of the lobes; apothecial margin
isidiose instead of phyllophorous, subnude below; spores 36—44 x
18-25 y.
This variety would seem to differ from L. papillosum in that
the isidia are in scattered groups or marginal, much more highly
[Vor 20
424 ANNALS OF THE MISSOURI BOTANICAL GARDEN
developed and more denuded below. A single sterile specimen
has been referred here.
San José: C. Zurquí, 2000-2500 m., Standley & J. Valerio 48292.
EvLrEPTOGrUM Tuck., Gen. Lich. 95. 1872.
Stephanophorus Fw., Linnaea 17: 290. 1843.
Type species: no species cited.
Thallus foliose, monophyllous, without rhizinae, upper and
lower cortex present, each of a single layer of isodiametric cells;
spores muriform, many-celled.
This subgenus is predominantly tropical with many species in
tropical America.
Leproarum OLIVACEUM (Hook.) Zahlbr., Cat. Lich. Univ. 3:
146. 1924, excl. syn.
Collema olivaceum Hook. ap. Kunth, Syn. PI. Aequinoct. Orb.
Nov. 1:38. 1822.
Type: Colombia, Cauca, between Popayan and Almaguer,
Humboldt 252.
Thallus isabella to light brownish olive, lobes broad and round-
ed, surface deeply reticulate-scrobiculate with sharp ridges and
margins granulate-isidiose, or ridges rarely microphylline; about
200 u thick, corticate above and below with an algal layer of
loosely tangled filaments of Nostoc corresponding to a medulla,
separated from the cortex by a palisade layer about 30 y thick.
Apothecia sessile, margin verrucose, concolorous, disc darker;
amphithecium thick, homogeneous with the thallus; parathecium
about 80 u thick, of very densely woven hyphae; hypothecium
about 25 y thick, very deeply staining; thecium about 100 y
tall; paraphyses filiform, about 1.5 u in diameter, ending in the
epithecial gel.
The apothecia are very rare and too immature in the one
specimen from Costa Rica to give all the characters, but it
seems quite distinct from L. reticulatum and L. foveolatum.
Cartago: R. Reventazón below Santiago, 740—750 m., Dodge 8010.
Var. granulosum Dodge, var. no
Type: Costa Rica, San José, R. Virilla below El Brazil, Dodge
8030.
Thallus similis speciei sed nigro-granulatum ambobus super-
ficiebus. Apothecia obscure brunnea, nigricantia, sessilia, basi
1933]
DODGE—LICHENS OF COSTA RICA. I 425
constricta, disco concolore; amphithecium 190 y crassitudine,
corticatum cellulis isodiametricis; parathecium 130-140 y. crassi-
tudine, ad marginem 40 y. attenuatus, hyphis dense contextum;
hypothecium 30 uw crassitudine; thecium 120-130 y altum;
paraphyses similes speciei; asci 12-16 y diametro metientes;
ascosporae imbricatim monostichae, octonae, fusiformes, 22-24 x
12-14 y.
Thallus similar to that of the species but black-granulate on
both surfaces. Apothecia dark brown, blackening, sessile,
constricted at the base, disc concolorous; amphithecium 190 y
thick, homogeneous with the thallus, corticate with a single
layer of isodiametric cells; parathecium 130-140 y thick, thinning
toward the margin to 40 u, of densely woven hyphae; hypothecium
30 u thick, so deeply staining that its structure is not clear;
thecium 120-130 y tall; paraphyses as in the species; asci cylindric,
12-16 y in diameter; ascospores imbricately monostichous,
8 per ascus, fusiform, 22-24 x 12-14 y.
San José: R. Virilla below El Brazil, Dodge 8030.
LEPTOGIUM FOVEOLATUM Nyl., Syn. Meth. Lich. 1:124. 1858.
Type: not stated, specimens from Bolivia, Weddell, and
Mexico, Fr. M üller, are mentioned.
Thallus mineral gray or darker, often discolored light yellowish
olive or darker, lobes rounded, smooth, surface deeply foveolate
with sharp wrinkles both above and below; very variable in
thickness up to 250 u, Nostoc very sparsely scattered throughout,
rather denser and tending to form a palisade layer just below the
surface, but this layer much thinner and less definite than in L.
olivaceum. Apothecia about 1.5 mm. in diameter, constricted
at the base, margin smooth, light buff, dise cinnamon rufous;
amphithecium about 130 u thick, similar to the thallus in structure
but algae much more abundant and closely tangled and cortex
pseudoparenchymatous, about 50 y thick; parathecium about
40 y thick below, thinning out and disappearing above, pseudo-
parenchymatous with small cells; hypothecium about 80 y. thick,
very deeply staining; thecium about 170 y» tall; paraphyses
filiform, about 2 y in diameter, tips clavate; asci 8-spored,
cylindrical, about 12 y in diameter; ascospores imbricately
[Vor. 20
426 ANNALS OF THE MISSOURI BOTANICAL GARDEN
monostichous, 30-40 x 12-16 u [only 20 x 10 u in Costa Rican
specimens, but obviously immature].
San José: R. Virilla below El Brazil, Dodge 7783.
LEPTOGIUM MicnosTICTUM Vainio, Dansk Bot. Ark. 4": 18.
1926
Type: not stated, specimens cited, Mexico, Palenque, Liebmann
7415, 7416, 7428; Papantla, Liebmann.
Thallus light glaucous blue or a little darker, lobes rounded,
surface smooth but irregularly impressed, margin thin and smooth;
about 225 y thick, corticate on both surfaces with cells about 4 u
in diameter, the filaments of Nostoc very loosely tangled, more
or less parallel to the surface and very scanty in the middle.
Apothecia 0.5-1.0(— 1.8) mm. in diameter, sessile, constricted
at the base, margin thin, whole, concolorous with the thallus,
dise pale to tawny; amphithecium about 60 u thick, cortex
pseudoparenchymatous, about 25 y thick below, thinning out
above to a double layer of cells, algal layer of tangled filaments of
Nostoc; parathecium pseudoparenchymatous, about 40 y thick,
thinning above to about three layers of cells; hypothecium 30 y.
thick, of densely tangled rather large hyphae suggesting pseudo-
parenchyma but cells much smaller than in the parathecium;
thecium about 100 y tall; paraphyses filiform, 1 u in diameter,
ending in the epithecial gel; asci clavate to cylindrical, 12-16 u
in diameter; ascospores distichous, 8 per ascus, fusiform, with
about 5 transverse septa, 20-30 x 8-10 u.
A single small specimen from Costa Rica is doubtfully referred
here. The thallus is somewhat more finely impressed and oc-
casionally near the margin it appears very minutely scrobiculate.
This specimen is from a greater elevation than most other mem-
bers of this subgenus.
San José: Guayabillos and Cabeza de Vaca, 2150-2350 m., Dodge & Thomas 7418.
LEPTOGIUM VESICULOSUM (Sw.) Malme, Ark. f. Bot. 198:
14-15. 1924.
Lichen vesiculosus Sw., Nov. Gen. Sp. Pl. Prodr. 147. 1788.
Lichen bullatum Ach., Lichenog. Suec. Prodr. 137. 1798.
Leptogium bullatum Mont., Ann. Sci. Nat. Bot. II, 16: 113.
1841.
Type: Jamaica, tops of mountains, O. Swartz. L. bullatum Ach.
based on the same material.
1933]
DODGE—LICHENS OF COSTA RICA. I 427
Thallus plumbeous, lobes rounded, more or less appressed,
longitudinally wrinkled with shallower cross ridges; about 100 y.
thick between the ridges, corticate on each surface with a single
layer of isodiametric cells, homoeomerous, of very loosely tangled
filaments of Nostoc running more or less parallel to the surface.
Apothecia sunk in the tips of podetiiform elevations of the
thallus, margin wrinkled but not isidioid or phyllophorous, edge
of parathecium conspicuous light buff, disc concave, rufous;
amphithecium continuous with the thallus without differentiation,
cortex a single layer of cells both above and below; parathecium
100 u thick below, thinning out to 40-60 u above at the margin,
pseudoparenchymatous; hypothecium thick, 80 y, of densely
woven slender hyphae; thecium 120-140 y tall; paraphyses
slender, filiform, cells almost isodiametric above, disintegrating
into the brown epithecial gel; asci cylindrical, 20 » in diameter,
8-spored; ascospores imbricately monostichous, broadly fusiform,
23-32 x 10-13 y, with 5 transverse septa.
This species seems to be widespread in the American tropics,
from 700 to 2100 m., rarely also below these levels. L. stipitatum,
with which this species is easily confused, seems confined to the
lower levels, reaching up to about 800 m., and has so far only
been reported from Guadeloupe and Costa Rica.
Limón: Waldeck, Dodge & Nevermann 7938; Marta, 20 m., Dodge 7427.
Cartago: Angostura, Polakowsky 456; Juan Vifias, 1000 m., Calvert 68; Santiago,
1140-1180 m., Dodge 4555; R. Birrís, 1220-1340 m., Dodge & Thomas 4628; Agua-
caliente, 1240-1460 m., Dodge & Thomas 7082; Carpintera, 1700 m., K. Danielson
101a.
San José: S. Pedro de Montes de Oca, 1200 m., Thomas 4717; S. Juan Tibas,
1000-1100 m., Dodge 4303.
Guanacaste: H. Santamaría, 640-680 m., Dodge & Thomas 6797.
Puntarenas: Boruca, 560 m., T'onduz 5380.
Var. DIGITATUM Eschw. ap. Martius, Fl. Brasil. 1: 238. 1833.
Collema bullatum Raddi, Atti Soc. Ital. 18: 36, pl. 4, f. 2.
Collema bullatum var. dactylinoideum Nyl., Flora 41: 338.
1858; Syn. Meth. Lich. 1: 129. 1858.
Type: Brasil, between Mandiocca and Morro do Frade,
Raddi, type of var. dactylinoideum not cited, but specimens from
Mexico, Fr. Müller, Colombia, Tolima, Goudot, and Bolivia,
Weddell, are mentioned.
This variety is stated by Eschweiler to have a thicker thallus,
[Vor. 20
428 ANNALS OF THE MISSOURI BOTANICAL GARDEN
deep green instead of lead color, densely wrinkled instead of
granular; podetia cylindrical instead of ventricose, 6-12 mm.
instead of 2-6 mm. tall, about 6 mm. in diameter.
From the description this variety does not seem distinct from
the species and I have seen no material referable here from
Costa Rica. It is possible that Eschweiler was attempting to
separate this species from L. phyllocarpum which he called
Collema (Leptogium) bullatum var. sertatum.
LEPTOGIUM STIPITATUM Vainio, Hedwigia 38: (255). 1899.
Type: Guadeloupe Island, near Gourbeyre, P. Duss 434.
Thallus plumbeous, soon discolored buffy citrine to Saccardo’s
olive, irregularly dichotomously branched, ultimate lobes 2-7
mm. broad, then crowded and confluent, rounded, acutely and
irregularly wrinkled above and below; about 200 » thick between
the ridges, with Nostoc filaments densely coiled and tangled near
the surface, mostly parallel to the surface within, much more
abundant than in L. vesiculosum, with cortex of isodiametric
cells on each surface. Apothecia on short thalline pustules,
up to 2.5 mm. in diameter, margin thick with periclinal folds
(never radial as in L. vesiculosum), disc chestnut; amphithecium
about 200 y thick, of the same texture as the thallus; parathecium
80 u thick, disappearing above, of large-celled pseudoparenchyma;
hypothecium 80 y thick, of more or less parallel hyphae continued
upward as a false parathecium; thecium about 140 y tall; para-
physes filiform, about 1.5 » in diameter, septate, with capitate
tips in the brown epithecial gel; asci cylindrical, 16-20 y in
diameter; ascospores distichous, broad-fusiform with acute ends,
24—32 x 10-12 y.
Ban José: cafion of R. Virilla, below El Brazil and Sta. Ana, oe 8019.
Puntarenas: Peninsula of Puntarenas, 3-5 m., Dodge
LEPTOGIUM PHYLLOCARPUM (Pers. ap. pie e Mont., Ann.
Sci. Nat. Bot. III, 10: 134. 1848.
Collema phyllocarpum Pers. ap. Gaudich. in Freycinet, Voy.
Uranie, Bot. 204. 1820.
Collema bullatum var. sertatum Eschw. ap. Martius, FI.
Brasil. 1: 239. 1833.
Type: Brasil, Rio de Janeiro, Gaudichaud.
Thallus plumbeous but frequently becoming black, especially
1933]
DODGE—LICHENS OF COSTA RICA. I 429
in specimens which have been dried slowly or repeatedly wetted
and dried after collection, lobes broad, surface deeply and acutely
wrinkled, the wrinkles predominantly longitudinal or radial,
the secondary wrinkles very irregular; more than 200 y thick
between the wrinkles, corticate on both surfaces by a single
layer of isodiametric cells, the filaments of Nostoc loosely tangled,
more or less parallel to the surface and much more coiled and
densely tangled just under the upper cortex. Apothecia about
2 mm. in diameter, margin with crowded periclinal wrinkles,
dise rufous, immersed in hollow protuberances as in L. vesi-
culosum; amphithecium appearing lobed in section up to 500 y
thick, homogeneous with the thallus but algae a little denser;
parathecium about 100 y. thick below, thinning out to about 20 y.
above at the margin, of large-celled pseudoparenchyma; hypo-
thecium about 30 y thick, of slender densely woven hyphae
continuing up to the margin at about the same width; thecium
about 140 y tall; paraphyses filiform, not dilated above, epithecial
gel brown; asci cylindric, about 20 y in diameter, 8-spored;
ascospores broad-fusiform with acute ends, monostichous, with
5 transverse septa, 22-30(- 35) x 10-12(- 14) y.
Apparently this species is very widespread and common in the
American tropics from low elevations in the southern United
States to 2500 m. in Perú. In Costa Rica it ranges from 500 m.
to about 1900 m., being characteristic in the moister fog-bathed
areas of the temperate region.
Cartago: Santiago, 1140-1180 m., Dodge 8000; R. Birrís, 1220-1340 m., Dodge &
Thomas 7994.
San José: Cañon of R. Virilla below El Brazil, Dodge 7782; Zapote, 1200 m.,
Standley 40273; La Palma, 1500-1700 m., Maxon & Harvey 7891; Quebradillas, near
Sta. María de Dota, 1800 m., Standley 42977; Sta. María de Dota, 1500-1800 m.,
Standley 42424; S. Marcos de Dota, 1200 m., Tonduz 5374; R. Naranjo, Tonduz
5375; S. Gabriel, Tristan 5229; S. José, Polakowsky 113.
Alajuela: R. Ciruela, 920-980 m., Dodge & J. Valerio 4858; Santiago de S. Ramón,
1000 m., Brenes; Viento Fresco, 1600-1900 m., Standley & Torres 47771.
? Puntarenas: Corozal, 5-50 m., Dodge 8025.
Var. MACROCARPUM Nyl., Syn. Meth. Lich. 1: 130. 1858.
Type: not cited but specimens from Aequatorial America,
Humboldt, Chile, Perú, and Venezuela, Lind. 1092, are mentioned.
Similar to the species but apothecia large, 5-9 mm., margins
lobulate, foliose. American material referred to var. daedaleum
[Vor. 20
430 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Nyl. probably belongs here. Distribution in Costa Rica es-
sentially similar to that of the species but not yet collected in the
higher elevations.
Cartago: Santiago, 1140-1180 m., Dodge 8001; R. Birrís, 1220-1340 m., Dodge &
Thomas 4632; Carpintera, 1700 m., K. Danielson 101b.
San José: R. Naranjo, Tonduz 5377.
Alajuela: La Palma de San Ramón, 1250 m., Brenes 91.
Guanacaste; H. Santamaría, 640—780 m., Dodge & Thomas 6776, 8002; Tilardn,
500—690 m., Standley & J. Valerio 44803, Dodge & Thomas 6562.
Puntarenas: Corozal, 5-50 m., Dodge 7529.
Var. CAMPESTRE Malme, Ark. f. Bot. 198: 13. 1924.
Type: Brasil, Bahia, R. Vermelho, Malme 29, 30; Matto
Grosso, Cuyaba, Malme 2715c, etc.; Serra da Chapada, pr.
Bocca da Serra, Malme 2245B.
Differs from the typical form by narrower crowded lobes,
margins crisped; apothecia 1-2 mm. in diameter, margin rugulose,
thin; amphithecium corticate with a single layer of cells; para-
thecium 60-70 » thick, reaching the margin above; thecium 100-
125 y tall, asci up to 20 u thick, 8-spored; ascospores irregularly
distichous, broad-fusiform, acute ends, 22-27 x 8-10 y, usually
with 5 transverse septa.
San José: Turrdcares, 540-600 m., Dodge & Thomas 7481.
Alajuela: R. Ciruela, 920-980 m., Dodge & J. Valerio 7921; Santiago de S. Ramón,
1000 m., Brenes.
LEPTOGIUM CORALLOIDEUM Vainio, Ann. Acad. Sci. Fenn.
A 67:110. 1915.
Leptogium diaphanum f. coralloideum Mey. & Fw., Nova Acta
Acad. Leopold. Carolin. 19: Suppl. 226. 1843.
Leptogium phyllocarpum var. isidiosum Nyl., Syn. Meth.
Lich. 1: 130. 1858.
Leptogium phyllocarpum var. coralloideum Hue, Nouv. Arch.
Museum Paris III, 10: 228. 1898.
The type of var. isidiosum is from Mexico, Orizaba, Fr. M üller.
Thallus as in L. phyllocarpum but densely isidiose, especially
along the wrinkles, somewhat thicker, semi-pellucid. Apothecia
large when present, very rare, margin densely coralloid-isidiose,
dise rufous; amphithecium highly developed, homogeneous with
the thallus; parathecium 140 y thick, disappearing above, of
large thin-walled pseudoparenchyma; hypothecium 60 y thick,
reaching the margin above, of densely woven slender hyphae;
1933]
DODGE—LICHENS OF COSTA RICA. I 431
thecium 200 y. tall; paraphyses filiform with clavate apices; asci
cylindrical; ascospores 8 per ascus, imbricately monostichous,
ellipsoid to fusiform, apices acute, 30-40(-45) x 12-17 u, with 7
transverse septa.
This species is perhaps only a large isidiose variety of L.
phyllocarpum, essentially similar in morphology but larger in
most dimensions. It occupies the same general region as L.
phyllocarpum. It is usually sterile and hence not easily distin-
guished from sterile L. marginellum var. isidiosellum. Material
has been seen from Mexico, Pert, and Brasil.
Limón: Hamburg, 20-30 m., Dodge 7420.
Cartago: Pejivalle, 650-900 m., Standley & J. Valerio, Dodge & Thomas 4337;
Cañon of R. Reventazón below Santiago, 920-1000 m., Dodge & Thomas 8018,
Dodge 8004; R. Birrís, 1220-1340 m., Dodge & Thomas 7954; C. Carpintera, 1320—
1700 m., Dodge 3758, Dodge & Thomas 4762, K. Danielson 98; Aguacaliente, 1240—
1460 m., Dodge & Thomas 8003; R. Reventado, 1460-1650 m., Standley & J. Valerio
614.
one José: R. Virilla below El Brazil, Dodge 6496; Zurquí, H^. d m., Standley
& J. Valerio 48255, 48266, 48146, Dodge, J. Valerio & Thomas 6042.
Alajuela: near Fraijanes, 1500-1700 m., Standley & Torres 47425, 47462
Guanacaste: H. Santamaría, 640—680 m., Dodge & Thomas 8005; Tilarán, 500-650
m., Standley & J. Valerio 44367, 44403.
LEPTOGIUM MARGINELLUM (Sw.) S. F. Gray, Nat. Arrang.
Brit. Pl. 1: 401. 1821.
Lichen marginellus Sw., Nov. Gen. Sp. Pl. Prodr. 147. 1788.
Collema vesicatum Taylor, London Jour. Bot. 6: 196. 1847.
Leptogium corrugatulum Nyl., Syn. Meth. Lich. 1: 182. 1858.
Type: Jamaica, Blue Ridge Mt., Swartz. The type of C.
vesicatum Taylor is from St. Vincents, West Indies. The type
of Leptogium corrugatulum is from Mexico, Jalapa, Galeotti 9630.
Thallus plumbeous, round-lobed, margin smooth or occasionally
microphylline, pellucid, surface of undulating longitudinal
wrinkles less acute than in L. phyllocarpum, of variable thickness,
about 160 x thick, corticate on both surfaces with small isodia-
metric cells. Apothecia minute, up to 0.3 mm. in diameter, margin
densely isidiose, disc rufous; amphithecium thick, with pseudo-
parenchymatous cortex below, a single layer of cells above, as are
also the isidia; parathecium 150 y. thick, pseudoparenchymatous;
thecium 150-180 y tall, epithecium tawny; asci 8-spored; spores
imbricately monostichous, ellipsoid-fusiform, ends acute, 25-35
(-40) x 10-13 y, with 5 transverse septa.
[Vor. 20
432 ANNALS OF THE MISSOURI BOTANICAL GARDEN
This species rarely matures spores although rudimentary
apothecia are nearly always abundant. ‘The microscopic details
of the apothecium are largely taken from Malme as I have seen
no mature asci in my Costa Rican material. There is consider-
able variation in the amount of isidia, as they easily break off
leaving minute foveoles on the margin. The species is quite
distinet and is widespread in the American tropies. I have seen
material from Florida, Alabama, Bermuda, various islands of
the West Indies, Mexico, Brasil, and the Galápagos Islands. It
seems to range from sea level to 1500 m.
From the nature of the isidia it seems likely that var. isidio-
sellum Riddle, Brooklyn Bot. Gard. Mem. 1: 115. 1918, should
be referred to L. coralloideum, although sterile isidiose states are
very difficult to place.
Limón: Marta, 20 m., Dodge & Nevermann 7986; Carmen, Dodge 7422; Waldeck,
Dodge & Nevermann 7421.
Cartago: Pejivalle, 600-900 m., Dodge & Thomas 8014, Standley & J. Valerio
46769; R. Birrís, 920-1100 m., Dodge 7988; Santiago, 1140-1180 m., Dodge 7987;
Aguacaliente, 1240-1460 m., Dodge & Thomas 7937; Carpintera, 1500 m., Dodge 3948.
Alajuela: Santiago de S. Ramón, 1000 m., Brenes 242; S. Pedro de S. Ramón,
700 m., Brenes.
Guanacaste: H. Santamaría, 640—720 m., Dodge & Thomas 6867, 7989; H. Grana-
dilla, above R. 8. José, 480 m., Dodge & Thomas 7936; H. Q. Azul, on lower slope of
V. Tenorio, 400—600 m., Dodge & Thomas 6653; Tilardn, 500-690 m., Dodge & Thomas
61.
Puntarenas: R. Terrones, 30 m., Dodge & Marks 7980; Corozal, 5-50 m., Dodge
7583.
LEPTOGIUM DENTICULATUM Nyl., Ann. Sci. Nat. Bot. V, 7:
302. 1807.
L. tremelloides v. leptophyllinum Mey. & Fw., Nova Acta
Acad. Leopold. Carolin. 19: Suppl. 228. 1843.
L. tremelloides v. azureum f. isidiosum Müll. Arg., Bull. Soc.
R. Bot. Belg. 29: 49. 1891, non Flora 65: 292. 1882.
L. leptophyllinum Zahlbr., Cat. Lich. Univ. 3: 136. 1924.
Type: Colombia, San Jil, 1300 m., Lindig.
Thallus mineral gray or darker, large, broadly lobed, lobes
round, edges smooth, somewhat crisped, surface slightly wrinkled,
100-120 u thick, microphylline, isidiose, corticate on both sides
with a single layer of cells, the short filaments of Nostoc with
cells 4-5 y in diameter mostly assembled just under the cortex,
1933]
DODGE-—LICHENS OF COSTA RICA. I 433
leaving the central portion relatively free of algae. Apothecia
relatively rare but when present usually numerous, scattered
over the upper surface, up to 2 mm. in diameter, margin granular,
thick, becoming isidiose in age, buff, disc rufous; amphithecium
220 y thick, with a layer of pseudoparenchymatous cortex below
about 160 u thick, not thinning out much above and merging
with the parathecium, algal layer 60 y thick below, thinning
out to 30 u above; parathecium 100 y. thick below, thinning out
to about 40 u above, of large thin-walled hyphae at times sug-
gesting pseudoparenchyma; hypothecium 20 » thick, of more
slender deeply staining hyphae; thecium 160 y tall; Bus
slender, ending abruptly in the epithecial gel; asci 10-12 y in
diameter, cylindrical, 8-spored; ascospores imbricately mono-
stichous, fusiform with acute ends and about 5 transverse septa,
about 24 x 10 y.
This species seems widespread in the American tropics, al-
though it is so rarely fertile that it is difficult to be sure of the
determination of much of the sterile material, especially to
separate it from L. cyanescens var. austro-americanum Malme.
As here described, this species is close to, although not identical
with, Malme's variety, while the L. denticulatum Malme seems
to be an isidiose state of or closely related to L. Tuckermant.
Only a careful comparison of all the fertile specimens from
tropical America in the various herbaria of Europe can settle
these problems.
Limón: Marta, 20 m., Dodge & Nevermann 7428; Hamburg, Standley & J. Valerio
48750; Castilla, 20 m., Dodge & Nevermann 7996; Waldeck, Dodge & Nevermann
7997, Dodge 8012.
rtago: R. Reventazón below Santiago, 740-750 m., Dodge 4626, 920-1000 m.,
Dodge & Thomas 7993; Santiago, 1100-1140 m., Dodge 7939; R. Birrís, 1220-1340 m.,
Dodge & Thomas 7992; Estrella, 1600 m., K. Danielson 162; near Cartago, 1500 m.,
K. Danielson 22; C. Carpintera, 1500 m., Dodge 3978
San José: San José, Tonduz 5247.
Heredia: C. Zurquí, 1600-1700 m., Dodge, J. Valerio & Thomas 8048.
Alajuela: C. Mondongo de il PE 750-800 m., Brenes 275; Alto de la Palma
de S. Ramón, 1250 m., Brenes 377.
Guanacaste: H. tannin, 680-780 m., —— E poe 6870, 7014; H. Grana-
dilla above R. Las Cañas, 500-600 m., Thom
Puntarenas: Osa, Corozal, 5-50 m., Dodii m.
LEPTOGIUM AZUREUM (Sw.) Mont. ap. Webb, Hist. Nat. Îles
Canaries 3!: 129. 1840.
[Vor. 20
434 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Lichen azureum Sw. ap. Ach., Lichenog. Suec. Prodr. 137.
1798.
Type: mountains of Jamaica, O. Swartz.
Thallus pellucid, greenish-glaucous-blue to puritan gray,
broadly sinuate-rounded lobes, smooth; margins smooth, 80-
100 uw thick, of loosely coiled and tangled filaments of Nostoc
with ellipsoidal cells 2.5 x 4 u. Apothecia scattered, 1-2 mm.
in diameter, margin smooth, light buff, disc rufous, sessile or on a
short stalk; amphithecium 320 y thick, below with a pseudo-
parenchymatous cortex about 100 » thick, above with a single
layer of isodiametric cells in the cortex; parathecium filamentous,
40 y. thick, poorly developed, of slender hyphae not reaching the
margin; hypothecium 20 y thick, deeply staining; thecium 120-
150 y tall; paraphyses filiform, 1.5-2 u thick, thickened at the
tips to about 4 u; asci clavate-cylindrical, about 18 y in diameter,
8-spored; ascospores imbricately monostichous or somewhat
distichous, fusiform ends acute, with 3 or 5 transverse septa,
22-28 x 8-10 y.
Only a careful study of all the types involved can settle no-
menclature of the group of species centering around L. tremel-
loides. ‘This species was originally collected in South Africa by
Thunberg, and very briefly described by Linné fil. 'The early
lichenologists of the west coast of Europe identified their material
as this species and later reduced Swartz’ Jamaican L. azureum
to synonymy. There is apparently a whole group of species
having approximately the same macroscopic appearance but
wholly different microscopic structure which are at present
referred to this species by various workers. As we have here
defined it (agreeing closely with L. tremelloides Malme), it is a
species of the lowlands coming up the river valleys under favor-
able conditions to about 1000 m. in Costa Rica. It seems
widely distributed at lower elevations in tropical America.
Limón: Marta, 20 m., Dodge & Nevermann 7414, 7426; Hamburg, 20-30 m.,
Dodge & Nevermann 7417; Castilla, 20 m., Dodge 7209, 7419, 7424; Carmen, Dodge
7415; Indiana I. near Siquirres, 60—70 m., Dodge & Thomas 6572; along R. Siquirres,
70-350 m., Dodge, Catt & Thomas 8006, 8007, 8008; Livingston, 80-100 m., Dodge,
Catt & Thomas 6573; Waldeck, Dodge & Nevermann 7412, 7413, 7416; Guápiles, 300-
500 m., Standley 37137.
Cartago: Turrialba, Ørsted; R. Pejivalle, 650-800 m., Dodge & Thomas 4414; R
Reventazón, 920-1000 m., Dodge & Thomas 4604.
1933]
DODGE—LICHENS OF COSTA RICA. I 435
Guanacaste: H. Santamaría, 760-900 m., Dodge & Thomas 6876; Tilarán, 500-690
m., Dodge & Thomas 8009.
Puntarenas: Osa, C. Guaca, 85 m., Dodge 7505; Corozal, 5-50 m., Dodge 7530.
Isla Coco: Snodgrass & Heller.
Leptocium Standleyi Dodge, sp. nov.
Leptogium tremelloides Auct., non Linné fil.
Type: Costa Riea, La Hondura, Standley 37870.
Thallus plumbeus, crassus, laevis vel minute rugulosus, lobis
rotundatis, latis, marginibus integris, 150-170 y crassitudine,
filamentis nostocaceis sub cortice dense implicatis, centro laxe
aut sparse implicatis, corticatum e serie simplice cellularum.
Apothecia ad 3 mm. latitudine, plana, margine integro, tenui,
laevi, fulva, disco rufo; amphithecium inferne 160 u, ad 100 y.
crassitudine superne in margine attenuatum, cortice pseudo-
parenchymatico, 90 v. crassitudine inferne ad una serie cellularum
superne attenuatum, filamentis nostocaceis dense contextum;
parathecium 60 y. crassitudine, hyphis tenuibus dense contextum;
hypothecium flavum, 20-30 y crassitudine, hyphis tenuibus
dense contextum; thecium ad 180 y altitudine; paraphyses
filiformes, tenues, apice non incrassata; asci cylindrici, 20 u
diametro metientes; ascosporae octonae, imbricatim monostichae
vel distichae, fusiformes, 24—28 x 12 y.
Thallus mineral gray or darker, thick, smooth or minutely
and shallowly wrinkled, lobes broad, rounded, margins smooth,
150-170 u thick, filaments of Nostoc densely tangled next the
cortex, loosely tangled and scattered in the center of the thallus,
cortex of a single series of cells on both surfaces. Apothecia
3 mm. broad, plane, margin smooth, thin, tawny, dise rufous;
amphithecium 160 » thick below, thinning to 100 y. above at the
margin, corticate below, 90 y thick, pseudoparenchymatous,
above becoming a single layer of cells, filaments of Nostoc densely
tangled; parathecium 60 y. thick, of slender more or less parallel
hyphae; hypothecium yellowish, 20-30 y thick, of slender densely
woven hyphae; thecium 180 y. tall; paraphyses filiform, slender,
apices not thickened; asci cylindrical, 8-spored, 20 y. in diameter;
ascospores imbricately monostichous or distichous, fusiform,
24-28 x 12 y.
This species is essentially close to L. azureum in its microscopic
characters but differs in its much larger dimensions, its thicker,
[Vor. 20
436 ANNALS OF THE MISSOURI BOTANICAL GARDEN
opaque thallus, and larger apothecia. Superficially it some-
what resembles L. pulchellum but is much less coarsely wrinkled
and differs in microscopic details. In Costa Rica it seems to be
confined to the higher elevations reaching 1700 m. It is probable
that material determined as L. pulchellum from Costa Rica by
Müller Argau belongs here, although I did not have time to
section it. It certainly is not typical L. pulchellum.
Cartago: Turrialba, Ørsted; Orosf, 1000-1100 m., Standley 39791.
San José: San José, 1130 m., Standley 41233; La Hondura, 1300-1700 m., Standley
$7870, TYPE.
Alajuela: Pods, Tonduz 5301.
Leprocium Tuckermani Dodge, sp. nov.
Leptogium tremelloides var. minor Tuck. in herb.
? Leptogium moluccanum Vainio, Étude Lich. Brésil 1: 223-224.
1890, non Collema moluccanum Pers. ap. Gaudich. in Freycinet,
Voy. Uranie, Bot. 203. 1826.
Type: Cuba, Monte Verde, C. Wright 56.
Thallus plumbeus vel obscurior, lobis confertis, rotundatis,
superne inferneque sat laevigatus, tomento isidiisque destitutus,
tenuissimus, 35-60 y crassitudine, strato nostocaceo 25 y, corti-
cibus pseudoparenchymaticis cellulis magnis. Apothecium minu-
tum, 0.5-0.9 mm. diametro metiens, peltatum, basi constrictum,
margine integro, pallescente, disco castaneo; amphithecium
corticatum cellulis pseudoparenchymaticis, 60 u crassitudine
inferne, attenuatum ad 12 y superne, strato nostocaceo 40 u
crassitudine; parathecium pseudoparenchymaticum tenue, 10-
12 » crassitudine, cellulis parvis; hypothecium 20 y crassitudine,
hyphis tenuibus dense contextum; thecium 90-120 y. altitudine;
paraphyses filiformes, 2 y diametro, apicibus incrassatis; epi-
thecium brunneum; asci cylindrici, 12 & diametro metientes;
ascosporae octonae, imbricatim monostichae, apicibus acutis,
murales, septis transversalibus 3-5, cellulis haud numerosis,
18-22 x 10-12 y.
Thallus mineral gray or darker, lobes crowded, more or less
elevated and crisped, rounded, smooth on both sides, 35-60 y
thick, algal layer of Nostoc about 25 u, with pseudoparenchyma-
tous cortex above and below of large cells. Apothecia minute,
0.5-0.9 mm. in diameter, constricted at the base, margin whole,
smooth, pale, dise chestnut; amphithecium with a thick pseudo-
1933]
DODGE—LICHENS OF COSTA RICA. I 437
parenchymatous cortex below, about 60 y, thinning to two rows
of cells about 10-12 u thick above; parathecium thin, pseudo-
parenchymatous, 10-12 u thick, of small cells, inconspicuous
and often reported absent; hypothecium 20 » thick, of slender
densely woven hyphae; thecium 90-120 u tall; paraphyses
filiform, 2 v in diameter, with clavate apices; epithecium brown;
asci cylindric, 12 y in diameter; ascospores 8 per ascus, imbricately
monostichous, ends acute, muriform, with 3-5 transverse septa,
cells not numerous, 18-22 x 10-12 y.
This species has long been confused with L. moluccanum, L.
Mariannum, and L. diaphanum. It differs from the two former
in habit and color and from the latter by its structure. It is
apparently widespread in the American tropics, being reported
from Paraguay, and from Brasil in the states of Rio Grande do
Sul and Matto Grosso by Malme, and in Minas Geraes by Vainio.
Limón: Hamburg, Dodge 7425.
Cartago: Pejivalle, 680 m., Dodge & Thomas 4556; Santiago, 1140-1180 m., Dodge
4555, 8044; Las Concavas, 1350-1450 m., Dodge 6842.
San José: hills above Sta. Ana, Dodge 7781.
Guanacaste: H. Santamaría, 680—780 m., Dodge & Thomas 8043; H. Granadilla,
480 m., Dodge & Thomas 8045.
LEPTOGIUM SIMPLICIUS Vainio, Ann. Acad. Sci. Fenn. A6’:
109. 1915.
Type: S. Domingo, La Cumbra, C. Raunkiaer.
Thallus between light drab and light grayish olive, with
irregularly incised lobes 5-7 mm. broad, not isidiose, with slightly
undulate elevated acute wrinkles on both surfaces, corticate with
a single layer of isodiametric cells. Apothecia broadly adnate,
constricted at the base, sessile, 1.5-2.2 mm. in diameter, disc
plane, rufous, margin thin, smooth, lighter than the thallus;
amphithecium 140-150 u thick, corticate with a single series of
cells above, pseudoparenchymatous below, 90-100 wœ thick;
parathecium 120 u thick below, thinning out to 20 u thick at
the margin above, pseudoparenchymatous; hypothecium fila-
mentous, 30 y thick; thecium 150-170 u thick; paraphyses
filiform, tips somewhat thickened in the brownish epithecial gel;
asci cylindric, 14-16 y in diameter, 8-spored; ascospores imbri-
cately monostichous, fusiform, thin-walled, with 5 transverse
septa, 22-30 x 11-13 y.
[Vor. 20
438 ANNALS OF THE MISSOURI BOTANICAL GARDEN
This species previously reported only from the type locality
has yet been found only at low elevations on the Pacific slope.
San José: Turrücares, 540—600 m., Dodge & Thomas 8047.
Guanacaste: Liberia, 100 m., Dodge, Alfaro & Thomas 6586, Dodge & Thomas 8028,
Tilarán, 500-690 m., Dodge & Thomas 8046.
Var. pichneoides Dodge, var. nov.
Type: Costa Rica, Guanacaste, H. Santamaria, Dodge &
Thomas 8029.
Thallus isidiosus, marginibus microphyllinis aut isidiosis.
Apothecium margine tenuiore, laevi, thecium 130-150 y altitudine.
Thallus isidiose, margins varying from dentate to short isidiose
or slightly microphylline. Apothecia with thinner, smooth
margins, thecium 130-150 y. tall.
Only the type has been found fertile and here there are very
few apothecia. The other collections cited probably belong
here, but it is always difficult to determine sterile isidiose speci-
mens in this genus.
Alajuela: R. Ciruela, 920-980 m., Dodge & J. Valerio 4894.
Guanacaste: H. Santamaría, 640-680 m., Dodge & Thomas 8029.
Besides the species cited above, a single small fragment was
found which seems to belong in Leptogiopsis, but more material
is necessary for a satisfactory disposition.
PANNARIACEAE
Thallus squamose or foliose, not gelified; hypothallus and
rhizinae usually highly developed, heteromerous, cortex of
erect, irregular or periclinal hyphae, usually more or less pseudo-
parenchymatic; medulla well developed, with Nostoc, Scytonema,
or Dactylococcus; lower cortex of thick-walled periclinal hyphae
often thin or wholly absent. Apothecia marginal or scattered
over the upper surface, biatorine or lecanorine; paraphyses
unbranched; asci 8-spored; spores hyaline, unicellular, rarely
2-4-celled ; spermatia short, straight.
'This family seems to be of southern origin with some genera
confined to the southern hemisphere, while a few species come
northward until they are cireumpolar. In general, the northern
species are much smaller and more depauperate than the southern
species.
In many ways the family seems very homogeneous, so that
1933]
DODGE—LICHENS OF COSTA RICA. I 439
genus distinctions are often rather arbitrary. Hue, in 1912,
reduced the whole family to a single genus, but his sections and
subsections correspond more or less closely to the genera of
earlier workers. Zahlbruckner has recognized twelve genera,
although the generic delimitation is not altogether clear. Fol-
lowing this treatment I have tentatively recognized seven genera
from Tropical America. Psoroma, with Dactylococcus algae, is
largely confined to the southern hemisphere, and by many authors
has been placed in the Lecanoraceae. Lepidocollema, with a thin
upper cortex and almost homoeomerous, is very close to the
Collemaceae and is known from a single collection in Brasil.
The other five genera are much better known and widespread in
the American tropics.
The morphology of the thallus is not altogether clear and the
nomenclature of the parts is rather confusing. In the ideal case,
the uppermost layer of the thallus, here called ‘‘tomentum,”’
is a mass of loosely woven hyphae formed by proliferation of
deeper layers. In Erioderma it is soft and spongy, like a loose
felt, while in Malmella and in some of the isidiose members of
Pannaria it has the appearance of a fine blotting paper, or in
Coccocarpia the hyphae are more conspicuously parallel and
conglutinate, suggesting a frayed cotton yarn. In some species
of each genus the tomentum is partly evanescent, or occasionally
it is conglutinated into spines or warts. Below the tomentum is
an upper cortex of pseudoparenchyma, usually formed by the
partial disintegration of a palisade of large thick-walled hyphae
which occasionally penetrate between the algal filaments. In
Coccocarpia the pseudoparenchyma is thin and scarcely distin-
guishable from the tomentum or even absent. The algal zone
varies from a dense palisade of filaments of Scytonema and
hyphae in some species, to ellipsoidal colonies of Nostoc sur-
rounded by hyphae. The medulla in most genera consists of
loosely tangled hyphae, occasionally not even well differentiated
from the algal layer, while in Coccocarpia it is formed of septate
periclinal hyphae, scarcely to be distinguished from the lower
cortex except in color and the size of the intercellular spaces.
The lower cortex may be completely absent or consist of one or
more layers of thick-walled periclinal hyphae which give rise to
[Vor. 20
440 ANNALS OF THE MISSOURI BOTANICAL GARDEN
tufts of rhizinae. The so-called hypothallus in most species is a
thick dense mat of rhizinae, which may be absent in Erioderma,
or occasionally only slightly developed in certain species of the
other genera.
