SIDA Sis" 


VOLUME 8 NUMBER 1 JUNE 1979 
CONTENTS 
Charles Mohr and plant life of Alabama. L. J. Davenport. ] 
Taxonomy and distribution of Gentiana (Gentianaceae) 
in Mexico and Central America. Il. Sect. Chondrophyllae. 
James S, Pringle. 14 


Revision of North and Central American Najas (Najadaceae). 
Robert R. Haynes. 34 


Bassia (Chenopodiaceae) in North America. Scott L. Collins 
and W7ll H. Blackwell, Jr. 57 


A new species of Erigeron (Compositae) from Coahuila. 
Guy L. Nesom. 


65 
Vascular flora of Cedar Gap Lake and environs, Webster and 
Wright counties, Missouri. Ralph L. Thompson. 71 
Taxonomic notes and new sections of Clitoria subgenus Bractearia 
(Leguminosae). Paul R. Fantz, 90 
New species of Clitoria subgenus Bractearia section Bractearia 
(Leguminosae) from Peru and Ecuador. Paul R. Fantz. 25 
Floristic and geologic aspects of indian middens in salt marshes 
of Hancock County, Mississippi. Lionel N. Eleuterius and 
Ervin G, Otvos, Jr. 102 
NOTES. Notes on three species of ee ssum from North eee 113—New state 
records for Ranunculaceae in the southeastern United States. 114— Cuphea carthagenensis 
(Jacquin) MecbHaee correct orthography 114—Zamia (Cycadaceae) new for Georgia. 
115—Galium species new to the southeastern United States. 116—Notes on Mitracarpus 


hirtus (M. villosus) and Scirpus Pai senitus in Mississippi. 117. 
Documented plant chromosome numbers 1979:1. 119 


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SMU Herbarium 

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Guidelines for contributors are available upon request beginning with 
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Sida, Contributions to Botany, Volume 8, Number 1, pages 1-121. 
Copyright 1979 
by Wm. F. Mahler 


CHARLES MOHR AND PLANT LIFE 
OF ALABAMA’ 


L. J. DAVENPORT 
Ecology and Systematics Section, Debartment of Biology, The University 
of Alabama, University, Alabama 35486 


Charles Mohr (1824-1901) was one of the foremost botanists of the 
southern United States during the late nineteenth century. His stature as an 
early southern botanist is overshadowed only by that of A. W. Chapman, 
author of numerous editions of the Flora of the Southern States. Like Chap- 
man, Mohr was both a resident botanist and, for the most part, an amateur 
botanist. His 40 years of study centered in Mobile, Alabama. These studies 
culminated in the publication shortly after his death of Plant Life of Alabama, 
which remains as the only published attempt to describe the flora of the 
entire state. 

During an investigation of the contents of the herbarium of the Geological 
Survey of Alabama (Mohr Herbarium), I realized that very little was known 
about either Mohr’s life or his scientific works. It is the purpose of this paper 
to examine the life of Mohr, his scientific works, and the circumstances lead- 
ing to the publication of Plant Life of Alabama. 

Facts contained in the following biographical sketch were drawn largely 
from Hoffmann (1887), Scribner (1893), Smith (1901b, c), and Tracy 
(1901). Supplementary facts were garnered from less extensive works and 
from letters exchanged between Mohr and Eugene Allen Smith, then State 
Geologist of Alabama. These letters are stored in the Special Collections of 
the Amelia Gayle Gorgas Library, The University of Alabama. The sources 
for other letters cited are included with the references. 

SKETCH OF THE LIFE OF CHARLES MOHR 

Charles Theodore Mohr (originally, Karl Theodor Mohr) was born De- 
cember 28, 1824 in Esslingen, Wiirttemberg, Germany, the son of Louis M. 
and Dorothea Mohr. He spent his youth in Esslingen and at the Cloister 
Denkendorf, where his father had obtained an interest in a chemical factory. 
His early interest in plants is linked to his great uncle, who was a pensioned 
forester of the district, and to a local pastor who was interested in nature 
studies. 

In the fall of 1842 Mohr entered the polytechnic school of Stuttgart, where 
his studies concentrated on chemistry and natural sciences. These studies 
were supplemented by weekend geological and botanical excursions, during 


‘This paper is adapted from a thesis submitted to The University of - labama in 
partial fulfillment of the requirements for the degree of Master of Scienc 


SIDA 8(1): 1-13. 1979. 


(Ae ced « foc Ae ; 
oo pa 


Fig. 1. Portrait of Charles Mohr, from Smith (190Ic). 


3 


which Mohr made notations of the geological formations and plant associa- 
tions observed (Hoffmann 1887). 

During the summer of 1845 Mohr was assigned to arrange the herbarium 
duplicates of the institute and to distribute these specimens to the secondary 
schools in the area. While pursuing this work he met August Kappler, who 
was engaged in the collection of natural history specimens from Dutch 
Guiana (Surinam). In November, 1845, he left with Kappler on an expedi- 
tion to explore the headwaters of the Surinam River. The party arrived at 
the coast of Guiana near the middle of March, 1846. Mohr was only able to 
spend seven months in South America, however, as an attack of dysentery 
and fever confined him to camp—the first attack of the ill health that would 
always plague him. According to Robinson (1903), Kappler “heartlessly 
left him behind, to live or die, while he [Kappler} pushed onward in his 
ventures.” 

In March, 1847, Mohr returned to Europe and soon found employment as 
a chemist in Brunn, Austria. However, the political turmoil of 1848 forced 
the business to close, while it also eliminated Mohr’s hopes for a higher 
education (Mohr 1879). At this time Mohr and his older brother immigrated 
to the United States, arriving in New York in October, 1848. After a short 
stay in Philadelphia, they moved on to Cincinnati, where Mohr again found 
employment as a chemist. 

In March, 1849, Mohr set off with about 50 other young men as a member 
of the “Cincinnati Gold Mining and Trading Company” to seek his fortune 
in the gold fields of California. After outfitting in Independence, Missouri, 
the group headed westward in mid-April with 10 wagons. The journey be- 
came increasingly difficult, and at Fort Laramie many belongings had to be 
abandoned. Ac this point Mohr traded his gun to a Sioux Indian in exchange 
for a pack horse to carry his books and plant collection (Hoffmann 1887). 

After splitting into parties of 10, the group reached Fort Hall in July. 
It was here that the wagons and baggage had to be given up, and Mohr 
dumped his books and plant collection into the Snake River. The parties 
reunited in Sacramento near the middle of August, after a trip totaling 110 
days. 

Mohr and his party met with some success at placer mining in California. 
However, again, in the spring of 1850 Mohr fell ill to mountain fever and 
decided to return east. He and some others left the mines in September, 
1850, for San Francisco, where they booked passage for Panama. While 
traveling across the Isthmus, the party’s pack mules—along with Mohr’s 
collection of California plants—were stolen, and Mohr again fell ill to fever. 
He finally reached New Orleans and traveled to Cincinnati, arriving toward 
the end of December, 1850. He and his brother purchased a farm in Clark 
County, Indiana, and he married Sophie Roemer on March 12, 1852. 

Various attacks of rheumatism soon convinced Mohr that he was not suited 
for a farmer's life, so he established himself in the drug business in Louisville, 


4 


Kentucky. Here he became acquainted with the bryologist Leo Lesquereux 
and resumed his botanical studies, concentrating on the mosses. However, 
again his health failed, and Mohr was advised by his physician to move to a 
warmer climate. He moved first to Louisiana, then to Vera Cruz, Mexico, and 
in early 1857 established himself in the highlands of Mexico between Cor- 
doba and Orizaba. Again, political turmoil forced him to move on, and he 
arrived in Mobile, Alabama, in the autumn of 1857 

Mohr remained in Mobile for the next 40 years, where he founded a suc- 
cessful drug business. This business suffered much during the Civil War, 
during which Mohr was employed by the Confederate government in the 
manufacture of drugs from native resources and the testing of medicines 
smuggled in from Europe. He remained in the drug business until 1892, 
when he turned the management over to his son, Charles A. Mohr. Mohr 
lived in Mobile for eight years following his retirement, and Schenck (1955 ) 
describes him during this time as “an old and broken man, who spoke Eng- 
lish with a Wurcemberg ian accent, although he had lived in the United States 
for forty years. 

In March, 1900, Mohr and his wife moved to Asheville, North Carolina, 
both to provide a change in climate and to give him the opporunty to work 
in the Biltmore Herbarium. He died in Asheville on July 17, 1901 


— 


SCIENTIFIC WORKS OF CHARLES MOHR 


The scientific works of Charles Mohr can be summarized as being scien- 
tifically accurate but leaning toward the practical and economic aspects of 
botany, agriculture, and geology. In his biographical sketch, Smith (1901c) 
states: 


From about 1878 the results of these [Mohr’ - scientific investigations began to be 
made public in a series of articles, at first practical and economic only, afterwards 
more strictly scientific and specialized, but a directed toward the imparting of 
useful knowledge to his fellowmen. As with his collections, primarily intended t 
illustrate some feature of our natural resources, they grew in breadth and comp 
ness until they became illustrative of monographs. 


Oo 


= 


fod Bi 


These interests and goals adequately sum up the scientific works of Charles 
ohr. 

Tracy (1901) states that Mohr’s first scientific paper was a lecture pre- 
sented in Vienna in 1847 on the geology of the Surinam region. Like most 
of his peers, Mohr was interested in other aspects of the natural sciences 
besides forest botany, and he included mineralogy, geology, and conchology 
among his interests. He contributed a short paper concerned with both 
geology and anthropology to the 1881 Smithsonian Institution Annual Report 
(Mohr 1883a 

Smith (1901b) states that during the summer of 1876 Mohr examined the 
gold reserves of the metamorphic region of Alabama, and that the floristic 
notes that he took during this study were published in Berney’s Hand Book. 


Smith further states that the collection of minerals from this trip was ex- 
hibited in Mobile in 1876 and in Atlanta in 1881, and that the ieee 
was eventually sent to the U. S$. Department of eis A report by 
Mohr on the economic geology of Alabama was issued in 1878. 

Mohr’s earliest contributions to North American botany involved collec- 
tions of mosses and ferns. As mentioned previously, Mohr met the bryologist 
Leo Lesquereux while at Louisville, and Scribner (1893) states that Mohr 
“greatly assisted Lesquereux and James by furnishing material for their work 
on the mosses of North America.” Smith (1901b, c) adds that Mohr’s col- 
lection of ferns of southern Alabama was sent to Daniel C. Eaton and incor- 
porated into his Ferns of North America. Mohr also published ae short 
papers on bryology in the Bulletin of the Torrey Botanical Club in 1874. 
Although he published no more on the subject, Mohr’s interest in ena 
must have continued throughout his life, as F. S. Earle read a paper by Mohr 
on the mosses of Alabama to the 1899 Columbus botanical meetings (Anony- 
mous 1899). 

Mohr’s interests always leaned toward the practical side of science, and 
these interests brought him to perform numerous investigations for the 
Grange on methods of improving the soils of Alabama. These investigations 
included the assessment of the value of various types of wood ashes, pine 
straw, leaves, and cotton seed hulls as soil builders. The results of these 
experiments were never published. 

Another primary interest of Mohr’s was the examination of plants intro- 
duced to North America on ships’ ballast. By living in the international port 
of Mobile, Mohr was able to comb the ballast grounds for newly introduced 
species. His preliminary findings were published in the Botanical Gazette 
in 1878, and many detailed notations of plant introductions and dispersals 
are found in Plant Life of Alabama. Many plants collected by Mohr and de- 
posited in the Mohr Herbarium are simply marked “ballast ground” and 
include the date of their first introduction. 

Botany, however, was a recreational activity for Mohr, as his training was 
that of a chemist and his business was based on drugs. He combined his 
vocation and avocation in a number of articles in the Proceedings of the 
American Pharmaceutical Association, as well as in the German Pharma- 
ceutische Rundschau. His “Medicinal Plants of Alabama” was published 
both in English and in German (Mohr 1890a, b). His knowledge of medic- 
inal plants was well known, and in 1897 Mohr was appointed to the 
subcommission of the Pan-American Medical Congress to study the Kener 
can medicinal flora (Anonymous 1897). 

A quick glance at Mohr’s publications indicates that his interests lay pri- 
marily in forestry and forest products. These interests led to his many popular 
articles published in Garden and Forest and in the Rundschau, as well as 

dresses to various meetings, such as the American Cotton Planters Asso- 
ciation (Mohr 1883c). He was in charge of several natural history exhibi- 


6 


tions at regional expositions, including the New Orleans exposition of 1884. 
Mohr (1883b) published a list of the natural resources displayed at the 
Louisville exhibition for the Louisville and Nashville Railroad which Scrib- 
ner (1893) described as “one of the few papers of its kind which possesses 
real scientific merit.” 

In 1880, Mohr was contacted by Charles Sprague Sargent to investigate 
the forests of the Gulf states. Although he expressed concern about receiving 
credit for his work (Mohr 1882a), Mohr’s results were published in the 
ninth volume of the Tenth United States Census. This work marked the be- 
ginning of a long association with the Department of Agriculture, which 
culminated in the writing of a series of extensive monographs on southern 
trees of economic importance. One such report (Mohr 1896a) treated the 
southern pines and included a study on wood anatomy by another author. 
“Notes on the Red Cedar’ was published posthumously. Smith (1901b) 
reported that at the time of Mohr’s death the monographs on Bald Cypress 
and Juniper were also in press, and that one on the oaks had been completed, 
but none of these papers has ever been published. 

Mohr was a member of many pharmaceutical and scientific associations, 
and he played an active role in most of them. As enumerated by Tracy 
(1901), Mohr was an honorary member of the state pharmaceutical asso- 
ciations of both Ohio and Louisiana, and ts listed as a member of the Ameri- 
can Pharmaceutical Association from 1871 onward (Anonymous 1885). 
Mohr's letters indicate his participation in two revisions of the Pharma- 
copeia (Mohr 1890c, 1900a). As for botanical associations, Mohr was a 
corresponding member of the Philadelphia Academy of Natural Sciences, 
the Massachusetts Horticultural Society, and the Torrey Botanical Club, and 
a fellow of the American Association for the Advancement of Science. He 
is known to have presented papers to the 1891 and 1892 meetings of the 
latter Association (Anonymous 1891, 189 

Mohr’s most active participation, however, was in the field of forestry, 
and especially as a member of the American Forestry Congress. At the 
Cincinnati meeting in April, 1882, he presented a paper, “The Southern Pine,” 
and was appointed to chair a committee “to report upon Forest Fires, and 
the Injuries to Forests by Cattle’ (Anonymous 1882a). At the Montreal 
meeting in August, 1882, he was appointed temporary treasurer and a mem- 
ber of three committees, and also presented a paper (Anonymous 1882b). 

Through his many activities, Mohr was able to meet and maintain contact 
with many of the prominent scientists of his day. He corresponded frequently 
with N. L. Britton, largely concerning questions of nomenclature and the 
identification of sedges, and these letters can be found in the archives of 
the New York Botanical Garden. The Smithsonian Institution Archives 
contain his many letters to G. R. Vasey, J. N. Rose, S. M. Tracy, and C. L. 
Pollard. In letters to Smith, Mohr (1882b, c) mentioned meeting A. W. 
Chapman while working in Washington and meeting George Engelmann 


5 


while staying with Sargent in Massachusetts. His correspondence and con- 
sultation with Chapman continued until the latter’s death and is related in 
a biographical sketch of Chapman (Mohr 1899a). In a different letter, 
Mohr (1898) mentioned that Gifford Pinchot hoped to meet him, and his 
association with another prominent forester, George B. Sudworth, is described 
in an earlier publication (Mohr 18922). 


PUBLISHED WORKS OF CHARLES MOHR 
The following is a list of the published works of Charles Mohr. Fifty-five 
entries are listed, and, although it is hoped to be complete, additional works 
may be found, especially in German journals. This list does not include re- 
printings or re-editions, nor does it include the many abstracts listed by 
Harper (1928). Journal title abbreviations follow Lawrence et al. (1968). 
An asterisk (*) indicates a work that I have not seen. 


mdeinons to the bryology of the ae States, I-V. 1874. Bull. Torrey Bot. Club 
-8, 12-13, 34-35, 49-50, 50- 
On Pynanthemum lintfolinm ae its ial constituents. 1877. Proc. Amer. Pharm. 
Assoc. 24:5 


Foreign ies edad into the Gulf states. 1878. Bot. Gaz. (Crawfordsville) 
3 :42-46. 


*Economic geology of Alabama: report on the minerals of Alabama, exhibited at the 
se annual fair of the Mobile agricultural, horticultural, and mechanical fair 
. 1878. Gulf ee 1: 1-537, 
The rte of Alabama, and their products. 1878. Pages 221-235 im Saffold Berney, 
ed. Hand book of Aiea. va Register Press, Mobile, Ala. 338 
The grasses, and other forage plants of Alabama: i een naturalized, nd culti- 
vated. 1878. Pages 236-247 in Saffold Berney, ed. Hand book of Alabama. Mobile 
Register Press, Mobile, Ala. 338 pp 
Preliminary list of the plants growing without cultivation in Alabama, from the 
collections made by Eugene A. Smith, Tuscaloosa, and Chas. Mohr, Mobile, Ala. 


Analysis of Eriodictyon See £880: Proc. Amer. Pharm. Assoc. 27:736-740. 

Rhus cotinoides, Nutt. 1882. Proc. Acad. Nat. Sci. Phil ae 1882:217-220. 

On the distribution of the more ee forest trees in the Gulf region. 1882. 
Amer. J. Forest. 1:78-88, 120-126. 

On distribution of the more oo forest trees in the Gulf region. 1883. 

J. Forest. 1:179-184, 200-2 

ees lt ee ay and aan ov in Alabama. 1883. Annual Rep. Smith- 
sonian Inst., 1881:617 

List of trees ee plants characters of each region of the state. 1883. Rep. Alabama 
Geol. Surv., 1881-82 :291-2 


The natural resources of hn displayed in the exhibit of the Louisville and 
Nashville Railroad Company, at the ee Exposition, Louisville, Kentucky. 
1883. Daily Register Press, Mobile, Ala. 22 

On Quercus durandii, Buckley. 1883. Proc. ee ar Sci. Philadelphia, 1883 :37-38. 

The lands of the Louisville and Nashville Railroad in Alabama 
the scttler. 1884. Roberts and Son, Birmingham, Ala. 16 pp. 

*Rare and little known trees and shrubs of Alabama. 1884. Trans. Mississippi Valley 

ort. Soc. 2:216-219. 
*Ueber die eee der terpentin liefernden pinusarten im suden der vereinigten 


as homesteads for 


staaten und ueber die gewinnung und Gece ue terpentin. 1884. Pharm. 
Rundschau (Berlin & New York) 2:163-166, 187- 

The forests in the vicinity of Mobile. 1885. Pages ns a in Sub Rosa [pseudonym 
for Paul Ravesies}, ed. Scenes and settlers of Alabama. No publisher. 120 

ae of forest ae — exas, Louisiana, Mississippi, Alabama, and Floridal: 

885. Pages 66-67 . V. Riley, ed. Fourth report of the United States Ento- 

eens jeans 

*Ueber drei vercinzelte burger ie — see der nordamerikanischen sudstaaten. 
1887. Pharm. Rundschau (Berlin & New York) 5:8- 

*Untersuchung der blatter von See triacanthos, L. 1887. Pharm. Rundschau 
Berlin & New York) 5:250. 

fae of the United States. 1887. Pap. New Orleans Acad. Sci. 1:19-39. 

Winter in Mobile. 1888. Gard. & Forest 1:16. 

The ea forests of the South. 1888. Gard. & Forest 1:34-35. 

Spring in Mobile. 1888. Gard. & Forest 1:88. 

The long-leaved pine. 1888. Gard. & ae 1:261- 

*Ueber die verbreitung der pflanzen durch thiere. ee Pharm. Rundschau (Berlin 
& New ae 6:177-181. 

The pecan tree (Hickoria pecan). 1889. Gard. & Forest 2:569-570 

The lz vTee}st aaduce to the shrubs of eastern North America. 1889. Gard. & Forest 


The medicinal plants of Alabama: systematic list of the medicinal plants occurring 
within the limits of the state, with notes on their distribution and oe ume 
of collecting the parts used. 1890. Mo - pe Press, Mobile, Ala. 17 

Vegetation in southern Alabama. 1890. . & Forest 3:140, 212 

Pinus glabra. 1890. Gard. & Forest 3.295. 

The Florida spruc e pine, 1890. Gard. & Forest 3:402-403. 

* Hickoria olivaeformis. 1890. Pharm. Rundschau (Berlin & New York) 8:56-59. 

*Die medicinischen pflanzen von Alabama. 1890. Pharm. cere (Berlin & New 
York) 8:240-243, 257-262. 

*Vegetation of Louisiana and adjoining regions, and its products, in relation to 
een and allied siddnstiies, 1891. Pharm. Rundschau (Berlin & New York) 
9:132-135, 


O= 


The general features of the vegetation of Louisiana and adjoining region, and its 
products in relation to pharmacy and allied industries. 1892. Proc. Amer. Pharm. 
Assoc. 39:76-84 

Our remaining hard-wood resources. 1892. Engineering Mag. 4:378-385. 

Variation in the leaves of Clematis reticulata and other notes. 1892. Bull. Torrey 
Bot. Club 19:308-3 

*Die gebirgsflo ora Mabe 1892. Pharm. Rundschau (Berlin & New York) 10:253. 

The mountain flora of Alabama. 1892. Gard. & Forest 5:507-508 

The distribution of some forest trees in the southern states. 1893. Gard. & Forest 
6:372-373. 

*Die walder des sudlichen Alabamas. 1894. Pharm. Rundschau (Berlin & New York) 

21 

*Die walder det alluvial region des Mississippi in den staaten eee Lo 
und Arkansas. 1895. Pharm. Rundschau (Berlin & New York) 13:1 

*Die walder des sudlichen Alabamas. 1895. Pharm. Rundschau (Berlin & eae York) 


jon 


*Ueber das vorkommen des balsams von Liguidambar styraciflua L. 1895. Pharm. 


The timber pines of the southern United States: with an introduction by B. 
Fernow, and a discussion of the structure of their wood by Filiberr Roth. U.S.D.A. 
Div. Forest. Bull. 13. 160 pp. 


y 


Notes on some undescribed and little known plants of the Alabama flora. 1897. 
Bull. Torrey Bot. Club 24:19-28. 

Report on the forests of Sand Mountain: 1898. The Forester 4:211-215. 

Notes on some ae ee little known plants of the Alabama flora. 1899. Bull. Torrey 
Bot. Club 26:11 

Alvin Wentworth ean 1899. Bot. Gaz. (Crawfordsville) 27:473-478 

Plant life of Alabama: an account of the distribution, modes of association, and 
adaptations of the flora of Alabama, together with a systematic catalogue of the 

#12921. 


Plant life of Alabama. 1901. Alabama Geol. Surv. Monogr. 5. 921 pp. 
*Notes on the red cedar. 1901. U.S.D.A. Div. Forest. Bull. 31. 37 pp. 


PLANT LIFE OF ALABAMA 


Mohr's major scientific work was Plant Life of Alabama, published July 
31, 1901—just two weeks after his death. This work was pro Hee with the 
assistance of the Geological Survey of Alabama and its head, Eugene Allen 
Smith, and represents an elaboration of the “Preliminary List’ from the 
collections of Mohr and Smith (Mohr 1880). 

Letters exchanged between Mohr and Smith fail to indicate when the deci- 
sion was made to begin work on a complete flora of Alabama. However, in 
1882, after completion of his field work for the Census, Mohr (1882d) 
wanted “to be left quietly at home’ until he had completed “our Alabama 
Flora.” Four years later, Mohr (1886) mentioned working on the Flora, and 
by 1889 he was ready to add the lower plants to it (Mohr 1889). 

Concentrated work on the project was not begun until 1891, when ap- 
proval of funding was granted by the governor of Alabama (Mohr 1891). 
Under the terms agreed upon, Mohr received $50 per month as a retainer, 
with the total amounting to $5 per finished page "eh: 1892b, 1896b). 
Mohr was able to devote most of his time to the Flora after his retirement 
from the pharmacy business in 1892, although commitments to the U.S.D.A. 
had to be met also. In 1896, Mohr asked for and was granted a temporary 
reprieve from his Department of Agriculture duties (Mohr 189Ge). 

Work on the Flora accelerated in 1896. In that year a botanist at Auburn, 
Alabama—P. H. Mell—published a bulletin describing the plants of Alabama 
(Mell 1896). In this work, Mell utilized data obtained from Mohr. The 
Botanical Gazette (Anonymous 1896a) accused Mell of undermining Mohr’s 
work, stating that “we cannot believe that this extensive information was 
obtained from our good friend with the expressed intention of anticipating 
his own flora. In other words, Dr. Mohr must have granted a favor that has 
been abused.” Although a later article (Anonymous 1896b) in that journal 
exonerated Mell from blame, a series of letters exchanged between Mohr 
and Smith indicate that suspicions of Mell’s abuse were valid (Mohr 1896c; 
Smith 1896b). Mohr soon after decided to treat the entire Mell affair with 
“silent contempt” (Mohr 1896d). 

Later in 1896, Mohr indicated additional pressure to complete and publish 
his work from such botanists as John K. Small and Thomas H. Kearney. 


10 


He stated (Mohr 1896f): “By the activity with which of late the younger 
northern botanists begin to explore these parts, they are apt to take the 
wind out of my sails on many points.” For this reason he decided to quickly 
publish his “novelties” in the Bulletin of the Torrey Botanical Club. He 
was further dismayed by the publication of Britton and Brown's Illustrated 
Flora, as he felt that it embraced a large number of southern plants (Mohr 
189G6h). 

Although Mohr relied heavily upon his own experience, knowledge, and 
herbarium in writing Plant Life of Alabama, many other experts and col- 
lections were consulted. An updated catalogue of the plants contained in the 
Geological Survey collection was used as a working model, and this catalogue 
was sent to N. L. Britton for revision to fit the Rochester rules of nomencla- 
ture (Mohr 1893a). Mrs. N. L. Britton asked to examine the portions 
of the manuscript concerning the mosses (Mohr 1899b). Specimens of the 
genus Aster were sent to Prof. Burgess in New York, while the grasses and 
Silphium were shipped to Columbia University for verification (Mohr 1897a, 
b). Mohr conversed with C. H. Merriam on plant distributions within the 
state (Mohr 1894). The Engelmann Herbarium at St. Louis was consulted, 
as well as the herbarium and libraries at Cambridge, Massachusetts (Mohr, 
1893b, 1895). 

Mohr constantly incorporated information from the most recent publica- 
tions, even those published while Plant Life was in press (Mohr 1901a). 
In a letter to Tracy, Mohr ( listed his last-minute revisions and 
consultation of the type material at the National | Herbarium, with all changes 
made at “no matter what sacrifice of time and trouble. 

Plant Life of Alabama was originally planned to be one of the publications 
of the Geological Survey of Alabama, and Mohr was partially supported by 
Survey funds from 1892 on. However, it became apparent to Smith that 
publication by the Survey would be far too costly and might necessitate the 
elimination of large parts of the work. He suggested that a publication 
arrangement be made with the Smithsonian Institution or with the Depart- 

ment of Agriculture, and that an extra edition with an altered title page be 
printed as an Alabama state document (Smith 1896a, c). Such an agreement 
was reached to print the work as a Contribution from the U. S. National 
Herbarium (Mohr 1896g). 

Publication by the U. S. National Herbarium, however, resulted in a great 
number of problems. In Smith’s opinion, the cost of publication became 
exorbitant, and he expressed regret at having the work published in Wash- 
ington (Smith 1900). Although Mohr was informed in July, 1897, that 
the Printing Office was ready for his manuscript, the work was not published 
until four years—as well as Mohr’s life—had elapsed. Mohr (1900c) could 
rightfully claim “to have been among the first (if not te first) to treat the 
flora east of the Rockies in respect to its ecological relations,” but delays in 
printing his work allowed others to assume this distinction. 


ll 


In its final form, Plant Life of Alabama is an expanded checklist of all 
plants known to occur in the state. Each species or variety of plant, from 
slime mold to angiosperm, is listed with a bibliographic citation, important 
synonyms, distribution, occurrence in the state, type locality, and the loca- 
tion of representative specimens. A tabular summary of the plants follows the 
checklist, as well as a list of cultivated species. 

Plant Life of Alabama was lauded as “the most One and philosophical 
local flora ever published...” (Tracy 1901). It is more than a mere check- 
list of plants, because the first 137 pages are oe These pages include 
accounts of the history of Alabama botany, physiographic features of the 
state, plant distributions and associations, the relation of the Alabama flora 
to other floras, and a discussion of introduced plants and their influences. 
The work thus includes sections covering most of Mohr’s interests. 

Plant Life of Alabama was planned to be the first of a two-volume work 
on the botanical resources of Alabama, with the second volume dedicated to 
the economic plants (Smith 1901la). While the economic volume was viewed 
as being more immediately valuable to the populace, it was felt that the 
systematic volume was needed first (Smith 1896a). 

The earliest mention of an Economic Flora of Alabama was made in 1887, 
while the first remittance for such work was forwarded in 1899 (Mohr 1887; 
Smith 1899). The first 350 pages of the work were to be devoted to trees 
and shrubs, with a sketch of forest botany, while short descriptions of plant 
families would be included to allow the work to be used as a systematic 
botany textbook (Mohr 1901b, c 

It is generally believed that Mohr’s work on an Economic Flora was prog- 
ressing well at the time of his death, although a search of Mohtr’s belongings 
by his son, Herman B. Mohr, failed to locate the manuscript. His son later 
implied that, due to his father’s failing mental faculties, actual work on the 
Economic Flora may never have begun (H. B. Mohr 1901a, b). If a manu- 
script was produced, it is possible that Harper used it as a basis for his very 
similar work (Harper 1928). However, an examination of the extensive 
Harper material at The University of Alabama, Gorgas Library, Special Col- 
lections failed to turn up Mohr’s manuscript. 


ACKNOWLEDGMENTS 


I wish to thank Dr. C. Earle Smith, Jr. and Dr. Robert R. Haynes for their 
help and encouragement in the completion of this project. 


REFERENCES 
a hela 1882a. The American Forestry Congress: Cincinnati meeting. Amer. 
J. Forest. 1:31- 
1882b. The American Forestry Congress: Montreal meeting. Amer. J. 
~ Forest. 1:64-68. 
1885. Roll of aN — a Pharm. Assoc. 32:561—-583. 
1891. Botanical club o . S. Bot. Gaz. (Crawfordsville) 16: 


a 


1892. Papers presented to the botanical club of the A. A. A. S. Bot. Gaz. 
(Crawfordsville) 17: ak Rg 


_, «1896a. New t. Gaz ules her 29 

E ; 1896b. Onn ene Bot. (Cr forse) 22 417-421. 

_ «1897. News. Bot. Gaz. ek omen 24:13 44, 

- 1899, A synopsis . proceedings of the bocanical organizations meet- 
ing | at Columbus, Ohio, August 17- 25, 1899. Pall tere Bot. Club 26:500-505. 

HARPER, R. M. 1928. fie botany of Alabama, part 2: catalogue of the trees, 


shrubs, and vines of Alabama, ise Ae economic neat and local distribu- 
tion. Alabama Geol. Surv. Monc Q.. 4 

HOFFMANN, F. 1887. Karl ae Mohe: ‘eine biographische skizze. Pharm. 
Rundschau (Berlin ae oe 5 :4-— 

LAWRENCE, G. H. SGC ERe G. S. DANIELS, and H. DOLEZAL, 
eds. ane Boni perce Huntianum. Hunt Botanical Library, Pittsburgh, 
Penna. 


)63 p 
MELL, P. 1896. The flora of Alabama, part 5. Alabama Agric. Exp. Stat. Agric. 


MOHR, C. T. 1879. er to E. A. Smith nas Mobile, Ala., 21 May 
- 1880. ice list of the plants growing ae een in Ala- 
bama, from the collections made by Eu ae A. aes Tuscaloosa, and Chas. Mohr, 
Mobile — No publisher a 


_.. 1882a. fe to E. A. Smith from Mobile, Ala., 30 ae 
_____. 1882b. Letter to 5 A. Smith from a D.C. 5 Jul. 
as . 1882c. Letter to E. A ree ith from Geor n, D. S 11 re 
—_.. 1882d. Letter to E. A. Smith fro Pave in omery, Ala., 21 O 
t: 1883a. Aenea soapstone quarry and shell-heaps ‘in Nise Annual 
Rep. Smithsonian eae 1881 :617- 
83b. The natural resources ‘of Alabama, displayed in the exhibit of the 
Louisville ma Nashville eee See at oe Southern Exposition, Louisville, 
ucky. Daily 


Register Press, Mobile 
eeene eT 1883c. Letter to E. A Smith fom ‘Mobile, Ala, 19 a 
= __. 1886. Letter to E. A. Smith fro Mobile, Ala., 26 S 


- —. 1887. Letter to E. A. Smith oan Mobile, Ala., 26 May 
_ . 1889. Letter to E. A. Smith from Mobile, Ala. 
1890a. The medicinal plans of pagers Die list of the medicinal 

plants occurring within the limits of the state, with es on their distribution 
proper ime of collecting the nee used. Mobile bovine Press, Mobile, Ala. 17 

_ ie medicinischen pflanzen von Alabama. Pharm. Rundschau (Be. 
“din & New See 8:240-243, 257-262. 

_ . 1890c. oe a E. A. ae from Mobile, 27 a 

: . ey Let o E. A. Smith from Mobile, iG 19 O 

7 892a. Variation in the ae oe Clematis reticulata a other notes. Bull. 
Torrey | on Club 19 :308—30 

- 1892b. Letter to E. A. van se Mobile, oe 2 May. 

a . 1893a. nes to E. A. Sm Mobile 28 May. 

a _. 1893b. Letter to E. A. Sm | om event Sea 6 Oct. 

1894, ee to E. . Smith from Mobile, ‘Ala., 2 Oct. 

________. 1895. Letter to E. A. Smith from Mobile, Ala., 22 Aug. 

= 1896a. The timber oe es of the southern United States: with an int 
duction ‘by B. E. Fernow, and a ere of the structure of their woods by Fili bert 


Div. Forest. Bull. 13. O pp. 

—_. 1896b. Letter to E. A. Sm th from Mobile, Ala., 4 Jan. 
189G6c. Letter to E. A. Smith from Mobile, Ala, 17 Apr 
1896d. Letter to E. A. Smith from Mobile, Ala., 23 Apr 

_, «189G6e. Letter to E. A. Smith fro obile, , 1O May 

_ 1896f. Letter to E. A. Smith from Mobile, Ala., 27 Oct 

_.. 1896g. Letter to E. A. Smith from Mobile, Ala., 19 Dec. 

«18 96h. te - E. A. Sm ie from Mobile, Ala., ur 
. 1897a. L A. Sm bile, oe la., 


anes _ o E. from Mo 2 
a . 1897b. ee . E. A. Smith ane Mobile, Ala., 6 TL 


13 


________. 1898. Letter to E. A. Smith from Mobile, Ala., 28 Jul. 
—_____. 1899a. Alvin Wentworth Chapman. Bot. Gaz. (Crawfordsville) 27:473- 
478. 


Seo 1899b. Letter to Mrs. N. L. Britton from Mobile, Ala., 27 Jan. Archives, 
New y York Botanical Garden. 
__,s«1900a. Letter to E. ‘A. Smith — Asheville, N. C., 25 Apr 
—_____. 1900b. Letter to S$. M. Tracy from Asheville, N. ve Jun. Archives, 
Smithsonian Aon ae Mills ag Papers, 1895-1914 
_.,s«21900c. Letter to E. A. Smith from ene N. C., 7 Oct 
__. «1901a. Letter to E. A. ae from Asheville, N. C., 22 Jan. 
___. 1901b. Letter to E. A. Smith from As Reville , N. C., 1 Mar. 
. 1901c. Letter to E. A. ei ne Asheville. N. C., 8 Mar 
MOHR, iL 1901a. ma to " A. Smith from Asheville, N. C 20 Ju. 
: Olb. Let o E. Smith from Asheville, N. 
1903. Dr. sae Mohr, the South’s oa Ties Democrat 
an. 
Fan Onae a Vs 1055. The Biltmore story. Minnesota Historical Society, St. Paul. 


22 
SCRIBNER, F. L. 1893. Southern botanists. Bull. Torrey Bot. Club 20:315—-334. 
SMITH, E. A. 1896a. Letter to C. T. Mohr from University, Ala., 2 : 
_______. 1896b. Letter to C. T. Mo . ae Univ ersity, Ala., 19 Apr. 


ate . Moh m Uni Jul. 
________. 1899. Letter to C. 'T. Mohr fees Sas Ala., 15 Nov. 
«1900. Letter to C. T. Mohr from University, Ala., 5 Apr 

1901a. Letter of aie Pages iii-iv in C. T. MGRE: Dian life of Ala- 


~ bama. Alabama Geol. Surv. Monogr. 5. 921 pp. 
Se ee Olb. Charles Theodor ore eee Pages v-xii 7 C. T. Mohr. Plant life of 
Alabama. Barca Geol. Surv. Mon 292 


= 
a Ae C. Biosenial oe “Of ‘Dr. Charles Mohr. Bull. Torrey Bot. Club 
28:59 604. 


oe 
TRACY, S. M. 1901. Dr. Charles Mohr. Pl. World 4:167-170. 


TAXONOMY AND DISTRIBUTION OF 
GENTIANA (GENTIANACEAE) IN MEXICO 
AND CENTRAL AMERICA. I. SECT 
CHONDROPHYLLAE 


JAMES S. PRINGLE 
Royal Botanical Gardens, Box 399, Hamilton, Ontario, Canada L8N 3H8 


This paper is the second of two monographs on Gentiana L. in Mexico 
and Central America. Delineation of the genus, terms pertaining to its 
morphology, and other introductory topics were discussed in the first paper 
(Pringle 1977), which also included a key to the sections of Gentiana in 
these regions. 


GENTIANA {sect.} CHONDROPHYLLAE Bunge, Nouv. Mém. Soc. Imp. Nat- 
uralistes Moscou 1:207. 1829. Lectotype species (Pringle 1978): Gentiana 
aquatica L. 


Sage Philippi, Florula Atacamensis p. 35. 1860. Type species (only species 
ted): Gentiana podocarpa Vee ae Griseb., as Varasia podocarpa Philippi 

Chondrophl (Bunge) A. Nels., Bull. Torrey Bot. Club 31:245. 1904 ar 
n.). 


Holubla Live & Live, Anales Inst. . Cavanilles 32:226. 1975. Type species: 
Sb pyrenaica L., as Holi ee (L.) Love & Léve. Non Holubia 

Hooker’s Icon. Pl. 15:59. 1884. 

Poesia: Live & Léve, Bot. se 131:385. 1978. Based on Holubia Live & 
Live, non Oliv 


© 


Plants monocarpic or perennial. Roots slender, branched; taproot persistent 
in annual species. Stems solitary or clustered, usually branched except in 
minute plants, decumbent to erect, slender, in many species less than 15 cm 
long, up to 30 cm in others. Basal and lower cauline leaves of most mono- 
carpic and some perennial species persistent, crowded, and broader than me- 
dian and upper cauline leaves. Leaf bases strongly connate-sheathing. Blades 
usually less than 1.5 cm long, scalelike to oblanceolate, oblong, or orbicular, 
usually carinate. Margins obscurely to prominently white-cartilaginous, en- 
tire or minutely denticulate. Flowers solitary or in nee condensed cymes, 
without involucral bracts, 5-30 mm long. Calyx tu or 15-veined, 
cylindric to funnelform, uncleft. Calyx lobes deltoid to oblong often carinate, 
with cartilaginous margins. Intracalycular membrane discontinuous or not 
developed. Corollas usually pale to deep blue, sometimes white, pale yellow, 
or violet, usually suffused externally with green and/or deep violet, often 


‘Contribution No. 31 from the Royal Botanical Gardens, Hamilton, Ontario, 
Canada 


~ SIDA 8(1): 14-33. 1979. 


ihe; 


photo-, thigmo-, and seismonastically closing. Corolla tubes cylindric to 
funnelform. Corolla lobes abruptly wide-spreading, shorter than tube, deltoid 
to ovate or oblong-ovate. Free portions of corolla appendages nearly sym- 
metrical, uncleft to shallowly or deeply bifid, or erose to fimbriate. Stamens 
equal, with straight filaments. Anthers free or loosely connate. Pollen grains 
oblate-spheroidal to prolate. Sexine striate, striato-reticulate, or nearly smooth. 
Ovaries compressed-ellipsoid, usually widest near middle, abruptly contracted 
at summit. Stigmas short, in most species sessile or subsessile, in some ele- 
vated by a well-developed style. Gynophores of some species greatly elongat- 
ing in fruit, remaining short in others. Carpels each with 1 dorsal and 
ventral veins. Ovules in 4 parietal zones corresponding to positions of ventral 
carpel veins. Seens ovoid to ellipsoid, often apiculate, wingless or with nar- 
row, incomplete wings. Testa striato-reticulate. Sectional description com- 
piled from Kusnezow (1895), Lindsey (1940), Nilsson (1967), Toyokuni 
(1963), and my own observations. 

Section Chondrophyllae occurs in the Pyrenees, the Alps, and the Carpa- 
thians; in the Caucasus, the Himalayas, Siberia, China, Japan, and Southeast 
Asia; in Indonesia, New Guinea, and the Philippines; from the Aleutians 
and the Brooks Range in Alaska through the Rocky Mountains to Arizona; 
in Mexico, Guatemala, and Costa Rica; and in the Andes from Venezuela 
to Tierra del Fuego. Throughout its range, this section is confined largely to 
mountainous regions, with many of its species occurring above treeline. 

The pollination of G. prostrata Haenke in Europe has been discussed by 
Kerner von Marilaun (1891), who concluded that although outcrossing 
was promoted by protandry, autogamy could readily occur within corollas 
prevented from opening by prolonged cloudy periods. Similarities in corolla 
morphology and in pistil and anther development indicate that these con- 
clustons are generally applicable to the species discussed in the present paper. 
The widespread phenomenon of thigmonastic corolla-closing in this section 
is usually assumed to be related to pollination, but its specific role has not 
been investigated. 

From the field aspect of open capsules of G. perpusilla Brandegee, i 
appears that the fruits of this and some other species of sect. Chondrophyllae 
function as splash-cups. In species with more elongate gynophores, seeds are 
probably dispersed as wind shakes the fruits (censer mechanism). Many 
species grow where rill action can further disperse the seeds after they have 
fallen. Long-distance dispersal evidently has also occurred, since the alpine 
habitats of the Mexican and Central American species are widely separated, 
are mostly of late Tertiary or See origin, and have no history of con- 
nection by continuous alpine vegetation 

Since its establishment by Bunge, sect. Chondrophyllae has been accepted 
as a section of Gentiana by most authors, except for those who restricted 
Gentiana to the five species usually Gare as its nominate section. Varasia 
Philippi and Chondrophylla (Bunge) A. Nels. were ae ae tO: SeGr. 


16 


Chondrophyllae only. Neither of these names has been used recently. 

Love & Léve (1975) divided sect. Chondrophyllae between two segre- 
gate genera. The “annual or biennial species with x = 9” were placed, 
along with sect. Crmnalis (Adans.) Dumort. (which consists of perennial 
species), in the genus Ciminalis Adans. emend. Holub (type species: 
Gentiana acaulis L., in sect. Ciminalis, as Ciminalis [sp.]). The “perennial 
species... with ...x = 13,” believed by Love & Love to comprise “about 
ten species of Eurasia, North and South iar ” formed Holubia Love & 
Love, later renamed Holucogentia Love Love. Weaver & Riidenberg 

1975), however, reported 2 = 20 1 Dee called G. sedifolia, but prob- 
ably an unnamed, closely related species, in Venezuela. In Love & Love's 
generic concept, this species, perhaps with certain Asiatic species having 
n = 10, will presumably be treated as a third genus. 

Little can now be postulated about relationships within sect. Chondro- 
phyllae in the Western Hemisphere, because its diversity has largely remained 
unrecognized, Existing information on chromosome numbers and_ pollen 
morphology indicates that phylogenetic studies will require more such data. 
Subdivisions of sect. Chondrophyllae are therefore not considered here. 

In this study, I examined specimens in the following herbaria: BM, DS, 
ENCB, F, GH, HAM, K, MEXU, MICH, MO, MSC, NY, TENN, US, and 
WIS (abbreviations from Holmgren & Keuken 1974). Among these are 
collections each comprising several to many plants, representing well the 
variation that may occur within a population. Specimens from South America, 
northern North America, and Eurasia were among those studied, and G. 
perpusilla and G. prostrata were observed in the field. Consequently, it has 

een possible to compare the Mexican and Central American plants with 
those in other regions, and to distinguish, with reasonable confidence, taxo- 
nomically significant differences among populations from minor variations 
among individuals. 

All of the species discussed here have sometimes been equated with G. 
prostrata. Comparisons with specimens of G. prostrata from Colorado (Figs. 
7, 13), Alaska (Fig. 19), and the type region in Austria, however, indicate 
that all of the Mexican and Central American plants are distinct from this 
species. Gentiana fieldiana, which resembles G. prostrata in its leaf shape 
and in its apiculate corolla lobes, differs in its wider white leaf margins, in 
ie more oblong shape and strict apical symmetry of its corolla lobes, in 

e greater adnation of its stamens, and in its usually cymose aa 
ee sedifolia resembles G. prostrata in its narrow leaf ma and in 
the long free portions of its filaments, but differs in its tome much 
narrower leaves, in its blunt corolla lobes, in its more copiously and more 
irregularly erose-serrate corolla appendages, and in its coarser sexine sculp- 
turing without well-developed longitudinal striations. The other species 
differ even more sharply from G. prostrata: G. beamanii in its narrower 
leaves and in the much shorter free portions of its stamens, G. perpusilla 


i? 


in its proportionately small corolla lobes and in its uncleft, nearly entire 
corolla appendages, and G. pumilio in its proportionately much broader 
eaves, in its wide and prominently denticulate leaf margins, and in its 
spheroidal pollen grains. 


KEY TO MEXICAN AND CENTRAL AMERICAN SPECIES 


la. moe pgs sonal a a spatulate to orbicular, many with 5 or more 
prominent primary veins 2-4 lesser ones; white leaf margins conspicuous, 
ca. 03 mm wide, those a upper leaves denticulate from below middle to apex; 
flowers often sessile in small clusters; calyx tube largely concealed by leaves; 
pollen grains spheroidal 4. G. 


lb. Leaves, except sometimes near base of stem, less densely crowded and_ propor- 
tionately narrower, usually with only 1 or 3 promin nent primary veins; white leaf 


solitary at end of main stems and branches; calyx tube not concealed; pollen 
grains subprolate to prolate. 


2a. Leaves linear-oblong, usually more than 3 times as long as ee cue! to 


strongly divergent from near base; stamens becoming free in | 2/5 of 
corolla, with free portions, including anthers, 3-7 mm long; throat “of corolla 
with race color demarcation sedifolia 


2b. Leaves oblanceolate to obovate, usually less than 3 times as nie as aes if 

proportionately longer then proximally appressed-ascending, only the distal 

portion spreading; stamens becoming ae near or above mide or corolla, 

with free portions up to 3 mm long, if diverging lower, then with free 

portions up to 2 mm long and with ae appendages cee throat of 
corolla with or without color differentiation. 

3a. Stems erect except at base, with strongly ascending branches; internodes, 

except near base of stem, mostly as long as or longer than subtending leaves; 

leaf blades mostly over 5 mm long; throat of corolla with distinct tates 

2. G. beamanii 

. Stem and branches variously spreading; internodes mostly much shorter than 

subtending leaves; many leaf blades less than 5 mm long; corolla throat 


Ww 
i 


4a, White leaf margins and keel well defined, those of upper aoe distinctly 


denticulate; corolla lobes oblong- eaneualar, abruptly rounded to apiculate 
tip; free portions of Ae appendages erose-serrate; stamens cee in 
free near middle of corolla woo... cccccccccececcceeeecces sevens veeees G. freldiana 


dX 
> 


. White leaf margins aad ke el “obscure, generally entire or ee Howly un- 
dulate; corolla lobes ovate-triangular, tapering gradually to subacute - 
free portions of corolla appendages entire or nearly so; stamens becom 
free below middle of corolla 5. G. See 


TAXONOMIC TREATMENT 


lis Sanit SEDIFOLIA H.B.K., Nov. Gen. Sp. - ae ae need 

ext). Type: without locality [probably ECUADOR: Napo or Pichincha}: 

Humboldt & Bonpland s.n., no date [probably ne Jun 1802] ee 
P, photo MSC! isotype K! ). Figs. 1, 8, 14. 


Gentiana cespitosa Willd. ex Schultes, Syst. Veg. 6:185. 1820. Type: Duplicate 
Humboldt et al., cited above (B-herb. Willd.). (Ex char. and fide Kunth in 


Humboldt et al., 1823.) 
Gentiana ebineb opacenars Willd. ex Schultes, Veg. 6:185. 1820. Type: 
SUADOR: Chimborazo: Chimborazo, Seen et al, s.m., no date (B-herb. 
Willd.). (Ex char.) 

Ericala sedifolia (H.B.K.) G. Don, Gen. Hist. A: ne “1838” [1837]. 

As Seak edifolta var. casapaltensis J. Ball, J. L Soc., Bot. 22:49. 1885 
PE ima: supra Casapalta [Casapalca], Ball m,, 22 Apr 1882 (holotype 
K! isotype GH!). 

Gentiana sedifolia var. [3 grandiflora Kusn., Trudy S.-Peterburgsk. Obshch. 
Fstestvoisp., Otd. Bot. 24(2):217. 1894. Syntypes: ECUADOR: Napo_ or 
Pichincha: Antisana, Lehmann, no. and date not cited (LE) and COLOMBIA: 
Magdalena: Santa Marta, Lehmann, no. and date not cited (LE); probable 
isosyntype, “Cordillera of Ecuador and Colombia,” Lehmann 623, 22 Dec 1880 
(K! ) 

Plants usually 1.3—4 cm tall, occasionally up to 6 cm. Stems 1-many. Well- 
developed stems decumbent at least at base and often for much of length; 
longer stems usually much branched. Leaves linear-oblong, 5-10 mm long, 
1.5-2.5(-3) mm wide, apiculate. White leaf margins inconspicuous and 
sometimes not sharply defined, mostly 0.02-0.06 mm wide distally on lower 
leaves, ca. 0.1 mm wide on upper leaves, entire or irregularly and minutely 
denticulate toward apex. Basal leaves persistent, crowded, less conduplicate 
than cauline leaves. Cauline leaves increasingly conduplicate and sometimes 
more or less arcuate On upper part of stem (except on very small plants), 
with midrib keels more distinctly developed (white portions up to 0.05 mm 
wide). Median and upper internodes often scarcely longer than sheathing 
leaf bases, sometimes up to 2—6 mm long. Calyx tube 5.5-8 mm long, 
ridged distally along median sepal veins. Calyx lobes ovate-triangular, erect 
proximally, sometimes more or less arcuate distally, conduplicate and nar- 
rowly carinate toward apex, (1.2—)1.8—4 mm long, acute, with white margins 
ca. 0.2 mm wide, widening near apex, and keel up to 0.05 mm _ wide; 
margins and keel entire. Corollas 9-20 mm long when closed; expanded 
limb 6-14 mm across. Corolla tube 6.5-17 mm long. Lobes broadly ovate- 
triangular, 1.8-4 mm long, as wide or slightly wider than long, obtuse to 
subacute. Free portions of appendages ca. 0.75 times as long and about as 
wide as lobes, broadly rounded to triangular, shallowly to rather deeply 
erose-serrate or shallowly several-cleft near apex, not distinctly bifid. Corolla 
tube yellowish-white; upper portion suffused with deeper yellow and often 
bordered and spotted with purplish-brown. Uppermost part of fused portion 
of corolla, lobes, and free portion of appendages pale to deep blue, with a 
narrow whitish zone next to yellowish eye, or occasionally de white 
throughout; exterior surface of lobes, except near inner margin, and adjacent 
portions of tube suffused with deep violet (scarcely or not at all in white- 
flowered forms) and also with green, especially near tips of lobes. Stamens 
becoming free at 1/4—2/5 total height of corolla; free portions (including 
anthers) 3-7 mm long. Pollen grains prolate. Sexine relatively coarsely 
striato-reticulate. Pistil tapering into style ca. 1 mm long. Capsule 3-5 mm 


on) 


ee i 2 eee arium specimens of oe spp. 
Ecuador). Fig. 2. G. beamanti aes 
rapes eo 3071, MSC, a ae large plant). 


G. sedifolia Cem io 
3899, MSC). Fig G. 


20 


long, elevated well above marcescent corolla by gynophore 15-18 mm long 
ac maturity. Seeds ellipsoid, light brown, striato-reticulate, ca. 0.85 mm long, 
mm in diameter. 


Central American specimens examined: 

COSTA RICA: Cartago (all from Cordillera de Talamanca, near Cerro de la 
Muerte): not more specifically cua Carlson 3630 (F); Ciénega “3 de Junio,” 
at km 73 SE of San José on Carretera Panamericana, Jiménez M. 1993 (F, NY), 
Jiménez M. 3372 (F), Lent 405 (NY), and Anderson 7 Mori 231 (BM, DS, F, 
WIS); near Ojo de Agua, Williams et al. 28281 (BM, F, WIS); La Trinidad, be- 
tween oe 60 and 77, Molina R. et al. 17855 (F, ed Border of aed and 
San José: Cerro de la Muerte, Hetthaus 318 (MO). San José: Cerro de Buena Vista, 
Tonduz (Pinter exped.) 3498 (US); Cerro de las Vueltas, Standley & Valerio 43640 
(F, US). PANAMA: Chiriqui: between Cerro Bine and IJtamut, Weston 10175 
(MO; first report of Gentiana in Panama). 


In Central America, G. sedifolia is known only from high elevations in 
the Cordillera de Talamanca in Costa Rica and the adjacent Cordillera Cen- 
tral in Panama (Fig. 20). It is widely distributed in the Andes of South 
America, from Col ne and Ecuador south at least to the Cordillera Central 
of southern Peru (Fi 

Herbarium data eae that G. sedifolia in Costa Rica and Panama grows 
in boggy depressions and similar wet sites in meadows from 2275 to 3335 
m elevation. Plants have been collected in flower from December through 


uly. 

In their original description of G. sedifolia, Humboldt et al. (1819) 
reported having seen this species “in montibus ignivomis Antisanae {Ecua- 
dor}, Puracé, Paramo de Almaguer [Colombia], etc., alt. 1600-1800 hex. 
[= 3072-3456 m}.” On the basis of minor geographic variations in this 
species, discussed below, it seems most likely that the type was collected on 
or near Antisana. 

Distinctive vegetative features of G. sedifolia throughout its range include 
the proportionately long-tapering leaves and the very narrow white margins. 
Distinctive floral features include the funnelform corolla tube, which widens 
more conspicuously from base to summit than the nearly cylindric corolla 
tubes of the other species described here, and the relatively low divergence 
and long free portions of the filaments. Pollen grains of G. sedifolia from 
Costa Rica closely resemble those of Ecuadorean specimens, the sexine in 
both being relatively coarsely reticulate, with poorly developed longitudinal 
striations. 

The two later species names cited above have generally been accepted 
as synonyms of G. sedrfolia, at least the first having been based on the same 
collection (Humboldt et al., 1823). Ten taxa, with 11 epithets, were de- 
scribed as varieties, subyacieries or forms of G. sedifolia during the 19th 
Century, but, except for the two cited above, their names seem to have been 
based. on specimens of other species. Because all of these names were based 
on South American plants, a thorough investigation of their status has not 


Zu 


been included in this study. More recent collections, however, have indicated 
that much of the variation in G. sedifolia occurs within populations. Con- 
spicuous variation exists in stem and internode length, but extremes can be 
found within a single population and even within a single collection, e.g., 
Couthouy in 1855 (GH), from Ecuador, and Pennell & Hazen 9996 (GH), 
from Colombia; some variation may occur among the stems of an individual 
plant. This variation, appearing to represent responses to seasonal factors 
and differences in microhabitat, is not recognized taxonomically here. Max1- 
mum flower size is greater in Ecuador than farther north, but there is 
considerable variation in all regions and no discontinuities are evident. 
Corolla color likewise varies within populations. 

Plants of G. sedifolia from the region of Antisana and Cotopaxi in Ecua- 
dor often have relatively long, much-branched prostrate stem bases, which 
appear brownish-black from the remains of old leaf bases, and from which 
adventitious roots arise. Such plants resemble the type collection and appear 
to be long-lived and probably recurrently flowering, corresponding to the 
description of G. sedifolia as perennial by Humboldt et al. (1819). Plants 
from other parts of the range of G. sedifolia generally have less well de- 
veloped prostrate stems, without blackish leaf remnants, and root systems 
comprising only the primary root and its branches. These plants appear to 
be relatively short-lived and monocarpic. Longevity and life cycles are, how- 
ever, difficult to determine in specimens from the tropics, and no year-round 
observations have been made on populations of this species. Therefore it is 
not certain whether these differences represent genetic differentiation, or 
whether climatic conditions near the Equator permit greater longevity of 
individual plants. Weberbauer (in Gilg, 1906), however, commented on the 
inadequacy of plant size and apparent longevity as bases for dividing G. 
sedifolia, noting that at a single locality in Peru plants of this species might 
exhibit all stages of intergradation from minute, unbranched, one-flowered 
forms with a single filiform root to large, much-branched, patch-forming 
plants. Therefore no variants based on apparent longevity are recognized here. 
2. GENTIANA beamanii Pringle, sp. nov. Figs. 2, 9, 15. 

plus minusve erecti, 3.5-16 cm alti, internodis vulgo longioribus quam 


Caules 
foliis. Folia oblanceolata vel anguste spachulata: 5-10 mm longa, 1.5-2 mm lata, 
proximale fere ewan et distale patentia, marginibus albis usque ad 0.05 mm 


latis, apicem versus minute denticulatis. Tubus calycis 5-8 mm longus. Lobi calycis 
triangulares, 1.8—3 Pion. acuti, marginibus albis circa 0.15 mm _ latis. Tubus 
corollae cylindricus, 8. 61 m longus. Lobi corollae late ovato-triangulares 1.5—3 


emarginatae, breviores quam lobi. Filamenta staminum sese liberantia aliquantum 
supra medium corollae totae, partibus libris antheribus inclusis circa 2.5 mm_longis. 
Pollinis grana subprolata vel prolata, sexinio striato-reticulato. Capsula 4-6 mm 
longa, in parte exserta. 

Type: GUATEMALA: aac: Sierra - los Cuchumatanes, aay ees 
N of Tojiah [Tojquia} at km 322 on Ruta Nacional 9N, Beaman 3899, 1 Aug 
1960 (holotype MSC, isotypes GH, US). Paratypes: GUATEM MALA: Huehuetenango: 


pas 


vicinity of Chémal [Xémal], Sierra de los Cuchumatanes, Steyermark 50287, 8 Aug 
19a) (i, GH). 

Plants 3.5-16 cm tall. Stems more or less erect throughout or basally 
decumbent, usually 1-6, each usually with 1-3 branches arising in basal half 
and reaching about equal height, sometimes with additional branches from 
upper axils. Leaves usually broader and more closely spaced near stem base; 
lowest leaves (sometimes shriveled at flowering time) elliptic-oblong or 
oblanceolate, 5-15 mm long, 1.8-4 mm wide, spreading or ascending, scarcely 
conduplicate. Median and upper cauline leaves oblanceolate to narrowly 
spatulate, 5-10 mm long, 15-2 mm wide, increasingly conduplicate up- 
ward; proximal portions (ca. 1/3 near middle of stem, ca. 2/3 near summit ) 
appressed-ascending, distal portions more or less arcuate-spreading. Leaf 
apices abruptly acuminate, apiculate. White leaf margins almost obsolete 
proximally, widening distally to ca. 0.08 mm, entire or sparsely and minutely 
undulate-denticulate above middle. Midrib keel of similar width, minutely 
denticulate above middle. Internodes distinct, all but lowest few usually 
1-2.5 times as long as subtending leaves. Calyx tube 5-8 mm long, slightly 
ridged along midveins. Calyx lobes triangular, 18-3 mm long, acute to 
acuminate, scarcely to moderately conduplicate, erect or slightly arcuate- 
spreading distally, with white margins ca. 0.15 mm wide, entire or nearly 
so, and keel up to ca. 0.08 mm wide, entire or sparingly denticulate. Corollas 
10-17 mm long when closed; expanded limb 6.5—12 mm across. Corolla tube 
8.5-14 mm long. Lobes broadly ovate-triangular, 1.5-3 mm long, slightly 
wider than long, subacute to acute. Free portions of appendages ca. 0.8 times 
as long and about as wide as lobes, triangular, sparingly erose-serrate, usually 
emarginate to shallowly bifid. Corolla tube mostly whitish, blue above, with 
dark purple streaks on interior surface of throat, in 2 lines in each petal 

roper and 1 in each appendage; lobes and free portions of appendages 
medium blue; exterior surface with deep violet pigment between relatively 
pale margin and overlapping = (when c ne of next lobe, giving striped 


— 
_ 
— 


orollas fred portions eer anthers) ca. 25 mm long. Pollen grains 
subprolate to prolate. Sexine relatively finely striato-reticulate. Pistil tapering 
into style ca. 1 mm long. Capsule 4-6 mm long, about 2/3 exserted from 
marcescent corolla by gynophore 7-10 mm long at maturity. Seeds not seen. 
Gentiana beamanti is known only from a small area in Los Altos 
Cuchumatanes, Guatemala (Fig. 23). Both collections cited were made along 
small streams in meadows at ae of 3200-3750 m. 
iis species is named for Dr. n H. Beaman, author of several papers 
on the alpine flora of Mexico and See whose notes and comments on 
differences among the Central American species of sect. Chondrophyllae have 
been very helpful in this study. 
1e elongate, erect stems and widely separated leaves of G. beamanii 
give this species an aspect unique among Central American species of sect. 


23 


Chondrophyllae. Its distinctness from plants treated here as G. fieldiana 
was first noted by Steyermark (in sched., Steyermark 50287), as follows: 
Corolla larger, broader, and with longer tube . . . corolla-lobes with deeper 
purple in outer lobes [sic] . . . also stems with more branching and with 
more flowers than in other smaller type.” Beaman (unpublished notes) like- 
wise contrasted these plants, noting that those here called G. beamanii had 
narrower leaves with less conspicuous white margins. 

Of the three Guatemalan species, G. beamantzi is the closest to G. sedifolia 
in floral morphology. It differs conspicuously, however, in its cylindric, 
abruptly flaring corolla tube and in the much greater adnation of its stamens. 


Figs. 4-7. Herbarium specimens — ems spp. Fig G. fieldiana ene 3071, 
MSC, a plant ca. average size). 5. G. pu Seite wie 2868, M) Fig. 6. 
G. perpusilla (Pringle aes Lae Fig. 7. G. prostrata ae Zoe MICH, 

from Colorado). Figs. 1-7 to same scale. 


24 


3. GENTIANA fieldiana Pringle, sp. nov. Figs. 3, 4, 10, 16. 


Caules ascendentes vel erecti, 1-4(—6.5) cm alti, vulgo pauciramosi. mae congesta. 
Folia basalia oblongo-oblanceolata vel elliptico-obovata, 4— = mm longa et 2-4 mm 


n 
saree marginibus albis circa 0.05 mm latis, integri . Tubus corollae cylindricus, 
4.5-10 mm longus. Lobi corollae oblongo-triangulares, ee 2 mm longi, apiculata, 
azurei aut albi. Partes librae plicarum triangulares, eroso-serratae et vulgo emarginatae, 
breviores quam lobi. Filamenta staminum sese liberantia circa medium corollae totae, 
partibus libris antheribus oS circa 2 mm longis. Pollinis grana prolata, sexinio 
striato-reticulato. Capsula 3—5 mm longa, in parte vel in toto exserta 

Type: GUATEMALA: beta etenango: Chéemal [Xémal], at km 318 on Ruta 
Nacional 9N, Beaman 3071, 4 Aug 1959 (holotype MSC). gee : GUATEMALA; 
Huchuetenango (all from Los Altos Cuchumatanes): vicinity of Tunima [Tuinima}], 
Steyermark 48368, 7 Jul 1942 (F, GH); 2.5 mi E of San ae Ixtatan, Steyermark 
49875, 31 Jul 1942 (F); near Paquix, Sharp 451025, 24 Dec 1945 (F, TENN); 
E of Tocquin {Tojquia}], Holdridge 2345, 27 Apr 1948 (US); Llano de Tierra 
Blanca, near oo to Todos Santos from Hane de San Miguel, near Chémal [Xémal], 
ca. 5 km W of km 311 on Ruta Nacional 9N, Beaman 3118, 5 Aug 1959 (MSC); 
3 km § of road ae Llano de San Miguel and Todos Santos, from a point 2.5 
mi W of Llano de San Miguel, Beaman 3975, 2 Aug 1960 (MSC); 7 mi N of 
Santa Eulalia along road to San Mateo Ixtatan, Breedlove 8596, 5 Feb 1965 (F); 
along road to San Pedro Soloma, 3 mi SW of San Mateo Ixtatan, Breedlove _ 
6 Sep 1965 (F, US). Totonicapan: near Cumbre del Aire, on road between Huehu 
tenango and Siga, Standley 65939, 30 Feb 1939 (F); Tecum Uman Ridge at me 
154 on Ruta Nacional No. 1, ca. 20 km of Totonicapan, Beaman 4205, 14 Aug 1960 
(MSC). 


Plants 1-4(-6.5) cm tall. Stems 1-12, ascending to erect, those of larger 
plants usually with 1-4 branches at various levels and further branched near 
summit, bearing flowers in small, leafy cymes. Basal leaves (sometimes more 
or less marcescent at flowering time) spreading, scarcely conduplicate, with 
midrib keel well developed only near apex, broadly oblong-oblanceolate to 
elliptic-obovate, usually 4-9 mm long, 2-4 mm wide. Cauline leaves some- 
what narrower, increasingly conduplicate and carinate upward; upper ones 
carinate for most of length. Leaf apices acute, apiculate. White leaf margins 
and keel sharply defined, both 0.05—0.13(-0.2) mm wide, or slightly wider 
toward apex, at least upper leaves with keel and usually margins minutely 
denticulate. Leaves of most plants densely crowded throughout; those of 
taller plants often more distant on lower part of stem, with lower internodes 
up to twice as long as subtending leaves, but crowded on upper part. Calyx 
tube 5-8 mm long. Calyx lobes triangular, 1-1.5 mm long, erect, scarcely 
conduplicate, acute to acuminate, with white margins ca. 0.05 mm wide, 
entire, and keels scarcely developed or up to 0.04 mm wide. Corollas 6-12 
mm long when closed; expanded limb 5—8 mm across. Corolla tube 4.5-10 
mm long. Lobes oblong-triangular, 15-2 mm long, apiculate. Free portions 
of appendages ca. 0.75 times as long and about as wide as lobes, triangular, 
erose-serrate, sometimes emarginate. Corolla tube mostly whitish; lobes, free 


Figs. 8-9 a-c, 10-13 b Cc 


b 


Figs. 8-9. Gentiana spp. G. sedifolia. Fig. 9. G. beamanii. a, interior surface 
of corolla, slit oe and pressed; ae exterior surface of calyx; c, pistil; 
d, intact open flower from above; e, closed flow 


0-13. Gentiana spp. Fig. G. fieldiana. Fig. 11. G. pumilio. Fig. 12. 
“perp Fig. 13. G. prostra ” dee awn from a specimen from Colorado, where 
species normally bears tetramerous flowers). a, interior surface of corolla, slit 
ia and pressed; b, exterior surface of calyx; c, pistil. 


ay 


portions of appendages, and uppermost portion of tube white to medium 
blue; exterior surface of lobes, except near margins, suffused with green, 
and occasionally also with deep violet-blue except near inner margin. Stamens 
becoming free at ca. 1/2 total length of corolla; free portions (including 
anthers) ca. 2 mm long. Pollen grains prolate. Sexine relatively coarsely 
striato-reticulate. Pistil tapering into style ca. 1 mm long. Capsule 3-5 mm 
long, 2/3 to entire length and up to 2 mm of gynophore exserted from 
marcescent corolla at maturity. Seeds ellipsoid, light brown, striato-reticulate, 
ca. 0.7 mm long, 0.35 mm in diameter. 

Gentiana fieldiana is native to the Sierra Madre and Los Altos Cuchu- 
matanes in Guatemala (Fig. 22) where it grows in moist sites in subalpine 
meadows (llanos) and openings in pine forests, from 2500 to 3500 m. Plants 
have been collected in flower from December through July. Its name honors 
the contributions of botanists now and formerly of the Field Museum of 
Natural History to the knowledge of the flora of Guatemala, especially Dr. 
Julian A. Steyermark and the late Mr. Paul C. Standley, who first collected 
this species. 

A conspicuous feature of all but the smallest plants of G. freldiana is the 
presence of 1-several short, leafy branches near the summit of the stem, 
with the flowers thus being borne in small cymes. In this trait, G. freldiana 
differs from G. sedifolia, G. beamanii, and G. prostrata, in all of which 
solitary flowers generally terminate longer branches or unbranched stems 
(one such branch occasionally present in G. sedifolia). Also, G. fieldiana 
differs from the other species discussed here in that its corolla lobes taper 
abruptly to an apiculate tip, rather than gradually from near the base. Its 
pollen is similar to that of G. sedifolia, but the sexine pattern is more dis- 
tinctly striato-reticulate, rather than simply reticulate, and the lirae are finer. 

Photographs of living plants of G. fieldiana were not available for this 
study, but Breedlove 8637 and Sharp 451025 contain some corollas pressed 
while open, with the colors well retained. From these specimens it appears 
that no guidelines or sharply contrasting corolla-throat patterns are present 
in this species. 

Some specimens of G. fieldiana were previously identified as G. pumulio. 
Both of these species, as well as G. beamanii, have been collected near 
Xémal, but even in this area they maintain their distinctness. Gentiana 
fieldiana is readily distinguishable from G. pumilio by its narrower, less 
densely crowded leaves with narrower, less prominently denticulate margins, 
and by the shape of its corolla lobes. 

Breedlove 8596 consists of usually large plants with relatively narrow 
leaves for this species. The general aspect of these plants consequently re- 
sembles that of G. sedifolia, but their well-developed leaf margins and keels 
and the position of their stamens clearly identify them as G. fieldiana. 


4. GENTIANA PUMILIO Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot. 
Ser. 23:76. 1944. Type: GUATEMALA: San Marcos: between San Sebas- 


28 


tian and summit of Volcén de Tajumulco, rel mark 35489, Feb 1940 
(holotype F!, isotypes MSC!). Figs. 5, 11, 


AL csasstinissestnsssssssisnsninasied 
0.0] mm 


Figs. tee Scanning-electron eee a a of pollen grains of Gentiana spp. 
Fig. an y. sedifolia (Mulroy 1099, HAM). ae a — beam eg (Beaman 3899, 
MSC) 16. aA sags saa (Beamon 3071, 17 ee aad erage 
2868, MSE). ae perpusilla ani ie cs “MICH Fig. 
prostrata (Sharp s ae from Alaska). Figs. 14-19 to same ree 


22 


Plants 0.7-4 cm tall. Stems usually 1-10, sometimes much branched form- 
ing dense tufts. Basal and lower cauline leaves persistent and green or the 
lowest pairs marcescent. Leaves ascending, densely imbricate-crowded on 
whole length of stem, broadly spatulate to orbicular, either tapering grad- 
ually to base or at least upper leaves more commonly abruptly contracted 
below middle. Larger, lower leaves 2.5-15 mm long, 0.3-1.2 times as wide 
as long; upper leaves smaller and more strongly ascending but similar in 
proportions. Apices abruptly acuminate-apiculate. All leaves prominently 
white-margined and keeled but not strongly conduplicate; margins and keels 
of upper leaves mostly 0.25-0.35 mm wide, minutely but copiously denticu- 
late at least above middle. Flowers solitary or in clusters of 2-3. Calyx tube 
largely concealed by leaves, 3-5.5 mm long, carinate distally. Calyx lobes 
ovate-triangular, erect, strongly carinate-conduplicate, 1-2. mm long, 0.5- 
1.5 mm wide, obtuse, with prominent white margins and keels ca. 0.2 mm 
wide, denticulate to base. Corollas 5-11 mm long when closed; expanded 
limb 4-7 mm across. Corolla tube 4-9 mm long. Lobes ovate-triangular, 
1-2 mm long, ca. 1.2 times as wide as long, obtuse. Free portions of appen- 
dages triangular, ca. 0.6 times as long and 0.8 times as wide as lobes, acute, 
entire or shallowly undulate. Corolla tube mostly greenish-white; lobes, free 
portions of appendages, and upper part of tube white to light blue; exterior 
surface of petals proper moderately to strongly suffused with green, and in 
blue corollas sometimes with deep violet, the suffusion strongest toward 
center. Stamens becoming free at ca. 3/5 total height of corolla; free por- 
tions (including anthers) ca. 2.2 mm long. Pollen grains spheroidal. Sexine 
relatively fine striato-reticulate. Style ca. 3 mm long, spiralling in fruit. 
Capsule 5-7 mm Jong, 1/2 to almost completely exserted from marcescent 
corolla. Seeds cylindric-ellipsoid, medium brown, striato-reticulate, ca. 0.9 
mm long and 0.3 mm in diameter. 