The morphology of the apothecium is equally confusing. In
Pannaria and Malmella an amphithecium is present, usually
with a pseudoparenchymatous cortex, and a thin algal layer
which may be separated from the parathecium by a thin medulla.
The parathecium is pseudoparenchymatous, formed from thick-
walled periclinal hyphae in Lepidocollema, Parmeliella, Cocco-
carpia, Erioderma, and the smooth species of Pannaria, while in
the isidiose species of Pannaria and in Malmella the parathecium
is filamentous, in the former of large thin-walled cells, in the
latter of slender hyphae. The hypothecium is differentiated
somewhat in the species of Pannaria but practically undiffer-
entiated in the other genera. Instead of the above interpretation
those with a filamentous parathecium might be conceived
as lacking a parathecium and having a somewhat highly developed
hypothecium, or perhaps the so-called parathecium is really only
a compact medulla of the amphithecium. Further study of the
morphology of the developmental stages, also more representa-
tives of some of the genera, are necessary before the correctness
of any interpretation can be assured.
KEY TO THE TROPICAL AMERICAN GENERA OF PANNARIACEAE
Thallus with prm apothecia lecanorine, rhizinae little developed
es a es oe KG eae a eee hee EE TELE LU Psoroma cinchonarum
Thallus with het rhizinae usually well developed.
Upper cortex thin, pseudoparenchymatous, algal layer occupying most
of the thallus; apothecia biatorine; Brasil.......... Lepidocollema carassense
Upper cortex well developed, peeudoparenchymat
Upper cortex of two or more rows of cells; bros etd to the
surface, medulla loosely woven.
Apothecia lecanorine.
m absent or very early evanescent, algae definitely
Nostoc, colonies — separate; parathecium usually
tou
Tomentum thin but kai algae Nostoc ? or perhaps Scytonema;
parathecium of slender hyphae................. 0000000 Malmella
Apothecia biatorine, pseudoparenchymatous from periclinal hyphae.
Tomentum absent, algae Nostoc; rhizinae well developed... .Parmeliella
Tomentum present and highly developed, algae Shiai or
Nostoc; lower cortex little developed; apothecia large, mar-
| e ee E err E ree rioderma
1933]
DODGE—LICHENS OF COSTA RICA. I 441
Upper cortex of septate periclinal hyphae which may simulate pseudo-
parenchyma, tomentum usually absent; apothecia biatorine, medulla
of periclinal conglutinate hyphae appearing pseudoparenchymatous,
des e e 5 5 ck enh s 8s cc res ceves ces oe aes Oe aR Coccocarpia
PANNARIA Del. ap. Bory
PANNARIA Del. ap. Bory, Dict. Class. Hist. Nat. 13: 20. 1828.
Type species: Pannaria rubiginosa (Thunberg ap. Ach.) Del.
ap. Bory.
Thallus granular, squamose to foliose with a well-developed
bluish black or black hypothallus, rarely with dark, more or less
tangled rhizinae below, heteromerous, upper surface corticate
with large-celled pseudoparenchyma formed from a palisade of
hyphae; algae Nostoc, medulla single or double, in the former
case arachnoid, in the latter the upper portion of more or less
parallel, thin-walled hyphae, loosely woven, and below of densely
tangled hyphae, without lower cortex. Apothecia at first sunk
in the thallus, finally sessile or peltate, superficial; amphithecium
pseudoparenchymatous with a few algae in the center; hypo-
thecium hyaline; asci clavate, 8-spored; ascospores hyaline,
elongate, ellipsoidal to almost fusiform with a somewhat thick-
ened and finely verrucose wall. Spermagonia in hemispherical
warts, spermatiophores septate with short, broad cells; spermatia
straight or very slightly curved, elongate, cylindrical.
This genus seems one of the most variable and widespread of
the family, occurring from the Arctic to the Antarctic, in Costa
Rica occurring from sea level to 1700 m.
In tropical America the genus separates easily into two sections,
the isidiose species with a filamentous parathecium, and the
smooth species with a pseudoparenchymatous parathecium.
KEY TO THE TROPICAL AMERICAN SPECIES OF PANNARIA
Hypothallus not well developed, pale with pale rhizinae below.
Thecium 80-100 x or less tall; parathecium filamentous; hypothecium not
differentiated; spores 11-12 x 4-5 u, more or less constricted at the
middle and appearing 2-celled; S. Afriea............... suus P. rubiginosa
Thecium 100-120 u or more tall; parathecium of septate hyphae; hypo-
thecium pseudoparenchymatous; spores 13-16 x 7-9 u; Rio Grande
do Sul, Brasil [P. rubiginosa Malme non aliorum]. ............... P. Malmei
Hypothallus developed, sometimes sparingly.
Hypothallus pale, thallus squamulose, pale olivaceous to rusty yellow,
margin cut-crenate, irregularly lobulate; spores 8-9 x 2.5-3 u; Mexico.
Ee ate ce s uet a iho whe 24 M Cee TOOT er e P. applanata
[Vor. 20
442 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Hypothallus black.
Thallus isidioid.
Margins and surface with cylindrical, coralloid branched c
spores 14-19 x 7-8.5 y; Antilles and Brasil............... . stylophora
Margins isidio-lacinulate; spores 8-10 x 5-6 u, with thick epispore;
Minos Gurus, BEAD... oee khao hmm ont ct COR ead P. isidioidea
Margins with verruciform isidia, lobes under 1 mm. broad; apothecia
1-1.5 mm. in diam.; spores 12-14 x 8-10 4, wall thin and smooth.
Thallus without isidia or soredia
Thallus livid fuscescent, adini broad-lobed; spores 14 x 6 4; hypo-
thallus less developed, center glebulose-lobed; São Paulo, Brasil.
Duns Notas EMO EE T E TESTEN EE . imbricatula
— pallid or ashy or pale glaucous, lobes subeuneate, up to
m. broad; spores 12-15(-18) — ig 10) u; parathecium and
iS »othecium pseudoparenchymatous.............6 2000.00: P. Vainii
Thallus coeruleo-ashy, lobes tna peat epithecium black
aeruginous; spores 12-20 x 7-8 yu, surface subplicate; São Paulo
en, Temi P. caeruleo-nigricans
Thallus livid glaucescent or lurid pallid, laciniae very Peeler
radiate-stellate, even subimbricate............. 00.00.00 eee . radiata
Thallus pale fuscescent, lobes 2-2.5 mm. broad and about 6 mm.
long; epithecium rufous; spores 12-19 x 8-10 yw; parathecium
Bici arose vndis hypothecium of conglutinate vertical
ivl: DM A ooeviséveré VC he E443 443 Axio ERRORES P. brasiliensis
PANNARIA ISIDIOIDEA Vainio, Ann. Acad. Sci. Fenn. A6’:
102. 1915.
Pannaria Mariana var. isidioidea Vainio, Étude Lich. Brésil
1: 206. 1890, excl. syn.
Type: Brasil, Minas Geraes, Sitio, 1000 m., Vainio 669, 983
(sterile). The following description is based on fertile specimens
from Costa Rica.
Hypothallus of dark green rhizinae extending only slightly
beyond the thallus; thallus dark olive buff, margins lighter,
surface very minutely tomentose, pinnatifid dissected, lobes
somewhat cuneate, about 1 mm. wide, ultimate lobules rounded,
0.25—0.5 mm. broad, toward center becoming microphylline and
subisidiose but idis somewhat flattened, rarely cylindric, 150-
160 u thick, upper 40 u of pseudoparenchymatous cortex, algal
layer of Nostoc about 40 y thick, and medulla 40 y thick, of
loosely woven hyphae with occasional small chains of Nostoc;
rhizinae thick-walled, black, 6-8 yw in diameter. Apothecia
crowded, peltate, constricted at the base, about 1 mm. in di-
ameter, margin prominent, crenulate, almost lobulate, disc
1933]
DODGE—LICHENS OF COSTA RICA. I 443
chestnut; amphithecium 200 y thick, of the same texture as the
cortex and algal layer of the thallus; parathecium 60 y thick,
filamentous, of large thin-walled periclinal hyphae, ending above
in a false pseudoparenchyma; hypothecium about 20 y thick,
of similar structure but more deeply staining; thecium 80-100 y
tall; paraphyses slender, filiform, somewhat clavate above in the
brownish epithecial gel; asci clavate, 12 » in diameter; ascospores
ellipsoidal with a thick epispore, 8-10 x 5-6 u (perhaps still
immature).
This species is somewhat variable in appearance and perhaps
should be separated into varieties and forms but I have not seen
sufficient material to do so properly. Sterile material has been
referred here solely on the structure of the thallus. Dodge &
Thomas 6558 from Tilarán has a white reticulate cortex.
Cartago: R. Birrís, 1220-1340 m., Dodge & Thomas 4559.
Heredia: C. Central de Zurquí, 1600-1700 m., Dodge, J. Valerio & Thomas 6048.
Alajuela: Piedades de S. Ramón, 900 m., Brenes 375.
Guanacaste: H. Santamaría, 640-680 m., Dodge & Thomas 6823, 6911, 6988;
Tilarán, 500-690 m., Dodge & Thomas 8059, 6558.
Var. pulvinata Dodge, var. nov.
Type: Costa Riea, Alajuela, C. de Pata de Gallo a S. Rafael de
S. Ramón, 1200-1250 m., Brenes.
Thallus ut in P. isidioidea sed isidiosissimus, marginibus
adscendentibus, cinerascentibus; isidia coralloidea, elongata,
pulvinum formantia.
Thallus as in P. tsidioidea but densely isidiose with ascending,
cinerascent margins; the isidia coralloid, repeatedly branched,
erowded, forming a cushion up to 2 cm. in diameter and 3-4 mm.
thick.
This variety might easily be mistaken for a sterile pulvinate
Stereocaulon or a Siphula, were it not for the occasional lobe at
the margin which shows its relationship.
Alajuela: C. de Pata de Gallo a S. Rafael de S. Ramón, 1200-1250 m., Brenes.
PANNARIA Moseni Dodge, sp. nov.
Pannaria rubiginosa Malme, Ark. f. Bot. 20%: 7-8. 1924,
pro parte.
Type: Brasil, São Paulo, Sororocaba near Santos, Mosen 3284.
Thallus albidus, subpinnatifidus, lobis 3-5 mm. longitudine,
0.8 mm. latitudine, apicibus rotundatis, marginibus loborum
[Vor. 20
444 ANNALS OF THE MISSOURI BOTANICAL GARDEN
verrucosis, vel etiam in centro thalli isidiosus, ad 140 y crassitu-
dine, cortex superior 30-35 y crassitudine, pseudoparenchymaticus
ex hyphis perpendicularibus formatus, zona nostocacea coloniis
subsphericis 35 y diametro metientibus, medulla ad 60 y crassi-
tudine hyphis laxe implexis, rhizinae brunneo-nigricantes, hyphis
nigris pachydermaticis 3-4 u diametro, conglutinatis. Apothecia
1-1.5 mm. diametro, marginibus tenuibus, crenato-lobatis et
subisidiosis; amphithecium inferne 120 y. crassitudine superne ad
60 y attenuatum, cortice 40 y crassitudine, pseudoparenchy-
matico; parathecium bene evolutum, inferne 40 u superne ad
60 u crassitudine, hyphis ad 4 y diametro leptodermaticis con-
textum; hypothecium 25 y crassitudine, hyphis periclinalibus
dense compactum; thecium ad 100 y altitudine; paraphyses
filiformes 1-2 u diametro apicibus clavatis; asci clavati, 25 u
diametro; ascosporae octonae, distichae, 12-14 x 8-10 u, epi-
sporio tenuiusculo.
Thallus pearl gray, subpinnatifid, lobes 3-5 x 0.8 mm., with
rounded tips, margins of lobes verrucose or even occasionally
cylindric, isidiose toward the center, about 140 y thick, upper
cortex 30-35 u, pseudoparenchymatous from the breaking up of
a palisade layer, algal zone of subspherical colonies of Nostoc
about 35 u in diameter, medulla about 60 u thick, of loosely
woven hyphae; rhizinae of brownish black thick-walled conglu-
tinate hyphae 3-4 y in diameter. Apothecia 1-1.5 mm. in
diameter, margins thin, crenate-lobulate or even somewhat
isidiose; amphithecium about 120 y thick below, thinning to
about 60 u above with a cortex about 40 u thick of pseudoparen-
chyma; parathecium well developed, about 40 u below, spreading
to 60 u above, of interwoven thin-walled hyphae about 4 u
in diameter; hypothecium about 25 y thick of dense periclinal
hyphae, staining very deeply; thecium about 100 y tall; para-
physes filiform, 1-2 y in diameter, with clavate tips; asci clavate,
25 y in diameter, 8-spored; ascospores distichous, 12-14 x 8-10 y,
with a somewhat thinner epispore.
It is with some hesitation that I have described this species as
new. ‘The thallus is very suggestive of the sterile P. stylophora,
although it has a somewhat different color and lobing and rather
larger dimensions of the parts. It does not seem to be closely
1933]
DODGE—LICHENS OF COSTA RICA. I 445
related to P. rubiginosa, originally described from Thunberg’s
South African collections, and is certainly not closely related to
the material referred to that species in the North Temperate
zone. So far I have only seen this species from the Atlantic
Coastal Plain of Costa Rica.
Limón: Hamburg, Standley & J. Valerio 48702, 48717; R. Siquirres, 70-200 m.,
Dodge, Catt & Thomas 8037.
PANNARIA STYLOPHORA Vainio, Ann. Acad. Sci. Fenn. A6’:
102. 1915.
Type: Antilles, Guadeloupe, Sofaga, P. Duss 1387 (sterile).
Hypothallus black, extending slightly beyond the thallus;
thallus with pale olive gray margins and olive gray center, lobes
long, slender, oblong, 0.5-0.8 mm. wide, and primary lobes
about 15 mm. long, irregularly more or less pinnatifid, smooth
above around the margin, the center with scattered to crowded
cylindrical isidia which are rarely dichotomously branched, and
concolorous, about 125 y thick, upper cortex 20-25 yu, pseudo-
parenchymatous from the development of a palisade layer; algal
layer about 40 y thick, consisting of ellipsoidal colonies of Nostoc
about 30 x 40 u; medulla about 60 y. thick, loosely woven; lower
cortex absent; rhizinae of black thick-walled hyphae about 3-
4 y in diameter.
Limón: R. Siquirres, 70-200 m., Dodge, Catt & T'homas 8036.
PANNARIA Vainii Dodge, sp. nov.
Pannaria rubiginosa Vainio, Étude Lich. Brésil 1: 204—205.
1890; Malme, Ark. f. Bot. 20%: 7-8. 1924, quoad spec. Dusenia-
num.
Type: Brasil, Minas Geraes, Sitio, 1000 m., Vainio 606.
Hypothallus black, extending 1-2 mm. beyond thallus; thallus
between olive buff and smoke gray, irregularly and repeatedly
laciniate, lobes 1-2 mm. broad, ultimate lobes short, broader
ones up to 4 mm. long, smooth above, not isidiose, about 160-
170 u thick, upper cortex 20-30 y thick of 2-3 layers of large
isodiametric cells; algal layer of spherical to ellipsoidal colonies
of Nostoc about 60-65 y. in diameter, algae rather large and some-
what angular by mutual pressure, medulla about 80 y thick of
large, thin-walled hyphae, loosely interwoven; rhizinae of black,
thick-walled, conglutinate hyphae. Apothecia peltate, 0.8-2
[Vor. 20
446 ANNALS OF THE MISSOURI BOTANICAL GARDEN
mm. in diameter, margin thick, crenulate, dise rufous; amphi-
thecium 130-440 y. thick, with cortex 20-30 » thick; parathecium
well developed, about 60 y thick of periclinal large, thin-walled,
hyphae simulating pseudoparenchyma; hypothecium about 20 u
thick, of similar structure but of more slender less frequently
septate hyphae; thecium 90-100 y tall; paraphyses 1.5-2.5 w
in diameter, filiform, slightly clavate at the tips in a yellowish
brown epithecial gel; asci 14-16 » in diameter, 8-spored; asco-
spores distichous, broadly ellipsoidal, 12-18 x 8-10 y, with thick
epispore.
This species is another of the group which in various herbaria
is often more or less indiscriminately determined as P. rubiginosa,
P. Mariana, or even P. nigrocincta, usually without much critical
attention to microscopic details.
Guanacaste: near Tilarán, both Atlantic ye Pacific slopes, 500-690 m., Standley
& J. Valerio 44810, Dodge & Thomas 6564, 6565
PANNARIA radiata (Vainio) er pn inei nov.
Pannaria Mariana f. radiata Vainio, Ann. Acad. Sci. Fenn.
A6': 101-102. 1915, excl. syn.
Type: Antilles, Guadeloupe, P. Duss.
Hypothallus brownish black, extending about 1.5 mm. beyond
the margin of the thallus, thick; thallus mineral gray with ends of
lobes pale smoke gray, dichotomously (rarely trichotomously)
divided, laciniae 1 cm. or more long, about 1 mm. broad, tips
rounded, reflexed, wholly discrete in outer centimeter, thallus
about 90 y thick, upper cortex of large thin-walled pseudo-
parenchyma 20-30 y thick, algal zone of Nostoc colonies ellip-
soidal, about 40 y in long diameter, medulla about 20 y. thick of
loosely woven hyphae; rhizinae of conglutinate black thick-
walled hyphae very closely woven at the lower side of the medulla
and performing the function of a lower cortex. Apothecia mostly
about 1 mm. in diameter, occasionally up to 2 mm., peltate,
margin crenate, of medium prominence, dise rufous; amphi-
thecium 160 y thick, cortex about 60 y thick, pseudoparenchy-
matous, Nostoc colonies smaller than in the thallus, separated by
strands of pseudoparenchyma; parathecium pseudoparenchyma-
tous, 60-80 y. thick, filamentous, of large very thick-walled hyphae
appearing pseudoparenchymatous; hypothecium 15-20 y thick,
1933]
DODGE—LICHENS OF COSTA RICA. I 447
of periclinal, thin-walled, slender hyphae; thecium 120-130 u
tall; paraphyses slender, filiform, septate, without clavate tips;
asci 10-12 y thick, 8-spored; ascospores irregularly distichous,
ellipsoidal, 15-16 x 7-8 u, with a very thick epispore.
This species shows little relationship to P. Mariana Fr. of the
Old World, a species to which it has been referred. It seems
closest to P. Vainii but differs in microscopic details and habit.
Limón: T Standley & J. Valerio 48768; R. Siquirres, 70-200 m., Dodge,
Catt & Thomas 5
MALMELLA Dodge
Malmella Dodge, gen. nov.
Type species: Erioderma physcioides Vainio.
Thallus squamosus aut microfoliosus, rhizinis nigris bene
evolutus, tomentum superficiei superioris parce evolutum tenue,
hyphis implexis sed non conglutinatis ut in Coccocarpia; cortex
superior pseudoparenchymaticus, algae nostocaceae; medulla
hyphis laxe implexis, cortex inferior tenuis hyphis nigris con-
glutinatis aut nullus. Apothecia lecanorina, sessilia, sparsa;
amphithecium adest; parathecium non pseudoparenchymaticum;
paraphyses filiformes; asci clavati; ascosporae octonae, ellip-
soideae, episporio crasso.
Thallus squamose or small foliose, with a well-developed
hypothallus of black rhizinae; tomentum of the upper surface
less well developed than in Erioderma, of slender hyphae loosely
woven, not conglutinate as in Coccocarpia; upper cortex pseudo-
parenchymatous, algae Nostoc ?, medulla of loosely tangled hyphae,
lower cortex of black hyphae as in Coccocarpia or wholly absent.
Apothecia lecanorine, sessile, scattered over the upper surface,
not marginal as in Érioderma; amphithecium present; parathe-
cium of slender hyphae, not pseudoparenchymatous; paraphyses
filiform; asci clavate, 8-spored; ascospores ellipsoidal, with a
thick epispore.
This genus seems intermediate between Pannaria, Erioderma,
and Coccocarpia, and collections have been previously referred
to each of these genera. In structure of the thallus it seems
intermediate between Pannaria and Erioderma, while in super-
ficial appearance it suggests the more laciniate species of Coc-
cocarpia.
[Vor. 20
448 ANNALS OF THE MISSOURI BOTANICAL GARDEN
KEY TO THE TROPICAL AMERICAN SPECIES OF MALMELLA
Thallus becoming densely isidiose.
Thallus olivaceous with isidia becoming pruinose and subsorediate, then
Done 3 o1 EET ORNETTE PERPE RSS DDR . Randii
Thallus white, more laciniate............. 00.000 nnn BIA M. albida
Thallus not isidiose, medulla not well differentiated from algal zone.
Thallus white, 120 u thick, thecium 140-160 y tall.............. M. Santamariae
'Thallus olive buff or darker
Thallus olive buff, 120 4 thick, thecium 100-120 à tall........... M. physcioides
Thallus dark olive with white pruina about margins, 200 y thick,
thocium 80-80 p Wl. s ioosasaso uae e aH rera 1. caestocinerea
MALMELLA Randii Dodge, n. sp.
Type: Maine, Mt. Desert Island, E. L. Rand.
Hypothallus rhizinis viridi-nigricantibus; thallus olivaceus,
lobis subdichotomis aut irregularibus, 1.0-1.3 mm. latis, margini-
bus crispatis elevatisque, verrucosis et etiam isidioideis, glaber in
lobis marginalibus, centro isidiosissimus, apicibus isidiorum
pruinosis et subsorediosis; thallus 140-160 u crassitudine, to-
mentum 12 y crassitudine hyphis laxe implexis, tenuibus, sub-
dilabentibus; zona nostocacea [vel scytonematica] 100 y crassi-
tudine, filamentis dense convolutis, in coloniis ellipsoideis,
cellulis sphericis vel angularibus; medulla hyphis laxe implexis
2-3 u diametro, aut plus minusve hyphis subparallelis ut in
Coccocarpia pellita, frequenter ad margines; cortex inferior
hyphis conglutinatis nigris tenuis.
Hypothallus of well-developed greenish black rhizinae extend-
ing about 0.5 mm. beyond the thallus; thallus ecru olive to buffy
olive, main lobes subdichotomous or wholly irregular, 1.0-1.3
mm. broad, margin crisped and elevated, becoming verrucose
and finally isidioid, surface appearing like surface of blotting
paper under microscope, smooth on marginal lobes, becoming
densely isidiose toward the center, the tips of the isidia soon
becoming pruinose and subsorediate, giving a puritan gray ap-
pearance to the plant; thallus about 140—160 u thick, the tomen-
tum 12 y thick of loosely appressed, somewhat tangled, slender
hyphae which more or less disintegrate; the upper cortex 20 u
thick, of about 2 layers of cells from the palisade; algal zone
about 100 » thick, of closely coiled filaments, forming ellipsoidal
colonies, cells spherical to angular from mutual pressure, Nostoc
or perhaps Scytonema; medulla of loosely woven hyphae 2-3 y
1933]
DODGE—LICHENS OF COSTA RICA. I 449
in diameter, in places more or less parallel as in Coccocarpia
pellita, especially at the margin of the thallus with about 1 row
of agglutinated black thick-walled hyphae functioning as a lower
cortex, giving off rhizinae either as single hyphae or as small
fascicles of hyphae; sterile.
This species has not been seen fertile and its systematic position
is uncertain. It has somewhat the appearance of an isidiose
Malmella physcioides but differs from the latter in a less well-
developed tomentum, a thinner upper cortex, and a thicker algal
zone. It differs much more from Coccocarpia cronia in the
loosely tangled tomentum layer which is present in Coccocarpia
but closely appressed and formed of longitudinal hyphae, in the
loosely tangled medulla (in Coccocarpia almost pseudoparenchy-
matous of periclinal hyphae), and the much less developed lower
cortex.
MALMELLA albida Dodge, sp. nov.
Type: Costa Rica, Limón, R. Siquirres, 70-200 m., Dodge,
Catt & Thomas 8060.
Hypothallus crassus, niger, bene evolutus; thallus albidus,
cinerascens, laciniatus, di- vel trichotomus aut irregularis, laciniis
linearibus, 0.2-0.5 mm. latis, et superficiei marginibusque dense
isidiosis, coralloideis, inferne niger, rhizinis nigricantibus; thallus
80-100 u crassitudine, cortex superior hyphis periclinalibus
5 u et cellulis pseudoparenchymatibus 20 y. metiens, zona nosto-
cacea (scytonematica?) 60 y crassitudine, filamentis contortis,
cellulis sphericis, 4-5 u diametro; medulla 40-45 y. crassitudine,
hyphis laxis implexis; cortex inferior 8-12 y. crassitudine, hyphis
nigris pachydermatibus conglutinatis compactus. Sterilis.
Hypothallus thick, black, well developed; thallus chalk-
white, becoming ashy from isidia, laciniate, di- or trichotomous
or wholly irregular, laciniae linear, 0.2-0.5 mm. broad, upper
surface and margins densely isidiose, coralloid, below black with
black rhizinae; thallus 80-100 » thick, upper cortex about 25 y
thick, the outer 5 v. of periclinal hyphae, below which are about
two layers of pseudoparenchymatous cells; the algal layer of
Nostoc (Scytonema ?) about 60 » thick, of contorted filaments of
spherical cells 4-5 y. in diameter; medulla 40-45 y. thick, of loosely
tangled hyphae; lower cortex 8-12 y thick, of thick-walled,
[Vor. 20
450 ANNALS OF THE MISSOURI BOTANICAL GARDEN
black, conglutinate hyphae; sterile. Known only from a single
small collection in Costa Rica.
Limón: R. Siquirres, 70-200 m., Dodge, Catt & Thomas 8060.
MALMELLA Santamariae Dodge, n. sp.
Type: Costa Rica, Guanacaste, H. Santamaría, 640-680 m.,
Dodge & Thomas 8049.
Hypothallus nullus sed rhizinae coeruleo-nigricantes, dense
implexi, ei simulantes; thallus albidus, minute tomentosus,
palmatim lobatus, lobis oblongis, 0.5-1 mm. latitudine, 120 y
crassitudine, cortice filamentosa, 15-20 y crassitudine hyphis
pachydermaticis periclinalibus; strato nostocaceo 40 u crassitu-
dine in medulla gradatim transeunte, rhizinis nigris pachy-
dermaticis 4-5 y. diametro metientibus. Apothecia 0.5-0.75 mm.
diametro marginibus crassisimis crenulatis inflexis, disco nigro;
amphithecium inferne 140 y. crassitudine superne ad 100 y
attenuatum, cortice 40 y crassitudine pseudoparenchymatico;
parathecium 20-30 y. crassitudine, hyphis tenuibus periclinalibus
dense implexis; thecium 140-160 y altitudine; paraphyses tenues,
filiformes, apicibus in gelatina dilabentibus; asci clavati, 12 y. dia-
metro; ascosporae distichae, octonae, 10-11 x 7-8 y, ellipsoideae,
episporiis crassis.
Hypothallus of densely tangled bluish black rhizinae; thallus
white, minutely tomentose, palmately lobed, lobes oblong,
0.5-1.0 mm. broad, 120 y thick; cortex filamentous, 15-20 y
thick, of thick-walled hyphae, ind layer of Nostoc qradunlly
merging into the medulla; rhizinae thick-walled, black, 4-5 y in
diameter. Apothecia 0.5-0.75 mm. in diameter, with very thick
inrolled margins and blackening disc; amphithecium 140 u thick
below, thinning to 100 & at the margin, cortex 40 u thick, pseudo-
parenchymatous; parathecium 20-30 y thick, of densely ap-
pressed periclinal hyphae not clearly differentiated from the
hypothecium; thecium 140-160 y. tall; paraphyses very slender,
filiform, degenerating into the dark brown epithecial gel; asci
clavate, 12 u in diameter, 8-spored; ascospores distichous,
ellipsoidal, with thick epispore, 10-11 x 7-8 y.
While eventually this species may be found to have a much
wider distribution, it seems fitting that the excellent H. Santa-
maría, with one of the richest lichen floras known to me, should be
1933]
DODGE—LICHENS OF COSTA RICA. I 451
commemorated in botanical literature. The hacienda in turn
preserves the name of the national hero of Costa Rica not far
from the scene of his famous exploits.
Guanacaste: H. Santamaría, 640-680 m., Dodge & Thomas 8049.
MALMELLA physcioides (Vainio) Dodge, n. comb.
Erioderma physcioides Vainio, Jour. Bot. Brit. & For. 34: 70.
1896.
Type: West Indies, St. Vincent, Boxwood, W. R. Elliott 252.
Hypothallus sparingly developed, strigose-pannose, greenish
black; thallus deep olive buff, radiately subpinnatifid, ultimate
lobes 1-1.5 mm. broad, short and convex, surface toward the
margin slightly spongy, very minutely verrucose, about 120 y
thick, upper cortex about 40 y thick, of large thin-walled pseudo-
parenchyma, the rest of loosely woven hyphae with very small,
separate colonies of Nostoc, the lower 10 u with more densely
woven, more or less parallel hyphae. Apothecia circular to
quite irregular by mutual pressure, crowded, about 1 mm. in
diameter, margin concolorous with the thallus, crenulate, disc
chestnut; amphithecium about 200 u thick, the outer 60 y of
tomentum consisting of slender, thin-walled hyphae more or less
perpendicular to the surface and somewhat tangled. Within
this is the true cortex about 30 y thick of large-celled pseudo-
parenchyma, thinning out to a single layer above, and the algal
layer of slender periclinal thin-walled hyphae and colonies of
Nostoc about 100 y thick below and about 80 v above; parathecium
about 40 yu thick, of densely woven periclinal hyphae, staining
very lightly and reaching the margin above; hypothecium deeply
staining, 15-20 y. thick, of densely woven hyphae; thecium 100-
120 y. tall; paraphyses filiform, 2 » in diameter, frequently septate,
disintegrating into the brownish epithecial gel; asci clavate,
6-8 u in diameter, 8-spored; ascospores broadly ellipsoidal, 10-
12 x 6-8 y, with a gelified wall, minutely roughened within.
Previously reported only from the type locality. I have
previously distributed duplicate material of this species under
the name P. imbricatula Müll. Arg., which was originally briefly
described from southern Brasil. Only a careful study of both
types can settle its identity and relationships.
Alajuela: Fraijanes, 1500-1700 m., Standley & J. Valerio 47651; S. Isidro de
Alajuela, 980-1300 m., Dodge, J. Valerio & Thomas 4883.
Guanacaste: H. Santamaría, 640—780 m., Dodge & Thomas 8034, 8035.
[Vor. 20
452 ANNALS OF THE MISSOURI BOTANICAL GARDEN
MALMELLA caesiocinerea (Vainio) Dodge, n. comb.
Pannaria rubiginosa f. cinerascens Vainio, Jour. Bot. Brit. &
For. 34: 70 [10]. 1896, non Nyl.
Pannaria rubiginosa f. caesiocinerea Vainio, Acta Soc. Sci.
Fenn. A6': 102-103. 1915.
Type: not stated although several specimens mentioned.
P. rubiginosa f. cinerascens Vain. based on Antilles, St. Vincent,
Mt. St. Andrews, 320 m., Elliott 15. The following description
is based on Costa Rican material.
Rhizinae bluish black, highly developed and tangled; thallus
buffy brown in the center, lobes dark olive with a white pruina
about the margins, tomentose, lobes rounded, 1-4 mm. broad,
sinuate, incised, the lobules oblong, about 0.5 mm. broad, convex,
margins thick, inflexed below, 200 u thick, the upper cortex a
palisade of thick-walled hyphae about 20 u thick, homoeomerous,
with coiled filaments of Nostoc distributed throughout a very
loosely tangled layer of hyphae, fraying out below without
lower cortex. Apothecia up to about 0.5 mm. in diameter,
margin very thiek and inrolled, white, crenulate, disc black;
amphithecium thick, continuous with the thallus, about 140 y
thick, with cortex about 60 » thick below and with algae much
more abundant and closely tangled than in the thallus; para-
thecium thin, filamentous, 20 y. thick, not differentiated from the
hypothecium; thecium 80-90 y. tall; paraphyses filiform, septate,
ending in the hypothecial gel which is deep brown and deeply
staining; asci clavate, about 20 u in diameter, 8-spored; asco-
spores distichous, ellipsoidal, 11-12 x 6-7 y, with a gelified
epispore about 2 u thick, sometimes appearing 2-celled or even
4-celled with 2 transverse septa and 1 longitudinal septum in the
middle, probably due to oil globules or other highly refractive
substances, as other spores in the same ascus seem to be definitely
unicellular.
This species suggests a close relationship with Lepidocollema
but the cortex is much better developed and the apothecium is
lecanorine.
Guanacaste: H. Santamaría, 640—680 m., Dodge & Thomas 8038.
PARMELIELLA Müll. Arg.
PARMELIELLA Müll. Arg., Mem. Soc. Phys. Hist. Nat. Genéve
16:376. 1862.
1933]
DODGE—LICHENS OF COSTA RICA. I 453
Type species: P. triptophylla (Ach.) Müll. Arg. [P. corallinoides
(Hoffm.) Zahlbr.]
Thallus squamose, becoming elongate and subfoliose at the
margins with a well-developed dark hypothallus, algae Nostoc;
upper cortex of thick-walled pseudoparenchyma; medulla loosely
woven; lower cortex an appressed layer of rhizinae. Apothecia
superficial; amphithecium absent; parathecium pseudoparenchy-
matous; asci 8-spored; spores hyaline, unicellular, elongate-
ellipsoidal with thin walls; spermatia short, straight.
This genus seems world-wide in its distribution, perhaps
predominantly of the southern hemisphere but widespread
in the North Temperate zone and in the tropics, preferring the
higher elevations. Only P. pannosa has yet been found in Costa
Rica. The other species have been incompletely described but
have characters which should make them recognizable if found.
KEY TO THE TROPICAL AMERICAN SPECIES OF PARMELIELLA
Hypothallus indistinct, thallus glauco-cinereous or pale, squamules c
fluent, crenate, granulate; spores small, 8-9 x 4-6 u; Chile... . P. scored likes
Hypothallus forming a black margin about the thallus
Hypothallus very thin, blue green; spores 15-19(-21) x 6.5-8 u; surface
smooth; thecium 100 u thick; Chile................ 0 ee ee eeeee . nigrocincta
Hypothallus blue black, densely woven, cortex a single layer of cells;
spores 11-17 x 5-8 u; thecium 110 y thick; Mexico.......... P. miradorensis
Hypothallus brownish black, densely woven and thick, cortex several
layers of pseudoparenchyma; spores 12-15(-18) x 6-8 u; thecium
ij, | rr NEU IU vos ir on P. pannosa
Hypothallus very thick; apothecia 1-1.5 mm. in diam.; spores 16-25 x
8-10 »; BOWER... 05 one cc ee ee n ie eee P. nigrocincta var. Weddellit
PARMELIELLA PANNOSA (Sw.) Miill. Arg., Flora 64: 86. 1881.
Lichen pannosus Sw., Nova Gen. Sp. Pl. Prodr. 146. 1788;
Fl. Ind. Occ. 3: 1888. 1806.
Type: Jamaica, O. Swartz.
Hypothallus brownish black, extending 1-2 mm. beyond the
thallus; thallus tawny olive to isabelline, irregularly somewhat
pinnatifidly divided in the margin, the central portion of the
thallus becoming appressed microphylline and even somewhat
isidioid, margins of lobes somewhat lighter in color, otherwise
smooth above; thallus about 100 y. thick, the upper cortex 40 y
thick, of large thick-walled spherical cells forming a pseudo-
parenchyma, algal zone thin, about 20 y thick, of small somewhat
[Vor. 20
454 ANNALS OF THE MISSOURI BOTANICAL GARDEN
confluent colonies, medulla about 40 y thick, of large thin-walled,
loosely woven hyphae, the lower cortex 10-12 u thick, of con-
glutinate, black, thick-walled hyphae. Apothecia abundant,
about 1 mm. in diameter, round to irregular in shape, the margin
lighter colored when young, appearing to be thalline but not so,
darkening in age but always slightly lighter than the disc and
not exceeding the epithecium in height, disc rufous; amphi-
thecium absent; parathecium 100-120 » thick, pseudoparenchy-
matous, of large thick-walled cells extending to the top of the
thecium; hypothecium about 40 y thick, of densely woven
hyphae, deeply staining; thecium 120-140 y thick; paraphyses
branched above, 1.5-2 y. in diameter, with thin walls; asci clavate,
about 15 y in diameter, 8-spored ; ascospores irregularly distichous,
ellipsoidal, 12-15 x 6-8 y, epispore thick.
This species seems to be the commonest one in tropical America,
in our area confined largely to higher elevations than Pannaria,
being most abundant above 650 m., but rarely found at lower
elevations.
Without locality: Ørsted 8.
Limón: Waldeck, Dodge 7403.
Cartago: R. Pejivalle, 600—750 m., Dodge & Thomas 4341, 8038; Cartago, 1200—
1500 m., C. Wercklé (Mus. Nac. 17218).
San dedi Sta. María de Dota, 1500-1800 m., Standley & J. Valerio 43394, 44057,
48444, 434406, 43472; La Hondura, 1300-1700 m., Standley 36458b.
Heredia: Barba, El Gallito, R. Torres 150; C. Central de Zurquí, 1600-1700 m.,
Dodge, J. Valerio & Thomas 8039.
Alajuela: 8. Ramón, La Palma, 1100 m., Brenes 64; Piedades, 900 m., Brenes 380;
Alto Calera, 750-800 m., Brenes 265.
Guanacaste: H. lantaa 680--780 m., Dodge & Thomas 6892; H. Granadilla,
500-600 m., Dodge & Thomas 6746.
Var. coralloidea Dodge, var. nov.
Type: Costa Rica, Heredia, C. Zurquí, 1600-1700 m., Dodge,
J. Valerio & Thomas 8061.
Thallus iteratim laciniatus sed non microphyllinus, isidiis
coralloideis marginalibus rare sparsis, obscurior, 140 y. crassitu-
dine, cortex superior 25-30 y crassitudine, duobus seriebus
cellularum, serie superiore fusca, inferiore hyalina; stratum
nostocaceum ad 60 u crassitudine, filamentis dense contextum;
medulla hyphis magnis laxe implexa; cortex inferior ad 12 y
crassitudine, hyphis nigris, pachydermaticis, conglutinatis, com-
pactus; sterilis.
1933]
DODGE—LICHENS OF COSTA RICA. I 455
Thallus repeatedly laciniate but not microphylline, isidia
coralloid, mostly marginal, rarely scattered over the surface,
darker, 140 y thick, the upper cortex 25-30 y thick, of two layers
of thick-walled cells, the outer layer fuscous, the inner hyaline;
the algal layer about 60 x thick, of densely woven filaments of
Nostoc; medulla of large loosely tangled hyphae; lower cortex
about 12 y thick of large black, thick-walled hyphae; sterile.
Heredia: C. Central de Zurquí, 1600-1700 m., Dodge, J. Valerio & Thomas 8061.
ERIODERMA Fée
ERIODERMA Fée, Essai Cryptog. Ecorc. Offic. 146. 1824.
Type species: Érioderma polycarpum Fée.
Thallus foliose, attached to the substrate by either marginal
rhizinae or those from the under surface; tomentum highly
developed, cortex pseudoparenchymatous, formed by a palisade
layer, the algal zone of short chains of Scytonema with thin
sheaths or Nostoc; medulla loosely woven of thin-walled hyphae;
no lower cortex, the lower surface somewhat veined and in some
species with a dense covering of rhizinae. Apothecia marginal
or superficial, peltate, constricted at the base; amphithecium
absent; parathecium well developed, of large thick-walled cells,
pseudoparenchymatous, with a loosely woven medulla; asci
8-spored; spores hyaline, unicellular, ellipsoidal, occasionally
fusiform or subspherical. Spermagonia marginal, small, black-
ish warts; spermatiophores septate; spermatia short, cylindric,
straight.
KEY TO THE TROPICAL AMERICAN SPECIES OF ERIODERMA
Dense blackish fuscous tomentum below, tier hypothallus.
Lobes narrow, deeply sinuate divided margins.................... E. Wrightii
Lobes broader, margin with irai caesious soredia below, and
white-villos6.......... rest re a a aa EOS . limbatum
Nearly nude and veined, white below, although rhizinae bluish black or
white, verruculose and villose above; Brasil.................. verruculosum
SUME m below with white veins and eee coarsely éhorlipiiien
e; apothecia up to 0.8 mm. in diam.; Jamaica.............. E. microcarpa
Bluish white with fleshy white veins and incidi, m tufted d giving
an net appearance when dry; apothecia 3-4 mm iam.;
E E a E E R iD . pulchrum
White Au thallus isidiose; apothecial margin isidiose, ashy virescent
UL [go oo oo 8 8 ba nce 4 oe 0g 24 eee ss > cw E. Leylandi
EnropERMA Wricuti Tuck., Amer. Jour. Arts & Sci. II, 25:
423. 1858
[Vor 20
456 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Type: Cuba, top of Loma del Gato, C. Wright.