Additional specimens examined: 

GUATEMALA: Huehuetenango (all from Los Altos Cuchumantanes): region of 
Chémal [Xémal]}, Standley 81107 (F); cumbre de la Sierra de los Cuchumatanes, 
between the first cumbre and La Pradera, Standley 81172 (F); near Calaveras, 
Williams et al. 21954 (F) and 21955 (F, NY, US); 3-15 km N of Chémal [Xémal], 
Williams et al. 22208 (F).San Marcos: between Sibinal and summit of Volcan Tacana, 
Steyermark 36156 (F); Volcan Tajumulco, Bunting 337 (F); between Comitancito 
{[Comitancillo} and Santa Rosa junction, Sierra Madre, Williams et al. 27058 (F); 
lower slopes of Volcan Tajumulco near San Paaeieee Plowman 3064 (GH). Solola: 


near Maria Tectn, Sierra Madre, about 10— cm NW of Los Encuentros, Williams 
et al, 27325 (F). Totonicapan : region of Esra Standley 62735 (F); region 
of Salvachan, mountains above Totonicapan, just before reaching Desconsuelo, 


Standley 84489 (F); slopes of Maria Tecin above Totonicapan, Williams & Williams 
18557 (GH, US); summit of Cerro Maria Tectin, about 12 km SW of Totonicapan, 
Williams et al. 25457 (F); Maria Tecin, Molina R. et al. 16376 (F, NY, US). 
MEXICO: Chiapas: Volcan de Tacana pico, Matuda 2868 (F, GH, HAM, K, MEXU, 
MICH, MSC, NY, US). 


Gentiana pumilio is native to the Sierra Madre of southwestern Guatemala 


T q 
ICARAGUA 


GENTLANA SEDIF 


OLIA 


PANAMA 


ch - 
, ’Matanes a 


GENTIANA FIELDLANA 


SALVADOR 


9 eee ) 
L | 


A GUATEMALA 


GENTIANA PUMIL1O 


Figs. 20-24. Documented distribution of Gentiana spp. 


at 


and adjacent Chiapas, Mexico, and to Los Altos Cuchumantanes in Guatemala. 
All collections of this species are from open habitats, mostly in subalpine 
or alpine zones, from ca. 2500 to 4062 m. Most collections are from moist 
sites, but some are from dry slopes and pastures. Plants have been collected 
in flower from November through May. 

Because of its proportionately wide leaves and the wide and conspicuous 
white margins and keels of its leaves and calyx lobes, G. pumilio is readily 
distinguishable from the other species described here. Its relatively long 
style and spheroidal pollen grains are also distinctive. Gentiana pumilio 
further differs from the three preceding species in that the flowers are often 
borne in clusters of two or three, rather than singly at the ends of branches. 


5. GENTIANA PERPUSILLA Brandegee, Zoe 5:181. 1904. Type: MEXICO: 
México: Ixtaccthuatl, Purpws 302, Mar-Jul 1903 (holotype US!, isotypes 
F!, GH!, MO!, MSC!, UC). Figs. 6, 12, 


Plants 0.8—3 cm tall. Stems 1-10, ascending, not branched except occa- 
sionally near base. Leaves short-acuminate, apiculate. Basal leaves mostly in 
good condition at flowering time, obovate to broadly spatulate, moderately 
ascending, scarcely to moderately conduplicate, 4-10 mm long, 1.8-3.5 mm 
wide. Cauline leaves densely crowded, increasingly conduplicate and more 
strongly ascending upward, those near summit of stem oblanceolate, 3-6 
mm long, 1-2.5 mm wide. White leaf margins and midrib keel inconspicuous, 
ca. 0.05 mm wide, entire or with a few shallow projections near apex. Calyx 
tube 4-6.5 mm long, carinate distally. Calyx lobes triangular, erect, 1—1.2 
mm long, 0.7-1 mm wide, acute, with white margins and keels ca. 0.08 
mm wide, entire. Corollas 5-9.5 mm long when closed; expanded limb 3.5— 
4.5 mm across. Corolla tube 4-8.5 mm long. Lobes ovate-triangular, 1-2 mm 
long, about as wide as long, subacute. Free portions of appendages triangular, 
ca. 0.67 times as long and about as wide as lobes, subacute to acute, entire, 
merely undulate, or shallowly few-toothed. Corolla tube mostly greenish- 
white; lobes and free portions of appendages pale to medium blue; exterior 
surface of lobes, except near margins, and adjacent portions of tube strongly 
suffused with green. Stamens becoming free at ca. 2/3 total height of corolla; 
free portions (including anthers) 1.5—-2 mm long. Pollen grains prolate. 
Sexine relatively finely striato-reticulate. Style almost obsolete. Capsule 2.5— 
3.5 mm long, 1/2 to entire length and up to 3 mm of gynophore exserted 
from marcescent corolla. Seeds ovoid-ellipsoid, brownish-white,  striato- 
reticulate, ca. 0.75 mm long, 0.5 mm in diameter. 


Additional specimens examined: 

MEXICO: México (all from the SW side of Ixtaccihuatl): Canada de Alcalican 
(Tlaltapitongo) below La Joya, Beaman 3534 (MSC); La Joya, Canada de Alcalican, 
Rzedowski 21594 (ENCE) and 25681 (ENCB, na MSC); in canyon below 
end of auto road, ca. 2.4 km N of microwave tower, J.S. Pringle 1749 (HAM). 
Veracruz: Cofre de han NW side of mtn., Bean 5197 (GH, MSC). 


es) 
i) 


| a 
ae ig 

=, HIDALGO } = 
| op — ; 

LS eee ; 

od ee ( 
fr» 
tiene) oe Me 
# \ A A, 
‘a \ ( m TLAXCALA be. 
ee 2 Ae 
Drak, * acinar é 

1 La 7 

/ e VERACRUZ 
|. f ~~ MORELOS < J 

ne P PUEBLA i we 
ae Is) Leo ? _ 
J — 
25 - 7) a 4 
_ : oe ee 
GUERRERO f \ eo OAXACA ‘ 
- ph 9 
| 


Fig. 25. Documented distribution of Gentiana perpusilla. 


The two localities known for G. perpusilla are on peaks in the Transverse 
Volcanic Zone of Mexico (Fig. 25). It grows in the alpine zone at ca. 
3850 m on Ixtaccihuatl and 3590 on Nauhcampatépetl (Cofre de Perote), 
in microhabitats that are wet at least in summer. 

Among the species described here, G. perpusilla has by far the smallest 
corollas, and its pollination appears probably to be largely hygrocleistogamous. 
It grows where fog and drizzle are nearly continuous during its flowering 
season, and consequently its photonastic corollas are infrequently and briefly 
open. Observation of pollen in preserved flowers from Ixtaccihuatl indicated 
that dehiscence of the anthers into the small space within a closed corolla 
can readily bring pollen into contact with the stigmatic lobes. In one corolla, 
however, the presence of a few alien grains indicated that some chasmogam- 
ous pollination does occur. 


ACKNOWLEDGEMENTS 


I express my sincere thanks to: Dr. John H. Beaman, for his comments 
and notes on the species described here, and for his review of the manu- 
script; Dr. Juliana C. Mulroy, for suggestions for the improvement of an 
earlier version, and for collections of G. sedifolia and related species; Mrs. 
Margaret Williams, for additional collections; Dr. Thomas Duncan, for his 


2) 


cooperation and assistance with field work; Messrs. James Darley and 

ouglas Brown, for SEM photographs of pollen grains, using facilities at 
McMaster University; the curators and staff of the herbaria listed in the 
introduction of this paper, for making specimens available for study; the 
Consejo Nacional de Ciencia y Tecnologia, for permission to collect speci- 
mens of Gentiana in Mexico; and the Board of the Royal Botanical Gardens, 
for its support of the publication of this paper. 


REFERENCES 


GILG, E. 1906. Beitrage zur Kenntnis der Gentianaceae III. Gentianaceae andinae. 
Repert. Spec. Nov. Regni Veg. 2:33-56. 

HOLMGREN, P.K., & W. KEUKEN ae Index Herbariorum. Part I: The herbaria 
of the world, ed. 6. Regnum Veg. 92. pp. 

HUMBOLDT, [F.H.} A. VON, A. [J.] BONPLA ND, & CS. KUNTH. 1815-“1823” 
{-1825]. Voyage aux Régiones Equinoctiales du Nouveau Continent, Fait en 1799- 
1804. Sixiéme Partie. Bota anique. Sect. 3. Nova Genera et Species Plantarum 
Libraria Graeco-Latino-Germanica, cee 7 vols. (Gentiana in vol. 3(10). 1819; 
supplementary note in vol, 5(23). 3, 

KERNER VON MARILAUN, ae 887-1891. me Bibliographisches 
Institut, Leipzig. 2 vols. (Gentiana prostrata in vol. 2 91.) ae translation 
by F.W. OLIVER ecu Henry Holt and canes New 

KUSNEZOW, N.J. 1895. Gentiana Tournef. Iz: ENGLER, [H.G.] i “& [A.E.] 
P L. Die aes Pflanzenfamilien . . . Verlag von Wilhelm foi 
Leipzig. 4(2) :8 

LINDSEY, me 1940 a anatomy in the Gentianaceae. Amer. J. Bot. 27 :640-652. 

LOVE OVE. 1975. The Spanish gentians. Anales Inst. Bot. Cavanilles 
32:22] "932. 

NILSSON, S. 1967. Pollen morphological studies in the Gentianaceae-Gentianinae. 

145. 


PRINGLE, J.S. 1977. Taxonomy of Gentiana aguas in Mexico and Central 

America. I. Sect. Bee une. Sida 7:174-217. 

__. 1978. Sectional and subgeneric names in Gentiana (Gentianaceae). Sida 
47. 

TOYOKUNI. H. 1963. Conspectus Sg eee Re re eee a general view of 
the Gentianaceae aa to Japan. J. Fac. . Hokkaido Imp. Univ., Ser. 5, 
Bot. 7:137-—259, pl. LIV. 

EAVER, R. - Tie ary RES. 1975. Cytotaxonomic notes on some 
Gentianaceae. a Arnold Arbor. 56:211 


REVISION OF NORTH AND CENTRAL 
AMERICAN NAJAS (NAJADACEAE) 


ROBERT R. HAYNES 
Ecology and Systematics Section, Department of Biology, 
The University of Alabama, University, Alabama 35486 


Najas is a cosmopolitan genus of submerged aquatic plants with its greatest 
diversity in tropical and subtropical regions. In North America, individuals 
of the genus can be separated from other aquatic genera by the plants of 
Najas possessing sub-opposite serrulate leaves and axillary see flowers. 
Although floristic treatments have been prepared for certain 
eastern United States (Clausen 1936) and Panama (Wentz and ie 
1973 )—at no time since Rendle (1901) have all the species, either world- 
wide or regional, been included in one revisionary treatment. Morphological 
variability within the group has been poorly understood. Because of the 
ack of a thorough revisionary treatment and a poor understanding of the 
taxa, the genus is, I believe, in need of revision. 

The genus Najas has been divided into two subgenera, Najas and Caulinia, 
by Ascherson (1864) with Najas being dioecious and Caulinia_ being 
monoecious. Magnus (1889) separated subgenus Caslinia into two sections, 
Americanae with sloping leaf sheaths and Ewvvaginatae with truncate to 
auriculate leaf sheaths. Rendle (1899) divided the subgenus into two more 
sections, Spathaceae with staminate and carpellate flowers enclosed in floral 
envelopes and Nwdae with no flowers enclosed in floral envelopes. According 
to Rendle (1899), the sections Americanae and Euvaginatae were composed 
of plants with staminate flowers enclosed in floral envelopes and carpellate 
flowers without such envelopes. Wilde (1961), in an investigation of Asiatic 
and Malaysian species of Najas, determined the presence of the envelopes 
to be too inconsistent to segregate sections. He thus proposed the abolition of 
Rendle’s sections. 

Variation in vegetative parts (plasticity) within individual taxa of aquatic 
plants has been recognized in Callitriche (Fassett 1951), Halodule (Phillips 
1967), Nymphaea (Williams 1970), Najas minor (Wentz & Stuckey 1971), 
and Potamogeton (Haynes 1974). These workers concluded that because 
of this plasticity, the taxonomy of the genera in question should rest_pri- 
marily on reproductive characters. As a result of my field work and the 
examination of several thousand herbarium specimens, I have found consider- 
able plasticity within the leaf sheaths of Najas. 1 have concluded, therefore, 
that the separation of sections based upon the shape of the auricles is super- 


"Paper No. 7 from the Aquatic Biology Program, The University of Alabama. 


SIDA 8(1): 34-56. 1979. 


—_~ 


3 


ficial. I agree with Wilde that four sections are unwarranted. Wilde (1961) 
also suggested that the two subgenera be reduced to sections because of the 
“great resemblance of N. marina to other species of Najas ....” I agree 
that N. marina resembles the other species. However, with N. marina being 
dioecious, having a testa several cell layers thick, and having prickly inter- 


have a testa three cell layers thick, and do not have prickly internodes and 
dorsal surfaces of the laminae—I believe the subgeneric category to be 
warranted. 

The nomenclature of Najas is difficult. Many of the taxa in North and 
Central America were named by Sprengel (1825), Braun (1864), and 
Magnus (1870, 1894). The specimens examined by these workers were de- 
posited at Berlin and later were destroyed during World War I. Gerloff 
(personal communication) informs me that no pre-World War II specimens 
of Najas (including the Willdenow collection) remain at Berlin. Therefore, 
I have had to resort to published descriptions and accepted usage of many 
names. In some instances (see Haynes and Wentz 1974) when only one or 
two taxa of Najas occur in an area, there is good evidence for my decision. In 
others, however, I have only the accepted usage and description to follow. 

The morphology, systematic relationships, and economic value of Najas 
were discussed by Haynes (1977). Evidence from field work has indicated 
that Najas is just as morphologically plastic as Callitriche (Fassett 1951), 
Potamogeton (Haynes 1974), and Ranunculus (Bostrack and Millington 
1962). Just as workers in these other genera have resorted to reproductive 
characters as a basis of a classification, I believe that a classification based 
mostly on reproductive structures is the only one workable for Najas. One 
should always attempt to collect specimens of Najas with seeds (Fig. 8). 
Just as identification of a sterile Aster may be nearly impossible, it is diffi- 
cult at best to determine a sterile Najas. 

The treatment that follows is based on field study and an examination of 
approximately 5000 herbarium specimens from 35 North American and 
European herbaria: A, ALU, AUA, B, BH, CAN, DAO, F, FSU, GA, GH, 
JEPS, K, KANU, KNK, LAF, LSUS, MEXU, MICH, MIN, MO, NCSC, 
NY, OS, PH, TENN, UC, UNA, UNC, UMBS, US, USF, UTC, VDB, and 
WIS (abbreviations according to Holmgren and Keuken 1974). Keys are 
based upon materials containing seeds; dimensions of leaves are taken from 
the fully expanded; longest leaves of a specimen; measurements of widths are 
taken approximately at the widest point of the leaf; and descriptions of the 
seeds are taken strictly from mature structures (Fig. 8). 


NAJAS Linnaeus, Sp. Pl. 2: 1015. 1753; Gen. PI. a 5. 449547 54. 
Caroliana Raf., J. Phys. Chim. Hist. Nat. Arts 89:259. 1819. 
Caulinia Willd., Mem. Acad. Roy. Sci. Hist. (Berlin) 1798. 
Fluvialis Adans., Fam. Pl. 2:472. 1763 
Hyas Dumort., Anal. Fam. Pl. 61. 1829. 
Ittnera C. C. Gmel., Fl. Bad. 3:590. 1808. 


36 


Plants glabrous, herbaceous, annual or rarely perennial, aquatic, submersed 
in fresh or brackish waters, monoecious or dioecious. Stems slender, much 
branched, rooting at the lower nodes, sometimes armed with prickles on the 
internodes. Leaves subopposite or appearing whorled due to reduced inter- 


and flattened, l—-nerved, sometimes dorsally armed with prickles on the 
midrib; margins usually serrulate with 5-100 teeth per side; apex acute to 
acuminate, with 1-3 teeth per side; teeth multicellular, formed by layers of 
cells decreasing in cell number outward terminated by a large, sharp-tipped 
cell, or unicellular. Sheaths variously shaped, each enclosing a pair of tiny 
hyaline scales; margins usually toothed with 1-15 teeth per side. Flowers 
pee axillary, sessile or short pedunculate, solitary or clustered, often 
subtended by an involucre; involucre clear, bronze, brown, light green, purple, 
or red-purple. Staminate flowers subtended by a membranous involucre or 
the involucre rarely absent, each consisting of a single stamen; peduncle at 
first short, elongating at anthesis, pushing the anther through the involucre; 
anther sessile, 1- or 4-loculed, dehiscing irregularly; pollen 3-celled, glob- 
ular or ellipsoid, densely filled with starch, monocol pate, the wall with 
shallow reticulations, thin, not divided into exine and entine. Carpellate 
flowers sessile, 1-loculed, 1-ovuled; involucre absent or rarely present; 
ovule basal, anatropous, with 2 integuments; gynoecial wall 2 cell layers thick, 
ending in a short style with 2—4 branches. Fruit 1-seeded, dehiscing by decay 
of gynoecial wall; gynoecial wall extremely delicate, closely enveloping the 
seed. Seeds without endosperm, areolate, with a basal raphe, fusiform to 
obovate, occasionally asymmetrical at apex or recurved; testa hard, brittle, 
3 or several cell-layered, pitted or smooth; areolae formed by outer two 
layers of testa, variously eae shaped, irregularly ope or in 

15-60 rows, the end walls often raised, giving the testa a papillose appearance; 
embryo elongate, without lateral enlargement of cotyledons. TYPE SPECIES: 
N. marina L. (From the Greek, naias, a water-nymph.)—Bushy-pondweed, 
Naiad, Water-Nymph. 


KEY TO THE TAXA 


1. Plants dioecious; testa pitted, 4 or more ae oS thick; internodes and Rais 
WN; 


surfaces of lamina with prickles; seeds 1.2 m e or wider 0.0.2... 

1. Plants monoecious; testa smooth or pitted, 3 cal ne thick; internodes and aa 
surface of lamina without ae seeds 1.2 mm or less wide, #f 1.2 mm wide, 
then with smooth testa 2 


. Seeds asymmetrical at apex or recurved 

3. Areolae broader than long, ladder-like; teeth multicellular ............ 7. N. minor 

3. Areolae longer than broad, never ladder-like; teeth unicellular ..... 7 
. Seeds strongly recurved; areolae in 10-20 rows; leaves in age stiff, recury 


ee 
4. vis asymmetrical at apex but not ae areolae in : 35 ‘5 ws; leaves 
n age limp and spreading to ascendin a gracillima 
2 Sale symmetrical at apex, not recurved 


oF 


5. Lamina teeth unicellular, (18—) 30-100 per side 
6. Testa smooth, glossy; seeds obovate, brown to yellow; anthers 1—loculed 
2.N 


eee . flexilis 
6. Testa pitted, dull; seeds fusiform, yellowish-white to greenish-brown; anthers 
1— or 4-loculed 


7. Leaf sheaths deeply auriculate; restricted in North America to California 
4, N 


7. Leaf sheaths rounded to truncate; widespread 


graminea 

8 

8. Seeds 3.3-3.8 mm long; areolae arranged in 
N 


DOGO OWS 2:4 anid aeeiueies 
dy, duets attests d. N. guadalupensis var. muenschert 
8. Seeds 2.5 mm or less ong: areolae in fewer than 


9. Teeth evident to unaided eye; anthers 1—-locu 
5) 


c. N. evadalapents var. floridana 
9. Pe invisible to unaided eye; anthers 4—locu 


aminae with 50-100 teeth per side; stem 


r less in di- 
meter 3a. N. gu anes var. guadalupensts 
10. (minde with less than 50 teeth per side; stem mm or more 

3b. N. 


epee. var. olivacea 


in diameter 
5. Lamina teeth multicellular, 5—25 per side 
11. Seeds 2 m 


m long or longer; areolae arranged in ca. 50 rows, elongate; 

teeth shorter than rest of the lamina is wide . _N. arguta 

. Seeds less than 2 mm long; areolae arranged in ca. 20 rows, Ssided, all 
sides about equal; teeth longer than rest of lamina is wide 


e 
—_ 


8. N. wrightiana 
NAJAS subgenus NAJAS 


Plants dioecious. Stem slender, much branched, rooting at the lower nodes, 
usually armed with prickles on the internodes. Leaves subopposite or appear- 
ing whorled, sessile; laminae coarsely serrate, the teeth multicellular, a 
midrib with prickles. Flowers solitary; anthers 4—loculed. Seeds 2.2-4.5 m 
long; testa several (more than 3) cell layers thick; areolae arranged es 
larly. Species 1. TYPE SPECIES: N. marina L. 


1. NAJAS MARINA L., PL 281019... 7 9: 


Ittnera major C. C. = , Fl. Bad. 3: a 1808. 
I. najas C. C. Gmel., Bad. 3:590. 1808. 


Najas gracilis vee Mea. Torrey Bot Club ae :61. 1893. 
. 40. 1903 


_ in Martius, Fl. Brasiliensis 3 (3) :804. 1894. 
marina Vat. as Gs. Br.) K. Schum. in Martius, Fl. Brasiliensis 
3G) :725. 1894. 


N. marina var. angustissima K. Schum. in Martius, Fl. Brasiliensis 3(3):725. 1894. 

N. mariana var. bollei K. Schum. in Martius, Fl. Brasiliensis 3(3) :726. 1894. 
7 a var. ree Rendle, Trans. Linn. Soc. London, Bot. 5:396. 1899. 

N. marina var. californica Rendle, Trans. Linn. Soc. onan Bot. 5:398. na 

vat. denticulata Rendle, Trans. Linn. Soc. London, Bot. 5:438. 

marina var. ebrenbergii (A. Br.) K. Schum. in Martius, Fl. eae 
3(3) a ees 

N. marin . genuina K. Schum. in Martius, Fl. Brasiliensis 3(3) :724. 1894. 

N. marina var. ae ie Dudley, Cornell Univ. Sci. Bull. 2:104. 1886. 


38 
N. marina var. latifolia (A. Br.) K. Schum. in Martius, Fl. Brasiliensis 3 (3) :725. 
N marin var. latior F. Muell. ex K. Schum. in Martius, Fl. Brasiliensis 3 (3) :725. 
ere var. mexicana Rendle, Trans. Linn. Soc. London, Bot. 5:398. 1899. 
marina a multidentata (A. Br.) K. Schum. in Martius, Fl. Brasiliensis 
1894. 


3(3) :72 


N. marina vat. muricata (Rafteneau-Delile) K. Schum. in Martius, Fl. Brasiliensis 
3 . 1894. 


S, 


N. pee var. Pig ii (A. Br.) K. Schum. in Martius, Fl. Brasiliensis 


marina var. recurvata Dudley, So Univ. Sci. Bull. 2:104. 1886. 
major All., Fl. Pedem. 2:221 
major var. angustifolia A. Br. | aa 2:275. 1864. 
major var. ebrenbergit A. Br., J. Bot. 2:275. 1864. 
major var. gracilis Morong, Boe Gaz. (Crawfordsville) 10:225. 1885. 
major var. intermedia A. Br., J. Bot. 2:276. 1864. 
major var. microcarpa A. Br., J. Bot. 2:276. eee 
m ilies var. multidentata A. Br. J. Bot. 2:275. 1864. 
vajor var. paucidentata A. Br., J. Bot. 2:276. 1864. 
ape ae ee Delile, ee Egypte, Hist. Nat. 281. 1813. 


ee . 


Plants dioecious. Stems 6-45 cm long, 0.5-4.0 mm in diameter, branched 
upward; internodes 0.3-11.0 cm long, with or without prickles. Leaves 
0.5-3.9 cm long; laminae spreading to ascending, 0.4-4.5 mm wide, acute 
with 1 tooth at apex, coarsely serrate with 8-13 teeth per side, the terminal 
teeth similar in size and structure to the lateral teeth, the teeth multicellular, 
the midrib with prickles; sheaths 2.0-4.4 mm wide, wider than the laminae, 
acute, the teeth similar in size and structure to those of the laminae. Flowers 
solitary, the involucre brown to slightly pu oe soar flowers 1.7—3.9 
mm long, the involucre beaks 0.3-0.7 mm long, 2—lobed, the anther 1.7—3.0 
mm long, 4—-loculed; carpellate flowers 2.5-5.7 mm iene. the style 1.2-1.7 
mm long, the stigma 3-lobed. Seeds 2.24.5 mm long, 1.2—2.2 mm wide, 
reddish brown, ovoid, the testa pitted, the areolae 3—4-angled, ca. 0.2 mm 
long, 0.1 mm broad, irregularly arranged, the end walls slightly raised (Fig. 

15s 


8A). Chromosome number, 27 = 12 (Viinikka : 
Brackish or highly mere waters of ponds and lakes, North and South 
Dakota to western New ; California to Utah, south to Panama; Florida 


and Caribbean islands (Fig. 

TYPE: Europe maribus (Hol orype: LINN; Microfiche #1156 IDC 1945, 
682:II 6, 7 

With its prickly internodes and prickles along the undersurface of the 
leaves, N. marina is the easiest of our Najas to recognize. The species is 
variable in the size of leaves, teeth, and seeds and, as a result, has been 
divided into numerous varieties. These differences usually can be found 
within single populations and, therefore, do not warrant taxonomic recog- 
nition. Viinikka (1973), however, observed two cytological races of the 
species, one race with B chromosomes and one without these accessory chro- 


Fig. of North America showing the documented distribution of Najas 
marina. Week see inserted. ) 


40 


mosomes. He examined one population from North America (Erie County, 
Ohio) and observed B chromosomes from those individuals (personal cor- 
respondence). He indicated (personal correspondence) that, in Europe, 
plants from the two races are morphologically different. Specimens with large 
seeds and leaves (no size given) invariably do not possess B chromosomes, 
whereas those with smaller seed and leaves do possess B chromosomes. He 
has not yet determined how to rank these apparent entities. 


NAJAS subgenus CAULINIA ( Willd.) Aschers., Fl. Prov. Brandenburg 1:670. 

1864 

Plant monoecious. Stem slender, much branched, rooting at the lower 
nodes, the internodes without prickles. Leaves subopposite or appearing 
whorled, sessile; laminae minutely serrulate to serrate, the teeth unicellular 
or area the midrib without prickles. Flowers solitary or clustered; 
anthers 1—- or 4—loculed. Seeds 0.7-3.8 mm long; testa 3 cell layers thick; 
nee arranged in longitudinal rows. Species 2-9. LECTOTYPE SPECIES: 
Caulinia flexilis Willd. = N. flexilis (Willd.) Rostk. & Schmidt. (Lectotype 
here designated. ) 


2. NAJAS FLEXILIS (Willd.) Rostk. & Schmidt, Fl. Sedin. 382. 1824, 
Caulinia flexilis Willd., Mem. Acad. Roy. Sci. Hist. (Berlin) 95. 1798. 
Fluvialis flexilis (Willd.) Pers., Synops. 2:530. 180 
Najas canadensis Michx., Fl. Bor. Amer. 2:220. 1803. 

N. caespitosus (Maguire) Reveal in Welsh, Atwood, & Reveal, 
uralist 35:357. 1976. 

N. flextlis subsp. caespitosa Maguire & Jensen, Rhodora 44:7. 1942 

N. flexilis var. congesta Farwell, Rep. Michigan Acad. Sci. 21:348. 1920. 

N. flexilis var. robusta Morong, Bot. Gaz. (Crawford suille) 10:255. 1885, 


Great Basin Nat- 


Plants monoecious. Stems 2.5-50 cm long, 0.2-0.6 mm in diameter, often 
profusely branched above; internodes 0.16-6.8 cm long, without prickles. 
Leaves 0.8-4.2 cm long; laminae spreading to ascending, 0.2-0.6 mm wide, 
acute with 1-2 teeth at apex, minutely serrulate with 35-80 teeth per side, 
the terminal teeth similar in size and structure to the lateral teeth, the teeth 
unicellular, the midrib without prickles; sheaths 0.7-1.6 mm wide, wider 
than the laminae, round, minutely serrulate with 8-10 teeth per side, the 
teeth similar in size and structure to those of the laminae. Flowers 1 (—2) 
per axil, the staminate in the upper axils, the carpellate throughout, the 
involucre bronze; staminate flowers 1.1-2.7 mm long, the involucre beaks 
0.7-1.2 mm long, 3-lobed, the anthers 1.1-2.7 mm long, 1-loculed; 
carpellate flowers 2.5-4.7 mm long, the style 1.5-1.7 mm long, the stigma 
3-lobed. Seeds (1.2—) 2.5-3.7 mm long, 0.2-1.2 mm wide, ee brown to 
yellow, narrowly to broadly obovate, the testa smooth, the areolae 5—6-angled, 
ca. 0.2 mm long, ca. 0.1 mm wide, in ca. 50 rows, without raised end walls 
(Fig. 8B). Chromosome number unknown. 

Lakes and rivers, Nebraska, Missouri, and Maryland, north to southwestern 


41 


Ontario and Newfoundland; Alberta and Saskatchewan south to western 
Oregon and central Utah (Fig. 2). 

TYPE: Pennsylvania (Holotype: B; microfiche # 17094 IDC 7440. 1224: 
III. 2.) 

In habit, N. flexilis is most similar to N. guadalupensis. However, when 
seeds are present, N. flexilis can be easily separated from the latter species 
by the glossy, smooth, yellowish seeds that are widest above the middle. 

Several different varieties have been proposed for North American indi- 
viduals of N. flexilis, e.g. var. robusta Morong, var. congesta Farwell, and 
var. caespitosa Maguire and Jensen. These varieties are based, for the most 
part, on differences in vegetative structures. Clausen (1936), in discussing 
some of these different forms, stated “Different forms of the species do 
occur, but the characters, such as width of leaves, size and shape of seeds, 
and habit, occur in all sorts of combinations and can not be correlated with 
geographical areas to give definite geographical races or varieties which are 
worth naming.” I have reached the same conclusion. 

Najas flexilis is probably the most common Najas in the northern United 
States. Wentz and Stuckey (1971), however, indicated that the species is 
becoming less common in Ohio. This decrease in abundance apparently cor- 
responds with the increase in turbidity and general decline in water quality 
of natural lakes and rivers. 


3. NAJAS GUADALUPENSIS (Sprengel) Magnus, Beitr. Gatt. Najas 8. 1870. 

Plants monoecious. Stems 11.0—90.0 cm long, 0.1-2.0 mm in diameter, 
profusely branched; internodes 0.1-9.0 cm long, without prickles. Leaves 
0.3-3.3 cm long; laminae spreading, 0.2-2.1 mm wide, round obtuse to 
acuminate with 1-3 teeth at apex, minutely serrulate with 18-100 teeth per 
side, the terminal tooth slightly larger than but similar in structure to the 
lateral teeth, the teeth unicellular, the midrib without prickles; sheaths 1.0- 
3.4 mm wide, wider than the laminae, round to slightly auriculate, serrulate 
with 4-8 teeth per side, the teeth similar in size and structure to those of 
the laminae. Flowers 1—3 per axil, the staminate in the upper axils, the 
carpellate in the lower axils, the involucre purple-tinged; staminate flowers 
1.5-3.0 mm long, the involucre beaks 0.2-1.3 mm long, 4—lobed, the anther 
1.0-1.7 mm long, 1- or 4—loculed; carpellate flowers 1.5-4.0 mm long, the 
styles 0.3-1.5 mm long, the stigmas 4—lobed. Seeds 1.2-3.8 mm long, 0.4- 
0.8 mm wide, purple-tinged, fusiform, the testa pitted, the areolae 4—G-— 
angled, 0.08-0.1 mm long, ca. 0.08 mm wide, in 20-60 rows, the end walls 
not raised. Chromosome number, 27 = 12, 36, 42, 48, 54, 60 (Chase 1947 
a,b). 

Najas guadalupensis has long been considered a highly variable species 
(Fernald 1923). Clausen (1936), Ooststroom (1939), Wentz and Haynes 
(1973), and Haynes and Wentz (1974) discussed variability within the 
species and suggested that the complex should possibly be considered as 


LAMBERT AZMUTHAL CQUAL-AREA PROJECTION 


Fig. 2. Map of North America showing the documented distribution of Najas 
flexilis. 


43 


several closely related taxa. Rosendahl and Butters (1935) named N. ol/vacea. 
It was said to differ from N. guadalupensis in its stouter habit, its larger 
seeds, and its testa. Clausen (1937) separated N. mwenscheri from 
ecial stay by its more slender habit and its seeds with smaller areolae. 

y field work and the examination of thousands of specimens of 
N. ee ee and most of these available of N. mwenscheri and N. 
olivacea, 1 have concluded that N. olivacea and N. muenscheri represent 
populations of N. gwadalupensis at the northern limit of its range. They are 
slightly different from most individuals of N. guadalupensis. However, inter- 
mediates do exist. Thus, I think they should be recognized but do not believe 
that specific rank is justified. I consider varieties as morphogeographic sub- 
divisions of a species that presumably reflect genetic differences (Kapadia 
1963). I have therefore chosen to recognize the taxa that have been called 
N. olivacea and N. muenscheri as varieties of N. guadalupensts. 

By having unicellular teeth on the margin of the lamina, N. gauadalupensis 
resembles N. flexilis and N. graminea. The seeds of N. guadalupensis are 
pitted; those of N. flexilis are smooth. The sheaths of N. gwadalupensis are 
rounded to slightly auriculate; those of N. graminea are deeply auriculate. 


3a. NAJAS GUADALUPENSIS var. GUADALUPENSIS 
Caulinia guadalupensis Sprengel, Syst. Veep 1220: 1824, 
Najas flexilis var. curassavica A. Br., J. Bot. 2:277. 1864. 
N. flextlis var. fusiformis Chapman, FIL. Seuthern U. S. 444. 1883. 
N. flexilis var. guadalupensis (Sprengel) A. Br., J. Bot. 2:276. 1864. 
N. microdon vat. curassavica A. Br., Sitzb. Ges. Naturf. Fr. Berlin. 1868. 
N. microdon var. guadalupensis (Sprengel) A. Br., J. Bot. 2:276. 1864. 
N. urbaniana O. C. Schmidt, Fedde Rep. Spec. Nov. 22:99. 1925. 


Stems 11-75 cm long, 0.1-0.8 mm in diameter. Leaves 0.3-2.8 mm long; 
laminae 0.2-1.8 mm wide, acute to mucronate, with 50-100 teeth per side; 
sheaths 1.0-1.9 mm wide, round, with 5-8 teeth per side. Flowers 1-3 per 
axil; staminate 1.5-2.5 mm long, the involucre beak 0.2-0.6 mm long, the 
anther elliptic, ce carpellate flowers 1.5-2.0 mm long. Seeds 1.2—2.5 
mm long, 0.4-0.6 mm wide, the areolae in ca. 20 rows (Fig. 8 

Lakes, rivers, a canals, southern Maine to southern Albena: and Wash- 
ington, south to Guadaloupe, Curacao, and Panama (Fig. 3). 

TYPE: Guadeloupe (B, destroyed during World War IT). 


3b. NAJAS GUADALUPENSIS var. Olivacea (Rosendahl & Butters) Haynes, 
comb. & stat. nov. 
Najas olivacea Rosendahl & Butters, Rhodora 37:347. 1935. 


Stems 15-40 cm long, 1.0—2.0 mm in diameter. Leaves 0.9-1.8 cm long; 
laminae 1.5-2.0 mm wide, acute, with 20-40 teeth per side; sheaths 2.5-3.4 
mm wide, round to slightly auriculate, with 4-8 teeth per side. Flowers 1 
per axil; staminate flowers 2.3-2.8 mm long, the anther ovoid, 44loculed; 
carpellate flowers 2.7-3.1 mm long. Seeds 2.3-2.5 mm long, 0.6-0.8 mm 


Fig. 3. Map of North America showing the documented distribution of Najas 
guadalupensis var. guadalupensis. (Caribbean islands inserted.) 


45 


wide, the areolae in 20-40 rows (Fig. 8D). 
Lakes and rivers, lowa to Manitoba, east to New York and Quebec (Fig. 


TYPE: UNITED STATES: MINNESOTA: Kandiyohi Co.: growing in tufts in 
1-3 feet of water, muddy bottoms; abundant in NE bay of Norway Lake, 
6 Sep 1933, C. O. Rosendahl and F. K. Butters 6446 (Lectotype, MIN}; 
isolectotype, MIN! ). (Lectotype here designated. ) 


3c. NAJAS GUADALUPENSIS var. FLORIDANA Haynes & Wentz, Sida 5:262. 
1974 


Stems 7-51 cm long, 0.1-1.7 mm in diameter. Leaves 0.9-3.3 cm long; 
laminae 0.5—2.1 mm wide, round obtuse to acuminate, with 18-42 teeth per 
side; sheaths 1.2-2.5 mm wide, round, with 5—8 teeth per side. Flowers 1 
per axil; staminate flowers 1.5-2.4 mm long, the anther elliptic, 1—-loculed; 
carpellate flowers 1.0-3.5 mm long. Seeds 1.6—2.2 mm long, 0.3-0.8 mm 
wide, the areolae in ca. 20 rows (Fig. 

In lakes, streams, and canals, Florida, ccna Alabama, and central Georgia 
(Fig. 4 

TYPE: UNITED STATES: FLORIDA: Dade Co.: abundant in brackish water 
of ditch along St. Rt. 41 at edge of Everglades National Park, ca. 40 mi. W 
of Miami, 5 Apr 1972, W. A. Wentz 670 (Holotype, US!; isotypes, GH!, 
MICH!, OS! ). 


3d. NAJAS GUADALUPENSIS var. muenscheri (Clausen) Haynes, comb. & 
stat. nOv. 
Najas muenschert Clausen, Rhodora 39:59. 1937. 


Stems 30-90 cm long, 0.8-1.0 mm in diameter. Leaves 0.9-1.3 cm long; 
laminae 0.5-1.6 mm wide, acute, with 50-100 teeth per side; sheaths 1.0-1.5 
mm wide, round, with 4-8 teeth per side. Flowers 1 per axil; staminate 
flowers 2.0-3.0 mm long, the anther elliptic, 1-loculed; carpellate flowers 
2.9-4.0 mm long. Seeds 3.3-3.8 mm long, 0.5-0.7 mm wide, the areolae in 
50-60 rows (Fig. 8F). 

Shallow water of the Hudson River, New York (Fig. 4). 

TYPE: UNITED STATES: NEw YorK: Greene Co.: Tidal mudflats of Hud- 
son River, Imbocht Bay, 3 Sep 1936, W. C. Muenscher and O. F. Curtis, Jr. 
5495 (Holotype, BH!; isotype, GH! ). 


4. NAJAS GRAMINEA Raffeneau-Delile, Descr. a Hist. Nat. 281. 1813. 

Caulina alagensis Pollini, Hort. & Prov. Veron. 26. 

Najas alagensis (Pollini) Pollini, Fl. Veron. 3:49. ry 

N. seminuda Griff. in Voigt, Hort. Suburb. Calc. 694. 1845. 

Plants monoecious. Stems to 35 cm long, 0.2-0.5 mm in diameter, spar- 
ingly branched; internodes 0.4-1.9 cm long, without prickles. Leaves 0.8—2.0 
cm long; laminae spreading to ascending, 0.5-1.0 mm wide, acute with 2—3 


10 r , 
j XS ; : - 

| ! mY 6) 
| 

as SCALE | 4 
LAMBERT ATBIUTHAL CQUAL-ARZA PROJECTION = 

of . 
Fig. Map of North America aS oie documented distribution of Najas 
Ps ines var. Fash ee va (solid dots), guadalupensis vat. muenscheri (solid 

eae ), and N. guadalupensis var. sees. a ots with open star). 


ra 


47 


teeth at apex, minutely serrulate with ca. 40 teeth per side, the terminal 
teeth similar in size and structure to lateral teeth, the teeth unicellular, the 
midrib without prickles; sheaths 1.0-1.5 mm wide, wider than the laminae, 
deeply auriculate, minutely serrulate with 8-15 teeth per side, the teeth 
similar in structure and size to those of the laminae. Flowers 1-2 per axil, 
the staminate in the upper axils, the carpellate throughout, the involucre 
clear in color; staminate flowers 2.0—3.0 mm long, the involucre beaks 1.0- 
1.5 mm long, 4-lobed, the anther 1.5 mm long, 4—loculed; carpellate flowers 
to 3.5 mm long, the styles to 1.0 mm long, the stigmas 2—lobed. Seeds 1.7- 
2.5 mm long, 0.4-0.6 mm wide, greenish-brown, fusiform, the testa pitted, 
the areolae 4—angled, less than 0.1 mm long, less than 0.1 mm broad, in ca. 
35 rows, the end walls slightly raised (Fig. 8G). Chromosome number, 27 
= 48 (Sharma and Chatterjee 1967 ) 

Rice fields, ayo (Fig. 5). 

TYPE: Egypt (M 

By the deeply ae sheaths, N. graminea is one of the easiest of 
the North and Central American Najas to recognize. A native of southeast 
Asia, the species apparently was introduced into California prior to 1946 
and now is evidently quite rare there. I have seen no recent specimens of 
the species from North or Central America. 

Rendle (1901) placed this species in the section Nwdae based upon the 
lack of any floral involucre. Wilde (1961), however, demonstrated the 
presence of a leaf-like involucre and used this as evidence to abolish the 
section. 


5. NAJAS GRACILLIMA (A. Br. ex Engelm. in A. Gray) Magnus, Beitr. Gatt. 

Najas 29 LO7 0; 

laa indica (Willd.) Cham. var. gracillima A. Br. ex Engelm. in A. Gray, Man. 

. (ed. 5) 681. 1868. 

Plants monoecious. Stems 4.5—-48 cm long, 0.2-0.7 mm in diameter, slightly 
branched; internodes 0.1—3.2 cm long, without prickles. Leaves 0.6-2.8 cm 
long; laminae spreading to ascending, 0.1-0.5 mm wide, acute with 2-3 
teeth at apex, minutely serrulate with 13-17 teeth per side, the terminal 
teeth similar in size and structure to the lateral teeth, the teeth unicellular, 
the midrib without prickles; sheaths 0.5-1.5 mm wide, wider than the 
laminae, truncate, minutely serrulate with 7-8 teeth per side, the teeth simi- 
lar in size and structure to those of the laminae. Flowers 1-3 per axil, the 
staminate in the upper axils, the carpellate throughout, the involucre bronze 
to purple; staminate flowers 1.5-2.0 mm long, the involucre beaks ca. 0.8 
mm long, 2—lobed, the anther ca. 1.3 mm long, 1-loculed; carpellate flowers 
0.5-2.7 mm long, the styles 0.3-1.5 mm long, the stigmas 2—lobed. Seeds 
2.0-3.2 mm long, 0.4-0.7 mm wide, light brown, fusiform with the style 
arising from one side at the apex, the testa pitted, the areolae 4—angled, 
less than 0.1 mm long, less than 0.1 mm wide, in ca. 40 longitudinal rows, 


0 
_ounene une cena Sena meee eee 


yoo MLS 
8 FO 400 00 te t000 00 OD RL OMETERS 


LAMBERT AZIMUTHAL EQUAL AREA PROJECTION 


Fi Map of North eae showing the documented distribution of Naj 
graminea (solid square), N. ancistrocarpa (dots with open star), N. arguta (solid 
triangles), and N. ee (solid dots). 


49 


the end walls raised (Fig. 8H). Chromosome number unknown. 

In soft water lakes, Nova Scotia to Alabama, west to Minnesota and Mis- 
souri; California (Fig. 

TyPE: Northeast United States (B, destroyed during World War II). 

Najas gracillima is most similar to N. minor, especially in vegetative 
condition (see discussion under N. minor). Najas gracillima, however, can 
be separated from the latter species by its style arising off-center at the apex 
of the gynecial wall and by its areolae being longer than broad. 

The species, once more common than it is now (Wentz and Stuckey 
1971), apparently cannot tolerate pollution and has become exceedingly 
rare due to the gradual degradation of lakes and streams in eastern United 
tates. 


6. NAJAS ANCISTROCARPA Magnus, Beitr. Gatt. Najas 7. 1870. 

Plants monoecious. Stems 7-22 cm long, 0.2-1.0 mm in diameter, pro- 
fusely branched; internodes 0.2-4.0 cm long, without prickles. Leaves 0.8— 
2.6 cm long; laminae usually recurved with es 0.1-0.7 mm wide, acute with 
1 tooth at apex, conspicuously serrulate with 5S—9 teeth per side, the terminal 
tooth slightly larger than but similar in structure to the lateral teeth, the 
teeth multicellular, the midrib without prickles; sheaths 1.0-1.3 mm wide, 
wider than the laminae, round to slightly auriculate, serrulate with 3-5 
teeth per side, the teeth similar in structure and size to those of the laminae. 
Flowers 1 per axil, the staminate in the upper axils, the carpellate in the 
lower axils, the involucre bronze colored; staminate flowers 2.0-3.0 mm 
long, the involucre beaks ca. 0.4—0.6 mm long, 4—lobed, the anther 1.5-2.0 
mm long, 4—loculed; carpellate flowers to 2.5 mm long, the styles less than 
0.4 mm long, the stigmas 4—lobed. Seeds 2.5-3.0 mm long, 0.5-0.7 mm 
wide, greenish-brown, recurved, the testa pitted, the areolae 4—angled, to 0.2 
mm long, ca. 0.05 mm broad, in ca. 20 longitudinal rows, the end walls 
raised (Fig. 81). Chromosome number unknown. 

n lakes, southern Georgia and northern Florida (Fig. 5). 

TYPE: JAPAN: Yokohama (B, destroyed during World War IL). 

Najas ancistrocarpa, according to Rendle (1901) and Miki (1935), 
exceedingly rare and has been known only from three Japanese islands, 
Honshiu, Shikoku, and Yokohama. The species is either not as restricted as 
it was once thought to be or it is spreading. Yang (1974) reported it new 
to Taiwan, and Haynes and Wentz (1974) recorded it for North America. 

The species is known from four localities in North America: Open Pond 
and Cane Water Pond in Decatur County, Georgia; Milton in Santa Rosa 
County, Florida; and Lake Jackson in Leon County, Florida. This species is 
another of the many found in southeastern United States and southeastern 
Asia (Graham 1972). 

The vegetative organs of N. ancistrocarpa resemble those of N. minor and 
N. wrightiana by the presence of large teeth scattered along the margin of 


50 


wo 


SCALE 
. 


ae} . HO Be aE FDO OME TED 
LAMBERT AZIMUTHAL EQUAL- AREA PROJECTION 


Fig. 6. Map of North America showing the documented distribution of Najas 
gracillima. 


51 


quite narrow leaves. However, N. ancistrocarpa is easily distinguished from 
the latter two species by its recurved fruits (sometimes crescent-shaped). I 
know of no other Najas with such curved fruits. 


7. NAJAS MINOR Allioni, Fl. Pedemon. 2:221.1785. 
Caulinia fragilis Willd., Mem. Acad. Roy. Sci. Hist. Sree ee 1798. 


Ger 

Fluvialis minor (Allinoi) Pers., Synops. 2:530. 

Ittnera minor (Allioni) C. C. Gmel., Fl. Bad. . oon. 1808. 

Najas australis Bory ex Cham., Linnaea 4:501. 

lichotoma Roxb., Hort. Beng. 71. 

N. fragilis (Willd.) Rostk. & Schmidt, . Sein 382. 1824. 

N. indica (Willd.) Cham., Linnaea 4:501. 1829. 

N. minor var. setacea A. Br., J. Bot. 2: oe 1864. 

N. setacea (A. Br.) Rendle, ans: Linn. oF London, Bot. 5:422. 1899. 

N. subulata Thuill., Fl. Paris (ed. 2) 500. 1800. 

N. ternata Roxb. ex. Gric., Notul. 3:183. ee 

Plants monoecious. Stems 11-120 cm long, 0.2-1.0 mm in diameter, pro- 
fusely branched near apex; internodes 0.5-5.8 cm long, without prickles. 
Leaves 0.5—3.4 cm long; laminae usually recurved with age, 0.1-1.2 mm wide, 
acute with 1-2 teeth at apex, conspicuously serrulate with 7-15 teeth per 
side, the terminal teeth similar in size and structure to the lateral teeth, 
the teeth multicellular, the midrib without prickles; sheaths 1.0—3.0 mm wide, 
wider than the laminae, truncate to auriculate, serrulate to lacerate with 5 
to 8 teeth per side, the teeth similar in size and structure to those of the 
laminae. Flowers 1-2 per axil, the staminate in the upper axils, the car- 
eee throughout, the involucre light green to purple; staminate flowers 

m long, the involucre beaks 0.4-2.1 mm long, 2-lobed, the anther 

ca. 03 mm long, 1—-loculed; carpellate flowers ca. 2.2 mm long, the styles 
1.0-1.2 mm long, the stigmas 2—lobed. Seeds 1.5-3.0 mm long, 0.5-0.7 mm 
wide, purplish, slightly recurved, the testa pitted, the areolae 4—angled, ca. 
0.01 mm long, ca. 0.03 mm wide in ca. 15 longitudinal rows, the end walls 
not raised (Fig. 8J). Chromosome number, 27 = 12 (Sharma and Chatterjee 
1967 


Ponds, lakes, and slow moving streams; New York to Illinois, south to 
Florida, Mississippi, and Arkansas (Fig. 7). 

Type: ITALY: Pedemont (Holotype, TO, photo, UNA! ). 

Najas minor, native of the Eastern Hemisphere, apparently invaded North 
America nearly 50 years ago (Merilainen 1968, Wentz and Stuckey 1971). 
With the gradual eutrophication of the waters of eastern United States, 
the species has become quite widespread and, in fact, has become the most 
abundant Najas in some areas (Wentz and Stuckey 1971). 

Najas minor, with its mature leaves recurved and with its areolae broader 
than long and arranged in longitudinal rows like the rungs of a ladder, is 
one of the more distinctive species of Najas. However, young sterile indi- 


| ° . eo wie Ms 
| “ oT oO LOMENE f 
| 7, 
| am | LAMBELAT AZIMUTHAL EQUAL- AREA PROUTCTION hse 


Fig. 7. Map of North America showing the documented distribution of Najas 
minor. 

Fig. 8. Najas sears SEM micrographs taken at 20 KV. A. N. marina (X 94 )- 
B. N. flexilis (X 18). CN, | ks var. gua dalicpens (X 284). 'f 
Seca Bilas var. eee (X ih uddalupensts . floridana (X 334). 
F, bee var. muenscher ae .N, wanen (X 23%). H. N. 
grace Mis (X 19). TN. anchitrocarha 204) . J. N. minor (X 193). KL. N., 
wrightiana (X 43). L. N. arguta (X 184). 


oO 


54 


viduals resemble N. gracillima. Merilainen (1968) indicated that N. minor 
can be separated from the latter species by the former having recurved 
leaves and the latter having leaves not recurved. Wentz and Stuckey (1971) 
demonstrated that the leaves of N. minor do not become recurved until late 
in the growing season. Therefore, during the early growing season, individuals 
of N. minor could be mistaken for N. gracillima when one utilizes vegeta- 
tive characters only. 


8. NAJAS WRIGHTIANA A. Br., Sitzber. Naturf. Fr. Berlin 17. 1868. 
Najas multidentata Koch, Ber. Schweiz. Bot. Ges. 44:341. 1935. 
N. wrightiana subsp. multidentata (Koch) See Bull. Torrey Bot. Club 73:365. 
16. 


Plants monoecious. Stems 9-45 cm long, 0.3-1.0 mm in diameter, pro- 
fusely branched; internodes 0.3-5.2 cm long, without prickles. Leaves 0.5- 
2.6 cm long; laminae ascending, 0.2-1.3 mm wide, acute with 1 tooth at 
apex, conspicuously serrulate with 8-22 teeth per side, the terminal tooth 
similar in size and structure to the lateral teeth, the teeth mulicellular, the 
midrib without prickles; sheaths 0.7-2.8 mm wide, wider than the laminae, 
round, serrulate with (1—) 3-5 teeth per side, the teeth similar in structure 
and size to those of the laminae. Flowers 1-2 per axil, the staminate in the 
upper axils, the carpellate in the lower axils, the involucre brown to red 
purple; staminate flowers 1.2-1.9 mm long, the involucre beaks 0.3-0.5 mm 
long, 2—lobed, the anther ca. 1.0 mm long, 4—loculed; carpellate flowers ca. 
2.0 mm long, the styles to 0.7 mm long, the stigmas 4-lobed. Seeds 0.7-1.5 
mm long, 0.3-0.5 mm wide, whitish, fusiform, the testa pitted, the areolae 
S—angled, ca. 0.1 mm long, less than 0.1 wide, in ca. 20 longitudinal rows, the 
end walls raised (Fig. 8K). Chromosome number unknown. 

Slow moving streams and ponds, Veracruz to Florida, south to Guatemala, 
Honduras, and Cuba (Fig. 5). 

TYPE: CUBA (B, destroyed during World War II). 

Najas wrightiana superficially resembles N. arguta. Clausen (1946) dis- 
cussed this similarity and suggested that the two species could be separated 
by teeth of N. wrightiana being longer than the rest of the lamina is wide, 
whereas those of N. argwta are shorter than the rest of the lamina is wide. 
I believe, however, the best characters are those of the seed. The seeds of 
N. wrightiana are short (1.0-1.5 mm long) and have 5—angled areolae 
arranged in ca. 20 rows. Najas arguta seeds, on the other hand, are 2.1-2.6 
mm long and have elongate areolae arranged in ca. 50 rows. 


9. NAJAS ARGUTA H.B.K., Nov. Gen. et Sp. Pl. 1:371. 1815. 

Caulinia tenella C. G. Nees in Neuwied, ae 2:345. 1824. 

Najas tenera Sched. Gort. Gel. Anz. 2:715. 1825. 

Plants monoecious. Stems 15-70 cm ae 0.3-1.0 mm in diameter, pro- 
fusely branched; internodes 0.5-13.5 cm long, without prickles, Leaves 1.2—2.8 


55 


cm long; laminae spreading, 0.6-1.9 mm wide, acute to acuminate with 1 
tooth at apex, conspicuously serrulate with 15—25 teeth per side, the terminal 
tooth slightly larger than but similar in structure to the lateral teeth, the 
teeth multicellular, the midrib without prickles; sheaths 1.2-3.4 mm wide, 
wider than the laminae, round to truncate, serrulate with 2—7 teeth per side, 
the teeth smaller than but similar in structure to those of the laminae. 
Flowers 1 per axil, the staminate in the upper axils, the carpellate in the 
lower axils, the involucre purple-tinged; staminate flowers 2.0-2.2 mm 
long, the involucre beaks ca. 0.5 mm long, 4-lobed, the anther to 1.6 mm 
long, 1-loculed; carpellate flowers 1.7—2.1 mm long, the styles ca. 0.5 mm 
long, the stigmas 3—4-lobed. Seeds 2.1-2.6 mm long, 0.6-0.8 mm wide, 
yellowish-white, elongate-fusiform, the testa pitted, the areolae sometimes 
inconspicuous, S—G-angled, to 0.2 mm long, ca. 0.05 mm broad, in ca. 50 
longitudinal rows, the end walls raised (Fig. 8L). Chromosome number 
unknown. 

In lakes, Panama and the Dominican Republic (Fig. 5) 

TYPE: Provenit in aquis prope momprox- Regno novogranatensi (P?). 


ACKNOWLEDGMENTS 


I am grateful to Dr. W. Alan Wentz for his help during the course of 
this work. Appreciation is also owed to Dr. Wentz and Dr. John W. Thieret 
for their reading the manuscript and to the curators of the herbaria from 
which specimens were borrowed. A portion of the field work was completed 
at the University of Michigan Biological Station, and a portion of the labora- 
tory investigations were carried out at The Ohio State University and at 
Louisiana State University at Shreveport. 


REFERENCES 
ASCHERSON, P. 1864. ae ae pea Brandenberg. I. Berlin. 1034 pp. 
BOSTRACK, J. M. and W. LLINGTON. 1962. On the determination of leaf 


m i an arene ie species of Ranunculus, Bull. Torrey Bot. Club 


89:1 
BRAUN. ne 1864. Revision of the genus ae e a J. Bot. 2:274—279. 
ar S. S. 1947a ‘oes studie enus Najas in oe United States. 
hesis, Gail Uni pp. 
1947b. Polyploidy in an immersed aquatic angiosperm. (Abstract.) Amer. 
aa Bot. 34: ae 582 


CLAUSEN, ale 1936, Studies in the genus Najas in the northern United States. 
Rhodora 38. es 345. 
1937. A new species ot Najas from the Hudson River. Rhodora 39: 57-60. 
eae ae - 1946. Najas ar ae in Central America and its relationship to N. wright- 
jana. Bull. Torrey Bot. 73 :363-3 
FASSETT, N. C. 1951. calivihe in the New World. Rhodora 53:137- 155, 161- 
182, 185-194, 209-22 
FERNALD, M. L. 1923. ae on the distribution of Najas in northeastern America. 
SGT 29% 105-1 09. 
HAM, A. 1972. Floristics aS ee of Asia and eastern North America. 
SEE: Publ. Co., New York. 
HAYNES, R. R. 1974. A revision of North American Potamogeton subsection Pasilli 
Re Oa Rhodora 76:564-649. 


56 


1977. The Najadaceae in the southeastern United States. J. Arnold Arbor. 
58: oi mn 
nd W. A. Wentz. 1974. Notes on the genus Najas (Najadaceae). Sida 


5: 259-264, 
HOLGREN, nd W. KEUKEN. 1974. Index Herbariorum I. The Herbaria of 
the i <a Bureau for Plant Taxonomy and Nomenclature, Utrecht. 


397 

KAPADIA, Z. J. 1963. nN aiieias and subspecies: a suggestion towards greater uni- 
eo ae 12:257-259. 
AGNUsS, 1870. io zur Kenntniss der Gattung Najas L. Berlin. viti ++ 


wed pp. 
1889. Dy esas In A. Engler and K. Prand Die natirlichen Pflanzen- 


familien a 17. Verlag con Wilhelm Engelmann, Leipzig 
1894, Ger die Gattung Najas. Ber. Deutsch. Bot. "Ges 14-224. 
MERILAINEN. J. 1968. ae. minor All. in North America. oe a 161-175. 
ie S. 1935. New water pla n Asia Oricntalis II. Bot. Mag. (Tokyo) 49:687-— 


693. 

OOSTSTR OOM, S. J. VAN. 1939. Some notes on a collection of aquatic phancrogams 
from the Netherlands West Tas Islands, and from Venezuela and Colombia. 
Recueil Trav. Bot. 36:705—-708. 

PHILLIPS, R. C. 1967. On ae of the seagrass, Halodule, in Florida. Bull. Mar. 
Sci. Gulf ets 17 :672- 

RENDLE, A. 899. A systematic revision of the genus Najas. Trans. Linn. Soc. 
London, Bot 5: ee 9-444. 

. Naidaceae. Pflanzenreich IV. 12(Heft 7) :1-21. 

ROSEND Ht, oC re ae _ K. BUTTERS. | 1935. The genus Najas in Minnesota. 
oo. 37 :345-3 

SHARMA, A. K., T. CHATTERJEE. 1967. aaa of Helobiae with 
special reference to the mode of oo Cytologia 32:28 

ger C. 1825. Caulinia, pv. 20. In C. Linnaeus ots “vegetabilium vol. 1. 
Gottin 

VIINIKKA. Y,. 1973. The occurrence - chromosomes and their effect of meiosis 
in Me marina. ae 75:207- 

WENT A., and R AYNES. 1933. Rupees re ae ee In R. E. Wood- 
son, : bane WW. Sch ae “elora of Panar i Bot. Gard. 60:1-5. 

and R. L. STUCKEY. ee The eee Mionribution of the genus Najas 
(Najadaceae) in Ohio. Ohio J. Sci. 71:292-302. 

WILDE, ne J O. DE. 1961. The morphological evaluation and taxonomic value 

of the sa in Najas, with descriptions of three new Asiatic-Malaysian taxa. Acta 
ot. Neerl. 10:164—170. 

WILLIAMS, G. R. 1970. Investigations in the white waterlilies (Nymphaea) of 
Michigan. hercam Bot. 9:72-86. 

YANG, Y. 1974. New records of Najas in Taiwan. Taiwania 19:106-108. 


BASSIA (CHENOPODIACEAE) IN 
ORTH AMERICA 
SCOTT L. COLLINS AND WILL H. BLACKWELL, JR. 


Herbarium, Department of Botany, Miami University 


Oxford, Ohio 45056 


ABSTRACT 


A survey of literature and herbarium specimens indicates that two species of 
Bassta (Chenopodiaceae), both introduced, have become established members of 
: : : 


has spread extensively in western North ee since its introduction in News 
around 1915, and has maintained a limited distribution in the East following its 
introduction there around 1930. 


INTRODUCTION 


The introduction of non-indigenous species into a geographic region may 
alter the structure and composition of local plant populations, in some cases 
presenting deleterious ecological consequences. The history of Evchhornia 
crassipes, water hyacinth, in North America clearly demonstrates this point. 
All too frequently botanists do not adequately document the spread and 
effect of adventive species (particularly their ability to compete with local 
species), even though such studies are usually feasible, interesting, and 
sometimes valuable. Many questions on immigrant species often remain un- 
asked, not to mention unanswered, including: Where did the plant come 
from? When and how was it introduced? Where has it spread since its 
introduction, and by what mechanisms? What habitats does it invade? Has 
it become a permanent member of the flora? Will it probably continue to 
extend its non-intrinsic range? 

In reference to the above questions, the genus Bassia (Chenopodiaceae ) 
can be taken as an example. Bassia, a rather well-known and _ significant 
Eurasian genus of annual herbs, has about 10 species (ca. 90 species if the 
Australian Austrobassia is included in it); however, little is known (at least 
by any kind of comprehensive study) of its adventive distribution, partic- 
ularly its North American occurrence. In addition to distributional questions, 
the genus presents nomenclatural difficulties. Several authors (cf. Ijin, 1936; 
Standley, 1916; Mason, 1957) used an alternate generic name, Echinopsilon, 
in preference to Bassia. To add to the confusion, some taxonomists employed 
the name Bassia for a genus of Sapotaceae (cf. Airy Shaw, 1973). Nomen- 
clatural problems exist at the species level as well. Authorship of the combi- 
nation Bassia hyssopifolia, for example, has been credited to Kuntze (cf. 


SIDA 8(1): 57-64. 1979. 


58 


Gleason, 1952) or to Volkens (cf. Davis, 1952; Aellen, 1964). In our 
study, in addition to providing distributional information, we attempt to 
clarify the nomenclatural aspect. 


NOMENCLATURE 


The generic name Bassia was “originally” published on two separate occa- 
sions. Allioni in 1766 (Mélanges de Philosophie et de Mathématique de la 
Société Royale de Turin, p. 177) assigned the name to a genus in the 
Chenopodiaceae. Independently, Koenig proposed the name for inclusion 
in Sapotaceae, but did not publish it. Linnaeus subsequently validated 

Koenig's manuscript name by yeaa: it in his Mantissa (Mantissa 
Plantarum 2; Appendix, p. 563; 1771). As both names are validly published 
under the rules of nomenclature, Bassia Koenig ex Linnaeus (Sapotaceae ) 
must be regarded as illegitimate and to be rejected as a later homonym. 
Bassia Alliont is the correct name of the genus. 

A commonly used synonym of Bassia Allioni is Echinopsilon Moquin- 
Tandon. However, publication of this name (Annales des Sciences Naturelles 
2,2: 127, 1834) was much later than that of Bassia Allioni, and there would 
seem to be no reason for selecting it over Bassia. Chenolea Thunberg (Nova 
Genera Plantarum 10, 1781), which has likewise been preferred in some 
cases over Bassia (e.g., Bentham and Hooker, Genera Plantarum 3: 60, 
1880), also has a later date of publication, and, in addition, probably con- 
stitutes a distinct genus (cf. Airy Shaw, 1973) 

Credit for the combination B. Ayssopifolia should go to Kuntze (Revisio 
Generum Plantarum 2: 547, 1891) rather than Volkens (in Engler and 
Prantl’s Die Natiirlichen Pflanzenfamilien 3,1a: 70, 1893) on the basis of 
priority. The correct citation is B. byssopifolia (Pallas) Kuntze, based on 
Suaeda hyssopifolia Pallas. 


DISTRIBUTION 


Only two species of Bassia, B. hirsuta and B. hyssopifolia, have been 
recorded in North America. Though introduced, both species are Mas 
established, permanent members of the flora. The distribution of each i 
discussed separately below. Discussions are based in part on specimens os 
the following herbaria: A, F, GH, JEPS, MO, MU, NY, OSC, PH 
TEX, UC, US, and UT. We are grateful to the curators of these herbaria 
for the loan of specimens. 

The two species of Bassia occurring in North America may be distin- 
guished as follows: 

Bassia hirsuta: Floriferous portion of usher with flexuous axis; three sepals bear- 


ing a short, non-hooked, dorsal proce n fruit, the other two sepals unappendaged. 
Bassia bysopiola Floriferous axis oe all five sepals bearing a a hooked 
prickle 


Dy 


BASSIA HIRSUTA 

Bassia hirsuta (L.) Ascherson, the first Bassia species to appear in North 
America, maintains today a limited though well-established distribution 
along the east coast of the United States. Native to Europe (type locality: 
along the seacoast of Montpellier, France), B. Airsuta apparently entered 
our flora around the turn of the century. Earliest specimens are from south- 
ern New Jersey (Anglesea, Cape May County, 30 Aug 1900, Brown s.n., 
PH; Barnegat Bay Beach, Ocean County, 21 Jul 1900, Harshberger s.n., PH) 
However, early floristic studies (e.g., Robinson & Fernald, 1908; Britton & 
Brown, 1913) neglected this species. In fact, we find no mention of the 
plant in literature until Taylor (1915) alluded to its presence along the 
borders of salt marshes around New York City. By this time, its distribution, 
according to herbarium records, reached from Massachusetts to New Jersey. 
Initial spread was apparently rapid, as it was recorded in South Boston by 
1908 (12 Sep 1908, Knowlton sn. GH). Knowlton commented (on the 
specimen label) on the abundance of B. hirswta on “made land, in tidal 
brook and elsewhere.” Standley (1916) wrote “adventive on sandy seashores, 
New Jersey, and at Boston, Massachusetts,” indicating a possible disjunct 
distribution. As indicated above, however, the plant was already present in 
New York State by the time of Standley’s writing. In fact, a specimen 
(Bicknell 3968, PH) was collected from Long Island in September 1915, 
and it is likely that B. Airswta actually invaded the New York flora several 
years prior to this. 

According to more recent floristic literature (e.g., Fernald, 1950; Gleason, 
1952), the distribution extends from Massachusetts to Maryland, in coastal 
sandy or saline habitats. Herbarium records confirm this habitat statement 
but indicate a range south to Virginia (Chincoteague Is., 23 Jul 1938, Travis 
679, PH). Delaware is the only state between Massachusetts and Virginia 
from which B. hirsuta has apparently not been recorded either by literature 
or herbarium specimens. However, it is not likely that disjunction is actually 
involved. A careful search would probably show a limited occurrence of 
the species within the state’s coastal islands. 

As to the future of B. Airswta in North America, a limited increase in 
range with time is probable, confined primarily to saline coastal areas. This 
statement is based on the documented gradual increase (since its original 
rapid spread) up to the present, the availability of additional coastal area 
for colonization i particularly south), and the local success of populations 
as suggested by information on herbarium specimen labels. Several collectors 
indicated that B. hirsuta was densely established on various beaches (e.g., 
Knowlton sm., South Boston, 4-18 Sep 1909, UC). Fernald (1950) stated 
that the plant “appears indigenous,” an eee of its success in naturaliza- 
tion and its potential for persistence. 

The method of entry and spread is uncertain; thus, we are limited to 
speculation. Considering the plant’s lack of economic value and its relative 


60 


unsightliness, it was in all probability not imported purposefully. The immi- 
gration of B. hirsuta to this continent must be considered accidental an 
mose likely due to man. As its early distribution (around 1908) in North 
America was possibly disjunct (Massachusetts and southern New Jersey), 
it is not improbable that the species was introduced from Europe more than 
once. After it initial introduction and establishment, shore animals (especially 
birds) and tidal activity certainly contributed to a gradual range expansion. 


Representative Specimens (one specimen from each county recorded in our study 
is included): UNITED STATES: MARYLAND: Worcester Co.: brackish flat border- 
ing Assawomen Bay, Ocean City, 16 Sep 1941, Herman 10769 (MO, NY, US). 
MASSACHUSETTS: Barnstable Co.: Monument Beach, Bourne, 18 Aug 1921, Blake 
8363 (TEX); Dukes Co.: Penikese Island, 31 Jul 1947, Mouwl ind sal 3247 (PH); 
Norfolk Co.: Wollaston, 21 Sep 1918, Blake 7195 (US); Suffolk Co.: South Boston, 
12 Sep 1908, Knowlton sn. (GH). NEW JERSEY: Atlantic Co.: Atlantic City, 
25 Sep 1906, Wheeler s.n. (US); Camden Co.: Cooper's Point R.R. Freight Yard, 
Camden, 19 Aug 1932, Witte s.n. (PH); Cape May Co.: Anglesea, 30 Aug 1900, 
Brown sm. Sr Cumberland Co.: Delaware Bay, Moore’s Beach, Robinson’s 
Beach, s. of Delmont, 22 Jun 1933, Long 40685 (PH); Essex Co.: Port Newark, 
23 Sep 1933, ined and Clausen 497 (NY); Ocean Coe Barnegat Bay Beach, 21 
Jul 1900, Harshberger sn. (PH). NEW YORK: Nassau Co.: Long Beach, East 
Point, Long Island, 19 Sep 1915, Bicknell 3968 (PH); Suffolk Co.: Point-O-Woods, 
Long Island, 9 Sep 1921, Ferguson 879 (NY); Queens Co.: Jamaica Bay Sa 

ueens, Long Island, 7 Sep 1958, Monachino 629 (NY). VIRGINIA: Accomack C 
Chincoteague Island, 23 Jul 1938, Travis 679 (PH). 


BASSIA HYSSOPIFOLIA 


Bassia hyssopifolia (Pallas) Kuntze apparently entered the flora of North 
America 15 to 20 years later than B. Airsuta. Like B. hirsuta, B. hyssopifolia 
must be regarded as introduced in North America; it is native to parts o 
Europe and Asia (type locality: Caspian Sea region). Unlike B. hirsuta, 
B, hyssopifolia appeared first in western North America. The earliest refer- 
ence we found to B. Sighs ia in sores (Blake, 1922) cites Fallon 
(Churchill County), Nevada (28 Jul 1919, Tidestrom 10755, US) as the 
site of earliest collection. A search of i floristic literature (e.g. Jepson, 
1923; Rydberg, 1906, 1917; Standley, 1916) yielded no mention of B. 
hyssopifolia. It is included, however, for Nevada by Tidestrom (1925) and 
in more recent western floras (e.g., Harrington, 1954, Kearney & Peebles, 
1960; Munz & Keck, 1959; Hitchcock & Cronquist, 1973; Weber, 1972). 
In general, the plant is said to be well-established, especially in saline or 
alkaline habitats varying in altitude from below sea level to several thousand 
eet. 

The earliest North American specimen in a herbarium is that of T. H. 
Kearney (US), 1917, from Fallon, N 
to Blake. As the plant was probably eae a year or so prior to collection, 
1915 would be reasonable as to the approximate time of introduction. After 
establishment in Nevada, B. byssopifolia spread rapidly in all directions. 


61 


Based on herbarium records, the following would be the sequence of estab- 
lishment: California, 1919; Washington, 1921; Utah, 1924; New Mexico, 
1926; Montana, 1927; Texas, 1928; Oregon. 1932; Arizona, 1936; Colorado, 
1936; Idaho, 1937; British Columbia, 1937; Wyoming, 1939; and South 
Dakota, 1959. It is probable that B. hyssopifolia entered Oregon considerably 
earlier than records show, thus providing access to Washington, Idaho, and 
Canada. According to Moss (1959), this species has been found in Alberta. 

The means of introduction and dispersal of B. hyssopifolia seems relatively 
clear. As the earliest plants were discovered in rangeland near the Fallon 
experimental Station in Nevada, it is conceivable that propagules entered 
originally from Eurasia as a seed contaminant. Bassia hyssopifolia grows well 
in soil too alkaline for crops (Blake, 1922) and thus finds a suitable niche 
in impoverished abandoned fields. The sepals persist around the ripened 
fruit, and each bears on its back a hooked spine (hence the common name 
“five-hook bassia”). These small hooks attach the one-seeded fruits readily 
to clothing, livestock, or wildlife, providing an effective dispersal mechanism. 
Establishment in dry, alkaline or saline environments where competition is 
minimal would seem to aid survival and necessitate regarding B. hyssopifolia 
as a permanent, and sometimes “weedy,’ member of the North American 
flora. As stated by Munz and Keck (1959), B. Ayssopifolia is “becoming a 
common weed in rather alkaline places through much of Calif. and in other 
w. states.” 

Some 15 years after its introduction in western North America, B. 
Ayssopifolia appeared in coastal eastern North America. The earliest her- 
barium specimens from the east are dated 1936 (Massachusetts, 5 Sep 1936, 
Bean sn., GH; and New York, 3 Sep 1936, Monachino 161, NY ). However, 
literature (Palmer, 1930; Bean, 1937) points to an earlier arrival. Palmer 
(1930) documented the spontaneous growth of B. hyssopifolia in an aban- 
doned quarry near the Arnold Arboretum in the Boston area. Later (1936), 
Bean discovered the plant near Fish Pier and the freight yard area of South 
Boston and also alluded to its occurrence several years earlier near Worcester, 
Massachusetts. Based on the evidence, 1929-1930 is the probable time of 
introduction of B. hyssopifolia in the east. There are no firm facts by which 
to confirm whether this appearance in the eastern United States constituted 
a second introduction from Eurasia or a transmittal from the western United 
States. Bean reported the Fish Pier locality as a vacant lot where wool-waste 
was thrown out and hinted at an introduction from southwestern Asia. 
However, this information provides an insufficient basis for making final 
judgment as to the source of the introduction. Regardless, B. hyssopifolia 
has maintained a limited distribution on the east coast. Seymour (1969) 
contended that the plant is rare in Massachusetts, being found in only four 
locations: Chelmsford, South Boston, Westford, and Worcester. The two 
New York counties in which it grows are both on the western end of Long 
Island. Thus, it seems safe to assume that B. hyssopifolia occurs in scattered 


62 


locations from Boston to New York City and that it will remain in the 
eastern flora, perhaps not extending its range to any appreciable extent. 


Representative Specimens (one specimen from each poe eeu in our study 
is cited): CANADA: BRITISH COLUMBIA: Cariboo Three Mile Lake, S of 
Clinton, 12 Aug 1956, a Parmalee and Taylor pee (GH. NY); oa 
Rupert Co.: 1 mi E of Inverness, 11 Aug 1958, Taylor and eck 3752 (UC 
Yale Co.: near alkaline sae below Sugarloaf Hills, Kamloops, 17 Oct 1965, Krajina 
65101713 (NY 

NITED STATES: ARIZONA: Maricopa Co.: Arlington, 12 Aug 1936, McLellan 
and Stitt 1009 (US); Navajo Co.: Kayenta, 11 Sep 1938, Eastwood and Howell 
6541 (US). CALIFORNIA: Alameda Co.: Newark, 21 Oct 1966, O’Brien s.n. (UC); 
Colusa Co.: Maxwell, 1 Oct 1929, Jepson a (UC); Fresno Co.: 1 mi W, 2 
mi S of Fresno, 23 Oct 1919, Hall sn. (UC, US); Glenn Co.: S of Willows, 17 
Jun 1934, Howell 12247 (GH); Imperial between Brawley and Imperial, 17 
Oct 1941, MacBride and Drouet 4515 (F, UC); Inyo Co.: Independence, 4 Aug 
1940, Kerr s.n. (TEX); Kern Co.: Bakersfield, 13 Nov 1926, Bauer s.n. (UC); 
Lassen Co.: Honey Lake Valley near Wendel, 12 Jul 1934, Howell 12568 (F); 
Los Angeles Co.: Long Beach, 12 Nov 1928, Bauer s.n. (UC, US); Merced Co.: 

os, 27 Sep 1921, Kennedy sn. (UC); Modoc Co.: Alturas, 

10 Oct 1935, Wheeler 4035 (GH, NY, US); Mono Co.: 4 mi SE of Fales Hot 
Springs, 9 Sep 1934, Crum 1712 (UC); Orange Co.: West Newport Beach, 27 
May 1932, Booth 1130 (UC); San Benito Co.: Silver Creek, 4 Oct 1966, Twissel- 
mann 12877 (OSC); San Bernardino Co.: Searles Lake at Trona, 20 Oct 1964, 
Twisselmann 10310 (OSC); San Diego Co.: San Diego, 18 Jul 1934, Youngberg 
sm. (UC); San Francisco Co.: § of Hunter’s Point, San Francisco, 3 Jun 1956; 
Howell 31466 (OSC); San Mateo Co.: E of Redwood City, 27 Aug 1949, Nobs 
and Smith 1617 (UC); Santa Clara Co.: Palo Alto Yacht Harbor, 12 Sep 1932, 
Ferris 8267 (UC); Siskiyou Co.: Shasta Valley 4 mi E of Grenada, 21 Oct 1934, 
Wheeler 3332 (F); Solano Co.: Suisun City, 9 Oct 1932, Booth 2071 (F); Stanis- 
laus Co.: W side of Modesto, 1 Oct 1928, Dudley s.n. (F); Tulare Co.: 1235, 
R24E, Sec 30, 2 mi SE of Stoil, 29 May 1940, Bradshaw 333 (TEX); Ventura Co.: 
Ocean beach near mouth of Ventura River, 29 Sep 1945, Pollard s.n. (UC). COLO- 
RADO: Alamosa Co.: Alamosa, 2 Sep 1936, Ramaley 15908 (RM); Bent Co.: 
Prowers, 3 Sep 1940, Cory s.n. (GH); Boulder Co.: just E of Boulder Reservoir, 
19 Sep 1965, Weber 12972 (TEX, NY, UT); Eagle Co.: Wolcott, 17 Sep 1960, 
Perdue 4062 (US); Montezuma Co.: Corker 29 Aug 1947, Durham s.n. (F). 
O: Ada Co.: 4 mi N of Grandview, 19 Jun 1940, Davis 2063 (F, NY, UC); 
Bingham Co.: Springfield, 4 Sep 1937, Christ 8946 (NY); Cassia Co.: mouth of 
Raft River, 24 Aug 1940, Christ 11803 (NY); Custer Co.: 1 mi NW of May, 
20 Aug 1944, Hitchcock and Muhlick 11304 (NY); Franklin Co.: ca. 15 mi NNW 
of Preston, 6 Sep 1961, Bright 61-73 (RM); Gem Co.: 12 mi W of Emmett, 6 
Aug 1937, Christ 8526 (NY); Lemhi Co.: 4 mi S of Tendoy, 18 Aug 1956, Baker 
14642 (NY); Owyhee Co.: Homedale, 18 Aug 1928, Haegete s.n. (NY); Power 

fo.: S mi W of Pocatello, 12 Jul 1937, Christ 8364 (NY); Twin Falls Co.: 2 mi 
E of Buhl, 14 Jul 1937, Christ and Ward 8179 (NY); Washington Co.: 5 mi S$ 
of Weiser, 25 Jul 1940, Davis 3003 (F). MASSACHUSETTS: Suffolk Co.: waste- 
land, South Boston, 5 Sep 1936, Bean s.n. (GH). MO ae Phillips Co.: Na- 
tional Wildlife Refuge, Bowdoin, 26 Sep 1940, Hotchkiss sn. (US); Sanders Co.: 
Perma, 17 Sep 1927, Kirkwood 2497 (GH, RM). NEVADA: eae Co.: Fallon, 
18 Aug 1917, Kearney sm. (US); Clark Co.: across from hospital, Boulder, 8 Jun 
1967, Mears 1942 (TEX); Elko Co.: 7 mi N of Currie, 10 Sep 1937, Train 969 
(F, MO, UC); Humboldt Co.: Ashdown Mine, Pine Forrest ns 22 Sep 1934, 
Train 189 (US); Lander Co.: 10 mi W of Austin, 30 Aug 1937, Goodner and 


63 


Henning 1294 (F, MO); Lyon Co.: T12N, R23E, Sec 36, Wellington, 20 Sep 1937, 
Johannsen 1279 (UC); Mineral Co.: S end of Walker Lake, 6 Sep 1938, Archer 
6963 (UT); Nye Co.: Golden Eagle Mill NW of Ione, 20 Aug 1940, Beach 1031 
(UC); Storey Co.: 4 mi E of Virginia City, 10 Sep 1937, ieee 524 (UC); Washoe 
Co.: Pyramid Lake, 7 Sep 1930, Abrams 12917 (F, GH, UC); White Pine Co.: 
TISN, RGS8E, Sec 19, 13 Aug 1964, Holmgren and ened oe (GH, NY). NEW 
MEXICO: Dona Ana Co.: Mesquite, Mesilla Valley, 7 Aug 1930, Fosberg 53831 
(UC); Sandoval Co.: Bernalillo, 11 Jul 1926, Arséne and Benedict 16799 (F, US). 
NEW YORK: Bronx Co.: Pelham Bay Park, N.Y. City, 10 Oct 1954, Monachino 
Y); Queens Co.: Newton Creek, Queens, Long Island, 3 Sep 1936, Mona- 

chino 161 (US). OREGON: Gilliam Co.: Olex, 15 Oct 1938, Cooke 12112 (OSC); 
Grant Co.: Canyon City, 31 May 1951, Farrell sn. (OSC); Harney Co.: Malheur 
Lake, 30 mi S of Burns, 13 Aug 2 7, Hansen sm. (OSC); Lake Co.: 1932, Hill 
sm. (OSC); Malheur Co.: Ontario, 7 Sep 1928, Larsen sn. (GH, OSC); Morrow 
Co.: Irragon, 1 Aug 1933, Peck 18100 (OSC); Sherman Co.: 29 Sep 1955, Thomp- 
son sm. (OSC); Umatilla Co.: Hermiston, Nov 1932, Best s.2. (OSC); Union Co.: 
North Powder, 15 Aug 1958, Sidor s.n. (OSC); Wheeler Co.: 3 mi W of Mitchell, 
20 Jul 1934, Peck 18641 (OSC). SOUTH DAKOTA: Jones Co.: 1 mi § of Murdo, 
28 Aug 1959, Wiggins 14974 (US). TEXAS: El Paso Co.: N of El Paso, Sep 1938, 
Kelly sn. (GH); Hudspeth Co.: Trans Pecos, 2 mi SE of McNary, 18 Aug 1942, 
Waterfall 3971 (GH); Reeves Co.: Pecos, 4 Sep 1939, Cory 32958 (GH); Ward 
Co.: Barstow, 20 Jul 1928, Cory 1454 (GH). UTAH: Box Elder Co.: Sulphur 
Springs, 15 Aug 1929, Garrett 5423 (UT); Carbon Co.: 10 mi E of Wellington, 
7 Aug 1940, van Cott 78 (NY); Davis Co.: Farmington Bay Waterfoul Manage- 
fate Area, 8 Jul 1966, Arnell sn. (UT); Duchesne Co.: waste places, 2 Se 
8, Harrison and Garrett 8886 (UC); Salt Lake Co.: saline flats, 26 Sep 1924, 
ee gece (US, UT); Sanpete Co.: Ephraim, 15 Sep 1934, Olsen sn. (UC); 
Sevier Co.: SW corner of airfield S of Richfield, 16 Sep 1944, Holmgren and Evans 


Utah Co.: Utah Lake, Provo, 6 Oct 1935, Larson 6678 (US); ua dan Co.: 

T43S, R16W, Sec 13, between fields and Virgin River at Bloomington, 9 Aug 1963, 
Christian 1112 (UT); Wayne Co.: Henry Mountains, 15 Aug 1966, Everitt Bee 

Weber Co.: Ogden Bay, 29 Jul 1955, McHugh s.n. (NY). WASHIN 

Grant Co S end of Soap Lake, 15 Aug 1943, Eyerdam 6389 (UC); Okanogan ae 
between Omak and Okanogan, 15 Sep 1933, Fiker 1426 (US); Whitman Co.: 
Pullman, 23 Sep 1925, Pickett sn. (GH, US); Yakima Co.: Yakima, 3 Oct 1921, 
Nelson 1002 (US). WYOMING: Albany Co.: Laramie, 31 Aug 1939, Meunscher 
and Meunscher 15931 (GH); Sweetwater Co.: Flaming Gorge Reservoir at Firehole, 
20 Aug 1968, Porter and Porter 10599 (RM); Washakie Co.: W of Worland, 31 
Jul 1962, Nichols 516 (RM). 


REFERENCES 
ee P. 1964. In: Tutin et al. Flora Europaea. Vol. 1. University Press, Cam- 


bri 
ae ‘SH HAW, H. K. 1973. Willis’ dictionary of the flowering plants and ferns (8th 
d.). University Press, Cambridge 
BEAN. R. C. 1937. Two rare eed Rhodora 39: 208 
BLAKE, S. F. 1922. Two new western weeds. Science ieee 455-456. 
RO : 3 


States, Canada, and the British possessions oe ed.). Vol. 2. Scribner’s ae 
DAVIS, R. J. 1952. Flora of Idaho. Brigham Young University Press, Provo, Utah. 
FERNALD, M. L. 1950. Gray’s manual of botany (8th ed.). American om Co., 

New York. 


64 


GLEASON, H. A. 1952. The New Britton and ae illustrated flora of the north- 
eastern a States and adjacent Canada. Vol. 2. New York Botanical Garden, 
New Yc 

HARRINGTON, H. D. 1954. Manual of the plants of Colorado. Sage Books, Denver. 

HITCHCOCK, C. L. and A. CRONQUIST. 1973. Flora of the Pacific Northwest, an 
cae manual. ees of eae Press, Seattle and London 

Kom and 


ILJIN, M. 1936. In omarov B. K. Shi shkin. Flora USSR. Vol. 6. Aka- 
ae Nauk SSSR: Nostow Lesiage ad. 

JEPSO 1923. A manual a the flowering plants of California. Associated 
ae Sot, Berkeley 

KEARNEY, T. nd R. H. ohio 1960. Arizona flora. University of California 
Press, Beikeley al Los Ange 

MASON, 1957. ora oF the marshes of California. University of California 
Press, Berkley and os Angeles 


MOSS, e H. 1959. Flora of a a. University of Toronto Press, Toront 

MUNZ A. and D. D. KECK. 1959. A California flora. University a California 
Press, Berke and Los Angeles 

PALMER, E. J. pol he eae flora of the Arnold Arboretum. J. Arnold 
ee ae 63-11 

ROBINSON, B. L. an M. L. FERNALD. 1908. Gray's manual of botany (7th ed.). 
American Book , New os rk. 

RYDBERG, P. A. 1906, F ora of Colorado. Bull. Colorado State Uni 

1917. oe of the Rocky Mountains and adjacent gis Pe iohed . 

the author, New Y 

sti , F.C. 1969. The flora of New England. Charles E. Tuttle Co., Rutland, 


STANDIEY, P. C 1916. In: North American flora. Vol. 21(1). New York Botanical 
Garden, New York. 
TAYLOR, N. O1S Flora of the vicinity of New York. Mem. New York Bot. Gard. 
By 


168 
TIDESTROM, 1925. Flora of Utah and Nevada. Contr. U. Herb. Vol. 25. 
WEBER, W. 1972. Rocky Mountain flora. University of Gils Pe Boulder. 


A NEW SPECIES OF ERIGERON (COMPOSITAE) 
FROM COAHUILA 


GUY L. NESOM 


Department of Botany, University of North Carolina, 
hapel Hill 27514 
Present address: Department of Botany, Ohio State University, 
Columbus, OH 43210 


ERIGERON chiangii Nesom, sp. nov. (Fig. 1) 


Ab affini E. folioso Nutt. acheniis discique flosculis brevioribus, pappi setis minu 
numerosis, ie basalibus spatulato-obovatis margine integris dentatisve incrassatis 
vel revolutis diversa 


Perennial herbs from a few, slender, woody roots, with a cluster of basal 
leaves and a single erect stem borne at the ends of short (1-4 cm), ascend- 
ing, leafless, more or less woody, caudex branches, basal leaves usually de- 
ciduous by mid-summer. Erect stems 11-40 cm high, often reddish at the 
base, simple or usually with 1 to numerous branches, mostly branching from 
the upper 1/2 of the stem; at least the upper part of the stem (sometimes 
only under the heads) conspicuously and densely stipitate-glandular wit 
dark orange-tipped, capitate, Type C trichomes,’ otherwise completely glab- 
rous if not glandular, rarely with tiny, viscid-appearing Type B trichomes 
and a very few, spreading Type A trichomes. Basal leaves hirsute or rarel 
pilose with fine Type A trichomes, pubescence of margins not different from 
that of the lamina, except longer at base of the petioles; lower cauline leaves 
moderately strigose-hirsute, eglandular, losing Type A trichomes and usually 
becoming densely stipitate-glandular higher on the stem, basal leaves spatu- 
late to obovate or oblanceolate, subcoriaceous, margins slightly thickened or 
revolute, entire or with 1 or 2 pairs of pinnate teeth or lobes, apex rounded 
or barely mucronulate, usually about 3 cm long and 7 mm wide but up to 
7.5 cm long and 14 mm wide, petiole when present about 1/3-1/2 as long 
as the leaf; cauline leaves numerous, alternate, sessile, markely different in 
shape from the basal leaves or sometimes changing gradually on the lower 
1/3 of the stem, usually narrowly linear but sometimes oblong-oblanceolate, 
7-55 mm long, 0.7-4.0 (6.0) mm wide, apex acute to rounded-mucronulate, 
entire, sometimes with reduced axillary clusters of smaller linear leaves. 


‘Illustrations of trichomes referred to as Types A, B, and C can be found in earlier 
studies of Erigeron (Nesom, 1976, 1978.). 


SIDA 8(1): 65-70. 1979. 


66 


67 


Heads at ends of erect stems or branches, not clustered, or stipitate-glandular 
penduncles 14 cm long; involucres narrowly companulate-hemispheric, 
about 6-12 mm wide, composed of 30-46 imbricated phyllaries in about 
3 series, reflexing sharply after maturation and release of achenes; phyllaries 
lanceolate or oblanceolate, long acuminate, up to 0.8 mm wide, the inner- 
most up to 6 mm long, the outermost about half as long, brownish-green 
with stramineous margins, sometimes purplish on the upper half, the inner- 
most with narrow midribs and chartaceous margins, densely glandular with 
long Type C trichomes, without other vestiture; receptacles 1.5-3.0 mm 
wide, very shallowly convex, slightly tuberculate with raised carpopodial 
attachments points. Ray flowers, pistillate, fertile, 16-40 (50) in number, 
in 1-2 series, corollas mostly white, without a definite midstripe but often 
tinged with lavender, pink, or blue, 34 (5) veined, apex rounded, acute, or 
slightly notched, 5.0-9.5 mm long, remaining straight with wilting or age, 
the tubular portion about 1/3 of the total length, sparsely pubescent with 
long Type C trichomes, 0.9-1.5 mm long. Disc flowers perfect, fertile, about 
30-100 in number, corollas yellowish, tubular to narrowly funnelform, 
glabrous or sparsely pubescent with Type C trichomes, sometimes slightly 
constricted in the lower 1/3, slightly indurated in the midregion, 2.6-4.0 
mm high, 0.5-0.8 wide, the lobes 0.4-0.6 mm long and sometimes purple; 
anthers with thecae 1.2-1.5 mm long, with lanceolate apical appendages 
0.2-0.3 mm long; style with branches 0.6-0.8 mm long, including the tri- 
angular to deltate collecting appendanges 0.17—-0.26 mm long. Achenes 
oblong, laterally compressed, 1.3-1.8 mm long, 0.3-0.5 mm wide, tan, with 
2 thin, orangish ribs, sparsely strigose with duplex trichomes; carpopodium 
a narrow, cartilaginous ring; pappus of ray and disc achenes similar, simple, 
of 16-20 tawny, barbellate bristles, equalling or slightly shorter than the 
height of the disc corollas, sometimes with a few short, outer setae or bristles. 
Chromosome eee known from one collection (Powell, Patterson, and 
Ittner 1602) n 

TYPE: eee Cee Canon de la Madera, Sierra de la Madera, 
29 Mar 1975, Wendt and Lott 842 (HOLOTYPE:LL! ). 


ADDITIONAL COLLECTIONS EXAMINED: MEXICO. COAHUILA: Mina El Popo, 
ca. 2 km S$ of Canon El Diablo on dissected E slope of Sierra del Carmen, 29 Jul 
1973, Johnston, Chiang, Wendt and Riskind 11922 (LL); higher elevations in the 
Sierra Jardin (in the Sierra del Carmen), 1 Sep 7 ae 1210 (SMU); canyon 
Seo eee E from high pass N of Sierra Jasdia 6 mi E of Rancho El Jardin, 23 

May 1968, Powell, Patterson, and Ittner oO 
n=—18 (TEX); middle and upper ee oF Canon de la Hacienda, Sierra de la 
Madera, 10 May 1973, Johnston, Wendt, and Chiang 10956 (LL); ibid., 6 Aug 


Fig. 1. Erigeron chiangit Nesom, ca. 0.5X. 


68 


1973, Henrickson 12000 with Wendt (LL- 2 sheets); ibid., 5 Aug 1973, Henrick- 
son 11950 with Wendt (LL); ibid., 21 Sep 1973, Chiang, men and Jobnston 
9451E (LL); higher part and N slope of Sierra de la Madera, SE and SSE of Rancho 
Cerro de la Madera, 20 Sep 1972, Chiang, Wendt, and Johnston 9430A (LL); 
about 10 km SW of R. San Miguel at Ejido Santa Eulalia in Canon de los Burros, 
NE side of Serranias del Burro, es m alt., 2 Jun 1972, Chiang, Wendt, and Jobn- 
ston 7519C (LL); along small, rocky canyon, about 1 mi N of “La Laguna,” Canon 
del Mulato, Serranias del Burro, aie ca. 3500 ft, 8 a 1963, Gould 10596 (TAES); 
Rancho Agua Dulce, E slope of oe Coe de San Manuel, 30 Jun ou Wynd and 
Mueller 387 (MO, ARIZ, US, : Cafién de la Gavia, S of Ran de la Gavia 
(Sierra la Gavia), 2-3 Aug mer Tepicsten, Wendt, Chiang, and ae 12035B 

L); Mina El Aguirreno, N side of Sierra de la Paila, 5 Jul 19 73, Johnston, Wendt, 
and ae 11700 (LL); Canon El Cono, Sierra de la Gloria, 6 Sep 1976, Wendt 
and Riskind 1621 (LL); Canén de Milagro, E side of the Sierra de los Guajes, 
about 12 km W - oo de la Encantada, 10 Sep 1941, Stewart 1513 (GH); 
Sierra de San Marcos, opposite Los Fresnos, NE-facing slope, 4-5 Apr 1969, Keil, 
Meyer, Lewis and "Pinkav a P6071 (ASU—several branches of E. chiangit, 1 plant 
and a branch of E. sp.); Sierra de San Marcos, opposite Los Fresnos, NE-facing 
slope, 4-5 Apr 1969, Keil, Meyer, Lewis, and Pinkava 6120 (NY -l plant of 
E. chiangit, 3 epee of E. sp.). 


Dr. B. L. Turner first recognized that these plants deserved specific dis- 
tinction and suggested the epithet. This commemorates Fernando Chiang, a 
recent graduate student at the University of Texas who was part of the 
group, with Dr. Marshall C. Johnston, which made several of the first col- 
lections of this taxon during their reconnaissance of the Chihuahuan Desert 
region. Erigeron chiangii is endemic to mountains and canyons of east- 
central Coahuila within a radius about 120 kilometers of Monclova. Most 
commonly it has been collected in calcareous gravel or limestone cliff crevices 
at about 750-2500 meters elevation. It grows in chaparral, izotal, or on 
limestone outcrops in oak-pine-fir forests at about 2700 meters. 

The new species is very distinctive and easily recognized by the following 
characters: in most plants, a growth habit of clusters of hirsute, basal leaves 
at the ends of slender, leafless, ascending, caudex branches, basal leaves 
subcoriaceous with slightly thickened or revolute margins, cauline leaves 
mostly linear; 3 series of imbricate phyllaries; conspicuous, stipitate, orange- 
tipped glands on otherwise glabrous phyllaries and upper stems and leaves; 
relatively few, broad, white to bluish or pinkish ligules; and an essentially 
simple pappus of 16-20 bristles. The holotype (Wendt & Lott 842, from 
the Sierra de la Madera, Fig. 1) was chosen because it shows with relative 
completeness the nature of the roots, ascending caudex branches, basal leaves, 
and erect stems. Most of the plants of Erigeron chiangii on other sheets 
examined are broken off from the root system with only a portion of the 
caudex, and on other sheets collected later in the season, the basal leaves 
have been lost. The cauline leaves on the illustrated plant are more or less 
intermediate in shape for the species—leaves from other collections are 
either more narrowly linear or more broadly oblanceolate. All but one of 
the extremely linear-leaved collections of E. chiangi are from northwest 


69 


of Muzquiz in the Sierra del Carmen and Serranias del Burro. In that area 
the plants grow at elevations between the low and high extremes for the 
species. The single chromosome count for E. chiangii is tetraploid (n=18 
from the Sierra del Carmen; Powell and Powell, 1978), and checks of pollen 
size and stainability suggest that plants over its whole range are probably 
also tetraploid 

Plants of Erigeron chiangii from the two collections cited from the Sierra 
de San Marcos (Keil, Meyer, Lewis, & Pinkava 6071 and 6120) are slightly 
atypical. The lower stems and leaves have moderately dense, spreading or 
retrorse pubescence in contrast to the normal glabrous condition. These 
plants are mounted on the same sheets with another species of Erigeron, 
yet unnamed, which is similar in general appearance to E. chiangii, at least 
at this locality. Other sheets from the same location have only plants of 
the other species. It seems likely that some gene flow is occurring between 
these taxa in the Sierra de San Marcos, but the plants of E. chiangii show 
little other evidence of hybridization. In characters of pappus, ligule width 
and coloration, glandularity, involucral pubescence and imbrication, and leaf 
texture and margin, they are typical. 

Although limited hybridization may be occurring between plants of 
Erigeron chiangii and the unnamed taxon mentioned above, the closest rela- 
tives of E. chiangii appear to be in the Erigeron foliosus-breweri complex 
of California, Oregon, and Washington. At least some plants this com- 
plex are strikingly similar in growth habit to E. chiangii, as well as in their 
linear to narrowly lanceolate or oblanceolate, often crowded, cauline leaves, 
their few ray flowers with broad ligules, and their strongly imbricated 
phyllaries. All of these taxa grow in dry, rocky habitats. The pubescence and 
glandularity characteristic of E. chiangii may also be found in plants of 
E. foliosus Nuttall, particularly in var. confinis (Howell) Jepson and in 
var. hartwegii (Greene) Jepson, but no single plant of these taxa has been 
observed in which the whole combination of E. chiangii characters occurs. 
According to Cronquist (1947) polymorphy a ae on make taxo- 
nomic segregation ae members of the E. folrsosus-breweri ae 
one of the most di t_ problems in the genus. donee plants of E. 
chiangii are not highly ae in morphology over the range of the species. 
Furthermore, the basal cluster of relatively broad basal leaves of E. chiangii 
apparently never occurs in E, foliosws. The pappus bristles of E. chiangii 
are fewer in number, and the achenes and disc flowers average considerably 
shorter in length. The range of E. chiangi is separated by a gap of over 
1600 kilometers from the nearest populations of its putative relatives. Thus, 
despite a close similarity to some plants of E. folzosws, the distinctive combi- 
nation of characters in E. chiangii which are rather consistent over the range 
of the species, the several characters distinct from those of E. foliosws, and 
the wide geographic disjunction of these two taxa, support the recognition 
of E. chiangii as a distinctive species. 


70 


In the genus Erigeron there are at present few known species with rela- 
tively wide geographic ranges which lack diploids somewhere within their 
range. If E. chiangii is composed only of tetraploid plants, as it seems, an 
allochthonous origin for this species is strongly implied. Diploids have been 
reported for E. foliosus and E. breweri, and despite the wide disjunction, 
the remarkable similarity of E. chiangii to forms of these species strongly 
suggests that they represent the primitive stock from which E. chiangii had 
its origin. 

Evolutionary convergence might account for the similarities between these 
disjunct population systems. However, if Erigeron chiangii and Erigeron 
foliosus do not share a relatively recent common ancestry, the origin of 
E. chiangii is obscure since there is apparently no species in north central 
Mexico which is sufficiently phenotypically similar to E. chiangi to warrant 
its consideration as an ancestor. 

Plants of Erigeron bigelovii A. Gray from Texas and northern Mexico 
have strongly imbricated, glandular phyllaries and relatively few ray flowers 
and might be mistaken for Erigeron chiangii. However, the former species has 
a woody base without caudex branches, highly branched and wiry stems, and 
spreading pubescence on the stems and leaves. The glands of E. bigelovi 
are uncolored, not conspicuously stipitate, and smaller than those of E. 
chiangii. No intergradation has been observed between these species, and 
they probably are not closely related. 


ACKNOWLEDGEMENTS 


Thanks go to Larry Dorr and Rupert Barneby for their assistance with 
the Latin diagnosis and to Betsy Birkner, Staff Artist, Department of Botany, 
U.N.C., for the excellent illustration of the holotype. Appreciation is ex- 
pressed to the curators of the following herbaria from which collections were 


borrowed or studied: LL, TEX, SMU, ASU, ARIZ, MO, GH, US, and TAES 


REFERENCES 


reg A. 1947. of the North American species of Erigeron, north of 
Mexico. Britonia 6: 121-302. 
NESOM, 1976. ‘, new species 7 Erigeron (Asteraceae) and its relatives in 
southwestern Utah. oles 28: cae 
978. Erigeron hessit sp. nov. al Erigeron kuschei Eastwood (Compositae ), 
two “closely aa narrow se from the southwestern United States. Brittonia 
30:44 5. 
POWELL, M. and S. A. POWELL. 1978. Chromosome numbers in Asteraceae. 
Madrono os 160-169. 


VASCULAR FLORA OF CEDAR GAP LAKE AND 
ENVIRONS, WEBSTER AND WRIGHT 
COUNTIES, MISSOURI 


RALPH L. THOMPSON’ 


Department of Botany, Southern Illinois University 
Carbondale, Illinois 62901 


ABSTRACT 

The vascular flora of Cedar Gap Lake and environs in the Oak-Hickory Forest 
Region of southern Missouri are the focus of this study. An annotated list of 504 
taxa from 87 families are recorded from Cedar Gap, the second most elevated point, 
1694 feet, of the unglaciated, dissected Ozark Plateaus Province = the Interior 
Highlands. Twenty-one percent of the taxa are nonindigenous plants. One hundred 
twenty-six Webster County and 134 Wright County ees records are 
documented. 


The Oak-Hickory Association, the principal upland forest type in Mis- 
sourl, 18 most extensive and highly developed in the unglaciated Ozarks 
Region (Braun, 1950). This association appears to have originated when 
the increased aridity of the Western Interior eliminated more mesic species 
of the Mixed Tertiary Forest from the western region of the Eastern 
Deciduous Forest Formation (Braun, 1955). The most elevated portion of 
the highly eroded Ozark Uplift in the Ozark Plateaus Province of the Interior 
Highlands is situated within the boundaries of Webster County and Wright 
County 1 in southern oes (Steyermark, 1959). Fenneman (1938) stated, 


“Aside from the Boston Mountains the domed Ozark Plateau reaches its 
Tee level near its center . . . at Cedar Gap where the upland surface is 
almost 1700 feet... in Wright County, Missouri.” Steyermark (1963) 


reported Webster and Wright Counties as being among the least worked 
counties in documentation of vascular plants of the Missouri Ozarks. The 
purpose of this study is to present the first descriptive account of the vascular 
flora from an unique upland area within these counties, Cedar Gap Lake 
and vicinity. 

A floristic study of this area was conducted semimonthly from June, 1974, 
through July, 1976. Four plant community types were recognized: A dis- 
turbed tall grass prairie strip restricted to the St. Louis-San Francisco Rail- 
road right-of-way; a wet sedge-grass meadow around Cedar Gap Lake; a 
small oak-hickory woodland adjacent to the lake; and a waste ground com- 
munity of general open disturbed habitats without any permanent structure. 


= 1 Former address: ee of Life Sciences, Southwest Missouri State Univer- 
sity, Springfield, Missouri 65802 


SIDA 8(1): 71-89, 1979. 


72 


The 300 acre study site is located in the SW'Y4 SY% Section 15 and the NW 
N'Y N% Section 22 of Wright County, and the SW'4 Section 16 of Web- 
ster County, T28N, R16W, of the Cedar Gap Quadrangle (Figure 1). 

The topography consists of gently rolling to pa sloping uplands 
with a mean gradient of two to twelve percent. The elevated, dissected 
Ozark Dome forms a part of the main dividing ae nee the es 

the Missouri River to the north and those of the White River to the 
south. This dividing ridge may be traced from the western limit of Webster 
ee along the main line of the St. Louis-San Francisco Railroad. The 

rock is composed of resistant cherty, crinoidal limestones belonging to 
the Burlington-Keokuk Formations of the Osagean Series of the Mississip- 
pian System (McCracken, et al., 1961). Two small limestone sink-hole de- 
pressions within the study area are a feature of Karst topography. 

ie representative forest soil series, Wilderness Cherty Silt Loams, con- 
sist of moderately well-drained soils with cherty fragipans from nineteen to 
thirty-one inches in depth. Acid soil pH reactions range from 6.0 to 
The surface layers are medium acid, brown, cherty silty loams and the sub- 
soils are strongly acid, dark brown, cherty silty loams. The fragipans are pale 
brown, cherty silty loams overlying strong brown, cherty silty clays. The 
layers beneath the fragipans are very strongly acid, red to yellowish-red, 
cherty silty clays (USDA, Soil Conservation Service, 1975 

The continental climate in southern Missouri consists of wide seasonal 
variations in weather with typically hot, humid summers and cold winters. 

e mean annual temperature is 57.2° F. with mean temperatures of 31.4° 
during January and 78.6° F. through July. Total annual precipitation approxi- 
mates 41.38 inches. The mean minimum precipitation of 1.96 inches occurs 
in January and the maximum precipitation of 5.28 inches falls in May. The 
mean growing season is 192 days with the first and last freeze occurring 
around October 23 and re 14 (McQuigg, 1969). 

A tall grass prairie strip was sampled along a two-mile stretch of the 
St. Louis-San Francisco Railroad extending through Webster and Wright 
Counties. Several perennial prairie grasses and forbs, many weedy invaders, 
and a few woody members characterize this disturbed right-of-way situation. 
Local colonies of pioneer woody plants along fence row thickets have de- 
veloped into the Sassafras-Rhus-Diospyros Associes of Steyermark (1940). 
In terms of species, most belong to the Compositae, Gramineae, Leguminosae, 
Rosaceae, Euphorbiaceae, and Asclepiadiaceae. Native perennial grasses in- 
clude Andropogon gerardi, A. vir ginicus, Schizachyrium scoparium, ae 
canadensis, Sorghastrum avenaceum, Tridens flavus, Eragrostis spectabilis, 
Koeleria pyramidata, and Panicum oligosanthes vat. Veen evianai 

Cedar Gap Lake, a 43.7 acre body of water, was created when the main 
line of the St. Louis-San Francisco Railroad was built in 1870. This eutrophic 
lake abuts the railway embankment and lies on the boundary line between 
Webster County and Wright County. Cedar Gap Lake formerly served as a 


Oo 
He eal 
Sale 


Figure 1. Cedar Gap Lake and environs along the St. Louis-San Francisco Railroad. 
Adapted from the Cedar Gap Quadrangle, Missouri, 7.5-minute series, USGS, 1951. 


reservoir for steam locomotives from 1923 until 1951. 

Vegetational zones around Cedar Gap Lake reveal generalized stages of 
hydrosere succession. Dominant emergent species are notably scarce except 
for extensive colonies of Nelumbo lutea, Persicaria coccinea, and Potamogeton 
pieces Amphibious members along the muddy lake margin include 
clumps of Alisma plantago-aquatica, Sagittaria latifolia, Typha latifolia, and 
several species of Carex, Eleocharis, Scirpus, and Juncus. Common shrubs 
around the lake border are Salix nigra, S. caroliniana, and Cephalanthus 
occidentalis, 

The extensive wet sedge-grass meadow surrounding Cedar Gap Lake is 
dominated by members of the Cyperaceae, Gramineae, and Juncaceae. Com- 
mon sedges include Carex lupulina, C. lurida, C. frankii, C. scoparia, C. 
vulpinoidea, Dulichium arundinaceum, Eleocharis quadrangulata, and Fim- 
bristylis autumnalis. Principal lowland prairie grasses interspersed through- 
out this wetland zone are Spartina pectinata, Panicum virgatum, P. rigidulum, 
Phalaris arundinacea, Leersia oryzoides, Cinna arundinacea, Glyceria striata, 
Elymus virginicus, Agrostis stolonifera, and Tripsacum dactyloides. Frequent 


74 


rushes consist of Juncus interior, J. acuminatus, J. tenuis, and J. brachycar pus. 
Common taxa of dense thickets include Hibiscus lasiocarpos, Ribes missouri- 
ense, Polygonum scandens, Rubus pensylvanicus, Apios americana, Sambucus 
canadensis, and Rhus radicans. Pastures, cultivated fields, and an oak-hickory 
woodland border the sedge-grass meadow zone. 

The oak-hickory woodland at the eastern border of the lake adjacent to 
the railroad right-of-way has been derived from old pastures and orchards 
abandoned since 1905. Distinctive old field species, such as Juniperus vir- 
gimana, Rhus aromatica, Andropogon virginicus, and Danthonia spicata, are 
intermixed throughout the woodland. Important high understory and sub- 
canopy species include Sassafras albidum, Rhus copallina, R. glabra, Diospyros 
virginiana, Gleditsia triacanthos, Cornus florida, Amelanchier arborea, Ulmus 
rubra, and Viburnum rufidulum, Major canopy trees are Quercus stellata, 
Q. marilandica, Q. alba, QO. velutina, Carya tomentosa, C. ovata, C. texana, 
Prunus serotina, Nyssa sylvatica, and Celtis occidentalis. A few fruit trees, 
Pyrus malus and Prunus persica, persist as orchard remnants. 

According to Steyermark (1940), this subclimax forest type may be 
classified under the Quercus marilandica-Quercus stellata Associes converging 
through secondary succession into the probable climax forest type of more 
shade tolerant oaks and hickories, the Quercus alba-Ouercus velutina-Carya 
Association. Steyermark (1959) described the Oak-Hickory Association as 
the most complex edaphic association in the Ozarks Region. 

The waste ground community is rich in species composition, but it lacks 
the structural organization found in the older, more stabilized communities. 
Several open situations include pastures, fence row borders, cultivated fields, 
a fragipan road from the lake, highway roadsides along the right-of-way, and 
railroad track ballast. These disturbed habitats are normally colonized by a 
high number of ubiquitous native and nonindigenous weeds 

Voucher specimens are on deposit in the Herbarium of Southern Ilinois 
University (SIU) for documentation of the annotated list of vascular plants. 
Duplicate sets of specimens for most species have been placed in the Herbaria 
of Northeast Louisiana University (NLU) and Southwest Missouri State 
University (S . Identification and nomenclature for all taxa follow 
Steyermark (1963), except for several more recent taxonomic changes pre- 
sented by Correll and Johnston (1970). County voucher records are derived 
from county distribution maps found in the Flora of Missouri (Steyermark, 
1963) 

The annotated list is arranged alphabetically by family, genus, and species. 
Frequency of occurrence for each taxon is based upon field observations and 
field collections. This sbandanes scale had pliers progression values o 
common, frequent, sla ines and rare. An aster before a taxon 
represents a new ebster County voucher, and a double asterisk ( 
represents a new Wright oe voucher. A triple asterisk (***) before 
a taxon indicates documentation with new representative voucher records for 


75 


both Webster and Wright Counties. Naturalized and introduced nonin- 
digenous plant classifications follow the definitions of Lawrence (1951, p. 
279). These appear after the community category and are represented by 
N or J, respectively. 

The annotated list is comprised of 87 families, 295 genera, and 504 
specific and subspecific taxa from Cedar Gap Lake and environs. One gym- 
nosperm, three pteridophytes, 133 monocots, and 367 dicots are represented. 

ve largest families are: Compositae 67, Gramineae 66, Leguminosae 
39, Cyperaceae 34, and Rosaceae 22. One hundred-seven taxa (21%) are 
nonindigenous plants. One hundred twenty-six Webster County and 134 
Wright County distributional records are documented. The total species num- 
ber, 504, constitutes fifteen percent of the vascular flora of Missouri based 
on Steyermark (1963). 

The author wishes to express his gratitude to several individuals for 
their cooperation and assistance during this study. Identification and veri- 
fication of certain groups was generously given by Dr. Robert H. Mohlen- 
brock, Southern Illinois University, for the Graminae and Leguminosae; 
an r. Dan K. Evans, Marshall University, for the Cyperaceae and 
Juncaceae. Appreciation is extended to Dr. John W. Voigt and Douglas 

. Ladd, Southern Illinois University, for final review of the manuscript. 
Special thanks are given to Dr. Milton S. Topping, Southwest Missouri State 
University, for securing aerial maps and descriptive data on Cedar Gap Lake 
and vicinity. Publication of this paper was made possible by the James E. 
Ozment Award in Natural Science through the Southern Illinois University 
Foundation. 


ANNOTATED LIST OF VASCULAR PLANTS 


ACANTHACEAE 
Raellia humilis Nutt. Wild Petunia. Infrequent, railroad prairie strip. 
ACERACEAE 
Acer saccharinum UL. Silver Maple. Rare, margin of Cedar Gap Lake. 
AIZOACEAE 
** Mollugo verticillata L. Carpetweed. Infrequent, waste ground areas, N. 
ALISMATACEAE 
Alisma plantago-aquatica L. Water Plantain. Frequent, lake ma 
** Sagittaria latifolia Willd. Oe ear Frequent, lake margin; sink hole pond. 
AMARANTHACEAE 
*** Amaranthus hybridus L. Green Amaranth. Common, waste ground of railroad. N. 
***Froelichia gracilis (Hook.) Moq. NE Common, railroad track ballast. 
AMARYLLIDACEAE 
Hypoxis hirsuta (L.) Coville. Yellow Stargrass. Frequent, railroad prairie. 


ANACARDIACEAE 
Rhus aromatica Ait. Fragrant Sumac. Common, open woodland; fence rows. 
Rhus copallina L. Dwarf Sumac. Common, edge of woods; fence row borders 


Rhus radicans L. Poison Ivy. Common, oak woods; sedge thickets; fence rows. 


APOCYNACEAE 
*Apocynum cannabinum L. Indian Hemp. Common, railroad prairie strip. 
ARACEAE 
Acorus calamus L. Sweet Flag. Frequent, lake spillway; sink-hole pond. N. 
ASCLEPIADACEAE 
** Asclepias hirtella (Pennell) Woodson. Green Milkweed. Frequent, prairie strip. 
*Asclepias incarnata L. Swamp Milkweed. Infrequent, wet sedge- grass meadow. 


*Asclepias purpurascens L. ae le Milkweed. Infrequent, open prairie ae 
*** Asclepias syriaca L. Common Milkweed, Common, railroad prairie strip. 
sie fae tuberosa L. var. interior (Woodson) Shinners. Butterfly Weed. Frequent, 
ad prairie strip; roadsides 
i verticillata L. Borsenil Milkweed. Rare, railroad prairie strip. 
BALSAMINACEAE 


Impatiens capensis Meerb. Spotted Touch-me-not. Frequent, lake border. 


BERBERIDACEAE 
Podophyllum peltatum L. May Apple. Common, oak woodland; moist thickets. 


BETULACEAE 
Corylus americana Walt. Hazelnut. Frequent, fence rows; woodland edges. 
BIGNONIACEAE 
**Campsis radicans (L.) Seem. Trumpet Creeper. Rare, mesic lake thickets. 
BORAGINACEAE 


Hackelia virginiana (L.) J. M. Johnston. Stickseed. Rare, prairie ose 
*Lithospermum arvense L. Corn Cromwell. oe equent, waste ground area 
ieee ee canescens (Michx.) Lehm. Hoary Puccoon. Common, oe strip: 
*** Myosotis virginica (L.) BSP. Scorpion fees Infrequent, cultivated fields. 


CALLITRICHACEAE 


Callitriche heterophylla Pursh. Water Starwort. Rare, spillway overflow. 
Callitriche terrestris Raf. Terrestial Starwort. Rare, prairie strip. 


CAMPANULACEAE 
Campanula ae L. Tall Bellflower. Rare, wet sedge-grass a. 
Lobelia spicata Lam. pase ae Lobelia. Infrequent, railroad a 
Triodanis bi fore (R. & P.) Greene. ae Looking Glass. Rare, pale strip. 


Triodanis perfoliata (L.) Nieuw. Venus’ Looking Glass. Comming, prairie strip. 


CAPPARIDACEAE 


***DPolanisia dodecandra (L.) DC. Clammyweed. Frequent, railroad ballast. 


CAPRIFOLIACEAE 
Sambucus canadensis L. Elderberry. Common, sedge meadow; lake margin. 
Sym phoricarpos orbiculatus Moench. Coralberry. Common, fence rows; woods. 


Triosteum angustifolium L. Yellow Horse Gentian. Infrequent, shaded woods. 
Viburnum rufidulum Raf. Southern Black Haw. Frequent, oak woodland. 


CARYOPHYLLACEAE 

* Cerastinm Seiten Thuill. Mouse-ear Chickweed. acces: waste ground. N. 

*Cerastinm vulgatum L. Mouse-ear Chickweed. Common, waste ground areas. N 
*** Dianthus armeria L. Deptford Pink. Frequent, roads discbed ground. N. 
ss ase: officinalis L. Bouncing Bet. Rare, railroad track ballas 

ene antirrhina L. Sleepy Catchfly. Frequent, eee ae ae 

Slee stellata (L.) Ait. Starry Campion. Rare, moist shaded woodland. 

Silene virginica L. Fire Pink. Rare, cherty wae woodland. 

*Stellaria media (L.) Cyrillo. Common Chickweed. Common, waste ground. N. 


CELASTRACEAE 
Celastrus scandens L. American Bittersweet. Rare, oak woodland thicket. 
CHENOPODIACEAE 
*** Chenopodium album L. Lamb's Quarter. Common, open waste ground. N. 
CISTACEAE 
Lechea tenuifolia Michx. Pinweed. Infrequent, railroad prairie strip. 
COMMELINACEAE 
*** Commelina communis L. Dayflower. Infrequent, railroad track ballast 


Commelina erecta L. var. enguifole (Michx.) Fern. Hierba del Pollo. 2 aan 
lake border by old pumping statio 
Tradescantia obiensis Rat. ances Common, railroad track ballast. 


COMPOSITAE 
* Achillea millefolium L. Yarrow. Common, railroad prairie; La a Pale 
Ambrosia artemisiifolia L. Common Ragweed. Common, wa ound areas. 


Ambrosia bidentata Michx. Lance-leaf Ragweed. Common, open waste ground. 
Ambrosia trifida L. Giant Ragweed. Common, lake border; moist ditches. 
Rp anes plantaginifolia (L.) Hook. Pussy’s Toes. Frequent, open woods. 
nthemis cotula L. Mayweed. Infrequent, rai a aes ae N. 
By ieee minus (Hill) Bernh. Burdock. Rare, cultivated field edge 
Aster eat Nutt. var. angustatus Shinners. ee pees peed oak- 
hickory woodland; fence rows 
Aster joe ie Spreading Aster. Frequent, oak-hickory woodland. 
Aster pilosus Willd. var. demotus Blake. White Heath Aster. Common, railroad 
prairie strip; pastures; roadsides. 
** Aster praealtus Poir. Willow Aster. Rare, railroad right-of-wa 
* Aster rt ais (Nees ) T. & G. Wants bon land aes Rate, ia strip. 
as 


** Bidens iS iL. Ueuckeiohts: ee ae sed ge-grass ee 
** Bidens polylepis Blake. Tickseed Sunflower. ee lake margin; oe meadow. 
Carduns nutans L. Musk Thistle. Infrequent, talleoad prairie; pastures = 
Centaurea maculosa Lam. Star Thistle. Infrequent, railroad track ballas 
Chrysanthemum ieieaninemun L. Ox-eye Daisy. Frequent, prairie ae a aaenee, 
N. 


Cic horium intybus L. Chicory. Frequent, roadsides; right-of-way strip. N. 


Coreopsis palmata Nutt. Prairie Coreopsis. Frequent, railroad prairie. 


78 


Echinacea pallida Nutt. Pale Coneflower. ap aaa railroad prairie. 
Ertgeron annuus ( Pers. Daisy Fleabane. Common, waste ground areas. 
Erigeron canadensi. rseweed. Common, ae disturbed habitats. 
Erigeron pu nee iviic ise Robin’s Plantain. Infrequent, cherty woodland. 
Erigeron strigosus Muhl. Whitetop. Common, roadsides; pastures 
**Eupatorinm je Seven L. Throughout. Rare, wet ped be erties meadow. 
**Enupatorinm serotinim Michx. Late Boneset. Frequent, scdge- Brass meadow. 
Gnaphalinm obtusifolium L. Sweet Everlasting. Frequent, prairie strip. 
*Gnaphalium purpureum L. Purple Cudweed. Infrequent, open oak woodland. 
* Heleastrum ihe N a (Ait.) Shinners. Heleastrum. Rare, prairie strip. 
Helenium flexuosum Raf. Purple-headed Sneezeweed. Common, prairie swales. 
** Helianthus annuus L. Common Sunflower. Rare, railroad track ballast. I. 
Helianthus grosseserratus Martens. Sawtooth Sunflower. Rare, lake margin. 
Helianthus hirsutus Raf. Bristly Sunflower. Common, prairie strip; fields. 
Helianthus mollis Lam. Ashy Sunflower. Common, railroad prairie strip. 
**Heliopsis helianthoides (L.) Sweet. Ox-eye. ee railroad praitte. 


ee erotheca eee (Nutt.) Shinners. Golden Aster. Rare, roadsi 
“Iva annua L. 1 Elder. Rare, fale border is i tracks. 
*Krigia biflora eo ) Blake. False Dandelion. Rare, cherty moist woods. 


*Krigta oppositifolia Raf. Dwarf Dandelion. Rate: spillway border 
**Kubnia eupatorioides L. False Boneset. Infrequent, railroad prairie ‘tan 


***Lactuca canadensis L. Wild Lettuce. Common, open waste ground. 
***Tactuca serriola L. Prickly ates Common, disturbed open napa. N. 
Liatris aspera Michx. Rough Blazing Star. Infrequent, prairie strip. 


Liatris pycnostachya Michx. Pranic Blazing Star. Infrequent, ie strip. 
Parthenium hispidum Raf. American Feverfew. Common, railroad prairie strip. 
Prenanthes aspera Michx. Rough White Lettuce. Rare, elie prairie. 
Pyrrhopappus carolinianus (Walt.) DC. False Dandelion. Picanene roadsides. 
Ratibida pinnata (Vent.) B Borne Gray-headed Coneflower. Rare, prairie strip. 
*Rudbeckia hirta L. Black-eyed Susan. Common, right-of-way; pastiires. 

udbeckia triloba L. Brown-eyed Susan. Tame svene oak woodland borders. 
j i i . Infrequent, right-of- fuge thickets. 


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Solidago nemoralis Ait. Old- field Goldenrod. Common, prairie strip; pastures. 
*Solidago petiolaris Ait. Goldenrod. Infrequent, field edges; woodland. 
Solidago rigida L. Stiff Goldenrod. Rare, railroad prairie strip. 
olidago ulmifolia Muhl. Elm-leaf Can aiea, eee ae WS 
***Taraxacum officinale Wiggers. Dandelion. Common, pastures; prairie ee N. 
**Tragopogon dubius Scop. Goat's Beard. Infrequent, pate track ballast. N. 
*Verbesina helianthoides "Michx . Yellow Crownbread. Infrequent, field edge. 
Vernonia arkansana DC. ee Ironweed. Infrequent, woodland thickets. 
Vernonia baldwinit Torr. Western Ironweed. Frequent, righ t-of-way. 
**Vernonia missurica Raf. Missouri Ironweed. Common, prairie strip; fields. 
Nanthium strumarium L. Cocklebur. Infrequent, cultivated field edges. 


CONVOLVULACEAE 
***Calystegia sepium (L.) R. Br. Hedge Bindweed. Infrequent, railroad thickets. 
** Convolvulus arvensis L. ame ‘Bindweed. Frequent, waste ground sites. 
Cuscuta gronovii Willd. Dodder. Rare, moist lake border on Solidago spp. 
Cuscuta pentagona Engelm. Dodder. Rare, prairie swale er 
***Cuscuta polygonorum Engelm. Smartweed Dodder. Precuiene lake margin; wet 
sedge-grass meadow on Polygonum spp. 


79 


* Ipomoea — (L.) Jacq. Ivy-leaved Morning Glory. Infrequent, waste ground 
habitats. 
Ipomoea a L. Small White yaameage Glory. Rare, pasture fence row. 
*Ipomoea pandurata (L.) G.F.W. Meyer. Wild Potato. Infrequent, field edges. 
** Ipomoea purpurea (L.) Roth. poate Morning Glory. Rare, railway ballast. I. 


CORNACEAE 


Cornus florida L. Flowering Dogwood. Frequent, subcanopy of oak woods. 


CRUCIFERAE 
Barbarea vulgaris R. Br. var. arcuata (Opiz) Fries. Yellow Rocket. Common, pas- 
ture fields; roadsides; right-of-wa 

** Brassica juncea (L.) Coss. Indian niece d; quent, waste ground. N. 

** Brassica Raber (DC.) L.C. Wheeler var. peers (Stokes) L.C. Wheeler. 

Charlock. Infrequent, railroad ballast. N. 

**Capsella bursa-pastoris L. Shepherd's Purse. Infrequent, waste ground. N. 

**Cardamine parviflora L. var. arenicola (Britt.) O.E. Schultz. Bittercress. Frequent, 
disturbed open ground; cultivated fields. 
Dentaria laciniata Muhl. Toothwort. Frequent, open cherty oak woodlan 

**Descurainea pinnata (Walt.) Britt. var. brachycarpa (Richards ) ree Tansy 

Mustard. Infrequent, railroad prairie strip. 
*Draba verna L. Vernal Whitlow Grass. Rare, roadside; cane! ballast. N. 

**T epidium campestre (L.) R. Br. Fieldcress. Rare, waste ground by tracks. N. 
Lepidium virginicum L. Peppergrass. Infrequent, pastures; en of-way. 
Sisymbrium officinale (L.) Scop. Hedge Mastard, Infrequent, railroad ballast. N. 

***Thlaspi arvense L. Field Pennycress. Frequent, roadsides; railroad ballast. N. 
*Thlaspi perfoliatum L. Perfoliate Pennycress. Rare, waste ground. N 


CUPRESSACEAE 


Juniperus virginiana L. Red Cedar. Common, oak woods; pastures; fence rows. 


CYPERACEAE 
**Carex annectans Bickn. Scdge. Frequent, moist ditches; sedge-grass meadow. 
** Carex artitecta Mackenz. Emmon’s Sedge. Rare, cherty open woods. 
** Carex bicknellii Britt. Bicknell’s Sedge. Infrequent, woodland border. 
*Carex bushii Mackenz. Bush’s Sedge. Frequent, railroad prairie strip. 
Carex cephalophora Muhl. Oval-headed Sedge. Rare, prairie swale. 
**Carex crinita Lam. Fringed Sedge. Infrequent, wet sedge-grass mcadow. 
Carex digitalis Willd. Sedge. Rare, oak-hickory shaded woods. 
Carex frankii Kunth. Frank’s Sedge. Common, lake margin; oe oo 
Carex gravida Bailey. Heavy Sedge. Rare, wetl foe sedge-grass mc 
Carex birsutella Mackenz. Hirsute Sedge. Infrequent, shaded oak ead 
*** Carex lupulina Muhl. Hop Sedge. Common, wet sedge-grass meadow. 
Carex lurida Wahlenb. Sedge. Common, lake margin; sedge-grass meadow. 
Carex meadii Dewey. Sedge. Common, railroad prairie strip. 
Carex molesta Mackenz. Sedge. Rare, roadside ditch along right-of-way. 
Carex muhlenbergii Schk. Muhlenberg’s Sedge. Rare, prairie swale by lake. 
Carex scoparia Beats Pointed Broomsedge. Common, sedge-grass meadow. 


*** Carex squarrosa L. Squarrose Sedge. Frequent, scdge-grass meadow. 
** Carex tribulotdes Wahlenb. Blunt Broomsedge. Frequent, sed ge- -grass meadow. 
m d 


Cyperus esculentus L. Yellow Nutgrass. Infrequent, cultivated field edges. 
Cyperus ovularis (Michx.) Torr. Hedgehog Clubrush. Frequent, prairie strip. 


80 


*Cyperus psendovegetus as Umbrella Sedge. Rare, wet sedge-grass meadow 
Cyperus strigosus L. Flatsedge. Common, lake margin; sink-hole pond border. 
**Dyalichlum a ela (L.) Britt. Three-way Sedge. Common, lake margins. 
-ocharis acicularis (L.) R. & 8. Wire Spikerush. Common, muddy lake margin. 

ikerush. Common, Ce margin. 


* * Eleocharis quadran gulata (Michx.) R. . Square-stemmed Spikerush. Common, 
muddy lake margin; sink-hole a 
*Eleocharis tenuts Se Id.) Schultes. var. verrucosa (Svenson) Svenson. Slender 
Spikerush. Frequent, lake margin; prairie swales 
Fimbristylis See (L & S. Autumn Sede: Common, sedge meadow. 
*Scirpus atrovirens Wil goes Bulrush. Infrequent, wet sedge meadow. 
*** Scirpus pendulus Muhl. Bulensh, Common, roadside ditches; sedge meadow 
**Scirpus validus Vahl. var. creber Fern. Great Bulrush. Frequent, tie spillway 
overflow; sink-hole pond margin. 


DIOSCOREACEAE 
Dioscorea villosa L, Wild Yam. Infrequent, shady oak-hickory woodland 


DIPSACACEAE 


Dipsacus sylvestris Huds. Common Teasel. Infrequent, waste ground. N 
EBENACEAE 


Diospyros virginiana L. Persimmon. Common, subcanopy of oak woods; fields 


EUPHORBIACEAE 
Acalypha gracilens Gray. Three-seeded Mercury. Rare, railway ballast. 
Acalypha virginica L. toe seeded M 

Croton capitatus Michx. Hogwort. Rar ste ground of right-of-way 
**Croton glandulosa L. var. leptentrionali Muell.-Arg. Sand Croton. 
cultivated field edge; railroad bal 
Common, waste ground habitats. 

e. Common, railroad prairie strip. 


Euphorbi : 
Tragia betonicifolia Nutt. Noseburn. Rare, moist woodland thicket by ioe 


FAGACEAE 
Quercus alba L. White Oak. Common, canopy of oak-hickory ear mers 
Common, oak 


Quercus marilandica Muenchh. Blackjack Oak. 
ang. Post Oak. Common, canopy of oak ann fence 


Quercus stellata 
Quercus velutina Lam. Black Oak. Common, canpy of oak woods; fence rows 


rows. 


GENTIANACEAE 
Gentiana puberulenta Pringle. Downy Gentian. Rare, railroad prairie strip. 
Sabatia angularis (L.) Pursh. Rose Pink. Infrequent, roadside. 


GERANIACEAE 
*Erodium cicutarium (L.) L’ Her. Storksbill. Infrequent, waste ground N. 
bill. Common, disturbed waste ground. 


Geranium carolinianum L. Crane 
Geranium maculatum L. Wild Chee Frequent, shaded oak woodland edges 


Infrequent, 


oods; fence rows. 


GRAMINEAE 


* Aegilops cylindrica Host. Jointed Goatgrass. Rare, railroad waste ground. N. 
Agropyron repens (L.) Beauv. Quackgrass. R 


ures. I. 

m. as mon, eee prairie strip. 
mmon, pastures; fields; rae strip. 

Ma Michx. Bove rtyarass. Sea ee right-of-way. 

** Aristida oe Poir. Three-awn Grass. Rare, railway ballast. 

* Aristida oligantha Michx. Prairie Three-awn. Frequent, anes waste ground. 

Rare, cultivated fields: railway balla 
*Bromus commutatus Schrad. Hairy Chess. Common, waste sae habitats. N. 


why Chess. Common, wast nd habitats; fields. N 
Cenchrus longispinus (Hack.) Fern. Sandbur. Rare, cultivated field edges 
nna arund Wo : 


Beauv. Poverty Oatgrass. Frequent, open ds. 
** Digitaria ischaemum (Schreb.) Muhl. Smooth ers Frequent, waste ground. 


<2 Dios sanguinalis (L.) Scop. Hairy Crabgrass. Common, open waste ground 
***Echinochloa crusgalli (L.) Beauv. Barnyard Grass. Frequent, moist waste ground 
Bian indica (L.) Gaertn. Goose Grass. Infrequent, area of tracks, N. 

‘El 


‘Ym Ue. pecan L. Canada Wild i oe railr 
MUS ViT gZinich cL. Virginia Wild R 


Glyceria striata (Lam.) Hitchce. 
deum 


** * Hor usillum Nutt. Little Barl ommon, waste ground; roadsides 

Sonam ae L. Barley. Rare, cultivated field edges; railroad ballast. I 

**Koeleria pyramidata (Lam.) Bea Junegrass. Infrequent, prairie stri 
Leersia oryzoides Sw. Rice Cutgrass. Common, wet sedge-grass meado 
Leptoc filiformis (Lam.) Beauv. Red Sprangletop. Rare, cultivated field 
*Lolium perenne 


e L. Perennial Paes Infrequent, roadside waste ground. I. 
Muhlenbergia schreberi J.F. 


mel. Nimblewill Muhly. ek roadsides. 


ommon, sedge meadow; prairie cules: 
m oligosanthes Schultes var. seribnerianum (Nash) Fern. Panic Grass. Fre- 
quent, railroad prairie strip. 

Panicum rigidulum Bosc. Munro Grass. Common, prairie swale; sedge meadow. 


mon, lake margin; prairie swales. 
** Paspalum aaa Poir. var. glabrum Vasey. Beadgrass. Infrequent, moist waste 
ground; roadsides 


82 


***Phalaris arundinacea L. Reed Canary Grass. Common, wet sedge-grass meadow. 
Phleum pratense L. Timothy. Frequent, pastures; cultivated fields. I 
Poa annua L. Annual Bluegrass. Rare, waste baa site. N. 
Com ie strip; pasture 
oo scoparium (Michx.) Nash. ‘Little Blu estem. Common, eis strip. 
Secale cereale L. Rye. Infrequent, roadsides; railroad = lat 
** * Setaria fabertt Herrm. Giant Foxtail. Co 


, open waste ground. N. 
** *Seraria glauca ( 


mo 

(L.) Beauv. Yellow Foxtail. Conca, waste ground. N. 

*Setaria viridis (L.) Beauv. Green Foxtail. Infrequent, railroad ballast. 

So rghastrum avenaceaum (Michx.) Nash. Indian Grass. Common, prairie strip. 
(L.) Moench. Sorghum. Rare, cultivated fields; right-of- “way. I. 


** Spartina pectinata Tink. Prairie Cordgrass. Piccie 
** Sporobolus eh ead (Torr.) Wood. aA Diopeed. Rare, pastures. 
Tridens flavus (L.) Hitche. Purpletop. oe 
*Tripsacum dali L. Eastern Gamagra vate requent, sedge meadow. 
*** Triticum aestivum L. Wheat. Rare, eaeae fields; railway ballast 
Vulpia octoflora ai Rydb. Six es Fescue. Frequent, peseaics: woods. 
**Zea mays L. Corn. Rare, railway bal nm 


, prairie strip; joallsides. 


HALORAGIDACEAE 


*Prosperpinaca palustris L. Mermaid Weed. Rare, sink-hole pond; lake spillway. 


HYPERICACEAE 
** Hypericum drummondii (Grev. & Hook.) T. & G. Nits-and-lice. Rare, prairie 


strip. 
Hypericum gentianoides (L.) BSP. Pineweed. Rare, railroad prairie strip. 
Hypericum mutilum L. Dwarf St. Johns-wort. Frequent, muddy lake margin. 
Hypericum perforatum L. Common St. Johns-wort. Frequent, prairie trip. N. 
Hypericum a. Lam. Spotted St. Johns-wort. Frequent, prairie strip. 


IRIDACEAE 
Sisyrinchium angustifolium Mill. Blue-eyed Grass. Rare, cherty woodland. 
** Sisyrinchinm campestre Bickn. Prairie Blue-eyed Grass. Common, railroad prairie. 


JUGLANDACEAE 
Carya ovata (Mill.) K. Koch. Shagbark Hickory. Common, canopy of oak woods. 
texana Buckl. Black Hickory. Frequent, canopy of oak-hickory woodland. 
Carya tomentosa Nutt. Mockernut Hickory. Common, canopy of oak woodland. 
Juglans nigra L. Black Walnut. Infrequent, open pasture lands 


Q 
as 


JUNCACEAE 
** Tuncus acuminatus Michx. Sharp-pointed Rush. Frequent, sedge meadow; swales. 
Juncus brachycarpus Engelm. Short-fruited Rush. Common, 


Juncus dudleyi Wieg. Dudley’s Rush. Rare, roadside ditches; sedge meadow 


Luzula bulbosa (Wood) Rydb. Wood Rush. Frequent, cherty open seed and, 


LABIATAE 
Cunila origanoides (L.) Britt. Stone Mint. Infrequent, cherty oak woods. 
edeoma hispida Pursh. Mock Pennyroyal. Rare, railroad prairie strip. 
Isanthus brachiatus (L.) BSP. False Pennyroyal. Rare, moist lake margin. 


***Lamium amplexicaule L. Henbit. Frequent, open ground; roadsides. N. 
***lLamium purpureum L. Dead Nettle. Infrequent, roadside embankments. N. 
"*“*Lycopus rubellus Moench. Water Horehound. Common, lake margin; sink-holes. 
“**Lycopus virginicus L. Virginia Bugleweed. Infrequent, sedge-grass meadow 
Monarda fistulosa L. Wild Bergamot. Frequent, woodland edges; prairie ste 
Perilla frutescens (L.) Britt. Beefsteak Plant. Rare, moist embarkment. N. 
Physostegia virginia ana (L.) Benth. False Dragonhead. Rare, shaded thicket. 
*Prunella vulgaris L. var. lanceolata (Bart.) Fern. Self-Heal. Common, field edges; 
pastures; r ae of-way; lake border. 
Pycnanthemum tenuifolium Schrad. Slender Mountain Mint. Frequent, prairie strip; 


border. 
Scutellaria lateriflora L. Mad-dog Skullcap. Infrequent, muddy lake margin. 
**Scutellaria ovata Hill. Heart-leaved Skullcap. Rare, ae oak woodland. 
Scutellaria parvula Michx. Small Skullcap. Infrequent, open prairie strip. 
Teucrium canadense L. Wood Sage. Common, coe edges; prairie strip. 


LAURACEAE 


Sassafras albidum oe Nees. Sassafras. Common, subcanopy of oak woodland; 
fence row thicket 


LEGUMINOSAE 


Amorpha canescens Pursh. Lead Plant. Rare, railroad prairie strip; fence row. 
Aptos americana a Ground Nut. ee wet Siceae’ Sa 


Cassia fasciculata Michx. eee Pea. Common, sar anes oo. 
Cassia nictitans L. Sensitive Pea. Infrequent, cherty open ds 
Cercis canadensis L. Red Bud. Infrequent, subcanopy of an hickory woods. 
*Crotalaria sagittalis L. Rattlepod. Common, railroad prairie strip. 
* Desmanthus illinoensis (Michx.) MacM. ex Robins. & Fern. Illinois Mimosa. 
Rare, prairie strip 
Desmodium ciliare UMuhl. ) DC. Hairy Tick Trefoil. Frequent, Prairie strip. 
*Desmodium illinoense Gray. Illinois Tick Trefoil. Frequent, ee strip. 
Desmodium marilandicum (L.) DC. Small-leaf Tick Trefoil. Rare, oak woods. 
Desmodium paniculatum (L.) DC. Panicled Tick Trefoil. Gonmion: prairie strip. 
Desmodium rotundifolium DC. Round-leaf Tick Trefoil. Infrequent, open woods. 
ens ae (Torr.) T. & G. Sessile-leaf Tick Trefoil. Frequent, rail- 
road right-of-way str 
Gleditsia pie LH Honey Locust. Infrequent, open oak woods; pastures. 
Lespedeza capitata Michx. Round-headed Bush Clover. Frequent, prairie strip. 
***Lespedeza cuneata (Dumont) G. Don. Sericea Lespedeza. Common, waste ground. 
I. 


Lespedeza procumbens Michx. Traili ing ae sae ae open woodland. 
***T eshedeza stipulacea Maxim. Korea spedeza. Frequent, waste ground. I. 
Lespedeza striata (Thunb.) H. & i. eave Lespedeza. Rare, roadsides. I. 
Lespedeza violacea (L.) Pers. Violet Prairie Clover. Frequent, railroad prairie. 
Lespedeza virginica (L.) Britt. Slender Bush Clover. Common, eae prairie. 
Medicago lupulina L. Black Medic. Infrequent, roadside waste grou N. 
*** Medicago sativa L. Alfalfa. Frequent, pastures; roadsides; field ee L 
Melilotus albus Desr. White Sweet Clover. Common, roadsides; pastures. I. 
*Melilotus officinalis (L.) Lam. Yellow Sweet Clover. Frequent, waste ground. I. 
Petalostemum candidum (Willd.) Michx. White Prairie Clover. Rare, prairie strip. 
Petalostemum purpurcum (Vent.) Rydb. Purple Prairie Clover. Rare, prairie strip. 


Psoralea psoralioides (Walt.) Cory. var. eglandulosa (Ell.) Freeman. Sampson's 
nakeroot. Common, railroad prairie stri 

Schrankia uncinata Willd. Sensitive Brier. Frequent, railroad prairie strip; open 
woodland. 

Strophostyles helvola (L.) Ell. Wild Bean. Rare, fence thicket; Sevan 

Stylosanthes biflora (L.) BSP. Pencil Flower. Common, open wooc ; prairic strip. 

Tephrosia virginiana (L.) Pers. Goat's Rue, Frequent, open woods; etre strip. 

*Trifolium campestre Schreb. Large Hop Clover. Common, waste ground, 

*Trifolium pratense L. Red Clover. Frequent, fields; roadsides. N. 

*Trifolium reflexum 1. Buffalo Clover. Rare, roadsides 

Trifolium repens L. White Clover. Common, dicnubed waste plac 

Vicia dasycarpa Tenore. Hairy-fruited Vetch. Infrequent, railroad ball ast. N. 


LILIACEAE 
Allium canadense L. Wild Onion. Frequent, pastures; railroad prairie strip. 
*Allinm sativum L. Garlic. Rare, railway ballast edge. I. 
inm vineale L. Field Garlic. Infrequent, roadsides; pasture fields. 
*** Asparagus officinalis L. Garden eran Infrequent, fielc edges. I. 
*Camassia scilloides (Raf.) Cory. Wild Hyacinth. Infrequent, prairie strip. 
Erythronium albidum Nutt. White Trout Lily. Infrequent, cherty oak woods. 
** Melanthinm virginicum L. Bunch Flower. Rare, wet sedge-grass meadow edge. 
acacia oe (L.) Britton. False Garlic. Common, prairie strip; fields; 


ww 
A. 


as 

™ 

~— 
a. 


odland e 
Poly, ee biflorum Sle Ell. Solomon’s Seal. Infrequent, shady oak woods. 
Smilacina racemosa (L.) Desf. False Solomon's Seal. Frequent, moist oak woods. 


Smilax bona-nox L. Catbrier. Infrequent, open woods; fence rows; roadside. 
Smilax hispida Muhl. Bristly Greenbrier. Infrequent, shaded oak woodland. 
LINACEAE 

Linum sulcatum Riddell. Yellow Flax. Rare, railroad prairie strip. 

LOASACEAE 
*** Menzelia oligosperma Nutt. Stick-leaf. Frequent, railroad ballast. 

LYTHRACEAE 

Cuphea petiolata (L.) Koehne. Clammy Cuphea. Frequent, open waste ee 

Rotala ramosior (1..) Koehne. Toothcup. Tafeequent: muddy lake margin 
MALVACEAE 

aun theophrasti Medic. Velvetleat. eae, railroad ballast; fields. N. 