Hypothallus none, but dense brownish-black rhizinae clothe
the under surface to within 3 mm. of the margin which is arach-
noid-tomentose, white; thallus deep olive buff to citrine drab,
sinuate-lobed, upper surface tomentose, margins with tufts of
dark brown rhizinae, thallus about 120 y thick exclusive of
tomentum on upper surface and rhizinae below; cortex pseudo-
parenchymatous, 30—40 y thick, of large cells, algal layer very
thin, 10-15 u thick, of densely coiled filaments of Scytonema;
medulla very loosely woven, similar to the tomentum above; no
lower cortex. Apothecia marginal with white densely pilose
exciple and black disc; amphithecium absent; parathecium
200 y. thick, corticate with pseudoparenchyma about 25 y thick
below, a little less above, formed from a palisade of hyphae
perpendicular to the surface, the cortex covered with tomentum as
in the thallus, the rest of the parathecium of large, loosely woven
hyphae similar to the medulla but somewhat more closely woven;
hypothecium not well differentiated from the parathecium,
rather more densely woven; thecium about 120 y tall; paraphyses
filiform, slender, branched above, ending in dense brown epithecial
gel; asci clavate, 8-spored, 14-16 y in diameter; ascospores
distichous, 12-14 x 6-7 y, ellipsoidal, with a thick epispore.
This very distinct species resembles the Peltigeraceae in habit
much more than the Pannariaceae, but it has the typical uni-
cellular spore with the thick epispore of the latter family. It
seems rather rare in tropical America but it is quite abundant
in the few localities where it has been found. In Costa Rica it
occurs from 680 to 1100 m.
Alajuela: La Palma de S. Ramón, 1100 m., Brenes 50.
Guanacaste: H. Santamaría, 680-780 m., Dodge & Thomas 6920, 6982.
ERIODERMA limbatum (Nyl.) Dodge, n. comb.
Erioderma Wrightw var. limbatum Nyl., Flora 52: 119. 1869.
Type: Brasil, Rio de Janeiro, Serra dos Orgãos, Glaziou 2004.
Thallus cinnamon buff to chamois, lobes rounded-sinuate,
slightly excised, 5 mm. broad, margins crenate, thick, upper
surface only slightly tomentose, lower surface with margins
caesious-sorediate, white; rhizinae greenish black, very dense
and thick; thallus about 300 » thick, upper tomentum not highly
1933]
DODGE—LICHENS OF COSTA RICA. I 457
developed; cortex 25-30 y thick, pseudoparenchymatous from
the disintegration of a palisade; algal layer about 60 u thick,
of irregularly tangled filaments of irregular somewhat angular
cells about 8 u in diameter (Scytonema ?); medulla about 60 y.
thick next the cortex, of rather densely tangled hyphae, gradually
becoming looser to form another indefinite layer 120 y. thick, of
very large and very loosely tangled hyphae 10-12 y in diameter,
with black hyphae of about the same size forming a more dense
mat which serves as a lower cortex. Apothecia 3 mm. in diam-
eter, stipitate, stipe 1-2 mm. tall, margin thin, erect or incurved,
densely tomentose without, the hairs tangled, giving a granular
appearance, dise chestnut, soon blackening; no amphithecium;
parathecium highly developed, cortex a palisade of thick-walled
infrequently septate hyphae proliferating to form the tomentum
and medulla, 200 u or more thick below, thinning out at the edge
of the thecium so that only the cortex about 100 u thick surrounds
the upper portion of the thecium at the margin; hypothecium
not differentiated; thecium about 90 y tall; paraphyses slender,
filiform, apex not enlarged; asci cylindrical, about 8 y in diameter,
8-spored; ascospores monostichous, at least when found, thick-
walled, hyaline, ellipsoidal, 10-12 x 5-6 p.
This species, from its habit and color, might easily be mistaken
for Sticta Weigelii or S. rufa except for a somewhat duller surface,
but is distinguishable on the color and density of the rhizinae and
the lack of cyphellae. Known from Costa Rica from a single
collection with scant data by Brenes . . . a, probably from the
region of S. Ramón in Alajuela Province.
COCCOCARPIA Pers. ap. Gaudich.
CoccocarPIA Pers. ap. Gaudich. in Freycinet, Voy. Uranie,
Bot. 206. 1826.
'Type species: Coccocarpia molybdaea Pers. ap. Gaudich.
Thallus squamose to foliose, with dark or light-colored rhizinae,
corticate on both surfaces, upper surface pseudoparenchymatous
from thin-walled large hyphae parallel to the surface, without
tomentum; algal layer of coiled chains of Scytonema in thin
sheaths; medulla of thin-walled somewhat conglutinate hyphae
not sharply differentiated from the lower cortex, of more or less
[Vor. 20
458 ANNALS OF THE MISSOURI BOTANICAL GARDEN
septate hyphae running parallel to the surface. Apothecia
superficial, sessile or somewhat constricted below; amphithecium
absent; parathecium corticate with large-celled pseudoparen-
chyma, within, of large septate periclinal hyphae, medulla
lacking; hypothecium either light or dark; asci 8-spored; asco-
spores hyaline, unicellular, spherical to ellipsoidal-fusiform, thin-
walled. Spermagonia in warts on the thallus, spermatiophores
frequently septate; spermatia straight, elongate-cylindric.
This widespread tropical genus has two very widespread species
or species groups and a number of rare and seemingly localized
species scattered over the tropical regions.
KEY TO THE TROPICAL AMERICAN SPECIES OF COCCOCARPIA
Under side reticulate, black-fibrillose; lobes obtuse, ciliate, margin of apo-
thecia crenulate, disc yellow..................ecececeeeees C. portoricensis
Under side densely black-tomentose, forming a hypothallus.
Lobes bipinnately divided, linear, ciliate, aeruginous-green........... C. subtilis
Lobes cuneate to flabelliform, bluish to lead color.
Thallus isidiose or microphylline in the center...................... C. cronia
Thallus not reddish ferruginous within.
Isidia concolorous, laciniae cuneate, 1.5-10 mm. broad.
Laciniae 5-10 mm. broad, isidia applanate to microphylline. . . .
TRCN SSM P SE ERKRRAJAKEC-E CREARE RE RR eons eh aan eh Ee Eas v. prolificans
Laciniae 1.5-5 mm. broad, isidia terete; apothecia eni or
rufo-fuscous, exciple white-ciliate................... v. isidiophylla
Isidia usually darker at the tips and blackening, laciniae narrower
and sublinear.
Laciniae 1-1.5 mm. broad; hypothallus little developed; apo-
thecia fuscous, blackening, exciple white-ciliate; Brasil. .v. camporum
— 0.5-3.0 mm. broad, narrow granulose to lobulate in the
er; apothecia black; hypothallus well developed. . . . v. granulosa
Thallus hee ferruginous within; Cuba................ v. erythrocardia
Thallus not isidiose in center... 2.2.0.0. .6. 00 ccc cece cee eee eee C. pellita
Laciniae cuneate or broad-spathulate
Hy ypot eg bie highly develaped; apothecia yellow to tawny
POE PTT COPE ROLXXXAXERS AERIS v. pannosa
es hs but much less well developed.
Apothecia black from the first, — ere v. parmelioides
Apothecia yellow, rufous, or fuscous.
Thallus = thin, lobes anal, approaching microphylline
C. cronia w. prolificans; apothecia finally
TOT, «desees PENES S an ac ef en v. lividorufa
Tui tiie lobes broad, 4-8 mm.; apothecia not dark-
NE MOL. ee ree: v. pyrrhichocarpa
Laciniae ststinene, up to 4 mm. broad, short, rounded.
Apothecia black; Jamaica.................. ccc cece nnn v. genuina
Apothecia fuscous, ciliate...........0000 0. cece eee ee eee v. strigdso
1933]
DODGE—LICHENS OF COSTA RICA. I 459
Under side of thallus white with black-fibrillose margins, laciniae dick
e. ri dM TT""—"-— C. fibrillosa
Under ade of thallus white, rhizinae white.
Thallus isidiose.
Laciniae very narrow, 0.1—0.2 mm. broad, adnate............. C. dominguensis
Laciniae broad and flabelliform, up to 10 mm. broad................ C. albida
Thallus not isidiose.
Laciniae 150—200 u broad, loosely adnate, ja rhizinae penicilloid,
projecting beyond margin, dichotomous.................+ C. tenuissima
Laciniae 250 u broad, tips ascending, "unt long white rhizinae,
palmate with lacinulae pinnatifid; apothecia carneous, rigidly
hi os54 00. P E E T E a ev . elegans
Laciniae broader.
Thallus greenish blue, rhizinae rigid, white; flattened apothecia
, MMC 55 hia ade 4a eR NAE Pen epiphylla
Thallus lead-color or slightly bluish, onea 28
bricate, pterygoid; apothecia testaceous, white, ciliate...... "C. asterella
CoccocaRPIA PELLITA (Ach.) Müll. Arg., Flora 65: 320. 1882.
Parmelia pellita Ach., Lichenog. Univ. 468. 1810, Sw., Lich.
Amer. 7, pl. 6. 1811.
Type: West Indies, O. Swartz.
Laciniae of thallus rather narrow, about 4 mm. broad, pin-
natifidly incised, either discrete or approximate, smooth above;
hypothallus well developed and extending beyond the thallus.
Apothecia deep fuscous.
This species seems widely distributed and very variable in the
tropics. Only a monographic study by one who has access to the
types of all the proposed varieties as well as considerable experi-
ence in the field in the principal floristic areas of the tropics can
finally decide the validity of the proposed varieties and forms
and provide adequate descriptions so that others may recognize
them. Nylander, Miller Argau, Hue, Vainio, and Malme have
each attempted more or less elaborate revisions but none seems
wholly satisfactory. While probably artificial the separation of
C. cronia for all the isidiose varieties of this species is relatively
easy to apply, and has been followed here.
Var. PARMELIOIDES (Hook. ap. Kunth) Müll. Arg., Flora 65:
320. 1882.
Lecidea parmelioides Hook. ap. Kunth, Syn. Pl. Aequinoct.
Orb. Nov. 1: 15.
Type: Colombia, Cumana, Bordones, and Nueva Barcelona,
Humboldt & Bonpland.
[Vor. 20
460 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Thallus light olive gray, surface smooth, lobes cuneate, about
mm. wide, sometimes once-cleft, closely appressed to the sub-
strate, held by a few dark brown to black rhizinae composed of
fascicles of hyphae, about 200 u thick; upper cortex about 25 u
thick, the outer 5 y formed from the disintegration of a very thin
layer of periclinal hyphae, the rest pseudoparenchymatous from
the shifting of cells in a palisade layer, cells 10—12 y in diameter,
rather thick-walled; algal layer a palisade of hyphae and filaments
of Scytonema about 100-120 u thick; medulla about 40 y thick,
of large septate periclinal hyphae 5-8 y in diameter; lower cortex
of dark, periclinal conglutinate thick-walled hyphae 3-4 y in
diameter. Apothecia immarginate, appressed and more or less
adnate, 2 mm. or more in diameter, disc black; no amphithecium;
parathecium of periclinal thick-walled hyphae, giving a pseudo-
parenchymatous appearance, 100-120 y thick, thinning out to
about 20 u at the edge; hypothecium not differentiated from the
parathecium; thecium about 160 u tall; paraphyses 2-3 y in diam-
eter, frequently septate, clavate tips black, about 4 u in diameter;
asci cylindric to somewhat clavate, 8-spored, about 60 x 8-12 u;
ascospores monostichous at first, gradually becoming distichous,
ellipsoidal with acute ends, very thick-walled, unicellular, often
with some deeply staining material in the middle with a slight
thickening of the cell wall at that point, giving the appearance of
a two-celled polarilocular spore with a wide isthmus, occasionally
two other constrictions near the tip slightly suggesting a 4-celled
condition, 11-12(-16) x 6-7 y.
'This variety seems widespread in the lower elevations although
in absence of apothecia it is practically indistinguishable from
var. pyrrhichocarpa. In Costa Rica var. parmelioides occurs in
suitable habitats up to about 800 m., while var. pyrrhichocarpa
occurs from 700 to 1340 m., but seen from Brasil up to 2000 m.
in the state of São Paulo.
Limón: Waldeck, 40 m., Dodge 7402.
Guanacaste: H. Santamaría, 680-780 m., Dodge & Thomas 4709; C. San José de
Libano, 500-960 m., Dodge, Hanckel & Thomas 6684; R. San José, 460—480 m.,
Dodge & Thomas 6581.
Puntarenas: Osa, Puerto Jiménez, Brenes 825a; R. Sándalo, Dodge 7742.
Var. PYRRHICHOCARPA Hue, Bull. Soc. Bot. France 48: lx.
1901 [1902].
1933]
DODGE—LICHENS OF COSTA RICA. I 461
Var. smaragdina Müll. Arg., Flora 65: 320. 1882, excl. syn.
quoad specimina americana; Vainio, Etude Lich. Brésil 1: 210.
1890; Malme, Ark. f. Bot. 20°: 19. 1924.
Type: Brasil, São Paulo, near São Paulo, Azevedo Sampaio.
Thallus between yellowish glaucous and light mineral gray,
lobing of thallus similar to var. parmelioides but much more
irregular and lobes rather smaller, smooth above, 150-160 y
thick, upper portion of cortex 4—6 y thick, of longitudinal hyphae
soon disintegrating and disappearing, leaving a palisade 25-30 y.
thick, which in turn becomes a somewhat irregular large-celled
pseudoparenchyma; the algal layer 40-50 u thick, of loosely
coiled and disintegrating filaments, perhaps Scytonema, but the
cells rounding up and suggesting Nostoc, about 8 y in diameter,
the medulla 50 y thick, of large septate periclinal hyphae, some-
what less compact than in var. parmelioides, the lower cortex
about 25 y thick, of black somewhat smaller conglutinate hyphae;
rhizinae much more abundant, smaller conglutinate fascicles of
hyphae similar to those of the lower cortex. Apothecia irregular,
3-3.5 mm. in diameter, carnelian red to vinaceous rufous, only
slightly darkening, never dark brown or black, immarginate;
amphithecium absent; parathecium adnate to the upper cortex,
about 100 » thick, of large periclinal hyphae appearing almost
pseudoparenchymatous at times; thecium 60 y. tall; paraphyses
2-3 y in diameter, filiform, apex not enlarged; asci clavate, about
12 y in diameter; ascospores 10-14 x 4-5 y.
This variety is occasional at the medium elevations of the
temperate zone in Costa Rica, ranging from 700 to 1400 m. It
should not be confused with C. smaragdina Pers. or C. molybdaea
Pers., both from the Old World tropics.
Cartago: R. Birrís near Santiago, 920-1340 m., Dodge 4708, 8050; Cartago, C.
Wercklé (Mus. Nac. 172186 p. min. p.).
Alajuela: Santiago de S. Ramón, 1000 m., Brenes 238.
Guanacaste: H. Santamaría, 680—780 m., Dodge & Thomas 6813.
Var. srRIGOSA Müll. Arg., Flora 65: 326. 1882.
Coccocarpia molybdaea v. cronia Nyl., Acta Soc. Sci. Fenn.
7:441. 1863, non Tuck.
Type: Colombia, Cune, 1200 m., Lindig 2663 p. p.
Thallus between pale olive buff and yellowish glaucous, lobes
cuneate but narrower than in v. parmelioides, slightly incised,
[Vor. 20
462 ANNALS OF THE MISSOURI BOTANICAL GARDEN
smooth and shining above, thallus about 100 y. thick, outermost
layer of hyphae about 4 y thick, soon disintegrating, upper
cortex pseudoparenchymatous, 10-12 u thick, of about two layers
of pseudoparenchymatous cells not part of a palisade, algal layer
32-40 y thick, of curved hyphae in a disintegrating palisade
(intermediate between v. parmelioides and v. pyrrhichocarpa),
medulla 25-30 y thick, of large periclinal hyphae, lower cortex
12-15 y thick, of large black periclinal hyphae, closely agglu-
tinated rhizinae abundant but hyphae less fasciculate. Apo-
thecia about 2 mm. broad, ochraceous tawny at first, becoming
Prout’s brown or darker, immarginate; no amphithecium;
parathecium about 100 y thick, of large periclinal hyphae, at the
margins many hyphae projecting beyond the general level as
stiff hairs; hypothecium not differentiated and not deeply
staining; thecium about 40 » tall; paraphyses filiform with acu-
minate tips which extend 1-2 y above the epithecial gel; asci
clavate, 30x 8 y, with 8 spores; ascospores fusiform-ellipsoidal,
8 x 4 y, thick-walled, probably still immature.
In the only collection from Costa Rica the thallus may not be
quite mature, as no mature ascospores were found, hence the
measurements may be a little small. Reported elsewhere only
from Colombia.
Cartago: R. Birrís, 920-1100 m., Dodge 8051.
COCCOCARPIA CRONIA (Tuck.) Vainio, Ann. Acad. Sci. Fenn.
A6': 103. 1915.
Parmelia cronia 'Tuck., Proc. Amer. Acad. Arts & Sci. 1: 228.
1848; Syn. Lich. New England, 36. 1848.
Coccocarpia molybdaea v. cronia Nyl., Syn. Meth. Lich. 2: 41.
1863.
Coccocarpia pellita v. cronia Müll. Arg., Flora 65: 321. 1882.
Coccocarpia parmelioides v. cronia Hue, Bull. Soc. Bot. France
48: lx. 1901 [1902].
Type: United States, Massachusetts, Lynn Hills, and West
Cambridge, on mossy rocks, T'uckerman.
Thallus mineral gray or darker, lobes rounded or short, broad-
linear, more or less appressed, growing over mosses, about 130-
140 y thick; upper cortex 15-20 » thick of large thin-walled
hyphae often appearing pseudoparenchymatous; algal zone
1933] i
DODGE— LICHENS OF COSTA RICA. I 463
30-40 y. thick, with loosely woven hyphae in which are imbedded
solid little groups 15-20 y in diameter of algal cells of closely
coiled Scytonema; medulla 30-40 y thick, partly of close peri-
clinal hyphae, partly more loosely woven, the lower cortex of
two layers of black thick-walled hyphae giving rise to rhizinae,
sterile; isidia terete, concolorous, or somewhat darkened at the
tips. ‘Tuckerman noted on a specimen from Alabama, ‘‘spores
9-12 x 3-5 y, 2-celled?"
Var. LIVIDORUFA (Mey. & Fw.) Zahlbr., Cat. Lich. Univ. 3:
287. 19265.
Parmelia lividorufa Mey. & Fw., Nova Acta Acad. Leopold.
Carolin. 19: Suppl. 222, pl. 4, f. 2. 1843.
Coccocarpia molybdaea var. tenuior Nyl., ap. Krmplhbr., Flora
59:76. 1876 (nom. nud.).
Coccocarpia pellita var. tenuior Müll. Arg., Flora 65: 321. 1882.
'Type: Brasil, Rio de Janeiro, Meyen (Bot. Mus. Berlin), type
of var. tenuior; same locality, Glaziou 2026 (Univ. Genéve).
'Thallus mineral gray, lobes small, mostly about 1 mm. broad,
quite irregular, smooth, about 100 y thick, structure as in the
previous variety. Apothecia immarginate, irregular, convex,
dise snuff brown to bister; amphithecium absent; parathecium
130—140 y thick; hypothesi not differentiated; theifiutu 80-85 u
tall; paraphyses filiform, ending in the buitisecin] gel; asci 8-
spored, clavate to cylindrical, 12 » in diameter, ascospores 11-12 x
5-6 y, ellipsoidal, thick-walled, distichous.
In this variety the whole thallus approaches a microphylline
squamose state. Evidently a lowland variety seen also from
British Honduras and Nicaragua.
Limón: R. see Sebi 70-200 m., Dodge, Catt & Thomas 8058.
Alajuela: La Palma de S. Ramén, 1100 m., Brenes 43.
Guanacaste: Tilarán, 650—690 m., Dodge & Thota 6557.
Var. isidiophylla (Müll. Arg.) Dodge, comb. nov.
Coccocarpia pellita v. wsidiophylla Müll. Arg., Flora 65: 321.
1882.
Coccocarpia pellita v. cronia Vainio, Étude Lich. Brésil 1: 209.
1890, et auct. recentior., non Tuck.
Coccocarpia cronia v. primaria Vainio, Ann. Acad. Sci. Fenn.
A6':103. 1915.
Type: Brasil, Rio de Janeiro, Glaziou 2025.
[Vor. 20
464 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Thallus mineral gray or darker, laciniate, flabelliform or
cuneate with rounded margin about 4 mm. broad, sometimes
somewhat incised along the margin, upper surface isidiose with
either simple or slightly branched isidia which are confined to
the central portion of the thallus and to a lesser extent of the
lobes; upper cortex of large thin-walled hyphae perpendicular to
the margins of the lobes, up to about 20 u thick, closely septate;
algal zone 60 y. thick of filaments of Scytonema about 12 y. thick
with cells about 8 u in diameter, in the lower portion the algae
less crowded, leaving some air spaces, medulla not differentiated
from the algal zone; lower cortex 30 y thick, of conglutinate,
black, thick-walled hyphae, giving rise to scattered rhizinae as
well as penicillate fascicles of hyphae. Apothecia yellowish-
rufous, becoming fuscous, somewhat lobulate, immarginate,
adnate; parathecium 55-60 y in the center, thinning out at the
margin, pseudoparenchymatous, of large somewhat periclinal
hyphae; hypothecium not differentiated, about 60 y thick;
paraphyses 2-3 y in diameter, filiform, with swollen tips in the
very dark epithecial gel; asci clavate, 8-spored, 8-10 y in diam-
eter; ascospores distichous, fusiform, immature.
This species is close to C. pellita and perhaps should be con-
sidered a variety of that species. It has approximately the same
distribution, reaching from sea level to about 1700 m. in Costa
Rica, 2700 m. in Pert.
Cartago: Aguacaliente, 1240-1460 m., Dodge & Thomas 7087.
Heredia: C. Central de Zurquí, 1600-1700 m., Dodge, J. Valerio & Thomas 6087;
C. de las Caricias, 2000-2400 m., Standley & J. Valerio 55282.
Alajuela: La Palma de S. Ramón, 1100 m., Brenes (fertile).
Puntarenas: Osa, between R. Sándalo and R. Tigre, 1-2 m., Dodge 8057.
Var. prolificans (Malme) Dodge, n. comb.
Coccocarpia pellita var. prolificans Malme, Ark. f. Bot. 20°: 19.
1924.
Type: Brasil, Rio de Janeiro, Regnell 65.
Thallus mineral gray or darker, lobes flabelliform, somewhat
zonate, 5-10 mm. wide, margin rounded, more or less lobulate
along the incised portions between lobes, upper surface lobulate
and more or less microphylline, lobules concolorous, about 70 u
thick, upper cortex about 16 » thick, of large thin-walled longi-
tudinal hyphae; algal zone about 30 y thick, of a loose palisade
1933]
DODGE—LICHENS OF COSTA RICA. I 465
of Scytonema filaments, medulla about 20 u thick, of large thin-
walled periclinal hyphae with some air spaces; lower cortex
about 15 y thick, of thick-walled black hyphae, giving off fascicles
of rhizinal hyphae. Sterile.
This variety is rather common at elevations from 700 to 1400
m. in Costa Rica.
Cartago: R. Birrís, 1220-1340 m., Dodge & Thomas 4710.
Alajuela: La Palma de S. Ramón, 1250 m., Brenes 38a; Piedades de S. Ramón,
900 m., Brenes 385.
Guanacaste: H. Santamaría, 680-780 m., Dodge & Thomas 6987, 6999.
Var. granulosa (Müll. Arg.) Dodge, comb. nov.
Coccocarpia pellita var. granulosa Müll. Arg., Flora 65: 322.
1882.
Type: Brasil, São Paulo, Apiahy, Puiggari 244, and Colombia,
Bogotá, 2700 m., Lindig 2538.
Thallus thicker, greenish-glaucous-blue or darker, lobes up to
3 mm. broad, the larger lobes more or less incised, obovoid to
almost linear, soon blackish granular to almost isidiose in the
center, about 140 y thick, upper cortex a single layer of longi-
tudinal hyphae 4-5 y in diameter, algal zone of Scytonema
filaments 40-50 u thick, forming a palisade interspersed with
occasional rows of fungal cells, medulla of large, septate, peri-
clinal hyphae about 60 y thick, the lower cortex 6-8 y thick,
of thick-walled dark hyphae, not well developed. Apothecia
less adnate and somewhat isidiose; no amphithecium; para-
thecium 150 y thick, pseudoparenchymatous, some of the outer
layer of cells growing downward to form a ciliate to tomentose
surface below; hypothecium not differentiated, thecium about
60 y. tall; paraphyses filiform, tips not enlarged, ending in a dark
brown epithecial gel; asci clavate, rather immature.
This variety as here defined includes some of Malme's var.
isidiosa and some material may have been distributed under that
name. In Costa Rica this variety is widespread in the temperate
regions from 100 m. to 1500 m.
Limón: R. Siquirres, 70-200 m., Dodge, Catt & Thomas 559.
Cartago: Santiago, 1140-1180 m., Dodge 4638, 8054; C. istud 1320-1500 m.,
Dodge & Thomas 4763; Cartago, C. Wercklé (Mus. Nac. 17202).
San José: Turrdcares, 540-600 m., Dodge & Thomas 8056; R. Virilla below El
Brazil, Dodge 7777.
Alajuela: C. Pata de Gallo a S. Rafael de S. Ramón, 1200-1250 m., Brenes 215;
S. Pedro de S. Ramón, 700 m., Brenes 459.
[Vor. 20
466 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Guanacaste: Liberia, 100 m., Dodge & Thomas 6582; H. Santamaría, 640—780 m.,
Dodge & Thomas 6900, 7007, 8055; Tilarán, 650-690 m., Dodge & Thomas 6556, 6559,
Standley & J. Valerio 44429, 44520; H. Granadilla, between R. Las Cañas and R. S.
José, Dodge & Thomas 6719.
CoccocaAnPrA albida Dodge, sp. nov.
Type: Costa Rica, Limón, Hamburg, Dodge & Nevermann 7401.
Thallus albidus, isidiosus, lobis rotundatis, flabelliformibus,
10 mm. latis, zonatus, inferne albus rhizinis albidis, apicibus
rhizinarum nigricantibus, thallus 120-130 y crassitudine, cortex
15 u, hyphis longitudinalibus septatis, 5-6 uœ diametro; zona
scytonematica ad 40 y crassitudine, filamentis laxis implexis,
medulla 40-60 y crassitudine hyphis periclinalibus compactis;
cortex inferior hyphis hyalinis dense compactis; rhizinis albidis
fasciculatis penicillatis.
Thallus mineral gray or lighter, densely isidiose in the center,
lobes rounded, flabelliform as in C. pellita v. parmelioides, 10 mm.
broad, more or less zonate, below white with white rhizinae, the
latter sometimes tipped brownish black; thallus 120-130 y thick,
cortex 15 u thick, of longitudinal septate hyphae 5-6 y in di-
ameter; algal zone 40 y thick, of loosely tangled filaments of
Scyionema, medulla 40-60 u thick, of compact periclinal hyphae;
the lower cortex of hyaline hyphae more or less agglutinated;
rhizinae white, fasciculate, and penicillate.
The lower surface resembles that of Sticta sp. quite closely,
but cyphellae are absent and the structure is typical of Cocco-
carpia. Perhaps it is an albino variety of C. cronia var. isidio-
phylla. It has much broader lobes than C. dominguensis.
Limén: Hamburg, 20-30 m., Dodge & Nevermann 7401.
CoccocaRPIA ELEGANS Müll. Arg., Flora 64: 507. 1881.
Type: Brasil, Sáo Paulo, Apiahy, Puiggari.
No hypothallus, thallus terre-verte, small, linear, dichotomous
(rarely trichotomous at the ultimate branches), larger laciniae
0.1-0.2 mm. broad, ultimate branches much narrower, dull and
smooth above, white with white rhizinae below, thallus 80-85 y
thick, upper cortex 8-12 y thick, decomposed, apparently of
slender longitudinal hyphae; algal layer about 40 y thick, ir-
regular, of loosely woven fungus hyphae supporting short irregular
chains of angular to subspherical cells 8-12 y in diameter, medulla
not well differentiated from the algal layer but of very loosely
1933]
DODGE—LICHENS OF COSTA RICA. I 467
woven slender hyphae; lower cortex 10-15 u thick of closely
woven hyaline, longitudinal hyphae, the rhizinae 40-50 y in
diameter, large fascicles of hyaline hyphae. Apothecia immar-
ginate, 0.1-0.5 mm. in diameter, chestnut, with long fascicles of
hyphae forming stiff bristles about the margin, with stipe about
200 u in diameter and 40-50 y tall; no amphithecium; para-
thecium 60 y thick, of thick-walled hyphae forming pseudo-
parenchyma; thecium 60 y tall; paraphyses slender, filiform,
not enlarged at the tips; asci clavate, 4—6 u in diameter, immature.
The above description is based upon material from Brasil, Rio
de Janeiro, Glaziou 18071, determined by Müller Argau, who also
reports it from Puntarenas, Boruca, 560 m., Tonduz 5462. As
this is a small epiphyllous species, a large thallus being only 1 cm.
in diameter, it is quite possible that it has been overlooked as my
epiphyllous material has not yet been examined carefully.
A NEUTRAL (?) STRAIN OF MUCOR
SPHAEROSPORUS FROM MISSOURI
MORRIS MOORE
Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany
of
Washington University
In the caves of the Ranken estate in Missouri, where the
atmosphere is moist and the temperature approximately
16° C., was found a segment of a twig matted with a growth of
mycelial threads, white in a mass, and hyaline when single. The
appearance of the latter was that of many fine root hairs. When
transferred to agar substrates, numerous sporangia, sporangioles,
oidia, and chlamydospores developed, with no evidence of zygo-
spores. The characteristics and description were similar to those
given by Lendner (’08) for Mucor sphaerosporus Hagem 1908.
This fungus has been reported chiefly from Europe, and in
this country by Waksman (’16) and Gilman and Abbott (’27),
as isolations from soil. As far as the author could determine,
this is the first report of the above organism from Missouri.
A culture sent to Dr. A. F. Blakeslee at the Carnegie Institu-
tion of Washington, Department of Genetics, was tested with
strains of presumably the same species, one being a minus
isolated by Waksman in 1915, and another a plus strain probably
from the Centralstelle, of unknown origin. No zygospores were
produced. An attempt was made to obtain imperfect hybridiza-
tion by treating this strain with some of the strong testers in the
above laboratory, but no sexual reaction resulted. Growth on
various media produced changes in the amount of mycelium,
size and number of sporangia, sporangioles, hyphae, oidia and
chlamydospores, and in chromogenesis.
Thanks are due Dr. Blakeslee and Miss Satina of the Carnegie
Institution of Washington for their interest and assistance.
BIBLIOGRAPHY
Gilman, J. C., and E. V. Abbott eile A summary of the soil fungi. Iowa State
Coll. Jour. Sci. 1: 225-343.
— A. ('08). Les Mucorinées fi la Suisse. Matériaux pour la flore crypto-
mique Suisse 3!: 1-177. pl. 1-3. 8.
Walsman. S. A. (16). Soil fungi and their activities. Soil Sci. 2: 103-156. 1916.
Ann. Mo. Bor. Garp., Vor. 20, 1933 (469)
A STUDY OF ENDOMYCES CAPSULATUS REWBRIDGE,
DODGE AND AYERS: A CAUSATIVE AGENT OF
FATAL CEREBROSPINAL MENINGITIS!
MORRIS MOORE
Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany
of Washington University
INTRODUCTION
The probable taxonomic relationships of the fungi known as
yeast-like organisms have for many years attracted much atten-
tion and discussion among mycologists, and to a certain extent
among medical men. Considerations of their general physio-
logical properties, modes of reproduction, their cytological
differentiations, and varied pathogenic abilities were involved.
The group comprises several families and many genera and
species, with subdivisions of these, which from time to time call
forth long and expounding dissertations on the mechanism of
reproduction (the perfect stage). The problem of classification
is unfortunately a very complex one, since it has been found, at
least in the present work and with other closely related fungi
which will be considered later, that several forms of development
may be present in the same culture. Such phenomena are rather
rare but render incorrect any means of classification based on one
phase only.
It is well known that the fungi are divided, on the basis of
their morphology, cytology, and sexual development, into three
major divisions: Phycomycetes, Ascomycetes, and Basidio-
mycetes, and to these is appended a fourth group, the Fungi
Imperfecti, the life history of which is not at all or incompletely
known. The yeast-like organisms are considered to constitute
a branch of both the Ascomycetes and the Fungi Imperfecti,
the former where the production of asci is found, and the latter
where no sexual development has been determined. More
specifically, the perfect forms are considered to be members of
the Endomycetaceae (Gaiimann and Dodge, ’28; Rewbridge,
1 An investigation carried out at the Missouri Botanical Garden in the Graduate
Laboratory of the Henry Shaw School of Botany of Washington University, and
submitted as a thesis in partial fulfillment of the requirements for the degree of doc-
tor of philosophy in the Henry Shaw School of Botany of Washington University.
Ann. Mo. Bor. GARD., Vor. 20, 1933 (471)
[Vor. 20
472 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Dodge, and Ayers, ’29; Moore, '33), and Coccidioideaceae
(Moore, '32), including such genera as Endomyces, Coccidioides,
Rhinosporidium, and Pseudococcidioides, while the imperfect
forms, as Monilia, Cryptococcus, Oidium, M ycoderma, and several
others have features which are somewhat similar to the above
groups. ‘This of course does not include the true yeasts which
are included under the Saccharomycetaceae.
It was because of the imperfect knowledge existing in the field
that the author undertook the study of the cytology of Endomyces
capsulatus, which had been isolated from a case of meningo-
encephalitis. During the course of the work it was found
desirable to learn more about the phenomena which might be
associated with the pathogen, and hence attention was paid to
its physiological and other properties. In addition, the occurrence
of other infections by closely allied organisms, E. capsulatus var.
isabellinus (Moore, '33), and FE. dermatitidis (Moore, 733a),
have yielded the fungi which have been used in à comparative
study, so that what may pertain to E. capsulatus has been found
to hold true for the latter, with some minor exceptions as chro-
mogenesis and cultural characteristics.
HisronicAL REVIEW
Occasionally debatable points in the field of medical mycology
are cleared up by the isolation of some fungus. Such an organism,
E. capsulatus, was cultured in 1928 from a case of meningitis.
The patient was a white, married male, 48 years of age, a furrier
by trade. He entered the Boston City Hospital in August, 1928,
complaining of having had continuous headaches in the frontal
and temporal lobes for a period of three and a half months.
His illness began with a hard, tender, slightly movable mass on
the medial surface of the lower third of the left thigh, about five
months before entry. This mass subsided within four weeks,
but a similar mass then appeared beneath the anterior surface
of the neck. He developed defects in memory, losing himself
on familiar streets, and could not keep track of time. He lost
interest and was unable to understand questions or conversation.
The cyst which had been present at the level of the thyroid
isthmus between the trachea and anterior border of the sterno-
1933]
MOORE—ENDOMYCES CAPSULATUS 473
cleidomastoid muscle was aspirated. Thirty cc. creamy, tena-
cious, yellow pus was obtained, from which E. capsulatus was
cultured. Similar organisms were obtained from the pus aspi-
rated from the nodule of the left thigh.
About a month later, the patient’s condition became worse.
His temperature increased rapidly, as did his pulse and respi-
ration, and he finally died. The clinical diagnosis was mycotic
meningo-encephalitis (?), and an X-ray diagnosis showed bi-
lateral pulmonary tuberculosis. It has been the author’s
experience that an infection of this sort which involves the lungs
is often mistaken for tuberculosis, and the diagnosis made here
may have been incorrect. The organism from this case has been
used in these studies for the greater part, having been kept as a
stock culture by alternate transfers on nutrient and Sabouraud’s
agar.
The second case due to the variety isabellinus (pl. 21) occurred
in St. Louis and was reported by MacBryde and Thompson.
The patient, a white male, 28 years of age, had been a plumber
at the time of the appearance of the skin lesions. He was first
admitted to the Barnard Free Skin and Cancer Hospital in
February, 1930, with a palm-sized annular ulceration on the left
forearm just above the wrist, which involved the dorsal surface
chiefly and showed clinical signs of blastomycetic dermatitis.
Endomycetes were isolated from the pus of this lesion. About
3 cm. to the right of the anus was another lesion about 3 em. in
diameter and involving the anal margin. On the upper part of
the right arm was a scar of a lesion which had healed spontane-
ously. All active lesions were pruriginous and had a foul smell.
The patient later, in January, complained of coryza, with a mild
cough, pain in the right temporal region, accompanied by a rise
in temperature. His headache became universal and intense,
posterior to the eyes and extending to the occiput and base of the
neck. There was a marked bilateral Kernig reflex action. He
had nauseation and vomiting spells. A lumbar puncture gave
a cloudy fluid under slight pressure, in which were seen the organ-
isms found in greater abundance with later punctures. At this
time he received a large dose of iodine solution which cleared the
spinal fluid and decreased the number of budding cells, improving
[Vor. 20
474 ANNALS OF THE MISSOURI BOTANICAL GARDEN
his health greatly. However, two weeks later he had a relapse,
became comatose, and three days before death developed three
small subcutaneous abscesses. One was on the right wrist, one
on the sacrum, and one on the left side of the chest, and all
contained pus and E. capsulatus on culture. The patient died
forty-six days after the onset of the meningitis symptoms and
about three years after the first appearance of the skin lesions.
The morphology and cytology of the organism of this case was
found to be the same as the first fungus, having a very light cinna-
mon color as contrasted with the hyaline or white, in a mass, of E.
capsulatus, and cultural characteristics similar in detail.
The third organism studied is the one which has gone under
many generic names, as Blastomyces, Oidium, Saccharomyces,
Cryptococcus, and Mycoderma, because its life history was in-
completely known and the sexual act was not definitely estab-
lished, or the presence of the ascus and the actual number of
spores determined. Historically this fungus is important be-
cause it causes a disease known as blastomycosis which has
become fairly widespread, both in this country and Europe.
The organism studied by the author was isolated from a case
which occurred at the Barnard Free Skin and Cancer Hospital of
St. Louis and has been dealt with in detail in à previous paper
(Moore, 33). Suffice it to say here that E. dermatitidis, the
organism causing a clinical condition known as blastomycosis,
is similar to the above organisms, differing in its chromogenesis,
being a dark cinnamon to brown in culture and having a some-
what different cultural reaction, to be mentioned later.
In addition to the above three forms, an organism obtained
from a case of the so-called European blastomycosis or general-
ized torulosis reported by Urbach and Zach (730) (pl. 19, figs.
13-25) has been studied to a certain extent. The case in brief
was that of a 27-year-old shoe-worker who had never left Europe.
He entered the University Clinie for Syphilology and Dermatol-
ogy at Vienna with a swelling of the gums of the lower jaw
which was excised, and a half year later he returned with a similar
swelling of the gums of the upper jaw. About one year later
the patient noticed an abscess-like swelling on the left side of
the abdominal region, two months later a similar disorder on
1933]
MOORE—ENDOMYCES CAPSULATUS 475
his left thigh, and eight weeks later the same thing was found on
his neck. The lesion on his left thigh was aspirated and from
it 200 cc. purulent substance was obtained. A short time after
that an inflammation of the lungs developed which healed
spontaneously, but left him with a cough and vomiting spells.
Yeast-like cells were cultured from the purulent secretion of the
left thigh.
The disease was clinically identical with syphilis, tuberculosis,
leukemia, and several others. Inoculation of an extract of the
yeast-like cells (blastomycin) brought forth strong local, focal,
and generalized reactions, giving evidence of the presence of
blastomycosis.
The lesions were somewhat superficially healed by treatment.
The patient later developed high temperatures in the evening,
oral pains, deafness, violent coughing at night, and ruby-red
expectorations. A tumor of the nasal septum, very suggestive
of rhinosporidiosis, had developed and a catarrh of the Eusta-
chian tube as a result of the tumor. There was an ulceration of
the right tonsil and other lesions close by, and finally a heavy
exudate in his lungs. Treatment gave him relief temporarily,
but he returned to the clinic a short time later with a continuous
headache, high temperature, and rapid spread of the mouth
lesions. Paralysis of the optic musculature set in, with un-
consciousness, and death ensued.
The organism from this case has been cultured and subcultured
and as yet has shown no final stage as is present in the three
above forms. However, some organisms require a long time
before the perfect stage is obtained. Furthermore, the histo-
logical and pathological condition of the patient as investigated
by Chiari (30), shows the identical tissue reaction present in
cases of E. dermatitidis.
With this brief review of the history of the organisms in-
vestigated, let us now turn to a study of E. capsulatus.
TECHNIQUE
E. capsulatus was kept growing by subculturing on nutrient
and Sabouraud’s agar as a stock culture. In the work done here,
the organism was grown on nutrient agar, a product of the
[Vor. 20
476 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Digestive Ferments Co., at pH 6.8. In cases where an abundant
growth was necessary, with a thick mycelium, Sabouraud's agar,
pH 5.6, was employed.
For studying the cytology with regard to the nuclear changes
in the sexual act, glycerine agar (beef extract agar plus 6 per
cent glycerine, pH 7.1) was used. This medium gave the
greatest number of asci in culture, in addition to a thick growth
and many important diagnostic features, as chlamydospores,
conidia, racquet mycelium, etc. Cultures on nutrient agar and
Sabouraud’s agar were also used, but the cells were not so good
for details as on the above medium.