Althaea rosea L.. Hollyhock. Rare, railroad ballast. I. 


anne digitata Nutt. Poppy Mallow. Rare, on cherty wor 
ee US eae oo Woolly Rose Mallow. Common, ire margin; sedge 


**Sida ante L. Prickly Mallow. Infrequent, cultivated field edges. N. 
MORACEAE 
** Morus alba L. White Mulberry. Rare, cultivated field edge; oak woods. I. 
NYCTAGINACEAE 
*** Mirabilis nyctaginea (Michx.) MacM. Four O'Clock. Common, railroad ballast. 


NYMPHACEAE 


**Nelumbo lutea (Willd.) Pers. American Lotus. Common, emergent in lake. 


85 


NYSSACEAE 
*Nyssa sylvatica Marsh. Sour Gum. Infrequent, subcanopy of oak-hickory woods 


OLEACEAE 
Fraxinus americana L. White Ash. Infrequent, oak-hickory woodland. 


ONAGRACEAE 
Gaura biennis L. Biennial Gaura. Infrequent, railroad prairie strip. 
* *Iudwigia alternifolia L. Seedbox. Common, margin; sink-hole pond border. 
Ludwigia palustris (L.) EL var. americana (DC.) Fern. Marsh Purslane. Com 
mon, muddy ae margin; sink-hole 
*Oenothera biennis L. Common Evening Primrose. Common, moist waste ground. 
** * Qenothera iat Hill. Ragged Evening Primrose. Infrequent, waste ground 
OPHIOGLOSSACEAE 
Botrychinum virginianum (L.) Sw. — Fern. Infrequent, moist oak wood- 
land; sink-hole pond embankmen 
ORCHIDACEAE 
*Spiranthes cernua (L.) Richard. Nodding Ladies Tresses. Rare, prairie strip 


*Spiranthes gracilis (Bigel.) Beck. Green-lipped Ladies Tresses. 


Rare, railroad 
prairie strip. 


OXALIDACEAE 
**Oxalis dillenii Jacq. Yellow Wood Sorrel. Frequent, waste ground. 
Oxalis violacea L. Violet Wood Sorrel. Frequent, open woods; railroad prairie 
PASSIFLORACEAE 


Passiflora lutea L. var. glabriflora Fern. Passion Flower. Rare, shaded woods. 


PHYTOLACCACEAE 
Phytolacca americana L. Pokeweed. Common, disturbed open ground. 
PLANTAGINACEAE 
Plantago avristata Michx. Bracted Plantain. Frequent, open disturbed areas. 
Plantago lanceolata L. English Plantain. infrequent! waste ground; ee N. 
i ds border. 
Plantago virginica L. Hoary Plantain. “Antreanent, railroad prairie ‘strip. 


POLEMONIACEAE 
Phlox pilosa L. Prairie Phlox. Infrequent, railroad prairie strip. 
POLY GALACEAE 
*Polygala incarnata L. Pink Milkwort. Rare, railroad prairie strip 
**Polygala sanguinea L. Field Milkwort. Common, railroad prairie strip. 
Polygala senega L. Seneca Snakeroot. Rare, moist shaded woodland by lake. 


POLYGONACEAE 
**Persicaria coccinea (Mubhl.) eas Water Smartweed. Common, 
sink-hole pond; Cedar Ga 
Ree enue (Michx.) Small. Mild Water Pepper. Common, lake 
margin rie swales; sedge-grass meadow 
Bocsicaia lapathitolia (L.) Small. Pale Smmareweed Frequent, lake margin. N. 


emergent in 


86 


Persicaria pensylvanica (L.) Sm Pinkweed. Common, lake margin; swales. 
Persicaria punctata (EI.) Small. eee Rare, lake border by spillway. 
**Polygonum aviculare L. Knotweed. Frequent, disturbed open waste ground. N. 
Polygonum scandens L. False Buckwheat. Frequent, wet sedge-grass meadow thick- 

et 


ets. 
Polygonum es eel L. Virginia Knotweed. Rare, moist shaded oak woods. 
Rumex acetosella L. Red Sheep Sorrel. Common, throughout waste ground. N. 
*Rumex see L: oe Dock. Common, waste ground habitats. N. 


POLYPODIACEAE 
Asplenium platyneuron (L.) Oakes. Ebony Spleenwort. Rare, cherty embankment. 
Pteridium aquilinnm (L.) Kuhn. Bracken Fern. Rare, cherty open woodland next 


PORTULACACEAE 
Claytonia virginica L. Spring Beauty. Frequent, open woods; pastures. 


POTAMOGETONACEAE 


Potamogeton diversifolius Raf. Pondweed. Common, emergent of Cedar Gap Lake. 


PRIMULACEAE 
Dodecatheon meadia L. Shooting Star. Infrequent, railroad aan strip. 
** ysimachia bybrida Michx. Loosestrife. Rare, lake margin thic 
Lysimachia lanceolata Walt. Loosestrife. Frequent, moist shaded ‘ceellal 


RANUNCULACEAE 
Anemone virginiana L. Thimbleweed. Infre ae cherty open woods. 
Anemonella thalictroides (L.) Spach. Rue Anemone. Rare, cherty open woods. 
*Delphininm ajacis L. Rocket Larkspur. Rare, noadide 
gas tial eh Mase Walt. var. crispum Perry. Wild Blue Larkspur. Infre- 
ue railrc ight-of-way. 
+ Delphiniuum tricorne Michx. Dwarf Larkspur. Rare, cherty moist woodland. 
Ranunculus abortivus L. Small-flowered Crowfoot. Common, waste ground. 
— fascicularis Muhl. var. apricus (Greene) Fern. Prairie Buttercup. Fre- 
uent, cherty oak woodland; prairie strip. 
ee laxicaulis (YT. & G.) Darby. Spearwort. Frequent, sedge meadow. 
ia asi micranthus Nutt. Small-flowered Buttercup. Rare, cultivated field. 
*Thalictrum revolutum DC. Waxy Meadow-rue. Infrequent, railroad prairie. 


RHAMNACEAE 
Ceanothus americanus L. var. pitcheri T. & G. New Jersey Tea. Frequent, railroad 
prairie strip; open woods. 
ROSACEAE 
Agrimonia parviflora Ait. Swamp Agrimonia. Rare, sedge-grass meadow edge. 
Amelanchier arborea (Michx. f.) Fern. Shadbush. Infrequent, oak-hickory woods. 
k 


Crataegus crusgalli L. Cockspur Hawthorn. Rare, fence row thicket. 

Fragaria virginiana Duchesne. Wild Strawberry. Common, prairie strip; woods. 

Geum canadense Jacq. var. camporum (Rydb.) Fern. White Avens. te 
71 } woods near lake. 


moist shade 
Gillenia stipulata (Muhl.) Trel. Indian Physic. Infrequent, cherty woodland. 
**Potentilla norvegica L. Rough ee Ae Rare, spillway _N. 
Potentilla recta L. Sulfur Cinquefoil. Common, disturbed g nine habita . 
Potentilla simplex Michx. Old-field Cinquefoil. Cute oak oe noe 


tairie strip. 
**Prunus hortulana Bailey. Hortulan Plum. Rare, fence row border. I. 
**Prunus mahaleb L. Mahaleb Cherry. Rare, oak woodland border. i 
Prunus munsoniana Wight & Hedrick. Wild Goose Plum. Rare, fence rows. 
*Prunus persica (L.) Batsch. Peach. Rare, remnant from old orchard. 
i y of Bak woods. 


> woods. I 
osa carolina L. Pasture Rose. Fre t, pastures; prairie strip; woods 
Rosa multiflora Thunb. Multiflora Rose frequent, fence rov ickets. I 


>, Infrequent, pastures; prairie strip. 
*Rubus flagellaris Willd. Northern Dewberry. Common, fence rows; prairie strip; 
woodland thickets. 
Rubus pa L. Black Raspberry. Rare, mesic shaded woodland thickets. 
**Rubus pensylvanicus Poir. Highbush Blackberry. Common, pastures; fields; rail- 
road prairie strip; roadsides; sedge thickets. 


RUBIACEAE 
Cephalanthus ee L. Buttonbush. 
Wa 


Common, lake margin; sink- aa pond. 
Diodia teres 


ugh Buttonweed. Common, waste ground habitat 
Galium aparine L. Sea Frequent, mesic oak woods; sedge ee 
**Galium obtusum Bigel. Blunt-leaf Bedstraw. Rare, shaded oak woods. 
**Galium tinctorium L. Stiff Bedstraw. Frequent, sink-hole pond border. 
**Galinm virgatum Nutt. Dwarf Bedstraw. Rare, railroad prairie strip. 
Hedyotis es Raf. Small Bluets. Common, pastures; prairie s 


tri 
Hedyotis longifolia (Gaertn.) Fosb. Long-leaved Bluets 


p. 
Infrequent, railroad 


prairie strip. 
Hedyotis nigricans (Lam.) Fosb. Narrow-leavec 


1 Bluets. Rare, cherty woods. 
SALICACEAE 
Salix uneasy Michx. Ward’s Willow. Frequent, lake margin. 
Salix humilis Marsh. Prairie Willow. eee arise strip; field edge. 


** Salix nigra Nich. Black Willow. Common, lake margin; sink-hole pond. 


SANTALACEAE 
Comandra richardsiana Fern. False Toadflax. Common, railroad prairie strip. 


SAXIFRAGACEAE 
*Heuchera richardsonii R. Br. Prairie Alumroot. Rare, 
*Penthorum sedoides L. Ditch Stonecrop. Common, lake m 


9 


railroad prairie 
argin; sedge Ge 
Ribes missouriense Nutt. Missouri Gooseberry. Frequent, Sik woods; fence rows. 
SCROPHULARIACEAE 
*** Agalinus tenuifolia (Vahl.) Raf. Slender False Foxglove. Frequent, lake margin. 
**Tindernia dubia (L.) Pennell. False Pimpernal. Infrequent, muddy lake margin. 
Mimulus ringens L. Monkey Flower. Frequent, sedge-grass meadow zone 
Pedicularis canadensis L. Lousewort. Infrequent, ence cherty woodland. 


SUS m : 
Veronica arvensis L. Corn Speedwell. Frequent, disturbed open ground 


**Veronica peregrina L. Purslane Speedwell. Infrequent, waste ground sites. 


88 


Veronicastrum virginicum (L.) Farw. Culver’s Root. Rare, railroad prairie. 


SOLANACEAE 
Datura stramonium L. var. tatula (L.) Torr. Jimson Weed. Infrequent, railway 
Oi er t; cultivated fields 
salis heterophylla Nees. Clammy Ground Cherry. Rare, railroad ballast. 
ven subglabrata Mack. & Bush. Smooth Physalis. Frequent, waste ground. 
**Solanum americanum Mill. Black Nightshade, Rare, cultivated fields. 
Solanum carolinense L. Horse Nettle. Common, pastures; roadsides; fields. 


TYPHACEAE 
Typha latifolia L. Cat-tail, Common, lake margin; prairie swales; ditches. 
ULMACEAE 
**Celtis occidentalis L. Hackberry. Infrequent, oak woods; sink-hole pond border. 
Ulmus alata _ chx. Winged Elm. Infrequent, open oak-hickory wood 
Ulmus americana L. American Elm. Infrequent, oak woods; pasture edges. 


Ulmus rubra L. ‘Slippery Elm. Common, oak woodland; “fence rows; pastures. 


UMBELLIFERAE 


Angelica venenosa (Greenway) Fern. Wood Angelica. Rare, prairie swale. 
*Chaerophyllum procumbens (L.) Crantz. Wild Chervil. Common, waste ground. 
Cicuta maculata L. Water Hemlock. Common, lake margin; sedge-grass meadow. 


*Pastinaca sativa L. Wild Parsnip. Rare, railway ballast. 
Polytaenia nuttallii DC. Prairie Parsley. Infrequent, Paileoad prai 

**Sanicula canadensis L. Black Snakeroot. Infrequent, shaded cae a oos 
Torilis japonica (Houtt.) DC. Hedge Parsley. Common, waste ground sites. N. 
Zizia aurea (L.) Koch. Golden Alexanders. Rare, wet sedge-grass meadow. 


URTICACEAE 
Boemeria handles (L.) Sw. False Nettle. Infrequent, damp sedge meadow. 
Pilea pumila (L.) Gray. Clearweed. Infrequent, shaded mesic lake border 


VALERIANACEAE 


Valerianella radiata (L.) Dufr. Corn Salad. Common, waste ground habitats. 


VERBENACEAE 
Phyla sous (Michx.) Greene. Frog-Fruit. Infrequent, lake marg 
*Verbena bracteata Lag. & Rodr. Prostrate Vervain. Rare, roadside to fe 
Verbena conden L. Rose Vervain. Infrequent, Pits d prairie strip. 
Verbena hastata L. Blue Vervain. Frequent, wet sedge-grass meadow 
Verbena simples ae: Narrow-leaved Vervain. Frequent, waste ground areas. 
Verbena stricta Vent. Hoary Vervain. Infrequent, pastures; roadsides. 
Verbena urticifolia L White Vervain. Common, lake margin; sedge meadow. 


VIOLACEAE 
**Viola missouriensis Greene. Missouri Violet. Rare, cherty woodland. 

Viola pedata L. Bird-foot Violet. Frequent, railroad prairie; oak woods. 
Viola pratincola Greene. Common Blue Violet. Frequent, fields; oak woods. 
Viola rafinesquit Greene. Field Pansy. Common, prairi i 
*Viola sagittata Ait. Arrow-leaved Violet. Infrequent, railroad prairi 


s. 


ot 
= 
uo] 
a) 
a 
.o 
c 
= 
oO 
wn 


89 


VITACEAE 
***Parthenocissus quinquefolia (L.) Planch. Virginia Creeper. Infrequent, oak- 
hickor 
Vitis aestivalis Michx. Summer Grape. Infrequent, oak-hickory woodlan 
Vitis cinerea Engelm. Grayback Grape. Frequent, oak-hickory woods; chicken 
Vitis vulpina L. Frost Grape. Infrequent, oak woodland thickets by lake. 


REFERENCES 
BRAUN, E. L. 1950. Deciduous forests of eastern North America. Hafner Publishing 
Sone New York. 596 pp. 
: 1955. The phytogeography of unglaciated eastern United States and its 
interpretation. Bot. Rev. 21: ae 375. 
CORR D. S. and M. C. NSTON. 1970. Manual of the vascular plants of 
pee pie Research Bee ates Renner, Texas. 187 
ae ee N. M. 1938. cae of the eastern United States. McGraw-Hill 
ook Company, ey York. 714 
LAWRENCE, G. M. ee Taxonomy of vascular plants. Macmillan Publishing 
Company, Inc., New York. 
McCRACKEN, M. H. et os 
logical Survey and W Mis 
MCQUIGG, J. D. ine Ciisopianhy of ae Unieed States, oe 60—23: Missouri. 
SDC, ae | Data Service, Washington, D.C. 
STEYERMARK, J. A. 1940. Studies ‘of the veer of MEE I. Natural plant 
associations and succession in the Ozarks of Missouri. Field Mus. Nat. Hist. Bot. 
475. 


Ser. 9:349— 


vations 
1961. Peg oe = of Missouri. Missouri Geo- 


: We sarge history of the Ozark Forest. University of Missouri 
~~ Press, s, Columbia, Missou pp. 
___.. 1963. Flora of Micur Iowa State ai Press, Ames, Iowa. 1728 pp. 
USDA, SOIL CONSERVA TION SERVICE. 1975. Southwest Missouri resource con- 
servation and development project plan, Columbia, Missouri. 131 pp. 


TAXONOMIC NOTES AND NEW SECTIONS OF 
CLITORIA SUBGENUS BRACTEARIA 
(LEGUMINOSAE) ’ 

PAUL R. FANTZ 
Fairchild Tropical Garden, Miami, Florida 33156 


Botanists have traditionally followed Bentham (1858) recognizing three 
natural groups in the genus Clitoria, treating them as sections, and ignoring 
Baker (1879) who elevated two of them to the level of subgenus. Species 
with woody habits (i.e. trees, tall shrubs, and woody vines) are usually in- 
cluded in section Clitorianthes Bentham. The name Clitorianthes is a super- 
fluous name for Bractearia Martius ex Bentham (1837). In recent mono- 
graphic studies of Clitoria, Fantz (1977) concluded that these three natural 
groups, on the basis of morphological and distributional data, supported b 
the scant cytological and developmental data available, should be treated at 
the subgeneric level. The subgenus of woody members contains half of the 
58 species in Clitoria, a number of which are endemic to certain refugia 
recognized by Prance (1973) and as yet undescribed. The 29 species of 
this subgenus can be segregated into four sections. This paper provides 
taxonomic notes on subgenus Bractearia and describes the new sections and 
two new species. 

In his revisionary treatment of Clitoria, Bentham (1847) recognized three 
sections. Section Bractearia Mart. was described for the first time by fruticose 
habit, trifoliolate leaves and bracteoles equal to or longer than the calyx. 
Bentham included five species in this section: C. amazonum Mart. ex Benth., 
C. acuminata Benth., C. racemosa Benth., C. poitaei DC. and C. Pe 
Benth. 

In article on plants collected by Schomburgk in British Guiana, 
Bentham (1839) substituted the name Dendrocyamus for Bractearia justify- 
ing the change by the fact that Bractearia was used as a generic epithet 
in the Rubiaceae and as a sectional epithet in the genus Chaetogastra 
(Melastomataceae ). 

Although Bentham (1858) continued to recognize three sections in 
Clitoria, he again substituted a new name for the woody section, now calling 
it Clitorianthes. No comment was made on the changed named, nor was 
reference made to any pre-existing names for this section. Bentham here 
changed the sectional diagnosis of section Clitorianthes to erect shrubs or 


— studies on Clitoria accomplished at the IFAS Herbarium, Dept. of 
ny, University of Florida, Gaines ville, Florida. Florida Agricultural Experiment 
Sater Journal Series No. 1344. 


SIDA 8(1): 90-94. 1979. 


o1 


tall climbers with three leaflets and a flat or slightly convex, coriaceous 
legume. He further divided the nine species in section Clitorianthes into 
two unnamed groups distinguished by the bracteoles. The first group in- 
cluded six species with bracteoles narrow or much shorter than the calyx. 
Of the six species included, only Clitoria arborescens (synonym: C. poitaei 
DC.) had figured in his original list of 1837. The second group was char- 
acterized by ovate coriaceous bracteoles subequal to the calyx. three 
species now placed in this group had been assigned to this section in 1837. 
Their names now were reported as C. amazonum Mart.. ex Benth. (synonym: 
C. acuminata Benth.), C. hoffmannseggii (a superfluous name for C. arborea 
Benth.) and C. racemosa Benth. 

Publications of the next century which included Clitoria were primarily 
of a floristic nature. Authors adopted Bentham’s treatment and used the 
name Clitorianthes for the section which included the woody species. Baker 
(1879), in a floristic treatment of legumes in India, elevated two of Ben- 
tham’s sectional names to the level of subgenus. Section Clitorianthes Benth. 
was not included in this revisionary change, presumably because none of 
its species occur in India. No other floristic work on Clitoria has recognized 
Baker’s treatment of subgenera, the genus always being divided into sections 
following Bentham’s treatment of 1858 

In the first monographic treatment of the genus in the past century, Fantz 
(1977) concluded that the three natural groups can be distinguished morpho- 
logically by the fruits and seeds, supported by differences in the leaves, 
calices, androecia, and gynoecia. Distributional patterns, presence or absence 
of cleistogamy, and the limited cytological and developmental data available 
all support the recognition of these three groups as separate entities, and 
at the level of subgenus in agreement with Baker. Section Clitorianthes 
Benth. is thus elevated to the level of subgenus and Bractearia is adopte 
as the oldest and only legitimate name published for this woody group. 
Bentham’s diagnosis of the woody group is modified to include additional 
characteristics which distinguishes it from the other two subgenera. 


CLIToRIA L. subgenus Bractearia (Mart. ex. Benth.) Fantz, stat. nov. 
Clitoria L. sect. Bractearia Mart. ex Benth., Ann. Wien. Mus. Natur. 2: 115. 1837. 


Trees, tall erect shrubs or treelets, and lianas. Leaves 3-foliolate, long- 
stalked (petiole 4-16 cm), large (leaflets commonly 8-28 cm long x 3-23 
cm wide); petiolules elongate (4-10 cm long). Inflorescence woody, panic- 
ulate, subpaniculate or racemose-nodose; usually several- to many-flowered, 
the flowers all chasmogamous. Calyx subcoriaceous, multi-nerved, persistent 
in fruit. Ovary elongate (1-2 cm). Staminal tube elongate (2.5-4 cm). 
Fruits flat or occasionally weakly convex around the seeds and depressed 
between them, ecostate, long-stipitate (stipe 12-33 mm), coriaceous, 6-25 
cm long x cm wide. Seeds smooth, compressed, orbicular or nearly so, 
7-16 mm in diameter; germination epigeal. 


oe 


LECTOTYPIC SPECIES: C. amazonum Mart. ex Benth. (LECTOTYPE: Brazil, Para, 
ad fluy. Amazonum prope Ponte de Mattary, Sep, Martins 2740, M). 


Bentham (1837) placed five species in section Bractearia when he first 
described the group. Of these, C. amazonum is designated as the lectotype 
because it was from the type collection that Bentham obtained the name 
Bractearia Mart. At Munich, six sheets of the type collection (M 12408- 
12413) are contained in one folder, sheet M 12408 bearing the identification 
Bractearta amazonica Mart. and annotated Clitoria amazonica by Bentham. 


KEY TO THE SECTIONS OF SUBGENUS BRACTEARIA 
1. Bracteoles 6-16 mm wide, coriaceous, 10-40 mm long and subequal to the calyx 
(it often hidden from view), rarely shorter; inflorescence terminal or axillary (ap- 
pearing with the leaves), paniculate to acacia the primary lateral aaa 


ie bear the pedicels either short (1-5 mm long) or elongate (to 30 mm long). 
2. Calyx strigulose with oe short Ane ant trichomes, to glabrate, or a 
pilose, its surface easily seen; inflorescence straight to slightly curved, paniculate 


to subpaniculate, ered Sements ascending in a nearly straight line (non 
flexuous); trees or occasionally tall erect shrubs, rarely with a climbing apex. 

1. Bractearta 

2. Calyx velutinous with short, eee a more or fe any echoes its surface 
hidden; inflorescence flexuous, at least toward the apex, racemose-nodose, inter- 
nodal segments weakly to strongly bent in the opposite direction; lianas, occa- 
sionally erect shrubs or treelets 2. Flexuosae 

1. Bracteoles 1-4, rarely 6 mm wide, giiceiae cis ena 2 12 mm ioue and short- 
r than the ie or rarely élongate and subequal to the calyx, but then always 
narrow (calyx conspicuous); inflorescence axillary, and then often appearing before 
the leaves and/or cauliflorous, racemose, the primary ase branches bearing the 
pedicels lacking or represented by a conspicuous knob to 1 mm long (=nodose). 
3. Calyx tube 6-13 mm long with minute lobes 1-3 (—5) mm long; legume slightly 
convexed around the seeds and ee aaa, ——e between them at maturity; 
trees or tall shrub- ae rarely lianas. 3. Brachycalyx 

3. Calyx tube 10-30 mm long with conspicuous lispes 4— 18 mm iba ; legume flat, 
sometimes raised around the seeds but not uci between them lianas or 
rarely shrubs-treelets . . ou Bats # ze 4, ‘Cualiiioras 


1. CLrroriA L. subgenus BRACTEARIA (Mart. ex Benth.) Fantz sect. BRAC- 
TEARIA 


Clitoria L. sect. Bractearta Mart. ex Benth., Ann. Wien. Mus. Natur. 2: 115. 1837. 
Clitoria L. sect. Dendrocyamus Benth., Ann. Ant. Hist. 3: 44. 1839; nom. on erfl. 
Clitoria L. sect. Clitorianthes Benth., Journ. Linn. Soc. 2: 41. 1858; nom. superfl. 


Abores vel interdum frutices alti. Inflorescentiae sansa ia vel subpaniculatae, 
lignosae; rami laterales primarii pedicellifert (1) 4-30 mm longi; axes centralis non 
flexuosus. Calyx pubescentia dispersa vel plerumque glabratus, aes brevibus, 
appressis, infrequenter pe Bracteolae coriaceae, calycem subaequantes et pler- 
um que ia. 6-1 m latae et (11-) 1 < mm longae. Legumen planum, 
coriaccum vel sublignosum, ee latum, (1.5-) 2-4 cm latum, suturis incrassatis; 
pubescentia glabrata plerumque vel ee Microscopicis uncinatis, magni- 
trichomatibus ribi repertis secus suturas. 


ee) 


Members of section Bractearia are commonly collected in forests along 
the Amazon River and its major tributaries. The section includes seven 
species (Fantz, 1977): C. arborea Hoffm. ex Benth.; C. amazonum Mart. ex 
Benth., C. fairchildiana Howard, C. nervosa Herz., and three to be described 
in a succeeding article. 


2. CLITORIA L,. subgenus BRACTEARIA (Mart. ex Benth.) Fantz sect. 
Flexuosae Fantz, sec. nov 


Frutices lignosi saepius volubiles raro arbores. Inflorescentia debilis vel valde 
flexuosa prope apicem, segmentis internodiorum abruptly flexuosis, subpanicula; ramis 
laterales primarii pedicelliferi inconspicuis, subsessilibus ad 6 mm _ longae. Calyx 
pubescentia conferta, trichomatibus brevibus, appressis, aliqguanto sericeis. Brae 
coriaceae, calycem subaequantes vel calyce paulo breviores vel raro calyce multo 
breviores, semper latae factae, (6—) 9-15 mm latae, 10-40 mm longae. Flores grandes 
6-8 (-9.5) cm. Tubes stamineus elongatus, 3.5-5 cm Bae antheris grandibus, 
2-3 mm longis. Legumen longistipitatum (stipes 2.5-4 cm lanum, coflaceum, 
1.5-2.5 cm latum, pubescentia appressa. Semina brunnea, incrassata, longitudine 
latitudine longiore 

OLOTYPIC SPECIES: C. flexuosa Fantz? (HOLOTYPE: Peru, Tarapoto, Feb. 1856, 
Spruce 4527, K-Hb. Bentham). 


ee of section Flexwosa have been collected from forest refugia in 
u and Eucador. The section includes three species (Fantz, 1977): C. 
oe Fantz, C. eae Macbride, and one to be described. 


3. CLITORIA L. subgenus BRACTEARIA (Mart. ex Benth.) Fantz_ sect. 
Brachycalyx Fantz, sect. nov 


rbores vel frutices alti, raro frutex lignosus saepius volubilis. Folia decidua, typice 
tempore florendi, superne ki uberula, rarius glabra. Stipulae deciduae vel caducae, 


lanceolatae, acutae, angustae, 1-2 mm latae. Inflorescentia nodoso-racemosa, axillaris, 
plerumque esa pay oe nye denudato visa praecox. Calycis tubus brevis, Le Dee 
cupulatis | tubularis), 7-15 mm longus, lobis late deltoideis minutis vel 
fere Lena 1-4 mm longis, 3-4 mm latis, latitudo longitudinem subaequalante 
vel ok ae ee ngiore. Bracteolae ane. 2-5 mm longae, 1.5-3 mm_ latae. 
Legumen pubescens, leniter vel valde depr rs inter semina tempore liberationis. 


ae ‘suborbiculat vel leviter longiora quam latiora, compressa, vulgo 7-13 mm 

diam —3 mm es data. 
HoLornc SPECIES: C. brachycalyx Harms (TYPE: Brazil, Rio Branco, Surumu, 
p de Serra de Maiacy Feb 1909, Ule 8398, B destroyed during war, photo at 


MO 675333. Isotype at K) 


Members of section Brachycalyx are collected from dry tropical forests or 
occasionally from open grassy areas of savannas at altitudes up to 900 m, 


* CLITORIA flexuosa Fantz, sp. A Clitoria aeecaeieee Macbride affinis a qua 
ee is et bracteis eset thas: eee longo-st ipitatis, tubo staminale breviore, 

oe revi-un rade s, et foliis plerumque infra subpilosis distinquitur. HOLo- 
cas : Peru, Tarapoto. pee volubil robust, flores rosei, Feb 1856, Spruce 4527 
(K- Hb ae Payee mae YPES: Hb. Hooker ne ee PARATYPES: Tarapoto, 
Spruc (K San Martin, aay Jui, Rio eee ca. 400 m, Se 
1934, oe eee coer 766344, GH, MO Tse & 1105521). 


94 


in northwestern South America and adjacent Panama. Isolated collections of 
one species have been made as far north as Chiapas, Mexico. The section 
includes seven species (Fantz, 1977): C. brachycalyx Harms, C. brachystegia 
Benth., C. dendrina Pittier, C. glaberrima Pittier, and three to be described. 


4. CLITORIA L. subgenus BRACTEARIA (Mart. ex Benth.) Fantz_ sect. 
auliflorae Fantz, sect. nov 


Frutices lignosi saepius volubtilis vel infrequenter frutices erecti. Inflorescentia vulgo 
cauliflora et axillaris, racemosa, plus minusve nodosa. Bracteolae amplitudine variae, 
| 


sed non grandae et calycem occultantes, typice calycis tubo breviores et angustae, 
1-3 (raro —6) mm latae. Calycis deed ne CONS picuos ferens, typice 4-13 (—27) 
mm longos. Legumen planum vel leniter depressum inter semina, pubescentia typice 


microuncinata cum ee ee dispersis, effusis vel eae interdum 
confertis cum trichomatibus uncinatis — iene vulgo suborbiculata, raro oblonga, 
3-5 mm incrassata, interdum com a. 

OLOTYPIC SPECIES: C. bees "Fante! (HOLOTYPE: French Guiana, Karouany, 
1857, Sagot 120, K-Hb. Bentham) 


Members of section Cawliflorae are found mostly in northern South 
America, one extending into Central America. The section includes twelve 
species (Fantz, 1977): C. arborescens R. Brown in Ait., C. coriacea Schery, 
C. javitensis (H.B.K.) Benth., C. leptostachya Benth., C. obidensis Huber, 
C. sagotii Fantz, C. selloi Benth. and five to be descri é 

Acknowledgement is made to Helen Correll for the Tata descriptions, 
and to Daniel B. Ward, Dana Griffin III, and Donovan Correll for reviewing 
the manuscript aa providing constructive criticisms. 


REFERENCES 


BAKER, J. G. 1879. es no. 76: Clitoria. In Hooker, Fl. Br. India, 208-209. 
L. Reeve & ae ndon. 

BENTHAM, G. 337, i de Leguminosarum Generibus. Ann. Wien. 
Mus. ee 2: oe 

839. Pameration of Pane eee by Mr. Schomburgk, British Gui- 
ana. re Wien. Mus. Nat : 4 
< er Synopsis - oe genus Cite Journ. Linn. Soc. 2: 33-44. 

FANT 1977. A monograph of the genus Clitoria (Leguminosae: Glycineae). 
08. bp Ph.D. Thesis, University of Florida. 

PRANCE, G. T. 1973. a, ae ae meek for a seg of Pleistocene forest 

bas 


refuges. in the Amazon ed on evidence from the distribution pattern in 
Caryocaraceae, Chrysobalanacea Sener ce Lecythidaceae. ACTA Ama- 
zonica 3 (3): 5- 


3 CLITORIA seein Fantz, sp. nov. A Clitoria javitensts Cee eae _ a 


qua calycis, leguminis, et ova rii trichomatibus prominentibus uncin pagina supra 
folit medionervum pipers um ferenti, eee subsessile, vexillo brevis Serene 
stylo ovario multo longiore et fructibus parvioribus, semina incras distinguitur. 


HOLOTYPE: Fr. Guiana, Karo ieee 1857, aes 120 (K-Hb. ee photo at S). 
IsOTYPES: GH,NY,S,U 37632 A,W 


NEW SPECIES OF CLITORIA SUBGENUS 
BRACTEARIA SECTION BRACTEARIA 
(LEGUMINOSAE) FROM PERU AND ECUADOR’ 
PAUL R. FANTZ 
Fairchild Tropical Garden, Miami, Florida 33156 


Members of Clitoria L. subgenus Bractearia (Mart. ex Benth.) Fantz sec- 
tion Bractearia are distinguished by the large coriaceous bracteoles which 
subequal and obscure the sparsely pubescent to glabrate calyx, (sub-) 
paniculate inflorescences in which the pair of pedicels are borne at the 
apex of the primary lateral branches, broad thick-sutured fruits, and by the 
tall, erect, woody habit (trees or shrubs commonly 5-30 m tall). Section 
Bractearia includes seven species (Fantz, 1977): C. arborea Hoffm. ex Benth., 
C. fairchildiana Howard, C. amazonum Mart. ex Benth., C. nervosa Herz., 
and three newly described below. 


1. CLITORIA juninensis Fantz, sp. nov. 

Frutex 2-4 m altus. Folia Fea subtus subvelutina; foliola obtusa, acumine 
brevi, 1-7 mm longo. Stipulae deciduae, 8-11 mm longae; stipellae 5-10 mm longae. 
inlorenennee subpaniculato-racemosae, axillares et terminales, multiflorae; rami 
laterales primarii pedicelliferi subsessiles ad 5 (-8) mm longi. Bracteolae amplae, 
coriaceae, oblongo- bee calycem subaequantes, 14-18 (—20) mm longae. Flores 
4-5.5 cm; vexillum extus fluvum, sericeum. Stylus 16-17 mm _ longae. Legumen 
sae planum, aces sutura incrassata. Semina fere orbiculata, anthracina, 

mpressa, laevis. 


Shrub, erect, 2—4 m tall. Branches elongated (vide Killip & Smith 23380), 
hollow, to 10 mm thick, juvenile branches subangular with dense, spreading, 
rufus trichomes, branches becoming terete and seemingly glabrate, with 
uncinate trichomes becoming more conspicuous (observe at 30X); bark dark 
brown, splitting in longitudinal strips, lightened beneath; axillary buds 1-2 
mm, scales glabrate. Leaves 3-foliolate, coriaceous, leaflets broadly elliptic, 
ovate-elliptic, or obovate, apex generally obtuse, rapidly narrowed to a short 
acumen of 1-7 mm, more or less mucronate, base broadly cuneate to rotund, 
midrib impressed above, glabrous or bearing short erect and/or uncinate 
trichomes, primary nerves of 10-16 (—18) pairs, upper surface dark green 
and glabrous, lower surface pale green and subvelutinous, lamina 8-22 cm 
long, 4-9 (-12.5) cm wide. Petioles subangular-terete, 4-11 cm ribbed, 
with moderately dense, spreading, short, rufus trichomes, rachis more com- 


ecuiices studies on Clitoria accomplished at the IFAS Herbarium. Dept. of 
Botany, University of Florida, Gainesville, Florida. Florida Agricultural F penmient 
Station Journal Series No. 1345. 


SIDA 8(1): 95-101. 1979. 


96 


pressed, (1.5—) 2-3.5 cm; petiolule subquadrangular, 6-10 mm, with dense, 
spreading, rufus trichomes. Stipules deciduous, lanceolate-deltoid, acute, 8-11 
mm long, 2-3 mm wide; stipels more persistent, linear, acute, 5-10 mm long, 
0.2-0.7 mm wide. Inflorescence subpaniculate-racemose, axillary and terminal, 
solitary, multiflowered, pubescence dense, trichomes short, spreading, rufus; 
central axis 5—23 cm long, lignose and nodose in fruiting stage, the primary 


Figure 1. Holotype of Clitoria juninensis (Killip & Smith 23380, NY). 


97 


lateral branches which bear the pedicels subsessile to ‘ (-8) mm _ long; 


pedicels paired, 4-7 mm. Bracts ovate, acute, reflexed in age, 4-7 mm long, 
2—3 mm wide, pubescence dense, trichomes appressed, eee ee large, 
coriaceous, oblong-elliptic, Oi bequal to calyx, 14-18 (- mm long, 7-10 


mm wide, pubescence appressed. Flowers all chasmogamous, 4-5.5 cm, white 
to pink with a bluish-tinge. Calyx sparsely pubescent, trichomes appressed, 
tube 13-17 mm long, 3-5 mm wide at the base expanding to 8-12 mm wide 
at the throat, lobes ovate-deltoid, acuminate, 5-7 mm long, 3-4 mm wide, 
ventral lobe narrow, 7-8 mm long. Vexillum pubescence dense, trichomes 
appressed, tawny, blade 3-3.5 cm wide, claw 8-9 mm. Alae white to pinkish, 
extending 6-8 mm beyond the carina, blade 18-23 mm long, 5-9 mm wide, 
claw 13-15 mm. Carina bluish, falcate, blade 9-13 mm long, 4 mm wide, 
claw 24-27 mm. Staminal tube glabrous, 27-32 mm long, free filaments 3-5 
mm long; anthers 1.5-1.7 mm long, 0.5-0.8 mm wide. Gynophore 3 mm; 
ovary ca. 14 mm long, 1.2 mm wide, pubescence dense, trichomes appressed, 
yellowish-white; style bearded, 16-17 mm long, geniculate 6-7 mm; stigma 
flattened, 1 mm diam. Legwme stipitate, slightly exerted beyond oe ae 
ish-brown, flat, coriaceous, glabrous, thick-sutured, 21 cm long, 2.5-3.2 cm 
wide; stipe 14-17 mm; beak to 4 mm; dehiscence causing valves to twist 

one-half of a turn. Seeds nearly orbicular, black, smooth, compressed, 1-2 
mm thick, ca. 10 mm long, 9 mm wide; 7 seeds per pod. Fig. 1 

TYPE COLLECTIONS: Pert. Shrub, 8—12 ft., with ae branches; standard and 
wings white, blue-tinged, keel blue, white ee La Merced, Dept. Junin, 700 
m, 29 May-4 Jun 1929 Killip & Smith 23380 (HOLOTYPE: NY. ISOTYPES: F 616214, 
US 1358082). PARATYPES: Clearing along Rio ae a neat “Hacienda 3”, Ciloata 
Perene, 600 m, Killip & Smith 25136 (NY); Pampa Whaley, Cpa Perené, 
prov. Tarma, 700-750 m, 14 Feb 1959, Ferreyra 13555 (WIS-2 sheets). 

Clitoria juninensis has close affinities with Clitoria andrei which is dis- 
tinguished by the elongated acumens of the leaflets, shorter stipules and 
tipels, somewhat smaller flowers with larger bracteoles, a nonsericeous 
vexillum, and by the tree habit. All known collections of C. jaminensis occur 
from the Dept. of Junin, Peru, where the species is an apparent endemic 
of highland scrub communities at elevations of 600-2000 m. 


2. CLITORIA andrei Fantz, sp. nov. 


ad 20 m alta. Folia — subtus subvelutina, foliolis abrupte acuminatis, 
l bre 


acumine eA, 1-2 cm longo. Stipulae caducae, 4 mm longae; stipellae 

—5 mm longo. Inflorescentiae elongatae, subpaniculato-racemosae, axillares et termi- 
nales, multiflorae; rami primarii latecales pedicelliferi a ad 2 mm longi. 
Bracteolae amplae, coriaceae, oblongae, obtusae, calycem subaequantes, 22-25 mm 


longae. Flores 4-4.5 cm; vextllum extus fuluum, cum PO Ne Secus nervos. 
Stylus 17-21 mm. Legumen ignotum. 


Tree to 20 m tall. Branches subquadrangular becoming terete, to 1 mm 
diam., hollow, juvenile branches with short appressed to spreading trichomes, 
becoming glabrate; bark light brown; axillary buds 15-2 mm, bud scales 


98 


ovate, acute, concave, strigose. Leaves 3-foliolate, coriaceous, leaflets ovate- 
lanceolate to elliptic lanceolate, apex abruptly acuminate, acumen 1-2 cm, 
mucronate, base broadly cuneate to rotund, midrib impressed to subimpressed 
above, often bearing short erect trichomes, primary nerves 13-17 pairs, 
upper surface dark green, glabrous, lower surface pale green, subvelutinous, 
lamina 7-20 cm long, 3-8 cm wide. Petiole subangular-terete to compressed 


Figure 2. Holotype of Clitoria andrei (Andre 4675, GH). 


oD 


above, 4-7 cm, pubescence of short, spreading trichomes; rachis subquad- 
rangular, 1-2.5 cm; petiolules quadrangular, 6-7 mm, trichomes. falcate. 
Stipules caducous, deltoid, acute, 4 mm long, 3 mm wide; stipels deciduous, 
linear, 2-5 mm long. Inflorescence elongate, subpaniculate-racemose, axillary 
and terminal, solitary, multiflowered, pubescence of axes dense, trichomes 
short, erect, rufus on juvenile portions becoming less dense and whitened 
with age; central axis 7-20 cm long, the primary branches which bear the 
pedicels subsessile to 2 mm long; pedicels paired, 6-8 mm. Bracts ovate, 
acute, reflexed in age, 4-G mm long, 1.5-2 mm wide, pubescence dense, 
trichomes rufus. Bracteoles large, coriaceous, oblong, obtuse, subequal to calyx, 
22-25 mm long, 9-12 mm wide, pubescence strigose. Flowers resupinate, all 
chasmogamous, pink with bluish tinge, 4-4.5 cm. Calyx pubescence sparse, 
trichomes short, appressed, tube infundibular, 14-16 mm long, 4-5 mm wide 
at the base expanding to 9-11 mm wide at throat, lobes deltoid, 5-7 mm 
long, 2-3 mm wide, ventral lobe 6-8 mm long. Vexillum indumentum of 
short, appressed, tawny trichomes confined primarily along the nerves, blade 
2.5-3 cm wide, claw ca. 12 mm. Alae extended 6-8 mm beyond the carina, 
blade ca. 16-18 mm long, 4-6 mm wide, claw ca. 15-17 mm. Carina falcate, 
blade ca. 9 mm long, 3-4 mm wide, claw ca. 24-27 mm. Staminal tube 
cue ae near apex, trichomes microscopic (observe 30X ), uncinate, 

tube mm long, free filaments 2-3 mm long, uncinate-trichomed; 

eee ee mm long, ca. 0.4 mm wide, connective apiculate. Gynophore 
3 mm, ovary ca. 14 mm long, 1.4 mm wide, pubescence dense, trichomes 
yellowish-white; style 17-21 mm, bearded, geniculate 6-7 mm; stigma discoid, 
0.5 mm diam. Legwme unknown. Fig. 2. 


— 


TYPE COLLECTIONS: Ec cuador. Inter Guayaquil and ee ce 60 m, July 1876, 
Andre 4675 (HOLOTYPE: GH. IsOTYPES: F 537681, 


Clitoria andrei has close affinities with Clitorta juninensis of Peru which 
is distinguished by the obtuse leaflets with a minute acumen, larger stipules 
and stipels which are more persistent, smaller bracteoles on somewhat larger 
flowers, a sericeous vexillum, and by the shrubby habit. The species is knowa 
only from its type locality in Ecuador. 


3. CLITORIA moyobambensis Fantz, sp. nov. 


Arbor. ee trifoliolata svbtus glabrescentia, foliolis acuminatis, acumine elongato, 
1.5-—3 cm lon Stipulae amplae, concavae, 10-12 mm longae, 4-6 mm latae; stipellae 
caducae. ee centiae axillares et terminales, subpaniculato-racemosae, multiflorae; 
rami laterales primariit pedicelliferi ae ae ad 2 mm longi. Bracteae 


coriaceae, lanceolatae, breviacuminatae, 8—/1 mm longae, 3—5 mm latae. Bracteolae 
amplae, coriaceae, oblongo-ellipticae, ee subaequantes, obtusae vel abrupte api- 
culatae, 12-15 mm longae. Flores 4—5 cm; vexillum extus fulvo-sericeum. Stylus 


brevis, 14 mm longo. Legumen ignotum 


Tree. Branches hollow, 4-7 mm diam., juvenile branches subquadrangular, 
longitudinally striated and sulcate, grooves shallow to deep, pubescence 


100 


prominent, trichomes uncinate (observe 30X) and short, falcate to appressed 
(conspicuous at 10X), branches becoming subterete and glabrous with age; 
bark medium gray, splitting into longitudinal strips, whitish-tan beneath. 
Leaves 3-foliolate, coriaceous, leaflets elliptic-lanceolate, tapering to acuminate 
apex, acumen 1.5—3 cm long, to 1.2 cm wide at broadly cuneate base, mid- 


Figure 3. Holotype of Clitoria moyobambensis (Mathews s.n., K-Hb. Hooker). 


101 


rib impressed above bearing uncinate trichomes, primary nerves 13-16 pairs, 
upper surface dark green, glabrous or with short trichomes along major 
nerves, lower surface light green, glabrous, lamina 12-15.5 cm long, 6-8 cm 
wide. Petioles conspicuously twisted at base, longitudinally striated and 
canaliculate, 4-5 cm long, pubescence scattered, trichomes minute, falcate to 
appressed; rachis similar, 4 cm; petiolules quadrangular, rugose, 8-9 mm. 
Stipules large, conspicuous, concave nearly halfway around stem, lanceolate- 
ovate, rapidly tapering above middle to acute apex, 10-12 mm long, 4-6 mm 
wide; stipels caducous, only one terminal stipel observed (2 mm x 0.4 mm). 
Inflorescence axillary and terminal, solitary, subpaniculate-racemose, multi- 
owered, trichomes subappressed; central axis twisting, longitudinally striated 
and caniculate, to 4 cm long (juvenile? ), primary lateral branches which 
bear the pedicels subsessile, 1 (—2) mm long; pedicels paired, 8-14 mm. 
Bracts large, conspicuous, lanceolate, short-acuminate, 8-11 mm long, 3-5 
mm wide, trichomes appressed. Bracteoles large, coriaceous, multinerved, 
oblong-elliptic, obtuse or abruptly apiculate, subequal to calyx, 12-15 mm 
long, 9-10 mm wide, pubescence ciliolate and strigose. Flowers resupinate, 
all chasmogamous, 4-5 cm. Calyx infundibular, glabrate, trichomes widely 
scattered, heii tube a ae eae multinerved, nerves eel 
13-15 mm long, 4-6 mm wide at the base expanding to 8-10 mm wide at 
throat, lobes deltoid, acute, 5-6 mm long, 4 mm wide, ventral lobe 7 mm 
long. 2 mm wide. Vexillum tawny-sericeous, blade 3-4 cm wide, claw 11 
mm. Alae and carina hidden by vexillum and/or damage, not clearly observed. 
Staminal tube glabrous, 27-28 mm long, free filaments 1-3 mm _ long; 
anthers 0.8 mm long, 0.5 mm wide. Gynophore ca. 5 mm; ovary 13 mm 
long, 1 mm wide, lateral surfaces densely pubescent, trichomes uncinate, 
sutural edges with 1 mm long, ascending trichomes; style 14 mm _ long, 
bearded, geniculate 6 mm; stigma dark, ca. 0.7 mm diam. with a ring of short, 
reflexed trichomes at its base. Legume unknown. Fig. 3. 

TYPE COLLECTION: Peru. Prov. Moyobamba, 1838. Matthews s.n. (HOLOTYPE: 
K-Hb. Hooker). 


Clitoria moyobambensis is easily distinguished from other species in the 
subgenus by the unusually large bracts and stipules, and the pubescence of 
the ovary, a type unique amongst those species with a tree habit. This species 
appears to have close affinities with Clitoria juninensis and Clitoria andrei 
which have similar floral structures. It is known only from the type locality. 

Acknowledgement is made to Helen Correll for the Latin descriptions, 
and to Daniel B. Ward, Dana Griffin II], and Donovan Correll for reviewing 
the manuscript and providing constructive criticisms. 


REFERENCES 


FANTZ, . R. 1977. A monograph of the genus Clitoria (Leguminosae: Glycineae). 
1066 pp. Ph. D. Thesis, University of Florida. 


FLORISTIC AND GEOLOGIC ASPECTS 
OF INDIAN MIDDENS IN SALT MARSHES OF 
HANCOCK COUNTY, MISSISSIPPI 


LIONEL N. ELEUTERIUS 
Botany Section 
and 
ERVIN G. OTVOS, JR. 
Geology Section 
Gulf Coast Research Laboratory 
Ocean Springs, Mississippi 39504 


ABSTRACT 


Sixty-two species of flowering plants were collected from Indian middens (shell 
mounds) located in the salt marshes of Hancock Co., Miss. The diversity of the 
flora depended on the height of the midden above the marsh surface and increased 
with increased elevation. The largest midden (Cedar Island), with an elevation of 

mS ' 


greatest diversity of species (47). Radiocarbon dating indicates that clam shells 
(Rangia cnneata) found on the surface of Cedar Island are over 2,600 years old. 
The geological history surrounding the establishment of the middens is discussed. 
Indicator plants for these Indian occupation sites 0.5 m or more above MLW are 
the calciphiles ail silicola, Erythrina berbacea, and Aesculus pav - These shell 
middens provide the only known habitat of Sageretia minutiflora, the first report of 
its occurrence in Misicibal 


INTRODUCTION 


Salt marshes were apparently utilized by prehistoric humans for a con- 
siderable length of time. Human occupation sites in the coastal areas of 
Louisiana were established as early as 12,000 B.P. (Gagliano, 1963). Pres- 
ently known cultural sites on the Mississippi Coast date back only to the 
last four millenia. The fact that these early people gathered shellfish as a 
food item from the estuarine waters and salt marshes is evident by the 
presence of numerous and extensive shell mounds, composed of the clam 
Rangia cuneata. Such shell deposits are “kitchen middens” (refuse heaps ) 
at periodic Indian occupation sites, located close to the source of food. These 
mounds are relatively enduring features in the changing coastal environment 
and represent the only remaining local sites of floristic disturbance by pre- 
historic man, not obscured by natural processes or European man. Over the 
past several years we have located five previously unreported and undisturbed 
shell middens in the salt marshes of Hancock County, Mississippi. Although 
Brown (1936) described the vegetation of 16 mounds and middens in 
Plaquemines and St. Bernard Parishes (delta areas of the Mississippi River ), 


SIDA 8(1): 102-112. 1979, 


103 


Louisiana, on the shell middens we have visited, we noted an array of plan 
species previously unreported in Louisiana or Mississippi. The Enea 
conditions that relate to the establishment of these middens are essential 
for the understanding of the flora’s development. The present paper is the 
result of a combined effort to study these middens floristically and geo- 
logically. 

he middens can only be reached by boat or pontooned helicopter. This 
isolation probably contributed to their preservation. A study of aerial photo- 
graphs indicated the location of suspected shell middens and was followed 
by a general survey of the salt marsh area where five middens in the tidal 
marsh east of the Pearl River and south of Cadet Bayou were selected for 
study (Fig. 1). Floristic surveys were conducted seasonally in 4, an 
periodically in 1975, 1976, and 1977. Geological information was taken 
from an ongoing, detailed geological study of the Mississippi coast includ- 
ing core drillings in the marshland and radiocarbon dating of the middens. 


GEOLOGICAL FRAMEWORK 


The geological evolution of southern Hancock County (Fig. 1) began 
after a long period during the Wisconsin glacial stage and the early Holo- 
cene epoch when the subject area was dry land. Transgressing mid-Holocene 
sea waters reached it at about 6,300-6,000 years ago. By about 4,000—3,500 
years ago, all of the present marsh area was inundated and a low dune ridge 
(Magnolia Ridge) formed along this new shoreline. With the first arrival 
of sea waters, silty-muddy and marsh deposits characterized by Ammotium- 
Haplophragmoides-Miliammina agglutinate foraminifers, formed in the low 
salinity nearshore waters influenced by fresh waters from the Pearl River. 
Oyster reefs developed in more saline waters further seaward, while the 
central and southern-southeastern parts of the present marsh area were cov- 
ered by the highest salinity waters. Open-Gulf nearshore-type, varied and 
rich foraminifer fauna (Elphidinm- Be ae Nonion-Ammonia) thrived 

Fi . The 


on the bottom sediments there (Fig. 2 igh energy conditions near 
the shore resuited in the development of ay shoals and in the subsequent 
emergence of low (1.5-4.0 m), narrow, elongated barrier islands: Point 


Clear and Campbell Islands. 

The sand for these developing islands was derived from the east along 
the Mississippi barrier island-shoal chain (via Cat Island—Square Handker- 
chief Shoal) and the Hancock barrier islands themselves become parts of 
another 50 km long barrier-shoal trend (Bayou Sauvage—Hancock County 
trend, Otvos 1973, Fig. 16) which extended deeply into the present New 
Orleans area. Growth of the large Mississippi River subdeltas (St. Bernard 
Parish) prevented further seaward progradation of Point Clear and Campbell 
Islands. This delta development south-southeast of the Hancock area event- 
ually cut off the sand supply. It severely restricted open marine water influx 
and also reduced wave energies. The decreasing salinities south of the two 


‘}X9] UT passndstp “suoneso] 


Udpprul UvIpUy O} JaJaI C-] sJoqUINNY ‘puryysseur AWUNOD YoourpYY YyINog "] sNITy 


miles 
oO 1 2 
ee 
ie) 1 2 3 
kilometers 

4 

? 

yi 

ee | 

wa 

ALIS 

=f Ss 

ue git oe 


89°30’ 
x , 


NG, ¢ 


HERON BAY 


s ; y Aft ae. Yh, 
ae Ja 
mn - ’ Cedar Island 
2 : * ape 


89°27'30" 
| 


ei 


. \ ers 


MISS. > 


New AA 
Orleans~) 


St Joseph Point AREA OF STUDY 


(ZZ SAND RIDGES 
[|] MARSH (inland 
ps) 


also swam 


vO! 


105 


islands are reflected by the less varied Asmmonia-Elphidium-Nonion- 
Nonionella fauna in the muddy, sandy-muddy nearshore deposits of the 
period. Immediately south of Campbell and Point Clear Islands, sediments, 
presumably mostly from the Pearl River, built the bottoms up to intertidal 
levels and marsh vegetation was established, protected by the Mississippi 
River subdeltas. At its peak of development, this marshland extended con- 
siderably beyond the present southern and eastern marsh shores. 

At this stage, Indian refuse middens started to dot the banks of creeks 
in the newly established marshland. Shells of a very low salinity bivalve, 
the edible Rangia cuneata almost exclusively form the mounds. This is 
especially so, at the largest midden, “Cedar Lsland”. Some of the other 
middens also contained lesser quantities of oyster shells (Crassostrea vir- 
ginica) and the snail (Littorina irrorata). “Cedar Island”, with a surface 
area of 50 by 150 m (Fig. 1, Site 1), extends about 1.0 m above mean sea 
level and, according to borings, to about 3.5 m below sea level. Most of the 
mounds were formed during the Tchefuncte and later cultural periods, 
characterized by pottery production (Table 1). Each mound represents long 
periods of intermittent occupation and shell accumulation during seasons 
of food gathering activities. Subsidence, caused mostly by the compaction 
of the underlying muddy Holocene sediments, clearly affects the mounds. 
This is shown by occasional, partially drowned oak trees and red cedars in 
the marsh along mound peripheries. Due to subsidence, some middens may 
have been completely buried under surface deposits. 

Marsh accretion ended between 2,000-1,700 years ago. Termination of 
the active flow through the protective Mississippi-St. Bernard subdeltas started 
their disintegration. Renewed coastal erosion increased the adjoining western 
Mississippt Sound water area and the strength of erosive wave energy on 
the shores. This initiated the present steady reduction of the south Hancock 
marshland area. 


Table 1. Radiocarbon dates of Rangia cuneata from Indian shell mounds, 
Hancock County salt marshes 


No. of Date 
Site (Fig. 1) Dating laboratory Analysis (yrs. B.P.) 
1 (Cedar University of Georgia, UGa-353 2650+70 
Island ) Geochronology Lab. UGa-354 2900+70 
2 Krueger Enterprises GX2653 1295+90 
3 University of Georgia, UGa-371 2680475 
Geochronology La 
4 University of Georgia, UGa-373 2035+65 


Geochronology Lab. 
5 Krueger Enterprises GX25671 1190+110 


5m 


Ikm Imile 


Magnolia Bayou PointClear Cedar Three Oaks St. Joseph 
Ridge Caddy Island Island Bayou Point 
| 


Sm{ 


sea level 


Pleistocene 


land surface my 


au 


901 


107 


BOTANICAL ASPECTS 
COMPOSITION AND DISTRIBUTION. 


The vegetation of the Indian middens contrasts conspicuously with the 
low profile, homogeneous, and rather monotonous vegetation of the sur- 
rounding, extensive salt marshes. A list of plants found on the middens is 
shown in Table 2. The marshes are composed primarily of Juncus roemerta- 
nus, Spartina alterniflora, and Distichlis spicata (Eleuterius, 1972). The most 
abundant trees on the middens are the live oak (Quercus nigra) and the 
southern cedar (Juniperus silicola). The prevalence of cedar on the largest 
midden studied is apparently the basis for its name (Cedar Island; USGS 
Quadrangle, Grand Island Pass, Miss.-La., 1956, photorevised, 1970). Here 
16 oaks with diameters at breast height (DBH) ranging from 6-62 cm and 
about 50 cedars, the largest with a 45 cm DBH. Other common trees found 
were Celtis laevigata, Diospyros virginiana, Morus rubra and Zanthoxylum 
clava-herculis, 

e understory of site 1, “Cedar Island” is composed of a mixture of 
shrubs, primarily Ywcca doi folia and herbs. This midden, the most diverse 
floristically, contained a dense vegetational cover forming an almost 1m- 
penetrable mass. Other shrubs of frequent occurrence on the top of the mid- 
dens were: Ilex vomuitoria, Sabal palmetto, Sambucus canadensis, Hypericum 
hypericoides, Rhus copallina, Erythrina herbacea, Aesculus pavia, and 
Sageretia minutiflora. 

There is no obvious zonation of plants on the midden and the general 
impression is one of over-crowding. Numerous trees and shrubs lean outward 
from the mound, this “overhang” shading much adjacent marsh. Many of the 
shrubs are quite tall and often reach about 6 m. One large oak blown down 
by winds of Hurricane Camille in the east-central island area in 1969 left 
a large opening that subsequently was invaded by Yucca alozfolia. The south- 
eastern portion of Cedar Island presently is vegetated primarily by small 
trees, shrubs, and herbs. Stumps indicate that larger trees grew here in the 
past. The prevailing southeasterly winds apparently have an effect on the 
vegetation here, sculpturing and damaging the vegetation, as on the eastern 
ends of the offshore barrier islands (Eleuterius, 1975; Penfound and O'Neil, 
1934; Lloyd and Tracey, 1901; and Miller and Jones, 1967). The periphery 
of the midden contained shrubs that are common on natural levees along 
the bayous, upland edges of salt marshes and dredged soil. There were 
Baccharis halimifolia, Iva frutescens, Borrichia frutescens, and Myrica cerifera. 


Figure 2. Geological cross section through Holocene deposits in south ock 
County ypacshland Symbols: 1-clay, mud, sandy mud with Hiphidiem. neds 
fauna (h ini deposits); 2-clay, mu baer) clay ae mud with Ammonia- 
Elpbidium. Nonion fauna Cae alnty yi uddy Rae vay mud, mud with 
Ammotium Ga ae -salt marsh deposits; 
5—dune and barrier ridge es va ae ei eee (shell mound). 


108 


Table 2. 


Plant species found on Indian middens in Hancock County marshes. 
Indicator species are those found on shell middens and not in 


the 


marshes. They are marked with an asterisk. 


Scientific Names 
TREES 
Celtis laevigata Willd. 


s L, 
*Juniperus silicola (Small) Bailey 
Morus rubra L. 

rcus virginiana Miller 
bai olan clava-herculis L, 


SHRUBS AND WOODY VINES 
* Aesculus pavia L. 
*Ampelopsis arborea (L.) Koehne 
Baccharis halimifolia UL. 
Borrichta frutescens DC. 
*“Bumelia lanuginosa (Michaux) 
Campsis radicans (L.) Seemann 
] ! Des Moulins 
DC. 


(Pers. ) 


Pypevcun hvpevicoides 
Ilex vomitoria Aiton 
1 frutescens L 


(L.) Crantz 


li 
*Matelea caroliniensis (Chapman) Woodson 


Myrica cerifera . 

Rhus copallina 

Sabal palmetto a ex Schultes 

* Sageretia Lishee ih (Michaux ) 
Sambucus canadensi 

Serenoa repens (Bartram) Small 
Sane bona-nox I. 

Wusterta Daca (L.) Poiret 
* ie dloifolia L. 


Trel. 


HERBS 
Amaranthus spinosus L. 
Ambrosia artemistifolia U. 
Chaerophyllum tainturieri Hooker 
Cuscuta pentagona Engelm 


philadelphicus L 
Eryngium yuccifolium Michaux 
Eupatorinm — Michaux 
Euphorbia maculat 

Ipomoea sagittata Caw 

Lepidium virginicum UL. 

Manisuris rugosa (Nucall) Kuntze 


Common Names 


Hackberry 
Persimmon 

Honey Locust 
Southern Red Cedar 
Red Mulberry 


Live Oa 
Hercules-Club, Prickly Ash 


Red aoe 

Pepper-Vin 

Groundsel- Tree 

Sea Ox-eye 

False a Chittumwood 
Trum e 

Marine ivy, Weahe vine 
Coralbeads 

Coral Bean 

St. Andrew's Cross 


sim Pod Milkweed, Climbing 
r Vining Milkweed 

Wax Myrt ‘le 

Dwarf or Winged Sumac 

Cabbage Palmetto 


Ww a 
Spanish Bayonet, Yucca 


Thorny Amaranth 
Ragweed 


Thoroughwort 
Forb 


Morning Glory 
Poor-mans Pepper 
Joint Grass 


Opuntia compressa (Salisbury) Macbride 


Passiflora lutea UL. 

Phragmites communis as 
Physalis angu Bee 
Phytolacca rigida Sm 

Rhus radicans L 

Samolus parv Bees Raf. 


109 


Prickly Pear 

Prickly P: 

Wood Sorrel, Sheep Sorrel 
Bitter Panicum, Beach Grass 


Common Rec Roseo Cane 
Ground Che 

Poke, Pokebery, Pokeweed 
Poison I 

Water Pimpernel 


Scutellaria ovata Hill Skullcap 

Solidago sempervirens L. Goldenrod 

Hog Cane 

Clumped Cord Grass 
Venus Looking-Glass 


(44 
Stellaria media eee Chickweed 
Tillandsia usneoide Spanish Moss 
Verbesina ee (L. ) Walter Crown Beard 
Verbesina virginica L Coffee Bea 


Vicia ludoviciana Nute, Vetch, Deer Pea 


There were also many woody vines on the middens such as: Cissus mecisa, 
Wisteria frutescens, Ampelopsis arborea, Smilax bona-nox Cocculus carolinus, 
Campsis radicans, and Rhus radicans, 


ELEVATION-TIDE RELATIONS. All the other middens are of relatively low 
profile, and less than one meter in elevation. The diversity of the herbaceous 
flora is reduced (Fig. 3) with reduced elevation. Those middens only slightly 
above the surface of the marsh (0.1 m above MLW ) had fewer species than 
those on more elevated areas. No tree species were found below 0.5 m 
elevation (above MLW). Locally tidal amplitude is about 0.3-0.5 m, thus 
tidal inundation is probably responsible for the reduction of species. 


PLANT-SOIL RELATIONSHIPS. Although the soil on the middens is generally 
thin, a very black soil layer 4-8 cm thick is often found and is apparently 
nutritionally rich, judging from the dense vegetation supported. Roots cia 
ently rework soil and shell; shell and soil were found mixed at 8-16 c 
depth. Although high phosphorous content of soils is characteristic of up- 
land Indian habitation sites (Diety, 1957; Zeiner, 1946), available phos- 
phorous was generally low, probably due to the relatively high pH of the 
soil (Hole and Heizer, 1965 


RARE SPECIES. Sageretia minutiflora. Mohr (1901) indicated the distribu- 
tion of Sageretia minutiflora as extending from the Carolinas to Florida and 
west to Mississippi. Recent floristic surveys have failed to locate this species 
on the coastal plain of Mississippi and Alabama. Radford et al. (1968) 


110 


state that it occurs in only two coastal counties of South Carolina and in one 
in Georgia. Long and Lakela (1971) reported it from Lee County, Florida 
and Clark (1971) from Mobile Co., Alabama. Specimens were taken from 
a large shell midden on Dauphin Island. Michael Lelong (personal commu- 
Sag collected the species from clam shell middens on Little Dauphin 

sland, Alabama. Shell middens apparently provide the only habitat for this 
oe and this is the first report of its presence in Mississippi. 


CULTURED RELATIONSHIPS. Several of the plant species apparently had cul- 
tural relationships with prehistoric man since identifiable plant parts have 
been found in burial mounds (personal communication, Dale Greenwell). 
Seeds of plants of economic value may have been purposefully introduced 
at the semi-permanent occupation sites. Cedars may have been propagated 
because of the fine tinder produced from the bark and used in fire making. 
Coral bean seeds were probably a source of pigment for paint and used as 
“bead” ornaments. The fruits of the buckeye are historically known (at least 
locally, from pioneer days) to make a powerful fish poison. Manisurus rugosa 
consistently found on the more elevated sites is cited by Mangledorf (1974) 
as an ancestor of modern corn Zea mays. There is no known basis for extend- 


50 
e 
Ww 
= 
Oo 
ud 
w 25 
" @ 
o 
ac 
Wi 
va) 
= 
> 
i e 
th hh ht t+ + 
0.! 0.5 1.0 1.5 
ELEVATION (Meters above MLW) 
Figure 3. Species diversity in ce to ae elevation of Indian clam shell grat 
in hes marshes of Hancock Co., Miss. The number of plant species increased with 
increase in elevation. (MLW = mean on water 


111 


ing such speculation to the other species. Chemical work (pharmaceutical 
and taxonomic) on these species, might be highly rewarding. 


CALCIPHILES AS INDICATOR SPECIES. Brown (1938) stated that many species 
found on Indian mounds and middens that he examined in south Louisiana 
may be common on alluvial soil elsewhere in that state. He thought that 
the mound vegetation may have represented a relict. The investigated Han- 
cock marsh mound vegetation, however, can not be relict, since most of 
the plant species, especially those considered indicators, are calciphiles 
Their presence is apparently favored and determined by the large amount of 
calcium contained in the clam shells. These calciphiles, paradoxically, have 
not been found on recent deposits of oyster shells, left by European man 
during the past three centuries. 

Indicator species (Table 2) are those species that represent consistent 
and reliable indicators of the shell deposits on the studied middens. This 
does not preclude their occurrence elsewhere in other habitats. Several of the 
indicator species are more frequently found on Indian occupation sites than 
others. They are also found on occupational burial sites (mounds) on the 
adjacent terrestrial mainland and may also serve as indicator species for both 
marsh and terrestrial occupation sites (Eleuterius, unpublished data). These 
are Juniperus silicola, Aesculus pavia, Erythrina herbacea, and Morus rubra. 

The presence of shell midden within the salt marsh ecosystem provides 
unique habitats for an unusual flora which increases the diversity of our 
estuaries. The effects of these specialized, isolated plant communities on the 
surrounding marsh is of considerable interest for further studies. 


ACKNOWLEDGEMENTS 


We thank Dr. Sidney McDaniel for identifying many of the plant species 
and his helpful comments. Dolores Smith and Darlene Jennings provided 
technical assistance. 


REFERENCES 

BROWN, C. 936. The vegetation of the Indian Hane ares and marshes 
in pdaeoia a St. Bernard Parishes. In Lower Mississippi r Delta, Reports 
on the Geology of Plein. and St. Bernard See hes. ree Geol. Surv 
Bull. No. 8: i ear 

1938. The flora of ore deposits in the western Florida pees 

West | Feliciana Padi sh and e aton Rouge Parish, Louisiana. Jn Contribu 
to ae eee History - ihe Florida Parishes of Louisiana. Louisiana Geo fil 
Surv. . No. 12: 59- 

CLARK, a "OTL. Woody plants of Alabama. Ann. of Missouri Bot. Gard. Vol. 58: 
9-2 242. 


DIETY, E. F. 1957. Phosphorus accumulation in soil of an Indian habitation site. 
mer. Antiquity 22: 405-409. 

ELEUTERIUS, L. N. 1972. The marshes of Missis ssippi. Castanea 37: 153-168. 

te O75 he plant life of a co Hele een nland, associated waters and bar- 

ne island of Mis ssissippi with Saag to cones as a natural resource. 
ae ide Mar. Resources of anes ae pe ‘Grane pa a 84-87. 

GAG ANO, S. M. 1963. A s in aoe of south 

Tadee and south Micdccipei. Fronds edo See 104-13 


1B, 


HOLE, F. and R. F. HEIZER. 1965. An introduction to prehistoric archeology. Holt, 
tas Le oe N.Y. 497 
LOY . TR 


L nd § RACY. 1901. a insular flora of Mississippi and Louisi- 
ana. “Bull Toney Boe ‘Club. 28: 68-10 

LONG, R. and O. LAKELA. 1972 Manual of the flora of tropical Florida. Univ. of 

Miami Press. Coral Gables, Florida. 692 

MANGLEDORF, P. C. 1974. meee origin, evolution and improvement. Harvard 
Univ. Press, ace Mass. 

MILLER, G. J. a JONES, ai 1967. The vascular flora of Ship Island, Missis- 
sippl. ee a eioe. 

MOHR, C. 1901. hei in Alabama. Geol. Surv. Alabama. Vol. 6: 921 pp. 

OTVOS, E. G, Jr., 1973. Geology of the oe eye Alabama coastal area and 
nearshore zone. Guido New Orleans Geol. Soc. 67 pp. 

PENFOUND, W. s M. E. O'NEIL. 1934. a vegetation of Cat Island, Missis- 


sippi. Ecology 13 

RADFORD, A. E., H. EE AHLES, and C. R. BELL. 1968. oo of the vascular flora 
of the oo Univ. of North Carolina Press. 1183 p 

ZEINER, H. M. 1946. poe survey of the Angel Mock site, Evansville, Indiana. 
Amer. J. Bot. 33: 83-90. 


NOTES 


NOTES ON THREE SPECIES OF OPHIOGLOSSUM FROM NORTH 
CAROLINA—While making a study of over 4200 sheets of Ophioglossum 
from 114 different herbaria, the senior author became aware that the only 
Ophioglossum common to North Carolina was O. vulgatum VL. var. 
pycnostichum Fern. Specimens of this fern were examined from 32 counties. 
The only other species from the state was O. crotalophoroides Walter, and 
it was known only from a collection made on Dec. 9, 1956 on the lawn 
of Duke University Marine Station in Carteret County. Since other species 
of Ophioglossum are much more common than the number of sheets in 
herbaria indicate, it seemed probable that others could also occur in the 
Coastal Plains Province of North Carolina. A field trip was made in March, 
1976 to look for these ferns in North Carolina. 

On March 4, 1976, Thomas collected specimens of O. nudicaule var. 
tenerum from Brunswick County. The locality is: Bennett's Cemetery just 
west of Singletree Creek, 1.5 miles west of U.S. 17 near Hickman Cross- 
roads (Thomas 48453). It is common at this site with O. crotalophoroides 
in sandy soil. No other collections of this fern from North Carolina were 
made 

Later, the same afternoon, Thomas collected O. petiolatum new to North 
Carolina. The locality is: Lawn of Letties Grove Pentecostal Free-Will Bap- 
tist Church, south of U.S. 17 east of Shalotte, Brunswick County (Thomas 
48454). Thomas and Marx spent March 6-7, 1978 searching for Ophio- 
glossum in eastern North Carolina. We found O. petiolatum from Craven 
(Thomas and Marx 48476), Carteret (48489), Beauford (48502), Wash- 
ington (48509), and Dare (48524) counties. 

Although Ophioglossum crotalophoroides had been known only from one 
county, the authors found it to be abundant in sandy, grassy areas such as 
cemeteries and school lawns. We made collections from Brunswick (48452), 
Craven (48475), Carteret (48482), Beaufort (48506), Pamlico (48503), 
Martin (48506), Washington (48512), Hyde (48516), and Dare (48525) 
counties. 

We searched for O. crotalophoroides in southeasternmost Virginia with no 
success. We were hampered by a cold, rainy, and windy day that made 
crawling on the ground very miserable and seeing small plants almost a 
possibility. No doubt O. crotalophoroides and O. petiolatum will eventually 
be found in the Coastal Plain area of Virginia. R. Dale Thomas, Biology 
Department, Northeast Louisiana University, Monroe, LA 71209 and Paul 
S, Marx, Botany Department, University of North Carolina, Chapel Hill, NC, 
27514. 


SIDA 8(1); 113, 1979. 


114 


NEW STATE RECORDS FOR RANUNCULACEAE IN THE SOUTH- 
EASTERN UNITED STATES—During continued studies of the Ranun- 
culaceae for the forthcoming Vascalar Flora of the Southeastern United States, 
the following localities constitute, so far as I know, new state records for 
the species indicated below. 


1. RANUNCULUS FASCICULARIS Muhlenberg ex Bigelow. GEORGIA: Dade 
Co., moist shady deciduous forest, SW of Trenton, 12 Apr 1964, Walliams 
sa. (UNA!). 