The cultures were fixed with a number of agents. Flemming’s
stronger solution shrunk the material rather noticeably, but
the weaker did not give very good results either. Bouin’s
picro-formalin solution, which has been advocated by Kater
(27) and other cytologists working with yeasts, was of no value
in this work. Benda’s fluid, which is a modification of Flem-
ming's stronger solution, caused the fungus to have too great an
affinity for the stains, and inasmuch as it is difficult to destain
the organism without causing some sort of damage, it had to be
used with caution. Hermann’s fluid, which is probably the
most expensive of the fixing agents, was also used. This is a
variation of Flemming’s fluid, the chromic acid being replaced
by platinie chloride, and although not particularly recommended
by Chamberlain (732), the author has found it to give the best
results. Various other fixing agents were employed, but the
results were not worthy of note.
The embedding proved to be a problem because the material
had to be fixed and embedded while on the agar. Paraffin was
used at first, but in the glycerine agar cultures it would not ad-
here to the agar substrate and sectioning could not be done
without damage to the material. Gradual and repeated changes
of paraffin did not remedy the condition. A medium was then
tried which contained very little protein and carbohydrate, but
no suitable growth could be obtained. Finally the cultures were
embedded in Dupont parlodion, using the Jeffrey technique.
This method has been recently outlined by Wetmore (32) and
with some modification was applied here. It consists in fixing
1933]
MOORE—ENDOMYCES CAPSULATUS 477
the material, in this case either with Hermann’s or Benda’s
fluid, pumping and dehydrating, then passing it through an
intermediate stage of ether-alcohol, and then through a series of
concentrations of the celloidin or parlodion. The first concentra-
tion was a 2 per cent solution and each series increased 2 per
cent until the final concentration was 12 per cent. The material
on the agar was taken from the test-tube after fixing, cut into
convenient pieces, and put through the above procedure. The
pieces were kept in a tightly plugged bottle in an incubator at
45° C. and changes were made daily. After embedding, sections
were cut to a thickness of 10 u. These were then stained and
mounted.
The best stain for this procedure was found to be iron-alum
haematoxylin, using Heidenhain’s haematoxylin, although Ehr-
lich’s gave just as good results. A combination of Benda’s
fixing agent and iron-alum haematoxylin showed the reticulated
network and the metachromatic material very well, whereas
Hermann’s fluid plus Heidenhain’s iron-alum haematoxylin was
best for nuclear structure. Methylene blue and Hermann’s
fluid brought out the vacuoles and volutin and the metachro-
matic material very clearly, as seen in pl. 23, figs. 15-18.
For morphological work, hanging-drop cultures were made of
2 per cent proteose peptone and 2 per cent bacto-peptone, as
well as lactose broth cultures. Material was also placed in a
drop of a 1 per cent solution of crystal violet (aqueous) in glyc-
erine, the dye being added to the desired intensity. The prep-
aration was allowed to stand from 15 to 30 minutes for a sufficient
clearing. Aman’s lactophenol was also applied, as was carbol
fuchsin and very dilute solutions of methylene blue, eosine, and
crystal violet. In addition, iodine potassium iodide (saturated
solution) was used for studying the glycogen contents of the
cells and chondriosomes, as advocated by Guilliermond. Osmic
acid, platinic chloride, and iodine green were also used, as well
as neutral red, but these materials and other methods of pro-
cedure will be explained later in the text.
CYTOLOGY
The cytology of the lower Ascomycetes, particularly the yeasts
[Vor. 20
478 ANNALS OF THE MISSOURI BOTANICAL GARDEN
and the yeast-like organisms, had been for a number of years a
topic of discussion among the older cytologists. The question
as to whether or not a nucleus was present was indeed a serious
one, judging by the numerous and lengthy dissertations on the
subject. As is customary, there were two sides to the argument.
One group maintained that the cells, of the yeasts in this case,
were made up of a mass of protoplasm and of nuclein, without a
true nucleus, and that the nuclein is differentiated at times in
the cytoplasm in the form of granules which assume a definite
color on staining. The other group favored the presence of a
definite nucleus.
The first reference to a nucleus in yeasts was perhaps made in
1844 by Nägeli, who decided that “ʻa little nucleus of whitish
mucus, lying on the membrane, regularly in each cell” (Wager,
’98), was often found in the yeast cell. Whether Nägeli actually
saw a nucleus in the fresh condition is rather doubtful, for an
oil globule has often been mistaken for the nucleus even to-day,
with our more advanced knowledge of cytology.
About five years later, in 1849, Schleiden, by treating cells
with ether-alcohol or potash, was able to find a rounded structure
with a clear cell wall which contained delicate granules either
singly or in groups, and in addition, a large flat body, which he
called a cytoblast.
Following this work, Brücke, in 1861, asserted that in living
material, as well as dead cells treated with iodine and acetic acid,
no definite nucleus could be seen and he reprimanded the former
workers by saying that no one was justified in taking bodies of
various sizes and numbers, such as often occur, for nuclei.
In 1879 Schmitz, using haematoxylin, was able to demonstrate
a nucleus in the cytoplasm of the cell near the vacuole, while
Strasburger, repeating this work in 1884 and 1889, using picric
acid and haematoxylin, was able to confirm these observations,
finding that the nucleus was not demonstrable in unstained
material.
Krasser, in 1885, contradicted these findings by saying that
granules existed in the yeast-cell, but no nucleus. His main
argument was that there was no specific staining reaction for
nuclei, and furthermore that the absence of a definite nucleus
1933]
MOORE—ENDOMYCES CAPSULATUS 479
in the yeast-cell was supported by the rapid growth of the
organism. For this reason he believed that there was nuclein
in the cell which was distributed through the protoplasm very
much as is generally held to be the case with bacteria to-day.
Other workers who held the view that a nucleus, or at least a
“corpuscle” as some called it, was present, the biochemical
properties and physiological functions of which were analogous
to those found in the cells of plants and of animals, were Hansen,
Strasburger, Zacharias, Moeller, Buscalioni, Henneguy, Dan-
geard (93), Janssens (02), and Janssens and Leblanc ('98). In
fact, the latter writers made the following statement, ''La
cellule de la levure peut étre considérée comme formé sur la type
général de la cellule. On y trouve en effet, un noyau, un proto-
plasme et une membrane." To these workers the nucleus had a
vacuolar appearance with a very differentiated structure, to
others it was a homogeneous body. Wager (’98), in a detailed
bit of work, described it as being a vacuole filled with chromatin
granulations (perhaps somewhat as is shown in pl. 23, figs. 17-18)
which had been taken for a nucleus by Janssens and Leblanc,
and furthermore that a spherical and homogeneous body, con-
sidered by some as the nucleus, was always adjacent to the vacuole
and could be compared with a nucleolus. He considered the
vacuole plus the eccentric nucleolus as a primitive stage in the
phylogeny of the development of the nucleus. This belief was
also based on the observation that the small body and the
vacuole divide simultaneously by budding.
Guilliermond, in 1902, demonstrated that a definite nucleus
was present, and that the vacuole was independent of the nucleus,
being filled with granulation products which at the present time
he holds to be nuclear decomposition substances (nucleic acid
derivatives with some unknown base). For the granules he
retained the name ‘‘corpuscules métachromatiques," a term
which had been applied previously and which is used even now.
Those who denied the presence of a nucleus but admitted the
presence of a nuclein substance as described above were Raum,
Roncali, Hieronymus, Macallum, and others.
In accordance with the views of the older writers, the author
has found that in Endomyces as treated here, when first isolated
[Vor. 20
480 ANNALS OF THE MISSOURI BOTANICAL GARDEN
from the tissue of the host, no nucleus exists, but simply a
distribution of the chromatin material or nuclein or metachro-
matic corpuscles throughout the cell, as may be seen in pl. 19,
figs. 13-14, pl. 20B, pl. 21, figs. 1-3, and several others. This
feature is a quite constant character of freshly isolated cells.
However, when kept on agar or an artificial substrate, the cells
not only change their form, passing through what may be termed
the secondary stage, that is changing from a yeast-cell to a hyphal
form, but also undergo a change in nucleoplasmic make-up.
It has been found that even though the yeast-like cells retain
the same morphological characteristics on agar media, their chro-
matin material or nuclein is converted into a definite nucleus.
This was clearly shown to be the case in E. dermatitidis (Moore,
'83). What the mechanism involved here might be, it is difficult
to say, but we may conjecture that it is linked up with the adap-
tation to a changed environment and a different mode of devel-
opment to which it must become accustomed, as shall be pointed
out later.
With such views being held in the past, it was of course to be
expected that the presence of any phenomena which might in-
volve the nucleus, as mitosis, would also be in dispute. In
yeasts the question of nuclear division has received considerable
attention, particularly by Guilliermond, Dangeard, and several
others, the former writing many and long papers on the subject,
and it has been said by some present-day workers, and the fol-
lowing is a direct quotation from the papers of one of them,
that ''the ideas of Guilliermond . . . which gain weight by the
mere bulk of his work on yeast, seem to meet with more favor."
The problem of the division of the nucleus in yeasts is perhaps
more complex than would seem offhand. "The same writers who
held that a nueleus was present in the cell first described nuclear
phenomena. Wager (’98) interpreted an amitosis in the yeast
with perhaps evidence of chromosomes. Janssens and Leblanc
(98), working with Saccharomyces cerevisiae and S. Ludwigii, as
well as Schizosaccharomyces octosporus and S. Pombe, described
a form of mitosis which, in the light of other works, seemed very
suspicious and they made the following statement, ‘‘On peut
dire que tout ce qui diminue la vitalité d'une cellule tend à réduire
la complication des phenoménes de division."
1933]
MOORE—ENDOMYCES CAPSULATUS 481
Dangeard (093) found that the nucleus elongated and the
nucleolus divided in two by an elongation to a thread-like process
within the nuclear wall. Swellengrebel (’05) and later Fuhrmann
(06) ascribed a definite mitosis to the nucleus, the latter studying
Saccharomyces ellipsoideus I. Hansen and presenting a typical
karyokinetie sequence with the formation of four chromosomes.
There were several authors who held to this view, and at present
there are those who believe that definite mitotic phenomena
are present. Kater (27), using a smear technique with S.
cerviciae [sic], demonstrated a mitosis, with the formation of
about eight chromosomes and a definite spindle, instead of divi-
sion by constriction. It should be pointed out here that smear
methods as applied to yeasts and particularly as carried out by
Kater have never yielded any results which might be considered
reliable. The act of smearing, no matter how good or how
careful the application of the fixative might be, usually allows
for some action on the cell wall that obliterates the correct
phenomena and substitutes artifacts, so that vacuolar constit-
uents or secretion products of the cytoplasm have been mis-
interpreted.
On the other hand, Guilliermond (17), in a summary of his
work, found amitosis to occur where budding was present. This
process was characterized by an elongation of the nucleus which
quickly divided by the resorption of the thread-like portion that
separated the two segments. Guilliermond also observed that
it was impossible to see the nuclear phenomenon clearly due to
the abundant products of secretion which covered the nucleus.
The observations of Kohl, of Wager and Peniston (710), and of
Pénau confirmed Guilliermond's results, and even went so far as
to actually claim definite amitosis.
Whether or not mitosis or amitosis actually occurs is still a
matter of dispute, despite the evidence that either side may
advance. "There are several factors which must be considered
in studying the process, or processes, in nuclear division. First,
does the nucleolus have a single morphological characterization,
such as exists in the higher forms, or does it constitute the total
nucleoplasmie material, as some call it, which has an affinity
for iron haematoxylin, for example, as in pl. 22, fig. 18? Second,
[Vor. 20
482 ANNALS OF THE MISSOURI BOTANICAL GARDEN
can the nucleus be seen so clearly that there is no mistaking the
change that may take place during the actual process? The
first question is difficult to answer because it has been found by
several workers studying the chemistry of the cell, yeast or
yeast-like or even that of the higher Ascomycetes, that certain
materials take the same dye. Then, the linin network with the
chromatin material or the chromomeres, as some call them,
usually cause confusion. The second question is really serious,
because we know that our most precise methods of technique
are still too crude for the minute nuclear make-up of these
lower Ascomycetes. Cell substances, or secretion products as
Guilliermond calls them, usually take stains which are peculiar
to the nucleus and mask any clear-cut pictures of nuclear phe-
nomena, and nucleic acid substances, derivatives of excretion
products of the nucleus, probably nucleophosphates and the
like, are also substances that must be considered in detail.
Besides, the size of the reactors in the process is too small for any
precise determinations with the means at hand.
On the other hand, according to the theories of modern genetics
there must be some mechanism whereby chromatic substance or
material is distributed through the agency of mitosis, whereas the
act of amitosis is simply a means of increasing the nuclear sur-
face or spreading the nuclear material through a cell and is
comparable to the fragmentation or lobulation of nuclei. The
latter is not at all reproductive in this respect. However,
there are many acts taking place in the daily life of a fungus,
particularly in these lower groups, which are impossible to explain
with our present knowledge of cytology or genetics, and theories
have been built up only to be torn down and built up again. This
does not mean that genetics is not standing on firm ground,
but that much more must be learned about mycological phenom-
ena before any generalizations can be made.
It is difficult to understand that the nucleus in the hyphae
of yeast-like organisms, E. capsulatus or E. dermatitidis, divides
by direct division, at least as far as can be made out from
the nuclear appearance. The process is evidently very rapid,
for an examination of a great many slides failed to show any
condition other than that shown in pl. 22, figs. 3-5. In the
1933]
MOORE—ENDOMYCES CAPSULATUS 483
ascogenous hyphae, or perhaps the antheridium and ascogonium
(pl. 22, figs. 6-9), the division is somewhat slower, and here
an elongation of the nucleus and nucleolus may be seen with
a deeper staining central portion indefinite as to character but
perhaps analogous to chromosomes.
In a discussion of nuclear phenomena, the next thing to be
considered is the mode of development or reproduction. Guillier-
mond (705, '05a, '05b, "08, "09, '09b, 710, "10a, "10b, TL '11a, 712,
'13,717,719,'20), Dangeard (793, '94, '94a, '94b, '97), Hansen (704),
and several others, in a long series of investigations, have de-
scribed the sexuality of the lower yeasts and some of the yeast-
like fungi. It is known, of course, that in the Ascomycetes,
particularly the lower forms, there may be three forms of devel-
opment, heterogamy, isogamy, and a reduction to partheno-
genesis. In the first case, two gametes, usually of unequal size,
a small one generally representing the antheridial cell and a
larger one the ascogonium (this is not a hard and fast rule), send
out one, or sometimes several, small tubes which copulate and
fuse. The cell contents of the antheridium then pass into that of
the ascogonium. The two nuclei, one from each gamete, fuse, and
finally an ascus develops through the subsequent division of the
fusion nucleus, the presumptive mother spore. In the case of
isogamy in the yeasts, two morphologically alike gametes fuse
in like manner. Finally in the series, there is a reduction to a
condition in which a cell may suddenly produce spores without
copulation. This condition has been found by Mangenot (’19)
for E. Lindneri and by Guilliermond for several other species, as
E. fibuliger and Zygosaccharomyces Pastori. It occurs also when
an ascogonial cell sending out a tube or tubes fails to copulate
and hence produces spores parthenogenetically.
In some species the above phenomena may be found separately,
or two processes or even all three may be present in the same
culture. This latter condition has been found to be true for E.
capsulatus.
In addition to the parthenogenetic formation of asci and asco-
spores, there is à non-sexual reproductive structure which has
not been given too much attention in the past. This is the
conidium (pl. 23, fig. 10), which is à non-nucleated structure
[Vor. 20
484 ANNALS OF THE MISSOURI BOTANICAL GARDEN
filled with chromatin granules or the metachromatic corpuscles
emphasized by Guilliermond. It occurs usually near a septum
of the hypha, and measures approximately 5 y in diameter.
Struetures of this sort may and usually do spring up from the
hyphae when there is a lack of nutrient material, or when the
hydrogen-ion concentration is fairly high (pl. 19, figs. 4, 7-8,
11-12). These may be pyriform (pl. 18, fig. 10) or round (pl.
18, fig. 7), sessile or on a short pedicel, with a thick wall (pl. 23,
fig. 10) ora thin wall. There is a heavy reticulated network with
the granules mentioned above usually occurring at the nodes of
the threads. These particular structures break off easily and may
serve as resting cells or chlamydospores which, when placed in
favorable media, germinate, form nuclei by an accumulation
perhaps of the chromatin material, and develop a normal growth.
They represent probably a degeneration from an ascus, or, on
the other hand, an advanced character of reproduction.
To get a clearer understanding of the life history of E. cap-
sulatus, it would be advisable to begin with the organism as it
is found in the parasitized host and explain the mechanism by
which it and related fungi propagate themselves.
In the tissue the organism grows as a yeast 6-8 y. in diameter,
and reproduces itself by budding. There is no nucleus present,
at least it could not be demonstrated in tissue sections or in
freshly isolated cells, but instead there is a distribution of nuclein
material throughout the organism, much the same condition
as exists in bacteria, Myxophyceae, or in the conidium shown
here. When isolated and grown on an artificial substrate as on
agar, the chromatin or nuclein material seems to become larger,
and there is usually formed a nucleus and many large granules
which may be presumptive nuclei. This latter condition is
based on circumstantial evidence (pl. 21, figs. 4-9; pl. 20B). The
cells then pass through this stage, the yeast-cells, to stage two
which consists of large irregular cells, attaining a condition as
seen in pl. 19, figs. 14-15, 20-25, and have been clearly shown for
E. dermatitidis. At this stage the chromatin material is spread
throughout the cell, but the reticulated network seen in the later
stages is not very clearly established. However, it seems to be
rapidly developing and nuclei are clearly distinct. The stage
1933]
MOORE—ENDOMYCES CAPSULATUS 485
following is probably the most complex of all, inasmuch as there
is à diversity in morphology and cytology, changes due probably
to hydrogen-ion concentration and temperature and several
other factors among which the changed habitat is outstanding.
In this third and final stage, we find that sexuality has developed
and asci with ascospores are produced. To follow the life cycle
on an artificial substrate, it would then seem best to follow the
development from the germinating ascospore. In doing this,
it is necessary to consider first normal growth on some favorable
medium, and second to correlate the nuclear changes in material
fixed in hanging-drop preparations (Van Tieghem cells) and
stained, with the nuclear phenomenon found in the material
fixed and stained as outlined in the technique. It was noticed
that from three to fifteen days were necessary for the life cycle
of the fungus to be completed, depending on the broth used in the
hanging-drop preparation. It was found that proteose peptone
broth gave good growth as did lactose broth, but that bacto-
peptone plus bacto-beef gave quickest development of asci,
from three to five days usually. The material was fixed by the
addition of two or three drops of a fixative described previously,
and after a period of from six to eight hours, or over night, the
fluid was drawn off with filter-paper and the material was care-
fully washed with distilled water and then mounted in lacto-
phenol plus crystal violet, or stained with iron haematoxylin.
By keeping several of these hanging-drop cultures growing, it
was possible to obtain different stages of growth, even in one
preparation. The method was of course not entirely accurate
cytologically, but it served the purpose as demonstrated here.
The single spore (2-215 u) is found to be made up of densely
staining granular material, but the granules are so small that
it is difficult to discern them. On germinating, a tube is sent
out or the spore simply elongates, taking with it the chromatin
material (pl. 22, figs. 1-2). When somewhat older, the devel-
oping hyphae produce nuclei which divide rapidly as pointed
2 Several solutions were used: lactose broth (product of Digestive Ferments Co.),
pH 6.8; 2 per cent bacto-peptone broth, adjusted to pH 7.0; 2 per cent proteose
peptone broth, pH 7.0; bacto-peptone plus 6 per cent glycerine; meat extract broth,
pH 7.1; 2 per cent bacto-peptone plus 5 per cent bacto-beef (dehydrated), pH 7.2.
[Vor. 20
486 ANNALS OF THE MISSOURI BOTANICAL GARDEN
out previously, are distributed in pairs very close together, and
then separate (figs. 3-4). At this time the reticulum is well
developed and the chromatin granules, or basophilie grains, as
Guilliermond calls them and which he believes probably rep-
resent albuminoid bodies playing the róle of products of nutri-
tion in the form of perhaps zymogen or reserve material, are
very evident and take a deep stain with haematoxylin. At the
same time, the wall increases in thickness and becomes fairly
evident as in fig. 4. The number of nuclei in the hyphae varies,
as many as thirty-seven having been counted on one hypha,
but this was apparently rare for the amount usually varied between
seven and fifteen with nine the most common, as in fig. 4. With
the elongation of the new hypha, septa are laid down (fig. 5)
and the formation of conidia as described previously (pl. 22,
fig. 5; pl. 19, fig. 4) may be and usually is associated with this act.
Following this, the hyphae, varying from 11% to 4 u on different
media, may grow out to form branches or reproductive structures,
usually lateral or terminal. However, on acid media or on
media where there is a definite lack of some suitable protein
or protein product in the form of peptone, peptides, or amino
acids, there is an excessive formation of chlamydospores, either
terminal (hypnospores), 415-6 x 9-12 u, or lateral, 5x 7 u, or
interealary, approximately 5 y in diameter, and conidia (pl. 18,
fig. 7; pl. 19, figs. 7, 11-12). With the apparent development of
a hypha, there is an accompanying formation of sexual cells.
This process may be slow or rapid, depending on conditions. On
glycerine agar it seemed to be the most rapid, these particular
organs being produced in three to five days. The antheridial or
male cell contains two or three nuclei as a rule (pl. 22, figs. 6-8),
but cases of four have also been seen. The ascogonium or female
cell usually contains three to four nuclei (figs. 9-11), but five
(fig. 12) have also been noticed.
When two sexual cells are ready for copulation, the antheridium
bends over to meet the ascogonium as seen in pl. 22, figs. 10-11.
The two gametes may be on the same hypha or on separate
hyphae. Both may be terminal (figs. 13-17), or both lateral
(fig. 16), or one may be lateral and the other terminal (fig. 12),
or there may even be variations of these. Nevertheless it is
1933]
MOORE—ENDOMYCES CAPSULATUS 487
quite apparent that the male cell goes to meet the female cell.
With the approach of the copulation branches, there is an action
which may be termed tropistic and which may be explained, if
analogies are acceptable, by a hormone action or physical stimulus
as is found in zoological specimens, which calls forth the produc-
tion of a beak, the copulating tube, as has been found in the lower
yeasts. These two beaks meet and fuse. In the meantime,
one of the nuclei proceeds to the tip of the beak. There is
probably no specific nucleus involved in this act, because as far as
can be made out all the nuclei are the same morphologically, and
any nucleus that happens to be nearest the beak becomes sensi-
tized to the fertilization reaction. With the progression of the
fertilization nucleus, there is an accompanying retrogression of
the remaining nuclei. Whether these disintegrate or not has not
been definitely determined. In several cases where fertilization
had taken place, it was noticed that the antheridial cell and
basal portion of the ascogonium merely existed very much like
functionless stamens in a flowering plant, which in the course of
time disintegrated or perhaps degenerated.
The next step in the cycle involves the fusion of the nuclei, or
syngamy perhaps. The walls between the two tubes are dissolved
or diffuse and the two nuclei copulate or fuse in the tube (pl. 22,
figs. 13-16). This is rather contradictory to Dangeard's ('04,
'04a, '94b, '97) observations that the two nuclei fused in the
ascus, and Harper's (96, ’97, '99, '00), that in the higher
Ascomycetes the fusion occurred in the archicarp after the
entrance of the contents of the antheridium. The fusion nucleus
is then transferred or migrates (whatever the process may be it is
analogous to that by which the nuclei copulate) into the asco-
gonium which is now known as the archicarp (pl. 22, fig. 17).
The fusion tube dissolves in some cases or breaks away in others,
with the broken portion being either resorbed by the cells or
degenerating. At any rate, the archicarp repairs its walls by
the resorption of the cellular contents, and the large fusion
nucleus is ready for division (fig. 18).
The nucleus divides by a process of amitosis as previously
mentioned. Although difficult to interpret theoretically, it
presents no clear-cut chromosomal formation. With the first
[Vor. 20
488 ANNALS OF THE MISSOURI BOTANICAL GARDEN
division, there is apparently the formation of a wall (pl. 22,
fig. 19). The daughter nuclei go through a synchronous division
(fig. 20) to form four nuclei. In several of the yeasts and in Æ.
Magnusii, the process halts here and each nucleus develops into a
spore, thus four spores. Here, however, the four nuclei undergo
a synchronous division, the second for the ascus, and eight nuclei
are produced, which number is à mean for the species of this
genus. In other species there may be twelve or sixteen nuclei.
In the higher Ascomycetes the problem of ascus formation is
rather confusing, and the reader is referred to the work of Atkin-
son ('15), Bagchee (725), Blackman and Fraser (05), Brooks
(10), Brown (710, '11), Carruthers (11), Guilliermond, Mangenot
and Plantefol (33), Faull (05), Fraser and Brooks (709), Fraser
and Welsford ('08), Gaümann and Dodge (28), Harper (796, '97,
'99, '00), and Maire (’04, '05), to obtain full information and
various opinions on the subject.
After the formation of the nuclei and the abjunction of the
future ascus (8-14 y. in diameter) from the basal portion of the
ascogonium, now a basal cell as seen in pl. 22, figs. 21-22, the devel-
opment of the spores begins. Associated with this phenomenon
are the secretion products which, as has been pointed out pre-
viously by Guilliermond, nourish the spores and produce the
spore wall.
It should be pointed out here that, in addition to this hetero-
gamous form of copulation, isogamy sind finally parthenogenesis
may also be present. The latter is quite apparent and has been
found time and time again. The process may begin by the
jutting out of what is apparently a conidium. "This in turn
develops and a nucleus becomes visible. The nucleus divides
as noted in the above reaction and an ascus with eight ascospores
forms (pl. 23, figs. 1-9). Also a terminal cell often produces a
pyriform ascus parthenogenetically as seen in fig. 11. In all
cases, the spores serve the same function and are equally as
capable of producing new hyphae. The sporogenous plasma of
the ascus, part of the cytoplasm, contributes greatly to the
formation of the mature spore.
The life cycle of E. capsulatus on an artificial substrate might be
graphically illustrated as in diagram 1.
1933]
MOORE—ENDOMYCES CAPSULATUS 489
Cellular contents.— The remainder of the ascus constitutes the
epiplasm which is made up of reserve products in the form of
lipoids, glycogen, the metachromatie granules of Guilliermond
or nuclear decomposition products as volutin, oil globules, and
nucleie acid substances and probably other protein derivatives
and carbohydrates. All of these materials have been shown
to be utilized by the spores in their growth. Several authors also
present evidence that during the maturation process of the spores,
£
antheridium |
bot — ,hyphac > Spscus
| pon E
ascogenous hypha
chlamydospores
hvpnospores —+ hyphae
‘conidia -——
Diagram 1.
the substances constituting the epiplasm are broken down, a
part being absorbed and clearly demonstrable in the granular
contents at maturity, and a part reserved for their germination.
Volutin.—The cellular contents will be considered in the order
studied. The first is what has been variously called ''Neisser's
granules" found by Neisser in bacteria; ‘‘sporogenous grains"
found by Ernst; ‘‘metachromatic corpuscles’ of Babés, who
found them in the diphtheria bacillus and so named them because
of their metachromatic action; ‘‘red grains" of Bütschli, so
named because they took on a red coloration with many of the
stains; and finally, one of the most common of all, volutin, as
named by Meyer.
In the fresh state these granules stain blue with methylene
blue (pl. 23, figs. 16-18), and when fixed their color is red to
violet. They are small at first, and then enlarge to assume various
forms and sizes. They may have a heavy staining outer portion and
[Vor. 20
490 ANNALS OF THE MISSOURI BOTANICAL GARDEN
a paler center, probably due to their refractive powers. They are
seen quite often in vacuoles as represented here and assume
fantastic features. ‘They appear in the cell along the reticulated
network, at the nodes, and stain heavily there. Volutin is very
common in the younger cells, but not so abundant in the older
hyphae. ‘The substance may also be stained in vivo with neutral
red. Wager and Peniston (710) found Gram's aniline violet to be
a useful stain for volutin.
Zikes (22), studying the nature of volutin and of the physio-
logical factors concerned in its production, found that it is present
in moderately large amounts in nearly all fungi. Peptone when
added to the culture media stimulated its production while
ammonium sulphate and asparagin were not so favorable. Phos-
phorus was found to be essential for its formation, and glucose
and fructose were more favorable than the carbohydrates of
higher molecular weight. He concluded that volutin is an
albuminous substance similar to the nucleo protein, as it contains
both phosphoric acid and nuclein.
The general opinion seems to be that volutin is a nucleic acid
substance with an unknown base, probably organic in nature,
that it exists as a colloid, and that, due to fixatives, it is pre-
cipitated in the vacuoles. It is supposedly a secretion from the
nucleus and is used in nourishing the spores.
Glycogen.—Another substance which supposedly acts as a
reserve material and is used up for the maturation of the spores
in the ascus is glycogen. This theory has been substantiated
by the fact that glycogen is used up in the cell, particularly in
maturing asci, as evidenced by its slow disappearance. Kohl
(07) considered glycogen not as a reserve material, but as a
regulator for the intake of sugar in the cell, in that it is the sub-
stance formed, not being able to diffuse out from the membrane.
He found it to be lacking in spores, and since reserve materials
should be present, he presented this as evidence that glycogen is
not a reserve material. Guilliermond, however, claims to have
demonstrated that glycogen is absorbed by the ascospores at the
time it is found to disappear from the epiplasm.
Glycogen is found in the organism practically from the begin-
ning of growth, first as small droplets perhaps, but later in
1933]
MOORE—ENDOMYCES CAPSULATUS 491
fairly large masses (pl. 23, figs. 19-30). It has been estimated as
constituting 32 per cent of the dry weight of yeast cells. It is
associated with the nutritive condition by some workers and
found to be almost entirely used up or greatly accumulated
according to the condition of nutrition and growth. According
to some workers it appears somewhat in the form of a vacuole
and it has thus been called a glycogen vacuole. It is not a true
vacuole, but perhaps a colloidal substance, as volutin is considered
to be, forit takes a stain much as thelatter does. It may be stained
in vivo with neutral red (figs. 26, 29-30), in which case it is not so
clear, at least in E. capsulatus, as with a saturated solution of
iodine potassium iodide, a modification of Gram's stain (figs.
19-25, 27-28).
This substance is a glucoside, a polysaccharide which, like
starch, is made up of n molecules of glucose to have the formula
(CeHiOs)n. It is often compared with starch since it requires
phosphorus for its formation. In fact, it has often been referred
to as an amylopectin, which makes up the superficial part of the
starch grain. By hydrolysis, glycogen is broken down to dextrin,
then maltose, and finally glucose. On heating a saturated
solution of glycogen stained with iodine potassium iodide it
becomes pale but regains its intensity on cooling. It is insoluble
in alcohol.
Vacuoles.—The question of vacuoles in fungi has been given
considerable attention in the past, and the author does not
intend to enter into a discussion of them. However, in view of
the many theories advanced as to their character and presence,
mention of them in E. capsulatus would not be out of place.
They are easily demonstrable with many dyes and present
varied characteristics. With methylene blue, they are easily
made visible (pl. 23, figs. 15, 17-18). They appear to be formed
by the reticulum and to take the stain. They may be demon-
strated fairly clearly with iodine green (fig. 14), and the standard
dye seems to be neutral red, in which case small crystalloid
bodies, often described as ‘‘dancing bodies," probably Brownian
movement, may be seen (figs. 26, 28). Haematoxylin also
brings them out distinctly by staining the surrounding network
(figs. 1-2). What the function of vacuoles may be is difficult
[Vor. 20
402 ANNALS OF THE MISSOURI BOTANICAL GARDEN
to understand. It is known that they are colloidal solutions of
cytoplasmic materials, formed by the absorption of water, that
they may enlarge or disappear in a cell, and Guilliermond,
Mangenot and Plantefol, say this of them, * Les vacuoles rep-
résentent la phase aqueuse du cytoplasme et jouent certaine-
ment le róle important en réglant la teneur en eau de la cellule."
Chondriosomes.— he bodies designated as mitochondria or
chondriosomes have been demonstrated in animal and plant
tissue. What their particular function may be is not definitely
known. In plants they have been associated with the elaboration
of chlorophyll and starch through the agency of the chloroplasts.
In animals they are supposedly associated with the elaboration
of secretions, but no definite evidence has been produced along
this line. They have been demonstrated in a number of lower
plants, lower fungi and the higher Ascomycetes, and in fact,
in practically all of the groups, as well as in animal cells. For
further information the reader is referred to Guilliermond ('11b),
who has traced the development of these particular bodies in
many organisms.
Chondriosomes (Cowdry, 17) have been shown in both animal
and plant cells to be almost completely soluble in alcohol, ether,
chloroform, and dilute acetic acid (organic solvents) and to be
insoluble by chromization and treatment with formalin, at least
in most cases. In the animal cell they do not stain with Sudan
III or IV and are sometimes blackened with osmic acid. In
plants, the whole cell blackens with osmie acid so readily that
it is impossible to know just to what extent the chondriosomes
themselves are affected.
In studying E. capsulatus, the author made use of the knowledge
that chondriosomes are very clearly demonstrated by the addition
of iodine potassium iodide to a preparation of an organism.
They are fairly evident as lightly staining rod-like and short
bodies (light yellow with iodine), but one must pay particular
attention to distinguish them (pl. 23, figs. 19-25). "There are
short and rod-like bodies in the terminal hypnospores and some
long forms too, as well as small point-like reflective granules.
The rods are longer in the hyphae (figs. 23-24). Chondriosomes
are scattered throughout the asci as fine, small, rounded bodies.
1933]
MOORE—ENDOMYCES CAPSULATUS 493
It can thus be stated that by the iodine potassium iodide method,
bodies comparable to those described as chondriosomes for other
fungi are demonstrable.
Fat, lipoidal substances.—In addition to the many substances
mentioned, there are also fats, lipoidal substances, and other
materials called reserve materials, secretion products, and also
excretion products. These materials are found in varying
amounts in many of the specialized portions of the organism,
being abundant in the asci and chlamydospores and in very
small amounts in young hyphae and younger elements. They
can be demonstrated comparatively easily in spores and in
yeasts particularly. The nature of many of these substances has
not been determined as yet, but it is generally known that fatty
acids, glycerides and sterides, glycerol, phospholipides and
phosphoaminolipides (complex lipides) may be present in certain
amounts.
Several agents were used for the study of these materials,
each giving some degree of difference in distinguishing them. A
2 per cent osmic acid solution which reduces fats and gives
them a black coloration was the first tried. When applied to the
living mycelium, the fats and lipoidal substances appear as small
refraetile bodies or droplets but in some cases they are very
much larger (pl. 23, figs. 31-36). "They are seen rather abund-
antly in old hyphae (fig. 36) and in very small amounts in young
hyphal tips (fig. 35). With platinic chloride (5 per cent solution)
they appear as blackened granules much the same as with osmic
acid (figs. 37-38). Iodine potassium iodide as applied for
glycogen and chondriosomes shows lipoidal substances, as oil
droplets, equally as well. These are very small, highly refractile,
and hyaline, and to be seen require careful focusing and adjust-
ment of the microscope (figs. 19-20, 23-25). Neutral red has
also been used, but not much attention was paid to the reaction
outside of what has already been mentioned with regard to
glycogen and the vacuoles. Iodine green also has an affinity
for lipoidal substances, but not enough attention was given to it
to prove its value here.
[Vor. 20
404 ANNALS OF THE MISSOURI BOTANICAL GARDEN
CULTURAL CHARACTERISTICS
In studying the cultural characteristics of E. capsulatus,
all the media available at the time were utilized. This was
considered essential, inasmuch as it has become an unfortunate
custom with some mycologists to name as new species organisms
which show a physiological variation on a different medium.
Since the fungus had shown on previous occasions of culturing that
it favored protein substrates, a variety of protein materials was
chosen. In addition, regular routine media were used, some with
much carbohydrate and some with very little. For some years
the question of hydrogen-ion concentration has become an im-
portant factor in work of this sort, and media with a wide range
of pH were chosen within which the possibilities of growth had
at one time or another been emphasized. All cultures were
grown at a temperature of approximately 25? C.
The following media arranged in the order of their decreasing
hydrogen-ion concentrations were used:
Raulin’s Solution Agar (Raulin’s solution plus 1.5 per cent agar,
pH 4.1).—(pl. 18, fig. 7; pl. 19, figs. 8, 10-12). Growth poor,
being present only around inoculum after 18 days and having a
diameter of only 0.7 em. and 1.6 em. at end of 30 days. Color
white. Hyphae grown heaped up from center in a loose fashion.
Microscopically, abundance of conidia, pedicelled or sessile,
5 y. in diameter; endo-chlamydospores 5-7 y. in diameter; terminal
hypnospores 4-6 x 8-11 u; hyphae 2 y in diameter; racquet
mycelium in small amount, not very noticeable; asci 9-10 v in
diameter.
Richards’ Solution Agar (Richards’ solution plus 1.5 per cent
agar, pH 4.3).—(pl. 19, figs. 5-7, 9). Growth slow, very sparse
and cottony, with long and narrow hyphae 2 y in diameter,
projecting loosely and irregularly from the edge of the colony
which attained a diameter of approximately 1.8 em. at end of 18
days and 3.5 em. at end of 30 days. Round conidia very nu-
merous, approximately 4 y. in diameter; pyriform conidia sessile or
on short stalks, several, 314 x 7 w in diameter; round, thick-walled
chlamydospores on short pedicels, 5 u in diameter; asci few,
7-9 y. in diameter; endo-chlamydospores approximately 6 y in di-
ameter.
1933]
MOORE—ENDOMYCES CAPSULATUS 495
Czapek’s Agar (pH 4.4).—Growth of very loose and sparse
mycelium which spreads over the surface of the agar with thin
hyphae. Macroscopically the culture is barely visible except
by reflected light. Colony approximately 3 cm. in diameter at
end of 18 days and 6.7 cm. at end of 30 days. Color white.
Hyphae 214-31% y in diameter; racquet mycelium present, 6-7 y.
in diameter at swollen portion and 214-3 y at thin portion;
terminal hypnospores 7 x 12 v approximately; conidia many, 5 y
in diameter; round chlamydospores 6 y in diameter; lateral
chlamydospores 6 x 11 yu; asci several, 10 » in diameter.
Wort Agar (Product of Digestive Ferments Co., pH 4.6).—(pl.
18, figs. 6, 8-10, 15). Growth at first slow, none on several of
the cultures, in general thin and loose. One of the cultures
showed good growth, attaining a diameter of 3.2 cm. at end of
18 days and 5 em. at end of 30 days. This was unusual and may
be ascribed to too great an inoculation and to a dissemination of
spores as a result of shaking. Color white. Terminal hypno-
spores many, 7 x 12 y; hyphae 314-4 y. in diameter; chlamydo-
spores 614-7 u in diameter; racquet mycelium abundant; conidia
numerous, 5-6 y in diameter; asci 10-11 y in diameter.
Malt Extract Agar (pH 5.1).—(pl. 18, figs. 1-3). Growth not
so abundant, colony 1.6 cm. at end of 30 days. Color slightly
brown, due to the malt extract. Mycelium of numerous swelled
cells; hyphae 3-4 x in diameter; terminal hypnospores 6 x 16 y;
hyphal swellings or chlamydospores 9 x 12 u; numerous conidia,
pyriform and round, the round being approximately 5 y in di-
ameter; asci very few, 10 u in diameter.
Malt Extract Broth (The above minus the agar).—Very little
growth at end of 30 days. Characteristics same as above.
Maltose Agar (pH 5.4).—Growth slow. Colony coremium-
like, with a diameter of 1.5 em. at end of 30 days. Hyphae short,
3-4 u in diameter. Many round chlamydospores 5 y in diameter;
terminal hypnosp numerous, 4-6 x 9-11 u; asci few, 8-10 y
in diameter.
Sabouraud's Broth (Sabouraud's dextrose agar minus the agar, pH
5.5).—(pl. 21, fig. 21). Culture consists of submerged colonies
of mycelium varying from V4 to 3 cm. in diameter at end of 18
days, with a great mass at the end of 30 days as a result of the
[Vor. 20
496 ANNALS OF THE MISSOURI BOTANICAL GARDEN
coalescence of all the colonies. The large flakes are grey in color
when moist, but with white mycelium on the surface, fairly dry.
Submerged hyphae 214 y in diameter with none or very few
morphological characteristics. Aerial or dry mycelium, however,
similar to that on the agar culture of the same medium.
Sabouraud’s Agar (pH 5.6).—(pl. 16, figs. 12-14; pl. 19, figs.
13-25; pl. 21, figs. 14, 18, 28). Thick cream-colored growth
with a diameter of 4.5 em. in 18 days and 7 em. in 30 days.
Hyphae 2-3'5 y in diameter, in needle-like projections from the
surface of the mycelium which appears very cottony. Color
white. Older cultures show a felt-like matting with a tendency
towards ridge formation, a condition found to a certain extent in
Microsporon Audouini Ota and Langeron. Racquet mycelium
present with swollen portions 4-6 » and narrow section 3 y;
many lateral chlamydospores, 5-7 x 10-12 u; terminal hypno-
spores 5 x 11 u; and many conidia, pyriform or round, approxi-
mately 5 u in diameter; numerous asci, 10-12 y in diameter, with
8 ascospores 2-216 y in diameter.
Oat- M eal Agar (Decoction of oat-meal plus dextrose and agar, pH
5.9).—Growth diffuse and slow, 5 em. in diameter after 30 days.