. RANUNCULUS ‘TRILOBUS Desfontaines. TEXAS: Harrison Co., in a field 
off Interstate 20 at Waskom, 15 Apr 1974, MacRoberts 982 (LSUS! ) 

THALICTRUM DASYCARPUM Fischer & Avé-Lallemant. ALABAMA: Greene 

Co., common in rich low woods 1 mi E of Boligee, 9 May 1969, Thomas 

2059 (UNA!) 

. THALICTRUM DEBILE Buckley. MIssissIPPI: Lowndes Co., common in 
oak-hickory-cedar ridge in the Black Belt region, 9 mi E of State College, 
25 Mar 1968, McDaniel 10397 (GA!). 

I am indebted to the curators of the various herbaria for their loan of 
the specimens during this study—Carl S. Keener, 202 Buckhout Laboratory, 
The Pennsylvania State University, University Park, PA, 16802. 

CUPHEA CARTHAGENENSIS (JACQUIN) MacBRIDE—THE COR- 
RECT ORTHOGRAPHY .—With the publication of Radford, Ahles, & Bell’s 
Manual of the Vascular Flora of the Carolinas (1964) a typographical error 
was introduced in the treatment of Cuphea carthagenensis ('C. carthagensis’) 
which was repeated in Correll & Johnston’s Manual of the Vascular Plants 
of Texas (1970) and Correll & Correll’s Aquatic and Wetland Plants of 
Southeastern United States (1972). The error now appears more and more 
frequently on herbarium labels and in print (e.g. SID 

In hopes of preventing perpetuation of a misspelling I began this note 
only to face the question of what the correct spelling should be. The original 
spelling suggested the epithet should be carthagenensis, ee the original 
Lythrum carthagenense Jacquin (Enumeratio Pl. Carib. 22. 1760) based on 
plants collected at Cartagena, Columbia, $. A. As a ot place name, 
however, the epithet would be correctly formed by the stem carthagin- and 
the adjectival suffix -ensis denoting place of origin (Stearn, W., Botanical 
Latin, p. 218. 1966). To clarify the matter I consulted Dr. Dan Nicolson, 
expert in botanical orthography at the Smithsonian Institution. His clarifica- 
tion is one which many may find useful, particularly since today so few 
taxonomists are well-versed in the Latin necessary for correct formulation 
of new epithets. 


i) 


ee) 


PN 


‘The problem is whether a latinization of the Spanish Cartagena should or can 
yield a stem cartagen- (directly Spanish), carthagin- (directly from classical Latin 


SIDA 8(1): 114. 1979. 


115 


Carthago, genitive carthaginis) or an intermediate carthagen-. Jacquin’s stem 
carthagen-, represents a latinization of Cartagena as Carthagena, treating it as if i 
were a First Declension name (genitive carthagenae, stem carthagen-). This is 
clearly not classical Latin which would be Carthago oa carthaginis, stem 
carthagin-) of Third Declension. Purists would object to such a neolatinization as 
Carthagena but it strikes me as a perfectly reasonable option. As such I believe 
carthagenensis should be maintained under the paragraph of Art. 7 (ICBN ) 
which states that when changes made in orthography by earlier ee who adopt 
geographic names are intentional latinizations, they are to be pre d. 

“In another case, Panicum chartaginense Swartz (Nov. Gen. Sp. Bs 22. 1788), 
Swartz ae made yet another neolatinization of . rtagena which he cites as 
Chartagena. A classical scholar would substitute carthaginense, ae under the 
Code I believe this would not be acceptable and that the original spelling should 
stand. 

“Obviously it is hard to draw a firm distinction between orthographic (in- 
cluding pee aphic) errors and intentional latinizations. One can say that 
carthagenense and chartaginense are errors to justify correcting both to cartha- 

c 


in cases like this. However, the Preamble to the Code tells us that, ‘Other con- 
siderations, such as absolute grammatical correctness, regularity or cuphony of 
names, more or less prevailing custom, regard for oo etc., notwithstanding 
their undeniable importance, are relatively accessory.’ Under this concept I favor 
accepting original spelling, in most cases” (Pers. comm., 1978) 


The correct epithet is thus Cuphea carthagenensis, not C. carthaginensis 
or C. carthagensis. Users of the floras cited above should correct the name 
in those works to include the omitted syllable—Shirley A. Graham, c/o 
Alan Graham, Dept. of Biological Sciences, Kent State University, Kent, OH, 
44242. 


ZAMIA (CYCADACEAE) NEW FOR GEORGIA—This is the first re- 
port for the natural occurrence of Zamia in Georgia. It is reported by Small 
(1933) for Florida, occurring the most abundantly in the peninsular portion 
of the state. Lawrence (1951) states that Zama in the United States is 
“restricted almost exclusively to Florida, with an additional station reported 
in adjoining Alabama.” There appear to be no specimens to support the 
natural occurrence of Zamia in Alabama. Dr. Robert Kral of Wanderbilt 
University, who has been working intensively on the flora of Alabama for 
several years, says in a recent communication that to his knowledge it does 
not and has not occurred naturally in Alabama, but that ‘there is every likeli- 
hood that someone could have reported it as an escape—.” 

In northeast Florida Zamia is documented by herbarium specimens north 
into St. John’s County. The Georgia records are from Camden and Glynn 
Counties, with Duval and Nassau Counties, Fla., intervening. The classifica- 
tion and nomenclature of Zama species in Florida are in confusion. Until 
these problems are resolved it seems best to refer to the Georgia material as 


SIDA 8(1): 115. 1979, 


116 


Z. umbrosa Small which Small (1933) reports for “Hammocks, sand-dunes, 
and shell mounds, NE Fla 

The first collection presumed to be from Georgia was sent in for naming 
to the University of Georgia Harbarium in 1928 by Gertrude Proctor. Within 
a year or so after my arrival at the University of Georgia in 1938 I corre- 
sponded with Ms. Proctor in regard to the Zamia. The specimen had been 
given to her for naming from near Woodbine, Camden County. Ms. Proctor 
could not remember or find out who sent the specimen, but thought it was 
collected in a hammock. She and I searched considerably for the species 
in Camden County without success. There seemed to be a strong possibility 
that the specimen had been collected in Florida by someone living near 
Woodbine. Hence, I did not report Zamia for Georgia at that time. 

My beliefs are now altered by my collection of Zama from St. Simon's 

Island, Glynn County, where, according to Albert Fendig, Sr., a local resident, 
it occurs in a few scattered natural localities, but nowhere abundantly. M 
collection was made from one of three plants under a Quercus virginiana 
tree in a woods dominated by this and other evergreen species. Pinus was 
scattered, Vit7s common. 

In view of the above, Zamia should be considered native to Camden 
and Glynn Co., Georgia. The collections reported are: Camden Co.: {with 
male cone} Apr 1928, GA18054. Glynn Co.: Duncan 26359. Vigorous plant 
with about 20 leaves and “fruiting.” —Elev. ca. 25 ft. 17 Sep 1971, GA100484. 
—Wilbur H. Duncan, Department of Botany, University of Georgia, Athens, 
GA, 30602. 


REFERENCTS 
tae ee G. ee M. 1951. Taxonomy of vascular plants. MacMillan Co., N.Y. 
SMALL, J. K. 1933. Manual of the eee flora. University of North ee 

Press Chapel Hill 1554p. 

GALIUM SPECIES NEW TO THE SOUTHEASTERN UNITED 
STATES—While preparing a treatment of Galivm for the Flora of the 
Southeastern United States, | examined specimens which proved to be Galiam 
tricornutum Dandy and Galium palustre L. Both species are previously un- 
recorded for the southeastern United States. 

Galium tricornutum is a Eurasian species which occurs sporadically in 
the eastern United States and is reported from California (Munz, 1959) and 
western Oregon (Hitchcock, 1959). This species superficially resembles 
G. aparine L. 

Specific collection data for specimens from the southeastern United States 
are as follows: ARKANSAS: Miller Co.: Red River bottom, E of Texarkana, 
3 May 1951, Moore 510145 (UARK). GEORGIA: Oglethorpe Co.: banks of 
artificial pond, just W of Dry Fork Creek between Lexington and Washing- 
ton, 17 May 1952, Duncan 13541 (GA). SOUTH CAROLINA: Cherokee Co.: 


SIDA 8(1): 116. 1979. 


Li? 


san ditch, 2.4 mi N of jet. SC Rt. #150 and US Re. #29 on SC Rt. 

50 (N of Gaffney), 13 Apr 1957, Ables 22713 (NCU). Greenwood Co.: 
ae SC #34, 2.5 mi SW of Ninety-Six, 30 Mar 1957, Radford 20103 
(NCU). 

Galium palustre L. is also a Eurasian species which previously has been 
introduced in the United States in New England, New York, Pennsylvania, 
Michigan, and Wisconsin (Fernald, 1950) and which closely resembles 
G. tinctorium L. 

Specific collection data for the one known specimen from the southeastern 
United States is as follows: WEST VIRGINIA: McDowell Co.: wet ditch, 
Anawalt, 17 Jun 1969, Music s.n. (WVA). Presently it is unknown whether 
this species is established in the area of collection or merely a waif—Cheryl 
A. Lawson, 960 Oakridge, Duncan, Oklahoma, 73533. 


— 


REFERENCES 
a M. L. 1950. Gray's manual of botany. 8th edition. American Book Co., 
New York. 
HITCHCOCK Go Ls A: ee & M. OWNBEY. 1959. ae plants of 
the Pacific Northwest. Part 4. Univ rsity of Washington Pres eat 
MUNZ, P. A. 1959. A California ae University of California Dee Bete 


NOTES ON MITRACARPUS HIRTUS (M. VILLOSUS) AND SCIRPUS 
CALIFORNICUS IN MISSISSIPPI—Mitracar pus birtus (L.) DC. (Nichol- 
son, 1977) has been previously reported in the continental United States from 
southern Texas (Correll & Johnston, 1970, as Miétracarpum hirtum (1.) 
DC.), central Florida (Ward, 1976, as Métracarpus villosus (SW.) DC.), 
and most recently from southern Louisiana (Wunderlin, 1979, as Mitracarpus 
hirtus). In the course of identifying a Rubiaceous collection from R. Dale 
Thomas, Northeast Louisiana University, | discovered an SMU collection of 
Mitracarpus hirtus misidentified as Spermacoce glabra from southern Missis- 
sippi. This is apparently the only known record of it from that state. Collec- 
tion data: Mississippi. Pearl River Co.: cultivated field with Crotalaria 
spectabilis, 2 mi N of Henleyfield, 15 Nov 1964, Sargent 8542 (SMU). 

his is about 12.5 mi NW of Picayune on hwy 43, which is around 15 mi 
NE of the Louisiana collection reported by Wunderlin. Wunderlin’s report 
from Louisiana is based on a later collection from St. ae Parish: 
waste place RR, Abita Springs, 3 Oct 1970, Thieret 32568 

I suspect the two collections to have their origin from one introduction 
possibly originating in Mississippi based on collection dates and localities. 


Scirpus californicus (C.A. Mey.) Steud. is presently known in the United 
States from Arkansas and Texas west to California. It is believed that this 
is the first known collection to be reported from Mississippi. 


SIDA 8(1): 117. 1979. 


118 


Mississippi. Hancock Co.: low wet areas, Bay Saint Louis, 10 Aug 1952, 
Demaree 32719 (SMU). This extends the known range of S. californicus 
about 275 mi eastward.—Barney Lipscomb, Herbarium, Southern Methodist 


University, Dallas, TX, 75275 
REFERENCES 
arene D. S. & M. ree ica 0. 


xas. Texas Sa See Ren 
5. typification of taxa: Proposals on 


ac ‘OL ge D. H. ypification ore names vs 
48 and oid pettin of Mitracarpus birtus vs. M. villosus (Rubiaceae). 
" 


Manual of the vascular plants of 


Article 
Taxon 26: 569- 
SMALL, J. K. 1933. Flora of the southeastern flora. Publ. by author, New York. 
WARD, D. B. 1976. Maracas rpus ieee genus new to Florida od. eastern 
North America. ee a 78: was — 
WUNDERLIN, R. 79, Not Spermacore and Mitracarpus (Rubiaceae) in 
313-316. 


southeastern United Bae Phyoloeta 41(5) 


SIDA 8(1): 118. 1979. 


DOCUMENTED PLANT CHROMOSOME 
NUMBERS 1979:1° 
JAMES S. PRINGLE 
Royal Botanical Gardens, Box 399, Hamilton, Ontario, Canada LEN 3H8 


Figs. 1-12 are tracings from photomicrographs. All figures represent 
mitosis in root tips. Figs. 1, 3, 5, and 7-12 show chromosomes at metaphase; 
Figs. 2, 4, and 6 show chromosomes at anaphase (2 sets per figure ). 


ALISMATACEAE 
ALISMA TRIVIALE Pursh.” (Fig. 1.) 2n=28. CANADA, Reg. Mun. Hamilton- 
Wentworth: Pringle 1286 (HAM). 
SAGITTARIA CUNEATA Sheldon.” (Fig. 2.) 2n=22. U.S.A., Mich., Presque Isle 
Co.: Pringle 1225, 1273 (illus.) (HAM). 
SAGITTARIA LATIFOLIA Willd.” (Figs. 3,4.) 2n=22. CANADA, Ont., Reg. 
Mun. Hamilton-Wentworth: Pringle 1285 (HAM). 


ARACEAE 
ARISAEMA MACROSPATHUM Benth." (Figs. 5,6.) 2n=28. MEXICO, Mexico, 
Mun. Temascaltepec: Duncan et al. 2486 (MICH). 
LILIACEAE 
CLINTONIA ery ae Raf.’ (Fig. 7.) 2n—=28. U.S.A., Mich., Cheboygan 
Co.: Pringle 1224 (HAD 
CLINTONIA Sane a Raf.’ (Fig. 8.) 2n=32. U.S.A., N.H., Coés Co.: 
Pringle 1277 (HAM). (Documentation tor report in Utech, F., & L.B. Thien, 
Michigan Bort. 12:122—123. 1973.) 
RANUNCULACEAE 
CLEMATIS INTEGRIFOLIA L.” (Fig. 9.) 2n=16. Cult.: Pringle 345 (HAM). 


COMPOSITAE 
SOLIDAGO CANADENSIS L. var. HARGERI Fern.” (Fig. 10.) 2n=18. CANADA, 
= : in. Hamilton-Wentworth: Pringle 1372 (HAM). 
SOLIDAGO OHIOENSIS Riddell.” (Fig. 11.) 2n=18. Cult. ex U.S.A., Mich., Che- 
boygan Co.: Pringle 1289 (HAM). 
SOLIDAGO RIDDELLII Frank.” (Fig. 12.) 2n=18. Cult. ex CANADA, Ont., Essex 
Co.: Pringle 1250 (HAM). 


® Contribution No. 32 from the Royal Botanical Gardens, Hamilton, Ontario, Can- 
a 
"Taxon for which the same chromosome count has been published previously. 


© Taxon for which a different chromosome count has been published previously. 


“Taxon for which no chromosome count has been published previously. 


SIDA 8(1): 119. 1979. 


120 


AP 


L >» eo \ x Z ” 
ies a Ate PM yen 
» ee as ( n \ % 7 a NS = S 
= 7 
i Vy = =a? 
3 ™~ , 3 
A 
) ‘Dv ad 
a ' oo 
ARS SEP 
aN hy 


wy i ab 
Nowe / xX) U ~~ 


‘ ~\ I 
ri | ty“ Sex 


NOTICE 
VASCULAR PLANTS OF NORTH AMERICA NORTH OF MEXICO 


The Flora North America project (FNA) was recently revitalized by 
the Man and Biosphere Program (MAB) as a binational effort between the 
United States and Canada to produce a conventional flora of the vascular 
plants of North America north of Mexico using traditional methods. Initial 
funding for the proposed five-volume work is being provided by the Na- 
tional Park Service of the Department of the Interior. The /FNA 
Program Council plans to coordinate the research which will be necessary 
to produce a floristic publication of high scientific quality pertinent to na- 
tional needs. 

he Program Council has appointed an Editorial Subcommittee consisting 
of Dr. Reed C. Rollins, Chairman, Gray Herbarium of Harvard University; 
Dr. Howard S. Irwin of the New York Botanical Garden; and Dr. Roy L. 
Taylor of the University of British Columbia Botanical Garden. Dr. James 
L. Reveal of the University of Maryland has been appointed Editor. Their 
function will be to stimulate and coordinate the efforts of the botanical 
community in the writing of the flora. Toward this goal, the Editorial Sub- 
committee is currently working on a proposed format for the flora. Initial 
efforts will be toward the production of a volume treating the monocotyledon- 
ous plants, with a volume on the sympetalous dicotyledonous plants to follow 
next. The remaining three volumes will be worked on in the future. 

Long-term funding for the flora project is being explored by the Program 
Council, chaired by Dr. Peter H. Raven of the Missouri Botanical Garden. 
It is hoped that the flora project will be completed by 1990. 

Individuals wishing additional information or interested in contributing 
to the project, and in particular treatments of the monocots and sympetalous 
dicots should write to: 


Dr. James L. Reveal, Editor 


College Park, eure’ 3072 4? 


SIDA 8(1); 121, 1979. 


| 


SIDA Susie" 


VOLUME 8 NUMBER 2 NOVEMBER 1979 


CONTENTS 
Sidus Sidarum—III. Sida rzedowskii sp. nov., including a preliminary 
discussion of the Sida elliottii species group. Paul A. Fryxell. 123 


Taxonomy of the Verbesina virginica complex (Asteraceae ). 
John Olsen. 128 


The vascular flora of Savage Gulf, Tennessee. B. Exgene eee 
Thomas S. Patrick, Loy R. Phillippe, and David H. We 135 


A new species of Centrosema (Leguminosae) from nea and a 
key to the species in Central America. Paul R. Fantz 152 


Halogeton (Chenopodiaceae ) in North Se Will H. Blackwell, 


J. Douglas Haacke, and Christine O. Hopkin. 157 
Changes in hesnaes eo of the southeastern United States. 

Wilbur H. Dunca 170 
A new species of Linum from southern Texas and adjacent Mexico. 

C. M. Rogers 181 
Rediscovery of Spiranthes parksii Correll. P. M. Catling and 

K. L. Melntosh. 188 
Clematis pitcheri T. & G. var. dict = caudal: Dennis, comb. nov. 

(Ranunculaceae). W. Michael Denni: 194 
Field notes on Nigerian Striga (Scrophulariaceae). Lytton J. eee 

Daniel L. Nickrent, Richard A. Mansfield, and James E. A. Ogbor: 196 
NOTES. Hyptis mutabilis (Labiatae) in southeastern United States. 202—Trillinm 

pusillum (Liliaceae) a oni 209—Range extensions of Brazoria pulcherrima 

dell Corea Rubus trivialis Michx. var. duplaris (Shinners) Mahler, 


cea rae note on the distribution of Styrax americana 


fern and flowering plant flora. 213—Neviusia Wee: (Rosaceae) in 
Mississippi. 215—-The chromosomes of Collinsonia canadensis L. (Labiatae). 216— 
Nicolletia aia (Compositae) in Baja California. 216. 


US ISSN 0036-1488 


SIDA, CONTRIBUTIONS TO BOTANY 
Founded by Lloyd H. Shinners, 1962 


Editor & Publisher 
Wm. F. Mahler 
SMU Herbarium 

Dallas, Texas, 75275 


Associate Editor Assistant Editor 
John W. Thieret Barney L. Lipscomb 
Northern Kentucky Universit SMU Herbarium 
Highland Heights, Kentucky, 41076 Dallas, Texas, 75275 


Guidelines for contributors are available upon request beginning with 


volume 8. 


Subscription: $6.00 (U.S.) per year; numbers issued twice a year. 


© 
Sida, Contributions to Botany, Volume 8, Number 2, pages 123-216. 
Copyright 1979 
by Wm. F. Mahler 


SIDUS SIDARUM—III. 
SIDA RZEDOWSKII SP. NOV., INCLUDING 
A PRELIMINARY DISCUSSION OF THE 
SIDA ELLIOTTII SPECIES GROUP 


PAUL A. FRYXELL 


Research Geneticist, U.S. Department of Agriculture, Texas AGM University, 
College Station, TX 77843 


Eight North American species of Sida show strong affinities with one an- 
other and merit recognition as a distinct species group, as was first noted 
by Small (1898). The characters that unite them include (1) distinctive 
conformation of the fruits (generally oblate) and of the mericarps (see 
Figure 1), which are usually about 8 in number; (2) leaf shape that varies 
from broadly elliptic to narrowly linear (never rhombic or cordate-ovate); 
(3) a tendency in some species to rose-colored flowers (at least on drying), 
reaching an extreme in S. rzedowskii with purplish flowers; and (4) a 
tendency to extreme shortening of the apical internodes in several species, 
giving rise to congested apical inflorescences 

The abbreviated apical internodes are well expressed in the newly de- 
scribed species, Sida rzedowskii, and are suggestive of a similar condition in 
Sida sect. Malachroideae (Clement, 1957; Fryxell, 1975). In the latter group 
of species, typified by S. anomala St-Hilaire, the condition is described by 
Schumann (1891) as “Flores petiolis bractearum foliacearum  adnati, 
saepissime apice ramulorum subumbellato-congesti” and by Kearney (1954) 
as ““Peduncles adnate to the petiole of the subtending leaf or bract.” Careful 
examination indicates that the adnation of parts characteristic of sect. 
Malachroideae is simply an extension and intensification of the shortening 
of the apical internodes found in S. rzedowskii, S. neomexicana, and S. inflexa 
and occasionally expressed weakly in S. ell/ottii. Thus, we find in the present 
group of species an intermediate condition between the extreme characteristic 
of sect. Malachroideae and the less specialized condition found in the re- 
mainder of the genus. It is plausible to suggest that this tie indicates a 
phylogenetic link connecting sect. Malachroideae with the balance of the 
genus. It should be noted that sect. Malachroideae also has a tendency to 
rose-colored corollas and elliptic leaves, similar to the tendency in the S. 
elliottii group. 

Iam explicitly not suggesting, however, that the members of the S. elliottii 
group should be included in sect. Malachroideae. They are distinguished 
from the latter by their erect (not procumbent) habit and by their more 
numerous mericarps that lack rugulose or muriculate ornamentation on the 


SIDA 8(2): 123-127. 1979. 


124 


dorsal walls, as well as by other characters. The S. e/liottii group 1s not now 
given formal taxonomic status, pending further studies leading to a better 
understanding of infrageneric groupings in Sida. It is simply recognized for 
the present as a coherent group, without rank, and the following key is 
presented as an aid to distinguishing the species. 

Species are sometimes difficult to distinguish within this group. For 
example, some eee of characters occurs between S. elliottii and 
S. lindheimeri (e.g. in Arkansas). In this instance it may be preferable, 
following more detailed study, to merge the species and distinguish the taxa 
as S. elliottii var. elliottit and S. elliottii var. texana Torrey & Gray. Similarly 
in northern Mexico, it is sometimes difficult to distinguish S. ed/votti and 
S. nomexicana. These taxa are retained in specific rank in the key, following 
prevalent usage. A detailed analysis of these problems is merited. 


KEY TO THE SPECIES OF THE SIDA ELLIOTTIIT GROUP 


1. Pedicels up to 15 cm long, usually more than twice the length of the subtending 


eaves 

2. Leaves narrowly lanceolate, 7-10(—15) times as long as wide; calyx 6-7 mm 
long . S. longipes 

2. Leaves broadly elliptic, 1.2—2 times as long as wide; ee 9-11 mm long 


; otosina 
1. ae oe no more than 6 cm long, usually shorter than the caheending lea 
. Apical congestion of flowers and fruits conspicuous, as a result of oa ener: 
re the apical internodes 
. Corolla rose or purple (with yellow center); leaves elliptic, 2-6 times as 


long as broac . S. rzedowskti 
4. Corolla yellowish, sometimes — rose; leaves Tanceolate to linear, usually 
0 (—20) times as lon oad 
5. Plants freely branched one i base; leaves narrowly lanceolate to linear, 
1 broad S. neomexicana 


pow 


on plane sparingly branched; leaves broadly elliptic, 10-15 mm a 
. S. inflex 
. Apical congestion of flowers and fruits slight 
“e Calyx 7-10 mm long; pedicel often 2-5 cm long, shorter than to slightly 
exceeding ian: leaf . &. lindheimert 
6. Calyx 5-7 mm long; pedicel usually 1-3 cm feng, shorter than subtending leaf 
7. Leaves eee often 15-20 times as long as wide; mericarps 8-11 
. S. elliotti 
7. Leaves elliptic, usually 1.5—-3 times as long as wide: ‘mericarps 5- 
.. . S. turneroides 


SIDA ELLIOTT Torrey & Gray, Flor. N. Amer. 1:231. 1838 (based on S. 
gracilis Elliott, 1822, non Richard, 1792). (S. rubromarginata Nash, 1896; 
S. leptophylla Small, 1898). North Carolina south to Florida and west to 
southernmost Missouri and Arkansas; eastern Mexico from Veracruz and 
Querétaro north to Nuevo Leén and the Coastal Bend of Texas. 
ida rubromarginata and S. leptophylla have been maintained by Kearney 
(1954) and others as distinct from S. ellottii, but they seem to be rela- 
tively broad-leaved and relatively glabrous variants (respectively) of it. 


125 


Although Elliott originally described S. elliottii as ane examination 
of Elliott's type reveals that this is incorrect (Weatherby, 2 

SIDA LINDHEIMERI Engelmann & Gray, Boston J. Nat. Hist. 5: 213, 1845. 
(Sida texana (Torrey & Gray) Small, 1903). Central Texas and Louisiana 
to Sinaloa (1 specimen); cited from Guatemala by Standley & Steyermark 
anaes and from Chiapas by Standley (1923), but these may refer to 
S. ellio 

SIDA LONGIPES A. Gray, Pl. Wright. 1:19. 1852. Trans-Pecos Texas and 
Coahuila. 

SIDA NEOMEXICANA A. Gray, Proc. Amer. Acad. Arts ee ae Trans- 
Pecos Texas, New Mexico, Coahuila, Chihuahua, and Duran 

SIDA POTOSINA Brandegee, Univ. California Publ. Bot. 4:184. 1911 San Luis 
Potosi and Tamaulipas. 

SIDA INFLEXA Fernald, Rhodora 42:463. 1940. A restricted endemic in south- 
eastern Virginia. 

SIDA TURNEROIDES Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22:90. 
1940. Hidalgo to Tamaulipas. 

SIDA rzedowskii Fryxell, sp. nov. 


Planta herbacea perennis, minute stellato-pubescens, internodiis apicalibus abrupte 
abbreviatis. Laminae foliorum ellipticae, dentatae, utrinque stellato-pubescentes. Pedi- 

celli in axillis foliorum summorum, solitarii, plerumque in apicem congesti. Calyces 

aepe in nervis hirsuti. Petala glabra, gene Fructus oblati, in apicem minute 
pare nee mericarpia 8-11, laevigata vel leviter reticulata, 1-seminalia, in apicem 
acuta vel rotundata. 


Herbaceous perennial 1-4 dm tall, the stems branched and suberect, in- 
vested with minute stellate pubescence, the internodes abruptly shortened 
at the apices of the branches. Leaf lamina elliptic or oblong, 1-3 cm long, 

—)3-6 times as long as wide, subacute or obtuse, dentate, 3(—5)-nerved 
from the base, the lateral nerves inconspicuous, with dense or sparse stellate 
pubescence above and beneath. Petiole densely pubescent, 4-10 mm long. 
Stipules 4-7 mm long, linear-lanceolate, eearaes Maes: the petiole. 
Pedicels 1-10 mm long, solitary in the axils of the upper leaves, mostly 
aggregated apically (forming inflorescences eee to ae of Side ciliaris 

Bs owers, reduced leaves, and stipules crowded together because of 
the shortened internodes). Calyx 4-6 mm long, with pubescence like that 
of the leaves except sometimes hirsute on the nerves, 10-nerved, 5-lobed, the 
lobes ovate-acuminate. Petals glabrous (including the claw), 6-8 mm long, 
markedly asymmetrical, rose to red-purple but yellowish at the base. Staminal 
column pallid, 2-3 mm high, slightly pubescent or glabrous; filaments 1-1.5 
mm long, arising from the apex of the column; anthers yellow, few (ca. 20); 
pollen yellow. Styles 8-11; stigmas capitate. Fruits oblate, 5-6 mm diameter, 
3-4 mm high, apically with minute stellate pubescence; mericarps 8-11, 
3-4 mm high, smooth or slightly reticulate on the lateral wall, apically 
acute or rounded and dehiscent, l-seeded (Fig. 1,D). Seeds 2 mm long, 


126 


Figure 1. Comparison of mericarps of the four species of Sida. A, Sida lindheimeri 
(Byrd 42); B, Sida neomexicana (Board s.n.); C, Sida elliottii (Hill 2841) 
Sida rzedowskii (Hilerio 72). Scale = 5 mm. 


glabrous, blackish 

YPE: MEXICO: Hidalgo: Cerro Ventoso, entre Pachuaca y Real del Monte; 
orilla del camino, matorral de Hechtia podantha; alt. 2500 m; flores de color 
morado oscuro; 29. viii. 1965, Rzedowski 20560 (holotype: ENCB). 


Additional Ae examined : 

Hidalgo: Mpio. Pachuca, 4 km al NE de Pachuca, sobre la carretera a Real del 
Monte, alt. 2650 sas Sage ee (ENCB). Mpio Tepeapulco, Cerro Tres Penas, 

alt. 2500 m, Ventutn 245 (ENCB, pf). 

Edo. México: Mpio. Phen 3 alrededores de la Presa de la Concepcion, alt. 
2350 m, Hilerio 72 (ENCB, pf); ca. 2 km al NW de Tepotzotlan, alt. 2350 m, Huerta 
41 (ENCB); alrededores de la Hacienda Lanzarote, alt. 2350 m, Rzedowski 35021 
(ENCB, pf). Mpio. tae sed as te W de Cerro Sincoahe alt. 
Rzedowski 34307 (ENCB); al de Huehuetoca, Cerro Ahumada oe del 

Rancho Nuevo, alt. 2350 m, Roses ski a 03 ag Mpio. ae 3kma 
de Atizapan de Zaragoza, alt. 3400 m [2400 m?], Cruz 633 (ENCB); ele a 
Tigre, al NW de Atizapan, alt. 2500 m, Rzedowski 32004 (ENCB, a 

Distrito Nees Pedregal de San Angel, cerca de San Angel, Rzedowski 1019 
(ENCB); al sur de Cerro Zacatepec, Rzedowski 283 (ENCB). 

Jalisco: Mpio. Be de Moreno, Paso de La Troje, Cerro La Campana, SW of 
Ojuelos, alt. 2100-2300 m, McVaugh et al. 16832 (ENCB); ia ae SW “of Lagos 
de Moreno, Waterfall 15659 (SMU-in part). 

Chiapas: Cerro San Cristobal, San CriscSbal de las Casas, alt. 7000 ft, Breedlove 
DS 


San Luis Pe ytosi: Mpio. de Mezquitiq [Mexquitic], km 65 de la carretera San Luis 
Potosi-Zacatecas; alt. 2090 m; Garcia 665 (CHAPA, pf). 


The new species is named in honor of Dr. J. Rzedowski, author of 
Vegetacién de México and coauthor of Flora Fanerogdmica del Valle de 
México, whose extensive Mexican collections, especially in the Valley of 


ry 


Mexico, have brought this species to light. It is especially noteworthy that 
this species occurs rather commonly in the Valley of Mexico at elevations 
of 2300-2700 m, higher than any other North American species of Sida 
is known to occur. Moreover, it appears to be confined to these high eleva- 
tions, extending from Chiapas to northern Jalisco and southwestern San Luis 
Potosi. 


REFERENCES 


CLEMENT, I. D. 1957. Studies in Sida tira Contr. Gray Herb. 180: 1-91. 

FRYXELL, P. A. 1975. are sidarum [lI]. a 6: 1-6. 

ore T. H. 1954. ae key to North American species of Sida L. 

eafl. W. Bot. 7: eae 

SCHUMANN, K. 1891. oe In: K. F. P. von Martius, Flora Brasiliensis 12(3): 
279-34 

SMALL, : K. 1898. eercice in the botany of the southeastern United States.-XIV. 
Bull. Torrey Bot. Club 25 84. 

STANDLEY, P. C. 1923. Sida ae Trees and Shrubs of Mexico. Contr. U.S. Nat'l. 

Herb. oe Rene 

_ d J. Steyermark. 1949. Malvaceae. In: Flora of Guatemala. Fieldiana, 

Boe Date. 324-38 

WEATHERBY, C. A. 1942, A list of type specimens in Elliott’s herbarium. Rhodora 
44: 249-262 


TAXONOMY OF THE VERBESINA VIRGINICA 
COMPLEX (ASTERACEAE) 
JOHN OLSEN 
Department of Biology, Southwestern at Memphis, TN 38112 


The Verbesina virginica complex contains three taxa, V. virginica L. var. 
virginica, V. v. var. laciniata (Poir.) Gray and V. microptera DC., in the 
widespread tropical section Ochractinia. The complex has been a source of 
taxonomic confusion for some time. This confusion has arisen over the 
segregation of the southwestern United States taxon, V. microptera, from the 
more widespread V. virginica and the segregation of the southeastern popu- 
lations of V. virginica at either the specific or the varietal levels. 

Verbesina microptera DC. was described from a south Texas Berlandier 
collection (no. 1442) in 1836. Gray (1883) and Stanford (1976) con- 
sidered this name synonymous with V, virginica, while Correll and Johnston 

1970) treated the two taxa as distinct species, though suggesting they are 
conspecific. 

These two taxa are commonly distinguished by the number of wings on 
the stem and the extent to which these wings continue into the inflorescence. 
Examination of herbarium material quickly shows these characters are com- 
pletely unsatisfactory; the number of wings and their extent varies with the 
robustness of the plant and ranges from none to six in all taxa of this 
complex. 


Coleman (1977) has performed experimental crosses on populations of 
all three taxa in this group and recognized V. virginica and V. microptera 
as distinct species while maintaining the southeastern United States popula- 
tions as a variety of the former. He also points out the differing number 

ray florets that distinguish between V. microptera (10-13) and V. 
virginica (1-5). 

My examinations of herbarium and living material supports the conclu- 
sions of Coleman (1977). Furthermore, V. sicroptera is geographically 
isolated from V. virginica, the former largely restricted to the region south 
of San Antonio, Texas, and extending into Coahuila and Nuevo Leon, Mexico, 
and the latter occurring from west of Austin, Texas, north to Kansas and 
east to the Carolinas (Figure 1). Correlated with this geographic break 
(Figure 2) is a distinct morphological boundary as noted by Coleman. 
Verbesina virginica usually has three ray florets (always less than seven) with 
10-13 disc florets, while V. mcroptera usually has 11 ray florets (always more 
than nine) with ca. 25 disc florets. No exceptions to this meristic difference 
have been detected in the specimens examine 


SIDA 8(2): 128-134. 1979. 


inica 
laciniata 


oa 


rd 


eudidnigadinaeee nn 
—_— 
_————— ead 
ES | : 
: 
virginica 
var. virginica 


li 


Wt 


a7, 


il 
\ | 


| 


¢ 
) 


ov 


< 


micropte 


V. 


Fig. 1. Distribution of the taxa in the Verbesina virginica complex. 


SA : 
ipauuiinecsaanae nee 
btm: 
—" ‘ 
Vie VLE 
var. 
7 <a 
——ee ° 
ane 6A : 


130 


g. 2. Distribution of Verbesina virginica and V. microptera in Texas. V. virginica 
var. virginica (squares); V. microptera (stars). 


The Atlantic coast populations of V. virginica differ in several characters 
from the typical variety. The key differences are the lobing of the lower 
stem leaves, the indistinct stigmatic lines in the disc floret style branches, 
acute pale apices and glabrous ray achenes. Long and Lakela (1971) have 
recognized these populations as V, lacmiata (Poir.) Nutt. while others 
(Radford et al., 1968) have accepted them as a variety of V. virginica. I 
am inclined to agree with the latter disposition because of the close morpho- 
logical similarities between the two taxa. The most consistently reliable 
character distinguishing the two, leaf morphology, varies continuously from 
the typical. strongly dentate or serrate form to the sinuately lobed condition 
found in the coastal areas. Coleman (1977) also supports the latter view- 
point based on hybridization data. The pollen fertility in hybrids between 


131 


Pao aa, 
oo HOAc oOo HOAc 
15% Sy 4 
TBA TBA 
Cc 
O an) HOAc 
157 
TBA 


2-D chromatographic patterns of flavonoids in the Verbesina virginica com- 
plex. ‘- Vy. microptera; 2. V. virginica var. virginica, 3. V. virginica var. lacintata. 


V. v, var. laciniata and V. v. var. virginica is 93% (compared to eastern and 
western var. virgimica crosses which yielded 95% good pollen). This may 
be compared to the 73% fertility in crosses between V. microptera and 
V. virginica var. virginica and 41% between V. mcroptera and V. virginica 
var. lacimiata. Clearly, the coastal United States taxon is more compatible 
with the typical genome than is V. cro ptera. 

Further support for this disposition of taxa can be found in their flavonoid 
chemistry. V. virginica var. virginica and V. virginica var. laciniata produce 
identical patterns (Figure 3); they each produce three glycosides of 
kaempferol. V. mcroptera produces those same three compounds and two 
additional kaempferol glycosides as well. Chromatographically, the two va- 
rieties are obviously more closely related to each other than either is to 
V. nucroptera. 

My conclusion, based on these examinations, is to recognize two species, 
V. microptera and V. virginica, the latter being comprised of two varieties. 


142 


KEY 
_ florets fewer than seven; disc florets fewer than 15. . . 3.3...) OV. virginica 
em leaves unlobed . Cee ee ee Va virginica 
Stem leaves lobed or pinsately. divi de d toe ee ee var. laciniata 
Ray florets more than 9; disc florets more ‘haa 20° . oe ee.) OV microptera 


VERBESINA VIRGINICA L. Sp. Pl. 2:901. 1753. Type: USA, Virginia. Clayton 

166. (Holotype:BM ). 

Herbaceous perennials, 0.5—3.5 m tall. Stem erect, striate, unbranched until 
the inflorescence, with (O—) 1-6 decurrent wings up to 5 mm wide, vestiture 
puberulent below to tomentose above. Leaves lanceolate to broadly ovate in 
outline, laciniately lobed to entire, margins smooth to serrate, upper surface 
glabrous to scabrous, lower surface tomentose; petioles winged, wings usually 
decurrent and extending below the base onto the stem. Capitulescence 
corymbose, ultimate tomentose peduncles less than 1 cm long. Heads to 2 cm 
in diameter (including the rays); receptacle short-conical. Involucre 2-seriate, 
bracts oblanceolate, pubescent, keeled by a prominent midevein, outer series 
short, innter series grading into the pales, apex acute. Pales oblanceolate to 
weakly trifid, conduplicate, keeled by the prominent midvein, apex acute or 
acuminate. Ray ie 1-5 (7), pistillate and fertile, corollas white, tube 
pubescent, 1.6-2.1 mm long, ligule glabrous, 4.4-5.3 mm long, notched at 
the apex; style ae linear, exserted from the tube. Disc florets 9-13, 
corollas white, tube short, 0.3-0.6 mm long, pubescent; limb narrowly 
cylindric, glabrous, 2.3-2.9 mm long; style branches acute, stigmatic lines 
distinct or indistinct; anthers black, apical appendages acute. Ray achenes 
black, winged, minutely tuberculate to sparsely pubescent, laterally com- 
pressed, 4.1-4.9 mm long, 1.1-2.1 mm wide; wings 0.2-1.2 mm wide; pappus 
of 2 barbed aristae to 2.5 mm long. Disc achenes black, winged, glabrous, 
laterally compressed, 4.3-5.7 mm long, 0.9-2.0 mm wide; wings and pappus 
as in the ray achenes. Flowering Aug.-Dec. Chromosome number, 7 = 17. 
VERBESINA VIRGINICA L. var. VIRGINICA—Phaethusa virgimica (L.) Britton, 

Wy, Pied. 2, S487. 1915 
a polycephala DC. Prodr. 5:616. 1836. TYPE: USA: Texas, Bexar County 

ejar’” (San Antonio). 1828. Berlandier 1830. (Microfiche G-DC!, See 

US! a MO! Sketch GH!). 

Variously known as ice plant, white crown beard, frost plant or ice stickweed, 
V. virginica var. virginica is a widespread north American weed (Figure 1). 
It is characterized by unlobed, ovate-lanceolate to broadly ovate leaves. Leaf 
margins are highly variable, ranging from entire to serrate or dentate. More 
technical characters of distinction include the acuminate apex of the pales, 
the distinct stigmatic lines in the disc florets style branches and the sparse 
pubescence along the ridges at the apex of the ray achenes. 

VERBESINA VIRGINICA L, var. LACINATA (Poiret) Gray, Proc. Amer. Acad. 

Arts 19:11. 1883. —Siegesbeckia laciniata Poiret, Enc. Meth. 7:158. 1806. 

Described from a specimen sent by M. Bosc. —Verbesina laciniata (Potr.) 


195 


Nutt., Gen. 2:170. 1817. —Phaethusa laciniata (Poir.) Small, Fl. Miami 
195:200. 1913 


Verbesina sinuata Elliott, Sk. 2:411, 1823. ropes ere at CHARL. Material indi- 
cated as sent to Muhlenberg at PH is no 
Verbesina virginica L. var. insularis Rob. z — enm., Proc. Amer. Acad. 34:560. 
TYPE: een: do Noronha. (an island in the Cape Verde Archipelago) 
1887. Ridley, Lea & Ramage sn. (Holotype GH!). 


Although other authors regard this as a separate Species, the present in- 
vestigation, as pointed out above, cannot support this view. This variety 
commonly occurs on sandy soils along the coastal southeastern United States. 

The variey insularis from the Cape Verde Islands is obviously an intro- 
duction from the southeastern United States. 


VERBESINA MICROPTERA DC., Prodr. 5:616. 1836. TYPE: USA: Texas, ‘be- 
tween Laredo and Bejar”. 1828. Berlandier 1442 (Microfiche G-DC!, 
Isotypes GH! MO! NY! PH!. Phototype US! ) 


Verbesina texana Buckley, Proc. Phila.. Acad. 13:458. 1861. TYPE: MEXICO: Nuevo 
Leon, near Tantoyuca, province Huasteca, 1858. Ervenberg 58. (Lectotype [here 
designated } : PH! GH!). 

Verbesina microptera DC. var. mollissima Rob. & Greenm., Proc. Amer. Acad. 34:560. 
1899. TYPE: MEXICO: Nuevo Leon, Valley nee - Monterey. 7 July 1888. Pringle 

1916. (Holotype GH! Isotypes F! GH! NY (2 


— 


Herbaceous perennials to 2.5 m tall. Stems erect, striate, unbranched unti 
the inflorescence; winged; glabrous to puberulent below to tomentose above. 
Leaves ovate-lanceolate to broadly ovate, margins serrulate, serrate or lobed, 
apex acuminate or rounded; base narrowing quickly to a winged petiole, 
the wings continuing down the stem; lower leaves up to 25 cm long, 15 cm 
wide; pubescence of the upper surface scabrous, lower surface tomentose. 
Capitulescence a corymb, ultimate peduncles tomentose, less than 1 cm long. 
Heads to 2.5 cm in diameter (including the rays); receptacle nearly flat. 
Involucre 2-seriate, bracts oblanceolate, pubescent, keeled by the prominent 
midvein, apex acute; inner series intergrading with the pales. Pales oblanceo- 
late, conduplicate, keeled by the prominent midvein, 6.0-6.5 mm long, apex 
acute. Ray florets 11-15, pistillate and fertile; corollas white, tube pubescent, 

1.1—-1.6 mm long; limb glabrous, 2.7-3.3 mm long, apex with 1 or 2 notches; 
style branches linear. Disc florets ca. 25; corollas white, tube pubescent, 
0.7-1.5 mm long, corolla narrow cylindric, 2.1-2.9 mm long; style branches 
narrow acute, stigmatic lines distinct; anthers black, apical appendages acute. 
Ray achenes black, glabrous, laterally compressed, 4.44.8 mm long, 1.5-1.8 
mm wide; winged by 2 broad wings, 0.3-1.0 mm wide; pappus of 2 shia 
aristae to 2 mm long. Disc achenes similar, 3.9-4.5 mm long, 1.4-1.7 mm 
wide; wings to 1 mm wide; pappus of 2 barbed aristae to 2.5 mm ion 
Flowering May-Nov. Chromosome number, 7 = 17. 

As emphasized above, this taxon differs from V. virginica in the number 


134 


of ray and disc florets, as well as being geographically separated. Plants of 
this taxon are further distinguished from V. virginica by a more deltoid 
leaf shape, whereas the leaves of the latter narrow much more gradually 
along the petiole. Verbesina microptera also has much narrower wings on 
the achenes than either variety of V. virginica. 

Verbesina microptera is closely related to the Mexican taxon V. ramicifolia 
Rob. & Greenm., from which it differs primarily in its leaf pubescence. 


EXCLUDED TAXA 
V. virginica var. palmeri Gray = V. rumicifolia Rob. & Greenm. 
ACKNOWLEDGEMENTS 
I thank the curators of the following herbaria for the loan of material 
examined during this study: F, GH, LL, , PH, 
Drs. J. Hendrickson and B. L. Turner are acknowledged for reading and com- 
menting on the manuscript. 


REFERENCES 

COLEMAN, J. R. 1977. A eel of experimental hybridization in Verbesina 
ee ag Rhodora 17-3 

CORRELL, D. S. and M JOLINSTON 1970. Manual of the vascular plants of 
Texas. 2 Research ene Ren 

GRAY, A. 1883. Contributions to North ies botany. Characters of new Com- 
positae, with revisions of certain genera, and critical notes Proc. Amer. cad, 
Arts 19:11. 

LONG, R. W. and O. LAKELA. 1971. A flora of tropical Florida, Univ. Miami Press. 
Coral ee 

RADFORD, A. E,, H. E. AHLES and C. R. BELL. 1968. Manual of the vascular flora 
of the € carolina Univ. North Carolina Press, Chapel Hill. 

STANFORD, J. W. 1976. Keys to the vascular plants of the Texas Edwards Plateau 
and ane areas. Howard Payne Univ. Brownwood. 


THE VASCULAR FLORA OF 
SAVAGE GULF, TENNESSEE’ 


B. EUGENE WOFFORD, THOMAS S. PATRICK, LOY R. PHILLIPPE 
and DAVID H. WEBB 


Department of Botany, University of Tennessee 
Knoxville, Tennessee 379 


Remnants of pristine forests are manifestly important to both present 
and future generations of mankind. Such areas offer unique aesthetic experi- 
ences, demonstrate balanced ecosystems, support a rich fauna and flora, and 
most importantly, demand the acute awareness of persons interested in their 
preservation. According to eee Hemmerly and Turner (1972), a 
relic stand in Savage Gulf“... appears to be the only virgin tract of typical 
mixed mesophytic forest remaining in Tennessee. 

Savage Gulf is within the proposed Savage Gulf Natural Area which 
includes the gorges and adjacent uplands of Savage Creek, upper Collins 
River, and Big Creek. The Tennessee Department of Conservation now owns 
a significant portion of this 10,000 acre Preserve and negotiations for addi- 
tional land acquisitions are underway. Now that a means for protection 
seems assured, it is anticipated that critical habitats will be properly managed 
and preserved. 

Past studies of the floristics and vegetation of the gorges of the Cumber- 
land Plateau of Tennessee are those of Caplenor (1955, 1965), Clark (1966), 
Quarterman ef al. (1972), and Sherman (1958). Except for occasional her- 
barium vouchers deposited at the University of Tennessee, Knoxville, and 
Vanderbilt University, no extensive collections of vascular plants had been 
made from Savage Gulf. 

This report on the flora of Savage Gulf represents a concerted effort to 
obtain detailed floristic information about a remnant virgin forest and 
immediate surroundings. Data were obtained on the species of vascular plants 
present, the occurrence of unusual habitats, and the status of rare plants. 
Although no vegetative sampling or quantitative analyses of plant communi- 
ties were undertaken, collections were made throughout Savage Gulf and the 
adjacent uplands (Figure 1 ) 


GENERAL PHYSIOGRAPHY 


Savage Gulf lies on the Cumberland Plateau in eastern Grundy County 


"Contributions from the Botanical Laboratory, The University of Tennessee, Knox- 
ville, N.S. No. 503. 


SIDA 8(2): 135-151, 1979. 


136 


ONE MILE 


Figure 1. Savage Gulf, Grundy County, Tennessee. The gulf is outlined at the 
1800 ft contour which approximates the rim. Numbers 1-6 refer to collection areas 
(see text for vegetation and topographic descriptions ). 


137 


approximately 35 miles northwest of Chattanooga, Tennessee. The Cumber- 
land Plateau is the southern extension of the Appalachian Plateau Province 
(Fenneman, 1938). Since prominent vertical escarpments are characteristics 
of the region, it is referred to as the “Cliff Section” by Braun (1950). 

Savage Creek, the primary stream within the Gulf, flows almost due west 
and drops from ca. 1740 ft near its headwaters to ca. 970 ft at its confluence 
with the Collins River for an average gradient of 150 ft per mile. The scenic 

orge is extremely steep and hazardous. The rim is roughly 1800 ft in 
elevation on both the north and south sides. The caprock, composed of a 
coarse Pennsylvanian conglomerate, is often exposed as vertical cliffs or over- 
hangs ranging from 70 to 140 ft nae From the base of the escarpments 
the gorge is typically terraced and slope ranges from 45° near the base o 
the caprock to 15° at Savage Creek. Boulder fields of conglomerate, sand- 
stone, and limestone bedrock fragments are common on both the north and 
south facing slopes 

DESCRIPTION OF THE VEGETATION 

Savage Gulf is divided into six areas (Fig. 1) to facilitate annotation 
of the checklist and to describe the vegetation. For the most part, these areas 
are natural plant communities that can be defined by their unique indicator 
species, soils, slopes, and exposures. 

rea | is an up!and swamp that remains relatively wet during the sum- 
mer and supports a rich flora throughout the growing season. Similar habitats 
occur infrequently throughout the Cumberland Plateau in Tennessee, but 
most have a depauperate flora during the drier summer months. Dominant 
woody species include Nyssa sylvatica, Acer rubrum, Liquidambar styraciflua, 
Itea virginica, and Viburnum nudum. The herbaceous layer is characterized 
by Osmunda cinnamomed, O. regalis, Woodwardia areolata, W. virginica, 
Carex intumescens, C. joorti, Lilium michiganense, Zigadenus leimanthoides, 
Platanthera ciliaris, P. es P. cristata, P. integrilabia, lsotria eae. 
Xyris torta, Rhynchospora corniculata, Juncus repens, Parnassia asartfolia, 
Prosperpinaca pectinata, and Aster umbellatus. Taxa from this area that have 
affinities with the Coastal Plain, although not restricted to it, include 
Woodwardia virginica, Juncus repens, Rhynchospora corniculata, Itea 
virginica, and Proserpinaca pectinata. 

Area 2, the north rim, and area 3, the south rim, include the relatively 
flat to rolling uplands of the surrounding Cumberland Plateau. Both areas are 
disturbed to varying degrees by logging an support a similar flora. In general, 
the vegetation of the regenerating forest is dominated by various species 
of Quercus and Carya. Other frequent trees and shrubs are Pinus echinata, 
Oxydendrum ee caked floridus, Nyssa sylvatica, Viburnum 
werifolium, R ndron nudiflorum, and Vaccinium spp. Occasional 
small streams that are Bac to the rim of the gorge support dense 
understory vegetation of Kalmuia latifolia, Tsuga canadensis, Hex opaca, and 


138 


Euonymus americanus, Several noteworthy herbs occur chiefly in the upland 

woods, namely, Chamaelirium luteum, Cypripedium acaule, Malaxis unifolia, 

and Monotropsis odorata, A small swamp on the extreme western end o 

south rim is dominated by Woodwardia virginica. The distribution of this 

infrequent fern in Tennessee is discussed by Wofford, Webb, and Dennis 
C 


Area 4 represents the south facing slope of Savage Gulf. Extensive boulder 
fields with thick entanglements of Parthenocissus quinquefolia and Rhus 
radicans characterize much of the upper slope. e herbaceous flora is 
meager on these drier sites, but is relatively rich on more mesic sites near 
Savage Creek. A few of the more common spring wild flowers found on the 
lower slope are Orchis spectabilis, Cyripedium calceolus var. pubescens, 
ge arifolia, Geranium maculatum, Sedum ternatum, Viola conspersa, 

Phlox divaricata. Quarterman et al. (1972) sampled the overstory on 
a. facing slope and found high importance values for Quercus prinus 
(85.4), Carya tomentosa (56.7), Quercus rubra (56.6), Liriodendron 
nes (23.3), Quercus alba (23.0), Carya ovata (17.6), Carya ovalis 
(12.0), and Pinus virginiana (11.0). Furthermore, they consider this forest 
to be an oak-hickory-tuliptree expression of the mixed mesophytic forest type. 

Area 5, the north facing slope of Savage Gulf, is the most floristically 
diverse area, The spring and early summer flora contain many elements of 


are Dryopteris intermedia, Disporum maculatum, Trillium erectum, Poly- 
gonatum pubescens, Aristolochia macrophylla, Claytonia caroliniana, Mitella 
diphylla, Viola blanda, and Chelone lyonii. Dryopteris goldiana, Athyrium 
eee ne bipinnatifida, and an occasional element more typical 
of the Interior Low Plateau and Central Lowland Provinces, such as Trilliam 
recurvatum and Stylophorum diphyllum, are other features of Area 5. Two 
particularly northern elements, Streptopus amplexifolins and Maianthemum 
canadense, are reported by Quarterman e¢ al. (1972), but were not observed 
in the present study and specimens have not been located. 
co-dominants occupy the overstory of Area 5. Quarterman ef al. 
(1g72) A high importance values for Tilia heterophylla (61.7), Tsuga 
canadensis (54.4), Acer saccharum (40.0), Liriodendron tulipifera (37.6), 
Aesculus octandra (23.1), Carya tomentosa (20.8), Fagus grandifolia (17.1), 
Quercus rubra (12.6), and Magnolia acuminata (12.2). Cladrastis lutea, an 
uncommon tree of the eastern U.S., occurs sporadically in the understory. 
In addition, Quarterman ef al. (1972) consider the north facing slope to 
more closely resemble Braun’s (1950) typical mixed mesophytic forest of 
the Cumberland Mountains than adjacent mixed mesophytic stands in 
Tennessee. 
Area 6 is defined as the mouth of Savage Gulf and consists of many 
habitats from flood plain woods to open, drier limestone outcrops. Savage 
Creek runs underground in the mouth of the Gulf, except for periods of high 


159 


rainfall when torrents may increase the water’s height above ground by 10 
to 15 ft. The flood plain woods is composed of cutover pines (Pinus echinata 
and P. virginiana), several oaks (Quercus falcata, Q. alba, O. rubra), Platanus 
occidentalis, Carpinus caroliniana, Crataegus spp., and Liquidambar styraciflua. 
Characteristic shrubs along the bouldery channel are Xanthorhiza sim- 
plicissima Dirca palustris, Ilex ambigna var. montana, Cornus amomum, 
Rhododendron arborescens, and Viburnum rufidulum. Herbaceous taxa that 
occur in abundance include Asplenium platyneuron, Pachysandra procumbens, 
Phlox glaberrima, P. maculata, Collinsonia canadensis, Mitchella repens, and 
Marshallia trinervia. Other open habitats support a more weedy flora of many 
legumes and composites. 

A noteworthy limestone site occurs at an elevation of 1250 to 1300 ft in 
a northwesterly exposure toward the mouth of Savage Gulf. Calciphiles of 
this site are Agave virginica, Quercus muehlenbergii, Bumelia lycioides, 
Fraxinus quadrangulata, Swertia caroliniensis, and Echinacea pur pured, 


RARE TENNESSEE PLANTS” 


Several rare Tennessee plants, designated as such by the Committee for 
Tennessee Rare Plants (1978), occur in Savage Gulf. These are listed 
below with their degree of endangerment in Tennessee and location within 
the study area. 

1. Woodwardia virginica (L.) Smith—SPECIAL CONCERN. Dense colonies 
in swamp on extreme western edge of south rim, 14 May 1977, Webb & 
Wofford 812; upland swamp on north rim, 15 May 1977, Webb &Wofford 
819 


2. Poa languida Hitchcock—sPECIAL CONCERN. Mouth of Gulf, flood 
plain, 26 Apr 1977, Phillippe & Patrick 3769. 

3. Carex pedunculata Muhl.—SPECIAL CONCERN. Savage Gulf below still, 
under hemlock, 4 May 1977, Kral 59910. 

4. Veratrum woodii Robbins—ENDANGERED. Savage Gulf, nw of Sewanee, 
Jul 1972, M. Rhinehart s.n.; north facing slope near western end of Gulf, 
23 Jul 1977, Patrick, Phillippe, & Simmers 289, This taxon is genuinely rare 
in the Southeast (Johnson, 1969) and the Savage Gulf population of about 
200 individuals is the only known Tennessee locality. 
in 


upland swamp on north rim, 22 Jun 1977, Wofford & Webb 77-90 

6. Platanthera integrilabia (Correll) Luer—ENDANGERED. Locally abund- 
ant in upland swamp on north rim, 26 Jul 1977, Wofford & Webb 77-219; 
swale along old logging road on south rim, 23 Jul 1977, Patrick, Phillippe, 
Walson, & Simmers 300. 

2 Trichomanes petersti Gray—ENDANGERED. Dr. David K. Smith recently collected 
specimens of this rare fern from the Gulf. Exact data and discussion of this report 
will appear separately. 


140 


7. Cimicifuga rubifolia Kearney—YHREATENED. Near base of caprock on 
north facing slope, 23 Jul 1977, Patrick, Phillippe, & Simmers 288 
8. Hydrastis canadensis L.—YHREATENED. Deciduous woods near flood 
plain on south side of Savage Creek, in mouth, 14 May 1977, Patrick & 
Phillippe 83. 

9. Panax quinguefolium L—YHREATENED, Uncommon on north facing 
slope, 23 Jul 1977, Patrick, Phillippe, & Simmers 281. 

10. Montropsis odorata Schweinitz in EIL—THREATENED. Well-rotted 
wood along trail on north rim of Savage Gulf, ca. 0.5 mi W of Meadow 

creek, elev. 1820 ft, 2 Apr 1977, M. Whitten s.n.; pine woods po: foot 
trail on north rim near edge of gorge, 26 Apr 1977, Wofford, Webb, & 
Rader 77-2 

In addition, Carex hitchcockiana Dewey was collected from the base of 
the south rim on limestone near Savage Creek, 8 Jun 1977, Patrick, Phillippe, 
& Malter 201, Although this is the first collection on record in a Tennessee 
herbarium, Fernald (1950) and Underwood (1962) include this taxon for 
Tennessee. Another noteworthy record is the first verification of Chelone 
lyonii from the Cumberland Plateau. 


jon 


BOTANICAL-ECOLOGICAL SIGNIFICANCE 


Savage Gulf is one of the more floristically diverse natural areas in Ten- 
nessee. Considering the size of the area (ca. 10 sq. mi.), it contains an 
extremely high number of taxa (680) with elements from the Interior Low 
Plateau, the Appalachian Highlands, and the southeastern Coastal Plain. 
Several Tennessee rare plants occur here, including the only known locality 
for etnies woodii in the State. Furthermore, as noted by Quarterman 
et al, (1972), the forest that occurs on much of the north facing slope ts 
an ae virgin relic of mixed mesophytic forest. 

The area is presently under the management of the Tennessee Heritage 
Program of the Tennessee Department of Conservation, as part of the Savage 
Gulf Natural Area. It is our recommendation to those responsible for main- 
taining the antiquity and botanical uniqueness of the area that any develo 
ment be limited to primitive use. All concerned citizens must shoulder the 
responsibility to preserve yet another small piece of our natural heritage. 

ACKNOWLEDGEMENTS 

The development of this report was financed through a planning grant 
from the Bureau of Outdoor Recreation, Department of the Interior, under 
provisions of the Land and Water Conservation Fund Act of 1965 (Public 


Law 88-578) in cooperation with The Nature Conservancy and the Ten- 
nessee Department of Conservation. 


— 


FLORISTIC SUMMARY 


Collections were made throughout the study area from mid-April through 


141 


October, 1977. Voucher specimens are deposited in the University of Ten- 
nessee Herbarium, Knoxville (TENN) and duplicates will be distributed 
at a later date. 

A total of 680 taxa from 360 genera and 111 families is represented in 
the flora. For the most part, nomenclature is that of Radford, Ahles, & Bell 
(1968). Taxa are arranged alphabetically by family, genus, and species 
under three major headings, as follows: PTERIDOPHYTES (ferns and fern 
allies), GYMNOSPERMS, and ANGIOSPERMS. Number(s) after each entry refer 
to the general locality or plant community from which the voucher was 
collected. Slight records are not included. These areas are delimited in a pre- 


an 


ceding section, as follows: 
1—Upland Swamp on North Rim 
2—North Rim Woods 
3—South Rim Woods 


— 


PTERIDOPHYTES 
ADIANTACEAE 


Adiantum pedatu 

Chetlanthes ee Cviehis) D.C. Eaton 
—4 

Pellaca atropurpurea (L.) Link—6 


ASPLENIACEAE 
Asplenium montanum Willd.—5 
Asplenium platyneuron (L.) Oakes—3, 


5, 6 
Asplenium rhizophyllum L.—5 
‘ee asplenioides (Michx.) A. A, 


3 
rece geen (Sprengel) Tide- 
strom— 
Athyrium thelypterioides (Michx.) Des- 
Vaux— 
Cystopteris bulbifera (L.) Bernh.—%s 
Cystopteris protrusa (Weatherby) Blas- 
dell—4, 6 
Dryopteris goldiana (Hooker) Gray—5 
Dryopteris intermedia (Willd.) Gray— 
5 


Dryopteri _ A, 
5 

Polystichum acrostichoides — ( Michx. ) 
Schott—4 

Thelypteris hexagonoptera  (Michx. ) 
Weatherby—5 


Thelypteris noveboracensis (L.) Nicuw- 
and—3, 5 


4—South Facing Slope 
5—North Facing Slope 
6—Mouth of Gulf 


BLECHNACEAE 
Woodwardia areolata (L). Moore—1 
Woodwardia virginica (L.) Smith—1, 

3 


DENNSTAEDIACEAE 


Dennstaedtia punctilobula — ( Michx. ) 
Moore—3, 


Preridium aquilinum L.—? 


HYMENOPHYLLACEAE 


Trichomanes petersii Gray—4 
LYCOPDIACEAE 
Lycopodium eae Michx.—5 
Lycopodium obscurum L.—2, 
Lycopodium ate ons Lloyd & Un- 
derwood—4 


= 


Lycopodium tristachyum Pursh—2, 3 
OPHIOGLOSSACEAE 
Botrychium dissectum Sprengel—3, 5, 6 
Botrychium virginianum (L.) Swartz— 
4,5 
> 


OSMUNDACEAE 
Osmunda cinnamomea 
Osmunda regalis ie ) Gray—l1 

POLY PODIACEAE 
Polypodium polypodioides (L.) Watt— 


Polypodium virginianum L.—5 


142 


SCHIZAEACEAE 
Lygodium palmatum (Bernh.) Swartz 
—2 


SELAGINELLACEAE 


Selaginella apoda (L.) Spring—4 
VITTARIACEAE 
Vittaria sp. (gametophyte )—5 
II]. GYMNOSPERMS 
PINACEAE 
nus echinata Miller—3 
Pinus taeda L.—: 
Pinus virginiana Miller—2, 3, 4 
Tsuga canadensis (L.) Carr.—5 
CUPRESSACEAE 
Juniperus virginiana L.—3 


Ill. ANGIOSPERMS 
ACANTHACEAE 

Ruellia caroliniensis ( Walt.) Steud —6 

ACERACEAE 

Acer negundo 3 

Acer pense L—5 

Acer rubrum 

Acer nee naenil eek —S5 
ALISMATACEAE 


Alisma subcordatum Raf.—2 


Sagittaria engelmanniana J. G. Smith 
ssp. longirostra (Micheli) Bogin—3 
AMARYLLIDCEAE 

Agave virginica L.—6 

Hypoxis hirsuta (L.) Coville—3, 6 
ANACARDIACEAE 

Rhus copallina L—3 

Rhus glabra L.—6 

Rhus radicans L.—4 


ANNONACEAE 
Asimina triloba ( 


APIACEAE 


Angelica venenosa (Greenway) Fern.— 


Chaerophyllum tainturieri Hooker—6 
Cicuta maculata 


Cryptotaenia oe (L.) DC.—6 
Daucus carota 
Erigenia bulbosa (Mich. ) Nutt.—5 


Osmorhiza claytoniti (Michx.) Clarke 
—5, 6 
Oxypolis rigidior (L.) Raf—l1, 3 
Sanicula canadensis L—3, 5 

Sanicula gregaria Bicknell—5 

Sanicula smallii Bicknell—4 

Sanicula trifoliata Bicknell— 

Taenidia integerrima (L.) Drude—6 
Thaspium barbinode (Michx.) Nutt.— 


t, 
Zizia aptera (Gray) Fern.—6 


AQUIFOI IACEAE 


Ilex ambigua Scare Torr. var. mon- 
tana (T. & G.) _— 3, 6 
=i decidua Walt.— 
x opaca Ait.—3, 4 
ARACEAE 
Arisaema triphyllum (L.) Schott—4, 5 
ARALIACEAE 
Aralia racemosa L.—6 


Aralia spinosa L.—3, 6 
Panax quinquetolium L—S5 


ARISTOLOCHIACEAE 


Aristolochia macrophylla Lam.—5 
Aristolochia serpentaria L.—5 
Asarum canadense L.— 

rece arifolia (Michx.) Small—2, 


ASCLEPIADACEAE 
Asclepias amplexicaulis Smith—3 
Asclepias quadrifolia Jacq.—4, 5 
Asclepias tuberosa a 


ata L— 
Matelea_ carolinensis a Woodson 
—6 


ASTERACEAE 
Ambrosia son ae L—3 
Ambrosia trifida L.— 
Antennaria Sia nraeaitels (L.) Hook. 
—3 


Antennaria ska Rydb.—4 
Aster cordifolius L.—6 
Aster ere L—3, 5, 6 


Aster dumosus L.—2 
Aster Soe Michx.—3 

Aster laevis L—6 

Aster mere (L.) ae 6 
Aster linariifolius L.— 

Aster lowrieanus ae 

Aster patens Ait. 

Aster paternus Cronquist—2, 3 
Aster shortii Lind].— 


Bidens polylepis Blake—3 

Cacalia atriplicifolia L—6 

Chrysanthemum leucanthemum L.— 

Cirsium ae (L.) Sprengel—2 

Cirsium muticum Michx.—6 

Coreopsis eae L.—6 

Coreopsis major Walt. var. stellata 
utt. 


foe album = 


Heterotheca graminifolia (Michx.) Shin- 
ners—3 


rs 
Heterotheca mariana (L.) Shinners—2, 
3 


143 


Hieracium gronovii L.—2, 3 
Hieracium paniculatum L. 
Hieracium venosum L.—3, 


Krigia biflora (Walt.) Blake—3, 6 
6 


Lactuca canadensis L.— 

solr floridana (L.) Gaertn.—6 

Liatris aspera Michx.—3 

Liatris microcephala (Small) K. Schu- 
mann—2 

a spicata (L.) Willd —3 

Marshallia trinervia (Walt.) Trelease 
ex ae & Coville— 

Parthenium integrifolium L.—3 


L—3, 
Prenanthes serpentaria Pursh— 
Pyrrhopappus carolinianus (Walt.) DC. 


Rudbeckia fulgida Ait. var. umbrosa 
on & Beadle) Cs 
fidbeclia hirta L—3, 6 
Rudbeckia laciniata L.—3, 6 
Senecio aureus L.— 
Senecio obovatus Muhl. ex Willd. —4 
3 


Solidago arguta Ait—3 
Solidago boottit Hook. var. bootti 
Solidago boottii Hook. var. Se iglae 
) Cronquist— 
Solidago caesia L.—6 
canadensis L. var.  scabra 
(Muhl.) T. & G—3, 6 
Solidago curtisii T. & G—3 
Solidago erecta Pursh—2, 3, 6 
Solidago flexicaulis L. = 
Solidago gigantea Ait 
Solidago -raminifolia “ys a var. 


Solidago rugosa 
Cronquist—2, 

Solidago liaifolia Muhl.—3, 6 

Taraxacum officinale Wiggers 

Verbesina alternifolia (L.) Brit—6 

Vernonia altissima Nutt.— 


—6 
Mill. ssp. aspera ( Ait.) 
6 


BALSAMINACEAE 
Impatiens capensis Meerb.—6 
Impatiens pallida Nutt.—5 


144 


BERBERIDACEAE Viburnum nudum L.—2, 3 


Caulophyllum thalictroides (L.) Michx. Viburnum rufidulum Raf.—2, 6 
—_ - CARYOPHYLLACEAE 
Podophyllum pelratum L.—5 : . ; _. 
Cerastium holosteoides Fries var. vulgare 
BETULACEAE (Hartman) Hylander— 
Alnus serrulata (Ait.) Wéilld.—2 Silene rotundifolia Nutt—4, 5 
Betula lutea Michx. f.—4, 5 Silene stellata (L.) Ait. f—3 
Silene virginica L.— 


Betula nigra L.—2 : 
Carpinus caroliniana Walt.—6 Stellaria pubera Michx.—5 


Corylus americana Walt.—6 
CELASTRACEAE 
Ostrya virginiana ( Mill.) K. Koch—%5 


Euonymus americanus ma) 
BIGNONIACEAE Fuonymus aceopun puree ice, —06 
Antsostichus capreolata (L.) Bureau— 
ae ain: 3 sae CISTACEAE 


4,0 
Campsis radicans (L.) Seemann—6 
es tomentosa (Thunb.) Steud. Sheen 
—6 


Lechea racemulosa Michx.—3, 6 
> 


Tradescantia subaspera Ker—6 
BORAGINACEAE 


Cynoglossum virginian CONVOLULACEAE 


Lithospermum toa Rugel ex DC. Calystegia spithamaca (L.) R. Brown— 

—4 6 

Cuscuta campestris Yuncker—6 
BRASSICACEAE Cuscuta compacta Juss.—3 
Arabis canadensis L.—6 Ipomoea purpurea (L.) Roth—6 
Arabis leavigata’ (Muhl. ex Willd.) 

Poir.—4, 6 CORNACEAE 
Cardamine hirsuta L.—6 Cornus alternifolia L. f.—5 
Dentaria laciniata Muhl.—5 Cornus amomum Mill.—6 

Cornus florida L.—5 
BUXACEAE 


CRASSULACEAE 
4 


Pachysandra procumbens Michx.—6 
Pa eee Sedum ternatum Michx.— 
Calycanthus floridus L.—4 CYPERACEAE 
Carex aestivalis M. A. Curtis—3, 6 


CAMPANULACEAE é . 
Carex albolutescens Schwein.— 


Campanula americana L.—6 Carex albursina Sheldon—3 
Lobelia cardinalis L.—6 Carex amphibola Steud.—2 
Lobelia inflata L—2, 3 Carex austro-caroliniana Bailey—4 
Lobelia puberula Michx.—6 Carex bail ey! Britt. —" 
Specularia perfoliata (L.) DC.—3, 6 Carex cephalophora Muhl. ex Schkuhr 
—3 
CAPRIFOLIACEAE Carex communis Bailey—2, 4 

Lonicera japonica Thunb.—6 Carex complanata Torr. & Hook.—3 
Lonicera sempervirens L.—3 Carex crinita Lam,—3 
Sambucus canadensis L.—6 Carex debilis Michx. var. rudgei Bailey 
Sambucus pubens Michx.—5 —l, 2, ° 
Triosteum perfoliatum L.—3 Carex emmonsi1 Dewey—5 

Viburnum acerifolium L.—2 Carex flaccosperma Dewey—1 
Viburnum cassinoides L.—2 Carex gigantea Rudge—l 


Viburnum dentatum L-3 Carex hitchcockiana Dewey—6 


Carex incomperta Bicknell—3 

Carex intumescens Rudge—l, 2, 3 

Carex jamesii Schwein.—6 

Carex joorit Bailey—1l 

Carex laxiflora Lam.—3, 5 

Carex lurida Wahl.—3 

Carex pedunculata Muhl.— 

Carex pensylvanica Lam. var. distans 
Peck—4 


Carex plantaginea Lam 

Carex purpurifera Mac Aenrie= 2: 4 

Carex retroflexa nee ex ere ee 

Carex rosea Schkuhr— 

Carex swanii noe ) Macca: 3, 
6 

Carex virescens Muhl. ex Schkuhr—2, 3, 

6 

Cyperus praclongatus Steud.—6 

Cyperus retrofractus ae Torr.—3 

Cyperus strigosus 

Eleocharis obtusa (Willd. ) Schultes— 

Rhynchospora capitellata (Mich) Vahl 

535 


Rigethossoe corniculata (Lam.) Gray 
—l1 


Scirpus atrovirens Willd. var. georgianus 
(H Fern.—3, 6 

Scirpus cyperinus (L.) Kunth—1 

Scleria oligantha Michx.—6 

Scleria triglomerata Michx.—3 


DIOSCOREACEAE 

Dioscorea batatas Dcne.—6 
Dioscorea villosa L.—4, 6 
EBENACEAE 


Diospyros virginiana L.—3 


ERICACEAE 
SEs meee (L.) Pursh—6 
Epigaea repens L.—3, 4, 6 
Kalmia latifolia L—2, 3, 4 
yonia ligustrina (L —3 


Monotropa uniflora L.— 
Mo ene: odorata Schwein. in Ell— 


ae arboreum (L.) DC.—3 
Rhododendron alabamense Rehder—l, 
Rhododendron arborescens (Pursh ) 


orr.—6 
Rhododendron calendulaceum (Michx. ) 
orr.— 


145 


Rhododendron cumberlandense Braun 
Rhodedend {ron maximum L.—5 
Rhododendron nudiflorum (L.) Torr.— 
Vacaniuin arboreum Marsh.— 
Vaccinium constablaei ey 4 
Vaccinium corymbosum L.—3 
Vaccinium stamineum L.—3 
Vaccinium vacillans Torr.—3, 4 


“aS 


EUPHORBIACEAE 


Acalypha gracilens Gra 

Acalypha rhomboidea Rat. 6 

Croton monanthogynus Michx 

pe es corollata L. var. ainnii Aone 
nall ) hles—3 

ic mercurialina Michx.—4 


FABACEAE 
Albizia julibrissin Dur 
Amphicarpa bracteata (L.) Fern.—6 
pios americana Medicus—3 
Baptisia tinctoria (L.) R. Brown—2, 3 
Cassia marilandica L.—6 
Cassia nictitans L—6 


Cercis canadensis L.—6 
Cladrastis ae (Michx. f.) K. Koch— 
5 


Clitoria mariana L.—3 

Desmodium eee (Muhl. ex 

W illc Loudon— 

Desmodium Ae (Muhl. ex 

Willd.) Wood—6 

Desmodium marilandicum (L.) DC— 
) 


jou 


jae 


ille 


Desmodium nudiflorum (L.) DC—2, 6 
ee sae ta (L.) DC. var. 
niculatum— 
eres ices Soe DC. var. 
dillenit (Darl.) Isely—3, 
Desmodium obtusum (Muhl. ex Willd.) 


=O). 
Desmodium pauciflorum (Nutt.) DC. 
—6 
Desmodium rotundifolium DC.—6 
Galactia volubilis x Britt.—6 
Lespedeza bicolor Turcz.—3 

Lespedeza cuneata pen G. Don 


iecpedens hirta (L.) Hornemann—2 aes, 

Lespedeza hirta X inte 

Lespedeza intermedia (S. Wea aie 
—3,6 


146 


Lespedeza repens (L.) Bart.—3 
aed steuvel as t— 
Melilotus alba Desr.—3 
fe oralioies ee Cory—4 
Robinia rid )— 
Robinia i. acacia L—4 
carla ee (Soland. ex Sm.) 
ride— 

Se ena ice (L.) BSP.—3 
Tephrosia virginiana (L.) Pers.—2, 3 
Trifolium campestre Schreb.—6 
Trifolium pratense L.—6 
Vicia caroliniana Walt.—6 

FAGACEAE 
Castanea dentata (Marsh.) Borkh.—3, 4 
Fagus grandifolia Ehrhart—2 
Quercus alba L 
Quercus coccinea Meunchh. —3 
Quercus falcata Michx.—3 
Quercus marilandica Muenchh.—3 
Quercus muehlenbergii Engelm.—6 
Quercus prinus L.—3 
Quercus rubra L.—3 
Quercus stellata Wz 
Quercus velutina pee 3 


FUMARIACEAE 
Dicentra cucullaria (L.) Bernh.—5 
GENTIANACEAE 


Bartonia virginica (L.) a 
Gentiana quinquefoli ia L— 
1 i 


Swertia caroliniensis (Wa It.) Kuntze— 
6 


GERANIACEAE 
Geranium carolinianum L.—6 
Geranium maculatum L.—4 


HALORATACEAE 


Proserpinaca pectinata Lam.—1 


HAMAMELIDACEAE 


Hamamelis virginiana L.—2 


Liquidambar styraciflua L—4, 6 
HIPPOCASTANACEAE 

Aesculus octandra Marsh.—5, 6 
HYDROPHYLLACEAE 

Hydrophyllum canadense L.—5 


Phacelia bipinnatifida Michx.—5 


HY PERICACEAE 
Hypericum ae Walt. var. 
recognitum Fern. & —3 
Hypericum gentianoides “(L) BSP.—2, 
3 


Hypericum mutilum L—3 
Hypericum nudiflorum Michx. ex Willd. 
—6 


Hypericum prolificum L—6 
Hypericum punctatum Lam.—2, 6 
Hypericum stans (Michx.) P. cer & 
Robson—3 
Hypericum stragalum P. Adams & Rob- 
son— 6 
IRIDACEAE 
Iris cristata Ait—4, 5 
Sisyrinchium a tbidugn Raf.— 
Sisyrinchium angustifolium Mill—6 


JUGLANDACEAE 


Carya cordiformis (Wang.) K. Koch— 
Carya glabra ( Mill.) Sweet—6 


g. 

Carya ovata (Mill.) K. Koch—5 

Carya pallida (Ashe) Engler & Graebn. 
—3 


Carya tomentosa (Poir.) Nutt.—6 
Juglans cinerea L—6 
Juglans nigra L—6 
JUNCACEAE 
Juncus canadensis J. Gay ex La Harpe 
—3 
Juncus coriaceus Mackenzie—3 
Juncus debilis Gray—1, ° 


Juncus effusus L.—1, 3 
Juncus marginatus Rostk.—3 


Juncus repens ch 
Juncus tenuis Willd —2, 3 
la acuminata 


—6 
Luzula bulbosa (Wood) Rydb.—3 


LAMIACEAE 
eoninrona canadensis 
m virginianum L.—6 
i 6 


Mosla dianthera Caamition) Michx.—6 
Perilla frutescens (L.) Britt.— 
Prunella vulgaris L.— 


Pycnanthemum incanum (L.) Michx.— 
Pycnanthemum muticum (Michx.) Pers. 


Salvia lyrata L— 

Salvia urticifolia L—6 

Scutellaria elliptica Muhl.—3, 
Scutellaria pseudoserrata Epling—6 
Trichostema dichotomum 


LAURACEAE 
Lindera benzoin (L.) Blume—2, 5 
Sassafras albidum (Nutt.) Nees—4 


LILIACEAE 
Aletris farinosa L.—3 
Allium canadense 
Chamaelirum ee CL 
Disporum lanuginosum Miche) Nic- 
holson—5 
Disporum maculatum (Buckl.) Britt.— 


Erythronium americanum Ker—6 
Lilium michiganense Farw.—1 
edeola virginiana L.— 
Polygonatum biflorum (Walt.) 
Polygonatum pubescens (Willd.) Patch 
5 


Smilacina racemosa i Desf.—4 

Smilax bona-nox L.—6 

Smilax ecirrhata (Engelm.) Watson var. 
hugeri (Small) Ahles—4, 5, 6 

Smilax glauca Walt.—3, 4 


Trillium erectum L.—5 
Trillium grandiflorum (Michx.) Salisb. 
5 


Trillium recurvatum Beck—5 
Uvularia grandiflora Smith—5, 6 
Uvularia perfoliata 

Uvularia sessilifolia L.—2, 3 
Veratrum i Robbins—5 
Zigadenus eed. Gray—1 


LINACEAE 


Linum striatum Walt.—2, 3 
Linum virginianum L var. 
chon 


Plan 


floridanum 


LOGANIACEAE 
Spigelia marilandica L—6 


147 


LORANTHACEAE 
Phoradendron serotinum (Raf.) M. C. 
ohnston—5 


MAGNOLIACEAE 
Lirodendron tulipifera L.—4 
Magnolia acuminata L.—4 
Magnolia tripetala L—6 

MALVACEAE 


Hibiscus syriacus L—6 


MELASTOMATACEAE 
hexia mariana L.—3 
Rhexia virginica L—3 
MORACEAE 
Morus rubra L.—G 


NYSSACEAE 
Nyssa sylvatica Marsh.—4, 5 


OLEACEAE 
Chionanthus virginicus L.—3 


Se 
(Beadle) J. Wright e ex rein 
Fraxinus quadrangulata Michx. 6 


ONAGRACEAE 


Circaea lutetiana L. ssp. canadensis (L.) 
6 


nis L. 
Oenothera ona Roth—3 


ORCHIDACEAE 
Aplectrum hyemale (Muhl. ex Willd.) 
Orr. 
Corallorhiza odontorhiza (Willd.) Nutt. 
2G 
Cypripedium acaule Ait 
eae ca mee, To var. Seas 
Willd.) Correll—4 
Goose pubescens (Willd.) R. Brown 


eae spicata (Walt.) Barnhart— 
Isotria verticillata (Muhl. ex Willd.) 
Raf.—1 
Malaxis unifolia Michx.—2, 3 
—4 


Orchis spectabilis L 
Platanthera ciliaris cn Lindely—1, 3 


148 

Platanthera clavellata (Michx.) Luer— 

Platanthera cristata (Michx.) Lindley— 
1 


Platanthera integrilabia (Correll) Luer 
—1,3 

Tipularia discolor (Pursh) Nutt.—6 

Triphora trianthophora (Swartz) Rydb. 
5 


OROBANCHACEAE 
Conopholis americana (L.) Wallroth— 
4 
Epifagus virginiana (L.) Barton—%S, 6 
OXALIDACEAE 
Oxalis di ee Jacq.—6 
Oxalis stricta L—6 
Oxalis aes L.—6 
PAPAVERACEAE 


Sanguinaria canadensis L.—5 
ba diphyllum (Michx.) Nutt. 


PASSIFLORACEAE 
Passiflora lutea L.—6 
PHRYMACEAE 
Phryma leptostachya L—4 


PHYTOLACCACEAE 


Phytolacca americana L.—6 


PLANTAGINACEAE 
Plantago aristata Michx.—3 
Plantago lanceolata L.—3 
Plantago rugelit Dcne.—6 
Plantago virginica L.—6 
PLATANACEAE 
Platanus occidentalis L—4, 6 


POACEAE 
Agrostis ia (Walt.) Tuckerman 
25D. 


eee gerardit, Vitman—3 
Andropogon scoparius Mi 
Andropogon virginicus L.—6 
Aristida dichotoma Michx.—3 
oo gigantea (Walt.) Muhl.— 


ene erectum (Schreb.) Beauv. 
—6 


Bromus purgans L.—6 


Bromus secalinus L—2 
Calamogrostis cinnoides (Muhl.) Bart. 
1,3 
Chasmanthium © latifolium ( Michx. ) 
ates—6 
Chasmanthium laxum (L.) Yates—3 
Cinna arundinacea L.—6 
Dactylis glomerata L.—: 
anthonia compressa Austin—3 
anthonia serivea Nutt.—2 
Danthonia spicata (L.) Beauv. ex R. & 
—)D 
Decchamibets flexuosa (L.) Trin.—2, 3 
Diarrhena americana Beauvy.—6 
Digitaria ischaemum (Schreb.) Schreb. 
ex Muhl. var. itschaemum—3 
Digitaria ischaemum (Schreb.) Schreb. 
uhl. var. violascens (Link) Rad- 


elesere 


ord—- 
Echinochloa muricata (Beauv.) Fern.— 
3 


jon 


Eleusine indica (L.) Gaertner—3 
Elymus virginicus L.— 
Eragrostis spectabilis (Pursh ) Steuc 


jon 


el|— 


Erianthus alopecuroides (L.) Ell—6 

Festuca elatior L.—3 

Festuca obtusa Bichler—6 

Holcus lanatus L.—3 

Hystrix patula Moench—6 

Leersia virginica Willd.—3, 6 

Lolium multiflorum Lam.—3 

Melica mutica Walt.—2, 4 

Microstegium “icine (Trin.) A. 

-amus—3 

Muhlenbergia schreberi J. F. Gmelin— 

3 


Muhlenbergia sylvatica (Torr.) Torr. ex 
Gray—6 
Panicum an 


ceps Michx.—3 


nicum dichotomum L.— 
Panicum laxiflorum oon —3, a 


are longifolum Torr.—1 


nm 


Panicum longiligulatum Nash—1 
Panicum philadelphicum Bernh.—3 
Panicum polyanthes Schult.—2 
Panicum sphaerocarpon Ell.—3 
Paspalum laeve Michx.—3 

Poa compre 5 
Poa oe ‘Hitche —6 
Poa pratensis L.—6 


Setaria faberi W. Herrmann— 

Setaria geniculata (Lam. ) Deaig: —3 
Sorghastrum nutans 3 
et nitida (Bichler) Scribner 


Sporobs vaginiflorus (Torr.) Wood 


sce avenacea L.—2, 6 
Tridens flavus (L.) te —6 


POLEMONIACEAE 
Phlox amoena eS 
Phlox carolina L.— 
Phlox divaricata L_4. 5,6 
Phlox glaberrima L—6 
Phlox maculata L. ssp. pyramidalis 
(Smith) Wherry—2, 6 
Phlox pilosa L.—6 


POLYGALACEAE 
Polygala curtissii Gray 
Polygala incarnata L.—3 
Polygala senega L.—4 
Polygala verticillata L—2, 3 


POLYGONACEAE 

Polygonum  cespitosum Blume ~ var. 
longisetum (DeBruyn) Stewart—6 

Polygonum persicaria L.—6 
Polygonum punctatum Ell.—6 
Polygonum scandens L.—6 
Rumex acetosella L.—6 
Rumex crispus L.—6 
Tovara virginiana (L.) Rat—G 


PORTULACACEAE 
Claytonia caroliniana Michx.—5 


PRIMULACEAE 
Lysimachia lanceolata Walt—6 
Lysimachia quadrifolia L.—3, 6 
Lysimachia tonsa (Wood) Kunth—6 


RANUNCULACEAE 
yeas pachypoda EI].— 
Anemone quinquefolia L.—6 
Ane irginiana L.—6 


pe canadensis L.—4 


Hepatica acutiloba 
Hydrastis canadensis L—6 
Ranunculus abortivus L—6 


149 


Ranunculus recurvatus Poir.—5 
Thalictrum eine DC.—4, ‘ 
bescens Purs iG 
valictrum Fane (L.) Boivin— 
, 6 
Xanthorhiza simplicissima Marsh.—2, 
5 
RHAMNACEAE 
Ceanothus americanus L.—6 
Rhamnus caroliniana Walt—6 
ROSACEAE 
Agrimonia parviflora Ait—6 
Agrimonia pubescens Wallroth—6 
Agrimonia rostellata Wallroth—6 
Amelanchier canadensis (L.) Medicus— 


3, 6 
Aruncus dioica (Walt.) Fern.—6 
Crataegus crus-galli L—6 


Crataegus flabell lata Raa K. Koch—3 
Crataegus flava Ait. 
C rataegus viridis L.— 


i = 
Gillenia stipulata (Muhl.) Baillon—o6 
oli (L.) Moench—3 
Physocarpus opulifolius (L.) Maxim.— 
6 


Potenulla canadensis L.—2 


Pyrus angustifolia Ait.—2, 3, i 

Pyrus arbutifolia (L.) L. f— 

Pyrus meclanocarpa (Michx.) ee 
) 

Rosa carolina L.—95 

Rosa multiflora Thunb.—6 

Rubus argutus Link 5 

Rubus betulifolius Small 

Rubus hispidus L.—2, 3 

Rubus occidentalis 

Rubus phoenicolasius | Maxim.—6 

Spiraca japonica L.— 


RUBIACEAE 
Cephalanthus occidentalis L—3 
Diodia teres Walt.— 
Diodia ee L—6 
Galium aparin 
Galium circaezans ie —3, 6 
Galium latifolium Michx.—5 
Galium pilosum Ait.—3, 6 


150 


Galium triflorum Michx.—3, 6 THYMELAEACEAE 
Houstonia caerulea L—2, 3 Dirca palustris L—6 
oustonia purpurea L.—6 
Mitchella repens L.—S, 6 TILIACEAE 
Tilia heterophylla Vent.—5 
SALICACEAE 
Salix humilis Marsh.—3 TYPHACEAE 
Salix nigra Marsh.—3, 6 Typha latifolia L—3 
SANTALACEAE ULMACEAE 
Pyrularia pubera Michx.—6 Celtis occidentalis L.—6 
Ulmus alata Michx.—6 
SAPOTACEAE Ulmus americana L—5 
Bumelia lycioides (L.) Pers.—6 Ulmus rubra Muhl.—4 
URTICACEAE 
SAXIFRAGACEAE 


Boehmeria cylindrica (L.) Swartz—2, 6 
Laportea canadensis (L.) Weddell—5, 6 

Heuchera americana L.—5, Pilea pumila (L.) Gray—6 

Heuchera praviflora Baltes 5 ~~ 

Heuchera villosa Michx.—5 VERBENACEAE 

Hydrangea arborescens L. ssp. discolor 
(Seringe) McClintock—5 

Itea virginica L.—1, 2 

Mitella diphylla L.—s VIOLACEAE 

Parnassia asarifolia Vent.—1, 3 

Penthorum sedoides L.—6 


Astilbe biternata (Vent.) Britt.—5 


Verbena simplex Lehmann—3 
Verbena urticifolia L— 


Hybanthus concolor (Forster) Sprengel 
5 


Tiarella cordifolia L.—5 Viola affinis LeConte—4, 5, 6 
Viola blanda Willd.—5 

SCROPHULARIACEAE Viola canadensis L.—5 
Agalinis tenuifolia (Wahl) Raf.—2, 3 Viola conspersa Reichenb.—4, 5, 6 
Aureolaria laevigata (Raf.) Raf.—6 Viola hastata Michx. 
Aureolaria virginica (L.) Pennell—3 Viola hirsutula Brainerd 
Chelone glabra L.—3 Viola pedata L—2, 3 
Chelone lyonti ae Viola pensylvanica Michx.—5 
Gratiola virginiana L.—3 Viola primulifolia L—2, 3 
Lindernia dubia (L.) Pennell—6 Viola rostrata Pursh.— 
Mimulus alatus Ait—6 Viola triloba Schwein.—5 
Pedicularis canadensis L—4 
Penestemon canescens (Britt.) Britt. ssp. VITACEAE 

brevisepalus (Pennell) Crosswhite—6 Parthenocissus quinquefolia (L.) Planch. 
Verbascum blattaria L—6 —4, 
Verbascum thapsus L.—3 Vitis aestivalis Michx.—3, 4, 6 
Vitis cinerea Engelm. ex es 
SIMAROUBACEAE Vitis rotundifolia Michx.—4 
Ailanthus altissima (Miull.) Swingle—5 XYRIDACHAE 
SOLANACEAE Xyris torta Smith—1 


Physalis heterophylla Nees—6 
Solanum carolinese L.— 
THEACEAE 
Stewartia ovata (Cav.) Weatherby—4, 
6 


151 


REFERENCES 


BRAUN, E. L. 1950. Deciduous of Eastern North America. The Blakiston Co., 
Philadelphia, Pennsylvania. 59 

CAPLENOR, D. 1955. An snorted list of the vascular plants of the gorges of the 

Fall Creek Falls State Park. Jou enn. Acad. Sci. 39: 93-108. 

fest ell 1965. The owe 5 ne gorges of the Fall Creek Falls State Park in 
ee Jour. Tenn. Acad. Sci. 40: 27-39. 

CLARK, R. C. 1966. The vascular loa of the Fiery Gizzard gorges in south central 
Tennessee. eae M.A. Thesis, The University of North Carolina, Chapel 
Hill. 89 p 

COMMITTEE FOR TENNESSEE RARE SS 1978. The rare vascular plants of 
Tennessee. Jour. Tenn. Acad. Sci. 53: 128-133. 

FENNEMAN, N. M. 1938. Physiography of Eastern United States. McGraw-Hill Book 
Co., New York. 691 

FERNALD, M. L. 1950. rages Manual of Botany. Eighth Edition. American Book 


JOHNSON, R. G. 1969. A taxonomic and floristic study of the Liliaceae and allied 
families in the southeastern a States. Unpublished Fh.D. Diss., West Virginia 
University, ia are 337 p 

QUARTER MAN, E., B. TURNER and T. E. HEMMERLY. 1972. Analysis of 
virgin mixed aa Honests in Savage Gulf, Tennessee. Bull. Torr. Bot. Club 
99: 228-232. 

RADFORD, A. E., H. E. AHLES and C. R. BELL. 1968. Manual of the Vascular Flora 
of the Carolinas. Univ. of North Carolina Press, Chapel Hill. 1183 p 

SHERMAN, H. L. 1958. The vegetation and floristics of five gorges of the Cumberland 
Plateau. Unpublished M.S. Thesis, The University of Tennessee, Knoxville. 103 PP. 

UNDERWOOD, J. K. 1962. Cyperaceae of Tennessee. Unpublished Ms. deposited in 
the University of Tennessee Herbarium, Knoxville. 80 

WOFFORD, B. E., D. H. WEBB and W. M. DENNIS. 1977. State records and 
other recent noteworthy collections of Tennessee plants II. Castanea 42: 190-193. 


A NEW SPECIES OF CENTROSEMA 
(LEGUMINOSAE) FROM NICARAGUA AND A 
KEY TO THE SPECIES IN CENTRAL AMERICA’ 


PAUL R. FANTZ 
Fairchild Tropical Garden, Miami, Florida 33156 
Present address: Dept. of Horticultural Science, Box 5216, 
North Carolina State University, Ralergh, NC 27650 


Seymour (1978a) reports four new species of Centrosema from Nicaragua, 
viz. C. angustifolium (H.B.K.) Benth. C. plwmiert (Turp.) Benth., C. 
pubescens Benth, and C. virginianum (L.) Benth. However, a specimen of 
this genus deposited at the herbarium in Paris (P) which was examined in 
the course of a monographic study of the genus Clitoria (Fantz, 1977), does 
not accord with these or any other previously described species reported 
from Central America as listed in revisions and New World floras (e.g. 
Bentham, 1837, 1839; Standley, 1928; Standley and Record, 1936; Standley 
and Steyermark, 1946). The new species has larger flowers and bracteoles 
similar to C. plamieri, but is easily distinguished by the leaflet shape, longer 
calyx lobes, and the failure of the plant to turn black when dried. 

Nicaragua, the largest country in Central America, has but one published 
flora (Ramirez Goyena, 1909-1911), which is reported to be of little value 
(Blake and Atwood, 1942). A retired minister and botanist, Frank C. Sey- 
mour, at the age of seventy-three, began a personal cataloguing of the Flora 
of Nicaragua in 1968. He made seven expeditions to Nicaragua to study 
the flora firsthand and to collect representative specimens, plus purchasing 
a number of Nicaraguan collections made by other scientists doing research 
in that country. This project was financed from Seymour's own personal 
funds and through the sale of duplicate herbarium specimens. Seymour 

8b) summarized his several publications on his expeditions and on the 
flora of Nicaragua. Now an octogenerian, Seymour's research is culminating 
in a manuscript which provides a checklist of the flora of Nicaragua. This 
new species of Centrosema is named in honor of Frank C. Seymour in recog- 
nition of his contribution to the botanical knowledge of Nicaragua. 


ou 


— 


CENTROSEMA seymourianum Fantz, sp. nov. 
Folia trifoliata, glabrata; foliola elliptico-oblonga, breviacuminata, infra sparsim 


Pverouic studies accomplished at the IFAS Herbarium, Dept. of Botany, Uni- 
versity of pen Gainesville, Florida. Florida Agricultural ee Station Journal 


Series No. 


SIDA 8(2): 152-156. 1979. 


1399 


puberula trichomatibus appressis secus nervos majories. Pedunculi axillares, 
floribus apicem pedunculorum congestis. Flores grandes, resupinati, 5.5-6 cm ae 
vexillum extus sericeo-villosum. Bracteolae ae ae calycem subaequantes, 10-20 
mm longae, 4-5 mm latae. Calyx late campanule atus, tubo 4—5 mm longa; dentes supert et 
laterales tubo vix longiores, 5-7 mm longi; dens eae elongatus, 13-15 mm longus; 
calycis pubescentia inconspicua, in dente ventrali densior, trichomatibus uncinatis. 
Legumen incognitum. 


Scandent herb; stems terete, faintly angular, longitudinally striated, hollow, 
2-3 mm thick, weakly twining, pubescence scattered, pilose and uncinate 
(observed latter at 30X). Leaves 3-foliolate, petiolate, glabrate; leaflets pin- 
nate, broadly elliptic-oblong, rapidly short-acuminate, apex mucronate, base 
rotund, primary nerves of 10-12 pairs, upper surface green, glabrous, lower 
surface concolorous, very sparsely pubescent with appressed trichomes along 
major nerves, lamina 12-13.5 cm long, 6-8 cm wide. Petioles subquangular- 
terete, densely pilose on pulvinus, sparsely so above, 5 cm long; rachis similar, 
ca 2.5 cm; petiolules quadrangular, pilose, 6-8 mm. Stipules ovate, striate, 
persistent, 4-6 mm long, 0.6-0.8 mm wide; stipels to 6 mm long. Inflo- 
rescence axillary, shorter than the petioles, several-flowered crowded at the 
apex of the peduncle; peduncles one or two per node, 2-2.5 cm. long, 
moderately densely pubescent, trichomes short, subappressed; pedicels paired, 
8-9 mm. Bracts deciduous, ovate, striate, 6-7 mm long, 2-3 mm _ wide. 
Bracteoles enclosing calyx and flower in bud, large, coriaceous, striated, ovate- 
oblong, ciliolate, 10-12 mm long, 4-5 mm wide. Flowers large, papilionace- 
ous, resupinate, 5.5-6 cm long. Calyx broadly campanulate, upper teeth 
widely separated from lateral and lower teeth, sparsely pubescent on tube 
but more densely on teeth, particularly the retell tooth, trichomes uncinate, 
inconspicuous (observe at 30X), to 0.5 mm long; calyx tube 4-5 mm long, 
4 mm wide at base expanding rapidly to 11 mm wide at throat; calyx teeth 
longer than the tube, upper teeth 5 mm long, 2 mm wide, lateral teeth 
7 mm long, 1.7 mm wide, ventral tooth elongated, complicate, 13-15 
long, 2.5 mm wide at the base. Vexillum broad, ca. 4.5-4.8 cm wide, compli- 
cate, densely sericeous-villosus on the outer surface, bearing a basal pouch 
ca 6 mm in diam. above the 4 mm long claw, and with a broad 1.8 mm 
sinus above the pouch and between it and a 3 mm long spur. Alae, carina, 
androecium, and gynoecium hidden within the complicate vexillum glued to 
the herbarium sheet, thus not observed. Legwme unknown 

TYPE COLLECTION: NICARAGUA. Bois, environs de Grenade, 40 m, Janvier 1870, 
Levy 419 (HOLOTYPE: P—plant “A” mounted on left side of ene: Fig 

The type specimen is mounted on a herbarium sheet along with another 
specimen labeled as “Herb. Mus. Paris, 1870, no. 20”, which is mounted on 
the right side and labeled "B” by this author. It is to be excluded as part 
of the type specimen of Centrosema seymourianum as this specimen belongs 
to the genus Phaseolus. The type specimen includes only the specimen 
mounted on the left-hand side of the herbarium sheet, and which has been 


Mayyy - WEARS, NES. Pads 


Kanter wcsrag unanes 


ede FPS 
: EX 7e MW 20 
re 1. ce of Sa Giese seymonrianum (Levy 419, P—plant “A” mounted 
on ae left side of the herbarium sheet; excluding plant “B”’, Levy [?] 20, ee 
on the right side of i. ca sheet. ) 


155 
labeled “A”. 


A key is Bene below to the species of Centrosema reported to occur in 
Central America 


1. Leaflets 1; petiole winged (Mexico to Colombia) 
Te sae 3; petiole not winged 
. Flowers small, 2—4 cm Tone: leaflets commonly small, 3—7.5 cm long; bracteoles 
—10 mm long. 
3. Upper calyx teeth much shorter than the tube; leaflets with main lateral 
nerves divergent at right angles from the midrib (Honduras to Brazil) 
Me eh che ta A he Me Msi cic ta a P a alln ins aCsa fce ene caa bese eee let eee 
3, Upper calyx teeth subequal to longer than the calyx tube; eee with ma 
lateral nerves ascending from the midrib. 
Lea ets anaes oblong or linear; legume short, weakly falcate (Costa Rica 
Cc 


a ee ete C. sagittatum 


to Brazil) oo cccccceeeee . paschorum 


pies ovate to oblong; legume long, straight. 

a Calyx teeth very unequal, upper and lateral teeth short, ca 2-3 mm long 
ay subequal to the length of the tube, lowermost tooth much longer 
than the tube length, 5-8 mm long; legume 10-20 cm long, 5-7 mm 
wide; leaflets ovate, pubescent on both surfaces at least in the juvenile 
state, veinlets somewhat inconspicuous (Mexico to Brazil, W. Indies) 


Be ct te laid s adh sida etttin ntidlntttns allen at sf a ee ae Tee . pubescens 
y: Calyx teeth acai all much longer than the length of the abe. teeth 
ca 6-9 mm long; legume 6-12 ong, 4—5 mm wide; ee ovate to 
oblong or linear- ae ae veinlets eonspiciously % sed, promi- 


nent eee United States to Argentina, W. Indies, topical Africa) 

ere hs C. virginianum 

2. Flowers large, 4-6 cm long; leaflets commonly large, (5) 7-14 cm long; 
bracteoles 10-20 mm long. 

6. Leaflets broadly ovate to rhombic-ovate, often drying black; flowers 4—5 cm 

long; bracteoles exceeding calyx by two to three times calyx length, ca. 15-20 

mm long; calyx tube 5-7 mm with obsolete to short teeth to 2 mm long; 

legume 9-10 mm wide (Mexico to Brazil, W. I Indies, a, Africa ) 

Aon plumiert 

6. Leaflets oblong to elliptic- oblong ¢ or ovate-lanceolate, not deine Giaee flowers 

5S-G cm long; bracteoles subequaling length of cal tube and lowermost 

tooth, 10-13 mm long; calyx tube 4-5 mm long with conspicuous teeth sub- 
equal to longer than the tube Icngth; legume 7-8 mm wide. 

7. Leaves ovate-lanceolate, subpilose below with trichomes spreading on the 
nerves; calyx sericeo-pilose, trichomes 0.5-1 mm long, conspicuous when 
viewed at 10X; vexillum gibbous (Mexico) .. C. galeottii 

. Leaves elliptic-oblong, glabrate with very sparsely appressed trichomes on 
the nerves; calyx Soeee with scattered uncinate trichomes (to 0.4 mm 
long), sparse on the tube and more densely compacted on hens al lobes, 
conspicuous when viewed at 30X; vexillum spurred (Nicar 


~~ 


ie seymourianum 


Acknowledgement is made to Helen Correll for the Latin description 


> CENTROSEMA galeotii Fantz, ‘ig nov, Bastonym: Clitoria grandiflora Mart. & 
Gal., Bull. Acad. Brux. 10: 189. se ee Galeotti 3284 (BR!). Non 
Geaerens ee Benth. of Brazi 


156 


and to Daniel B. Ward, Dana Griffin III, and Donovan Correll for reviewing 
the manuscript and providing constructive criticisms. 


REFERENCES 


BENTHUM, o 1837. ae entationes de Leguminosarum Generibus. Ann. Wein. 
Mus. Natur. 117-120. 
ie De (ae Generibus Commentations. Ann. Wein. Mus. 
Natur. 2: 117-120. 


BLAKE, S. F. and A. C. ATWOOD. 
aoe: 1: ae ce States Government Printing Office, Washington 
FANTZ, P. R. 1977. A monograph of the genus Clitorta Cesuminose: 

1066 pp. aaa Thesis, University of Florida. 
RAMIREZ GOYENA, M. 1909-1911. Flora Nicaraguense. 2 vol. 1064 pp. Managua. 
SEYMOUR, F. C. 1978a. A checklist of the vascular plants of Nicaragua. (Unpub- 
lished manuscript and personal communication ). 
978b. we in Nicaragua. Phytologia 39(6): 500. 


STANDLEY, C. 28. Flora of the Panama Canal Zone. Lae U.S. Nat. Herb. 
27: 215- 


1942. Geographical guide to the floras of the 


Glycineae ). 


ae Flora of Costa Rica. Pub. Field. Mus. Nat. Hist. Bot. Ser. 18(2): 
and S. J. REC — Bie ce = and flora of British Honduras. Pub. 
Field. Mus. Nat. Hist. Ser. 12 
and J. 


Se ee a Ce of Guatemala. Fieldiana Bot. 24(5): 
178-181. 


HALOGETON (CHENOPODIACEAE) 
IN NORTH AMERICA’ 


WILL H. BLACKWELL, J. DOUGLAS HAACKE, 
AND CHRISTINE O. HOPKINS 


Department of Botany, Miami University, Oxford, Ohio 45056 


The history, nomenclature, and geography of Halogeton glomeratus 
(Stephan ex Bieberstein) C. A. eyer in Ledebour in North America have 
been investigated with attention given to both literature and herbarium 
specimens. Halogeton received intensive study in the United States between 

1942 and 1961 because of its role in the mass mortality of sheep (oxalate 
poisoning) on western rangelands. Although it is apparently no less com- 
mon today in the Intermountain Region, sheep mortality has been reduced 
through educational efforts, and interest in this plant has declined somewhat 
in recent years. Confusion has existed in the literature, but a herbarium 
study indicates only one species of halogeton present in the United States 

now or historically and its identity is confirmed as Halogeton glomeratus. 
Halogeton was introduced into the United States (probably from  south- 
central Russia) earlier than previously reported (ie. before 1930), live 
animals (Karakul) being the probable source of introduction. A number 
of the unnoticed halogeton infestations in the United States were likely 
initiated by migrating sheep. This species usually does not self-spread over 
long distances a should only gradually, at most, extend its present range. 

t will probably not become as widespread, nor represent as great a threat 

to western sheep flocks, as once was thought. 


HISTORICAL PERSPECTIVE 


Halogeton, an annual Eurasian chenopodiaceous genus, might well have 
remained an obscure weed but for the alarm it brought when large numbers 
of sheep were poisoned in the western United States. Although present in 
the United States at an earlier date, substantial awareness of halogeton did 
not come until 1942 when 160 head of sheep were fatally poisoned near 
Wells, Nevada (Erickson et al., 1952; Flemming et al., 1944). Stomach 
contents of these dead animals were composed almost entirely of this plant 


"This work was supported in part by a grant (W. H. Blackwell, Principal Investi- 
gator) from the American Philosophical Society (No. 7991—Penrose Fund) and 
from the Willard Sherman Turrell Herbarium Fund of Miami University. We seed 
fully acknowledge curators of the herbaria listed for loaning eee and/or pro- 
viding information. We also wish to acknowledge Dr. Charles Perino for his assistance 
in gathering some of the specimen and literature information alluded to in this paper. 


SIDA 8(2); 157-169, 1979. 


158 


material. Subsequent feeding tests performed at the University of Nevada 
confirmed the strong circumstantial evidence of the poisonous nature of 
this plant (Flemming et al., 1944). The mass poisoning near Wells led to 
first attempts (in 1943) to control the weed, a mixture of furnace oil and 
detergent being employed (Erickson et al., 1952). This and subsequent con- 
trol measures were largely unsuccessfu 

Whereas halogeton was known (at least by a few people) to be present 
in the United States before 1942, it was not thought to be harmful (Kings- 
bury, 1964). In fact, it was even considered by some (Erickson et al., 1952) 
as fair forage for livestock. The casual regard in which halogeton poisoning 
was initially held in this country was doubtless due to the fact that there 
were apparently no reports of its toxicity in the Old World (Dayton, 1951), 
and thus no special reason to suspect it. However, suspicious small losses 
of livestock in the western United States had actually preceded the Wells 
incident (Kingsbury, 1964). 

After the major poisoning at Wells, numerous minor losses of both sheep 
and cattle were soon reported in which halogeton was implicated (Erickson 
et al., 1952). However, a wider recognition of this taxon as a poisonous 
range plant was to wait until 1945 when 1620 head of sheep perished in a 
single day near Bridge, in south central Idaho (Morton et al., 1959a). Other 
severe losses in this Raft River area (Cassia County) included 750 head of 
sheep belonging to a Mr. John Ward of Almo, Idaho (ZoBell and Silcock, 
1950). In fact, the problem became so serious in the mid 1940's that a 
dozen major sheep ranchers in the Raft River Valley were forced out of 
business (Stoddart et al., 1951b), finding it necessary in some cases to 
abandon their ranches. These and other events brought the noxious weed 
to the attention of the public by 1950, and generated an expansion of re- 
search by personnel of the U. S. Department of Agriculture. Popularized 
reports often took the form of alarming accounts of the spread of this 
“invader,” sometimes with not so subtle innuendoes as to the country of 
native origin. The following are typical of the frightening titles or inflama- 
tory headlines of the early 50's: “Sheep- Killing Weed,” Life Magazine, Jan. 
15, 1951; “Poison Rides the Range, ’ Reader's Digest (condensed from the 
Denver Post), December 1953; “Another Russian Invades US.,” the Na- 
tional Wool Grower, 1950; and “Soviet Weed Kills Sheep,” Oakland Tribune, 
June 28, 1950. The Life Magazine article contained the statement that more 
than one-third of the flocks of sheep in the United States were in danger. 
It was understandably at this time that several agricultural colleges, univer- 
sities, and range extension agencies in the western states took an active 
interest in halogeton, initiating various studies into its ecology and possible 
control. Professional interest was stimulated when the United States Congress 
appropriated over four-and-a-half million dollars for study in legislation 
(Halogeton Act, Velie, 1953). Co-operative projects and committees were 
established in Idaho (Erickson et al., 1952) and Utah (Stoddard et al., 


— 


159 


1951b) to investigate its biology and possible eradication. Warnings of the 
danger of halogeton appeared in agricultural extension service bulletins of 
several western states in the early 50's, including Arizona (Armer, 1952) 
where its occurrence has yet to be documented. Besides the usual admonitions 
of its effect on sheep, of possible habitats in which it might occur, that it 
is not controlled selectively by 2,4-D or similar herbicide sprays, and recom- 
mendations for trailing sheep so as to avoid it, these bulletins usually de- 
scribed the plant. Particular emphasis was given to means of distinguishing 
halogeton from the less dangerous Russian thistle (Salsola kali L.), another 
introduced chenopodiaceous plant with which it was sometimes confused. 
For example, Durrell (1951) stated: “[halogeton] resembles Russian Thistle, 
which is common throughout the country. However, it has a softer appear- 
ance, its leaves are sausage-shaped, succulent and fleshy. The bristle at the 
tip is softer than the spine on Russian Thistle leaves.” 

The toxicity and precise effects of halogeton on animals received addi- 
tional attention during the late 1940's and early 1950's. Flemming et al. 
(1947), who had earlier (1943) reported halogeton as poisonous (contain- 
ing oxalic acid), indicated the toxic substances to be “salts of oxalic acid 
which are elaborated and stored during the growth of the plant to maturity,” 
present as two oxalate salts, water soluble and insoluble. Vawter (1950) 
subsequently described the mechanism, symptomology, and pathology of 
halogeton poisoning. Ingestion of only nine ounces will kill a large, mature 
sheep (weighing 135 to 165 pounds) within six to 10 hours due to “rapidly 
overwhelming loss of inorganic serum calcium.” Vawter (1950, 1951) 
listed the symptoms as dullness, prostration and coma. Kidney sections 
examined under polarized light show large crystal aggregates of calcium 
oxalate in the tubules. Anderson and Huffman (1957) indicated these changes 
resulted in acute renal failure. According to Vawter (1950) there “seems to 
be no practical way of saving sheep after eating a lethal dose of the plant.” 
The toxic form of oxalate was determined by Dye (1956) to be the soluble 
form, insoluble calcium oxalate being relatively harmless. Binns and James 
(1961) discussed the mechanism of sodium oxalate in the blood in pre- 
cipitating calcium (with resultant alkalosis) and stated that the feeding 
of alfalfa pellets, supplemented with dicalcium phosphate, was helpful in 
preventing (not curing) poisoning in sheep. 

More detailed study of the physiology of halogeton, particularly oxalate 
content and metabolism, was the concluding phase of an intensive two- 
decade study. Morton et al. (1959b) determined the oxalate content of 
plants to range from 18 to 21 percent of dry weight (about the usual range 
reported in the literature), declining somewhat during the growing season. 
Most of the oxalate was contained in the leaves, with relatively small amounts 
in the stems, and high oxalate concentration was associated with a high 
sodium content. Williams (1960) observed that soluble oxalate content of 
the plant, as well as growth and succulence, increased when sodium (as 


160 


NaCl) was added to nutrient solutions. In contrast to Morton et al., Williams 
found that the concentration of oxalate in the leaves continued to rise during 
the summer, attaining a peak in September. According to Walliams, the 
leaves of vigorous plants often contain in excess of 30 percent soluble oxalate 
on a dry weight basis, with an additional three or four percent of the dr 
weight of the plant accounted for by insoluble oxalate. A dry-weight soluble 
oxalate content of 34.5 percent was reported by Cook and Gates (1960), a 
high percentage compared to most other oxalate-containing plants. Kings- 
bury (1965) indicated a 10 percent soluble oxalate content to be high 
for beet tops (considered potentially dangerous forage), well below the 
usual estimates for halogeton. 

Both popular and scientific liceracure on halogeton decreased after 1961, 
although Kingsbury (1964) devoted attention to it in his comprehensive 
work on poisonous plants of the United States and Canada. Sheep losses 
from oxalate poisoning also declined, apparently from an awareness among 
sheep ranchers. Nonetheless, occasional reports of severe loss, particularly 
in Utah, continued to surface in various newspapers. As recently as 1971 
(Associated Press, Salt Lake City, January 23), 1,180 head from a flock of 
2,400 died near Garrison, Utah, close to the Nevada border. This report 
observed that “there’s halogeton all over that area.” Thus, a “slow down” in 
the literature probably did nor reflect a decrease in populations, but rather 
indicated better education concerning the plant, a lessening of the earlier 
somewhat faddish interest in the weed, and the realization that halogeton 
would not actually kill one out of three sheep in the United States (as 
suggested by Life Magazine). 

Questions left unanswered or inadequately answered include: When did 
halogeton enter the North American flora and by what means? What has 
been the pattern and the mechanism of its spread? Is 1t more common now 
than it was say 30 years ago? What is its probable future in the flora? Is 
more than one species of Halogeton present in the United States? And, per- 
haps surprisingly, what is actually the correct scientific name(s)? By taking 
a close look at the history of the literature on halogeton, and by examination 
of actual specimen records, this paper attempts to deal with these questions. 


NOMENCLATURE 


Since its appearance in the flora, three different names for halogeton in 
North America have found their way into the literature: Halogeton sativus, 
H. souda, and H. glomeratus. These three names are to be found among 
specimens in North American herbaria (although H. glomeratus is the 
common determination ). 

In 1936, the 77th Intermountain Forest and Range ee Station 
collection was forwarded to the Washington Office of the United States 
Forest Service (Dayton, 1951). These plants included two specimens (S-43 
and §-44, collected in 1934 by Ben Stahmann, a Forest Service officer, one 


161 


mile northeast of Wells, Elko County, Nevada) which were the first records 
of halogeton for the U.S. These two specimens were not immediately identi- 
fied. However, similar material (Field Museum, F 780344 and F 815074) 
collected by Mr. Stahmann from Wells in 1935 was subsequently deter- 
mined by Dr. Paul Aellen, an authority on the Chenopodiaceae, as “Halogeton 
sativus’ (based on Salsola sativa L., 1762), native to Spain and Algeria. 

Standley (1937) first published on halogeton in North America, accept- 
ing Aellen’s determination of Stahmann’s 1935 material as Halogeton sativus 
(L.) C. A. Meyer. Standley, seemingly unaware of Stahmann’s 1934 collec- 
tion, commented that the genus was first collected in “1935” at Wells, and 
speculated that “it will be interesting to learn whether this new addition 
to the United States flora thrives as well as Bassia hyssopifolia ( Pall.) Kuntze, 
a related plant of somewhat similar appearance, which, if I remember cor- 
rectly, also was found in the United States first in Nevada.” Standley further 
stated that “it [halogeton} is reported to be spreading rapidly.” Confusion 
persisted in the literature as to whether the initial collection of halogeton 
was made in 1934 or 1935 (e.g. ZoBell and Silcock, 1950); however, her- 
barium records clearly show that 1944 is correct. 

The question of the initial year of collection has not been the only con- 
fusion enshrouding halogeton in North America. Morton (1941) correctly 
observed that the publishing author of the combination Halogeton sativus 
was actually Moquin-Tandon (1840), rather than C. A. Meyer (as suggested 
by Standley and presumably also by Aellen). Although Meyer (1829) 
mentioned Salsola sativa, he never employed the combination Halogeton 
sativus (as did Moquin-Tandon ). Therefore, the correct citation is Halogeton 
sativus (L.) Moqguin-Tandon, not (L.) C. A. Meyer. However, Morton 
(1941) went on to discuss the existence of an earlier name for this species, 
viz. Halogeton souda (Loefling) Macbride (based on Salsola souda Loefling, 
1758). Our examination of photographs of the type specimens of both 
Salsola sativa and S. souda in the Linnaean herbarium (LINN) confirms that 
they are probably the same species. Thus, if the adventive American taxon 
were considered to be the Spanish-Algerian species, Halogeton souda would 
be the correct name (on the basis of priority), with H. sativus relegated to 
synonymy. Fosberg (1940) used the name H. sowda for halogeton in Nevada. 

The question of Halogeton sativus vs. H. souda is relatively moot to our 
considerations, however, because Morton (1941) further indicated that 
North American specimens are not Halogeton souda (H. sativus) at all but 
belong instead to a Russian species, Halogeton glomeratus. It 1s apparent 
that Morton examined Old World specimens of both H. sowda and H. 
glomeratus, although probably not the type specimen of either. Dayton 
(1951) subsequently offered a brief review of these taxonomic and nomen- 
clatural problems, concurring with Morton. However, Dayton stated that 
the type of H. glomeratus was behind the “Iron Curtain,” presumably in 
Moscow or Leningrad, and not available oe examination. Thus, ultimate 


162 


certainty of the identity of North American specimens must have eluded 
Morton and Dayton. 

Through the auspices of Dr. V. I. Grubov, we obtained a loan of speci- 
mens from the Leningrad herbarium (LE), along with photographs (nega- 
tives) of the type specimen of H. glomeratus. Prints made from these nega- 
tives matched well with the specimens collected in the United States. Thus, 
North American specimens can now be classified with reasonable assurance 
as H. glomeratus, not H. souda (H. sativus). 

Dayton (1951) and others indicated that the combination Halogeton 
glomeratus is traced to C. A. Meyer (1829) when he described the genus 
in Ledebour’s Flora Altaica. Dayton, however, noted that C. F. Stephan 
(author of a Flora of Moscow) was actually the first to use the epithet (in 
connection with herbarium material) but that Stephan himself did not pub- 
lish it. Stephan’s specimen, however, is the type of Halogeton glomeratus, 
and is the specimen in the Leningrad herbarium. This specimen (lacking a 
date) is labeled “Salsola glomerata’ in Stephan’s handwriting. The name 
written by Stephan is a herbarium or “manuscript” name. However, Stephan’s 
epithet “glomerata’ was later validly published by Marshall von Bieberstein 
(a colleague of the well-known Russian botanist, Pallas) as Anabasis 
glomeratus (1806). Meyer (1829) subsequently transferred Bieberstein’s 
glomeratus to Halogeton. The most complete and accurate author citation 
of the name (which we have understandably not seen published ) would be 
Halogeton glomeratus (Stephan ex Bieberstein) C. A. Meyer in Ledebour. 

Tutin (1964) considered Halogeton glomeratus and H. sativus (= H. 
souda) to be closely related species. In addition to the geographical separa- 
tion, the Spanish species is to be distinguished by longer leaves, a more 
robust habit, longer perianth, and a greater number of stamens per flower. 
Our examination of Specimens and phototypes lends support to those who 
eae these two species. However, if a future monographer were to com- 

ne H. glomeratus and H. souda, the name would then become H. souda 
on the basis of priority. 


—_ 


Halogeton, as is the case with many other chenopodiaceous genera, is in 
need of revision on a worldwide basis. This is evident from the variation 
in estimates as to the number of species of which the genus is composed: 
Ulbrich (1934) indicated that there are three species; Ijin (1936) called 
for four; and, somewhat inexplicably, Dayton (1951) suggested that there 
are 12. Regarless of whom one believes, there is agreement that more than 
one species exists. However, we consider that only one species (H. glomera- 
tus) has been and is in the flora of the United States. We find no evidence 
to support Dayton’s (1951) speculation that “it is perhaps only a question 
of time when one or more of these other species will emigrate to this 
country.” 


163 


INTRODUCTION, SPREAD, AND DISTRIBUTION 

Neither the manner nor the date of introduction of halogeton into the 
United States has been definitely established (Tisdale and Zappetini, 1953). 
Several references (Bellue, 1949; Standley, 1937; ZoBell and Silcock, 1950) 
indicated halogeton first entered the North American flora in 1935, although 
it was already common in the Wells area by 1934 (Dayton, 1951). Ic is 
apparent in the case of several “first records” for particular regions that 
infestations were extensive at the time of discovery (Bellue, 49; Velie, 
1953). Even the nature of various of the livestock losses reported, i.e. severe 
and sporadic, is an indication that it became abundant in certain areas prior 
to recognition. Stoddard et al. (1951b) set the time of entry back somewhat 
in suggesting the probable date of introduction from southern Russia “in 
about 1930.” Binns and James (1961) likewise favored 1930 as a likely 
date of introduction. 

Regardless of the exact time of introduction, it appears that halogeton 
became more common in western North America in the early 1930's. During 
this period an extremely dry climatic circumstance prevailed throughout 
most of the West. It is probable that the drought tolerant halogeton extende 
its populations (at least locally) while many native species suffered decline 
(Tisdale and Zappetini, 1953). This species does not compete well with 
healthy native perennial forbs (Fenley, 1952), yet it enters in force when 
soils are denuded (Erickson et al., 1952). As many as 50,000 winged seeds 
per plant (Morton et al., 1959a) are Ae locally by the wind (Kings- 
bury, 1964). This and a wide ecological tolerance range permits se 
to survive in habitats unfavorable to competitors (Erickson et al., 1952) 
and to “take over” local disturbed or barren areas quickly. The “Dust Bowl” 
of the western United States in the 1930's assuredly contributed to its spread. 

Although undocumented, halogeton must have been present in North 
America before 1930 (Erickson et al., 2). Erickson and his coworkers 
suggested three possible sources of Pees 1) impurity in crop seed, 
2) imported wool, or 3) importation with breeding stocks of sheep. Durrell 
(1951) and an anonymous writer in Life Magazine (1951) supported the 
introduction into this country as a contaminant in imported seed. Erickson 
et al. (1952) favored wool pelts or live animals as the vector, pointing out 
that new infestations coincide with sheep movements. Bellue (1949) simply 
commented that “Halogeton seed seems to have an affinity for and readily 
sticks to the wool of sheep,” and that in Nevada, Utah, Idaho and Wyoming, 
the weed has “followed sheep trails and bedding grounds.” Karakul ed 
among other possibilities, has been mentioned as a potential source of i 
troduction and the most likely vector from the Soviet Union. In contrast to 
other breeds (including Angora), the natural range of the Karakul corre- 
sponds closely to that of halogeton. Karakul are native to desert-like areas 

e Bokhara Province of southern Russia, and sheep with Karakul blood 
extend as far westward as the Caspian and Black Seas (Vaughan, 1931). 


164 


Halogeton Ses is well known from this general area (Ijin, 1936). 
Erickson et al. (1952) mentioned that both Karakul and ae. breeding 
stocks were oe into the United States between 1908 and 1925. Went- 
worth (1948) also alluded to the importation of Karakul early in the 
twentieth century. Vaughan (1931) identified “three lots” of Karakul that 
were imported (by Dr. C. C. Young), in 1908, 1912, and 1914 respectively. 
If Karakul were the vector, then 1908-1914 would be more likely the time 
of introduction than 1930 to 1934. 

Circumstantial evidence for the ovine origin of halogeton infestation 1s 
found in the implicating comparison (Figure 1) between sizeable halogeton 
populations and major sheep trails of the early part of the century (these 
trails were in heavy use until about 1915). Several primary routes were 
used in sheep drives out of California (Wentworth, 1948), including south- 
ern (leaving Bakersfield), middle (leaving Fresno), and northern (leaving 
Red Bluff) passages. The southern and middle routes passed along the 
eastern flank of the Sierras, then across Nevada, proceeding through Wells 
(a major crail junction), and up into the Raft River Valley of southern 
Idaho. Tisdale and Zappetini (1953) pointed out that the Raft River Valley 
area was one of the principal winter ranges for sheep in the early 1900's. 
The northern route passed directly through the well-documented area of 
infestation in Lassen County, California, on the way to Wells and southern 
Idaho. Thus, the correlation between major sheep trails and a number of 
significant halogeton infestations is, in our Opinion, not coincidental (Figure 
1). Localities plotted are based on specimens examined from the following 
herbaria (Holmgren and Keuken, 1974): A, ARIZ, DAO, F, GH, ID, MU, 
NY, OSC, PENN, PH, RENO, RM, SMU, UC, US, USFS, UT, and WTU. 
Additional information on specimens was provided by: CAN, COLO, TEX, 
and UNM. 

We consider, by virtue of the native home of the Karakul, its long fur 
(Persian lamb’s wool) in which seeds might easily become entangled, the 
time of introduction of Karakul into the United States, and the apparent 
relationship between old sheep trails and the distribution of the plant, that 
halogeton was both introduced and subsequently dispersed by live sheep. 
Introductions by means of fur pelts or grain shipments are conceivable, but 
not as probable. Karakul skins were mailed in bundles of 500 each from 
Asia directly to New York City where they were processed (Vaughan, 1931), 
and therefore had no connection with halogeton growing in the western 
states. Grain shipments contaminated with halogeton seed could have ac- 
counted for a few populations of halogeton. Seed contamination has played 
a major role in the introduction of certain other chenopodiaceous weeds 
(Blackwell, 1978; Collins and Blackwell, 1979). But if the majority of 
infestations had actually been caused by contaminated grain, there should 
be no correlation between its occurrence and well-known sheep trails. Also, 
railroads for shipping grain were not established early in the 1900's through 


Figure 1. Distribution of Halogeton glomeratus in North America. Stars with 


Halogeton populations based on herbarium specimen records. Sheep trails (double 
broken lines) are modified from E. N. Wentworth, 1948, America’s Sheep Trails. 


166 


some of the areas of heaviest infestation. Thus, live animals, in particular 
the Karakul, would seem to have been the vector, and we believe halogeton 
to have been present and growing in the United States by 1915. It is some- 
what ironic that the type of animal (sheep) which has been most adversely 
affected by its toxicity also appears to have been the source of introduction 
and dispersal. 

The assumption (e.g. Stoddard et al., 1951b; ZoBell and Silcock, 1950) 
that halogeton was first introduced into Nevada, from there spreading to 
the other states (Utah, Idaho, Wyoming, etc.), may be invalid. Even though 
it spreads locally in great numbers, there is no evidence that it is dispersed 
across large distances without the aid of an external vector (such as sheep ). 
Kingsbury (1964) indicated that wind is primarily effective only in local 
dispersal. Thus, a majority of present extensive areas of infestation probably 
correspond to the original sites of introduction by migrating sheep. While 
Wells, Nevada, was one of those early sites, it is mot possible to be sure 
that it was the earliest. It is reasonable to say only that it was first collected 
there. It is conceivable, for example, that one or more of the Raft River 
Valley localities in Idaho might predate the Wells, Nevada, occurrence. 
There is no proof one way or the other. Since halogeton was present for a 
number of years, probably in a number of locations, prior to its discovery, 
it is not likely that the first site of occurrence in this country will ever be 
conclusively identified. But even if Wells were accepted as the initial locality 
(as it is in much of the literature), an autogenic spread of halogeton from 
there to other widely separated localities is less than plausible. 

Bellue (1949) inferred that halogeton migrated from northwestern 
Nevada (located across the state from Wells) over the border into Lassen 
County, California. Yet, this rather isolated occurrence in northeastern 
California may have had nothing to do with its presence in Nevada. The 
California infestation could just as well have been initiated in situ in the 
early 1900's by a sheep drive. Bellue stated that “a survey made subsequent 
to our first California record in Lassen County in 1946, indicated that Halo- 
geton had become scattered over several thousand acres.” This extensive 
infestation did not develop overnight and it seems as feasible to us to specu- 
late that halogeton had been there for some time (the introduction having 
taken place at or near the site) as to postulate a recent invasion from 
adjacent Nevada. 

We believe that, whereas it is possible to determine a pathway of infesta- 
tion (and a probable cause), it is not possible to establish meaningful 
chronology. Halogeton was, we think, independently and separately intro- 
duced by sheep into scattered localities, sooner or later becoming locally 
common in some of them; and since these introductions probably took place 
well before discovery of the plant, the order in which they transpired can- 
not be determined. If it were possible to establish the chronology of in- 
festation, however, it is unlikely that it would follow an orderly geographic 


167 


sequence. Based on herbarium specimen records, and without any necessary 
implication of the sequence of spread, we can state that the first collections 
known in the following states were in the following order: Nevada, 193 
Utah, 1943; Wyoming, 1943; Idaho, 1946; Montana, 1951; California, 1951; 
Colorado, 1951; Oregon, 1954; and New Mexico, 1973. Halogeton was ob- 
served five years prior to collection in California (Bellue, 1949) and at least 
one year before collection in Idaho (Morton et al., 1959a). Thus, specimen 
records offer only a rough gauge of the time of awareness of its presence in 
the various states, and little or nothing about the actual time of its appear- 
ance in the state's flora. 

Opinion has varied as to the prognosis for halogeton in North America. 
Holmgren (1950) stated that “it’s migration into new areas is continuin 
at an alarming rate,’ and such a sentiment is echoed in popular articles in 
Life Magazine (anonymous, 1951) and Reader’s Digest (Velie, 1953). More 
conservatively, Erickson (1952) stated that “the future distribution of halo- 
geton cannot be forecast with accuracy at this time.” Erickson did mention 
the possibility that it could eventually spread over the entire Intermountain 
Shrub Zone (Cold Desert region) and even into the western margins of the 
grasslands lying to the east. Perhaps it was simply too early, and with too 
little information at hand, to forecast accurately the future of halogeton in 
the United States, except to postulate that it would certainly persist (Stod- 
dart et al. 1951la) and probably would not become any less common with 
time. 

It appears that the present major areas of halogeton infestation are 
essentially those aca in publications of the 1950's. Simple maps pre- 
sented by Erickson et al. (1952) and the U. S. Poe of Agriculture 
(anonymous, 1958), outline these areas rather well. To the USDA map, 
new localities can be added primarily in southern Oregon ia northwestern 
New Mexico. According to herbarium records, no new “halogeton states,” 
except for New Mexico, have been added to the list since 1954. Thus, it 
would seem that whereas halogeton can and does spread locally at a rapid 
rate, it does not readily establish new widely disjunct areas of infestation 
by its own devices. Bellue (1949), for example, noted that “the range o 
Halogeton is still confined to the original general location in the southeastern 
corner of Lassen County.’ And in the summer of 1977, two of us (Blackwell 
and Hopkins) carried out an extensive search throughout adjacent Modoc 
County, California, without turning up a single specimen. Suitable habitats 
were apparently abundant in Modoc County, yet halogeton had not sprea 
there in more than 30 years of opportunity from either Lassen County or 
adjacent Nevada. Morton et al. (1959a) indicated that the total area of 
halogeton infestation was 10.5 million acres (though it is not clear how this 
figure was derived). It does not seem that the area of infestation today is 
much larger than this, although an accurate estimate is lacking. en com- 
pared with the USDA map of 1958, major areas of infestation do not 


168 


appear to have increased strikingly. This is not to imply that there have 
not been numerous instances of local increase, or numerous rather closely 
spaced small new infestations in various parts of the Intermountain Region. 
Undoubtedly, the fact that major sheep drives have drastically declined as a 
means of transporting sheep great distances is related to the present minor 
spreading of the plant. New areas of halogeton infestation now perhaps most 
commonly originate in disturbed habitats along roadways (personal observa- 
tion, Blackwell, southern Oregon, 1977). Infestations in eastern Utah and 
western Colorado, New Mexico, and Oregon apparently developed in this 
manner. Thus, spread (not necessarily rapid) along roadsides allows move- 
ment into poorly managed rangeland. 

Control of halogeton remains a concern. Of the various alleged methods, 
biological control has come to be among the more common suggestions. In 
particular, the use of crested wheatgrass (Agropyron cristatum Gaertn.) as 
a competitor was recommended (Fenley, 1952; Stoddart et al., 195la) 
However, this, as various chemical control measures, produced dubious re- 
sults (Morton et al., 1959a). Halogeton and crested wheatgrass were observed 
growing successful ly together in Lake County, Oregon (Blackwell, personal 
observation, summer 1977). Perhaps Morton and his collaborators (1959a ) 
summarized it best in stating that “the successful control of halogeton does 
not depend upon any single control method . . . Ultimate control of halogeton 
depends upon the restoration of vigor and productivity to good range forage 


plants 


REFERENCES 


ANDERSON, W. A. and W. T. HUFFMAN. 1957. Halogeton poisoning in a ewe. 
J. Am. Vet. Med. Assoc. 130: 330-331. 

ANONYMOUS. 1951. Sheep- killing weed. Life Magazine, Jan. 15: 55-56. 

ANONYMOUS. 1958. Reducing livestock losses from halogeton poisoning in the 
western states. U.S. Dept. Agri. Bull. PA-321 

ARMER, W. . tac a stock- poisoning weed, Ariz. Agri. Exp. St. Bull., 
Ext. Circ. 197. 

BELL i M. K. ioe Halogeton invades California. Calif. Dep. Agri. Bull. 38(1): 


32-306. 

BIEBERSTEIN, F. A. M. VON. 1806. ieee Societe Imperiale des Naturalistes 
de Moscov, Memoires, vol. 1 (ed. 2): 108-1 

BINNS, W. and L. F. JAMES. 1961. a ee and other oxalic acid poisonings. 
Proc. Am. Col. Vet. Toxicol. 1960: 5-8. 

BLACKWELL, W. H. 1978. The history of Russian pigweed, Axyris amaranthoides 
(Chenopodiaceae, Atripliceae), in North America. Weed Sci. 26: 82-84. 

COLLINS, S. L. and W. H. BLACKWELL. 1979. Bassia (Chenopodiaceae) in North 
America. Cg 8: 57-64. 

COOK, C. W. and D. H. GATES. 1960. a of site and season on oxalate content 
ie ie. J. Range Manage. 13: 97— 

YTO W.A. 1951. cae nar o barilla (Halogeton glomeratus). J. 

eee Manage. 4(6): 375-38 

DURRELL, L. W. 1951. Ee ee a new stock-poisoning weed. Colo. Agri. Exp. 
St. am 170-A. 5 pp 


169 


DYE, W. B. 1956. Chemical studies on Halogeton eahitrs Weeds 4: 55-60. 
ERICKSON, L. C., H. L. MORTON, E. W. TISDALE, and G. ZAPPETINI. 1952. 
The present status of the halogeton problem. Weeds 1(4): aoe 328. 
FENLEY, J. M. 1952. How to live with halogeton while limiting its spread and_re- 
ducing loss. Nev. Agri. Ext. Serv. Bull. 106. 20 pp. 
FLEMMING, C. E.,, M. R. MILLER, L. R. VAWTER, and W. NEILSON. 1943- 
1949. Poisoning range plants. Nev. Agr. Exp. St. Ann. Rpts. 
FOSBERG, F. R. 1940. Chenopodiaceae of Nevada (in part). Contrib. Toward a 
Flora of Nevada, No. 8: 1-18. Mimeographed. 
oe A. 1950. Halogeton ee eastern Utah winter ranges. Utah Agri. 
, St., Farm Home Sci., 11(4 
foe P.K. and W. carat er Index herbariorum (6th ed.). Regnum 
Veg., Vol. es Oosthoek, Scheltema and Holkema, Emmalaan 27. Utrecht, Nether- 
lands, 397 
i M. 1936, Sree In V. L. Komarov and B. K. Shishkin. Flora of the 
R, 6: 4-272. (English Translation, 1970, by N. Landau. ) 
ane J. M. 1964, Poisonous plants of the United States and Canada. 
Prentice Hall, Inc., Englewood Cliffs, New Jersey. 626 pp. 
er ctale J. M. 1965. Deadly harvest. Holt, Rinehart and Winston, New York. 


eon e A. 1829. Halogeton In C. F. Ledebour. Flora Altaica, 1: 378-380. 
MOQUIN-TANDON, C. 1840. Cheaopodearn monographica enumeratio. 182 pp. 
MORTON, C. V. 1941. A note on u paloget Leafl. West. Bot. 3(2): 45-46. 
MORTON, H. L., R. H. HAAS, and L. C. TFRICKSON. 1959a. Halogeton and _ its 
control. Idaho Agri. Exp. St. Res Bull. 307. 
TON, H. L., R. H. HAAS, and L. C. aa om 1959b. Oxalate and mineral 
contents of Halogeton glomeratus. Weeds 7(3): 255-264. 
STANDLEY, P. C. 1937. Studies of Arete plants—VIII. Field Mus. Nat. Hist. 
Publ., Bot. air hee 225-284. (Publ. 395 ia 
STODDART, G. T. BAIRD, G. STEW RT, R. S. MARKHAM, and 
ses oan ‘the halogeton problem in ce Utah Agri. Ext. cee Bull. 050 


cnn L. A., H. CLEGG, B. 5S. ee se G. rears 1951b. The 
halogeton problem on Utah’s ranges. J. Range Manage. 4 ( 223-2 

TISDALE, E. W. and G. eee 1953, ae a on 4 aho ranges. 
J. Range Manage. 6(4): 225-236. 

TUTIN, T. G. 1964. ee. In Tutin et al. Flora Europaea, 1: 90-108. 

ULBRICH, E. 1934. Che pee In Engler and Prantl. Die Natiirlichen 
Pflanzenfamilien (ed. 2), 16c: 379-584. 

VAUGHA N, H. ue ee ee of livestock in America. R. G. Adams and Co., 


cattle. “Calif. Vet. 316). 

VAWTER, L. R. 1951. Teor poisoning in sheep. Natl. Wool Grower 41(2): 
24-25. 

VELIE, L. 1953. Poison rides the range. me Reader’s Digest. December: 79-82. 
Soles from the Denver Post, Nov. 8, 1953) 

WENTWORTH, E. N. 1948. America’s sheep Gaile: Iowa State Univ. Press, Ames. 


667 pp. 

WILLIAMS, M. C. 1960. Effect of sodium and Fea salts on growth and oxalate 
content of halogeton. Plant Physiol. 35: 500-505. 

ZOBELL, R. S. and ae W. SILCOCK. 1950. nes Russian invades U.S. Natl. Wool 
Grower 40(4): 


CHANGES IN GALACTIA (FABACEAE) 
OF THE SOUTHEASTERN UNITED STATES 


WILBUR H. DUNCAN 
Department of Botany, University of Georgia 
Athens, Georgia 30602 


ABSTRACT 


ifferent names and realignments are presented for several species of Galactia. 
It is shown that G. volwbilis (L.) Britt. and a macreei M, urtis sensu most 
recent authors should be called G. regularis (L.) BSP and G. Were (L 


respectively. G. glabella Michx. is supported as the sean name for what has 
B 


. reg G. glabella is shown to 
of three partly sympatric forms but studies are insufficient to decide what 
if ¢ uld be assigne : 


taxonomic status, if any, sho be igned to diagnostic key for eight 
species and distribution maps for six species are presented. More detailed diagnoses 


y are presented for better understanding of G. regularis 
glabella, volubilis, and mollis, specimens of which have been abundantly misidentifie 


Galactia in the Southeastern United States has interested me for at least 30 
ears. Intermittently I have attempted to solve some of the many complex 
problems inherent in this genus, but only recently have sufficient data and con- 

aoe been accumulated to justify their presentation. First I described one 
new species (Duncan, 1977). I am now presenting my remaining con- 
clusions, fully aware that some may rot hold up under future studies but 
also realizing the unlikelihood of my making further studies of this subject. 
Taxa confined to peninsular Florida are omitted due to insufficient study of 
materials from there. A key to all other species of the Southeastern United 
States as I interpret them follows. 


KEY TO SPECIES 

1. Leaves with 5—9 leaflets Coe ee eee eG. ebliottii 

1. Leaves with 3 leaflets. Se we © ay eee ee ER Se ee 
2. Plants erect, pps sulseeseile G. erecta 
2. Plants twining or trailing, inflorescences pedunculace 3 

. Calyx lobes ana to reddish brown on inner surface when dry; ‘corollas dark 
colored when dry, persisting after ha sometimes partly present wl 
fruit is mature G. mollis 
Calyx lobes preenich: cele to tan on inner bern ace when ays gonollas an 
pare when dry, falling as they wither or soon thereafte 

4, Stems with fine close mostly ieee hairs, the ee. 0. 7 ge or enge: 


S>) 


7. floridana 
4, Stems with fine antrorse, retrorse, or spreading haits under mm 
rarely a few longer, but hairs not dense 
5. Hairs on stems antrorse . 
6. Internodes only a little longer to usua lll suis than thet Renede leaflet 
of adjacent nodes, hairs on stems 0.05—0.25 mm long G. 


> 


minor 


SIDA 8(2): 170-180. 1979. 


171 


6. Several to most internodes (especially those toward the base) much 
longer than the oe t leaflet of uae nodes, hairs on the stem 
lon G 


.1-0.8 mm long : : . glabella 
5. Hairs on stems retrorse to i. gereadine ee es ae a al OF 
Mature flowers 7-9 mm long . . woe ee. GG. regularis 


7. Mature flowers 10-18 mm long 
8. Longest inflorescences 3-15 cm long, nodes. near the tip ‘close and the 
6-9 G. 


flowers congested; ovules 6— glabella 
8. Longest inflorescences 5-55 cm Tong A dower in separated ns and 
not congested near tip; ovules 10-13 : G. volubilis 


DISCUSSION OF SPECIES 


G. ELLIOTTI Nutt. and G. ERECTA ( Walt.) Vail 

These species have distinct features and are rarely confused with others in 
contrast with all other species, which are frequently misnamed. 
G. MOLLIS Nutt. 

This taxon js often confused with G. floridana. In addition to the characters 
in the key the legume of G. mollis has dense divergent hairs prominently 
visible macroscopically while in the former species the hairs are scarcely or 
not visible macroscopically and are retrorse-appressed to retrorse-spreading to 
a few scattered hairs being divergent. 

G. FLORIDANA T. & G. 

r the present this species is best treated as two varieties, floridana and 
microphylla Chapm. The latter has shorter internodes, smaller leaflets, shorter 
petioles, and the inflorescences are shorter than the leaves. Inflorescences are 
mostly longer than the leaves in var. floridana. Additional studies are needed 
to test this separation more thoroughly. Growing plants under the same con- 
ditions is likely to be helpful 
G. MINOR Duncan 

Additional aids to the identification of this taxon are: stems geniculate (un- 
commonly not), largest leaflets 14-28 mm long, inflorescences with 1-3 (4) 
flowers, the longest inflorescence little if any longer to shorter than the sub- 
tending leaf, and flowers 11-17 mm long. 

G. REGULARIS (L.) BSP. 

r many years there seems to have been little disagreement among tax- 
onomists concerning the application of the name G. regwlaris. Descriptions, 
although differing in some details, indicate that each author had the same 
taxon in mind. These authors include Small (1933), Fernald (1950), Gleason 
and Cronquist (1963), Wilbur (1963), and Radford et al. (1964). Under 
this name until now, as indicated by the descriptions, have been included 
individuals that were prostrate or rarely twining; had inflorescences shorter 
or longer than the leaves and with one to many flowers, nodes close in the 
upper part of most inflorescences, and flowers congested or few; flowers 10-18 
mm long and with a calyx 6-10 mm long; and stem hairs 0.05—-0.8 mm long 
and usually retrorse, but frequently antrorse and sometimes spreading [descrip- 
tion mine}. 


172 


G. MOLLIS 


pode tase 


é 


oo 


FLORIDANA 
MICROPHYLLA ~ 


Tee 
DN INK 


o 


G. MINOR 


Wor 1-3. 