Culture loose and cottony. Hyphae 2-214 y in diameter;
numerous thick-walled, round resting cells or chlamydospores,
4—6 p in diameter; terminal hypnospores 4 x 7 u; conidia numerous,
4 y in diameter; asci several, approximately 8 u in diameter.
Corn-Meal Agar (Product of Digestive Ferments Co., pH 6.0).—
(pl. 16, figs. 8-9; pl. 21, fig. 24). Growth very loose and thin,
with the hyphae projecting from the colony, appearing as threads
of silk. Colony barely visible except from a lateral view. Hyphae
long and thin, 2-214 y in diameter, with colony attaining a
diameter of 5 cm. at end of 21 days and 6 cm. at end of 30 days;
chlamydospores 6 y in diameter; conidia many, 4144-5 y in di-
ameter; terminal hypnospores 5 x 7 u; racquet mycelium present
but reduced in size, 3-4 y. at swollen portion; asci several, 7-8 y.
in diameter.
Potato-Dextrose Agar (Decoction of potatoes plus dextrose and
agar, pH 6.2).—(pl. 21, fig. 22). Colony 3.6 em. in diameter
at end of 30 days, edge smooth and round, no striations or other
cultural changes as ridges. Growth thick and cottony at in-
1933]
MOORE—ENDOMYCES CAPSULATUS 407
oculum, with a thin periphery about 2-3 mm. in width. Color
white. Hyphae 2-214 y in diameter and long; conidia numerous,
5 y in diameter; terminal hypnospores few, 4x 7 v; chlamydo-
spores rare; asci des: 8-10 y in diameter.
Two Per Cent Aqueous Bacto-Peptone (Hanging-drop culture, pH
6.2).—(pl. 21, figs. 1-8, 25, 32). Mass of thick-walled hyphae
ramifying and branching, 2-215 y. in diameter; racquet mycelium
not evident; numerous chlamydospores, 4 y. in diameter; asci 4—6 y.
in diameter; few terminal hypnospores, 3.5-5 x 6-7 y.
Two Per Cent Aqueous Bacto-Peptone Plus Five Per Cent Meat
Extract (Hanging-drop culture, pH 6.2).—(pl. 19, figs. 1-4; pl.
21, figs. 9-13). Growth profuse, covering the drop in 4 days.
Hyphae 2-214 y in diameter, with an abundance of racquet
mycelium intertwining and branching; thick-walled cells, chlamy-
dospores in abundance, as well as asci 10 » in diameter.
Two Per Cent Proteose Peptone Plus Six Per Cent Glycerine
(Hanging-drop culture, pH 6.2).—(pl. 16, fig. 7; pl. 17; pl. 21,
figs. 15, 30). Growth similar to that on bacto-peptone broth,
but with an abundance of chlamydospores and racquet mycelium.
Lactose Broth (Product of Digestive Ferments Co., pH 6.8).—
(pl. 21, fig. 17). Growth of submerged large flakes of colorless
mycelium measuring approximately 2 cm. in diameter at end of
18 days. These later coalesced or intertwined into a mat which
grew up the sides of the flask to form a white mycelium. Sub-
merged hyphae approximately 3 u in diameter, branching, inter-
twining, with cross-walls. Swellings, chlamydospores, terminal
hypnospores, and asci few in number and reduced in size as
compared with those on agar. The aerial mycelium above the
surface of the broth showed an increased number of characteristics
which simulated those found on the agar.
Lactose Agar (The above medium plus 1.5 per cent agar, pH,
6.8).—(pl. 18, fig. 11; pl. 19, fig. 20). Growth rapid, thick and
cottony, colony attaining a diameter of 3.6 cm. at end of 18 days
and 6.2 em. at end of 30 days. Culture showed 4 ridges radiating
from a thick inoculum to a thick cottony circular periphery
surrounded by a thin growing rim of hyphae. Culture similar
to that on nutrient agar, macroscopically. Hyphae 214-3) y
in diameter; conidia many, 414-5 y. in diameter; terminal hypno-
[Vor. 20
498 ANNALS OF THE MISSOURI BOTANICAL GARDEN
spores 4 x 715 u; racquet mycelium present in abundance; many
chlamydospores, varying from 6 y in diameter (round) to 5x 8 u
(pyriform); asci many, 8-9 y in diameter.
Nutrient Agar (Product of Digestive Ferments Co., pH 6.8).—
(pl. 21, fig. 19). Growth good, with the colony having a diam-
eter of 3.5 em. at the end of 20 days and 7.0 cm. after 30 days.
Growth loose and cottony, showing concentric circles, evidently due
to a periodical formation of asci and the liberation of the spores
which germinated. Single spore colonies 2 cm. in diameter
after 15 days. Colony flat with age, and white. Hyphae 2-3 u
in diameter; conidia many, sessile or pedicellate, 5 y in diameter;
chlamydospores several, 6 ». in diameter when intercalary, 5 x 7 y
when lateral, 6 x 11 u when terminal (hypnospores); racquet
mycelium present, 5 x 3 y.
Nutrient Broth (Meat extract, pH 6.8).—(pl. 21, fig. 31). Cul-
ture similar to that on lactose broth.
Uschinsky’s Protein-free Medium (pH 6.8).—No growth.
Beef Extract Agar (Liebig's extract of beef, pH 7.0).—(pl. 16,
fig. 15). Growth similar to that on lactose agar. Radiating
ridges present.
Eosine-Methylene-Blue Agar (Product of Digestive Ferments
Co., pH 7.0).—(pl. 21, fig. 27). Growth good, attaining a
diameter of 5 em. after 26 days. Color pink in younger portion
of culture (periphery), darker pink to blue towards the inoculum,
this being due to an absorption of the dyes by the mycelium.
Growth ceased and colony became flat. Hyphae in growing
culture 2-316 y in diameter, with morphological characteristics
similar to those on Sabouraud's agar.
Glycerine Agar (Nutrient agar plus 6 per cent glycerine, pH 7.1).—
(pl. 16, figs. 1-3, 11; pl. 21, figs. 16, 23, 26, 29). The colony
on this medium grew as a cerebriform, very thick, creamy culture,
having a diameter of 6 cm. at the end of 30 days. Medium best
for study of organism because of its nutrient constituents, pres-
ence of abundant protein and carbohydrate, and an abundance
of endo-chlamydospores, 515-6 x 11 v; intercalary chlamydo-
spores 4-5 x 5-7 v; terminal hypnospores 4-6 x 9-12 u; hyphae
2V5-4 u in diameter; round chlamydospores, either terminal or
lateral, approximately 6 y in diameter; racquet mycelium abund-
ant; asci numerous, 10-14 y in diameter.
1933]
MOORE—ENDOMYCES CAPSULATUS 499
Blood Agar (pH 7.2).—(pl. 16, figs. 4-6). Growth similar to
that on nutrient agar.
Chocolate Agar (Blood agar heated to about 75? C. until the blood
became chocolate colored, pH 7.2).—Growth similar to the above.
Serum Agar (Beef extract agar plus 10 per cent dehydrated blood
serum, pH 7.2).—(pl. 18, figs. 4-5, 12-14, 16). Growth moist,
showing a diameter of 1.4 cm. at end of 10 days and 3.5 em. at
end of 30 days. Colony flat and even. Many yeast-like cells
seen; hyphae few and several large cells. Center of colony
cerebriform. With age the culture shows prickly forms of
mycelium (dry) which covers the greater part of the culture.
This evidently is a reversion to the yeast form, to a certain
extent.
Calcium Carbonate Agar (pH 7.4).—(pl. 16, fig. 10). Growth
slow at first, then rapid, with a diameter of 7.4 cm. in 30 days.
Colony compact, with a cerebriform central portion, and a loose,
cottony outer zone. Color white. Hyphae 2-214 y. in diameter;
conidia numerous, 414-5 y. in diameter, being spread throughout
the entire culture, mostly round, several pyriform on short
peduncles, others sessile; round chlamydospores many, 6 y in
diameter, and terminal hypnospores 4-5 x 7-9 v; racquet my-
celium not very evident; asci many, approximately 8 & in di-
ameter.
Endo's Agar (Product of Digestive Ferments Co., pH 7.5).—
Growth slow, not quite so rapid as that on eosine-methylene-blue
agar. Colony white at first, then, due to an absorption of the dye
from the agar, pink to red, growing in coremium-like masses of
straight hyphae with very few or no conidia. Hyphae short
and thick-walled, 315-4 » in diameter. Few conidia in culture,
6 u in diameter; hypnospores very few, as well as round chlamydo-
spores; asci several, 10 u in diameter.
Litmus Milk.—Milk was heated in a flask for fifteen minutes
in steam, then set away over night in the ice-chest to allow the
cream to rise. The cream was then siphoned off and the milk
diluted in the ratio of 1 part milk to 4 parts water, with enough
litmus as an indicator. Tubes of this solution were sterilized by
steam, then inoculated, and kept at 25° C. No growth resulted
after 20 days.
[Vor. 20
500 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Reaction to Temperature.—In studying an organism, it is
necessary, especially if quantitative results are desired, to deter-
mine conditions under which optimum growth can be obtained.
One factor which enters into such considerations with fungi is
temperature. It is known that various organisms grow best at
certain temperatures and when placed in other conditions,
growth will either be inhibited or retarded. With this point in
mind, it was decided to grow a number of cultures on the same
medium and same pH at various temperatures. As a medium,
Liebig’s beef extract agar at pH 7.2, which on previous occasions
had shown a qualitatively and quantitatively good growth,
was employed.
E. capsulatus, when first isolated from the lesions in its yeast-
like form, showed very good growth at body temperature,
37.5? C. After having been kept in culture in its filamentous
form for about two years, the optimum temperature was shown
to be approximately 30? C. Continued growth on an artificial
medium again reduced its optimum temperature, to approximately
25? C., as may be seen in table r, or figs. 1-2.
Several cultures in petri dishes were grown at the various
temperatures indicated in table r. The experiment was per-
formed at three different times, and due to the fact that daily
measurements required handling of the cultures, several of the
plates showed contamination. A procedure such as this usually
brings in some foreign spores, especially in a large laboratory where
many open cultures may be found.
'The plates of the first two temperatures, —0.7? C. and 8.0? C.,
were kept in a Kelvinator, the former in the freezing unit where
a fairly constant temperature was maintained, and the latter in
a compartment some distance from the cooling device. The
third set of plates, 16.0? C., was kept in a glass jar with running
water around it. The temperature fluctuated somewhat, but
not enough to affect the experimental data seriously. The
plates of the other temperatures were kept in regulated incuba-
tors.
All the plates were inoculated with a loopful of a suspension
mashed up with a sterile needle and contained 5 cc. of a beef
extract broth solution of pH. 7.2. In some cases the inoculum
proved greater than others.
1933]
MOORE—ENDOMYCES CAPSULATUS 501
TABLE I
MEAN DIAMETERS IN CENTIMETERS OF n GROWN AT
VARIOUS TEMPERATURE
Beef extract agar (Liebig's extract of beef) pH 7.2
Temperature in degrees Centigrade
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Readings were made daily for 31 days, at approximately the
same hour. The mean results of a representative series are
given in table r. "Three plates were kept in the final table as
representive of the experiment although several additional
plates were used.
[Vor. 20
502 ANNALS OF THE MISSOURI BOTANICAL GARDEN
An analysis of the results of this experiment shows that no
growth takes place at —0.7° C. and at 40.0? C. At 8.0? C. growth
does not begin until the twenty-third day and then proceeds
slowly, with à maximum diameter of 1.2 cm. which would rep-
resent a total growth of 0.9 em. At 16.0? C. the culture does
not show growth until the fourth day, and grows slowly but
faster than that at 8.0? C., to show a final diameter of 3.9 cm.
or 3.6 em. actual growth. At 21.0? C. there is a final diameter
of 6.7 em., with a total growth of 6.4 em. At 25.0? C. the di-
ameter of the colony reached an optimum of 8.3 em. and an actual
growth of 7.9 em., and at 31.0? C. the diameter of the colony was
6.2 em. with a total growth of 5.8 em. The culture at 37.0? C. grew
fairly well for the first ten days but after that no further growth.
Transfers from these last plates to fresh medium produced no
growth, hence it could be concluded that the heat finally killed the
organism.
In fig. 1, the diameter of the colony in em. was plotted against
the time in days to obtain the rate of growth for each temperature.
It may be seen here how the growth was affected. At optimum
temperature, after the initial lag period of 1-2 days, growth
proceeded in practically a straight line. With a deviation from
the optimum temperature there is accordingly a decrease in the
growth rate which corresponds to the temperature affecting it.
At 16.0° C. we find a regular sigmoid curve which represents an
initial lag period followed by a period of increased activity and
finally a return to the lag period.
When the maxima points of total growth are plotted, fig. 2
(diameter of colony against temperature), we have the relation-
ship of the total growth to the various temperatures outlined.
These results seem to indicate that the optimum growth for
this organism has changed from approximately 37.5? C. in its
parasitic condition to 25? C. in a saprophytic condition. This
latter fact will be demonstrated in the animal inoculation experi-
ments. It also shows that the longer an organism is kept in
culture the greater will be its change in physiological phenomena
to an optimum point for its changed environment.
Hydrogen-Ion | Concentration.—The problem of hydrogen-ion
concentration has been of interest for several years and its
1933]
MOORE—ENDOMYCES CAPSULATUS 503
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consequences especially emphasized by von Mallinckrodt-Haupt
(32). It has been demonstrated that the change of pH in a
medium plays an important part in the cultural characteristics
[Vor. 20
504 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Temperature in degrees Centigrade
Fig. 2 Maximum growth at various temperatures on beef extract agar with same pH.
rr ra) + M
"moO UT 89[uoTo9 JO si939uverq
of the fungi grown. These changes may be brought about in
their nutrient medium through the formation of acids which are
1933]
MOORE—ENDOMYCES CAPSULATUS 505
present in large amounts, by the breakdown of the sugar molecule
in its utilization by the organism, by the decomposition of the
proteins or the products, and by the cracking of fats or lipoids as
a result of growth. As is perhaps fairly evident in the case of the
carbohydrate reactions to be explained later, we have the forma-
tion of alkaline bases which are observed in the course of the
utilization of the proteins.
From a study of the cultural characteristics mentioned pre-
viously and from further observations, a decided difference in
growth on acid and alkaline media may be pointed out. In
strongly acid media there is abundant budding and a formation of
numerous conidia, and on strongly alkaline media a tendency
towards shorter, thicker cells and a yeast-like cell formation. On
the other hand, on weakly acid or alkaline media there is a favor-
able condition established for mycelial and hyphal formation.
The effect of alkalinity and acidity on the development of fungi
was noticed for a number of years. Marantonio (93) found that
a more acid medium favored a greater quantity of mycelium.
These observations were confirmed by Concetti (’00) and were
extended to include a large number of media. Strains of an
organism isolated from cases of thrush were used. Fineman
(21), working with Monilia albicans apparently, found that
mycelium grew better in media under low surface tension and oxy-
gen tension while the yeast form of the organism predominated on
solid media, simple carbohydrates, and a high pH. — Milochevitch
(29) found that varying the hydrogen-ion concentrations of media
had no effect on the same sort of a fungus. Talice (30) found the
best development of hyphae on dilute potato decoction, the hydro-
gen-ion concentration being perhaps fairly high.
several workers have pointed out that fungi possess a certain
buffer action, that is, they have the ability to regulate their
growth according to the reaction of the medium, and it is found
to exist only under external changes. This means, then, that
acid media become less acid during the course of growth and
alkaline media become less alkaline.
What has been found to be true for the effect of various tem-
peratures on the growth of E. capsulatus may also be said to hold
with regard to hydrogen-ion concentration. For testing this,
[Vor. 20
506 ANNALS OF THE MISSOURI BOTANICAL GARDEN
beef extract agar made up with Liebig’s extract of beef but
adjusted to the various pH’s shown in table rr, was inoculated as
described previously. The same procedure was carried out as in
the above experiments, namely, daily diameter measurements
and occasional microscopic examinations. Here, as before,
TABLE II
MEAN DIAMETERS IN CENTIMETERS OF COLONIES GROWN IN
VARIOUS HYDROGEN-ION CONCENTRATIONS, AT A
TEMPERATURE OF 26°
Beef extract agar (Liebig’s extract of beef)
Days
iue pH values
inocula-
tion 2.1 | 3.3 | 42 | 5.2 | 5.5 | 6.1 | 6.4 | 7.0 | 7.4 | 7.7 | 8.2 | 9.3 [10.4
1 0.2 | 0.8 | 0.3 | 0.3 | 0.3 | 0.3 | 0.3 | 0.4 | 0.3 | 0.4 | 0.3 | 0.4 | 0.4
2 g» 0 0.3 | 0.3 | 0.3 | 0.3 | 0.3 | 0.4 | 0.3 | 0.4 | 0.3 | 0.4 0
3 0 0 0.3 | 0.4 | 0.4 | 0.4 | 0.4 | 0.6 | 0.5 | 0.6 | 0.4 | 0.4 0
4 0 0 0.3 | 0.5 | 0.5 | 0.6 | 0.7 | 0.9 | 0.8 | 0.9 | 0.6 | 0.5 0
5 0 0 | 03] 0.7 | 0.8 | 0.9 | 0.9 | 1.2 ]1.2] 1.2108 10.7] 0
6 0 0 0.3 | 1.0] 1.1 | 1.2 ]1.2 | 1.4116] 1.6] 1.1 | 0.9 0
7 0 0 0.31}1.8/1811.6118) 1.7 | 20,19] 14] 1.1 0
8 0 0 0.311.411. | 1.18]1.9|2.1|]2.3|22]|1.8]| 13 0
9 0 0.|0,88.1171108]|3.151 22:1 3.8 |.3/77 | 3812.0) 1.8) 0
10 0 0 0.4 | 2.0 | 2.2 | 2.4 | 2.4 | 2.6 | 3.0 | 2.8 | 2.4 | 1.8 0
11 0 0 0.5 | 2.3 | 2.5 | 2.7 | 2.7 | 2.9 | 3.3 | 3.2 | 2.7 | 2.0 0
12 0 0 0.6 | 2.6 | 2.8 | 3.0 | 8.1 | 3.2 | 3.7 | 3.5 | 2.9 | 2.2 0
13 0 0 0.8 | 2.9 | 3.1 | 3.3 | 3.4 | 3.6 | 4.0 | 3.8 | 3.2 | 2.8 0
14 0 0 | 10] 381 | 3.4] 3.5 | 3.7 | 38148 | 411]356)124/ 0
15 0 0 1.1 | 3.3 | 3.6 | 3.8 | 4.0 | 4.1 | 4.6 | 4.4 | 3.8 | 2.4 0
16 0 0 13|3.7|3.99|42|4.2|4.5|4.9|4.7| 41 0 0
17 0 0.4 | 1.6 | 4.0 | 4.3 | 4.5 | 4.6 | 48 | 5.3 | 5.0 | 44 0 0
18 0 |} 06] 1.8]43 | 45148 |50 | 5.2 |56] 54)471] 0 0
19 0 | 0.8] 2.1 | 46) 4.8] 51153 | 5.5 )]59] 5.7] 5.1] 0 0
20 0 | 0.9 | 2.3 | 4.9 | 5.1 | 5.4 | 5.6 | 5.9 | 63 | 6.0 | 54) 0 0
21 0/|[09/|[25|51/|53/|57/|60/|62/|606/|6.4|56]| 0 0
22 0 0.9 | 2.7 | 5.3 | 5.5 | 6.0 | 6.3 | 6.5 | 6.9 | 68 | 5.9 0 0
23 0.|090|2,8154|56|62]8.5|068|72|7.1|02]| 0 0
24 0 1.0 | 3.0 | 5.7 | 5.9 | 6.4 | 6.7 | 7.0 | 7.4 | 7.2 | 6.4 0 0
25 0 1.0 | 3.1 | 5.8 | 6.1 | 6.6] 6.9 | 7.2 | 7.6 | 7.4 | 6.5 0 0
26 0 0132/|58/|63/|068|7.0|7.4|]7.8|7.0|6.7| 0 0
27 0 0 0 5.9 | 6.4 | 6.9 | 7.1 | 7.5 | 8.0 | 7.8 | 6.9 0 0
28 0 0 0 6.1 | 6.5 | 7.0 | 7.2 | 7.6 | 8.2 | 8.0 | 7.0 0 0
29 0 0 0 6.2 | 6.6 | 7.1 | 7.8 | 7.7 | 83 | 8.1 | 7.1 0 0
30 0 0 0 568-167 | L1] T4 | 7.1] 84 182 71 0 0
*0 indicates no growth
1933]
MOORE—ENDOMYCES CAPSULATUS 507
contaminations were present, so that only three plate measure-
ments were incorporated in the table. The mean results in
table 1 represent a typical set of data in this work.
An analysis of the data shows that no growth took place at a
pH of 2.1 and 10.4. At pH 3.3, growth was not evident until
the seventeenth day, when the diameter was 0.4. "The total
growth was 0.7 cm. on the twenty-fourth day, after which it
stopped. A transfer of the culture to a medium of pH 7.2 gave
normal good growth. With an increase in pH to 4.2, growth was
at first slow, but the colony attained a diameter of 3.2 em. on the
twenty-sixth day and then growth stopped, showing a total increase
of 2.9 em. In like fashion, by subtracting the initial inoculum
measurement from the final colony diameter, we find that at pH
5.2 total growth for thirty days was 6.0 cm.; for pH 5.5, 6.4 em.;
for pH 6.1, 6.8 em.; for pH 6.4, 7.1 cm.; for pH 7.0, 7.3 em.; for
pH 7.4, we find the optimum pH with a total growth of 8.1 em.;
for pH 7.7, 7.8 cm.; for pH 8.2, 6.8 cm., which is the same as that
for pH 6.1; for pH 9.3, total growth of 2 em. was acquired on the
fifteenth day, with the colony showing no additional growth.
By plotting the diameter of the colony against the time in days,
fig. 3, we obtain the rate of growth for the optimum pH which
shows, except for the initial lag and a final lag, the latter being
due perhaps to external factors as slight drying and probable
utilization of most of the nutrients, an almost straight line.
This varies with the pH, showing a decreased growth rate from the
optimum, with an increased lag and decreased period of activity.
By plotting the maxima points of growth or total growth for
each pH against the pH values, we obtain a clear relationship of
the growth at various hydrogen-ion concentrations, with the
point of optimum growth forming a peak, as seen in fig. 4.
In addition, the pH of several of the media after growth was
determined. No definitely accurate results were obtainable due
to the dried-up condition of several of the media, but indications
seemed to be that there was an increased pH where growth was
best, with no change in the strongly alkaline and strongly acid.
The results may be interpreted by considering the amount of
growth as an index of the production of alkali due to the utiliza-
tion and decomposition of the protein substances in the medium.
[Vor. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
508
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1933]
509
MOORE—ENDOMYCES CAPSULATUS
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[Vor. 20
510 ANNALS OF THE MISSOURI BOTANICAL GARDEN
SUMMARY OF CULTURAL WORK
The results of the cultural studies described above for the three
organisms, Hndomyces capsulatus, E. capsulatus var. isabellinus,
and E. dermatitidis, but more especially for E. capsulatus, may
be summarized as follows:
1. Ability of the organisms to grow on a fairly wide variety of
media.
2. No growth or very little growth on media lacking or having
very little protein or carbohydrate.
3. An increase in the number of conidia and decrease in diam-
eter of hyphae on acid media as compared with fewer conidia
and increased thickness of hyphae on media with a high pH.
The number of conidia and size of cells may be an index of the
hydrogen-ion concentration of the medium.
4. E. capsulatus and its variety isabellinus show an almost
direct change from a yeast-like to mycelium growth macroscopi-
cally, as evidenced by the change from a moist colony to a
filamentous colony, whereas E. dermatitidis shows a macro-
scopically evident secondary stage in the form of a definite, prickly
or coremium-like growth. The secondary stage is identical
microscopically, for all the organisms.
5. E. capsulatus has a white color, isabellinus a light chamois or
isabella, while E. dermatitidis has a dark brown or cinnamon color.
6. Size and number of various morphological characteristics
differ on various media.
7. Reversion to the yeast-like form by growth on serum agar.
8. No growth on litmus milk.
9. The organism shows an optimum temperature of 25° C.
with no growth occurring at temperatures above 37° C. and below
8.0? C.
10. The optimum pH is shown to be approximately 7.4 with
deviations from the mean, and no growth occurring in a pH
higher than 9.3 or lower than 3.3.
BIOCHEMICAL REACTIONS
Carbohydrate Reactions.—There have been many conflicting
results in the past with regard to the carbohydrate reactions of
yeasts and yeast-like organisms. Systems of classification have
1933]
MOORE—ENDOMYCES CAPSULATUS 511
been established whereby an organism was placed in a definite
taxonomie position because of its production of either acid or
gas, or both, or whether or not it had an effect on a carbohydrate.
Such a system has been established by Castellani for Monilia.
Whether any confidence can be placed in a method of that sort
is doubtful, first, because it is problematical, at least in some
cases, whether absolutely pure sugars are obtainable, and small
amounts of impure substances may bring about an altered
reaction which may not coincide with the specified phenomenon.
That is, a reaction may be indicated as slight or weak, which
actually should have been negative. Dekker in 1931 outlined
the various methods of sugar fermentation and the faults to be
found with each, and for further information, the reader is referred
to her work. In the second place, it has been evidenced in the
past and frequently noticed at present, that pathogenic organisms
of the sort mentioned above have often lost their fermentative
abilities on standing in artificial media. The constant sub-
culturing has resulted, as in the case of many fungi, in a change
from a pathogenic and virulent parasite to a non-harmful sapro-
phyte, and while this may not necessarily be an index of the sugar
reactions of an organism, still we must consider such occurrences
as of some significance. It would seem, therefore, that not much
reliance must be placed on a system of that sort.
In the work carried out here, stress was laid not so much on
the desire to organize and establish a new method of classification,
such being altogether too numerous, but to determine whether a
yeast-like organism, such as E. capsulatus, could produce any
acid or gas. For this purpose, Pfanstiehl sugars, which have
been found to be considerably above the average, were used.
The sugar was added, at the rate of one per cent, to a beef extract
broth as prepared previously, to which had been added phenol
red as an indicator. The phenol red was made up as a 0.02 per
cent solution in distilled water and added in the amount of 1.5
cc. of indicator to 10 cc. of medium. This gave a fairly deep
color which proved satisfactory in the work. This particular dye
was chosen because preliminary experimentation had shown
that the reaction tended more towards alkalinity. Litmus was
also used in several series, but the results were not so clear and
definite as with phenol red.
[Vor. 20
512 ANNALS OF THE MISSOURI BOTANICAL GARDEN
A 12-day-old culture on beef extract agar was mashed up with
a sterile needle and 5 cc. beef extract broth of the same pH were
added to form a suspension. A loopful of this supension was
inoculated into Smith fermentation tubes which contained the
above medium. All the tubes were grown at 25? C.
The carbohydrates used were those listed in table rri. For
the sake of clarity, they may be outlined as follows:
Monosaecharides.
Pentoses:
Aldoses;
1-arabinose, 1-xylose
Hexoses:
Aldoses;
rhamnose, dextrose, d-galactose, d-mannose
Disaccharides.
lactose, maltose, saccharose
Trisaccharides.
raffinose
Colloidal polysaccharides.
dextrin, starch (soluble), inulin
Glucosides.
amygdalin, salicin
These carbohydrates represent a fairly wide range and include
those sugars which are used in the regular routine sugar reactions,
as xylose, dextrose, maltose, and lactose.
There will be no attempt at present to explain the work in
detail or to offer any complex chemical formulae for possible
decompositions, but the reactions and a possible empirical reason
for the observations will simply be pointed out.
An examination of table mr in which the carbohydrates are
arranged in the order of their increasing groups, shows that when
compared with the control (the same medium minus the carbo-
hydrate) in the first column there was a changed reaction due to
the presence of a carbohydrate. It will be noticed that the
control tube showed a change on the eleventh day and a period
of reactivity for six days, reaching its maximum on the seven-
teenth day after inoculation. It is further seen that 1-arabinose,
l-xylose, dextrose, d-levulose, lactose, maltose, saccharose, and
soluble starch showed the same initial period of definitely de-
513
y
MOORE—ENDOMYCES CAPSULATUS
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1933]
SNOILOVEY ALVHOAHOHNVO
III @IaVL
(Vor. 20
514 ANNALS OF THE MISSOURI BOTANICAL GARDEN
monstrable reactivity, by the color changing to red, and that
rhamnose showed a lag and hindered the reaction greatly before
the color change, this being demonstrable on the nineteenth
day. Amygdalin had a somewhat less harmful effect than
rhamnose, with the reaction occurring on the sixteenth day,
and salicin even less than that, having the initial color change on
the fourteenth day. In contrast with the normal, we find that
d-galactose and dextrin were very favorable towards color pro-
duction, inducing a reaction on the fourth day, whereas d-
mannose, raffinose, and inulin took effect in perceptible color
changes on the ninth day.
It might be added here that three series of tubes were used in
each experiment, two series being used to obtain the changes in
pH, which were measured by a potentiometer and also a set of
colorimetric standards. A set of tubes was prepared with phenol
red at various hydrogen-ion concentrations. 'This method
proved very satisfactory for further work, being much more
rapid and fairly accurate since readings were made only to the
first decimal place. It was found in practically every case that
& change in hydrogen-ion concentration occurred two days
before a color change, except in the case of d-galactose and dextrin
where the reaction was more rapid, taking place on the third day.
This latter reaction may be explained somewhat by the fact that
the two carbohydrates are very favorable to growth, since the
color change is due to the growth of the organism which brings
about a reaction in the medium as a result of the utilization of the
nutrients.
A striking feature of these reactions is the varied range of
reactivity as may be seen in fig. 5. No explanation, outside of
the rate of utilization of the carbohydrate, can be given at the
present time for this phenomenon. As compared with the
control, the range for dextrin is very large. By the range of
reactivity is meant the period over which the color changes and
the decrease in hydrogen-ion concentration take place, coming to
an end point of an approximate pH of 8.1. The various carbo-
hydrates present various ranges which are clearly evident in fig. 5.
Soluble starch, added to the solution at the rate of 0.2 per cent,
showed a partial hydrolysis on the nineteenth day as evidenced
1933]
515
1
MOORE—ENDOMYCES CAPSULATUS
'SsejurpAqoqr99 JO APIATZOVaI JO oguv] o “BLT
[Vor. 20
516 ANNALS OF THE MISSOURI BOTANICAL GARDEN
by a reddish-brown color on addition of a dilute iodine solution.
Fehling's solution reduced to cuprous oxide as shown by a faint
reddish-violet color. There was no complete hydrolysis at the
end of the experiment.
It may also be noted that in its production of alkalinity the
reaction proceeded in every case from the aerobic portion of the
Smith tube, or rather from the bulb, to the arm of the tube, or
the closed portion. In other words, there was a diffusion of ions
from the aerobic to the anaerobic. This of course was to be
expected, since the organism is strictly aerobic, as will be shown
later, and utilized all of the little material available, before growing
towards the anaerobic portion. In every case, however, no matter
how marked the reaction, the color change in the arm was compara-
tively slow.
Proteins are broken down very easily by reacting substances,
and the cultural experiments have shown that E. capsulatus
requires some protein, its derivatives, or carbohydrate, for growth.
It is also known that these proteins and their derivatives may be
further reduced or broken down to amino acids, peptones, pep-
tides, and hexone bases and other substances, whereas higher
carbohydrates may be reduced to simpler sugars. Ammonia is
often found to be a product in the hydrolysis of proteins. In
the presence of sugars, amino acids have characteristic reactions,
particularly in an alkaline solution, and these have been dealt
with by several authors, recently by Watanabe (732). This
author lists many reactions and covers the literature of the field
rather well, and the reader is referred to his paper for detailed
information. Euler and his associates, in some work carried
out in 1926 and 1927, found that the reaction between amino
acids and sugars which were reversible, took place at an equimo-
lecular ratio and that their velocity became greater in an alkaline
medium. ‘This may account for the rate of activity as noticed
in table rm. The work of several others seems to point to the
conclusion that a reaction between amino acid and sugar requires
a neutral reaction and a fairly high temperature, whereas Wald-
schmitt-Leitz and Rauchalles observed that the optimum hydro-
gen-ion concentration for the reaction between dipeptides and
glucose was pH 8.1. This seems to coincide with the results
obtained here.
1933]
MOORE—ENDOMYCES CAPSULATUS 517
It is clearly evident from the experimental data that, instead of
acid or gas or both, the carbohydrates here used induced the
production of an alkaline condition. In addition to the protein
decomposition substances as mentioned above, ammonia may be
an end product. This substance, where present in fairly large
amounts, brings about a strongly alkaline condition. To test
for it, several cc. of the culture broth were poured into a test-tube,
and a pinch of anhydrous sodium carbonate (Na,CO;) was added.
'The tube was shaken and a rather strong odor of ammonia was
given off. It is possible that, besides the evident ammonia,
hexone bases, as arginine, lysine, and histidine, which are alkaline
in solution, are formed as products of the protein derivatives in
the medium and that in addition to the sugar reaction they
cause a decrease in the hydrogen-ion concentration.
Indol and Skatol.—Reactions which have long been used in
bacteriological technique but which have not received much
attention until lately, due to the additional knowledge and better
methods which are applied, are those of indol and skatol. The
work of such investigators as Pittaluga (08), Goré (21), Holman
and Gonzales (23), Morelli (09), Sasaki (10), Zipfel (12),
Krumwiede and Pratt (113), Fellers and Clough (25), Koser and
Galt (26), and several others has greatly added to the methods
of technique in investigations. Tests have been advocated
from time to time, the importance of which lay in their specificity
for the reaction. Fellers and Clough list about thirteen tests
which have been applied, some successful some not.
Indol and skatol are protein derivatives and their determination
rests on the presence of tryptophane. From it they are broken
off by a deamidization to form first indol proprionie acid, then
indol acetic acid, and finally indol or skatol. The medium to be
used in such a reaction must therefore contain a supply of tryp-
tophane sufficient to determine whether the organism would
actually be capable of breaking it down to indol or skatol or both.
For this purpose, it was necessary to use peptone plus 0.1 per
cent casein in water to insure good growth and also to comply
with the requirements as stated. The broth was inoculated and
the organism was allowed to grow at 25°C.
In the first test the Gnezda (799) oxalic acid principle was
[Vor. 20
518 ANNALS OF THE MISSOURI BOTANICAL GARDEN
applied. Strips of filter paper were placed in a warm solution
(saturated) of oxalic acid. On cooling, crystals of oxalic acid re-
mained on the strips. The paper was then dried well and inserted
in the mouth of the flask under aseptic conditions, so that it was
pressed against the side of the flask and just above the surface of
the broth. Ifindolis formed it volatilizes and colors the oxalic acid
pink. In these tests for E. capsulatus, a very faint pink was barely
discernible. This test has been advocated rather strongly by the
Society of American Bacteriologists for regular routine work in
bacteriology.
A test that was applied successfully is the Ehrlich-Bóhme
technique which consists of two solutions. Solution 1 contains
1 gm. para-dimethyl-amino-benzaldehyde, 95 ec. ethyl alcohol
(95 per cent), and 20 cc. hydrochloric acid, concentrated. Solu-
tion 2 is a saturated solution of potassium persulphate (K;S,0;).
In the experiment, 5 ec. of solution 1 are added to 10 cc. of the
culture fluid, then 5 cc. of solution 2, and the mixture is shaken.
A faint red color appearing about five minutes later indicated a
weakly positive reaction. This method has proven rather
successful for indol. Steensma (06, 068) substituted vanillin
for the para-dimethyl-amino-l lehyde, but this was not
applied here.
The Salkowski ('83) nitroso-indol test was negative for indol.
Skatol was tested for by the Sasaki (10) methyl alcohol test.
This method consisted of adding to the broth to be tested about
four drops methyl alcohol and an amount of concentrated sul-
phuric acid equal to the broth. The acid contained a trace of
ferric salt. A reddish-violet color indicated a positive skatol
reaction.
Under the proper conditions, indol and skatol may be produced
by E. capsulatus.
Relation to Free Oxygen.—Aerobiosis has often been considered
important in experimental work. Organisms are classified
as strict aerobes, facultative aerobes, or facultative anaerobes
and strict anaerobes. In order to find the position of E. cap-
sulatus in this classification, tubes of broth were prepared and
inoculated by the Liborius method (Zinsser, ’29) and kept at
25? C. Cultures were kept at the same temperature and in the
1933]
MOORE—ENDOMYCES CAPSULATUS 519
same broth, but were plugged only, allowing oxygen to enter.
No growth occurred in the former while a luxuriant growth
occurred in the latter. Agar stab cultures were negative for
growth in the stab, but positive on the surface. The relationship
to free oxygen was also demonstrated in the carbohydrate re-
actions as mentioned previously.
We may conclude, therefore, that E. capsulatus is a strict
aerobe, at least in so far as its growth on agar after a period of
time has demonstrated. When in the yeast form, anaerobic
cultures showed slight growth, a condition of facultative anaero-
biosis. Continued growth on an artificial substrate has con-
verted it into a strict aerobe.
Reduction of Nitrates.—The tests for nitrate reduction as
advocated by the American Society of Bacteriologists were
inconclusive and unsatisfactory. Ammonia was found to be
present, but that was apparently due to the breakdown of the
peptone and other protein decomposition products. ‘Tromms-
dorf's method was also indefinite, being negative in general, but
at times gave a slightly colored reaction.
Production of Hydrogen Sulphide.—The formation of hydrogen
sulphide in culture has often been concerned with a toxic action
of the lead salt or lead acetate in the medium toward certain
bacteria. Whether hydrogen sulphide would be produced here
was of course problematical. However, a lead acetate agar put
up by the Digestive Ferments Co., ‘‘Bacto lead acetate agar,"
which is claimed to contain too little lead salt to be toxic and yet
was favorable for the demonstration of the hydrogen sulphide,
was used. Making stab cultures was of course illogical, for
reasons pointed out before, and so surface inoculations were made
and the cultures incubated at 25? C. A slight brownish coloration
was found to occur directly around the colony after thirty to
forty days, and increased somewhat on standing.
No definite method has been advocated for the above experi-
ment, and since the results were slightly evident even though the
period was a great deal longer than for bacteria, the reaction
might be considered positive.
Gelatine Liquefaction.—Tubes of 15 per cent meat extract
gelatine were inoculated by stabbing and incubated at 25°C. A
[Vor. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
520
slow liquefaetion took place between 30 and 40 days, and in some
cases even longer than that.
and proceeded downward.
growth of the fungus.
Liquefaction began at the surface
This may be correlated with the
REACTION To LIGHT
A great deal of experimental work has been done in the past,
and much is being attempted at the present, with respect to the
AALL LTT
LLL.
UP RARA AARAA LR 2
CLL LL LLL SLA LAA ho
rr 8 A AFF Bm
SS ff WAR ma
SAA AeA A AAAA ART O
DSL AR AARAA ITTI anon
SIF TD A A T
Fig. 6. Explanation in text.
It is
action of light on higher plants and also microbiology.
accepted as a fact by many that plant growth and vitality can be
greatly altered, either for better or worse, by continued treat-
ment under variously colored lights.
No great amount of work,
however, has been done on the action of the primary colors on
Several workers have found a destructive
fungi or bacteria.
1933]
MOORE—ENDOMYCES CAPSULATUS 521
action of light on the lower organisms, and Rahn has found that
red and green light have no effect while blue light has a very
harmful action, with the degree of destruction decreasing in the
violet and ultra-violet portion of the spectrum.
Effect of White Light on Growth.—To determine the differential
action of light and darkness on EF. capsulatus, series of plates were
e (|
Fig. 7. Explanation in text.
covered with opaque black paper of the sort used in covering
photographic plates, as illustrated in fig. 6, the portions with the
crossing lines representing the covered areas. The plates were
treated as follows: plate A was inoculated in the center and
completely exposed to the light; plate B was covered as shown
and was inoculated at a point about 1.5 em. from the rim of the
plate, or about 5 em. from the uncovered area. "This plate was
used to determine whether the organism would grow towards the
[Vor. 20
522 ANNALS OF THE MISSOURI BOTANICAL GARDEN
light or not. The plates covered as in C were inoculated in the
center of the portions covered, and plate D was inoculated as in
plate A and served as the complete darkness plate. Nutrient
agar was used. ‘These plates were placed in a light-proof con-
tainer illustrated in fig. 7, at room temperature. A General
Electric mazda bulb of 50 watts and 115 volts was used as a
source of light. A flow of water was kept constant beneath the
source of light to prevent the heat from reaching the interior of
the container. A sheet of colorless glass (A) was kept between
the flowing water and the cultures. Several cultures of each
series were kept in the container and the experiment was per-
formed on several different occasions. Plates prepared as above
were kept at room temperature and were subject to diffuse day-
light and darkness. Growth at the end of 24 days showed no
cultural or microscopical differences in any case, except for a
slight increase in diameter of three plates kept in the freely
circulating air. This small number of attempts seemed to
indicate that light had no effect on growth.
Effect of Red Light on Growth.—It has been pointed out that
red light stimulated growth in higher plants but, according to
Rahn, had no effect on lower organisms. Shrewsbury found
that when cultures of Willia were placed at a distance of 4 inches
from a Wratten safelight No. 1 screen, illuminated by a 60-watt
gas-filled lamp, with the temperature of the dark room ranging
from 23 to 28° C., growth was more rapid and more luxuriant
than in the dark or diffuse daylight, as expressed by a hastening
of spore formation. He attributed this increased activity to a
reflex of increased growth energy, with red light acting as a
stimulant. However, he adds that much the same growth was
obtained in cultures grown at 30° C. as in those grown at room
temperature, so that it was very likely that the heat and not
the red light was the activating agent.