‘ ag 

Ls i i we 

gy a A Os 

BPpeayt Pps 
=} L | ' Z 


am. 


ii) 


Although at first glance the type specimen (Virginia — Clayton — ex. 

Herb. Gronovii in BM) in some respects looks like a specimen of G. regularis 
sensu recent authors, it differs significantly from the above description in 
several details. The calyx is only about 5 mm long, the flower only 8 mm 
long, the upper flower clusters are evenly spaced and not congested, and the 
hairs on the stem are soft and spreading which is very unusual for G. regalaris 
as it has been interpreted. These data fit in all respects what has been treated 
variously as G. volubilis (L.) Britt. (Small, 1933, in part; Fernald, 1950; 
oe 1963; Gleason and Cronquist, 1963, in part; Radford et al., 1964), 

G. volubilis var. mississippiensis Vahl, and G. mississippiensis (Vahl ) Rydb. 
Also the description by Linnaeus (Se Pl. I: 726. — no. 8, DOLICHOS 
regularis) can as easily be applied to this taxon as to what has been passing 
as G. regularis. Therefore, the name G. regularis should be used for G. volu- 
bilis sensu Fernald (1950) et al. 

During the checking of hundreds of herbarium specimens, I noted that 
G. regularis and G. mollis often had been confused. The hairs on the stem 
do seem to intergrade in size and abundance, with G. mollis generally hav- 
ing the longer and more abundant hairs. A series of measurements of hairs 
on specimens of known identity probably will show that almost all speci- 
mens of these two species can be identified by hair characters. Until this is 
done, except for sterile specimens, there are other means of identification as 
follows: 

Calyx greenish-yellow to tan on inner surface when dry; corolla light colored when 
dry, falling as it withers or soon afterwards; legume Note scattered diver - nt to 


antrorse-appressed hairs scarcely or not visible macroscopica . G. regularis 
Calyx brown to reddish brown on inner surface w as dry; ar a anu co ied when 
dry, persisting after withering, sometimes partly present when fruit is gs 
with dense divergent hairs prominently visible oe : i 7 ollis 


A partial synonomy of G. regwlaris should be useful in adjusting to the 
new application of this name and follows: 
Galactia regularis (L.) BSP. Prel. Cat. N-Y. 14. 1888. 


Dolichos regularis L. Sp. Pl. 726. 

?Hedysarum volubile L. Sp. PI. a en 

?Ervum volubile Walt. Flr. Car. 187. 1788. 

Galactia mollis Nutt. Gen. 2: 117. 1818. Not Michx., 1803. 

Galactia pilosa Ell. Bot. S.C. and Ga. 2: 238. 1824 _ Not Nuttall, 1818. 

Galactia villosa Eaton & Wright. N. Am. Bot. 248. 1840. Not Wight and Arnott, 
Prodromus Fl. Ind. Orient I. 1834. 

Galactia volubilis (L.) Britt. Mem. Torr. Club 5: 208. 1894. Sensu:—Small, 
933, in part; Gleason & Cronquist, 1963, in part; Fernald, 1950; Wilbur, 
1963; Radford et al., 1964. 

Galactia ia (L.) Britt. var. misséss7ppiensis Vail. Bull. Torr. Bot. Club 22: 
500. 189 


Galactia ee Vail. Bull. Torr. Bot. Club 22: 500. 1895. 


G. GLABELLA Michx. 
In order best to understand the application of this name it is useful to 


174 


form of 
GLABELLA 


GLABELLA 


, o 
@ w 
: | < aN oe , 
. Ro oo 
: 4 o U8 : 
° = 
nM * oc q 
un8 Fal @ Pog 
=e 7 a 
Pe ee ; 
esr oe “54 
+ eae — ee ed Ge ee | 
ee aaa ; 
ert. f : v\e 
T jt is { eS, } ‘a 
4 4 is 1 ek ss 
STALL st gN \ ope : 
ee OR ae ; 
__¥ fa N ~ aS a 5 
N NSN 1 N nS “+ 
PaL-t—2] Ee 7 ; 
entlizike ¢ ki, 
EAL wu 
y, # AR pee “psd 
” - i b a 
a 


175 


discuss the variability of the plants that have long been designated as G. 
regularis. Actually there are two and maybe four taxa involved. One is G. 
minor which has been discussed. The remaining plants of the former G. 
regularis consist of at least three types of plants. One has retrorse or very 
rarely retrorse-spreading hairs. The other two types have appressed antrorse 
or very rarely partly antrorse-spreading hairs. No plants have predominately 
divergent hairs. The three types have definite distributional patterns which 
coincide only partly. They have some characteristics in common which are 
different from those of G. minor, including having the stems straight, not 
geniculate; several to most internodes much longer than the longest leaflet 
of the adjacent nodes; the longest inflorescences longer than the subtending 
leaves, uncommonly shorter, and with several to many flowers, rarely as few 
as 4, the upper nodes close and the flowers congested in the larger inflo- 
rescences. It is interesting that Small (1933) reserved the name G, regularis 
for those individuals having “minutely retrorse-pubescent” stems. However, 
none of the other species he includes can be the antrorse-haired G. minor. 

The “retrorse” population is much more abundant than the “antrorse” 
ones. Its characteristics essentially are those recognized in publications for 
those plants until now designated as G. regularis. The next oldest name that 
possibly could apply to this “retrorse” taxon is apparently ERVUM? volubule 
(Walter Flr. Car. 187. 1788). However, Linneaus (Sp. Pl. II: 750. 1753) 
had used the epithet previously describing the plants as having “caule volu- 
bile,” which is unusual for this taxon. Furthermore, I shall show later that 
voluble applies to another taxon. 

I believe, however, that Michaux’s Galactia glabella (Fl. Bor. Am. 2: 62. 
1803) can be used for the “retrorse” taxon with reasonable confidence. 
Michaux’s description of the plants as being (translation) “almost glabrous: 
leaflets oblong-oval, both ends shallowly notched, obtuse: calyx glabrous” 
is unlikely to apply to any other Galactia, and especially to those taxa known 
to occur where the type specimen was collected, namely G. volubilis and the 
“retrorse” plants. At flowering the calyx of G. volubilis is nearly always 
appressed hairy and only rarely nearly glabrous. On the other hand, the 
calyx of specimens of the “retrorse” plants is often glabrous or nearly so. 

The type specimen of G. glabella (Michx. Herb—Col. Co.—S. Carolina 
—in P) neither supports nor rejects application of the epithet to either of 
the above taxa. The specimen is apparently from Colleton County, which 
included at that time all of Charleston County southwest of the Ashley River. 
The specimen consists of three separate partly twining pieces, all sterile. 

one of its characters strongly favors either taxon other than the twining 
which is much more common in G. volwbilis. However, rather than creating 
a new name because of this possible discrepancy, I favor using G. glabella 
on the basis of the strong implications involved in Michaux’s describing the 
calyx as glabrous. His description most likely is based on a specimen from 
the “retrorse” population. 


176 


A detailed description of G. glabella (G. regularis, in the sense of previ- 
ous authors, minus G. szévor and the other “antrorse” populations) should 
be helpful and follows: 

Perennial herb. Stems prostrate, straight or sometimes twining, bearing 
appressed to rarely spreading—retrorse hairs 0.02-0.45 mm, or rarely a few 
scattered hairs to 0.7 mm long; some internodes, if not all, longer than the 
longest leaflet of the subtending nodes. Leaves compound, 28-92 mm long; 
leaflets 3, elliptic to narrowly elliptic to narrowly ovate to ovate, usually 
retuse, entire, largest per plant 7-20 mm wide and 20-45 mm long, thinly 
antrorse-appressed hairy beneath, glabrous or sometimes antrorsely scabrous 
above. Inflorescences axillary, longest 2-15 cm long, longer than the leaves, 
rarely shorter. Longest pedicels at flowering 2-5 mm long. Flowers 5-13 
per inflorescence, 12-18 mm long. Calyx 5.5-9 mm long, glabrous or with 
antrorse appressed hairs. Longest dehiscing anthers usually 0.88-1.20 mm 
long (1:17 is less than 0.88 mm long). Legume 35-52 mm long and 4.5- 
5.7 mm wide, bearing appressed antrorse hairs. Maximum number of ovules 
or seeds (including aborted ones) per specimen 6-9(10) per legume. 


Representative specimens include: VIRGINIA: Southampton Co.: Fernald and Long 
8744 (DUKE); NORTH CAROLINA: Pender Co.: Wilbur 4136 (FSU, GA); Scotland 
Fox and Godfrey 2412 (DUK es SOUTH CAROLINA: Orangeburg Co.: Ables 
31751 (NCU). GEORGIA: Chatham Co.: Duncan 21128 (NCU), Duncan 21229 
(GA). FLORIDA: Brevard Co.: Kral 5066 (FSU) 


\ 
a 
OQ 


I have no strong feelings as to how the “antrorse” population should be 
treated, as a separate species, as a variety of G. minor or G. glabella, or other- 
wise. It seems useful to others to point out that the Florida material and that 
from the coastal and adjacent counties into North Carolina have leaflets 
averaging broader than those of the more inland populations of NC, SC, and 
G he narrow aspect of the leaflets of this latter group is quite evident 
in comparison, width-length ratios being mostly around 1:3(4) as compared 
to mostly 1:2. I have other commitments which will keep me from attacking 
these and other problems involved for at least several years and so am leaving 
them for others who I know are interested in the problem. Until further 
studies can be made, it seems best to include these as forms of G. glabella, 
providing them with no names. 


Representative specimens of the antrorse forms include: NORTH CAROLINA: Bladen 
Co.: Rogers and Blomquist 3120 (DUKE). Moore Co.: Fox and Godfrey 2424 
(GA). Richmond Co.: Correll 1079 (DUKE). Scotland Co.: Ahbles 28568 (NCU). 
SOUTH CAROLINA: ee Co.: Radford 27337 (NCU). Kershaw Co.: hee 23549 
(NCU). Marion Co.: Bell 13677 (NCU). GEORGIA: Clinch Co.: Norris 1132 
Charlton Co.: Duncan A (GA). McIntosh Co.: Bozeman 2824 (NCU). Set DA: 
Lee C oldenke 951 (DUKE). Nassau Co.: Godfrey 64135a (FSU), Godfrey 
and Lindsey 56855 (FSU 

Useful synonomy for G. glabella: 

Galactia glabella Michaux. Fl. Bor. Am. 2 oe 1803. 

Galactia purshii Desv. Am. Sci. Nat. 9: 413. 1826. 


Ee 


177 


BOGE 
KB LQ A 


~pae 

we ONS 
s, 

LA 


n 
a 
ad 
ea 
2 
Oo 
WwW 
ox 


rsa} 


178 


— regularis (L.) Britt. Sensu Small, 1933. In part sensu: Fernald, 1950; 
Gleason & Cronquist, 1963; Wilbur, . Radford et al., 1964. 


G. VOLUBILIS (L.) Britt. 

As indicated earlier, G. regularis has been treated by recent authors as 
G. volubilis. These decisions were apparently based largely on the description 
by Linnaeus and an eies he pe (Dill. Elth. 173. t. 143. f. 170). 


Fernald (1950), ai Ibur (1963), and Radford et al. (1964). It is also 
quite doubtful which of these species the illustration represents. The leaf 
shape and relatively long pedicels are probably of the latter. The retrorse- 
spreading hairs on the stem and the antrorse-spreading hairs on the calyx 
are likely of G. regwlaris. However, I have seen each of these characters on 
specimens’ of G. macreei sensu above. Although the apex of the standard of 
the latter is rounded or nearly so and that of G. regularis is retuse, the illus- 
trations (Dill. Elth., above) of the flower are inconclusive as characteristics 
of both taxa are displayed. On these bases application of the epithet volabilis 
is clearly in doubt. Fortunately help is provided by a specimen in the Dillenius 
Herbarium labeled “Hedysarum trifoliatum scandens.” From 1:1  photo- 
copies of the specimen and other data provided by F. White, Curator of 
the Herbaria at Oxford University, Great Britain, the following diagnostic 
characteristics were derived: Hairs on stems retrorse-appressed, longest in- 
florescence 29 cm long with well separated nodes, longest pedicels at flowering 
ca. 4 mm long, longest flower 13 mm long, longest calyx ca. 6 mm long, 
the corolla light colored when dry. The specimen is clearly of G. macreei 
sensu the authors indicated earlier. On the basis of priority, however, G. 
volubilis (L.) Britt. should be used for the taxon. 

G. volubilis and G. glabella are similar in several ways and have been 
confused. Both twine, the former occasionally and the latter usually, but when 
the latter is growing in bare areas it cannot climb and is prostrate like the 
former. They may be separated as follows: 


Longest inflorescences 3-15 cm long, upper flower clusters congested; longest flowers 
12-18 mm long; longest dehiscing anthers usually 0.88—1.2 


less than 0.88 mm long); maximum number of — or seeds (including shared 
ones) per specimen 6-9, rarely 10(1:26) per legum G. glabella 
Longest inflorescences 5-55 cm long, all floriferous ane well separated: longest 
Sl 10-14 mm long ; longest diadas anthers 0.65—0.88 mm long; maximum 
mber of ovules or seeds (including aborted ones) per specimen usually 10-13, 

ee 9(1:43) per legume. ee owe . G. volubilis 


Numerous plants of G. volubilis have also been identified as G. regularis 
sensu Duncan and also the reverse. This can be prevented by checking the 
following differences: 


ee Co., SC; H. E. Ahles 15677; NCU 103886 + 103876.—Long Co., Ga.; 
J. R. Roseman, 2114; GA 94661. 


179 


Hairs on the stems usually spreading, to occasionally retrorse-spreading to uncommonly 
retrorse appressed; leaflets mostly elliptic to ovate; longest inflorescences usually 
3 : 


5—7, occasionally 4 or 8, or rarely 3 or 9? (1:133) per legume. regularis 
Hairs on the stems usually retrorse appressed to uncommonly retrorse- redid 
leaflets narrowly pe to narrowly ovate or approaching the above; longest in- 
florescences 5-45 cm long, longest pedicels at flowering 3.0-4.0 mm long; longest 
calyx (5.5) 6-10 mm long; longest flowers 10-14 mm long; longest dehiscing 
anthers 0.65—-0.88 mm ue hairs on legume antrorse-appressed or rarely antrorse- 
spreading; maximum number of ovules or seeds (including aborted soar per speci- 
men usually 10-12, teens 13, or a 9? (1:43) per legume . volubilts 
Partial synonomy for G. vol#bilis follow 
oe volubilis (L.) Britt., Mem. Tor rr. Club 5: 208. 1894. Sensu: Small 
33) in part, Gleason & Cronauist eae in part. 


Cina pilosa Nutt. var. macreei Carey T. & G. Fl. N. Am. 1: ae 1838. 

Galactia pilosa Nutt. var. oa sg T. & G. Fl. N. Am. 1: 287. 1838. 

Galactia macreet M. A. Curtis, Bost. Jour. Nat. Hist. 1: 120. 1837. Sensu Fernald 
(1950), Wilbur (1963), Radiord et al. (1964). 

Galactia volubilis (L.) Britt. var. intermedia Vail, Bull. Torr. Bot. Club 22: 
08. 1895. 


DISTRIBU TION 


During this study distribution maps were prepared for Galactia minor, 
the retrorse form of glabella, the antrorse forms of glabella, regularis, 
volubilis, mollis, and floridana. The specimens were mostly from DUKE, 
FSU, GA, NCU, VDB, and VSC. A few were from FLAS, GH, NY, and US. 
These distribution data were essential during my study, and are useful in 
understanding my interpretations of these taxa. They are presented here for 
the Southeastern United States. 

Appreciation is expressed to my colleague, Dr. Samuel B. Jones, for sug- 
gestions involving the study; to Dr. G. Taylor, Keeper of Botany, of the 
British Museum, London, who many years ago provided details about the 
type specimen of Galactia regularis, and to F, White, Curator of the Herbaria 
at Oxford University, Great Britain. The loan from Duke University Library 
of the Ph.D. dissertation, “The genus Galactia in the United States,” by 
Hollis J. Rogers and the loans of specimens through the courtesy of curators 
of the several herbaria are appreciated. 


REFERENCES 
DUNCAN, W. H. 1977. A new species of Galactia (Fabaceae) in the southeastern 
United States. Phyilosia 37: 59-61. 


? The only specimen with 9 seeds is from Yell Co., Ark. 
3 The only specimen with 9 seeds is from Iberia Parsich. La. 


180 


FERNALD, M. L. 1960. Gray’s manual of botany. 8th ed. American Book Co. N. Y. 

1632 
GLEASON, H. A. and A. CRONQUEST. 1963. Manual of vascular plants of north- 
oa 


eae United States and adjacent Canada. D. Van Nostrand Co. Princeton, 


ote: A. E., H. E. AHLES, and C. R. BELL. 1964. Manual of the vascular 
flora of the Carolinas. Univ. of North Carolina Press. Chapel Hill. 1182 p. 
LL, J. K. 1933. Manual of the southeastern Flora. Univ. of North Carolina 
Press. Chapel Hill. 1554 p 
WILBUR, R. L. 1963. The leguminous plants of North Carolina. Tech. Bull. No. 
151. North Carolina Agr. Exp. Sta. Raleigh. 294 p. 


A NEW SPECIES OF LINUM FROM 
SOUTHERN TEXAS AND ADJACENT MEXICO 
C. M. ROGERS 
Wayne State University, Detroit, MI 48202 


In an earlier publication (Rogers, 1968) one of the yellow-flowered 
flaxes, Linum rigidum Pursh, was described as comprised of four varieties. 
Of these the typical variety and var. compactum are plants of northern Texas 
and northward. The var. berlandieri, with some reason sometimes considere 
a separate species, is distributed nearly throughout Texas. It is rather readily 
distinguished from the other varieties by its comparatively thick-walled 
fruit and coarse sepals. These three varieties, although displaying some 
variation which may require further study, are reasonably well defined. The 
fourth variety, var. filifolimm Shinners, however, was described as a “variable 
population .. . more study is necessary to determine the relationship of the 
western Texas plants to those of southern Texas, as well as some anomalous 
collections included here from northern Mexico.” In order to help clarify 
these relationships, the author has collected additional material of var. 
flifoliwm and attempted a number of crosses between plants from different 
parts of its range. Herbarium material from the New York Botanical Garden, 
Southern Methodist University and the University of Texas, including the 
Lundell Herbarium, has also been reexamined and thanks are expressed to 
the respective curators. 

Within this “variable population,” in addition to Linum rigidum var 
filifolium, which, as now interpreted, is a plant of western Texas and nearby 
Coahuila, two taxa, mostly of southern Texas and adjacent Tamaulipas and 
Nuevo Leon, can be distinguished. These are L. elongatwm (Small) Winkler, 
a species proposed many years ago (Small, 1907), and previously regarded 
as a synonym (Rogers, 1968), and L. lwndellii, a previously undescribed 
species. The latter is named for C. L. Lundell, long a student of the Texas 
flora and co-collector of the type specimen. 

The three taxa may be distinguished from one another in the following 
way: 


1. Styles 7-8 mm long; petals mostly ca. 15 mm 
an lanceolate, acute; petals diffusely a - at the base; annual... L. 
ea Cie var. Alo 
2. Sepals lanceatenaat — ee reddish to wine-colored band below the 
aidale mostly perennial . L. elongatum. : 
. Styles 3-4 mm ie pe aie ca. 10 mm long... . L. landellii 


In the following descriptions, petal and anther coloration is taken from 


SIDA 8(2): 181-187. 1979. 


182 


a limited number of living plants, supplemented by scattered information 
from herbarium labels. 


LINUM RIGIDUM Pursh var. FILIFOLIUM Shinners in McVaugh, Field & 
Lab. 17: 136. 1949. Type: McVaugh 7798, 4 Apr 1947, near abandoned wax 
factory on Rio Grande at mouth of Big Canyon, Brewster Co., Tex. (Holo- 
type: SMU; isotype: TEX). 

More or less stiffly branched, essentially glabrous annual herb, 15-35 cm 
tall; leaves alternate, linear, 10-30 mm long, 0.5-2.0 mm wide; stipular glands 
mostly present (80% of the specimens examined have stipular glands, but 
see the discussion which follows the species descriptions); sepals lanceolate, 
sharply acute, 6-9 mm long, grayish; petals narrowly to broadly obovate, 
13-17 mm long, yellow to orange or salmon, brick-red near the base, not 
banded (Fig. 4a); stamens 5-6 mm long; anthers 1.5—2.5 mm long, yellow- 
ish; styles 6.5-8.0 mm long; stigmas dark-wine to black; fruit ovate, ca. 
3.5-4.0 mm high, 3.0 mm diameter; seeds reddish-brown, ca. 3.0 mm long, 
1.25 mm wide; chromosome no. 2 = 15 (chromosome voucher: Rogers 
13494, 29 Aug 1976, 4 mi SE of Del Rio, Val Verde Co., Tex. WUD 
About 70 collections have been examined. The following citations include 
the Mexican collections and a specimen from each of the counties in Texas. 


MEXICO. Coahuila: ca. 100 mi NW of Muzquiz, 12 May 1968, Latorre s.n. (TEX 
Santa Rosa Mts., a Jul 1938, Marsh 1386 (TEX); S of Sabinas, 29 Mar i 
Rinehart 289 (WU 

exas. a ee Co.: 23 mi on Eunice Highway, 8 es 1960, Scuddy s.n. 
(nea rest cs australe but apparently with some features of var. filifolinm) (LL). 
Brewster Co.: 26 mi E of Marathon, 23 Aug 1947, Warnock “6719 (TEX). Crane 
Co.: ca. 10 mi NW of Crane, 9 May 1966, Correll 32773 (appears to combine 
traits of var. filifolinm and var. rigidum) (LL). Culberson Co.: 51 mi W. Bal- 
morrhea, 29 May 1967, Mears 15654 (TEX). Jeff Davis Co.: SW éf Mt. Livermore, 
Davis Mts., 15 Jun 1926, Demaree & Palmer 212 (SMU). Kinney Co.: 
of Del Rio, 29 Aug 1976, Rogers 13493 (WUD). Maverick Co.: 6 mi N of Que- 
mado, 4 Apr 1959, Correll 20727 (LL). oa Co.: 10 og S of Ft. Stockton, | a 
1948, Warnock 7691 (LL). Presidio Co.: ca. 11 mi of Valentine, 25 Jun 1948, 
York 48185 (TEX). Terrell Co.: 3 mi W of sre 30 Aug 1976, Rogers 13498 

D). Val Verde Co.: near Del Rio, 30 Mar 1963, Correll 27114 (LL). 


distribution is shown in Fig. 1 


LINUM ELONGATUM (Small) Winkler, Nat. Pflanzenfam, ed. 2, 19a: 116. 
1931. Type: Reverchon 3776, 21 Mar 1903, Laredo (Webb Co.), Tex. 
(Holotype: NY 


Cathartolinum elongatum Small, North Amer. Flora 25: 82. 1907. 


More or less diffusely branched, annual (or merely flowering first year? ) 
or perennial, essentially glabrous herb, 15—30 cm tall; leaves alternate, linear, 
5-25 mm long, 0.5-1.0 mm wide; stipular glands present throughout or at 
bases of upper leaves only; sepals mostly lance-attenuate, 6-11 mm long; 
petals broadly obovate, 14-18 mm long, yellow-orange to salmon or brown- 


183 


° r 


1 L. rigidum 


var. filifolium . a 


@ 
> 
@ 
@ 
e > @ 
®e e* 
Wy ° 
@ 
op doe ee elongatum : 3 L. lundellii 


Figs. 1-3. Distributions of L. rigidum var. filifolium, L. elongatum, and L. lundellii. 


184 


ish-red, with a prominent wine-colored band below the middle (Fig. 4b); 
stamens 5-6 mm long; anthers 1.5-2.5 mm long, brick-red; styles 7.0-9.5 
mm long; stigmas mostly grayish to wine-colored; fruit ovate, ca. 4 mm 
high, 3 mm in diameter; seeds reddish-brown, ca 3 mm long, 1.2-1.3 mm 
wide; chromosome no, 7 = 15 (chromosome vouchers: Rogers 13475, 27 
Aug 1976, farm road 2895, 5 mi N of jct. with Tex. 359, Webb Co., Tex.; 
Rogers 13482, 28 Aug 1976, 1 mi SE of San Ygnacio, Zapata Co., Tex., 
both WUD). About 65 collections have been examined. The following 
citations include the Mexican collections and a specimen from each of the 
counties in Texas. 


on 


EXICO. Tamaulipas: 7 mi S of Nuevo Laredo, 12 oa 1964, Fields 45 (TEX); 
16 mi S of Nuevo Laredo, 7 Mar 1962, Garza 35 (TEX); 14 mi S of Nuevo Laredo, 
24 Mar 1944, Heard & Barkley 14604 (TEX); 10 mi SE of Nuevo Laredo, 8 2 
1964, Ibarra 103 (TEX); 12 mi S of Nuevo Laredo, 26 Jun 1963, Rogers 1284. 
(WUD); 2 mi S of Nuevo Laredo, 26 Jun 1963, Hagens 12846 (WUD); 7 mi d 
of Nuevo Laredo, 28 Aug 1976, Rogers 13476 (WUD); Highway 2, 5 mi E of jet. 
with a 85, S of eee Laredo, 28 Aug 1976, Rozen: 13477 (WUD); 4 mi 
f Nuevo Laredo, 14 Nov 1958, Rollins & Tryon 5802 (LL); Arroyo Coyote, 
7 km. Carretra Nacional, 1 Apr 1964, Vazguiz 30 (TEX). 
U Texas. Cameron Co.: US. 83, 12 mi N of Prowasyls 18 Apr 1965, 
Cavazos 262 (LL). Dimmit Co.: 5 mi N of Carrizo Springs, 16 Mar 1963, 
fue SO (TEX). Duval Co.: Realitos, 17 Mar 1934, oe 781 (NY). 
Jim Hogg Co.: 8 mi S of Hebbronville, 20 Mar 1969. ules 36801 (LL). Jim 
Wells Co.: 6 mi S of Alice, 20 Mar 1952, Jones 699 (SMU). LaSalle Co.: near 
Encinal, 27 Aug 1976, Rogers 13470 (WUD). Webb Co.: oe 11 Mar 1944, 
Crockett 5417 (LL). Zapata Co.: 13 mi N of oa Ygnacio, 31 Jan 1954, Shinners 
17656 (SMU). The distribution is shown in Fig. 


Bes 


LINUM lundellii Rogers, sp. nov. L. elongato affine, sed semper annuum, 
glandulis stipularibus foliorum oe nullis, petalis circa LO mm gen 
et stylis 3-4 mm longis. Type: C. L. & A. A. Lundell 9894, 2 Apr 1941, « 
gravelly hill, off U.S. 83, W of Sullivan City, Starr Co., Tex. (Holotype: tL . 

nnual, essentially glabrous herb, 10-40 cm tall; leaves alternate, linear, 
5-30 mm long, 0.5-1.5 mm wide; stipular glands absent above, moderately 
developed below; sepals linear-lanceolate to lanceolate, acute to acuminate, 
4-12 mm long; petals obcordate, 7-12 mm long, yellow to orange-salmon, 
faintly banded near the base (Fig. 4c); stamens 4-5 mm long; anthers 1.0- 
1.5 mm long; styles 3.0-4.0 mm long; stigmas mostly dark-wine; fruit ovate, 
3.3-4.0 mm high, 2.6-3.1 mm in diameter; seeds reddish brown, 2.5-2.7 
mm long, ca. 1.1 mm wide; chromosome no. 7 = 15 (chromosome vouchers: 
Rogers 13137, 2 Jul 1964, 2 mi N of Roma, Starr Co., Tex.; Rogers 13471, 
27 Aug 1976, 5 mi E of Laredo, Webb Co., Tex., both WUD). In addition 
to the type and the specimens cited for cnomescme number, the following 
collections have been examined. 

EXICO, Nuevo Leon: ca. 15 mi SW of Galeana, 19 May 1934, Mueller and 


M 
Mueller 470 (TEX): 4 mi S of China, 6 Aug 1964, Rogers 13189 (WUD). 
Tamaulipas: 5 mi SW of Reynosa, 29 Feb 1944, Painter & Barkley 14412 (LL); ca 


185 


mi SW of Reynosa, 6 Aug 1964, Rogers 13190 (WUD); 2 mi W of Nuevo 
eat 28 Aug 1976, Rogers 13478 (WUD). 

Texas. Dimmit Co.: 2 mi N of Carrizo a 3 Jul 1964, Rogers 13142 
(WUD); 3-4 mi NW of Carrizo ia 29 Aug 1976, Rogers 13486 (WUD); 
= mi NW of Carrizo Springs, 29 Aug 1976, Roger 13488 (WUD). Hidalgo Co.: 

mi W of LaJoya, 28 Aug 1976, Rowers 134 (WUD); La Joya, 9 Feb 1942, 
o alker 48 (TEX). Jim Hog Co.: Arroyo ee eae 15 Mar 1964, 
Correll 29000 (LL). Starr Co.: ca. 1.5 mi N of Roma, 17 Mar 1966, Correll 32270 
Glee): pees Falcon Dam, 17 Mar os Ramos et al 7864 Sas X); near Falcon 

Dam, 2 July 1964, Rogers 13138 (WUD); U.S. Highway 83, 3 mi SE of pape 
with aes road 2 098, 28 Aug 1976, sles 13480 (WUD); 3 mi N of Roma, 31 
Jan 1954, Shinners 17702 (SMU); Falcon State Park, 14 Mar 1968, Wood 83 8 
(TEX). The distribution is shown in Fig 


DISCUSSION 


Linum lundelli, except for vegetative features such as stature, amount of 
branching, and leaf size, is a very uniform species and may be readily dis- 
tinguished from L. elongatunz, the range of which overlaps that of L. landellii, 
as well as from L. rigidum var. filifolium by style length, which is well 
correlated with other differences such as petal size and shape, anther size, 
and the absence of stipular glands at the bases of upper leaves. Its distinctive- 
ness is supported by the results of attempted crosses with L. elongatum (128 
crosses) and with L. rigidum var. filifolinm: (164 crosses), none of which 
produced functional seeds. 

In L. landellii the anthers are placed at about the same level as the stigmas 
(Fig. 4c). In garden-grown plants, by the time the flowers were well opened, 
pollen had already been deposited upon the stigmas. In these plants. self- 
pollination was certainly the rule. In L. elongatum and L. rigidum var. 
fulifolium the styles extend well beyond the anthers (Fig. 4a,b), and there 
are ordinarily a few hours (depending upon cloudiness and wind condi- 
tions) during which the anthers have dehisced, but pollen has not been 

shed. In the Michigan garden no insect pollinators were observed; these may 
be present in the natural range of the species. There is at least the oppor- 
tunity for outcrossing. If cross-pollination fails, however, self-pollination ts 
assured, since the intact corolla, before falling, ordinarily slips upward past 
the open anthers like an ascending collar, depositing pollen on the stigmas 
above. 

Linum elongatum and L. rigidum var. filifolium appear to be closely re- 
lated. It may be that some hybridization takes place between the two in the 
field. The results of experimental crosses between the two reflect this possi- 
bility; one or more rather weakly developed but mature seeds were pro- 
duced in 14 of 136 crosses attempted. Of a series of features that differentiate 
the two, possibly none holds for every comparison of individual specimens. 
For example, most collections of L. elongatum are perennial. This is a very 
unusual character in the “L rigidum complex” of about twelve species. Linum 
subteres (Trel.) Winkler, of Utah and Nevada, is the only other which ts 


tal, stamens and a of L. rigidum var. filifolium (a), L. elongatum 
Pas a : lundellii (c). All x 2.5. 


consistently perennial. However, L. elongatum flowers the first year, and a 
number of herbarium specimens appear to be or are annual. In addition, 
two collections of L. rigidum var. fulifolium are quite clearly perennial. 

The petal coloration of L. elongatum makes this one of the most attractive 
of the flax species. The author's collections and the several dozen plants 
grown from their seed were uniformly colored in the manner described. 
Dried specimens, because of the fading of the pigments and because petals 
have frequently fallen if collections are made late in the day, do not always 
show petal color. In garden-grown plants, one individual (of about 40 
plants), which otherwise had the characters of (and was identified as) 
L. rigidum var. filifolium, had the petal coloration of L. elongatum. Other- 
wise the two taxa were easily distinguished on this character alone. The 
attenuate sepals, which were the basis for the selection of the specific epithet 
for L. elongatum, can be used for the identification of most specimens, but 
there is enough variation in both taxa that some specimens would be difficult 
to place using that character alone. 

Linum elongatum uniformly possesses stipular glands at the bases of the 
upper leaves, L. /wndellii uniformly lacks them there, although they may be 
moderately developed at the bases of the lower leaves. Most collections of 
L. rigidum var. filifolium have stipular glands also, but with the exception of 
a single collection from near Del Rio, none of the specimens from Val Verde, 
Kinney, or Maverick Counties have stipular glands. This may provide another 
distinguishing feature between L. rigidum var. filifolium and L. elongatum, 
if one is needed, in the region of closest geographical proximity of the two 
taxa. Baesd upon the specimens thus far examined, the range of L. rigidum 

var. filifolium is distinct from those of either L. elongatum or L. lundellii, 

being restricted in Texas to northwestern Maverick Co. and northwestward, 
while L. elongatum and L. lundellii are found only from Dimmit County 
southeastward. 


187 


REFERENCES 


ROGERS, C. M., 1968. Yellow-flowered ae of Linum in Central America and 
western North America. Brittonia 20: 107— 
SMALL, J. K., 1907. Linaceae. In: North ae Flora 25: 67-87. 


REDISCOVERY OF 
SPIRANTHES PARKSIT CORRELL 
P. M. CATLING AND K. L. McINTOSH 


Department of Botany, University of Toronto 
Toronto, Ontario, MSS 1Al1 


Spiranthes oS Correll was described in 1947 on the basis of specimens 
collected by H. B. Parks along the Navasota River (Democrat Bridge) in 
Brazos County, Texas (Correll, 1947). It appears that the late Dr. Parks was 
the only person to have seen a living plant. Correll (1950) and Correll and 
Johnston (1970) reported that this species was endemic to Brazos County 
but recent attempts to find it there have been unsuccessful (Luer, 1975; 

. Correll and M. C. Johnston, pers. comm.). Spiranthes parksii is the 
only North American orchid not illustrated with a photograph in Luer's 
(1975) recent work on the North American orchids. It is one of the 20 
species listed by Ayensu (1975) in an attempt to identify those North 
American orchids in serious danger of extinction. Spiranthes parksii is not 
only one of the rarest North American orchids, it is also one of the least 
well known. 

While collecting data from the AMES orchid herbarium (Harvard Unt- 
versity) in 1975, I noticed a sheet labelled “S. cernwa, Hy College—Navasota, 
10 miles west of Navasota R. bridge on hwy no. 6, H. B. Parks, 27 Oct 1945” 
(AMES 63043). This sheet had 12 plants mounted on it, some referable 
to S. cernua, others to S. parksii. The type of S. parksii (AMES 63039) was 
available for comparison. 

On 25 Oct 1978, we had an opportunity to explore the Post Oak Savanna 
northwest of Navasota. Local residents reported that it had been a dry sum- 
mer and perhaps for this reason Spiranthes spp. were not easily found. 
S. cernua (L.) L. C. Rich. occurred sparingly along margins of ponds and 
streams. Both the normal white open-flowered plants and more or less peloric, 
yellow closed-flowered plants were seen. A few S. lacera (Raf.) Raf. var. 
gracilis (Bigel.) Luer were found on dry sandy banks, all past anthesis. 

Exploring the open banks of a temporary stream surrounded by scattered 
oaks (Quercus stellata, Quercus marilandica) and Beauty-berry (Callicarpa 
americana), several Spiranthes orchids were found among the open cover 0 
grasses and forbs (Fig. 1). Some of these were closed-flowered S. cernua 
but others scattered among them had open greenish-white flowers with 
relatively short rounded petals and an ovate, distally truncate lip. The floral 
bracts of these plants also differed in having distinctly whitened tips. 
close examination and dissection of a few flowers indicated that these plants 


SIDA 8(2): 188-193. 1979. 


189 


Fig. 1. Open grassy meadow habitat of S. parksii along a temporary stream. 
Dominant plants include Schizachyrinm scoparinm var. freguens, Sporobolus juncens, 
and Eupatorium compositifolium,. Photographed 25 Oct 1978, ca. 19 km NW of 
Navasota, Brazos Co., Texas. 


190 


were beyond any doubt referable to S. parksii, the description and original 

illustrations of which we had with 

he dominant plant associates here included Schizachyrium scoparinm 
var. frequens, Panicum brachyanthum, Aristida longispica, Sporobolus junceus, 
Expatorium compositifolium, and Linum medium var. texanum. Chrysopsis 
pilosa and Ascyrum hypericoides var. hypericoides were also present. 

A few hundred yards away we found more plants of S. parksii on the 
banks of another temporary stream (Fig. 2). The aa here was again 
an open oak woodland (Q. marilandica, Q. stellata, Q. nigra) but with 
scattered thickets of Forestiera ligustrina, Callicarpa americana and Ilex 
vomitoria. Spiranthes parksii occurred both on the tops of the banks in open 
sand with a sparse cover of grasses, and on the sides of banks sometimes in 
the shade of thickets. Dominant associates here included Andropogon 
ternarins, Andropogon virginicus, Aristida longispica, Mublenbergia capillaris, 
Linum medium var. texanum, Eupatorinm compositifolium, and Smilax 
bona-nox var. hastata. Paspalum setaceum var. stramineum, Ascyrum byperi- 
coides var. hypericoides and Drosera annua were also recorded as associates 
at this location. 

In all, seven plants were found at the first locality and 13 at the second. 
Soil collected from about the roots of several plants was found to have a 
pH range of 4.0-5.8, with most readings between 4.5 and 4.9. 

Since the flower color of S parksii has not been described, various floral 
parts were compared with the Royal Horticultural Society Colour Chart 
(1966). The general flower color (7.e. the conspicuous perianth parts) varies 
from white (155A) to light yellow-green (154D). The dark green color 
of the ovary (144A-B) extends onto the basal perianth parts for 0.5—2.0 
mm. Lateral petals vary from whitish (155A) to yellow-green, (144B-C, 
154C). The midvein region is slightly greenish (145C) in white lateral 
petals and areal green (144A) in greenish lateral petals. The central por- 
tion of the lip varies from yellow (2D) to yellow-white (158B—C) and 
yellow-green ea 151C 154C). The calli are white (155A-D). The stig- 
matic surface is green (144B, C) but the proximal stalk of the column is 
white (15SA-D) and the pollen masses are yellow (9B) to light yellow- 
orange (14C). The darkest green color in the flowers occurs in the basal 
perianth parts, in the lateral petals and on the stigmatic surface. The remain- 
der of the plant is green (144A) except for the floral bracts which are 
white 1.5—3.0 mm from the tip. 

Although no insects were observed pollinating the flowers, the pollinia 
are easily detached, and the stigmatic surface is viscid, the pollen readily 
adhering to it. A drop of glucose-rich liquid is secreted near the base of 
the lip. These characteristics suggest insect pollination, but the ovaries can 
enlarge and develop seed without pollination. Brown and withered flowers 
on several plants had the pollen masses intact and the stigmatic surface had 
not received any pollen. The ovaries of these withered flowers contained 


Fig. 2. Habitat of S. parksit (lower center). Several plants were growing on the 
high banks of a temporary stream in the open with Andropogon ternarius, Andro pou 
VIN SINICHS, Mud aber capillaris, and others (see text). The surrounding trees 
Ouercus marilandica, and eee stellata. Photographed 25 Oct 1978, ca. 19 km 

W of Navasota, Brazos Co., Texa 


Fig. 3. Flowers of S. parksii showing relatively short lateral petals and white-tipped 


red 25 Oct 1978, ca. 19 km NW of Navasota, Brazos Co., 


floral bracts. Photograp 
Texas 


Oe) 


immature seeds of which 80-90% were polyembryonic. The occurrence of 
polyembryony in Spiranthes is associated with adventitious embryony 
(Swamy, 1948; Catling, unpublished data). 

With its rounded or oval lateral petals (4.5-6.7 mm long) much shorter 
than the sepals (5.0-8.0 mm long), and a distally truncate, erose-margined 
lip (5.0-7.0 mm long), S. parksii appears to be a very distinctive species. 
The loosely flowered spike and absence of leaves at flowering time also hel 
to separate S. parksii from other sympatric Spiranthes spp. These features 
are well illustrated in the drawings provided by Correll (1947, 1950) and in 
the accompanying photograph (Fig. 3). The green marking along the center 
of the lateral petals and the white-tipped floral bracts are helpful in field 
identification. S. parksii keys out readily in keys provided by Correll (1950) 
and Correll and Johnston (1970). 

Correll (1947) thought that S. parksii had no close allies in our flora, 
its affinity being with several Mexican and Central American species. Its 
taxonomic status and evolutionary relationships are indeed a matter of great 
interest. 

In view of the general rarity of the plant, ic seems desirable to document 
any future discoveries with photographs, measurements and detailed fie!'d 
notes rather than a large series of collected specimens. 

We thank D. S. Correll and M. C. Johnston for commenting on the 
manuscript. 


REFERENCES 


AYENSU, E. S. 1975. ee and threatened orchids of the United States. Amer. 
Orchid Soc. Bull. 44(5): 94, 

CORRELL, D. S. 1947. A new cS ranibes from Texas. Amer. Orchid Soc. Bull. 16: 
400 

1950. Native orchids of — America north of Mexico. Chronica 
Botsaies Co., Waltham, Mass. 399 
-and M. C. Johnston. 1970. von of the vascular plants of Texas. Texas 
Research Foundation Renner, Texas. 1881 pp. 

LUER, C. A. 1975. The native orchids of the rae States and Canada excluding 
Florida. New York Botanical Garden, New Y pp. 

ROYAL HORTICULTURAL SOCIETY. 1966. Colour chart (202 colours each with 
4 tints, printed in solid colour). Royal Horticultural Society, Vincent Square. West- 
minister, S.W.1, England. 

SWAMY, B. G. L. 1948. Agamospermy in Spiranthes cernua. Lloydia 11(3): 149- 
162 


CLEMATIS PITCHERI T. & G. VAR. DICTYOT'A 
(GREENE) DENNIS, COMB. NOV. 
(RANUNCULACEAE).' 


W. MICHAEL DENNIS’ 


Department of Botany, University of Tennessee 
Knoxville, Tennessee 37916 


Clematis pitcheri 'T. & G. is a highly variable, wide-ranging species dis- 
tributed on a northeast to southwest clinal axis extending from western 
Indiana, Illinois, and eastern Iowa south to Hidalgo and Queretaro, Mexico 
(Dennis, 1976). It is distinguished from other closely related taxa by its 
non-plumose achene tails. In Gray's Synoptical Flora of North America 
(1895), Robinson recognized four varieties within the C pitcheri complex. 
More recently Erickson (1943) recognized only two varieties: C. pitcheri 

. & G. var. pitcheri, which includes those plants with pinnate leaves having 
leaflets 4-10 cm long and ovate sepals, and C. pitcheri var. filifera (Benth. ) 
Robinson, which is distinguished from the type variety by having pinnate- 
ternate leaves having leaflets less than 4 cm long and lanceolate sepals. 
Clematis pitcheri var. pitcheri was ascribed a distribution from Indiana to 
eastern Nebraska and south to Texas and var. filifera was reported to occur 
along banks of streams and in canyons of western Texas, New Mexico and 
Mexico. 

In a recent biosystematic study of Clematis subsection Viornae (Dennis, 
1976) it was concluded that Erickson’s (1943) recognition of two varieties 
of C. pitcheri on the basis described above is taxonomically unsound. These 
characters (leaf size, degree of leaf division and flower shape) vary con- 
siderably within all members of the subsection suggesting polymorphism 
and ecotypic variation rather than variation indicative of speciation. How- 
ever, within C. pitcheri var. filifera sensu Erickson there is a morphologically 
recognizable element that was treated by Erickson (1943) as a synonym of 
var. filifera. This element is C. dictyota Greene, which is herein recognized 
as a variety of C. pitchert. 


CLEMATIS PITCHERI var. dietyota (Greene) ge stat. et comb. nov. Basionym: 
Clematis dictyota Greene, Pittonia 5:133. 1903. E: US, Texas, caiee Canyon, 
at Apr 1902, Tracy ey Earle 256 (Holotype, oo as photo MO!; Isotypes, TEX! 

US!). [Viorna dictyota (Greene) Heller.] Leaflets generally as pe and 
more divided than var. pitchert. Sepals light to brownish purple without, tips recurved 

‘Contribution from the Botanical Laboratory, The University of Tennessee N.S. 

12 


2 Present address: Tennessee Valley Authority, EDB, Muscle Shoals, AL 35660. 


SIDA 8(2): 194-195, 1979. 


195 


and slightly ae eee and connectives (including extended apex) essen- 
tially glabrous or occasionally w a few erect trichomes just below the anthers or 
along the connective or its oe apex. 


Clematis pitcheri var. dictyota is restricted to soil accumulations among 
boulders, crevices of rock formations, and stream banks of the Trans Pecos 
region of western Texas and adjacent New Mexico and Mexico. It is dis- 
tinguished from var. pitcheri by its glabrous filament and anther connectives. 
Erickson (1943) did not note this character and referred all small leaved 
plants of Texas and Mexico with both glabrous and pubescent filaments and 
connectives to C. pitcheri var. filifera. Correll and Johnston (1970) used 
pubescence on filaments as a key character to distinguish west Texas popula- 
tions as C. filifera Benth. However, examination of the type specimen of 
C. filifera Benth. [Mexico. Prope Leon, 1839. Hartweg 1590 (Holotype, K!; 
Isotype, LD! )} revealed that its filaments were pubescent. Since the most 
consistent taxonomic character distinguishing the Trans Pecos populations 
is the absence of trichomes on the filaments, the name C. filifera cannot be 
applied to these plants. Review of previously published names and_ study 
of type specimens indicate that C. dictyota Greene is the appropriate element 
upon which to base a taxon representative of the Trans Pecos populations. 
It was described from the region and its type specimen has glabrous fila- 
ments. Varietal status for the plants of the Trans Pecos region based on the 
name C. dictyota is therefore proposed. All small-leaved plants in this com- 
plex occurring in New Mexico, Mexico and Texas that have non-plumose 
achene tails and pubescent filaments, (ie. C. filifera Benth. sensu stricto ) 
are referred to C. pitcheri var. pitcheri. 

REFERENCES 
CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the vascular plants of 

Texas. Texas Research Foundation, Renner, Texas. 1881 
DENNIS, W. M. 1976. A a las study of Copa: section Viorna subsection 

Viornae. Ph.D. dissertation, Unive of Tennessee. 176 p. 

ERICKSON, R. O. 1943. eee Clematis section Viorna. Ann. Missouri Bot. 


GRAY wn "1895. Ranunculaceae. In B. L. Robinson 1895-1897. Synoptical 
flora of North America, Vol. 1, pt. 1. American Book , New York. 


FIELD NOTES ON NIGERIAN STRIGA 
(SCROPHULARIACEAE) 


LYTTON J. MUSSELMAN, DANIEL L. NICKRENT 


Department of Biological Sciences, Old Dominion University 
Norfolk, Virginia 23508 


RICHARD A. MANSFIELD, JAMES E. A. OGBORN 


Weed Science Section, Institute for Agricultural Research 
Ahmadu Bello University, Zaria, Nigeria 


Striga is a palaeotropical genus of perhaps as many as 50 species, although 
the genus has not been monographed. Like many Scrophulariaceae, Striga 
is root parasitic. In fact, several species are of great economic importance, 
particularly in the semi-arid tropics. In the rain fed areas of subsaharan 
Africa S. hermonthica (Del.) Benth. is an ubiquitous weed in guinea corn 
(Sorghum bicolor) fields. In some areas it is the single most serious para- 
site of this main crop of subsistence farmers. Millet (Pennisetum typhoides ) 
may also be attacked by this species. Striga gesnerioides causes — to 
a variety of crops, including legumes and tobacco. Witchweed (8. aszatica) 
is the most widespread species in the genus, extending across Africa, the 
Indian subcontinent, and into Indonesia. It parasitizes corn, rice, sugar cane, 
and other members of the Poaceae. Striga angustifolia (Don.) Saldanha 
(= Striga euphasioides Benth.) is an important parasite on sorghum in 
India. These and a few other species are briefly discussed in Hosmani (1978). 

Two species have been accidentally introduced to the New World. In 
the 1950s S. asiatica (L.) Kuntze (= S. lutea Lour.) was discovered in the 
Carolinas. It is presently known from several counties in southeastern North 
Carolina and adjacent South Carolina. It is now the object of a federal 
quarantine and iota program. Very recently S. gesnerioides ( Willd.) 
Vatke was discovered in Polk Co., Florida (Wunderlin, Musselman and 
Shuey, 1979). The extent of this infestation is being investigated. 

The most useful aid to identification of African material is found in 
Hepper (1963). Characters of diagnostic value in separating species include 
number of ribs in the calyx, indumentum, relative size of calyx and of corolla 
tube, size of the bract subtending each flower, and flower color. Like several 
other genera of parasitic angiosperms that exhibit reduction of vegetative 
parts, floral characters are the main basis of taxonomic differences in Striga. 
Plants are often confused in the field, however, particularly S. aspera and 
S. hermonthica. 


Despite its agronomic importance, very little information of a descriptive 


SIDA 8(2): 196-201. 1979. 


OF 


nature is available for the genus, especially for those species that are not 
serious pathogens. The purpose of this paper is to record field observations 
on several species of Striga observed in Nigera in October and November 
O78: 

Specimens were collected from Kazaure in northern Nigeria near the 
border with Niger and southeast along the main road south to Mokwa. 
This transect crosses several rainfall isohyets from 750 mm/year in the north 
to 1500 mm/year near Mokwa and corresponds approximately to the vege- 
tational zones known as Sudan Savanna, Northern Guinea Savanna, and 
Southern Guinea Savanna. 

All species of Striga are considered noxious weeds in the United States 
and are under a federal plant quarantine. Therefore, all dried specimens 
from Nigeria were devitalized by treatment with ethylene oxide at 25 
Ibs/1000 cu. ft for 2 hours at 70°F or above by the Animal and Plant 
Health Inspection Service, Plant Protection and Quarantine Programs. These 
specimens are deposited in the Old Dominion University Herbarium (ODU). 
Drawings were prepared from FAA preserved material and from Koda- 
chromes. Silhouettes are from actual specimens. 


~ 


STRIGA HERMONTHICA (Del. ) Benth. 

This is a strikingly beautiful species with large pink flowers. Of the six 
species examined, only 8. bermonthica has more than two corollas open per 
inflorescence branch and flowers that last more than one day (Fig. 1). 
Widespread throughout the drier regions of Africa, it is quite variable. A 
cursory examination of populations parasitizing sorghum at Musasa and 
Zaria (Both Kaduna State, Nigeria) revealed the following corolla variations: 
upper lobe entire to almost bifid, orifice of throat white or pink, and lower 
lobes deeply toothed to entire. The extent of this variation among popula- 
tions remains to be determined as does its source. Of greater agronomic 
importance is the physiological variation within the species. Preliminary 
work by C. Parker (personal communication ) of the Weed Research Organi- 
zation indicates that S. hermonthica may exist as two strains—one attacking 
millet, the other sorghum. In the present study “millet” S. bermonthica was 
examined at Mokwa and appeared morphologically identical to that growing 
on sorghum. Like other species of Srriga, little is known regarding floral 
biology despite the potential value of such work on a parasite that repro- 
duces entirely from seed. However, preliminary work by Parker (personal 
communication) suggests that at least some populations are distinctly out- 
crossing. 


— 


STRIGA ASPERA ( Willd.) Benth. 

Striga aspera and S. hermonthica are superficially quite similar and may 
be found growing together. They may be readily separated, however, on 
the basis of bract size and corolla pubescence. The bract subtending the 


198 


=p” 
Pe 


ee 


= Mam 
~— 


Z Lint is 
- cpio wh 
ZEESEES 
cases Sw, 


bre 
ne § ee ' 


=a Ue. 


(Scale for silhouettes equals 10 cm, for drawings 1 cm) 


ae : Striga hermonthica, portion of inflorescence. 
2. Calyces of S. hermonthica. Figure on right shows lateral bract. Smaller 

structures are bracteoles ; 

Fig. 3. S. aspera indowescenice: Musselman and Mansfield 5524 

Fig. 4. S. aspera, diminutive plant from Kuffena Rock; i te and Mansfield 
5530. 

Fig. 5. Same as Fig. 4, showing as 

Fig. 6. S. astatica, Tegina; Masel wa Mansfield 553 

Fig. 7. American strain of S. astatica trom U.S.D.A ee Laboratory. 

Fig. 8. Flowers of S. astatica; ee and Mansfield 5531. 


199 


flower of S. aspera is almost as long as the calyx (Fig. 4) while the bract of 
S. hermonthica is much shorter than the calyx (Fig. 2). The entire external 
portion of the corolla of S. aspera is covered with glandular hairs (Fig. 2) 
while S. hermonthica lacks glands (Fig. 2). Additional differences include 
the more abrupt angle of the corolla tube of S. hermonthica (Fig. 2, left) 
and the larger lower leaves. The two species are also somewhat different in 
their ecology, S. aspera being more commonly found in natural grasslands 
and S. hermonthica as a weed in sorghum. However, S. aspera has also been 
found to parasitize sorghum but has been invariably assumed to be S. 
hermonthica in such cases. Plants of S. aspera from Kuffena Rock, a massive 
granite Inselberg near Zaria, were diminutive but possessed the characteristic 
bracts and pubescence (Fig. 5). 


STRIGA ASIATICA (L.) Kuntz 


The most widely distributed of all species in the genus and a serious 
pathogen throughout most of its range, S. asatica is not a problem in Nigeria 
despite the planting of corn (Zea mays) on a large scale in some places. 
Plants were growing on an ironstone outcropping near Tegina, Kaduna State, 
in a stand of native grasses. The overall form of these plants was quite 
different from that of the strain introduced to the United States (cf. Fig. 6 
and 7). The Nigerian plants were much more pubescent than the American 
strain (Fig. 8). The American strain has recently been shown to be autogam- 
ous (Nickrent and Musselman 1979). While more careful studies are neces- 
sary, it appears that plants in the Tegina population were not autogamous. 


STRIGA GESNERIOIDES ( Willd.) Vatke 

Unlike the other species in this study, S. gesnerioides does not parasitize 
grasses. It is a serious parasite of cowpeas (Vigna unguiculata). In other 
parts of its range it is known to parasitize tobacco (Hosmani 1978). Two dis- 
tinct strains were observed in this study. The first is the widespread cowpea 
strain (Figs. 9-12), the second was seen only at Mokwa, Niger State and was 
parasitizing a weedy legume Tephrosia pedicellata (Figs. 13, 14). To our 
knowledge, there is no formal taxonomic difference between these two strains. 
They are, however, quite different in some aspects of their morphology and 
growth habit. The flowers of the Mokwa strain were more pink than the 
cowpea strain, the size of the bract was also different (cf. Fig. 9 with 13). 
Most, striking, however, is the general habit of the plant. The Mokwa strain 
only rarely branched (Fig. 14), while the cowpea strain branched repeatedly 
at the ground level (Fig. 9). 

The strain of S. gesnerioides recently introduced into Florida most closely 
resembles the Mokwa strain (Wunderlin, Musselman and Shuey 1979). 


STRIGA MACRANTHA Benth. 


Unlike the four species discussed above, S. macrantha and S. klingii are 


200 


Fig. 9. Cowpea strain of Striga gesnerioides, 


La. 


GRA 


(Scale for silhouettes equals 10 cm, 


Inflorescence of cowpea strain 


a 


nd 


of beate relative to calyx. 
Fi 


ig. 13 
Fig. 14 
Fig. 15. 
Fig. 106. 
Fig. 17. 
Fig. 18. 


. Mokwa strain of S. gesnerioides, 
, cL. strain of S. aleneieee showing a: 
Inflorescence of S. macrantha, 


of S. klingii, 


Inflorescence 


Tegi 


Mokwa; 


Flower and bract of S. &lin 


Face view of corolla, 


S. 


Mac aan 


na; 
Musselman and Mansfield, 


Kazure; 


Musselman and Mansfield, 


spect. 
Musselman and Mansfield, 


Musse 


for drawing 1 cm) 


+ 
Has 
A Nh) 
[Lad 
Woy \ 
Vy \ 
iA. 
ad hy 
oh ae 
LV 
Iman and Mansfield, 


2. Individual flowers of cowpea strain of S. gesnerioides show size 


5534. 


D225 
5533 


201 


not known to be of any economic importance. Striga macrantha was collected 
near Tegina, Kaduna State, on an ironstone outcrop where it was growing 
with S. astatica in rocky soil amongst grasses. It was up to 1.5 m tall and 
had large white flowers, apparently the only Nigerian species characterized 
by that corolla color. The flowers are crowded on a dense spike (Figs. 15, 
18) and the bracts and calyces are densely pubescent with both glandular 
and non-glandular hairs. 


STRIGA KLINGU (Engl.) Skan 

Great variation in the size of plants were observed in this species. It is 
apparently a characteristic plant of the dense Hyparrhenia (Poaceae) grass- 
lands of the Southern Guinea Savanna. When growing amongst tall grass, 
the plants are quite short while those on the margin of the grasslands are 
considerably more robust. This species is characterized by small tg flowers 
in crowded spikes and hispid hairs on the large bracts (Fig. 16, 1 

A workshop was held in Khartoum, Sudan in November eae on Striga 
and Orobanche. Of major concern were S. asiatica, S. angustifolia, and S. 
hermonthica although some time was devoted to the S. gesnerioides problem. 

he emphasis of the meeting was on control in its broadest sense including 
breeding for resistance/tolerance in host species. At present one of the more 
promising means of control is the use of germination stimulants that cause 
seeds of the parasite to germinate without a host—thus committing “suicide”. 
A group of compounds developed at Sussex University are now being tested 
for their efficacy in this regard. A summary of this meeting and suggestions 
for future work will be published in PANS. One critical area includes research 
on taxonomy and host specificity. 

Thus, much opportunity exist for taxonomic research in the genus Striga. 
Unfortunately for American botanists interested in conducting a biosystematic 
study, the species are under federal quarantine and cannot be grown in the 
United States without special permission. 


REFERENCES 


HEPPER, F. N. 1963. Scrophulariaceae. In: J. Hutchinson and J. M. Dalziel. Flora 
of West Tropical Africa. Second Edition. Crown Agents for Overseas Governments, 
London. 

HOSMANI, fe - 1978. Striga (a noxious root parasitic weed). Published by the 
author, Dhar 

NICKRENT, D. L ad L. J. MUSSELMAN. 1979. Autogamy in the American strain 
of witchweed. Hee 31: 253-256. 

WUNDERLIN, P., L. J. MUSSELMAN and A. G. SHUEY. 1979. Striga ges- 
Deregiees cei 2 Vatke first record of the species in the New World. Plant Dis 
Rep. 61: 251-25 


NOTES 


HYPTIS MUTABILIS (LABIATAE) IN SOUTHEASTERN UNITED 
STATES.—In September 1977 I collected, in far southeastern Louisiana, a 
member of the Labiatae that was not at the site when I was there in summer 
from 1963 to 1972. After some struggle through a scant and not-too-helpful 
literature, I finally identified the species as Hyptis mutabilis (L. Rich.) Briq., 
a weedy plant widespread in warm America and new to Louisiana. 

This paper is the result of my curiosity concerning H. muwtabilis. It sum- 
marizes literature on the species, provides illustrations of the species (there 
apparently are no good, easily accessible illustrations in the literature ), and 
gives data on its occurrence in southeastern United States. 

By the most recent monographer of Hyptis (Epling 1949), H. mutabilis 
was recognized as a single, highly variable species not divisible into infra- 
specific taxa (Epling 1949) although in older works it was treated as a 
species with several varieties or even as several species (Epling 1933, 1936). 
Its representatives in the United States were treated as two species or as two 
varieties of one species. Such distinctions fail when many specimens are 
examined. 

In Epling’s firs’ work on Hyptis (Epling 1933), he used “var. spicata 
Brig.” for those representatives of H. mutabilis in the West Indies and the 
Caribbean coastal regions. (Presumably var. spicata is what today would be 
called var. matabilis, although this point is not resolvable from Epling’s 
prose; he never was able to locate the holotype of H. muwtabilis.) In his 
synopsis of South American Labiatae, Epling (1936) continued to use “var. 
spicata Brig.” in the same way. However, he commented: “I have sought to 
indicate here the principal modes of variation within this puzzling com- 
plex [H. mutabilis],; however, it is impossible to refer numerous specimens 
with certainty to any one group [variety], and I hesitate to cite specimens 
within the divisions [varieties] I have indicated.” In Epling’s last work on 
Hyptis (Epling 1949), he reduced spicata and other varieties recognized 
in 1933 and 1936 to synonymy under H. mutabilis. Abandoning efforts to 
distinguish infraspecific taxa, he wrote: “. . . parece que no posse razas bien 
definidas” (he did, however, refer to his 1936 work where “La indiqué.. . 
las variantes principales”). In all his papers, he recorded H. mutabilis only 
from Florida in southeastern United States. 

The author of the epithet mwtabilis is frequently and incorrectly given 
as “A. Rich.” (e.g., in Epling 1933, 1936; Small 1933). ia actual author, 
Louis Claude Marie Richard (usually cited as “L. Rich.” or “L. C. Rich.” ), 
published Nepetha [sic] matabilis in 1792. Louis eee pea his atten- 
tion from Nepetha and fathered Achille Richard (“A. Rich.”), who was not 
born until 1794. 


SIDA 8(2): 202, 1979. 


203 


Hyptis mutabilis was recorded from southeastern United States as early 

130 years ago. Bentham (1848), under H. spicata, included the note 
“Florida (h. Torrey! ),” evidently based on a collection (or possibly two) 
he had received from the herbarium of John Torrey. Two undated sheets— 
one at NY, one at GH—appear to be duplicates of the specimen(s) Bentham 
saw; they bear, among equivocal collection data and in handwriting, the 
annotation H. spicata and the notes “fide Bentham” (NY) and “Benth. in 
lit.” (GH 


Gray (1878) ascribed H. spicata to “S. Florida.” Chapman (1889) re- 
ported it from “Tampa and Jacksonville, Florida’; he gave the same dis- 
tribution later (Chapman 1897), but the word “introduced” was added with- 
out explanation (and possibly without justification). Small (1903) recog- 
nized, under the name Mesosphaerum, two species in the mutabilis complex: 
Mesosphaerum spicatum and M. mutabile. The former was ascribed to Florida 
and Alabama, the latter to Florida. (I cannot verify, through herbarium speci- 
mens, the presence of H. mutabilis in Alabama that early.) Small later 
(1933) withdrew the Alabama record: using the name Hyptis, he cited H. 
mutabilis only from Florida and said, of H. spicata, “not now definitely known 
rom our range.” 

More recently, a good account of H. mutabilis is that by Standley and 
Williams (1973), who gave a detailed description of the species but did 
not ascribe it to conterminous United States at all. Long and Lakela (1971) 
gave the range as “Fla. to Va. 

The specimens of H. mutabilis | have seen suggest that, in southeastern 
United States, this species was originally 
only from Florida. From that state it has, in the last 3.5 decades, spread 
northward and westward, reaching Louisiana very recently. 


The earliest specimens of H. mutabilis available to me from southeastern 
United States (herbarium of John Torrey, NY, ) were, as indicated 
above, collected in Florida before 1848. All other pre-1900 specimens are 
from Florida. Indeed, the earliest non-Florida specimen I saw was collected 
in southeastern Virginia in 1939 by Fernald and Long. That this specimen 
represents a non-persistent introduction is suggested by the facts that (1) 
it is the only Virginia collection I saw and (2) the collection locale is sepa- 
rated by 500 miles from the main U.S. range of the species. The northward 
and westward spread of H. mutabilis can be seen in exsiccata: the earliest 
western Florida (te. the Panhandle) collection I saw is from 1897 (the 
species was only rarely collected there until the 1940s and later); the earliest 
Georgia collection, 1940; Alabama, 1952; Mississippt, 1968; and Louisiana, 
1977 (my collection was made just a few hundred feet west of the Louisiana- 
Mississippi border, which the species had obviously just crossed in its west- 
ward push. The documented distribution of H. mutabilis in southeastern 


SIDA 8(2); 203. 1979. 


204 


Fig. Hyptis mutabilis. Documented distribution in southeastern United States 
ee locale omitted ) 


United States is shown in Fig. 1. 

Available specimens of US. H. mutabilis suggest that the species has 
become much commoner in recent decades. The decade-distribution of speci- 
mens seen by me is as follows: pre-1848, 1 collection (possibly 2); 1850s, 
none; 1860s, none; 1870s, 2; 1880s, 1; 1890s, 9; 1900s, 5; 1910s, 1; 1920s, 
1; 1930s, 2; 1940s, 7; 1950s, 16; 1960s, 43; and 1970s, 34 (through 1976 
except my 1977 Louisiana specimen ). 

It is, of course, possible that specimen-based data on spread and abundance 
of H. mutabilis represent vagaries of collection and collectors rather than 
biological verities. But the data nevertheless strongly support my speculations. 

Of the 127 conterminous U.S. collections of H. mutabilis | have seen, 93 
are from Florida, 21 from Georgia, 6 from Alabama, 5 from Mississippi, 1 
from Louisiana, and 1 from Virginia. 


SIDA 8(2): 204. 1979. 


205 


Fig. lyptis mutabilis. Flower (left) and fruiting calyx (right) (based on 
as eee Curtiss 5112, Jacksonville, Florida, NY) The vertical line 
represents 0). 


Hyptis mutabilis occurs mostly in disturbed places (e.g., roadsides, rail- 
road yards, orchards, gardens, lake shores, river banks, fallow fields, ballast 
ground, thickets, and grazed areas). Some label data, though, suggest that 
it grows also in less disturbed habitats (e.g., stabilized dunes, hardwood or 
softwood forests, and hammocks). It is at home in moist or dry soil and 
in sun or shade. The method of its dissemination is unknown. No label data 
or literature reports suggest that H. mwtabilis is a troublesome weed. 

My Louisiana collection (St. Tammany Parish: 8 mi SE of “Slidell along 
hwy. 190, 8 Sep 1977, Thieret 50223, KNK) was made from a colony of 
about 25 plants in a roadside weedy area bordering a channel of the Pearl 

iver. Associates were mostly grasses, including Paspalum spp., Cenchrus 
incertus, and Eragrostis oxylepis. | especially noted two characteristics of 
the specimen I collected. First, its corollas, though small, were relatively 

showy, being oe purple with white blotches; and second, its foliage, 
crushed, ha oma so delightful that H. ables is certainly one of 
Louisiana’s eat mints, perhaps second only to Satureja georgiana. 

The best identifying characteristic for H. mutabilis is the calyx (Figs. 2, 

3). About 1.5-2.0 mm long, and campanulate in flower, it enlarges consider- 


SIDA 8(2): 205. 1979. 


206 


Fig. 3. Hyptis mutabilis. Section of fruiting inflorescence (Mitchell 350, Jackson 
Co., Florida, FSU). The vertical line represents 1 cm 


ably and quickly in fruit, becoming as much as 7.5 mm long and tubular. 
The 10 longitudinal veins increase in prominence, and strong cross veins 
develop to connect them. The accrescent calyx was first noted by Richard 


SIDA 8(2): 206. 1979. 


207 


(1792), the describer of the species, who wrote, “calycibus defloratis elon- 
gatus’; the characteristic was probably the basis for the epithet »utabilis. 

The following description of H. mutabilis is based primarily on specimens, 
secondarily on literature. 


Fig. 4. Hyptis mutabilis. Section of plant with dense inflorescences (Duncan 
17187, Wayne Co., Georgia, GA). The vertical line represents 4 cm 


SIDA 8(2): 207. 1979, 


208 


Plants herbaceous, erect, to 2.2 m high, often much branched, the branches 
glabrous to villous. Leaves membranaceous, 2.0-18.0 cm long, narrowly to 
broadly ovate to rhombic (rarely subrotund or even reniform), acute to 
acuminate at the apex, attenuate, cuneate, truncate, or subcordate at the base, 


Hyptis mutabilis. Section of plant with sachs ae inflorescences (Shuey 
eae oa Co., Florida, USF). The vertical line represents 4 cm 


SIDA 8(2): 208. 1979. 


209 


finely to coarsely crenate to serrate, sometimes doubly so, variously pubescent 
to glabrate, passing abruptly or gradually into bracts; petioles 0.5-8.0 cm 
long. Flowers sessile to very short pedicelled, in bracteate verticels, these 
arranged in dense or interrupted (Figs. 4, 5), leafy or naked, terminal, 
spiciform inflorescences 3.0—20.0 cm long; bracts ovate to elliptic, acute to 
acuminate, prominently nerved. Calyx tubular to narrowly campanulate and 
5-2.0 mm long at anthesis, tubular, reticulate-veined, and to 7.5 mm long 
in fruit, the teeth subulate to setaceous, erect, 0.7-1.2 mm long. Corolla 
3.0-5.0 mm long. 

I am grateful to the curators of the following herbaria for loan of speci- 
mens: A, FSU, GA, GH, MISS, NCU, SMU, TEX, US, and USF. Dr. Carroll 
E. Wood, Jr., gave help with the literature—John W. Thieret, Faculty of 
Biological Sciences, Northern Kentucky University, Highland Heights, 41076. 


REFERENCES 

BENTHAM, G. 1848. Labiatae. DC. Prodromus 12:27-603. [H. spicata, 121.} 

CHAPMAN, A. W. 1889. Flora of the southern United ao 2nd ed. Ivison, Blake- 
man, & Company, New York. 698 pp. [H. spicata, 637. 

= . 1897. Flora of the southern United States. 3rd ed. American Book Com- 

ny, New York, 655 pp. [H. spicata, ee 

EPLING, C. 1933. Synopsis of the genus Hyptis in North America. Repert. Spec. 
Nov. Regni Veg. 34:37-130. [H. nn 103-106. 

1936. peas of the South American Labiatae, 3. Teil. Repert. Spec. 
Nov. Resa Veg. Beih. 85:193-288. [H. mutabilis, 259-262.} 
1949. See del género Hyptis. Revista Museo La Plata, Secc. Bot., N.S. 
7:153- 497. [H. mutabilis, 317-318.} 

Bee A. 1878. Synoptical flora of North America, Vol. 2, Part 1. Gamopetalae 
after Compositae. Ivison, Blakeman, Taylor, & Company, New York. 402 pp. [H. 
spicata, 350; account repeated in ed. 2, 1886.} 

, R. W., an LAKELA. 1971. A flora of tropical Florida. University of 
Miami Press, Coral Gables, Florida. 962 pp. [Hyptis, 748-749. 

RICHARD, L. 1792. Catalogus plantarum, ad societatem, ineunte anno 1792, 
Cayenna missarum a Domino le Blond. Actes Soc. Hist. Nat. Paris 1(1) :105— 114, 
[Nepetha mutabilis, sp. nov., 110. 

SMALL, J. K. 1903. Flora of the southeastern United States. Published by the 
author, New os 1370 pp. [Mesosphaerum, i.e., Hyptis, 1052-1053; account re- 
ee in - 25 

023): Tae of the southeastern Flora. University of North Carolina 
Press, Chapel Hill. 1554 pp. Deda 1180-1181.] 

STANDLEY, P. C., and L. O. AMS. 1973. Labiatae. Mint Family, 

317. In a of Coens ee Bot. 24, Part IX, Number 3. [H. mutabilis, 
254.} 


TRILLIUM PUSILLUM (LILIACEAE) IN MISSISSIPPI—Trillivm pusil- 
lum Michx. is apparently one of the rarer species of the eastern United 
States. Roe (1978) considers the species to consist of four more or less 
disjunct populations probably of varietal status. His map shows a wide dis- 
tributional gap between South Carolina and Texas, which has in part been 


SIDA 8(2): 209, 1979, 


210 


filled by recent collections from Sumner Co., Tennessee and Madison Co., 
Alabama (R. Kral, pers. comm.). We here report T. pusillum from Missis- 
sippi, thus further bridging this disjunction. 

1978 we first collected plants of an unknown Trillivm with 
pedicellate fruit from Jones Co., Mississippi. It was initially noted as being 
somewhat different from all previously known species of the genus in 
Mississippi, but was not definitely identified until comparison with material 
at US. It is apparently most similar to material from South Carolina and 
North Carolina. The purpose of this paper is not to clarify the status of 
the varieties, but the material would therefore be referable to var. pusillum. 
The later collection of flowering material further confirmed the identity of 
this plant with T. pasillum. 

The habitat in which Trillium pusillum was collected by us is a rich 
alluvial floodplain dissected by several small creeks which empty into Bouge 
Homa Lake. Inundation occurs probably at least once a year. The first time 
we collected this species, it was necessary to wade through floodwater to 
get to the area. The second collection (in flower) was made only one day 
after the entire bottomland had been flooded. 