To avoid this condition, the light-proof container as described
above (fig. 7) was again used. This time a red bulb of the same
intensity as before was used and a plate of red glass replaced the
colorless glass. The same medium of the same pH was used here
too. Cultures were grown at room temperature in the dark and
diffuse daylight as controls. After 27 days it was found that
1933]
MOORE—ENDOMYCES CAPSULATUS 523
those cultures under red light displayed no cultural or micro-
scopic differences from those grown in the dark and daylight, and
showed no change from those in white light and total darkness,
except for a small increase in diameter which may be ascribed
to the extra time of 3 days.
REACTION TO DYES
The róle of certain dyes in medical work has received much
critical and experimental attention for a number of years. Their
application for therapeutic purposes has resulted in clinical cures
in various cases and has had no effect in others. Particular dyes,
as methylene blue, gentian violet, and crystal violet, have been
used locally or intravenously for skin lesions in cases of blasto-
mycosis, coccidioidal granuloma, sporotrichosis, monilial infec-
tions, and other diseases both of mycological and bacteriological
etiology. Some of these have yielded favorable results as
mentioned, but the value of the dyes applied is empirical since
dye therapy rests on no definite scientific basis, according to
Spring (’29). Faber and Dickey (’25) treated 15 infants afflicted
with thrush, with local applications of a 1 per cent aqueous
solution of gentian violet and found that 50 per cent showed an
apparent cure in one day or less; 36 per cent in from 2 to 3 days;
and 14 per cent in from 4 to 5 days. In several cases, however,
the lesions returned at various intervals, and again some dis-
appeared after one application while others resisted for several
applications. In general, the lesions seemed to become smaller
after continuous applications and to disappear. As a result,
Faber and Dickey advocate a trial of the dye for prophylactic
measures and also for therapy. Churchman (’20d), working
on chemotherapy with gentian violet in vitro, advised careful
considerations of the amount of bacteria affected, before any
conclusion as to the therapeutic value of the dye be evaluated.
Sanderson and Smith (’27) postulated the possibility of the
utilization of gentian violet dye for the treatment of blastomy-
cosis.
It has been known and recorded since the days of Koch that
certain aniline dyes have a ''bacteriostatic" effect on the cultur-
ability of various bacteria. The viability of the organisms is
[Vor. 20
524 ANNALS OF THE MISSOURI BOTANICAL GARDEN
diminished to such an extent that they lose their power to multi-
ply. This selective action of the dyes is very remarkable,
certain organisms growing very abundantly and luxuriously on a
medium containing gentian violet in a rather high concentration,
whereas other organisms which are very resistant to harmful
factors do not grow at all. This high selectivity or specificity
was attributed not only to gentian violet, but also to others, as
erystal violet and various members of the tri-phenyl methane
series of dyes.
In early work on dyes, smears and broth cultures were used,
but it was found that the bacteria ceased growing or were killed
in the smears and that various concentrations of dye in the broth
had practically the same effect. Later work (Churchman, 712)
was performed on the divided agar plate to determine whether
the reaction would be the same. This method consists of in-
serting a metal strip (Halsted’s aneurysm metal) across the
diameter of the petri dish and then pouring the agar on one side
and allowing it to cool. The agar containing the dye is poured on
the other side ofthe strip. When theagar hardens, the metal strip is
lifted out with sterile forceps. Thus one half of the plate con-
tains plain agar and the other half the dye-containing agar. "This
method has been used by several investigators, including Sander-
son and Smith.
Churchman (’12,’ 20, '20a, '20b, ’20c, '20d, '20e, '21) found that
& parallelism with the reaction of the Gram stain could be made.
That is, considering as violet-positive those organisms which
were inhibited in growth by a certain concentration of a dye,
gentian violet for instance, and as violet negative those upon
whose growth the dye had no effect, then it was found that a great
number of Gram negative organisms were also violet negative.
This is a general rule, with a few outstanding organisms being
exceptions.
The same author also found that the selective action of the dye
may be expressed in one of two ways. The dye may be toxic
to the violet-positive organisms when a direct application is
made, or a very strong inhibitory action is expressed when it
is incorporated in the medium. On the other hand, the dye is
not toxie to violet-negative organisms, and further, no inhibitory
1933]
MOORE—ENDOMYCES CAPSULATUS 52b
action is expressed except in very strong concentrations when it is
incorporated in the medium. His experimental evidence pointed
to a dilution of 1 : 1,000,000 as capable of stopping growth, and
1: 2,000,000 as possibly retarding it.
This work was confirmed by Krumwiede and Pratt (13), who
used dahlia agar, and also they confirmed the belief that the
action was quantitative. They further observed that several
dyes, in addition to those already mentioned, had this specific
action. This work then called forth several applications of this
principle to culture media, as that of Petroff (15) which was used
by that author to isolate tubercle bacilli from the sputum and
feces, and that of Farley (’20) as a restrainer in the isolation of
pathogenic molds. Farley found that a dilution of 1:500,000
inhibited the growth of Gram positive bacteria.
Later work on this problem showed that there might exist
two types of organisms within a single strain, one which might
grow vigorously on gentian violet media, and the other not at all,
yet both showing the same stain and cultural characteristics.
Also, heavy and repeated inoculations of violet-positive organisms
would give fair and even good growth, and on this point Church-
man (’20e) made the following statement: “This would indicate
that bacteria do not, as is commonly supposed, act as isolated
individuals; they possess the power, in numbers, of accomplishing
effects which, alone, they are incapable of. The nature of this
community of action it is at present impossible to guess at.”
The same author (Churchman, ’21) placed dead bacterial bodies
between living bacteria and gentian violet media and found that
the Gram positive organisms grew, the dye having no effect. He
attributed this occurrence either to a filtration or a stimulation
of growth. The same year Churchman and Kahn (’21) found
that a number of cells could accomplish more than a single
cell, hence the communal action belief was further emphasized.
Work on dyes was not confined to bacteria alone, for Lewis (’30)
found that a certain number of dyes failed to inactivate the virus of
chicken tumor while many had a harmful effect in certain concen-
trations. Several papers have been published showing the action of
certain of these dyes on various fungi, but only in a qualitative way;
Greenbaum and Klauder (22) tested the action of gentian violet
[Vor. 20
526 ANNALS OF THE MISSOURI BOTANICAL GARDEN
as advocated by Farley (20a); Sanderson and Smith (27) found
that dilutions up to 1:500,000 inhibited E. dermatitidis in its
yeast-like form; Clark (727) showed that no growth occurred in a
yeast-like organism and its mycelial form in dilutions from
1: 100 to 1 : 25,000 and that it gradually increased from 1 : 50,000
to 1:1,000,000, which gave good growth. No quantitative
determinations of the colony growth were made. Stearn and
Stearn (29) found that several fungi, including E. dermatitidis,
were inhibited at various concentrations of gentian violet, with
the organism being affected at 1 : 500,000.
The later work of Churchman indicated that the ‘“bacterio-
static properties” of dyes ascribe to them the ability to inter-
fere with the reproductive mechanism, which he terms “genesis-
tasis," without killing or in any way interfering with their other
properties. This same phenomenon has been found bot
qualitatively and quantitatively to hold for E. capsulatus.
Since the author-could find that no particularly quantitative re-
sults had been obtained for ascomycetous fungi, at least for the
hyphal form, it was decided to investigate the problem of the dye re-
action. For this purpose, several dyes, picked at random, were
used. ‘These are as follows:
Dyes of the nitro, - "— oxyquinone groups.
Dyes of the azo
orange G acid (N TT Aniline and Chemieal Co.)
Sudan III—Nweakly acid—(E. Merck)
poids d = oxyquinone group:
aliz —acid—(E. Merck)
The quinone-imide dyes.
The thiazins:
mn blue—basic— (Coleman and Bell Co.)
The a
Kaioak or eurhodin
hakai — Psi dias and Bell Co.)
Safra
aie A—basic— (Coleman and Bell Co.)
The phenyl-methane dyes.
Tri-phenyl methane spurte
Di-amino tri-phenyl methan
light green S. F. Ee A E and Bell Co.)
Tri-amino tri-phenyl methanes (rosanilins);
basic fuchsin—basic— (Coleman and Bell Co.)
crystal violet—basic— (Coleman and Bell Co.)
aniline blue—acid—(National Aniline and Chemical Co.)
1933]
MOORE—ENDOMYCES CAPSULATUS 527
The xanthene dyes.
Fluorane derivatives:
eosine B—acid—(Coleman and Bell Co.)
Phenolphthalein and the sulphonthaleins:
phenol red or phenol-sulphonphthalein—acid—(Dr. 'T. Schuchardt, G. m.
b. H. Chemische Fabrik)
The natural dyes
Brazilin and haematoxylin:
haematoxylin (Coleman and Bell Co.)
These dyes were incorporated in the media, nutrient agar
(produet of Digestive Ferments Co.) at a pH of 6.8, in the
following concentrations: .00001; .000025; .00005; .000075;
.0001; .00025; .0005. Several plates were used for each concen-
tration of the dye, but only three plates for each are recorded
in the tables for reasons stated previously. These plates were
inoculated in the same manner as were those in the temperature
and pH experiments, a broth of pH 6.8 being used here. The
plates were kept at room temperature, which showed slight
changes from day to day but not enough to affect the results
seriously. The diameters of the colonies were measured daily at
approximately the same hour each day. Inasmuch as the
colonies may show slight irregularities in peripheral growth,
these measurements may not be considered absolutely correct.
However, if the shape of the colony was irregular, several diam-
eters were measured and the mean taken as representative of
that colony. Where contaminations occurred, as has been
explained previously, the plates were discarded.
It is to be noted that these experiments were carried out on
several different occasions and that the figures in the tables
denote the mean of a representative series. Since preliminary
experiments with these dyes had given some seemingly strange
results, microscopic examinations were made daily, in the case of
the crystal violet and methylene blue series, and then, some time
after growth had started, in the case of eosine B.
Several of the dyes produced no effect on the growth of the
organism. These dyes were orange G, Sudan III, alizarin
(table 1v), safranine A, light green, basic fuchsin, aniline blue,
phenol red, and haematoxylin. Neutral red was also used, but,
as was expected, the results showed no difference from the normal
control, the same being true for the other dyes.
[Vor. 20
528 ANNALS OF THE MISSOURI BOTANICAL GARDEN
If normal conditions are maintained for the plates, that is, if
there is no drying out and no change in temperature the colonies
grow at a rate of speed which is fairly constant as long as there is
sufficient nutrient material for the organism. However, when
the plates are kept in an atmosphere which allows drying out of
the agar and when the amount of nutrient material is relatively
small for the particular organism, the colony will show a decrease
Fig. 8. Mycelium on .00025 per cent crystal violet agar.
in growth rate. Thus, in the former case, the growth curve
would be a straight line for a considerable portion, depending of
course on the rapidity of growth of the organism compared from
day to day, which in this case was sufficient to cover the plate
almost completely before drying of the substrate. An initial lag
on artificial substrates, as agar, must be taken into consideration.
In that case the curve would tend to be sigmoid, that is, there
would be an initial lag due to an acclimatization of the organism
to the medium, a portion of increased activity which would be a
sharp rise, and then a portion of decreased activity due to factors
mentioned previously.
The data of the growth on the several non-affecting dyes
529
.0005
0.4
«o
oo
N
=)
xn
i
er
5 2-
. 00025
. 0001
6
6.7
.000075
5
6.6
TABLE IV
Percentage concentration of dye
.00005
. 000025
MOORE—ENDOMYCES CAPSULATUS
TEMPERATURE OF APPROXIMATELY 22? C.
VARIOUS CONCENTRATIONS OF ALIZARIN, AT A
Nutrient agar (product of Digestive Ferments Co.) pH 6.8
.00001
6
6.6
MEAN DIAMETERS IN CENTIMETERS OF COLONIES GROWN IN
tion
1933]
inocula-
[Vor. 20
530 ANNALS OF THE MISSOURI BOTANICAL GARDEN
d
E E
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EA
showed that the rate was practically the same in each case, and if
the diameters of the colonies in em. were plotted against the time
in days, for each concentration of the dye, one would find this to
be the case. This is illustrated in fig. 9, the growth rate for
1933]
MOORE—ENDOMYCES CAPSULATUS 531
alizarin, results representative of these dyes. The differences in
the curves are within the range of error, so that in general we may
.9005
000075 +0001
Percentage concentration of dye
Oa
.00005
-00001
Fig. 10. Maximum growth at constant temperature and same pH on nutrient agar with various concentrations
+ ™
*uo uj sejuo[oo Jo StozPouetd
say that the growth rate is the same. Except for the initial lag
and a slight curve due perhaps to utilization of nutrient materials
of alizarin.
[Vor. 20
532 ANNALS OF THE MISSOURI BOTANICAL GARDEN
or drying of the agar, the line is about straight. When the
maximum points of growth for each concentration is plotted,
that is, diameter of colony against percentage concentration of
dye (fig. 10), it is found that a straight line results, with perhaps
some excusable error. In other words, growth is identical for
all the concentrations of dye, at least in the observations made
here.
In dealing with the other dyes, as crystal violet (table v),
methylene blue (table vr), and eosine B (table viz), a striking
difference in growth was observed. The cultures of crystal
violet grown under the same conditions as those above presented
a growth which was proportional, within a range of limits, to
the concentration of the dye. Like results were found to be true
for methylene blue, but not quite so well marked.
Colony measurements were made daily, and, as may be seen
in table v, the growth was inhibited in the higher concentrations
to the point of complete cessation. Along with the measure-
ments, microscopic examinations were also made, and it was
found that on the media of higher concentration of dye there was
a corresponding decrease in production of reproductive parts.
In other words, the number of asci with the ascospores gradually
diminished on the increasing concentration of dye media as
compared with the normal growth. A well-marked feature was
the development of thick walls in the hyphae, as may be seen in
fig. 8 which illustrates variously thick-walled swellings, chlamydo-
spores and hyphae, taken from a culture of .00025 concentration,
forty days after inoculation. The cellular material stained
heavily, and showed granular cytoplasm and a reduction in the
number of nuclear divisions. An increase in the number of
swellings, cellular contents, and thickness of walls, as well as the
number of chlamydospores and terminal hypnospores, is a
condition which is usually associated with unfavorable phenom-
ena. Similar characteristics were present approximately three
months later. Macroscopically, the colonies on the higher
concentration of crystal violet showed a growth upwards on the
inoculum and not until 15 days after inoculation did any growth
appear on the agar and then very little. In the case of methylene
blue, however, growth was flat and took on a blue color very
soon after inoculation.
1933]
533
MOORE—ENDOMYCES CAPSULATUS
TABLE V
MEAN DIAMETERS IN CENTIMETERS OF COLONIES GROWN IN
VARIOUS CONCENTRATIONS OF CRYSTAL VIOLET, AT A
TEMPERATURE OF APPROXIMATELY 22? C.
Nutrient agar (product of Digestive Ferments Co.) pH 6.8
Percentage concentration of dye
. 0005
eo et
ES eo OD c» co e$ OD OD ~H i
A M
)w vir Rise id . 1*5 00o0o0o0o0ooooooooo
e Co oo oc O OOO O O OQO
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2 e eoo occ OOOO oo oo © @
ha
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Te]
S eo C» c9 =H oo © ort by 00 OQ»: OQ» OQ» O Q r4 O1 «t i Or-AAOAOCr Oo «f
2 D D O © DOO OOOO SOC CH HH HHH S aa aa an aa EN EN | NN
a
S © OOO O m m - —- N AN ANNMAN CÓ CÓ SH H H BO H oH 10 10
—
e Co x» rr NNOD - 09150 «D O0 OC» r4 C$ «M (D b. C» OQ Q HAW A n <+ «o
o OOO m m on NN NAN OD C9 OD Hi oo © c coc - e cm om om ST 00000
[e] .
eo
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S S ooo c e — — tien! — ON C OD cO cO (CO H H oH
5 e
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=
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‘wo UT so]uoioo jo sLelewetd =
author for eosine-methylene-blue agar where the mycelium had
taken on a pink to blue color and ceased growing. ‘These cultures
when transplanted to fresh agar grew just as luxuriantly as
[Vor. 20
538 ANNALS OF THE MISSOURI BOTANICAL GARDEN
though they had never been affected. What may have happened
is that the dye, being acid, may have been absorbed by the fungus,
.o00075 +0001 -90025 .0005
Percentage concentration of dye
.00005
.000025
-00001
1 Y) N
*uo ug Sojuo[oo Jo 3434əƏwLe ad
Fig. 12. Maximum growth at constant temperature and same pH on nutrient agar with various concentrations o
crystal violet.
as evidenced by the color of the mycelium, and was strong enough
to counteract the metabolic activities of the cell. It would seem
1933]
ion
MOORE—ENDOMYCES CAPSULATUS
ouo[4q3our JO suorjurjuoouoo SNOBA qjr« 1e29 juotrjnu uo Hd eures pus o1njwroduro] 31uvjsuoo 49 q14013 jo NLY “ET “BI
"eni
— 8z əz tz zZz o2 SI 9l Yi rad or Q 9 * zZ
$0000°
$20900*
TO De wwe SR e
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+ rz
‘wo UT Se[uo[oo Jo sileQeuvIT
O
Ko
from a study of the pH of the medium, which changes from a
slightly acid to alkaline in normal growth, that this could be
detected. However, the medium was already covered with
[Vor. 20
540 ANNALS OF THE MISSOURI BOTANICAL GARDEN
«000075 00025
Percentage concentration of dye
000025
uw t re) N
*"u0 UT Ssojuoioo Jo S4ə}əwvyd
Fig. 14. Maximum growth at constant temperature and same pH on nutrient agar with various concentrations of
sufficient growth to cause an alkaline condition, so that no direct
evidence could be obtained from that direction. There are no
definite facts to uphold this belief, and the statement should be
considered more as a conjecture for the present. "There have
methylene blue.
1933]
c
NN
E
MOORE—ENDOMYCES CAPSULATUS
Time in days
is "n M N
*wo ug Sojuoloo jo s4Iəwerd
been many theories proposed, but none have had any final
acceptance.
The action of eosine B on the organism is illustrated in figs.
15-16. The former shows the rate of growth of the cultures on
Fig. 15. Rate of growth at constant temperature and same pH on nutrient agar with various concentrations of
eosine B.
[Vor. 20
542 ANNALS OF THE MISSOURI BOTANICAL GARDEN
a
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various concentrations of dye. Growth on the concentration
.0005 was inhibited on the nineteenth day, and that on the con-
centration .00025, on the thirty-fourth day. Fig. 16 represents
the maximal points of growth corrected for the inoculum. "The
inhibition of growth is clearly distinguishable here.
eosine B.
1933]
MOORE—ENDOMYCES CAPSULATUS 543
A further phenomenon is noted here, not because it was abso-
lutely specific for this experiment but because it had appeared
most frequently on crystal violet media of concentrations .0001
and .00025. This is the formation of crystals in the media,
appearing to emanate from the growing zone or peripheral
region of the colony and found in conditions which have not been
very favorable towards growth. They have occurred on beef
extract agar of pH 7.2, 7.0, and 6.8. No crystals of this sort
were found on media where there was no growth or where growth
was very luxuriant. They appear to resemble very closely
those of ammonium magnesium phosphate (NH,MgPQ,.6 H;O)
as was reported also by Scudder (26). They were insoluble in
cold water, completely and immediately soluble in hydrochloric
acid, and were reprecipitated with sodium hydroxide, tests which
are applied to crystals of the above composition. They were
apparently the result of fungal activity, and Scudder has asso-
ciated this condition with bacteria which “do not ferment the
ordinary carbohydrates and intensify the alkalinity of the
medium rapidly," a fact which has been clearly demonstrated in
the carbohydrate reactions carried out in this paper.
ANIMAL ÍNOCULATIONS
From time to time papers appear in the literature pointing out
that some particular organism which had been kept in culture over
a long period and then inoculated in experimental animals, had
retained its pathogenic properties. On the other hand, it is quite
well known that this property may be lost on subculturing. It
is also known that by growth on certain media at a specific
temperature, toxicity may be retained over a long time. It has
also been found that certain microorganisms may be made to
regain lost toxie powers by repeated inoculations.
In order to determine whether E. capsulatus had undergone any
such changes since it was isolated in September, 1928, and kept
in culture at different temperatures, on different media, and at
various hydrogen-ion concentrations, several animals were
inoculated with a twelve-day-old suspension of the organism in
10-ce. physiological saline solution. The culture was stirred up
thoroughly with a sterile needle to eliminate the clumps.
[Vor. 20
544 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The animals and inoculations were as follows:
Rabbits: Two females—intracerebral (0.2 ec.) and intravenous
(0.5 cc.). Two males—subcutaneous (2 ec.) and intratesticular
(1 cc.).
Guinea-pigs: Two, one male and one female—intracerebral
(0.2 cc.). Two males—subcutaneous (1.5 ec.) and intratestic-
ular (1 co.).
Mice: Two, one male and one female—intracerebral (0.2 co.).
One male—subcutaneous (1 cc.) and intratesticular (0.5 cc.).
The results of these inoculations were that one of the rabbits
receiving à subcutaneous and intratesticular inoculation died
twenty-three days later. Autopsy revealed no infection or
lesion, except an intestinal obstruction. "Three mice, healthy,
with no loss of weight or appetite, were killed fifty days after
inoculation. The organs were completely negative. Cultures of
the heart’s blood also negative. The rabbits and guinea-pig were
healthy, with no loss of weight, three months after inoculation.
The remaining mouse, with an intracerebral injection, died
sixty-two days after inoculation as a result of pneumonia. Au-
topsy showed no complication of the other organs. Cultures of
infected lung tissue and brain negative for E. capsulatus, the
lung in a hemorrhagic condition.
It may be concluded, therefore, that the organism had lost its
power to infect after four years’ growth on artificial substrates.
That it failed to produce a toxic action in an animal may be
attributed to the changed environment and habitat of the fungus.
It has been pointed out previously that the optimum temperature
has been reduced from 37.5° C. to approximately 25° C. In
addition, experimental data has shown that growth at the former
temperature was very slow and slight, with complete cessation
and death after a short time. It is very possible that the organ-
ism was killed here, as in the in vitro experiments, as a result of
the body temperature of the experimental animals. fructose> glucose >
sucrose. A year later, Quednow (30) published his observations
on a more extended list of sugars. He found the order of excel-
lence to be glucose > fructose > sucrose > maltose > mannite >
galactose > lactose. Smith (’32) used sucrose, glucose, and
maltose, singly and in all sorts of combinations, and observed no
apparent difference in the growth of the seedlings.
Unusual facilities for research have enabled the writer to extend
the list of sugars which has been used for orchid germination to
include the rarer and more expensive forms. ‘This opportunity,
together with the discrepancies between the work of La Garde,
Quednow, and Smith, led to the present work. ‘The sugars were
added to three different complete mineral nutrient solutions in
amounts to give 7 gms. of carbon per liter. The compositions of
these solutions were as follows:
[Vor. 20
572 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Knudson's (722) Solution Shive's (15) Solution La Garde's ('29) Solution
Mg80,.7H;0 .250 gm. MgSO,.7H;O 4.930 gm. MgSO,7H;O 1.00 gm.
Ca(NOs3)2.4H20 1.000 gm. Ca(NOj,4H;O 1.228 gm. Ca(NOj,4H;0 1.00 gm.
(NH480, .500 gm. KH;PO, 1.960 gm. KH;PO, 1.00 gm.
K;HPO, .250 gm. CaCl, 1.00 m
NH4NO;
.50
(NH4)CO;. H:O .500 n
Iron was added in all cases as 10 cc. of à M/200 suspension of
FePO, prepared according to Livingston (19) in a liter of nutrient
solution, Both Knudson and La Garde added iron in such
quantities as to cause a heavy precipitate of iron phosphate.
La Garde states that this precipitate was filtered off before the
final sterilization and contained, besides iron and phosphate ions,
calcium and potassium. It seems unwise to cause this bulky
precipitate because it tends to adsorb other ions which are
removed with it in filtration. The quantity of FePO, added by
the author does not cause appreciable precipitation at the hydro-
gen-ion concentration used. This reduced amount is undoubtedly
sufficient in quantity, since it is ten times that originally rec-
ommended by Livingston. La Garde designates the iron com-
pound used by him as Fe;(PO4),.8H;,O. Knudson (722), Qued-
now (730), and Smith (732) also added iron as ferrous phosphate.
Since it is well known that the ferrous ion in the presence of
oxygen reduces nitrate to nitrite, the ferric ion was used in the
present work.
The solutions were made up in liter flasks, and adjusted to
pH 4.00 with HCl. All precipitate dissolved at this acidity, but
there was a slight opalescence due to ferric phosphate. The
solutions were then titrated by means of the quinhydrone elec-
trode to such pH (see tables) that the values after sterilization
were 4.8-5.1. Aliquots of 100-ec. portions were placed in 200-cc.
Erlenmeyer flasks and 1.75 grams of Merck's Reagent Powdered
Agar added. Sterilization was by autoclaving at twenty pounds
pressure for twenty minutes. The medium was allowed to
solidify in a slanting position.
It is extraordinarily difficult to maintain sterile cultures in
warm moist atmospheres over long periods of time, and after
many preliminary failures, the following plan was adopted. "The
solutions were added to the culture flasks through a funnel, care
1933]
WYND—CARBOHYDRATE FOR ORCHID SEEDLINGS 573
being taken not to moisten the necks of the flasks. The flasks
were then closed with cotton plugs. A duplicate set of cotton
stoppers was carefully and tightly rolled, sterilized in empty
flasks in the autoclave, and then immediately transferred to the
dry-air oven until thoroughly dry. The seeds were shaken
vigorously for thirty minutes in a small vial of calcium hypo-
chlorite prepared as recommended by Wilson (15). "This vial was
then clamped in a sloping position so that contaminating sub-
stances might not fall in from the air. A culture flask and an
empty flask containing the especially prepared cotton plug were
held in a horizontal position in the left hand. The temporary
cotton plug was withdrawn from the culture flask and dropped.
A platinum-loop inoculating needle, held in the right hand,
was quickly flamed and a loopful of seeds transferred directly
from the hypochlorite solution to the drop of moisture that al-
ways exudes from the solidified agar. The neck of the flask was
then flamed and the especially prepared stopper quickly drawn
from the blank flask and inserted in the culture flask. This
procedure is advisable as it involves a minimum of exposure of
the cotton plug that is finally used in the culture flask, and
insures its perfect dryness.
The drop of moisture containing the seeds on the edge of the
agar was then distributed around the entire margin by rotating
the flask carefully. This even distribution of seeds was main-
tained by placing the flask in a rack so constructed that the agar
surface was perfectly level, thus preventing the liquid drop from
draining to one side and carrying the seeds with it. After a
convenient number of flasks had been inoculated, the necks of
the flasks were again flamed and the cotton plug well charred on
the surface. The plugs and the outer surface of the necks were
then moistened with saturated HgCl, solution. Heavy waxed
paper was then dipped in the HgCl, solution, tightly wrapped
around the stoppers and the upper part of the flask, and held
firmly in position by rubber bands. The writer has found that
unless these precautions are taken, fungi will frequently grow
along the surface of the flask and penetrate tbe stopper, con-
tamination usually not appearing until three or four months after
inoculation. Bernard, certainly a well-trained and experienced
[Vor. 20
574 ANNALS OF THE MISSOURI BOTANICAL GARDEN
mycologist, has commented on the extraordinary difficulty of
maintaining orchid cultures sterile in the moist warm atmosphere
desirable for germination. By taking the above precautions,
the writer has maintained sterile cultures as long as three years
in moisture-saturated atmosphere at 25-35? C.
In every case, the cultures were prepared in triplicate. "The
seeds for the entire series were from a single pod of Cattleya
Trianae Linden & Rchb. f. The flower was pollinated November
TABLE I
DATA OF KNUDSON'S SOLUTION
pH adjusted pH at time of | pH after support-
Sugar before steri- planting ing growth
lization 8 months
d-glucose.............. 4.12 5.1 4.2
d-fructose............. 4.12 4.9 3.8
d-galactose............ 4.12 5.0 No growth
d-mannose............. 4.13 5.0 4.2
l-xylose. i ois dao 4.15 5.0 No growth
l-arabinose . 4.10 5.0 No growth
Maltose............... 4.15 5.1 4.4
l-rhamnose 4.15 5.0 No growth
oo LL io intr 4.15 5.1 4.4
BD. 665.666 64048445 4.12 4.8 4.4
TABLE II
DATA OF SHIVE’S SOLUTION
pH adjusted pH at time of | pH after support-
Sugar before steri- planting ing growth
ization 8 months
d-glucose. ............. 4.12 5.1 4.4
d-fructose............. 4.12 4.9 4.0
d-galactose............ 4.12 5.0 No growth
d-mannose 4.13 5.0 4.5
Ixylose. .............. 4.15 5.0 No growth
l-arabinose............ 4.10 5.0 No growth
Maltose............... 4.15 5.1 4.6
]-rhamnose 4.15 5.0 No growth
NONE Lco ed dex, 4.15 5.1 4.5
Raffinose............-. 4.12 4.8 4.4
1933]
WYND—CARBOHYDRATE FOR ORCHID SEEDLINGS 575
TABLE III
DATA OF LA GARDE'S SOLUTION
pH adjusted pH at time of | pH after support-
Sugar before steri- planting ing growth
lization 8 months
OAEPUCON. Lco oa Lese 4.25 4.9 3.5
d-fructose............. 4.24 4.8 3.8
d-galactose............ 4.22 4.8 No growth
d-miBnnose. er 4.24 4.9 3.8
LX WIGNN oco dor dai purs 4.24 4.9 No growth
l-arabinose............ 4.24 4.9 No growth
alto8e.. Ie eean tn eas 4.25 4.9 4.2
l-rhamnose. ........... 4.22 4.9 No growth
IUCTOSO avi oe ees 4.24 4.9 4.1
Rafünose e o e: 4.51 4.9 4.2
27, 1930, and the matured pod harvested March 11, 1932, after
a developmental period of over sixteen months. The inoculation
of the culture media was made June 15, 1932, and the observa-
tions were recorded February 25, 1933, after a growth period of
about eight months.
In the younger stages of development the diameter of the
protocorm is an accurate basis for comparative determinations
of growth, but after the seedling has developed leaves the growth
is largely vertical rather than mere enlargement of the nearly
round protocorm. Accurate measurements of the height of the
young plant are difficult to obtain, since it is too large to measure
by a microscope micrometer and too small for any less accurate
means. For these reasons, seedlings as old as eight months can
best be rated with the eye by comparing different culture flasks
and sorting them into a few groups. The results of such a
comparison after the growth period of eight months are shown in
table rv. The relative excellence of the cultures is designated
by the number of X's.
'The results show that d-mannose produced definitely the best
growth. This is followed by the group glucose-maltose-fructose,
and this in turn by the third group sucrose-raffinose, and then
l-xylose. No growth was obtained on d-galactose, arabinose, or
rhamnose. A comparison of these results with the molecular
[Vor. 20
576 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE IV
GROWTH DATA
,
Kn
Description Mineral Solution Mineral Solution Mineral Solution
of plants
Sugar |Growth| Sugar | Growth| Sugar | Growth
Group I
Exceptionally
good growth — |d-mannose| xxxx* |d-mannose| xxxxx |d-mannose | xxxxx
and chlorophyll
development
roup II
Moderately d-glucose | xxx d-glucose | xxx
good growth ^ |maltose XXX maltose XXX maltose XXX
and chlorophyll |d-fructose | xxx d-fructose | xxx d-fructose | xxx
development sucrose XXX
Group III
Poor growth ^ |sucrose XX sucrose XX
an
chlorophyll raffinose XX raffinose XX raffinose XX
development
l-xylose x l-xylose x
Group IV d-galactose| 0 d-galactose| 0 d-galactose| 0
No growth l-arabinose | 0 l-arabinose | 0 l-arabinose | 0
l-rhamnose | 0 l-rhamnose | 0 l-rhamnose | 0
*The number of x's denotes the relative excellence of the cultures.
configuration of the respective sugars indicates that in but one
physical characteristic do they show any consistent physiological
action—and that is that the pentoses do not allow the germination
and growth of orchid seedlings. It is not impossible that their
quality of being levo-rotatory is related to their physiological
reaction. ‘The two instances in which l-xylose allowed growth
are of doubtful authenticity. Only two or three seeds germinated
in each flask, and these never developed beyond a very rudi-
mentary protocorm. Such rudimentary development was oc-
casionally noted even on sugar-free agar cultures, but in those
cases also development never proceeded beyond a rudimentary
protocorm. It is interesting to note that rhamnose, although it
1933]
WYND— CARBOHYDRATE FOR ORCHID SEEDLINGS 577
has six carbon atoms, is structurally a methylated pentose and
reacts physiologically to orchid seedlings as a pentose. Galac-
tose, although an aldo-hexose, as is also d-mannose, supported
no growth. These results are in approximate agreement with
those of Quednow, cited above.
From the following considerations the author believes that
the conspicuous superiority of d-mannose is of especial signifi-
cance. Mannose, in the form of mannan, is known to be widely
distributed as a constituent of the cell wall of many plants. This
is particularly true of seeds (Onslow, ’23). For example, various
complex mannans have been found in the seeds of palms, aspara-
gus, clover, coffee bean, onion, and various Leguminosae, Coni-
ferae, and Umbelliferae. Mucilages are particularly rich in
mannans, as, for instance, those obtained from lily bulbs (Parkin,
701) and tubers of various genera of Orchidaceae. Pringsheim
and his coworkers (’24, '28) succeeded in isolating and studying
mannan from orchid tubers. In this instance, it was water
soluble and was precipitated by alcohol as a white powder.
Klein (’32) gives a procedure for isolating mannan from salep
itself.
Salep, a product of orchid tubers, has been used as a substrate
for the orchid fungi and for germinating orchid seeds in the
presence of some symbiotic fungus from the earliest days of
orchid research. It therefore appears fairly certain that the
fungal element of the orchid mycorrhiza is able to hydrolyze
mannan to soluble mannose. In this way the symbiotic fungi
could make available any mannose which might be present in
the woody and mossy substrate of epiphytic orchids in their
natural habitat. The extraordinary slowness of the develop-
ment of orchid seedlings would be supported by the presence of
only very small quantities of mannan.
The Missouri Botanical Garden has produced seedlings by the
symbiotic method that surpassed in quality anything that the
author has seen obtained by asymbiotic methods. In many
instances the agar substrate consisted of the usual mineral
substances and finely shredded cocoanut fiber. An appropriate
fungus was inoculated on this medium some time before the seeds
were sown. Seedlings grown on fungus-inoculated cocoanut-
[Vor. 20
578 ANNALS OF THE MISSOURI BOTANICAL GARDEN
fiber substrate always surpassed those grown on La Garde’s
maltose media. Bultel (25) also found that symbiotic cultures
gave superior results with all genera tested by him except Pha-
laenopsis. The nature of his substrate was not indicated.
The writer has already reported (33) on the superior value
of the La Garde mineral solution over other solutions, and it
seems scarcely probable that symbiotic cultures could owe their
superiority to any inorganic constituent. It is much more
probable that the carbohydrate relationship is the important
feature. Knudson has shown the effectiveness of orchid fungi in
hydrolyzing starch to available sugar. Might not the superiority
of the symbiotic cultures be due to the mannose produced by
hydrolysis from the cocoanut fiber?
In order to determine the actual presence of mannans in
cocoanut fiber, an analysis was carried out according to the
method of Haegglund and Klingstedt (’24, '27). Ten gms. of
cocoanut fiber were allowed to stand in 150 ce. of 72 per cent
H.SO, for 244 days. The mixture was then diluted with water
and the acid neutralized with CaCO;. The precipitate was
filtered off, the residue washed on a suction filter, and the com-
bined filtrate and washings were evaporated on the steam bath
to 150 cc. Then HSO, was added to give a 2 per cent solution
and the mixture boiled 2 hours. The acid was again neutralized
with CaCO;, then weakly acidified with HC,H;O., and evaporated
to. 100 ec. After cooling, 10 cc. of phenylhydrazine plus 20 ce.
water were added and allowed to stand several days. A very
definite precipitate of the insoluble phenylhydrazone was obtained,
indicating the presence of mannose.
The above method is scarcely quantitative because of the
difficulty of obtaining complete hydrolysis of mannans without
causing their oxidation at the same time. The very heavy
precipitate of CaSO, which is produced when the acid is neutral-
ized with CaCO; is bulky and difficult to wash thoroughly.
The seedlings grown on La Garde’s maltose solution equalled
those of the same age originally obtained by La Garde. Hence
the mannose cultures might be regarded as distinctly superior
to those yet obtained by the use of a purely synthetic medium
and approach in quality the best of the symbiotic cultures. It
1933]
WYND—CARBOHYDRATE FOR ORCHID SEEDLINGS 579
is to be regretted, however, that this could not be verified posi-
tively by comparison, since seedlings of the same age grown
symbiotically were not at hand.
The glucose cultures were also of high quality, particularly
as to their green color. No trace of yellowness or of inferior
chlorophyll development was observed in seedlings grown on this
sugar.
It must be noted that superiority of symbiotic cultures may
depend not only on a carbohydrate relation, but also on a favor-
able degree of acidity. Knudson has pointed out that satis-
factory fungal symbionts maintain a favorable pH of the media
for the germination of the seeds. The pH of asymbiotic media
can, of course, be adjusted artificially, but the initial favorable
acidity is difficult to maintain over long periods of time.
The author hopes that some worker equipped to carry both
symbiotic and asymbiotic cultures simultaneously will test
further the suggestion that symbiotic cultures owe their superi-
ority to mannose and to the constantly favorable pH relation-
ship.
SUMMARY
1. The growth of orchid seedlings over a period of eight
months was observed on a series of sugars, each added to three
different inorganic media in amounts to give seven grams of
carbon per liter. The order of excellence of growth on the
different sugars was: d-mannose > d-glucose > maltose > d-
fructose > sucrose > raffinose.
2. No growth was obtained on d-galactose, and the pentoses,
larabinose, l-rhamnose, and l-xylose. This inability of the
pentose sugars to support growth may be related to their levo-
rotating property. Galactose occupies an anomalous position.
3. Mannose gave conspicuously the better growth. This may
be related to the fact that symbiotic cultures containing cocoanut
fiber, an effective source of mannan, produce seedlings superior
to any that the author has seen produced asymbiotically.
4. The mineral nutrient medium of La Garde plus d-mannose
is regarded as the best asymbiotic culture medium for orchids, as
shown by the reaction of Cattleya Trianae Linden and Rchb. f.
seedlings.
[Vor. 20
580 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ACKNOWLEDGMENTS
The author wishes to thank Dr. E. S. Reynolds, Physiologist
to the Missouri Botanical Garden, for helpful suggestions through-
out the work, and Dr. G. T. Moore, Director of the Missouri
Botanical Garden, for the supply of orchid seeds.
BIBLIOGRAPHY
Ballion, G., and Ballion, M. (24). The non-symbiotie germination of orchid seed
in Belgium. Orchid Rev. 32: 305-309. 24.
——,——— —,(328). Asymbiotie germination of orchid seed. Ibid. 36: 103-
2. 1928.
Bernard, N. (00). Sur quelques germinations difficiles. Rev. Gén. Bot. 12: 108-
120. 1900.
—— — ——, (02). Études sur la tubérisation. Ibid. 14: 5-25, 58-71. 1902.
— — ——, (08). La germination des orchidées. Comp. Rend. Acad. Sci. Paris
137: 483- 485. 1903.
— — ——, (04). Recherches expérimentales sur les orchidées. Rev. Gén. Bot.
16: 405-451, 458-476. 1904.
'00). Symbiosis d’orchidées et de eia champignons endophytes.
Com ips Rend. Acad. Sci. Paris 142: 52-54. 1906.
Bultel, G. (24-25). Germinations aseptiques Fac Cultures symbiotique
et asymbiotique. Rev. Hort. Paris 96: 208-271. 1924 ; 97: 318-321, 359-363.
1925.
— — ——,(26). Lesorchidées germé hampig t des plantes normales.
Ibid., 98:155. 1926.
Burgeff, H. ('09). Die Wurzelpilze der Orchideen, ihre Kultur und ihr Leben in der
Pflanze. Jena, 1
— E. (24a). Germination of Mm and other seed without fungal
Orchid Rev. 32: 233-238. 192
— — ——, (24b). The non-symbiotic ue of orchid seeds. bid. 359-365.
— — ——, (26). The non-symbiotic and symbiotic germination of orchid seeds.
Ibid. 34: 165-169. 1926.
— ———, (29). Non-symbiotic and symbiotic germination of orchid seed. Ibid.
37: 68-75. 1929.
— ————, (32). Raising orchid seedlings. Ibid. 40: 195-206.
Costantin, J. ('26). La vie asymbiotique des orchidées. Ann. ‘ai “Nat. Bot. 8:
TK = H. (55). Recherches sur les tubercules de l'Himantoglossum hircinum.
id. IV. 3: 253-291. 55.
— — ——, (56). Dela germination des ophrydées et de la nature de leurs tubercules.
Ibid. IV. 5: 163-186. 1856.
Haegglund, E., und Klingstedt, F. W. (24). Cellulose-Chemie 5: 58. 1924.
————-, ——, (27). Zur Charakterisierung von Celluloseprüparaten mittels
der Pri eina Ann. der Chem. 459: 26-38. 27.
Klein, G Ar ). Organische Stoffe. Membranstoffe. Handb. d. Pflanzenanalyse
21: 48. 1932.
1933]
WYND—CARBOHYDRATE FOR ORCHID SEEDLINGS 581
Knudson, L. (16). Influence of certain carbohydrates on green plants. Cornell
Agr. Exo. Sta. Mem. 9: 1-75.
Non symbiotic suininétion of orchid seeds. Bot. Gaz. 73: 1-25.
a~ Ta
Y NI
MU Mee
— o . Further observations on non-symbiotie germination of orchid
seeds. Ibid. 77: 212-220. 1924.
, 25). Physiological study of the symbiotic germination of orchid seeds.
, (26). Pissiclogisul investigations on orchid seed germination. Inter-
nat. Cong. Pl. Sci. Ithaca, Proc. 1183-1189. ;
—, (27). Symbiosis and asymbiosis relative to orchids. New Phytol. 26:
328-336. 1927.
La Garde, R. V. (29). Pi Rod germination of orchids. Ann. Mo. Bot.
Gard. 16: 499-514.
Livingston, B. E. (19). A a for cooperative research on the salt requirements of
representative agricultural plants. 1-54. 2nd ed. Baltimore, 19
Onslow, Muriel Wheldale (’23). Practical plant biochemistry. Camibridge, 1923.
Parkin, J. (01). On a reserve carbohydrate which produces mannose, from the
bulb of Lilium. Cambridge Phil. Soc., Proc. 11: 139-142. 01.
Prillieux, E. (56). De la structure anatomique et du mode de végétation du Neottia
nidus avis. Ann. Sci. Nat. Bot. IV. 5: 267-282.
(60). Observations sur la germination du Miltonia spectabilis et de
mud: autres orchidées. bid. 13: 288-296. 1860.
, et Rivière, A. (56). Observations sur la germination et le développement
d'une orchidée. Ibid. 5: 119-136. 1856.
Pringsheim, H., und Genin, A. (24). Uber die fermentative Spaltung des Salep-
ns. VI. Mitteilung über Hemicellulosen. Zeitschr. f. physiol. Chem.
140: 299-304. 1924.
— — ——., und Liss, G. (27). Über das Salep-Mannan. Liebig’s Ann. der Chem.
460: 32-42. 1927
Quednow, K. G. (32). Beitrüge zur Frage der Aufnahme rd Kohlenstoffver-
bindungen und andere Pflanzen. Bot. Archiv. 30: 51-108. 1930.
Ramsbottom, J. (22). The germination of orchid seed. Orchid Rev. 30: 197-202.
1922.
Shive, J. W. (15). A three salt nutrient solution for plants. Am. Jour. Bot. 2:
157-160. 1915.
Smith, F. E. V. (82). i: Meer seedlings asymbiotically under tropical
conditions. Gard. Chron 932
ares W. (’86). Batas. zur v h dn Orchideenwurzelpilze. Bot. Zeit.
1886.
81-488, 497—505.
"d $ K. (15). Calcium hypochlorite as a seed sterilizer. Am. Jour. Bot. 2:
420—427. 1915.
Wynd, F. L. (33). Nutrient solutions for orchids. Ann. Mo. Bot. Gard. 20: 363-
372. 1933.
SUPER OPTIMAL AND THERMAL DEATH
TEMPERATURES OF THE COTTON PLANT AS
AFFECTED BY VARIATIONS IN RELATIVE HUMIDITY:
DOROTHY MEGOWEN BERKLEY
FormerlyJ essieR. Barr Fellow inthe HenryShaw School of Botany of Washington University
AND EARL E. BERKLEY
Research Fellow 4n the Henry Shaw School of Botany of Washington University?
HISTORICAL REVIEW
In 1863 Sachs reported the results of an attempt to determine
the effects of high temperatures on the sensitivity of Mimosa
pudica. 'Transitory insensitivity, he found, was caused by an
exposure of one hour to a temperature of 40° C., and at 45° C.
for a half hour and 49? C. for a very brief time the same effect
was produced. When permanent insensitivity was attained, at
higher temperatures, death invariably followed. Sachs also
reported (64) on the effects of high temperatures on tobacco,
pumpkin, corn, nasturtium, and rape, exposed for various
periods of time. All tbe plants were able to withstand tem-
peratures of 49-51? C., but none survived 51? C. for more than
10 minutes without injury. The power of resistance to high
temperatures was found to vary at different ages. Developing
leaves, stems, and roots were more easily killed than older ones.
Ewart ('03) noted a decrease in the rate of protoplasmic
streaming in Elodea, Tradescantia, Chara, Spirogyra, root hairs,
pollen tubes, etc., depending upon the height of the temper-
ature above the optimum and upon the length of exposure.
Pfeffer (03) made the generalization that all turgid plants
ultimately die when the temperature reaches from 1° to 2° C.
above the maximum where the plant will grow indefinitely,
although growth may continue for a time, and that at temper-
atures of 10? C. above this maximum all flowering plants seem
to be rapidly killed. He noted that plants, which at first
1 Portions of this work, together with certain microscopical studies of the treated
plants, were submitted by Mrs. Dorothy Megowen Berkley as a thesis in partial
fulfillment of the requirements for the degree of master of science in the Henry Shaw
School of Botany of Washington University.
? A fellowship established by the American Creosoting Co.
Ann. Mo. Bor. GARD., Vor. 20, 1933. (583)
[Vor. 20
584 ANNALS OF THE MISSOURI BOTANICAL GARDEN
appeared fresh and unharmed after a short exposure to fatally
high temperatures, frequently died later as an after-effect, even
under the best external conditions.
Fung ('11) emphasized the necessity of considering the relative
humidity in determining the effects of high temperature on
plants. He found the maximum temperature for growth of
cotton to be 113° F. in a relative humidity of 90 per cent and the
optimum 85-90^ F. in a relative humidity of 70.6—72.2 per cent.
Cotton plants treated for four hours in a saturated atmosphere
at a variety of high temperatures also gave interesting results.
At 42-45" C. the stems and leaves were badly wilted but re-
covered; at 44-48? C. the stems and leaves were browned but
new leaves appeared after one week (probably secondary growth
from uninjured nodes); at 49—55? C. the plants were killed. The
degree of injury was determined after the plants had been trans-
ferred to ‘‘proper conditions," which were not described. The
value of Fung's results is limited by the fact that he worked with
an insignificant number of plants and used only very young
seedlings. Bose (013), who gave 60? C. as the average fatal
temperature for plants in general, found the death point to be
lower in young plants, which confirmed the earlier statement of
Sachs.
Collander (24) determined the temperatures at which death
occurred in individual cells of various plants. T'radescantia
discolor was killed at 65? C. within an average of 1.8 minutes,
Brassica oleracea at 60? C. within an average of 0.8 minute, Beta
vulgaris at 60° C. in 0.7 minute, Draparnaldia glomerata at 55? C.
in 0.32 minute, and Pisum sativum at 55? C. in 0.095 minute.
He found that these plants could live at slightly lower temper-
atures for some time, thus demonstrating that the thermal death
point is suddenly reached. Lepeschkin (725) discussed the
effects of optimum, maximum, and thermal death temperatures
on bacteria, fungi, and higher plants. He stated that most
plants died at 60-70? C. in one minute, although some died at
40-45? C. in that time.
Gilbert (’26) found that cotton grew better at 80? F. in a
relative humidity of 50 per cent than it did in a relative humidity
of 85 per cent. Wallace (31) tested the effect of 1-24-hour
1933]
BERKLEY AND BERKLEY— TEMPERATURE AND HUMIDITY 585
exposures to temperatures ranging from 15 to 60? C. on the sensi-
tivity of Mimosa pudica. Above 45? C. injury or death resulted,
depending upon the length of the exposure. He reported that
the relative humidity had little or no effect upon the sensitivity
of the plant.
Baker (29), in studying the effects of excessively high tem-
peratures on conifer seedlings 1-3 months old, also emphasized
the suddenness at which the thermal death point was reached.
The living tissues were quickly killed at 54° C. but were un-
injured after à prolonged exposure to a temperature only a few
degrees lower. Just below the thermal death point he noted a
region of no growth where photosynthesis was apparently unable
to keep up with catabolic changes, the chlorophyll decomposing
faster than it was made and the leaves becoming yellow or
withered. On prolonged exposure the plants died. He also
found the age of the plant to be a factor in resistance, due to
the development of protective tissues as the plant grew older.
The indications were that there was no increased protoplasmic
resistance with age.
APPARATUS
Four glass cases of identical design and size (60 in. long by
32 in. wide by 40 in. high) were used for the experimental work.
Three of these cases were variously used for growing plants, for
germinating seeds, and for plants under observation following
treatment. Additional plants were grown on benches in the
experimental room.
The fourth case was converted into the electrically controlled
temperature chamber (fig. 1). The experimental plants were
placed in a flat of sand which rested on a lattice rack supported
12 inches above the bottom of the case. The heating units
consisted of three heaters, two of which were controlled by
ordinary switches, and were placed on the floor of the case, one in
either half about midway between the center line and the end.
They were composed of resistance wires wrapped on racks which
extended practically the width of the case. With these two
heaters there were three possible combinations: when the two
were in parallel maximum heat was obtained, when one was used
alone approximately one-half of the maximum could be had, and
[Vor. 20
586 ANNALS OF THE MISSOURI BOTANICAL GARDEN
when the two were in series one-fourth of the maximum. A
further reduction in temperature was obtained by turning off
these heaters altogether, leaving only the third heater which
was composed of resistance wire wrapped on the guard-wires of
an electric fan. This heater was controlled by a Thyratron! and
relay combination which in turn was controlled by a thermostat
with à mercury make-and-break contact.
EM uu
M ~ "4
Thyratron ms ec
7 " —
F Ea eee e
a ee |
l
Ay
relay | » s
' C
\ T
M L —
3 N toluol
/ \ mercury
H
VLA
Fig. 1. Diagrammatic cross-section through the center of the controlled tem-
perature chamber in which the plants were treated. C, capillary; E, electric fan
r
The fan, which was located on the floor at one end of the
chamber, was hooked directly to the current so that it ran
continuously, blowing the warm air over the heaters and through
! Behmitt, F. O., and Schmitt, O. H. A. A vacuum tube method of temperature
control. Science N. S. 73: 289-290. 1931.
1933]
BERKLEY AND BERKLEY—TEMPERATURE AND HUMIDITY 587
humidifying cloths made of cheese-cloth. In this way the heat
and moisture were uniformly distributed throughout the com-
partment. Good ventilation was assured by boring small holes
in the frame of the compartment on either side of and in back
of the fan.
The thermostat consisted of a glass tube extending the full
length of the chamber, one end of which was shaped into a U
terminating in a capillary. The main body of the tube had a
capacity of about 200 cc. and was filled with toluol; the U tube,
with a capacity of about 20 cc., contained mercury. The
contact was made in the capillary in the usual manner. Due
to the large capacity of this tube and the fact that it extended
across the case at a level with the tops of the plants, it was
possible to control the temperature within 0.5 of a degree.
The temperature of the chamber was determined by means of an
incubator thermometer inserted through the top of the case, with
its bulb reaching down to the level of the plants in the center.
For experiments run at high humidity, a practically saturated
atmosphere was obtained by hanging from four pans at the
top of the chamber strips of cloth which were kept supplied with
water by tubes running from a tank above. These cloths
drained into three pans below, one of which was placed in the
flat of moistened sand and the others located on the floor. Addi-
tional cloths, through which the warm air from the fan was
blown, extended the entire width of the temperature chamber
below the rack and were moistened by constant sprays of water.
An overflow in the bottom pans, which received most of the run-
off water, was avoided by the use of constant-level siphons
which carried off the surplus through drains in the floor.
Still other cloths were hung above either end of the pan in
which the plants were placed, so that the pots of plants were
not only standing in water but were more or less surrounded
by moist cloths. Under these conditions the relative humidity
was so high that the entire inner surface of the temperature
chamber was covered with a film of water which dripped from
the top and ran down the glass sides in streams.
For the low-moisture experiments, all humidifying devices
were removed from the chamber.
[Vor. 20
588 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Since there is no method devised at the present time for
measuring the relative humidity at temperatures above 50? C.,
it was decided to measure the evaporating power of the atmos-
phere. This was done by the use of atmometers which consisted
of two porous cups of the cylinder type connected to burettes
on the outside of the chamber, all connections being made under
water to eliminate air bubbles. These atmometers were care-
fully filled with boiled water, and placed on either side of the
flat on which the plants were to be set. Notched corks were
placed in the top of each burette to prevent evaporation at that
point. After the chamber had been adjusted to a desired
temperature, readings were made on the burettes at regular
intervals. ‘lhe results of these readings are shown in fig. 2.
At the high relative humidity, after the humidifying apparatus
had been thoroughly adjusted, there was very little evaporation
from the atmometers. Even at the higher temperatures, with
no plants in the temperature chamber, there was less than 0.001
cc. evaporation per minute. "These results give some indication
of the conditions existing within the chamber, although it is
not intended to imply that the plants lost water at exactly the
same rate as the atmometers.
At temperatures below 50? C. an approximately saturated
relative humidity for the higher moisture experiments and an
average of 69 per cent for the lower moisture experiments were
calculated from the readings of wet and dry bulb thermometers.
When readings were made with the plants in the chamber, the
relative humidity varied with the time of day, being as low as
55 per cent at night when transpiration was cut to a minimum
and as high as 78 per cent at mid-day. Accordingly, most of the
plants at the lower humidity were treated during the day.
Since transpiration raised the relative humidity of the chamber,
care was taken, when watering the plants in preparation for
treatment, to keep the foliage dry and to allow the water to soak
into the soil before placing them in the temperature chamber.
PROCEDURE
These experiments were made in order to determine the effects
of humidity of super optimal and thermal death temperatures on
1933]
BERKLEY AND BERKLEY—TEMPERATURE AND HUMIDITY 589
the cotton plant. They were suggested by the previous work of
Berkley (31) and the unpublished studies of Fung (11). Cotton
plants (variety Upland Big Ball) were grown in pots containing
a mixture of sand and loam in a room of the greenhouse having a
usual temperature of 25-30? C. Since individual variations
made it necessary to use large numbers of plants of various ages,
seeds were planted at frequent intervals from September, 1931,
until the following September.
Two distinct series were run, one at a low, and the other at
a high, relative humidity. The low-humidity experiments were
made at temperatures between 42 and 84? C. inclusive, and those
at the high humidity, between 40 and 65° C.
Plants of various ages (their ages being computed from the
day the seeds were planted) were exposed to a particular temper-
ature and humidity combination. After the chamber had been
adjusted to the desired temperature, the plants were quickly
passed through one of the glass doors, the slight drop in temper-
ature occasioned by their entrance being quickly adjusted.
They were removed after treatment to a compartment at room
temperature which had a relative humidity similar to that of the
treating chamber. 'The subsequent behavior of all plants
treated was watched and the results noted. A minimum of 24
hours was allowed to elapse before the plants were pronounced
dead. When the leaves and the growing tip of a plant were
killed, it was called “dead” but was kept watered for some days
to allow secondary growth to take place at the nodes in case the
entire plant had not been killed. Such plants as did put forth
secondary growth are listed in the tables. It will be noted that
this was characteristic of the higher humidity experiments only.
SERIES I
Low humidity—The plants treated at the lower relative
humidity wilted to some extent before they were removed from
the temperature chamber. When not too severely injured by
excessive treatment, they revived and became turgid again im-
mediately after removal. When treated for a longer time than
was necessary to kill them, the wilted leaves and cotyledons
temporarily regained their turgidity and to all appearances were
[Vor. 20
590 ANNALS OF THE MISSOURI BOTANICAL GARDEN
unharmed, but the petioles of the younger leaves, the stems just
below the growing tips, and the hypocotyls of the seedlings were
withered beyond recovery. After some hours, the time de-
pending upon the intensity of the treatment, the injured portions
of all plants, including those that temporarily recovered, dried
up, only the cotyledons and leaves remaining green and turgid.
These living organs, connected to the main stem of the plant
only by dead tissue, remained in the green condition from 3 to
14 days before showing signs of death.
A microscopic study revealed no living tissue in the withered
portions of the petioles and hypocotyls, but in the leaves and
growing tips, which had recovered, the tissue was found to contain
a number of mitotic figures. In the meristematic regions of
the plants treated at times and temperatures just sufficient to
kill them, the cell contents were disorganized.
Very few of the plants whose growing tips were killed put
forth secondary growth at the nodes.
Tables 1 and m and fig. 3a give the detailed results of the lower
humidity experiments. The thermal death point for the older
plants treated at the lower relative humidity is in the neighbor-
hood of 63? C., as shown by a sudden break in the line when the
temperature is plotted against the time (fig. 3a). The time
required to kill all of the plants at this temperature was about
7 minutes. The evaporation rate of the atmometer at 63° C.
was 0.26 cc. per minute (fig. 2 and table 11). If the plants were
evaporating at a similar rate the cooling effect of the evaporation
would be appreciable and would account for the greater length
of time necessary to kill the plants at this temperature than at
slightly higher temperatures.
The seedlings 10 days old or less had a slightly lower and more
variable critical temperature (table 1). The cotyledons and
plumules showed about the same resistance as the older plants
but the hypocotyls were permanently injured when treated for 8
minutes at 60° C. This injury to the hypocotyls ultimately
caused the death of the entire plant.
In the older plants the petioles of the succulent young leaves
were similarly affected but the plants were not killed unless the
treatment was prolonged until the younger portion of the stem
1933]
BERKLEY AND BERKLEY— TEMPERATURE AND HUMIDITY 591
85
Temperature in Centigrade Degrees
malo .15 .20 .25 30 35 .40
Cubic Centimeters of Water Evaporated per Minute
Fig. 2. The number of cubic centimeters of water evaporated per minute from
an atmometer, plotted against the temperature in Centigrade degrees. The deter-
minations were made at 10° intervals. The figures in table 1 relating to the evap-
oration rate were taken from this graph
[Vor. 20
592 ANNALS OF THE MISSOURI BOTANICAL GARDEN
and growing tip was injured. ‘Therefore, the lower critical
temperature of the seedlings can be attributed to the lack of
resistance of the hypocotyl and not to the injuring of meriste-
matic regions.
TABLE I
COTTON SEEDLINGS TREATED AT THE LOWER RELATIVE HUMIDITY
Temp. in ; Age of | Number | Number | Number | Evap. rate
degrees C. | Time ants | of plants | living dead (cc. per
days) min.)*
42 72 hr. 10 12 12 .14
45 45 hr. 24 6 .15
47 26 hr. 9 1 15 .16
2 hr. 10 1 12 .19
: 10 min. ; 3 32 21
J 15 min. 1 1 5
20 min. ; 1 4 6
45 min. ; 2 20
215 min. , 1 11 .22
5 min. > 1 10
7 V$ min. ) 1 10
d 10 min. 8
min. 4 1 39 22
7 20 min. 1 2 8 12
7 35 min. 6 66
min. 1 15 25
( min. i 55
( 5 min. ( .26
( min. 1 17 .27
( 2V$ min. 1 ) 6 9
( 5 min. 1 29 29
( 1 min. y 18 3 .28
68 1 min. ) 10 .29
68 2 min. 40 40
69 1 min. 10 2 .29
) 2 min. ) ) 9
1 min. ) ) 1 5 .30
2 min. ) ) 4 2
7 min. ) 8
74 1 min 1( 2 8 32
74 1% min ) 2 21
5 1 min ) 4 4 .93
[7 1 min. ) 11 1 10 .94
18 1 min 6 6 .95
80 1 min. ] 11 3 8 .96
*This column shows the evaporation rate of water in cubic centimeters per minute
from an atmometer, as described under “Apparatus,” for the temperatures indicated.
In order to determine the cause of this differential resistance of
the hypocotyls and cotyledons an experiment was made to find
the amount of water lost during treatment and during various
periods of time following treatment. This was done by deter-
1933]
BERKLEY AND BERKLEY—TEMPERATURE AND HUMIDITY 593
TABLE II
COTTON xin FROM 16 TO 180 DAYS OLD TREATED
THE LOWER HUMIDITY
Temp. in A Age of Number | Number | Number | Évap. rate
degrees C. | Time plants | of plants | living dead (cc. per
(days) .
42 24 hr. 25 € .14
42 48 hr. 16 16 12 6
42 72 hr. 53 € 2
45 2 hr. 76 € 6 .15
47 18 hr. 77 3€ 36 .16
47 26 hr. 1 11 3
47 26 hr. 7 6
1% hr. 1 10 2 18
i 21í hr. ‘ ] 3 9
? 334 hr. [ 4 48 .19
6 hr. , ] 9 3
1 hr. 28 28 .20
hr. 12 E
30 min ) 19 3 16 .21
30 min l , 2
45 min ( 10 8 z
hr. , 18 18
( 3 min 1 4 7 24
) 15 min. 25 5 20
) 20 min. 27 2 25
) 25 min. 55 55
( min. 16 4 12 .25
( 20 min. 128 128
( | min 7 3 4 .25
( ; min 10 2 8
( 10 min 5 50
63 ) min 1 2 12 .26
( / min. 1 18
( 5 min. 8 82 .26
( 1 min. 4 8 197
2 min. 40 4
4 min. 4 11 1138
3 min. 40 T rt .29
2 min 70 16 2 1 .90
2V$ min. € 20 2
3 min. ^ 1 1
E 1 min. E 1 6 Ax
74 1 min ' T 2 :
/4 14% min 4 1 1 10
4 2 min 4 10 10
1 min 4 1 2 ) .94
J 1 min ^ 3
] 1 min 120 2
; 1% min 40 2 2t
) 1 min d 3 2 ( .96
) 1 min í 10 5 i
) 1% min ( 32 3:
4 1 min ' 3 2 6 .97
t l6 min f f 5 2 .88
| 1 min. 7 11 1 10
t 114 min. 30 10 10
*This column shows the evaporation rate of water in cubic centimeters per minute
from an atmometer, as described under “Apparatus,” for the temperatures indicated.
[Vor. 20
594 ANNALS OF THE MISSOURI BOTANICAL GARDEN
mining the average percentage of moisture in the hypocotyls
and cotyledons of treated and untreated seedlings (table rr).
The treated seedlings had been exposed for 15 minutes to a
temperature of 60? C., a treatment known to leave the cotyledons
of most of the seedlings in a living condition for a week or ten
days.
In all, five sets were run: in one, which was used as a control,
the plants were weighed before treatment, in one they were
weighed immediately after treatment, in one, after 2 hours, in
one, after 24 hours, and in another, 96 hours after treatment.
Before weighing, the seedlings were cut about one-half inch above
the surface of the soil and again immediately below the junction
of the cotyledons and the hypocotyls. The hypocotyls and
cotyledons were weighed separately, dried, and reweighed.
The results (table 111) show that the hypocotyls and cotyledons
had approximately the same water content before treatment,
but the cotyledons lost considerably more during the treatment
(3.3 per cent loss in the cotyledons as compared with 0.2 per
cent in the hypocotyls). During the first 2 hours the cotyledons
still showed a slightly more rapid rate of loss than the hypocotyls,
but after 24 hours they had regained more than 3 per cent,
while the hypocotyls still continued to lose. After 96 hours the
hypocotyls were very much withered, having an average of only
78.3 per cent moisture, while the cotyledons still retained an
average of 87.4 per cent and were more or less turgid.
TABLE III
LOSS OF MOISTURE IN SEEDLINGS DURING AND AFTER TREATMENT
AT 60° C. FOR 15 MINUTES AT LOW RELATIVE HUMIDITY
oo of Per cent loss Per cent loss
l of moisture in | of moisture in
plants
hypocotyls cotyledons
400 po BE. osos RERERI 92.2 92.1
200 Treated & weighed immediately. . 92.0 88.8
200 Weighed 2 hours after treatment. 91.1 87.7
200 Weighed 24 hours after treatment 88.2 90.8
200 Weighed 96 hours after treatment 78.3 87.4
1933
BERKLEY AND BERKLEY— TEMPERATURE AND HUMIDITY 595
SERIES II
High humidity.—Under the high humidity conditions also the
seedlings showed a somewhat lower and more variable critical
temperature than did the older plants. When first removed
from the temperature chamber, neither the seedlings nor the
RS
o2
Temperature in Centigrade Degrees
L L 1
ie] lo 20 30 40 50 60 TO
Time in Minutes
Fig.3. The time in minutes necessary to kill all plants, plotted against the
temperature in Centigrade degrees. The line (a) shows the results of the lower
relative humidity experiments and (b) the results of the higher. Note the sharp
break in (a). The lines in this graph are not drawn through any definite set of
points but are plotted from tables m and v and are based upon the results obtained
from the treatment of more than 4000 plants.
[Vor. 20
596 ANNALS OF THE MISSOURI BOTANICAL GARDEN
older plants showed any evidences of injury unless they had been
left in the chamber for a period of time considerably longer than
was necessary to kill them. In such a case, the young leaves
and cotyledons appeared as if scalded. In the seedlings, injurious
effects were later evidenced by the wilting or withering of the
TABLE IV
aiii p iege TREATED AT APPROXIMATELY
ER CENT RELATIVE HUMIDITY
Temp. in ] Age of Number Number Number
degrees C. Time plants of plants living dead
(days)
41 24 hr. 12 22 4 18
41 48 hr. 13 22 22
42 9 hr. 10 6 6
45 2 hr. 10 6 6
50 30 min. 14 12 2 10
50 45 min. 14 21 4 17
50 60 min 14 24 8 16
52 10 min 8 11 9
53 5 min 15 6 6
53 10 min 9 6 6
54 3 mi 8 21 20 1
54 5 min 8 15 6 9
55 5 min 8 48 25 23
55 7 min 8 23 23
57 2 min 8 14 14
57 3 min 8 15 8 7
57 4 min 8 15 2 13
58 2 min 8 25 15 10
58 3 min 8 16 3 13
58 5 min 8 20 20
59 3 min 8 23 1 22
59 5 min 8 14 14
60 1 min 8 8 8
60 2 min 8 22 12 10
60 21% min 8 40 40
61 2 min 8 79 79
62 Ló min 8 9 9
62 34 min 8 14 14
62 1 mi 8 37 3 34
62 2 min 8 24 24
63 34 min 8 45 45
64 4% min 8 54 54
65 14 min. 8 28 28
1933]
BERKLEY AND BERKLEY—TEMPERATURE AND HUMIDITY 597
cotyledons, followed either by their abscission or by the drying
up of the whole plant, the amount of injury and the time of its
appearance depending upon the severity of the treatment. In
older plants, injury appeared in the form of flaccidly wilted leaves
and blackened growing tips which proved to be dead. In all
cases, the growing tips were more resistant than the leaves, and
even when the growing tips were killed secondary growth often
appeared at the nodes. There was no withering of the hypo-
cotyls and petioles (the cotyledons and leaves remaining green)
as described for the lower humidity experiments.
Tables rv and v and fig. 3b give the detailed results of the
higher humidity experiments. It will be noted from the tables
and particularly from the figure that plants of any given age
were much less resistant to high temperatures in the saturated
atmosphere than at the lower humidity. The thermal death
temperature evidently lies in the neighborhood of 55° C., if a
definite point can be determined from this curve. This is 8° C.
below that shown by the plants treated at the lower humidity.
TABLE V
COTTON PLANTS FROM 18 TO 120 DAYS OLD TREATED
AT APPROXIMATELY 100 PER CENT RELATIVE HUMIDITY
Temp. in ] Age of Number | Number | Number |Secondary
degrees C. Time lants | of plants living dead growth*
days)
40 7 hr. 18 8 5 3
40 10 hr. ] 30 7 23
40 12 hr. ) 6 6
4] 24 hr. 50 30 24 6
4 72 hr t 6 2 4
4 9 hr 3 26 26
4 2 hr. 4 18 18
4 2 hr. 12 12
4 20 min. 6 6
4 40 min 6 5 1
48 1 hr. 6 2 4
435 114 hr. 6 3 3
48 134 hr. 38 38
) 30 min. 14 11 a
) 45 min. 77 19 13 6 6
hr. hae 38 17 19 19
10 min 12 11 11
min 26 6 1 5
min 36 3 3
10 min 35 102 102
min 72 1 11
min 70 4 2 2
[Vor. 20
598 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE V—Continued
Temp. in x Age of Number | Number | Number |Secondary
degrees C. | Time P E of plants | living dead growth”
5
3 min. 1 2 16
1
2
—
© =
= to
O e en GU is i PRE 00
t2
DIDI CSO:-i
bt
z
=
5
—
=
M pi
h ^
C.b2b20D-—-——Z
L
L
114 min. 60
Angra ih RR
DAD
BEEBE
PRPPPRPE SES
PRIUS, cc a a a
*This column shows the number of plants that put forth secondary growth at the
nodes after the growing tips had been killed.
DISCUSSION
When the data of the two sets of plants were compared, it was
found that the cotton plant was much more resistant to high
temperatures under the lower-humidity conditions. Further-
more, the nature of the injury and the parts of the plant first to
be affected were entirely different in the two sets. At the lower
relative humidity, the petioles, young stems, and the hypocotyls
were the first to be killed, whereas at the higher relative humidity
they were the last to be affected, the leaves and cotyledons
1933]
BERKLEY AND BERKLEY— TEMPERATURE AND HUMIDITY 599
dying first. "The seedlings treated at the lower relative humidity
were invariably killed by injury to the hypocotyls.
Experiments showed that the cotyledons of the seedlings
treated at the lower relative humidity lost moisture at à much
greater rate than the hypocotyls during the treatment. Im-
mediately following their removal from the temperature chamber
the hypocotyls lost moisture rapidly while the cotyledons re-
gained a large percentage of that lost during treatment. This
would indicate that the evaporation of the water immediately
utilized enough of the heat energy around the leaves and cotyle-
dons to protect them until the other portions became injured
and even killed. This assumption is further substantiated by
the fact that all plants were so readily killed in the saturated
atmosphere where transpiration was reduced to à minimum.
These experiments tend to prove the statement first made by
Sachs (764) and later by Ewart ('03) and Baker (729) that reduced
transpiration allows a more rapid concentration of heat in the
plant. This phenomenon has not been emphasized by sufficient
experimental data to show its true significance, and apparently
not in any case has it been previously demonstrated under the
conditions of thermal death temperatures. Clum (’26) claimed
that this cooling effect was greatly over-estimated, but his
experiments on thermal death temperatures were not sufficiently
well controlled to justify his statement.
In the light of the facts shown by the present experiments it is
evident that the thermal death point has not as yet been clearly
defined. It will be necessary to limit the term to the death of
the protoplasm alone and not to the entire plant. The death of
the plant might be caused by the injuring or killing of some
portion or organ which would prevent normal functioning,
as shown by the seedlings treated at the lower relative humidity.
Local injury which is not outwardly evidenced may occur at
temperatures below that designated as the thermal death tem-
perature. Litardiére (25), working with onion root tips, found
that the nuclei were affected at 48? C. after 24 hours and that the
injury increased with higher temperatures even for shorter
periods of time. Yamaha (’27) gave 38? C. as a critical temper-
ature for the root tips of the bean, and Milovidov (32), working
[Vor. 20
600 ANNALS OF THE MISSOURI BOTANICAL GARDEN
with a number of different plants, found that temperatures
around 45-47? C. were injurious to the plants studied if the
treatment were prolonged. The cursory microscopic studies
made on the cotton plants in the present work showed that
temperatures of 55° C. and above, even for very short periods
of time, were injurious to the protoplasm, causing plasmolysis
and deformation of the nuclei. Since these were not uniformly
distributed in the plant it would be difficult to estimate their
ultimate effects.
Blackman’s idea of the “extinction temperature"! (’05)
necessitates the setting of an arbitrary time limit which, in the
opinion of the writers, is not justified since the time required for
the protoplasm itself to assume the thermal death temperature
would vary with atmospheric conditions under which the plants
were treated.
At present no entirely satisfactory definition of the thermal
death point can be given, but the following may serve until more
data is available: The thermal death point is that temperature
which, at a given relative humidity, will kill the protoplasm im-
mediately upon its assumption. Were it not for the indication
that humidity has an effect in addition to preventing cooling
caused by the retardation of the transpiration rate, it would be
possible to eliminate from the definition the qualifying phrase
dealing with humidity. If the humidity had no other effects
it would merely vary the time required for the plant cells to
assume the temperature necessary to kill the protoplasm. This
does not appear to be the fact, however, since there is so much
difference between the low- and high-humidity experiments. It
is likely that the extremely high relative humidity has an addi-
tional effect, that of smothering the plants. When plants from
an atmospheric temperature of 25-30? C. were abruptly trans-
ferred into a temperature of 50-60? C. and a practically saturated
atmosphere, condensation immediately took place on the surface
of the leaves and stems. By reducing the transpiration stream
1 . we ought to find a temperature at which the earliest estimation that
could be actually made would give no measurable assimilation. The lowest
temperature to give this result might be called the ‘extinction temperature’
(say in 100 seconds, for the accepted specific extinction temperature would an course
have to be arbitrarily defined in time units).’’
1933]
BERKLEY AND BERKLEY— TEMPERATURE AND HUMIDITY 601
and the usual exchange of gases, this film of water would naturally
limit the oxygen supply of the plant. Since plants in the higher
temperature would demand more oxygen than could be obtained,
it is possible that the protoplasm was killed by a combination
of factors. The evaluation of the extent and full importance of
this smothering effect will necessitate further experimentation.
In regard to the time limit as a criterion of the thermal death
point, we must consider the factors influencing the rapid change
of temperature of the plants from that of the atmosphere in
which they were grown to that of the treating chamber. Some
of these factors are transpiration in the plant, condensation of
moisture on its surface, and temperature gradients immediately
around the plants. Temperature gradients can be prevented by
keeping the air thoroughly mixed, but it is very difficult to
determine the exact relations of transpiration or condensation
and the change in temperature of the protoplasm.
An examination of fig. 3a shows a sharp break in the line between
60 and 65° C., which marks the critical temperature for the cotton
plant at the low relative humidity. A thermal death temper-
ature of 63° C. may then be chosen, but that is the temperature
of the atmosphere. It will be noted that it took from 5 to 7
minutes to kill the plants at 63° C. and progressively shorter
intervals of time as the temperature increased, which would
indicate that the protoplasm died immediately upon assuming
the temperature of the atmosphere. The time lag, caused
primarily by the cooling effect of transpiration, makes it neces-
sary either to determine the exact temperature of the protoplasm
or, more simply, to plot the temperature against the time and
determine from the graph thus obtained the approximate thermal
death point.
It is true that prolonged temperatures below 63° C. will kill
the plants, but it is also evident from the slope of the upper
portion of the line that it did not take, for example, 25 minutes
for the protoplasm to assume a temperature of 60° C. when
treated at this temperature. Then they were killed gradually,
and in such a case the reactions set up by the high temperatures
may have led to the death of the protoplasm.
It may be concluded that no definite temperature can be set
[Vor. 20
602 ANNALS OF THE MISSOURI BOTANICAL GARDEN
as the thermal death point of the cotton plant without stating
first the atmospherie conditions under which the plants were
treated and giving the ages of the plants. Transpiration plays
a definite role in the cooling of the leaves and cotyledons under
low relative humidity conditions.
SUMMARY
1. T'wo series of cotton plants 5 to 180 days old were exposed
to super optimal and thermal death temperatures for periods
of time ranging from 1% minute to 72 hours. In Series I the
plants were exposed to temperatures of 42-84? C. at an average
relative humidity of 69 per cent at temperatures below 50? C.
The evaporation rate was substituted for relative humidity in
this series. In series II the plants were exposed to temperatures
of 40-65* C. at an approximately saturated atmosphere.
2. Seedlings were less resistant to high temperatures at any
given relative humidity than older plants.
3. The plants were less resistant to high temperatures at
the higher relative humidity.
4. At the higher relative humidity, the leaves and cotyledons
were the first parts of the plant to be affected.
9. At the lower relative humidity, the hypocotyls of the seed-
lings and the petioles and young stems of the older plants were
the first to be affected.
6. During treatment at the lower humidity, the cotyledons
lost water rapidly whereas the hypocotyls lost very little. After
removal from the treating chamber the cotyledons regained
a large percentage of the water lost and became turgid again,
whereas the hypocotyls continued to lose moisture until they
were completely withered.
7. The more rapid rate of transpiration in the cotyledons
apparently utilized sufficient heat energy to protect them from
being noticeably injured until the hypocotyls were killed. "These
data tend to substantiate the theory that the cooling effect of
transpiration is of great value to plants in preventing an accu-
mulation of heat energy.
8. The saturated atmosphere of Series II appeared to have
an additional effect, that of smothering the plants. This was
1933]
BERKLEY AND BERKLEY— TEMPERATURE AND HUMIDITY 603
apparently due to the retardation of the gas exchange caused by
the condensation of moisture on the surface of the plants, thus
reducing the oxygen supply.
9. Due to the two-fold effects of humidity the following is
given as a tentative definition of the thermal death point: The
thermal death point is that temperature which, at a given
relative humidity, will kill the protoplasm immediately upon
its assumption.
10. No definite temperature can be given as the thermal death
point of the cotton plant without stating the humidity of the
atmosphere and the age of the plant.
The writers wish to express their appreciation to Dr. E. S.
Reynolds, Physiologist to the Missouri Botanical Garden, for
suggestions and criticisms given during the progress of the work.
BIBLIOGRAPH Y
Baker, F. S. 29). Effects of excessively high temperatures on coniferous reproduc-
tion. Jour. Forest. 27: 192
Berkley, E. E. (31). Studies of the effects of different lengths of day, with varia-
tions in temperature, on vegetative growth and reproduction in cotton. Ann.
Mo. Bot. Gard. 18: 573-604. :
Blackman, V. H. (05). Optima and limiting factors. Ann. Bot. 19: 281-295.
1905.
Bose, J. C. (13). Researches on irritability of plants. Longmans, Green and Co.,
192
Clum, H. H. (26). The effect of transpiration and environmental factors on leaf
temperatures. I. Transpiration. Am. Jour. Bot. 13: 194—216.
—— ———, (26). The effect of transpiration ie environmental faoltas rs on leaf
temi pow bns II. Light intensity and the relation of ins nici to the
thermal death point. bid. 13: 217-230.
Collander, R. (24). Beobachtungen über die quantitativen Beziehungen zwischen
Tótungsgeschwindigkeit und Temperatur beim Würmetod pflanzlicher Zellen.
Soc. Sci. Fennica, Comment. Biolog. 1: 1-12. 1924.
Ewart, A. J. (03). On the physics and Mas as of protoplasmic streaming in
plants. pp. 59-68. Oxford, 1903.
Fung, H. K. (’11). An ecological study of the American cotton plant with incidental
reference to its possible adaptability in China. An unpublished doctor's thesis
of Cornell University. June,
Gilbert, B. E. (26). The response of certain photoperiodic plants to differing
emperature and humidity conditions. Ann. Bot. 40: 315-320. 1926.
Lepeschkin, W. (25). Pflanzenphysiologie. Julius Springer, Berlin, 1925.
Litardiére, R. de (25). Les diverses étapes de l’ “agonie” des noyaux sous l'in-
fluence d'une température élevée. Soc. Biol, Compt. Rend. 92: 796-798.
1925.
[Vor. 20, 1933]
604 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Milovidov, P. F. (32). Einfluss von Wasser hoher Temperatur auf den Kern der
Pflanzenzellen im Lichte der Nuklealreaktion. Protoplasma 17: 32-88. 1932.
Pfeffer, W. F. P. ('03). Physiology of plants 2: 224-231. Eng. ed. i
Sachs, J. (63). Die vorübergehenden Starre-Zustünde periodisch beweglicher und
reizbarer Pflanzenorgane. Flora 46: 449—459. 1863
————, » 64). Über die obere Temperatur-grünze der Vegetation. Ibid. 47:
5-12. 1864.
Wallace, R. H. (31). Studies on the sensitivity of Mimosa pudica. III. The
effect of temperature, humidity and certain other factors upon seismonic
sensitivity. Am. Jour. Bot. 18: 288-307. 1931.
Yamaha, G. (27). Experimentelle zytologische Beiträge. II. Mitteilung. Uber
die Wirkung des destillierten Wassers auf die Wurzelspitzenzellen von Vicia
Faba bei verschiedenen Temperaturen. Imp. Univ. Tokyo, Jour. Fac. Sci.
2: 215-290. 27.
STUDIES IN THE APOCYNACEAE. IV
THE AMERICAN GENERA OF ÉCHITOIDEAE
ROBERT E. WOODSON, JR.
Research Assistant, Missouri Botanical Garden
Instructor in Botany, Henry Shaw School of Botany of Washington University
INTRODUCTION
Because of their diverse and highly complicated floral mech-
anism, the American genera of the subfamily Echitoideae are at
present perhaps the most imperfectly understood of Apocynaceae.
As in pre-Linnean and early post-Linnean times laticiferous herbs
of several distinct affinities were grouped indiscriminately under
the name ‘‘Apocynum,” so even to-day practically any echitoid
liana indigenous to the western hemisphere may pass as a species
of the inclusive genus Echites P. Br., although several excellent
genera have been segregated from that amorphous and hetero-
geneous aggregate by Bentham, Robert Brown, Alphonse de
Candolle, Mueller-Argoviensis, and other eminent systematists.
To the less exacting botanical publie, however, Echites has re-
mained a convenient catch-all and species of very dubious
congenericity have been described and redescribed under that
name until the literature has become so involved that it is a
dangerous task to attempt routine determinative work, and much
more so to essay the description of novelties.
The Apocynaceae of the western hemisphere attain their
greatest complexity in tropical South America. Although mono-
graphic work hinting of any degree of finality upon most groups of
tropical American plants had best be deferred for the future
because of the incomplete state of our knowledge concerning the
flora of that region, constant exploration in behalf of varied
interests is bringing to the attention of science increasing multi-
tudes of plants which must be critically identified by the system-
atic botanist. In order to facilitate that activity and to render
the results more sure with respect to the numerous representatives
of Apocynaceae, it has been considered highly desirable to under-
take at this time a tentative revision of the troublesome Echit-
oideae.
Issued December 19, 1933.
Ann. Mo. Bor. Garp. Vor. 20, 1933. (1) (605)
[Vor. 20
606 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The most perplexing problem, as well as that first encountered
in the course of such a revision, as has already been intimated,
concerns generic relationships. The first requirement of a re-
vision of the American Echitoideae should be the careful delimita-
tion of the genera most deserving of recognition, and their
epitomization in the form of an identificatory key. The second
problem, of necessity wholly dependent upon the first, is that of
a separate revision of each genus maintained within the sub-
family.
In order to judge and to correlate more competently the
numerous genera which have been proposed from time to time,
the morphology of certain critical vegetative and floral organs
as been investigated and interpreted in separate reports which
are in preparation for publication in the near future. Only a
brief account of the structural features employed as classificatory
criteria is included here as an aid in the use of both generic and
specifie keys. It may be well to point out at this time that the
limits of the subfamily Echitoideae as accepted in this instance
are those established in the first paper of this series! to the
exclusion of the genera of Apocynoideae characterized by pollen
maintained within persistent tetrads.
For the purely taxonomie portion of the revision, recourse has
been had to the extensive collections of most of the principal
herbaria of America and Europe. Since it is believed that such
inclusions generally enhance the value of taxonomie studies, ex-
siccatae have been freely cited, particularly in the instance of
difficult or little-known species. The herbaria where specimens
have been examined or obtained for study, together with the
symbols employed in their citation, are as follows: Arnold Arbo-
retum of Harvard University, Jamaica Plain (AA); Botanischer
Garten zu Berlin-Dahlem (B); Museo Nacional de Buenos Aires
(BA); Herbier Boissier, Institut Botanique de l'Université,
Geneva (BB); British Museum (Natural History), London (BM);
Jardin Botanique de l'État, Brussels (Bx); Botanisk Museum,
Copenhagen (C); California Academy of Sciences, San Francisco
(CA); Lindley Herbarium, Botany School, Cambridge, Engl.
(Camb.); Delessert (D) and de Candolle (DC) herbaria, Con-
! Woodson, R. E., Jr. Ann. Mo. Bot. Gard. 17: 9. 1930.
(2)
1933]
WOODSON—STUDIES IN THE APOCYNACEAE. IV 607
servatoire Botanique, Geneva; Field Museum of Natural History,
Chieago (FM); Gray Herbarium of Harvard University, Cam-
bridge, Mass. (G); Royal Botanie Gardens, Kew (K); Linnaean
Society of London (Linn.); Dudley Herbarium, Leland Stanford
Junior University, Palo Alto (LS); Botanisches Museum, Munich
(M); Missouri Botanical Garden, St. Louis (MBG); Museo Com-
ercial de Venezuela, Caracas (MC); Muséum National del’ Histoire
Naturelle, Paris (MP); New York Botanical Garden, New York
City (NY); Academy of Natural Sciences of Philadelphia (PA);
Pomona College, Claremont (PC); Naturhistoriska Riksmuseum,
Stockholm (S); Botanisch Museum, Rijks Universiteit, Utrecht
(U); University of California, Berkeley (UC); United States
National Herbarium, Washington (US); Naturhistorisches Mu-
seum, Vienna (V). To the curators and members of the staff
of these institutions, as well as to many other friends, the writer
would express his gratitude for innumerable instances of aid and
good will. He is also greatly indebted to the Board of Trustees
and to the Director of the Missouri Botanical Garden for the
privilege of a trip to Central America in 1930 for observation and
study.
HisroricaL REVIEW
None of the species which are now comprised within the
American genera of Echitoideae was known to Linnaeus at the
time of publication of the first edition of the ‘Species Plantarum’
in 1753 or of the fifth edition of the ‘Genera Plantarum’ the year
following. In 1756, however, Patrick Browne? inaugurated the
genus Echiles, citing rather full distinguishing characters, but
failed to assign a binomial species. The single polynomial
referred to the genus by Browne, ''Echiles foliis ovatis nitidis
venosis; floribus herbaceis," was founded upon a plant well illus-
trated by Sir Hans Sloane; and quite clearly establishes the
common species of the Greater Antille, the Bahamas, the
peninsula of Yucatan, and southern peninsular Florida, Æ.
umbellata Jacq., as the type of the genus as we shall deal with it
here.
2 P. Br. Hist. Jam. 182. 1756.
3 ** Apocynum scandens majus, folio subrotundo," Sloane, Nat. Hist. Jam. 1: 207.
pl. 181, fig. 2. rfe
(3)
[Vor. 20
608 ANNALS OF THE MISSOURI BOTANICAL GARDEN
The monotypic genus Echites as defined by Browne was inter-
preted by Linnaeus‘ in the tenth edition of the ‘Systema’ merely
as constituting a species of T'abernaemontana, and the former
generic name was employed as a substantive adjective.
Four years after its first publication, the genus Echites was
expanded by Jacquin® to include ten species: E. biflora, E.
umbellata, E. agglutinata, E. trifida, E. quinquangularis, E.
suberecta, E. torosa, E. repens, E. spicata, and E. corymbosa. In
1763, Jacquin® maintained the species which he had described
three years previously, illustrating each in an admirable manner.
Of the ten enumerated, which are now distributed among nine
well-known genera, all but three were original. For E. umbellata,
Jacquin’ was able to cite four pre-existing polynomials, two for
E. suberecta,’ and one each for E. torosa,? and E. biflora.1
In the second edition of the ‘Species Plantarum’ Linnaeus"
included Zchites within the ‘“Pentandria Monogynia," and
appended the ten species of Jacquin. Two years subsequently
the genus also appeared in the sixth edition of the ‘Genera
Plantarum.’!
It would be a weary task to record in detail the great enlarge-
ment of the genus Echites, which almost immediately followed
its original publication, to include scores of Apocynaceous species
of the subfamily Echitoideae native to nearly all parts of the
* L. Syst. Nat. ed. 10. 945. 1759.
5 Jacq. Enum. Syst. Pl. Carib. 13. 1760.
ë Ibid. Select. Stirp. Am. Hist. 1: 30-35; 2: pls. 21-30. 1763.
7“ Echites scandens foliis ovatis nitidis venosis; floribus herbaceis,” P. Br. Hist.
Jam. 182. 1750; sprit scandens folio cordato, Tesi albo," Catesb. Nat. Hist.
Carol. 1: 58. pl. 68. 1754; “Apocynum scandens majus, folio subrotundo," Sloane,
Nat. Hist. Jam. 1: 207. pl. 131, fig. 2. 1707; ^ Periploca alia, floribus ug god cir-
cinatis & crispis, seu Nerium scandens, radice Bryoniae tuberosae," Plum. Pl. A
210. pl. 216, fig. 2. 1759.
8 “Apocynum scandens, amplo flore villoso, luteo, siliquis angustissimis," Plum. Cat.
Pl. Am. 2. 1703; “Apocynum erectum, fruticosum, flore luteo maximo specio-
sissimo,” Sloane, Nat. Hist. Jam. 1: 206. pl. 130, fi edd . 1707.—The former appears
to apply somewhat more obviously to E. biflora Ja
“Nerium sarmentosum, scandens, ramulis fepe P folliculis gracilibus torosis,"'
P. Th Hist. Jam. 181. pl. 16, fig. 2.
10 ** Apocynum scandens, ips nerit albo,” Plum. Descr. Pl. Am. 82. pl. 96. 1693.
u L. Sp. Pl. ed. 2. 307. 1762.
2 L. Gen. Pl. ed. 6. 117. 1764.
(4)
1933]
WOODSON—STUDIES IN THE APOCYNACEAE. IV 609
world.? The result was soon so obviously conglomerate that in
1811, in his treatise ‘‘On the Apocineae," Robert Brown" rec-
ommended the limitation of the generic name to the species of
the western hemisphere, distinctly citing E. umbellata Jacq. as
theoriginalspecies. Although, unfortunately, he did not attempt
to redefine the characters of Echites, Brown described in the
same work" the genus Prestonia, which is therefore the second of
the American genera of Echitoideae in point of age.
Since 1811, the publication of American genera of the sub-
family Echitoideae has constantly augmented. Summarized
chronologically in tabular form, the appearance of genera from
1756 until the present has been as follows:
1756 Echites P. Br. Hist. Jam. 182. 1756; Jacq. Enum.
Syst. Pl. Carib. 13. 1760.
1811 Prestonia R. Br. Mem. Wern. Soc. 1:69. 1811.
1818 Forsteronia G. F. W. Mey. Fl. Esseq. 133. 1818.
1819 Thenardia HBK. Nov. Gen. 3: 209. 1819.
1825 Haemadictyon Lindl. Trans. Hort. Soc. Lond. 6: 70.
1825.
1828 Syringosma Mart. ex Rchb. Consp. 134. 1828.
1838 Exothostemon G. Don, Gen. Hist. Dichlam. Pl. 4: 82.
1838.
1840 Mandevilla Lindl. Bot. Reg. N. S. 3: pl. 7. 1840.
1841 Odontadenia Benth. in Hook. Jour. Bot. 3: 242. 1841.
Thyrsanthus Benth. loc. cit. 245. 1841.
1844 Malouetia A. DC. in DC. Prodr. 8: 378. 1844.
Anisolobus A. DC. loc. cit. 395. 1844.
Robbia A. DC. loc. cit. 444. 1844.
Secondatia A. DC. loc. cit. 445. 1844.
Laseguea A. DC. loc. cit. 481. 1844.
Dipladenia A. DC. loc. cit. 1844.
Laubertia A. DC. loc. cit. 486. 1844.
1849 Cycladenia Benth. Pl. Hartw. 322. 1849.
1855 Cylicadenia Lem. Illustr. Hort. 2: Misc. 9. 1855.
1860 Heterothrix Muell.-Arg. in Martius, Fl. Bras. 6': 133.
1860.
13 Of. Stadelm. Flora 24!: Beibl. 1-13. 1841.
M R. Br. Mem. Wern. Soc. 1: 59. 1811.
1$ [bid. loc. cit. 69. 1811.
(5)
610
1878
1897
1905
1909
1917
1920
1924
1927
1931
1932
[Vor. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Macrosiphonia Muell.-Arg. loc. cit. 137. 1860.
Amblyanthera Muell.-Arg. loc. cit. 141. 1860, not
Blume.
Mesechites Muell.-Arg. loc. cit. 150. 1860.
Rhodocalyx Muell.-Arg. loc. cit. 172. 1860.
Rhabdadenia Muell.-Arg. loc. cit. 173. 1860.
Stipecoma Muell.-Arg. loc. cit. 175. 1860.
Elytropus Muell.-Arg. Bot. Zeit. 18: 21. 1860.
Prestoniopsis Muell.-Arg. loc. cit. 22. 1860.
Urechites Muell.-Arg. loc. cit. 22. 1800.
Chariomma Miers, Apoc. So. Am. 110. 1878.
Eriadenia Miers, loc. cit. 117. 1878.
A ptotheca Miers, loc. cit. 150. 1878.
Rhaptocarpus Miers, loc. cit. 151. 1878.
M icradenia Miers, loc. cit. 158. 1878.
Homaladenia Miers, loc. cit. 163. 1878.
Angadenia Miers, loc. cit. 173. 1878.
Perictenia Miers, loc. cit. 182. 1878.
Temnadenia Miers, loc. cit. 207. 1878.
Mitozus Miers, loc. cit. 217. 1878.
Streptotrachelus Greenm. Proc. Am. Acad. 32: 298.
1897.
Bracea Britton, Bull. N. Y. Bot. Gard. 3: 448. 1905,
not King.
Orthechites Urb. Symb. Ant. 6: 36. 1909.
Belandra 8. F. Blake, Contr. Gray Herb. N. S. 52: 78.
1917.
Neobracea Britton, in Britton & Millsp. Bahama Fl.
335. 1920.
Codonechites Mgf. Notizblatt 9: 80. 1924.
Macropharynx Rusby, Mem. N. Y. Bot. Gard. 7: 327.
1927.
Salpinctes Woodson, in Gleason, Bull. Torrey Bot. Club
58:453. 1931.
Allomarkgrafia Woodson, Ann. Mo. Bot. Gard. 19:
45. 1932.
Asketanthera Woodson, loc. cit. 46. 1932.
Fernaldia Woodson, loc. cit. 48. 1932.
(6)
1933]
WOODSON—STUDIES IN THE APOCYNACEAE. IV 611
Galactophora Woodson, loc. cit. 49. 1932.
Peltastes Woodson, loc. cit. 375. 1932.
An interesting if frequently perplexing feature shared in
common by many species of the majority of genera enumerated
is their persistent popular association with the parent genus
Echites within which they were once included. Perhaps the
highest attainment of the research upon the Apocynaceae of
Alphonse de Candolle and particularly of Mueller-Argoviensis
was the distillation of the American representation of the in-
clusive genus Echites into numerous more natural entities. Un-
fortunately, the painstaking studies of those eminent systematists
was discredited to a certain degree by the contemporaneous
treatment of John Miers which has discouraged the employment
of the valid segregates by confounding with them a miscellaneous
assortment of incongruous species and genera.
The revision of the Apocynaceae under the authorship of K.
Schumann" in Engler & Prantl’s ‘Natiirlichen Pflanzenfamilien’
has helped to reclarify the status of the American Echitoideae,
but has suffered as a result of the acceptance of Miers's mistaken
interpretations in several instances. Schumann unfortunately
profited little by the excellent natural order given the genera of
the subfamily by Mueller, and as a consequence in his enumer-
ation one finds Eriadenia Miers and Mandevilla Lindl., considered
as synonymous by Markgraf!” and in the present revision, sepa-
rated by such discrepant genera as Macrosiphonia Muell.-Arg.,
Rhodocalyx Muell.-Arg., Cycladenia Benth., Dipladenia A. DC.,
Odontadenia Benth., Elytropus Muell.-Arg., Rhabdadenia Muell.-
Arg., and Laubertia A. DC., in the order named. This situation
is the result of the key characters employed by Schumann,
particularly that of gross habit, which not only separate closely
related genera but even exclude generically certain species from
others which are manifestly their congeners.
MORPHOLOGY OF THE TAXONOMIC CRITERIA
The danger of basing generic distinctions within the Echit-
oideae upon habit already has been observed. As a group, the
16 K, Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 109-189. 1895.
17 Mgf. Notizblatt 9: 85. 1924.
(7)
[Vor. 20
612 ANNALS OF THE MISSOURI BOTANICAL GARDEN
subfamily is predominantly composed of extensive lianas so
characteristic of the tropical American rain-forests. Forsteronia,
Prestonia, Odontadenia, and Echites (sensu stricto) are familiar
examples of genera with exclusively volubile habit among all
known species. Neobracea is fruticose, while Malouetia is the
only known genus attaining arboreal proportions. The species
comprising Macrosiphonia, Salpinctes, Galactophora, and Rhodo-
calyx might with almost equal aplomb be called suffrutescent
herbs or subherbaceous undershrubs. Cycladenia is unique with-
in the entire family because of its low, subsucculent, herbaceous
habit, and its subalpine habitat. Various combinations of habit
render that character an extremely fallible guide in particular
instances. Mandevilla is probably the outstanding example in
this regard, including among its many species lianas, suffruticose
shrubs, and suffrutescent herbs in a wide range of intergradation.
Although predominantly terete, the stems of certain American
Echitoideae display a conspicuous alate development, as in
several shrubby species of Mandevilla subgen. Exothostemon (§
Eriadenia Mgf.). Such development of the stem appears to be
caused by the activity of the phellogen rather than of the vascular
cambium, however, and thus bears only a superficial resemblance
to the polydesmic stems of certain other tropical lianas.
Phyllotaxy is relatively uniform throughout the subfamily in
America, the distichous arrangement being almost invariable.
Mandevilla Benthamii (A. DC.) K. Sch. constitutes a striking
exception in its ternate or quaternate foliage. The leaves of
species of Laubertia and Macrosiphonia are occasionally ternate
as well.
The foliar glands of several genera are easy clues for identi-
fication. "These organs are small, occasionally quite inconspicu-
ous, multicellular, aculeolate emergences borne upon the ventral
surface of the midrib. In Allomarkgrafia, Mesechites, Macro-
siphonia, and most species of Forsteronia, they occur at the base
of the midrib in shapes, positions, and numbers generally char-
acteristic of genera or subgenera and to a somewhat less extent
of species. In Allomarkgrafia, Macrosiphonia, and Forsteronia
the glands are fusiform and indistinctly gathered into groups of
few to several. In Mesechites, however, the two component sub-
(8)
1933]
WOODSON—STUDIES IN THE APOCYNACEAE. IV 613
genera are rather nicely distinguishable by means of the glands,
those of subgen. Humesechites assuming a rather flat or laminate
shape and occurring in groups of two to five clustered concentri-
cally along the base of the midrib, while those of subgen. Antille-
chites are less conspicuous, fusiform bodies which are predomi-
nantly found in pairs radially placed at the very base of the midrib.
n Mandevilla subgen. Eumandevilla, the foliar glands are
fusiform and are grouped at the base of the midrib as in Allo-
markgrafia, but in subgen. Exothostemon the same structures
occur in variable numbers along the entire length of the rib.
Were it not for three common species supplying an indubitable
link between the two subgenera, which also have technical
differences in the reproductive organs, the characteristic position
of the foliar glands of subgen. Exothostemon would render it one
of the most easily recognized genera of the subfamily. Species of
the latter subgenus, however, have long been popularly associated
with those of Humandevilla, and in view of this consideration,
together with that of the intermediate characteristies of M.
funiformis (Vell.) K. Sch., M. callista Woodson, and M. congesta
(HBK.) Woodson, it has been thought desirable to maintain their
unity for the present.
The Apocynaceae are popularly known as an exstipulate
group. However, immediately subtending the petiole of many
species of that family, usually forming a definitely interpetiolar
girdle when the phyllotaxy is opposite or verticillate, are more or
less conspicuous appendages, variously arranged, which should
probably be interpreted as stipular vestiges. In the Echitoideae
of the western hemisphere, these structures are usually glandular
in function, although in Odontadenia anomala (Heurck & Muell.-
Arg.) Macbr. they are somewhat foliaceous, this departure from
the predominant condition prompting Miers to establish for its
inclusion the genus Perictenia.
3
ERE
ns e
214; parallela, 157, 218; parvi
140, var. latifolia, 142; paucifolia, 140;
avoniana, nduliflora, 194,
216; p hyteumoides , 194; inc
nifolia, 185; plantaginea,
diflora, '176, var. lat
ma, A , var. congesta, 200,
216 nC 174; pusilla, 182, 216;
811
recurva, 161, 214; Hage a ie. rhyaco-
phila, 168, 214, V. oda 2,170,
var. pr rocumben n 170; M ion 190,
216; Rothrockii, 161; Rusbyi, 9,
216; scapiformi. Schiedeana,
172, 214; Schultzei, 151; Shannonii,
177, 214, f. compa p. 178; sain
3,
4
195, 216; spatulata, 183, 216; s
cola, 201; , stellarioides, 1152; Stuebelii,
in
So
E
heimii, 145; :
valerianoides, 182; Ente, 153,
d ys cetoxieoides, 1 viridis,
eberbaueri, 189,
Worldellizna, 202, 216
Helleranthus quadr angulati us, 313
Heterothrix, 645; pycnantha, 668; Van-
heurckit 771
Hieracium carneum, 798
| tei a 70
Holoc s Leprieurii, 397
seus dumosus 800
, 645; "br. evifolia, 680; lin-
ariaefol ia, 6 » pastoru m, 680; pedunc
ularis Jhon puberula, 680; tonuifolia,
680; 8
Gr Frente, Super optimal and
thermal death temperatures of the
gg plant as affected by variations
S omi ion Mcr ne of nutrient
solutions in relat o growth of
i pelle 27, 579; various, reaction of
myces capsulatus to,
myces capsulatus,
Hymenopappus ie 798
Hypopitys sanguinea, 799
I
Indol, reaction of, to Endomyces capsu-
latus, 517
Iron in nutrient solutions for orchids,
226, 365, 572
Isoetes, 74; Butleri, 75; Engelmanni,
74; melanopoda, 75
J
Juniperus flaccida, 796
K
Keerlia, 37; bellidifolia, 37; effusa, 38;
linearifolia, 33; mexicana, 31; ramosa,
4, 25; skirrhobasis,
Knudson's solution for growing orchids,
365, 572, 574
Kujala, Dr. Viljo, lichens collected by, 7
812 ANNALS OF THE
L
La Garde's solution for growing orchids,
226, 234, 363, 572, 578
sther L. Astranthium and re-
kan ap 23
645; en 673
poe ie d 6 f
E Jrisco-fusca, 674, .
— E aa. OU
"n 'aniennacea, 741;
bracieta, 671; emarginata, "d erecta,
673, Guille eminiana, 673,
olivacea, Mie 2. €o-f
obliquinervi A id.
griseo- aa a, obovata, 673,
» 0 tr psi 673; ô. acutifolia, 673,
673;
scabri 673, ‘
foliosa, 658; glabra, 673; s Quilleminiana,
; ookeri, 6 Mandoni, 671;
obliquinervia, 673; Pentlandiana, 671;
subspicata, 740; inuidia. 740; villosa,
74
4
Lecanora yg pa, "e ‘gy timate 14;
symmicta, 14; umbr
cidea goniopl r^ 19;' lavicide: 19;
E:
3
E
opsis os
Lempholemma, 404; compactum, 404;
— , 406; Dussii, 405; obli ique-
peltatum, 40 5
Lepidoc na carassense, 440
reir vt americanum, 404; Finkii,
Leptogidium byssoides, 403
Leptogium, 404, 416; E bi 418
e
"0 O
o'a
3EE
UL
coralloideum, 419, 430; corrugatulum,
131; denticulatum, 419, 432; diapha-
4. diaphanum f. cor
0;
8 ; foveolatum, 418,
495; fusisporum, 418; granulare, 419;
Hide ii var. osum, 2;
ynum, 419; inflexum, 418,
a var. isidiosulum, 418, 423; laevius,
420; Lafayetteanum, 420; lepto phyl-
linum, 432; subg. Mallotium, 418, ^92;
—— um, 408, 419, = matto-
; mega 18;
centem di , 419; se Aamar 418,
426; moluccanum, 436; olivaceum, 418,
[Vor. 20
MISSOURI BOTANICAL GARDEN
424, var. granulosum, 424; pachy-
cheilum, 420; papillosum, S 422;
phyllocarpum, 419, 428 cam-
pestre, , 43
reg 420; Bos hellum
latum, 421; resupina
oides, 419, ; Standleyi,
490, 435; allan, 420; jstipitatum, 418,
428; tremelloides 17, 416; t melloides,
, Var. azureum . isidios Min "132,
it pressopunctatum, 421, var. leptophyl-
lim , var. minor, 436; T
Tieng: Mar a 11, f. compacta,
No rth niin "Contri-
n to de. 7
— ,asureun 434; MN be
, 408; diaphanum, 421;
Fe 431; melanocarpus , 394; Loa:
sus, 453; velut linus, 403; vesiculosus,
6
a sr he of wae Rica, The foliose and
F
Light: white, effect of, on growth of En-
omyces capsulatus, 521; of red, 522
Linum Schiedeanum, ' 79
Lithospermum viride, 797
Lobaria linita, 17; oregana, 17; pul-
Lobarina verrucosa, 17
Loeselia reggii,
icera arizonica, 797
eria areolata, 69
Bo ^W 12; complanatum flabelli-
orme, 73; lucidulum, 73, var. poro-
philum, 73; paeen v A 73
M
MODUS, TEL lansae, 788;
Berlandieri, 783; Éraohysipbon, 780;
784; Guar-
v
788, var. typica, 787; pinifolia, 788, f
glabr 88, var. i ia, . f
pedu aris, 788, f. b :
prostrata, 787; subg. Telosiphonia,
778; Velame, 788; Velame B. goyazen-
sis, 788; verticillata, 786, a. petraea
787, B , 787, y.
zr var. Missionum, 785; Wrightit,
81
1933]
INDEX 813
Magnesium, relation of growth of orchid
22
alme lla, 440, 447; albida, 448,
caesiocinerea, 448, 452; die
448, 451; Randii, 448; Santamariae,
448, 450
Maltose, growth of orchid seedlings in
5
ace nsis, 663; ca İsta, 700; cau-
rensis, 770; cercophylla, 669; cereola,
12; coccinea, 734; congesta, 675;
convolvulacea, 687; crassinoda, 703;
cuspidata, 730; denticulata, 764; Di jel-
J "D mithii
Exothostemon, 737; Fendleri, 758;
fluminensis, 764; foliosa, 658; fragilis,
. 13; funiformis, 697;
fun niform peduncularis, 697;
750; laxa, 695; sect. laxae, 682; lepto-
phylla, 750; linearis, 738; linearis, 756;
Loesneriana, a luc ida, 718; Lue tzel-
burgii, 698; oni, 693; Mar a n
702, var. S bei. "203, var. typica, 702;
mexicana, 660; minor, 711; mollissima,
na, 665; sect monta e, 661 ri-
can a, 705, var. bahiensis, ded var
typiea, 705; Moritziana, 768; Muel-
um, 68 pei
pital 644, 86; oblongifolia, 708;
palus ris, 764; parvifolia, 753; Pavonii,
749; pend la, 725; Pentla ndiana, 671;
permix 71 5; pla tydactyla, be
polyantha, 769; polyantha, 769;
sina, 777; pycnantha, 668; Rai
Schlimii, 759; Sc humanniana, 6
scutifolia, 663; Sellowii, 719; sert ae
gera, 649, 654; spigeliaeflora, 736;
splendens, 707; Spruceana, 776; suav-
colens, 695; subcarnosa, 644, 772;
subcarnosa, 772; subcordata, 693;
subpaniculata, 678; subsagittata, 644,
645, 743; subscorpioidea, 653; sub-
Vahleana, 764; torosa, 644, 656; Meri
torosae, 655; Trianae, 760; tubiflora,
648; sect. tubiflorae, 647; iana,
695; Ulei, 771; urophylla, 722; Van-
heurckii, 771; velutina, 731, var.
glabra, 732, var. typica, 732; velutina,
Wie venulosa, 723; a asensis,
690; veraguensis, 690; vi villos
Td sulphate in eara Aih solu-
226
Mannose, growth of ongie seedlings in
solutions containing, 574
Maua nodulosa, 65; Struthiopteris,
Media, growth of Endomyces capsulatus
in various, 494
Meningitis, fatal cerebrospinal, Endo-
of 471 capsulatus as a causative agent
o
Mentzelia asperula, 799.
Mesechites, 629; acumi Deer An-
drieuxti, 645, 688; ongkeaix, 644, 775;
angustifolia, 640; bicorniculata, 634;
brevipes, 638; rownei, , 656; citri-
folia, 637; dichotoma, 644; subg
dymadenia, 638; dis , 4081;
subg rore 30; Guayaqui-
lensis, 6 4; nsis, 645, 7
hast ata, 644, Jui “hirtella, 644, 744:
hirtellula, 6 4, 686; japurensis, 630:
jeter lg 644, 744; lanceolata, 639;
linearifo lia, "EUH 636;
minima, 643; myrtifolia, axa-
e 644 686; repens, 639 nll 641;
Micradenia, 645; atroviolacea, 724; cras-
sinoda, 703; fragrans 714; oo.
rliana, 7 oricandia
706; Riedelii, 740; Sellowii, 719.
Missouri, Ferns and fern allies of, 45;
A neutral (?) strain of Mucor sphaero-
sporus
Mitozus brachysta achyus, 753; brevipes,
638; ds "owed 753; exilis, 697; funi-
formis, 697; Guanabaricus, '697; epto-
phyllus , 756; micr ronila. € rugosus,
755; symp looarpus 755; tenuicaulis,
753; versicolor, 752
814
Monilia, 82, 103
Moore, John Adam, Steyermark, Julian
and. Report of a botanical can
tion into the mountains of western
Texas, 791
Moore, Morris.
prt p qe report
of a case, with a s of a
etiologic
yers: a causa-
tive agent of fatal bend men-
ingitis,
Mucor sphaerosporus m Missouri,
A neutral (?) strain of, 46
Mycoblastus melinus, 19; sanguinarius,
, var. Dodgeanus, 19
Mycoderma, 474
My pait nim dermatitis, 103
Nama ea 799; xylopodum, 799
Nephrod
Heskeuntun "Tasting se, 17; laeviga-
var. papyraceum, 17; 'subparile,
Nephromopsis ciliaris, 13; platyphylla,
Nerium sarmentosum, 642
Nitrates, reduction of, in Endomyces
capsulatus,
— in re to growth of orchid
seedlings, , 368.
N we pen E 6
Notholaena, 60; de albata 60; Greggii,
796; nivea « lealbat
Nutrient solutions for silo 225, 363,
Nutritional ions, The sensitivity of or-
chid seedlings to, 22:
Nine am A new ‘yellow, from tropical
A
eel iy (§ Ne yy erat "im 1, 6;
Stuhlmannii, 3; sulphur
Obleti
Ockrolckia upsaliensis, 14
Odontadenia cuspidata, 730; glandulosa,
Oidium, 82, 102, 474
On oclea, 61; sensibilis, 61; Struthiopteris,
Oosporaceae, 104
Ophioglossum, 46;
vulgatum, 46
Orbilia, 396
Orehid seedlings, Nutrient solutions for
; The sensitivity of, to nutritional
ions, 223; Sources of carbo ydrate for
germination and growth of, 5
Engelmanni, 47;
or. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Osmotic concentration, panty of, to
growth of orchid seedling -
Osmunda, 71; cinna siad y-
toniana, 72; regalis NET p?
Struthiopteris, 64
Ostrya Baileyi "79 7
Oxygen, free, relation of Endomyces
capsulatus to, 518
P
Pannaria, 440, 441; euler 441; bras-
iliensis, 442; uleo by m ans, 442;
imbricatu la, 442; isid var.
2, f. cinerascens, 452;
8 tlopory, "rm 445; Vainii, 442, 445
Pannariaceae
13; cronia, 462;
tula ana, 14; enteromor-
pha, 13; exasperatula, 13; hyperopta,
i 463; lophyrea, 13;
ata, vagans, = vitta , 13
Parmeliella, 440, 452;
mierophylloides,
o 61; atropurpurea, 62, va
ushii, 62, var. cristata, 62; Tiao.
6l; pe a
Peltigera puoi 16; canina, 16, va
membranacea, 16, var. oce 10;
NR NN 16; le phora, 16;
malacea, 16; Maurizii, 16; microphylla,
16; Nylasi i, 16; perfida, 16; rufes-
cens, 16; suomemsis, 16; variolosa, 16;
enosa,
TN growth of orchid seedlings in,
Pentstemon baccharifolius, 801
i, 798
Por, Lil rat
Perry . A revision of the North
n species of Verbena, 239
ye oat r amara, 14; faginea, 14; multi-
puncta, 14; pertusa, 1 xanthostoma,
argh 66; heragonoptera, 68
Phlyctis argena, 14
Phosphorus, growth of orchids in solu-
tob containing, 233, 367
Phryma caroliniensis,
Physcia, 15; adglutinata var. pyrithro-
1933]
INDEX
cardia, 15; astroidea, 15; caesia, 15;
a, 15; pyri-
rele na 15; stellaris, 15, var. rosu-
lata, 15
Physma, 404; Boryanum, 407; byrsinum,
08; pruinosum, 408
Pilophoron aciculare, 17
Pinaropappus parvus, 798
e M. Elizabeth. Ferns and
allies of icy
Placodium chrysodetum, 15; citrinum,
td aak Sinn a. "15; ferrugineum, 15;
ostigmoideum, 15
Platy, tysma seplentrionale, 12
Polychidium, 403
Po pen Timulicola, 800
Polyp m, 62; incanum, 63; polypodi-
oldda "63; "virginianum, 63; vulgare, 63
Polystichum, 63, 66; acrostichoides, 63,
incisum, 64, Hen Schweinitzii, 64
sel ohoi.
otamogeton clystocarpus, 802; diversi-
folius, 802
Potassium i in relation to growth of orchid
234, pe
est = hirsuta, 749; ana, 692
psis Fendleri, 670; y cese 676;
red venosa 7
H. and Robert E. Wood-
on, Jr. new yellow Nymphaea
opical Africa, 1
Pteretis, 64; nodulosa, 64
er borg 65; pese aa 65; latiuscu-
var. pseudocaudatum, 65
Perlini té, new or rare, in Texas, 796
Pterigyopsis atra, 403
Pieri ilina,
Pterospora Andromeda, 799
Quercus hypoleuca, 798
R
Raffinose, growth of a seedlings in
solutions containing,
Ramalina calicaris, E geniculata 11;
Roesleri, ori thrausta,
ä A B Contribution to
ains of western Texas, 791
Rhabdadenia campestris,
Rhamnose, of orchid seedlings
in solutions containing, 574, 576
Rhamnus fasciculata
Rhodocalyx coccineus, 734; hypoleucus,
815
781; lanuginosus, 782; ovatus, 735;
35
81; Tweedianus, 7
Rosa mirifica, 800
S
Saccharomycetaceae, 104
Saccha: rem 82, 103, 474; hominis, 103
EAM
tpe cent, relation of growth of
d id seedli ings to, 371
jum, 408
59
: apoda, 74; apus, 74;
Pringlei, 796; E estris, 74
rie ed x. chid seedlings to nu-
tional io
Shive's solution for growing orchids, 572,
Shuttleworthia, 251
iphu E =
Sisymb , 798
Skatol, empet e of, to ee cap-
Sodium borate i in ovde solutions, 226
98
Sorghastrum nutans, 801
ea aa new or rare, in Texas,
9
Sphaerophoraceae, 393
Sphaerophorus globosus var. lacunosus,
20; melanocarpus, 394; Tuckermanii,
Spheno Pp obtusata, 801
ripe
Spiranthes s saltensis, 802
Sporotrichum, 8
quamari pum placa, 14
Stau retails circinata,
Stephanophorus
Stereocaulon tomentosu um, 14
Steyermark, Julian A., John Adam
oore port of a botanical
expedition into the mountains
western Texas, 79
Sticta scrobiculata,
Styleurodon carolinianum, 309
Mgr rw in
Giu"; grow
lutions oleh ted Tikica, 226,
Sulphate, relation of growth of orchid
Mu ngs to, 368
816
Super optimal and thermal death tem-
peratures of the cotton plant as
affected by variations in relative
humidity,
Swertia, 138; — 162; asclepiadea,
wnis, 140; cornioulata, 162;
140; deflexa, 162; acilis,
M ichaun ciana,
14, y angusti-
] t lali 142; pingfolia,
185; plantaginea qu adricornis,
urva, 161; R4 LA ln 191
Symp oriearpos rotundifolius, 797
Synechoblastus, 413
e
T
— 646; cordata, 686, glauces-
686; lasiocarpa, 750; Lo bbiana,
750; poliidifora, 764; palustris, 764;
ndi or 744; tomentosa, 764;
redi 1,
uds chamber for treating cotton
plants, 585, 586
ny st Ar bay reaction of Endomyces
capsulatus to, 500; Super optimal and
thermal dead: of the cotton plant as
affected by variations in relative hu-
midity,
vana. 138; alatus, 139; Bren-
tonianus, 167; brevicornis, 140; Peta
-" " 178; pi dn 163, var. Brentoni-
var.
r
heterantherus ayt p 163;
longicorn 178; meri, 171; rein
florus, Lu ps hc m 140;
gineus, 175; DEN 161; Sc Phiede-
anus, 172; Wh
T! Report of ^ botanical bei
o the moun "^ ' of western, 7
mess new or r NUN + of, 795,
additional MN. of interest, 802
Thamnolia — 14
Thelypteris, 66; onoptera, 68;
marginalis, 67; E pubescens, 66;
spinulosa, 67, var. intermedia, 67; Thel-
ris, 66
Thermutis, 403; velutina, 403
Tillandsia recurvata, 801
Trachelopsermum stans, 659
Tradescantia Wrightii, 801
Trentepohlia, 396
U
Usnea wr z comosa, 9, var.
stuppea, 9; dasypoga, 9, var. subsca-
rede hirta p a e, 9; M
in cu: a, 9; pro ostrata, 9 rugulosa
milis, 9; o tain sa, 9
ule, 251
(Vou. 20
ANNALS OF THE MISSOURI BOTANICAL GARDEN
V
UM texana, 801
uelinia angustifolia, 800
Ventures sinensis,
Verbena, A revision of the North Ameri-
can species. of,
Verbena, 251; affinis, 257; Alopecurus,
285; brosse 326, i eglandulosa,
amoena, 341; An t
Aubletia var. Lamber
342; bipinnatifida, 323, v
325; eti da ta, 268; bonariensis, 254;
onariensis var. litt ar.
DING 253; bracteata, "304, $58;
304, var. albiflora
se: Hearn PRA 304; raat lands 255;
brevibracteata, 338; anadensis, 315,
nadensis var. Zone rgii, 318,
subsp. elegans, 318, var. Lamberti,
316; canescens, 301, var. Roemeri riana,
302; canescens, 304, var
296; caracasana, 257;
polystachya, 268, f. or E
33
e flor ora, 342;
207, ed punica 278, f.
mite 278,
suta, 268; imbricata, 304;
2; i wa, 338; UE
342; Lamberti, 316, var. rosea, 316;
lastostachys, 290; d 205;
lilacina, 340; litoralis, 257
rosella, 288; macrodonta, 289, 358,
360; maritima, 320; Ma ines 342;
menithsefalla, 263; eras 208, 280;
mexicana, 296,
v . 298, rt xylopoda, 297;
T 315; Obletia, 315; officinalis,
262; officinalis var. hirsuta, 296;
9; tifida,
ak UP ot 268; prostrata, 290;
1933]
INDEX 817
bera, AE puce 332; pumila, 338;
pumila albiflora, 318; quadrangu-
laris, i: quadrangulata, 313; race-
repens, 342; rigens, 286; rigida, 252;
riparia, 267; ' robusta, 292; Roemeriana,
302; rubra, 316; rudis, 304; rugosa,
263; simplex, Ran. sphaerocarpa,
262; squarrosa, ad str iste,
a
5;
Wrightii, 328, 358; Xo.
W
Woodsia, 68; obtusa
Woodson, Robert E., Pu Studies in the
Apocynaceae. IV, 605; George H.
Pring, and. A new nor Nymphaea
f tropiodl Africa
Woodwardia, 69; pea, 69; areo-
Sources of carbohydrate for germina-
tion and growth of orchid Lbs.
569
Xanthocephalum, i linearifolium, 25
Xanthoria polycarp
Xylose, growth of i chid seedlings in
solutions containing, 574, 576
Y
Yeast-like organisms, 471; cytology of,
477
Z
Zapania bracteosa, 304
Zygadenus elegans, 801
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T NEM SUES UT EE EON Il A? PRT? EE er oA QM 8 Pee), 2
_ OF THE MISSOURI BOTANICAL GARDEN .
= È à ; : L ^ : :
.. GEORGE T. Moogs.
' Assistant to the Director,
: Karnerine H. LE:GH. |
TIR VON Bonmewx, di Carnot W. Dover,
: pagine. j i4 57 "oMyeologist. ©
~ JESSE M. es aie. Rowann V. La Ganon,
Curator of the Herbarium, — Research Assistant.
ge S, Revi, | 1t. RoBERTE. WooDsoN, JR.,
Physiologist: ce oe eee Assistant. =
Newt C. Horner, -
Librarian me Editor of Publications,
J^ BOARD OF TRUSTEES $
E THE SECUR BOTANICAL GARDEN
President,
GxoncE C. HrrCHCOCK. -
Vice-President,
Samuzt C. Davis.
SSS o ded : Second. Vice-President,
ue eee E BON E Case 3
L. Ray ore o E -— ALBERT T. Paniers. |
| THowAs B. Mayers. ~~ EUGENBPETPUM i
GroRaE T. MOORE. . Ersan A. H. Buspust.
Ande P TORNA Frep G. ZEIBIG.
SS es BX-OFEICIO MEMBERS:
3 Gana Paucar, Bernard F, Dickmann,
SS Chancellor of Washington University. ZE Mayor of the Clty of Bt: Louis.
Eo Me Fumpzatcx F. JOHNSON, - SH SCC Be ALBERT Kuntz, .
1:5... Bishop of of the Diocese of Missouri. | President of The Academy of sd
ric eor Se. ‘Louis. -
X m Davi» C. Tob
itn oto nds tvm of bobus
: Gron F. Barrmnan, Secretary,