The bottomland forest where Trilliam pusillum was collected has been 
disturbed by selective logging, but is still reasonably well-shaded by uncut 
hardwoods. Dominent species are Pinus glabra Walt., Fagus grandifolia Ehrh. 
and Magnolia grandiflora L. associated with many other hardwood species 
such as Ulmus americana L., Fraxinus caroliniana Miller, Caprinus caroliniana 

alt., Quercus lyrata Walt, QO. michauxii Nutt. and Q. nigra L. Associated 
herbaceous species include ‘Sons parviflorus Raf., Sisyrinchium angusti- 
foliwm Miller and various species of Viola, Carex, Panicum and Polygala. 

There is a combination of factors which may have been responsible for 
Trillium pusillum having been overlooked in Mississippi and possibly other 
areas before now. First, it flowers very early in the spring, in fact is one 
of the earliest plants of the area. Also in some years the plant may be 
nearly inaccessible due to flooding. Certainly the habitat is not unique and 
sites similar to that where T. pwsillum was collected by us are scattered 
across the southern portions of Mississippi, Alabama, and Georgia. 

Specific data for the collections mentioned above are given below. 

MISSISSIPPI. Jones Co.: 7 m E Laurel, disturbed beech-magnolia-spruce 
pine woods, wet ground near ne road, 9 May 1978, Morgan & Me- 
Daniel 468 (IBE, MISSA, duplicates to be distributed), 7 Mar 1979, Morgan 
1589 (IBE, MISSA, duplicates to be distributed )—David Morgan and Sid- 
ney McDaniel, Dept. Biological Sciences, Mississippi State University and 
Institute for Botanical Exploration, Box EN, Mississippi State, MS 39762 


REFERENCE 
ROE, G. F. 1978. Additions to the Range of Trillium pusillum. Castanea 4: 191-195. 


SIDA 8(2): 210. 1979. 


211 


RANGE EXTENSIONS OF BRAZORIA PULCHERRIMA LUNDELL 
(LAMIACEAE ). —Brazoria pulcherrima Lundell is now known from four 
counties growing in disturbed loose white sand as a pioneer species. In 
addition to collections from Leon County, collections from the other three 
counties are (SMU): Freestone County—Mahler 8552, Anderson County— 
Mahler 8553; Houston County—Mabler 8545. 

This taxon was one of the proposed taxa for the Federal List of Endangered 
Species (Federal Register 41 (117): 24549. 1976) since it was known 
only from a local area. Funds for this field study were provided by the 
office of Endangered Species, U.S. Fish and Wildlife Service, Albuquerque, 
New Mexico. 

The range of distribution of the species has not been greatly increased 
but its role as an invader species of recently distributed loose white sands 
enables the taxon to apparently co-exist with man and his activities even 
though it is a rare endemic to Texas—IWm. F. Mahler, SMU Herbarium, 
Dallas, TX 75275. 


RUBUS TRIVIALIS MICHX. VAR. DUPLARIS (SHINNERS) MAH- 
LER, COMB. NOV. (ROSACEAE ).—Based on Rubus duplaris Shinners, 
Field & Lab. 22: 27. 1954. Holotype: Freestone County, 13.6 mi S of Fair- 
field, 2 May 1953, Shinners 14465 (SMU!). 

Rubus duplaris Shinners was one of the proposed taxa for the Federal 
List of Endangered Species (Federal Register 41(117): 24562. 1976) since 
it was known from only two locations (counties) in Texas at that time. 
Subsequent field work has resulted in expanding the known distribution of 
this taxon and observations of the variability of the amount and type of 
pubescence warrant the recognition of this taxon at the varietal level. 

This variety commonly grows in the deeper sands (Carrizo) of the oak- 


red, glandular hairs on the upper and lower leaflet surfaces of both primo- 
canes and floricanes. In adjacent areas of shallower sands and sandy loam 
soils, the amount and type of pubescence varies on the primocane and flori- 
cane leaflets. The specimens from Angelina County are intermediate and 
possess few red, glandular hairs on the leaflet surfaces. In some locations, 
Rubus trivialis var, trivialis and var. duplaris grow together with varying 
amounts of abundance and type of pubescence on the leaflets of the primo- 
canes and floricanes on the same plant. 

The Rare Plant Study Center of the University of Texas reported that 
TEX did not have any collections of Rubus duplaris (A preliminary survey 
of the a of proposed endangered Texas plants (FR 6/16/76). 
19772 U.S: PWS): 

The range of the distribution of this variety is given in Figure 1 and 
documentation includes more than thirty specimens (SMU) collected by 


SIDA 8(2): 211. 1979. 


ya Wes 


Trans—Pecos 


Gale cree oo 
Fayette Pra 
Gulf Cone ere 
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14 Big Thicke VEGETATION ZONES OF TEXAS 


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1. Documented distribution of Rvbuws trivialis Michx. var. duwplaris (Shinners ) 


Fig 
Mahler. 


Shinners, Kral, Cory, McVaugh, Lipscomb, and Mahler. The vegetation zones 
in Figure 1 are a compilation of past researchers. 

I am indebted to Barney Lipscomb and Larry Lodwick for their assistance 
and to ee se of Endangered Species, U. S. Fish and Wildlife Service, 
er ee , New Mexico for providing travel funds—Wm. F. Mabler, 

MU a, Dallas, TX 75275. 


SIDA &(2)2 212. 1979, 


215 


A NOTE ON THE DISTRIBUTION OF STYRAX AMERICANA 
(STYRACACEAE) IN TENNESSEE.—The range of Styrax americana Lam. 
var. americana, the American Snowbell, is primarily confined to the Atlantic 
and Gulf Coastal Plain, Piedmont, and Mississippian Embayment as indi- 
cated by the distribution map of Gonsoulin (Sida 5: 191-258, 1974). The 
rarity of this taxon in the Interior Low Plateau in general and the apparent 
absence of it from this Province in Tennessee or from other provinces east- 
ward in Tennessee is striking. This absence was also noted by Shanks in his 
lists of Tennessee woody plants (Journ. Tenn. Acad. Sci. 27: 27-50, 1952, 
28: 1958-1959, 1953; Castanea 17: 90-96, 1952). In fact, Shanks included it 
as an indicator species of the lowland forests of the Mississippian Embay- 
ment in western Tennessee (Journ. Tenn. Acad. Sci. 33: 195-210, 1958). 
However, recent collections from Stewart County (Schibig, 514, APSU, 
VSCC, Chester 3288, APSU) definitely adds this species to the ILP Province 
in Tennessee. 

Another fact of significance is that the Hamilton County collection cited 
in the Gonsoulin text is not indicated on the distribution map; hence the 
addition of this species to the eastern Tennessee flora may be overlooked 
if only the map is consulted—Joe Schibig, Volunteer State Community 
College, Gallatin, TN 37066 and Edward W’. Chester, Austin Peay State 
University, Clarksville, TN 37040. 


——t 


NOTES ON PLANTS OF MISSISSIPPI II. ADDITIONS TO THE FERN 
AND FLOWERING PLANT FLORA—In the course of field work on the 
flora of Mississippi, in 1978-79, I have found some ferns and flowering 
plants which appear not to have been previously collected in Mississippi. 
None of the species of ferns reported here were included in the most recent 
treatment of pteridophytes of Mississippi, Evans (SIDA 7(3): 282-297. 
1978). 

Herbarium specimens are in the University of Tennessee, Vanderbilt Uni- 
versity, and Museum of Natural Science (Jackson). 


PTERIDOPHYTA 
Asplenium resiliens Kunze 

Tishomingo County: Pickwick Lake: Wooded bluffs between Eastport 
Beach and entrance to Yellow Creek. Crevices of shale. Rare. 4-5 Oct 1978, 
Rogers 46125, 46195, 

Athyrium thelypteroides Desv. 

Hinds County: About 4.0 mi N of Edwards from I-20: Mesic woods in 
ravines in Loess Hills region. Rare. 20 Oct 1978, Rogers 46264, Tishomingo 
County: Yellow Creek peninsula, cove N of Cooks Landing. Rare in mesic 
woods, 19 May 1979, Rogers 46710. 


SIDA 8(2): 213. 1979. 


214 


Cheilanthes alabamensis Kunze 

ilkinson County: Clark Creek natural area: dry border of mixed woods 
on a road cut. Rare. 7 Aug 1978, Rogers 45417. 
Cyrtomium falcatum Presl 

Adams County: About 10 mi NW of Natchez just N of Hwy 555, T8N, 
R2W, Sect 27; wooded ravines in Loess Hills region. Frequent locally. 10 
May 1978, Rogers 45178. 

Adams County: Loess Hills just N of Quitman Road at the delta, T8N, 
R3W, Sect. Several plants growing on steep ravine slopes. 16 Aug 1978, 
Rogers 45540, 

Adams County: Loess Hills just N of Hwy 552, about 3 mi W of Mt 
Zion Church, T12N, R13E, Sect 32. Several plants on ravine slopes. 16 
Aug 1978, Rogers 45544. 

Pellaea astropurpurea Link 

Pohoniees County: Yellow Creek peninsula, T1S, R16W, Sects 24 and 
30; in second-growth hardwood-pine on calcareous shale bluffs. Rare 5 Oct 
1978, Rogers 45544. 

Selaginella uncinate Spring 

Jefferson County: About 1/4 mi W of Alcorn State University farm and 
campus: ravine slopes and bottom, in full sun. A single large population 
seen, numbering several hundred individuals. Rare and local. 5 Apr 1978, 
Rogers 45062. 


SPERMATOPHYTA 
Carex grayi Carey 

Tishomingo County: Yellow Creek Quad, T1S, RI1E, Sect 30, first cove 
N of Whetstone Branch. Rare and local in rich mesic woods, 15 May 1979, 
Rogers 46640. 

Dicentra cucullaria (L.) Bernh. 

Tishomingo County: Yellow Creek Quad, T1S, RIOE and RIIE, Sects 24, 
25, and 30. Rare and local in rich woods and on shale bluffs. 10 Apr. 1979, 
Rogers 40443. 

Hydrophyllum appendiculatum Michx. 

Tishomingo County: Yellow Creek peninsula, T1S, R1I1E, Sect 30; first 
cove north of Whetstone Branch. Rare in moist cherty deciduous woods. 
15 May 1979, Rogers 46638. 

Hydrophyllum macrophyllum Nutt. 

Tishomingo County: Yellow Creek peninsula, T1S, R11E, Sect 30; first 
cove N of Whetstone Branch. Rare in moist rocky deciduous woods. 15 May 
1979. Rogers 46637. 

Kochia scoparia (L.) Roth 


SIDA 8(2).: 214. 1979. 


219 


Hinds County: Jackson: along railroad tracks W of Travelodge Motel at 
Amite and Adams streets. 9 May 1979, Rogers 46621. 

Phacelia bipinnatifida Michx. 

Tishomingo County: Yellow Creek Quad, T1S, RIOE and R1I1E, Sects 24, - 
25, 30. Rare in rich mesic woods and wooded shale bluffs. 10 Apr 1979, 
Rogers 46442. 

Philadelphus hirsutus Nutt. 

Tishomingo County: Yellow Creek Quad, T1S, R10E, Sect 14. Very rare 
on calcareous shale bluffs, in pine-oak woods. 15 May 1979, Rogers 46628. 
Scutellaria ovata var. versicolor (Nutt.) Fern. 

Tishomingo County: Yellow Creek peninsula, T1S, R11E, Sect 30. First 
cove N of Whetstone Branch. Rare in rocky, mesic deciduous woods. 15 
May 1979, Rogers 46643. 

Taenidia integerrima (L.) Drude 

Tishomingo County: Yellow Creek Quad, T2S, R11E, Sect 31. Whetstone 
Branch; dry rocky south-facing slopes; several plants occurring locally. 11 
Apr 1979, Rogers 46503. 

Tradescantia ernestiana Anders. and Woodson 

Tishomingo County: Yellow Creek Quad, T1S, R11E, Sect 30; Boo Hollow. 
Rare in mesic woods. 15 May 1979, Rogers 46650. 

Trautvetteria caroliniensis (Walt.) Vail 

Tishomingo County: Yellow Creek Quad, T2S, R11E, Sect 6. Very rare 
and localized along Short Creek, on calcareous shale, in seepage areas E of 
road leading to Cooks Landing. 9 Apr 1979, Rogers 46502. 

Viola pensylvanica Michx. 

Tishomingo County: Yellow Creek Quad, T1S, R10E, Sect 24; Meat Hol- 
low. Several plants locally in cherty moist deciduous woods. 14 May 1979, 
Rogers 46629—Ken Rogers, Mississippi Museum of Natural Science, 111 
North Jefferson Street, Jackson, MS 39202. 


NEVIUSIA ALABAMENSIS (ROSACEAE) IN MISSISSIPPI—Neviusia 
alabamensis A. Gray, commonly known as Snow Wreath, is a low shrub in 
the Rose Family. It has been known previously from Tuscaloosa County in 
Alabama, Conway County, Arkansas, and Butler County, Missouri (Clark, 
1971; Small, 1933; Steyermark, 1963). 

In October 1978 I found a large colony of Snow Wreath, in the hilly 
woodland of northeast Mississippi, in Tishomingo County, growing on 
calcareous shale bluffs overlooking the Tennessee River and Pickwick Lake. 
A second colony was found in November in the same stretch of bluffs. 

Sun Wreath was growing in full sun on the bluffs which lie in a narrow 
belt from Eastport northeast to Yellow Creek. Several species of plants found 


SIDA 8(2); 215, 1979. 


216 


here are either unique to the bluffs or occur rarely elsewhere in Mississipp1. 
A few plants associated with Snow Wreath were Dodecatheon meadia, 
Pellaea atropurpurea, Woodsia obtusa, Solidago sphacelata, Heuchera villosa 
var. macrorhiza, Delphininm tricorne, Cladrastis lutea, Euonymus atropur- 
pureus, Fraxinus quadrangulata, Quercus mublenbergii, Quercus prinus, 
Aesculus glabra. 

Herbarium specimens (Rogers 46149, 46652, 46670) are in The Univer- 
sity of Tennessee, Vanderbilt University, and Museum of Natural Science 
(Jackson). —Ken Rogers, Mississippi Museum of Natural Science, 111 
North Jefferson Street, Jackson, MS 39202. 

THE CHROMOSOMES OF COLLINSONIA CANADENSIS | L. 
(LABIATAE).—The following preparations represent the first published 
illustrations of the chromosomes of Collinsonia, a genus of mints confined 
to eastern North America. The mitotic chromosomes (27 = 50) are shown 
in Figure 1 and the meiotic chromosomes (7 = 25) in Figure 2—all derived 
from Collinsonia canadensis L. (PA, Centre Co.: C. S. Keener 1978, PAC). — 
Carl §. Keener, Department of Biology, The Pennsylvania State University, 
University Park, PA 16802. 


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NICOLLETIA OCCIDENTALIS (COMPOSITAE) IN BAJA CALI- 
FORNIA!—A specimen at SD (Valle de San Felipe, southeast of Santa Clara, 
ca. 31°03'N, 115°13'W, with Larrea, Fouquiera, Simmondsia, and Cercidium 
at ca. 400 m., 3 Apr 1977, Fred Sproul s.n.), brought to my attention by 
Dr. R. Moran, documents the occurrence of Nicolletia occidentalis in Baja 
California. This collection extends the known range of the species some 300 
km. south to within ca. 100 km of the nearest known population of N. trifida 
(cf. my review of Nicolletia, Sida 7:369-374. 1978). The Parishes are not 
known to have collected in Lower California (cf. ibid.)—John L. Strother, 
Department of Botany, University of California, Berkeley, CA, 94720. 


SIDA 8(2): 216. 1979. 


SIDA susie" 


VOLUME 8 NUMBER 3 JUNE 1980 


CONTENTS 
The systematics of Hedeoma (Labiatae). Robert S. Irving. 218 
Historical account 219 
Generic limits and relationships 220 
Chromosomal studies ae 
Breeding habits 223 
Natural interspecific hybridization 224 
Specific concepts 725 
Ecology, distribution, and possible origin 225 
Systematic treatment 229 
Excluded names 292 
Acknowledgements 294 
References 294 


Taxonomy of Tetragonotheca ( Asteraceae-Heliantheae ). 
B. L. Turner and Deborah Dawson. 296 


New species of Clitoria subgenus Bractearia section 
Brachycalyx (Leguminosae) from Colombia, Venezuela, 
and Brazil. Pawl R. Fantz. 304 


NOTES. A chromosome count for Juniperus ashei (Cupressaceae) and additional 
chromosome numbers for Hedeoma (Labiatae). 312—-New and. te- aes 
combinations in Guwtierrezia (Compositae: Astereae). 313—He sper aloe funifer. 
(Agavaceac) in Texas. 314—Cwueurbita digitata (C ucarbitnceae ) in 
Phyllanthus polygonoides (Euphorbiaceae) new to Arkansas. 315. 


US ISSN 0036-1488 


SIDA, CONTRIBUTIONS TO BOTANY 
Founded by Lloyd H. Shinners, 1962 


Editor & Publisher 
Wm. F. Mahler 
SMU Herbarium 

Dallas, Texas, 75275 


Associate Editor Assistant Editor 
John W. Thieret Barney L. Lipscomb 
Northern Kentucky University SMU Herbarium 
Highland Heights, Kentucky, 41076 Dallas, Texas, 75275 


Guidelines for contributors are available upon request beginning with 
volume 8. 


Subscription: $6.00 (U.S.) per year; numbers issued twice a year. 


Sida, Contributions to Botany, Volume 8, Number 3, pages 218-315. 
Copyright 1980 
by Wm. F. Mahler 


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Frontispiece. Hedeoma microyhyllum Irving. 


THE SYSTEMATICS OF HEDEOMA 
(LABIATAE) 


ROBERT S. IRVING 
1422 Summit, Little Rock, AR 72202 


The genus Hedeoma (Labiatae) is a group of annual and perennial herbs 
found throughout southwestern United States and Mexico and in scattered 
localities in South America. In this treatment it is composed of 38 species 
and 5 varieties. 

Hedeoma is a member of the subtribe Melissinae of the tribe Stachyoideae 
of Briquet (1897). Its generic boundaries have undergone widely divergent 
modifications during the last 170 years, as indicated by the publication of 
Persoon (1807), Bentham (1832-1836, and 1848), Gray, (1878), Briquet 
(1897), and Epling and Stewart (1939). The intent of this work, although 
by no means definitive, is to present a coherent delimitation of Hedeoma 
based on extensive field and experimental data. 

In addition to the problems of generic limits and phyletic position of 
Hedeoma, there are numerous problems at the specific and intraspecific levels. 
In previous works, including the most recent inclusive treatment of the group 
by Epling and Stewart (1939), there was little or no attempt to explore 
questions of intra- and interspecific relationships or phylogeny. Such an 
exploration is one of the primary goals of this monograph, and it is believed 
that the present treatment to a great extent, places the taxa of Hedeoma in 
natural groupings not only from the standpoint of morphology, ecology, and 
geography but phylogeny as well. 

A list of representative specimens is available on request. 


HISTORICAL ACCOUNT 


The genus Hedeoma was proposed by Persoon in 1807 to accommodate 
three species: H. thymoides, H. glabrum, and H. pulegioides. In Bentham’s 
(1836) treatment of Hedeoma, H. thymoides and H. glabrum were excluded 
from the eae H. pulegioides was retained as the type species and seven 
other newly described or previously unrecognized species were added. He 

also divided the genus into two sections, Mosla and Hedeoma. Between 
1848 and the publication in 1876 of the Genera Plantarum, Bentham 
elevated the section Mosla to generic status and added four additional 
species to Hedeoma proper bringing the total to 12. Gray’s treatment 
in 1878 resulted in a circumscription very similar to the present one. 
Gray recognized eight additional taxa as belonging to Hedeoma, and 
divided the genus into the sections Exhedeoma, which includes most of the 


SIDA 8(3); 218-295, 1980. 


220 


currently accepted species, and Stachydeoma. The latter section was elevated 
to generic rank by Small (1903); later its two species were disaggregated 
into the monotypic genera Rhododon and Stachydeoma (Epling and Stewart, 
199?) 

In Briquet’s (1897) treatment of Labiatae, Hedeoma was expanded to 
include Keithia Benth., Poliomintha Gray, and other related genera. How- 
ever, Epling and Stewart (1939) returned to the earlier generic delineation 
of Bentham and of Gray. They believed that Bentham’s account was a more 
natural, logical, and consistent interpretation of the relationships involved 
than that of Briquet, who, for lack of a single clearcut difference between 
these genera lumped them together. 

The present treatment is essentially that of Epling and Stewart and | 
am in full agreement with their arguments against Briquet’s expanded 
generic concept. My treatment differs in that three of the four North Ameri- 
can representatives of Epling’s genus Hesperozygis have been incorporated 
into Hedeoma and several new taxa have been added. 


GENERIC LIMITS AND RELATIONSHIPS 


There are 11 other genera which, on morphological grounds, are con- 
sidered closely allied to Hedeoma. Conceivably, all of these genera, including 
Hedeoma, could be considered to constitute a single very large genus, in 
fact seven genera, Mosla, Rhododon, Stachydeoma, Poliomintha, Eriothymus, 
Hoebnea, and Rhabdocaulon have at one time or another been merged with 
Hedeoma. 

The genus Z7z/phora, of the Mediterranean region, can be considered the 
Old-World counterpart of Hedeoma as its overall habit and floral morphology 
are similar. In Bentham’s treatment of the Labiatae (1832-1836, and 1848), 
Ziziphora was placed in the tribe Monardeae, thus suggesting considerable 
morphological divergence of Hedeoma from Ziziphora. Briquet (1897), on 
the other hand placed Ziziphora next to Hedeoma in his subtribe Melissinae. 
Ziziphora and Hedeoma can be distinguished by several morphological fea- 
tures. Z/z/phora possesses a tubular calyx with connivent, subequal, narrowly 
triangular teeth—a feature very reminiscent of the calyces of Poliomintha. 
Further, Z7z/phora is separated by a technical character of the anthers; unlike 
Hedeoma, the lower anther-sacs of Ziziphora are completely or partially 
aborted 

The diverse genus Cumila, despite an obvious morphological affinity with 
Hedeoma, has always been placed distantly from Hedeoma by both Bentham 
and Briquet. The features which distinguish the genera are subtle. The 
stamens of Cwnila are seated above the middle of the corolla tube, ascend 
from the orifice, surpassing the upper lip. The tubular or turbinate calyx has 
10-13 nerves, and is rarely more than 5-6 mm long; the teeth are subequal. 
It should be noted that within Hedeoma, H. floribundum has characteristics 
of the calyx similar to those found in this genus, perhaps suggesting an even 


221 


closer relationship than previously supposed. 

Pogogyne is a small California genus which both Bentham and Briquet 
reckoned it to be a close relative of Hedeoma. However, careful comparison 
of the two genera reveals a number of striking differences. The principal 
features of Pogogyne which distinguish it from Hedeoma are as follows: 
habit, annual; inflorescence a dense terminal “spike” of cymes; calyx deeply 
cleft and 15-nerved; stamens 2 or 4; anther sacs parallel; style pubescent; 
and, chromosome number, 72 = 19 (Ornduff, 1967) 

The genus Glechon was recognized and treated as a close relative of 
Hedeoma by Bentham (1832-36 and 1848). Briquet (1897), on the other 
hand, placed Glechon into a different subtribe. Epling and Stewart (1939) 
again aligned the genus with Hedeoma. Since the principal distinguishing 
feature resides in the corolla, it is not surprising that Briquet, who strongly 
emphasized corolla morphology in his treatment, should fail to indicate close 
relationship between the two genera. Briefly, Glechon differs from Hedeoma 
in being composed of South American fruticose herbs with sessile cymules; 
calyx tube 13-15-nerved, turbinate, with subequal deltoid or deltoid-subulate 
teeth; corolla ample, abruptly expanded, the upper lip manifestly longer than 
the lower; stamens genticulate at the base and hairy. 

The Asiatic genus Mosla was originally placed in Hedeoma, later, as more 
species became known, it was separated from Hedeoma and given generic 
rank, Mosla differs from Hedeoma primarily in having a 10 rather than 
13-ribbed calyx and in having a racemose inflorescence. 

Rhododon ciliatus was included in Hedeoma sect. Stachydeoma and later 
Stachydeoma. In 1939 Epling and Stewart elevated it to generic rank. 
Rhododon is readily distinguished from Hedeoma. Rhododon is a coarse 
annual with sessile, long pilose-ciliate leaves and has a dense terminal ‘spike’; 
a 15-nerved, deeply cleft calyx; no staminodes; a pubescent style; and a 
chromosome number of 2 = 

The genera, Hoehna, Eriothymus, and Rhabdocaulon are all believed to be 
offshoots of the diverse genus Kerthia. Bentham treated Keithia as very close 
to Hedeoma, but Briquet (1897) included the whole of Kesthia into 
Hedeoma. Epling not only recognized Keithia as distinct but further divided 
the group into three genera, a treatment with which, after careful examina- 
tion of many representatives of these groups, I am in complete accord. 

Stachydeoma graveolens was included in the genus Hedeoma by both Gray 
(1878) and Briquet (1897). In 1903, Small created Stachydeoma as a dis- 
tinct genus based on H. graveolens and H. ciliatum. Unlike Rhododon ciliatus, 
with which it was previously grouped, Stachydeoma cannot be kept easily as 
a distinct genus. It is very Hedeoma-like in the details of the calyx and 
corolla and in the important features of the nutlets. The separating features 
reside in the pilose, sessile leaves and in its chromosome number, 7 = 9 

Poliomintha and Hesperozygis are also allied very closely to Hedeoma. As 
the differences are more quantitative than qualitative, the genera are con- 


222 


trasted with Hedeoma in the tabular form below. 


Hedeoma Poliomintha 
soccer semishrubs, usually her- Shrubs. 
eous perennials or annuals. 

Calyx tube gibbous or saccate. Calyx tubular and symmetrical. 

Calyx teeth usually acuminate and well Calyx teeth subsequal, convergent to 

differentiated into upper and lower sets. close the orifice. 

Calyx annulus in a well-defined ring. Calyx annulus absent or in an irregular 
ring. 

Nutlets oblong or orbicular in outline, Nutlets oblong, o 1.5 mm long, not 

ca. 1.5 mm long or less, usually Reem becoming een when moistened. 

ing mucilaginous when moistened. 

2n — 34, 36, 44, 72, 144. 2n = 36. 

Hedeoma Hesperozygts 

ses N. American Primarily S. American 

Semi-shrubs, of more co ommonly peren- Shrubs with leathery leaves. 

re or eas herbs, leaves various but 

not leathery. 

Calyx 13-nerved. Calyx 13-16 nerved. 

Calyx tube various but not funnelform. Calyx tube funnelform. 

Calyx teeth various. Calyx teeth ae ae deltoid lanceolate, 
erect a par 

Calyx annulus seated at or near the Calyx annulus usually seated near the 

juncture of the upper and lower teeth middle of the tube 

at the tube’s summit. 

2n = 34, 36, 44, 72, 144. 2n = 44, 


The genus Hesperozygis in both its original circumscription (Epling, 
1936; Epling and Stewart, 1939) and in subsequent additions (Epling, 1940 ) 
houses a bewildering array of seemingly unrelated taxa. It is rendered some- 
what more homogeneous by removing to Hedeoma three of its former species, 
but the remaining species are problematical still and are likely to remain so 
until a more comprehensive study is undertaken. 

All of the above-cited genera are closely interrelated. Though they all 
have two stamens, they are probably related to the four-stamen groups 
through the genera Satureja and Melissa. Indeed, Bentham (1832-36) stated 
that, “Hedeoma scarcely differs from Miécromeria except in the abortion of 
the upper stamina.” 


CHROMOSOMAL STUDIES 


Chromosome numbers for Hedeoma and related genera are reported else- 
where (Irving, 1976). In summary, 27 = 36 was the most frequently en- 


225 


countered number occuring in 67% of the taxa reported. Section Alpine 
was consistently 27 = 44. Aneuploidy (2 = 34) occurred in several taxa 
distributed along the eastern edge of the generic range: a cluster of popula- 
tions of H. drummondii in east-central Texas, H. hispidum, H. acinoides and 
both varieties of H. reverchonii. Polyploidy was found in the southernmost 
populations of H. costatum (2n = 72) and in H. multiflorum (2n = 72) 
and H. apiculatum (2n = 144). Poliomintha longiflora was 2n — = 36; 
Hesperozygis marifolia, 2n = 44; Rhododon ciliatus, 2n = 26 and Stachy- 
deoma graveolens, 2n =18. Based on an analysis of the distribution of 
chromosome numbers and the ae of S. graveolens, the genus Hedeoma 
is judged to be dibasic x = 9, 1 


BREEDING HABITS 


Inbreeding, in a variety of forms, is found in all species of ee 
The lower fertile pair of anthers, typically unseparated, dehisce and th 
style, which lies behind the anthers, recurves sharply at its tip into the ex- 
posed pollen mass. The time between the dehiscence of the anthers and 
the recurvation of the style, however, provides reproductive variability among 
the species. 

A few of the annual species (H. hispidum, H. medium and H. pulegioides ) 
are primarily cleistogamous and only occasionally are chasmogamous plants 
encountered. In the small to medium flowered species, which are the majority 
of Hedeoma species, the flowers open and the dehiscence of the anthers and 
the recurvation of the style occur almost simultaneously. A few of these 
species (H. acinoides, H. nanum, and H. costatum) can be — 
The remainder are chasmogamous inbreeders and their populations are often 
subtly and consistantly distinct from one another. Hedeoma Dei 
H. costatum, H. nanum are good examples of this phenomenon 

Among the larger flowered species (H. hyssopifolium, H. pulcherrimum, 
and H. ciliolatum) the opening of the flower and the maturation of the 
anthers and style are separated by periods of time. The anthers open approxt- 
mately 12-24 hours after anthesis and the style requires 24-48 hours, after 
anthes’s, to recurve into the exposed pollen. Insects (bees, moths and butter- 
flies) are commonly observed visiting the large flowered species and pre- 
sumably outbreeding is the predominant mode of reproduction. Occasionally 
through a special mechanism some plants of the largest flowered species, 
such as H. ciliolatum, are obligate outbreeders. Following anthesis in these 
plants, the anther pair separates and is no longer in the path of the recurving 
style. 

In general the mode of reproduction has determined in large part my 
thinking as to the evolutionary development of the genus, and is well cor- 
related with other plyletic characteristics. Large flowers and outbreeding are 
considered primitive traits and are typically associated with the long-lived 
perennial or semi-shrub habits and very restricted and non-weedy distribu- 


> 


224 


tion patterns. Small flowers and inbreeding are believed to be derived char- 
acteristics and most commonly occur in weedy, widely distributed annuals 
and short-lived perennials. 


NATURAL INTERSPECIFIC HYBRIDIZATION 


From both herbarium and field studies there is evidence of interspecific 
hybridization involving at least seven species in the genus Hedeoma. The 
putative hybrids have been documented both morphologically and through 
the use of secondary plant constituents (Irving, unpublished ms. ) 


H. DRUMMONDII var. DRUMMONDIT X H. NANUM var. NANUM 

Both H. drummondii var. drummondii and H. nanum var. nanum occur 
in pure form in the mountains of southern Nevada and adjacent California, 
and for each this area represents the extreme western range of their distribu- 
tion. Usually, these two taxa are quite distinct, but numerous specimen (cited 
below ), although determined as H. nanum, are actually intermediate between 
H.d, var. drummondii an m. var, nanum. Acceptance of their hybrid 
status follows directly from a chromatographic survey of the specimens cited 
which revealed both chromotographically distinct parents and a summation 
of phenolic patterns for the intermediates (Irving, unpublished ms.). 
Representative putative hybrids: 

ALIFORNIA: San aes Co.: Keystone Canyon, 14 May 1940, Alexander & 


Kellogg 1483 (DS, GH). NEVADA: Clark Co.: Lee Canyon, 22 Jul 1939, Clokey 
8476 (UC). Clark ae Harris Spgs. Rd. 10 Jun = Clokey 8633 (NY). Clark 
Co.: Kyle Canyon, 14 Jun 1939, Clokey 8478 (DS, GH, IA, MICH, NY, ae 


POM, US). Clark Co.: Deadman Canyon, 30 May oe Alexander & Kellogg 1614 
(DS, Se TEXAS: Val Varde Co.: junction of rts 90 and 277, 4 Jun 1967, Irving 
fo) CTEXs 


H. DRUMMONDII var. DRUMMONDII & H. PULCHERRIMUM 

These two closely related species occur in sympatric populations in the 
mountains of southern New Mexico around Cloudcroft. A visit to one such 
population 20 miles northeast of Tularosa revealed the presence of numerous 
intermediate plants. The latter were assumed to be hybrids since paper 
chromatography displayed an additive pattern for these intermediates and 
counts of pollen stainability revealed ca 20% stainable pollen. Both parental 
species consistently displayed 100% stainable pollen (Irving, unpublished 
ms. ) 

aid MEXICO: Otero Co.: 20 mi ne of Tularosa, 5 Aug 1967, Irving 835 a & b 
(TEX 
H. DENTATUM X H. HYSSOPIFOLIUM 

The occurrence of these hybrids were detected solely from morphological 
analysis of herbarium specimens deposited at LA. The hybrids are similar to 
H. hyssopifolium but possess reduced gibbous calyces and dentate-entire leaves 


bo 
i) 
i) 


(the calyces of H. hyssopifolium are usually large and nongibbous and the 
leaves entire). 
a eee hybrid specimens: 
NA: Cochise Co., Huachuca Mts, 5 Sep 1936, C. Epling and W. Stewart s.n. 

(LA) “0 sheets ) . 
H. DRUMMONDII &* H. REVERCHONII var. REVERCHONII * H. R. var. 

SERPYLLIFOLIUM 

Each of these taxa come together on the arid limestone outcroppings of 
the Edwards Plateau of Central Texas. Here all three form large sympatric 
populations on the same chromosome level (27 = 34). Although inbreeding 
and microhabitat differences work to maintain the integrity of each taxon, 
hybridization in all possible combinations occurs and much of the inter- and 
intra-populational variation appears to be the result of past and present gene 
exchange. Gas chromatography, morphology, and pollen fertility were em- 
ployed in the detection of hybrids (Irving and Adams, 1973). 


SPECIFIC CONCEPTS 


The taxa of Hedeoma are delimited by combined information taken from 
morphology, cytology, ecology, and geography. I have tried to be consistent 
in my recognition of species, subspecies, and varieties; that is, I have at- 
tempted to accord formal names to comparable categories. In several cases, 
as in H. palmeri and H. acinoides, various intraspecific races have been 
acknowledged but were not accorded formal taxonomic recognition. In regard 
to the species concept itself, I shall refer to discussions presented by Raven 
(1962) and Lewis and Lewis (1955). The species concept presented in 
these papers most accurately reflects my own beliefs 

With the categories subspecies and variety, the term “variety” is reserved 
for divergent subspecific populational units which display genetic and geo- 

raphic integrity. The term sages: is used as a grouping category for 
closely allied varieties, much as the subgenus is used to group closely allied 
sections. This usage is as interpreted by Turner (1965), and Kapadia 
(1964) 

ECOLOGY, DISTRIBUTION, AND POSSIBLE ORIGIN 


As a prelude to a discussion on the ecology, distribution, and possible 
origin of the species in Hedeoma, 1 would like to cite what I consider to be 
primitive and derived characters for the genus (Table 1). In an effort to 
present a phyletic picture, I have adopted the “Wagner Divergence Index 

Fig. 1) as developed by W. H. Wagner, Jr., and illustrated by Hardin 
(1957). To derive a divergence index for a given species the characters 
were ranked 0, 1, or 2, depending on their assumed primitive, intermediate 
or derived condition, respectively. A numerical total, which might be termed 
a phyletic index, is obtained by summing the value for each of these char- 


CALIATUM & 
= as 


TOD 
5 1) Ss) 


MS SATUREJOIDES 


ACIN 
Sect. Alpine wy, 
POLIOMINTHOIDES PATR c 
MOL AN 


5 .@) 15 


1. Divergence in the genus Hedeoma. Concentric arcs represent levels of 


ig 
specialization. Open circ 
taxa (see text). 


| 


es suggest hypothetical ancestors; and solid circles are extant 


20 


9C¢ 


Pa 


acter states for any given taxon. The phyletic indices are then plotted upon 
concentric circles and arranged so as to form a phyletic “tree” or dendrogram 


ie these calculations there have been no weighting of characters; for 
Seaeaple, the characters of the leaf margin carry as much weight as characters 
relating to corolla size. In many cases “intermediate” states were not found; 
ee they are not listed in the tabulation. 

m Table 1, the primitive Hedeoma would be an entire-leaved shrub 
an ae pubescence and solitary flowers; the calyx tube would be radially 
symmetrical with erect, united teeth and no annulus; the corolla would be 
large with a flat upper lip and possessing a definite annulus; nutlets ae 
be brown, oblong, not becoming mucilaginous when moistened; and there 
would be 36 large, symmetrical chromosomes. No single taxon has af of 
the criteria for one state or the other, although several approach the purely 
primitive or derived condition. In particular, three species emerge as primi- 
tive for the genus: H. céliolatum, H todsenti and H. palmeri (phyletic indexes 
7, 8, and 8, respectively; Fig. t the other end of the spectrum the 
annual, H. Deda. is the most advanced, possessing 13 of the criteria for 
the derived condition (phyletic index 24); it is closely followed by 
acinoides (phyletic index 23) 


PRIMITIVE AND DERIVED C HARACTERS _ 


DERIVED (2) 


PRIMITIVE (0) INTERMEDIATES (1 - 


— 


bo 


. Shrubby or ‘suffrutescent 


. Leaves entire 


3. Hairs simple, plants 


Awa) 


relatively glabrous 


. Flowers solitary 


. Calyx tube radially 


symmetrical or nearly so 


6. Calyx teeth erect and o 


CO 


slightly eeu into 
upper and lower s 


. Upper calyx teeth scarcely 
. Calyx annulus absent 


. Corolla large bn 12 


mm) and wit 
expanded ee 


Herbaceous perennials 
Intermediate 

Hairs simple; plts 
pubescent 

Cymules 3—5 flowered 


Tube gibbous or sigmoid 


Intermediate 


Teeth united for less than 
length 


4 their 


A broken or undulating 
ring 


Less than 12 mm or long 
tubular 


Anau 
Variously toothed 


Hairs branched 


Cymules 5 or more 
flowered 


Tube saccate 


Teeth variously 
reflexed and well 
differentiated into 
upper and lower sets 
Teeth united over 4 
their length 


A definite ring 


Minute and bar 
exserted from e iene 


228 


10. Delayed autogamous/ Intermediate Autogamous and/or 

allogamous in breeding cleistogamous 
abit 

11. Nutlets not becoming Mucilaginous when 
mucilaginous when moistened 
moistened 

12. Nutlets oblong in outline Orbicular in outline 

13. Chromosome number 2” 2m — 34, 72, or 144 
= 36, 44 

14. Chromosome large and Small and 
symmetrical asymmetrical 

15. Plants of mesic, montane Intermediate Open-arid habitats 


habitats 


The genus Hedeoma is divided into 4 subgenera and it is my impression 
that each developed early in the evolution of the genus and subsequently 
each followed its own evolutionary course. Ciliatum and Poliominthoides are 
believed to be the oldest of the sub-generic lines, with Satwrejoides and 
Hedeoma occupying the intermediate and youngest ae respectively. 
No firm evidence is available with which to determine the time of divergence 
of these phylads, but it is suggested that prototypes for ee. ee de- 
veloped out of the Madro-Tertiary Geoflora during the Oligocene, or perhaps 
earlier. As the extant species of Hedeoma are associated with both the 
Madro-tertiary and Arcto-tertiary floras, the ancestral stocks may have de- 
veloped in the ecotonal region between these two floras (Axelrod, 1950). 

he subgenus Ciliatwm is composed of five narrowly endemic species 
which are spatially and morphologically well-separated. This fact, along with 
their large number of primitive traits, speaks for a rather lengthy evolutionary 
history for the group. The species are confined to relict communities of the 
Sierra Madrean woodland of the dissected Madro-Tertiary flora (Axelrod, 
958). Hedeoma ciliolatum and H. todsenti are the most primitive, occuring 
in mesic pine forests near Galeana, Nuevo Leon, Mexico and the forest 
communities of the San Andres Mountains of New Mexico respectively. 
Hi. apiculatum and H. pilosum, are more advanced species occurring in the 
open, more xeric rock crevices in the mountains of Trans-Pecos Texas. 

The subgenus — is believed to have had an evolutionary 
pattern similar to that of Czliatum. Its four taxa, while also very distinct 
morphologically, are a wider ranging. To the north, H. patrinum, H. 
montanum, and H, molle are associated with the drier elements of the Sierra- 
Madrean woodland and to the south H. palmeri, occurs principally in the 
pine-oak woodlands of the Sierra Madre Oriental. 

The subgenus Saturejoides can be divided into two geographical and 
morphological groups: (1) section Alpine in which the members are all 


229 


montane species of the Sierra Madre Occidental; and (2) section Saturejoides, 
the species of which occupy a wide variety of habitats. 

he ecology and distribution of section Alpine suggest a strong affinity 
with the Arcto-Tertiary Geoflora and the section may well have had its origin 
within it. The pine forests in which the species of Alpine now occur are be- 
lieved to be extensions of the Arcto-Tertiary flora which extended southward 
with the uplifting of the Rocky Mountain chain, The lack of transitions with 
other species and the disjunctive distribution pattern of its species suggest 
that the former ranges of species belonging to this section cael have been 
contracted with the spread of aridity during late Tertiary times. 

Section Saturejoides is composed of taxa of varying Kae and habitats 
and can be considered the core of the genus. Xerophytic annuals and peren- 
nials dominate the section and, for the most part, the North American taxa 
can be aggregated into four species complexes: 1) H. costatum-plicatum- 
tenuipes-microphyllum, 2) H. ices aa inltiled op me a6 aUOn a aut: 
dentatum-hyssopifolium, and 4) H. d dii-reverchonit-4 
These complexes typically have wide distribution Paes and display combi- 
nations of relatively specialized morphological attributes. Each complex in- 
cludes a less specialized and distributionally restricted montane species, H. 
costatum, H. diffusum, H. hyssopifolium, and H. pulcherrimum respectively. 
The xerophytes presumably evolved with the expansion of arid areas and 
dissection of the Madro-Tertiary Geoflora. 

The subgenus Hedeoma has been the most difficult with which to contend 
in terms of origin and phylogeny. It seems to be a recent line, on morpho- 
logical grounds, but its connection with the other subgenera of Hedeoma is 
obscure. Further, the species are well separated and do not merge one into 
the other as might be expected in a youthful phylad. 


SYSTEMATIC TREATMENT 

HEDEOMA Pers., Syn. Pl. 131, 1807 

Annuals or caespitose, suffrutescent, or herbaceous perennials, forming 
semi-shrubs, tufts, clumps, or mats. Shoots solitary to numerous, strict, 
ascending, or decumbent and often producing adventitious roots from the 
lower nodes; shoots variously branched; arising from a slender or thickened 
taproot, woody caudex, or trailing rootstock; variously pubescent with hairs 
simple and spreading or retrorsely curving (tomentose, in H. molle with 
hairs profusely branched). Leaf pairs crowded or well-spaced. Leaves mem- 
branous or coriaceous; ascending, spreading, or cernuous; oval, ovate, ellip- 
tical, oblong, lanceolate, linear or spatulate; entire, crenate, serrate, or dentate 
(often obscurely so); apex obtuse or acute; base sessile, subsessile, or petio- 
late; blades tomentose (HL molle), canescent (subgenus Poliominthoides), 
or variously pubescent with ciliate margins; nervation indistinct to con- 
spicuous with the nerves forming raised costae on the undersurfaces (H. 
plicatum, H. pusillum, and H. ciliolatum), the leaf surfaces smooth, glandular- 


250 


dotted, or glandular-punctate to varying degrees. Flowers solitary or dis- 
posed in axillary cymes (cymules) 1-31 flowered, well-spaced along the 
axis or aggregated into a terminal “spike” of cymes; peduncles inconspicuous 
to several millimeters long; pedicels conspicuous, usually several millimeters 
long and variously pubescent with spreading or retrorsely curving hairs, 
bracteoles typically subulate or lanceolate, either shorter or longer than the 
pedicels they subtend. Calyx 3.0-15.0 mm long, highly zygomorphic to 
nearly actinomorphic (subgenus Poliominthoides); tubular, tubular-sigmoid, 
or tubular-funnelform; usually gibbous or saccate below for 14 to over 34 
of the tube’s length, variously pubescent and usually with the pubescence 
diminishing in size and density toward the teeth apices (evenly tomentose 
in H. palmeri and H. molle); the calyx teeth markedly or scarcely (sub- 
genus Poliominthoides) differentiated into upper and lower sets; the three 
upper teeth variously connate forming an upper lip; the connate portion 
of the lip either short or expanded, and either glabrous or pubescent on its 
inner face; the lobes semi-ovate, deltoid, narrowly triangular or subulate, 
glabrous or hirtellous to hirsute-ciliate, laterally spreading or erect and 
convergent (H. multiflorum, H. drummondi:); the two lower teeth generally 
not coalesced, narrowly triangular or subulate, and typically recurved and 
surpassing the length of the upper lip (short, equal to the upper lip in the 
subgenus Hedeoma), ciliate, or, occasionally glabrous on the margins; calyx 
with a well-defined ring of simple hairs (the annulus) within which tend 
to be projected outwardly and partially seal the calyx orifice, occasionally 
upwardly extended onto the inner face of the upper lip (H. piperitum, 
H. medium, and H. multiflorum), 0.5-1.5 mm. wide and seated at the 
juncture of the upper and lower teeth or below it, included or exserted 
from the calyx tube. Corolla white, blue, lavender, pink, or orange-red, 
extremely variable, 3.0-45.0 mm long, always zygomorphic with the limbs 
forming upper and lower lips, glabrous or pubescent within in two decurrent 
lines along an abaxial groove or channel running along the throat floor or 
simply pubescent in the throat, also commonly pubescent within the tube, 
occasionally the hairs forming a well-defined annulus; the tube gradually 
or abruptly expanded upwardly; the upper lip obcordate to ligulate, concave 
and subgaleate or flat, straight or reflexed away from the axis, scarcely to 
deeply emarginate; the lower lip spreading to erect (subgenus Hedeoma), 
equally three lobed or more commonly with the central lobe exserted beyond 
the laterals and bilobed. The abaxial stamen pair arising from the middle of 
the corolla (upper %4 in. H. martirense) and ascending upwardly under 
the upper corolla lip or surpassing it, the anther sacs nearly parallel to widely 
divaricate on well-developed connectives; the adaxial pair reduced to stami- 
nodia, 1.0-2.0 mm long, arising opposite the fertile pair on the adaxial 
surface, occasionally the staminodia bear abortive anther sacs at their apices 
(H. pulegiodes). Style long and bifid at the apex with the abaxial segment 
markedly longer than the adaxial one, glabrous. Nutlets orbicular, ovate, or 


251 


oblong in outline, ovate or ovate-triangular in cross section, brown or black 
at maturity, the surface areolate, foveate, or ruminate, weakly to strongly 
glaucous, usually mucilaginous when moistened. Chromosome numbers, 27 


= 34, 36, 44, 72, or 144 (x =9 


Type species: Hedeoma pulegioides (L.) Pers. 
Distribution: North and South America. Centers of distribution appear to 


be in the mountains of far northwestern Mexico, and the mountains of 
southern Arizona and New Mexico; also in the Sierra Madre Oriental of 
east central Mexico. 


1. 


— 


— 


KEY TO THE SUBGENERA OF Hedeoma 


pane caespitose, often forming dense mats, 4.0-12.0 cm high; leaves coriaceous, 

tire and sessile; flowers solitary or in three-flowered axillary cymules; corolla 
large - oe 10.0—-32.0 mm long; western Texas, ee Mexico and northern 
J. Subgenus CILIATUM 


, Plant. ieee eta annuals or perennials or semi-shrubs; leaves various; flowers 


in three or usuall ore flowered axillary cymules; corolla 3.0-45.0 mm long, 

plants variously Babar: 

2. Suffrutescent, canescent perennials, usually semi-shrubs, 15.0-100.0 cm. tall; 
leaves canescent; axillary cymules commonly but not always aggregated into 
terminal “spikes”; calyces tubular, nearly radically symmetrical with the teeth 
only slightly differentiated into upper and lower 


Oe I. Subgenus POLIMINTHOIDES 

2. Plants usually herbaceous annuals or perennials; ce variously pubescent but 
never canescent; calyces bilaterally symmetrical with the teeth markedly differ- 
entiated into upper and lower sets. 

3. Leaves and calyces various; lower calyx teeth usually longer than upper; 
nutlets brown to dark brown at maturity, commonly glaucous, oblong or 
occasionally obicular in outline, mucilaginous when moistened, plants vari- 

Ill 


ously distributed. . Subgenus SATUREJOIDES 
3. Leaves ovate or elliptic, toothed; calyx saccate, lower teeth short; nutlets 

black at maturity, never glaucous, orbicular in outline or nearly so, 

at all mucilaginous when moistened. Plants of South America and eastern 

North America. V. Subgenus HEDEOMA 


KEY TO THE TAXA OF THE SUBGENUS CILIATUM 


Leaves with regularly spaced costae on the undersurface and finely ciliate margins; 
calyces densely pubescent on the inner faces of the teeth; »lants of Mexico 

2. Plants 5.0-10.0 cm tall; leaves elliptical-lanceolate, 5.0-10.0 mm long: corolla 

bright orange-red, large and showy, over 25.0 mm long. 
1. H. ciliolatum 
2. ae 3.0-5.0 cm tall; leaves ovate, 3.5-5.0 mm long, corolla lavender, less 
n 15.0 mm lon 
H. pusillum 


2. 
. Leaves smooth or tuberculate on the undersurface but without regularly spaced 


costae, margins pilose or tuberculate-ciliate; calyx teeth glabrous on their inner 

faces; plants of U.S. 

3. Plants ca 4.0 cm high; leaves ovate, ca 5.0 mm long, calyces pilose. 
3. H. pilosum 

3. Plants 10.0-15.0 cm high, leaves, lanceolate-elliptic, ca 10.0 mm long, calyces 

hispidulous 


252 


. Leaves obtuse or acute at apex; corolla ca 30.0 mm long, — re 
as 
4. Leaves apiculate at apex; corolla ca 20.0 mm long, pink. 
5. H. apiculatum 


KEY TO THE TAXA OF THE SUBGENUS POLIOMINTHOIDES 


1. Plants with sim seca hairs; Mexico 
il 


a" 


ple c 
2. Tall shrubs, with axillary 15-31 flowered cymules, commonly aggregated into 
dense terminal spikes. 


ay 


meri 
2. Spindly shrubs or suffruticose perennials with 1-5 flowered enue not usually 
ageregatec 
3, Spindle shrubs with the upper calyx teeth narrowly triangular and reflexed 
away from the axis; calyx annulus more or less included. ...... 7. H. patrinum 
3. Suffruticose perennials with the upper calyx teeth subulate and, at least par- 
tially, convergent with the lower at maturity; calyx annulus well-exserted. 


montanum 


. Plants covered by a tomentum of highly branched hairs; Texas. .......... 9. H. molle 


KEY TO THE TAXA OF THE SUBGENUS SATUREJOIDES 


. Plants perennial herbs with the shoots decumbent and rhizomatous in some species; 


leaves ovate, elliptical, oblong, or lanceolate, margins entire or obscurely toothed, 
base tapered and subsessile; upper calyx lip broad; corolla showy, the upper lip 


wide; plants of the Sierra Madre Occidental of western Mex XICO. ....., A. sect. Alsine 
2. Plants with numerous ascending and decumbent se ee adventitious 
roots from the lower nodes; nutlets spherical or ne so. 

3. Leaves oblong or oblanceolate; are eet toward the branch apices; 
calyx large, tubular-funnelform, ca 8.0 mm long, 2.0-2.5 mm wide at the 
summit, the tube not at all gi 
4. Leaves crowded with the ea aks uniformly 1.0 mm long, leaves 6.5— 

7.0 mn pi 


cote 0. H. woliedlnetolum 
4, Leaves well-spaced, 10.0-17.0 mm long, the bases not decurrent; calyx 
hirtellous; upper calyx lobes deltoid or semi-ovate; plants . - Am. 


3. Leaves elliptical, ovate or subrotund; cymules well-spaced ae the axis; 

calyx zygomorphic, 5.5-7.0 mm long, 1.5 mm wide at the summit, the tube 
pibbo ous or saccate. 

5. Leaves ovate to narrowly elliptical; the calyx ia Behcon gibbous, the 

upper calyx lobes triangular; corolla 11.0-12.0 mm long. ......00000.00........ 

. . jucundum 

5. Leaves ovate to sub-orbicular; the calyx tube ee bia accate, forming 

a =u pouch, the upper calyx lobes semi-ovate or deltoid; oa la 7.0- 

8 13. £ ee 


2. Plants with ane ascending; nutlets ovoid; leaves with odor of oregan 
6. Cymules well-spaced along the upper 3 of the stems; calyx 4.0-6.5 mm 5 eee: 
corolla 7.0-9.0 mm long. 14. H. patens 
6. Cymules aggregated toward the branch apices forming a sn terminal 
“spike” of cymules; calyx 3.0-3.5 mm long; corolla 5. ae . mm long 
. aH. Fae ei 


259 


1. Annuals or aoa the shoots and leaves various; upper calyx lip not usually 


broad, or if so, t plants annual; corolla various, the up Ee “Pp os or if 
redered markedly a plants auaual: nutlets oblong, ca tw s long as wide; 
plants variously distributed, but not in the Sierra Madre Orci dental of. western 

exico. t. Saturejoides 


7. Plants perennial, or annual with the calyx teeth oe convergent and closing 
the calyx orifice, or with the corolla tbe markedly 
8. Upper and lower calyx teeth convergent, closing a o orifice at maturity, 
or if only slightly so, then plants with entire, long, elliptical-oblong leaves 
and large corolla, 10.0-14.0 mm long, the lower lip 7.0 mm wide 
9. Calyx tube gibbous but not saccate, not tapered upwardly; upper calyx 


the mountains of south central New Mexico. ............ . . pulcherrimum 
ee tube conspicuously saccate and tapered into a narrow neck; the 
upper calyx teeth, clearly convergent with the lower; ae of various 
crap and distribution. 
0. Upper calyx teeth ununited, subulate; nutlets areolate: plants of N. 
Am 


xe 


11. Plaats annuals or herbaceous perennials, occasionally somewhat 
t the a0) 


hirsute, the upper and lower teeth usually Sones convergent; 
corolla 7.0-11.0 mm long; widespread; odor of peppermint. 
i 


drum mondii 

11. Plants suffruticose typically averaging over 25.0 cm in height; 
leaves r dark green, usually less than three times longer 
than ; calyx coarsely hirsute or hoary villous, the upper and 


lower teeth incompletely closing the orifice; corolla 8.0-15.0 mm 
long; west and central Texas to south-central Oklahoma; odor of 
] 


camphor or lemon. 18. H. reverchonti 
10. Upper calyx teeth united, triangular; nutlets foveate; as of S. Am 
19. H ; ee 


8. Calyx teeth not as above, the Bees mae generally spreading and reflexed a 
maturity leaving the calyx tube 
Leaves ovate and entire, a ee cere short and gibbous-saccate, 
the distended region 4 of the length of the tube. 
13. Leaves ovate, acute; corolla often dilated upwardly; nutlets areolate; 
plants of N. Am. 

14. Plants with numerous decumbent stems, forming circular mat 
calyx sania but not saccate, the upper lip pubescent on its inner 
face; corolla ca 9.5 mm long, restricted to the ee forests of 
north-central eee (Blapstath) soe Dare, 20. diffusum 

14. Plants of various habits but not as above; calyx saccate; oe upper 
lip ee rous within; corolla 8.0-9.0 mm long, southwestern U.S. 
and Mexico. 

15. ee entire and smooth; widespread. .............. 21. H. nanum 
15. Leaves denticulate, costate; endemic to central Mexico. ............ 
2 : ie 
13. mas linear-elliptical; corolla not dilated upwardly; nutlets foveate; 
plants of S. Am 23. H. medium 
12. Leaves pen eatned (entire and ovate in the two s species from Baja, 
Caltiorna) calyx tubular or sigmoid, the distended region less than 
1 the length of the tube. 


234 


16. Tufted perennials with the shoots freely branching; axillary leaf buds not 
usually occurring; leaves oval, ovate, linear or elliptic, plants of central and 


Mexico. 
17. Leaves serrate or dentate; ovat, elliptic, rhombic or oblanceolate 
aves ovate, oval or elliptic; leaf nerves distinct but not dlievared: curved 
and branched; calyx sanulus seated at the juncture of Upp er and lower 


lips. 24. H. costatum 

18. Leaves ovate, oblanceolate, elliptic or rhombic; leaf nerves elevated and 
straight; calyx annulus seated below the jmenire of upper and lower 
li 


Ips. 
19. Leaves ovate-oblanceolate, dentate; calyx well over 10 mm ; 
corolla showy, over 30 mm long; known only from the Sierra jardin 
H, Risiee 


oO n Mexic 
19. Leaves elliptic or hone crac serrate; calyx - 8.0 mm long; 
corolla 7.0-10.0 mm long; widespread. .................... oo ee 


17. Leaves entire or obs ae crenate; ovate, dlise or ie 
20. Pouce aes or ovate; oe of Baja California. 
. Leaves 9.0-13.5 long; internodes elongated; cymules 1 
flowere 27. H. tenuiflorum 
21. Leaves 5.0-6.0 mm long; internodes short; flowers solitary in the 
upper leaf axils. 28. H. martirense 
20. Leaves lanceolate (at least the upper); plants of eastern Mexico. 
22. Axillary cymules 1-3 flowered; calyx 7.0-8.0 mm long; corolla ca 


mm long, well exserted from the calyx. ........ 29. H. quercetorum 
22. Axillary cymules many-flowered; calyx 5.0-6.0 mm long; corolla ca 
5.5 mm long, barely exserted from the calyx. ...... 30. H. tenutpes 


16. Tall robust plants with long ascending, usually anes hed shoots; axillary 
eav ieldi 


a 
plants of western New Mexico and eastern Arizona, occasionally in northwestern 


Mexico. 
23. Leaves linear or narrowly elliptical, entire, and subsessile; cen i yy 
flowered; bal slightly gibbous; tien large and showy, 


long, ca 7.0 wide across the lower lip. ................ 31. H. hysopiolm 
23. Leaves elliptical, dentate or cones shortly petiolate or bsessile; 
ymules 1-15 flowered; calyx m markedly gibbous; corolla 8.0-11 0: mm long, 


24. Basal leaves ovate, glabrous, entire. .....0..0.0000000000.. 32. H. ee 
24. Basal leaves ovate-rhomboid, yee pubescent, dentate. 200.000.0000... 
 -anbabain 


7. Plants annual, the calyx teeth not convergent or the corolla markedly Pie 
25. Leaves ovate, sUipecal or rhombic-elliptical, dentate, or entire, petiolate; calyx 
saccate for ca $+ of the tube’s length; corolla only occasionally cleistogam mous. ... 
34. H. acinoides 


25. eaves linear or linear-elliptic, at least five times longer than wide, entire 
sessile; calyx saccate for over 3 of the tube’s length; the upper lip ee the corolla 
straight, corolla often cleistogamous. 35. H. hispidum 


KEY TO THE TAXA OF THE SUBGENUS HEDEOMA 
1. Plants annual; leaves elliptical and dentate; plants of N. Am. .... 36. H. pulegioides 
1. Plants perennial; leaves ovate, obovate, spatulate, crenate or entire; plants of S. Am. 


2. Leaves sary crenate; calyx markedly saccate, ca 4.0 mm long; corolla ca 3.0 
m long, Braz 37. H. crenatum 


23 


2. Leaves obscurely crenate, mostly entire; calyx gibbous or ed slightly saccate, 
4.0-5.0 mm long, corolla 5.0-8.5 mm long, Bolivia and P 


8. H. er es 


I. SUBGENUS CILIATUM, Irving, subg. nov. 

Herba casespitosa vel tegeta, 4-12 cm alta; caules herbacei numerosi simplices; 
folia conferta, coriacea, ovato vel elliptico—lanceolata, integra, sessilia, 3.5-15.0 mm 
longis, 1.5—3.5 mm lata, margine conspicue ciliata; cymae triflorae vel flores singulares; 
calyx infundibuliformis vel oe: tubularis; corolla grandis speciosa; mericarpium 
oblongum madefactum mucilaginum; chromosomatum numerus 2” = 36, 144. 


Plants casespitose, often forming dense mats, 4.0-12.0 cm in_ height. 
Stems herbaceous, numerous, unbranched, arising from a woody caudex or 
trailing rootstock. Leaf pairs crowded, the internodes shorter than 2.0 mm; 
leaves coriaceous ascending, usually folded with drying, ovate to elliptical- 
lanceolate, entire, sessile, 3.5-15.0 mm long, 1.5-3.5 mm wide; margins 
ciliate, the upper and lower leaf surfaces glabrate to puberulent, nervation 
prominent or inconspicuous; undersurface to varying degrees glandular- 
punctate between the costae, the upper surface smooth. Flowers typically 
solitary in the upper leaf axils or in three-flowered cymules (H. apiculatum). 
Calyx tubular-funnelform, not at all gibbous below, distinctly bilabiate in 
both the tube and disposition of the teeth, 5.0-10.0 mm long; the upper 
teeth united for 4-2 their length, ee an upper lip, the lobes tri- 
angular or deltoid; the lower teeth narrow, triangular; the annulus dense and 
included. Corolla lavender, pink or red, usually large and showy, 10.0—32.0 
mm long, pubescent within. Nutlets oblong in outline, at least twice as 
long as wide, ovate-triangular in cross section, brown at maturity, the sur- 
face areolate, glaucous, becoming mucilaginous when moistened. Chromo- 
some number, 27 = 

Type species: HEDEOMA CILIOLATUM Epl. 


1. HEDEOMA CILIOLATUM (Epl.) Irving, Brittonia 22: 345. 1970. 
Hesperozygis ciliolata Epl., Rep. Spec. Nov., Beihefte 115: 13-14. 1939. 


Somewhat robust perennials forming dense clumps 7.0-12.0 cm (10.0 
cm) high. Stems numerous, unbranched or only sparsely so, canescent with 
short retrorse hairs. Leaves coriaceous, folded with drying, elliptical-lanceolate, 
entire, sessile, 1.7-2.5 mm (2.0 mm) wide, 5.0-10.0 mm (84 mm) long; 
tapered to a narrowly obtuse or acutish apex, margins distinctly ciliate; 
upper and lower surface glabrous; nerves of 4-5 prominent pairs, leaves 
smooth above, punctate below. Flowers in the axis of the upper leaves, soli- 
tary or rarely in two-flowered cymules, congested somewhat towards the 
apex; peduncles inconspicuous, ca 1.0 mm in length; pedicels 2.0-4.0 mm 
long, canescent; bracteoles short, oblong. Calyx coriaceous, 8.0-10.5 mm 
(9.9 mm) long, the tube tubular-funnelform, 6.5-8.0 mm (7.3 mm) long, 
2.0-2.2 mm wide above, densely puberulent; the upper teeth scarcely connate, 


236 


forming an upper lip, 1.5-2.1 mm long, the lobes triangular ca 2.6 mm wide 
at the base, 1.0-2.0 mm long, ciliate; the lower teeth lanceolate, ascending, 
1.5-3.0 mm long, ciliate, the annulus dense, included, 0.7-1.0 m wide but 
extended upwardly into the calyx lobes, seated 1.5-2.0 mm below the junc- 
ture of the upper and lower teeth. Corolla brilliant, orange-red, 30.0-32.0 
mm (31.0 mm) long, pubescent within on the palate, densely puberulent 
within the tube, forming annulus ca 4.5 mm wide; the tube, 25.0-27.0 mm 
long dilated upwardly, 5.0 mm wide above; the upper lip broadly ligulate 
5.0 mm wide, ca 6.0 mm long, distinctly emarginate; the lower lip ca 12.0 
mm wide, ca 7.5 mm long. Nutlets ca 0.6 mm wide, 1.2 mm long, the sur- 
face weakly glaucous. Chromosome number, 27 = 36. 

Distribution (Fig. 2). Endemic but abundant on gypsum outcrops in 
Open, mature, pine forests in the vicinity of Galeana, Nuevo Leon. 4,000-— 
6,000 ft. Jul-Sep. 

TYPE: Nuevo LEON. 15 mi southwest of Galeana, 18 Jul 1934, C. H. 
& M. T. Mueller 1062 (Holotype, LA!; isotypes, F! GH! MICH! TEX!). 

There are several features of H. ciliolatum which suggest an alliance to 
Poliomintha. Its large, brightly colored corolla is scarcely distinguishable 
from the corolla of Poliomintha longiflora, and its inner calyx pubescence is 
suggestive of the P. longiflora—P. glabrescens complex. However, its habit 
and clear alliance to H. pusillum, strongly support its inclusion in the sub- 
genus Ciliatum. In no way can I see any alliance to the South American 
representatives of the genus Hesperozygis nor to its sole North American 
representative, H. marifolia, as was suggested earlier by Epling (1939). 


2. HEDEOMA PUSILLUM (Irving) Irving, Brittonia 22: 345. 1970. 
Hesperozygis pusilla Irving, Brittonia 19: 245-247. 1967. 


Small, perennial, caespitose herbs, 3.0-5.0 cm high, forming mats. Stems 
numerous, unbranched, canescent with short retrorse hairs. Leaves coriaceous, 
folded with drying, ovate, entire, sessile, 1.5-2.1 mm (1.8 mm) wide, 
3.5-5.0 mm (4.7 mm) long, tapered to a narrowly obtuse or acutish 
apex, margins finely ciliate, both surfaces evenly puberulent with short 
appressed hairs; nerves of 4—5 prominent pairs; leaves smooth above, 
punctuate below. Flowers solitary in the upper leaf axils, congested toward 
the apex; peduncles short, less than 1.0 mm in length; pedicels 1.0-2.0 
mm long, canescent; bracteoles short, ovate-oblong. Calyx 5.2-7.0 mm (6.2 
mm) long, coriaceous; the tube tubular-funnelform, 4.5-5.5 mm (5.0 mm) 
long, densely puberulent; the upper teeth connate for 1/4-1/3 of their 
length, forming an upper lip, the lobes 1.0-1.5 mm wide at the base, ca 
0.7 mm long, the central lobe commonly reduced to about 1/2 the width and 
height of the two adjacent lobes, the lobes slightly laterally spreading and 
reflexed; the lower teeth lanceolate, short, ascending, ca 1.0 mm long, densely 
puberulent on the inner face; the annulus included, dense, ca 0.5 mm wide, 


257 


extending into the calyx lobes, seated ca 0.5 mm below the juncture of the 
upper and lower teeth. Corolla lavender, 10.0-11.0 mm (10.5 mm) long, 
pubescent within but not annulate; the tube, 8.5-9.5 mm long, only slightly 
dilated upwardly, ca 2.5 mm wide above; upper lip ligulate, subgaleate, 
straight, 1.55 mm wide, ca 1.5 mm long, distinctly emarginate, the lower 
lip ca 3.5 mm wide, ca 2.0 mm long. Nutlets ca 0.5 mm wide, 1.0 mm long, 
the surface, weakly or not at all glaucous. Chromosome number 27 = 36. 

Distribution (Fig. 2). Known only from the pine forests south of Galeana, 
Mexico; often sympathic with H. crliolatum. 

TYPE: Nuevo LEON. 2.5 mi south of Galeana near junction of road to 
Linares, 5-7 Aug 1965, Irving 471 (Holotype, TEX! isotypes, LA! SMU! 
TEX! MONTU! ). 

In general, H. pusillum is a diminutive counterpart of H. ciliolatum. The 
quantitative differences being reflected to some degree in all of the mor- 
phological features. The most conspicuous departure of H. pusillum is in 
its small (10.0-11.0 mm.) lavender corolla which is in marked contrast to 
the long (30.0-32.0 mm), orange-red corolla of H. ciliolatum. 


3. HEDEOMA PILOSUM Irving, Brittonia 22: 340-342. 1970. 


A small perennial, caespitose herb ca 4.0 cm high, probably forming mats. 
Stems extremely short, unbranched above, profusely branched below, pube- 
scent with short spreading hairs. Leaves coriaceous and sharply ascending, 
appressed to the stems, ovate, entire, sessile, 5.0 mm long, 2.5-3.0 mm (2 
mm) wide; tapered to a narrowly obtuse or acutish apex, margins short 
pilose-ciliate, both upper and lower surfaces glabrous; nerves of 1-2 slightly 
raised subopposite pairs. Flowers solitary in the uppermost leaf axils; ped- 
uncles short; pedicels 1.0—2.0 mm long, canescent; bracteoles short, oblone: 
ciliate. Calyx 7.5 mm long, somewhat coriaceous, the tube tubular-funnelform, 
dilated upwardly, ca 5.0 mm long, pilose, the hairs long and spreading; 
the upper teeth connate for ca 1/2 of their length, forming a conspicuous, 
broad upper lip, ca 2.4 mm long, the lobes triangular, ca 0.5 mm wide at 
the base, ca 1.2 mm long, laterally slightly spreading and reflexed, pilose 
ciliate; the lower teeth narrowly triangular-lanceolate, ascending, ca 2.5 mm 
long, pilose-ciliate; annulus dense, included, ca 1.1 mm wide but seated deep 
within the tube at ca 2.0 mm below the juncture of the upper and lower lips. 
Corolla, nutlets unknown. Chromosome number, unknown. 

Distribution (Fig. 2). Known only from the type locality. Open, exposed 
limestone, 5,000-6,000 ft. Jul. 

TYPE: TEXAS. Brewster Co., near top of Baldy Peak, Glass Mountains, 13 
Jul 1940, Warnock 352 (Holotype GH! ) 

Hedeoma pilosum seems to be intermediate between H. pasillum and H. 
apiculatum but due to the incompleteness of the specimen this position is 
tentative. Unfortunately a revisit to the type locality did not reveal any trace 
of H. pilosum and it may well be extinct. 


238 


Fig. 2. Distribution of Hedeoma ciliolatum (closed circles), H. pusillum (open 
circles), H. pilosam (open square) and H. apiculatum (closed square), H. todsenti 


(star 


239 


4. HEDEOMA TODSENII Irving, Madrofio 26: 184-187. 1979. 


Suffrutescent caespitose perennials 10.0 to 18.0 cm high. Stems numerous, 
simple, canescent or puberulent with hairs retrorsely curling. Leaves corta- 
ceous, entire, lanceolate-elliptic, subsessile, 8.0-14.0 mm long, 2.0-4.0 mm 
wide; apex obtuse to slightly mucronate; midrib and margins slightly his- 
pidulose-tuberculate; lower surface deeply glandular punctuate, the upper 
less so. Flowers solitary and well-spaced in the upper portions of the stems; 
peduncles ca 1.5 mm long; pedicels ca 3.5 mm long, canescent; bracteoles 
lanceolate longer than pedicels. Calyx 11.0-12.0 mm long (base to tip of 
teeth), tubular-funnelform, not gibbous, minutely scabrous on the nerves 
and glandular punctate between; upper teeth partially connate forming a 
conspicuously reflexed upper lip, the lobes lanceolate, ca 1.0 mm wide, 1.5 
mm long, sparsely ciliate; lower teeth ascending, deltoid-aristate, 3.0—3.5 mm 
long, sparsely hirsute-ciliate; annulus ca 1.0 mm wide, included and seated 
ca 2.0 mm below the juncture of the upper and lower teeth. Corolla orange- 
red, ca 30.0 mm long, 6.0 mm wide at the throat, sparsely pubescent in the 
tube but not annulate; the upper lip deeply obcordate, erect, up to 5.0 mm 
long, red in color on the interior margins; lower lip streaked with red, 
reflexed, the similar rounded lobes ca 3.5 mm long, 3.0 mm wide; anthers 
exserted. Nutlets ca 2.0 mm long, 1.0 mm wide, dark brown in color and 
weakly glaucous. Chromosome number, unknown. 

Distribution (Fig. 2). Known only from the type locality. Open, exposed 
limestone slopes. Aug—Sept. 

TYPE: New MEXxIco, Sierra Co., San Andres Mts., On steep north-facing 
gravelly gypseous limestone hillside in open pinyon, 2 km s of Hardin 
Ranch, 18 Aug 1978, T.K. Todsen SA 1-78 (aiouee NY; isotypes, NMSU 
US! TEX!). 

Hedeoma todsenii is most closely allied to H. apiculatum. The suffruticose 
habit; the overall patterns of pubescence; the coriaceous, entire, lanceolate 
leaves and the large tubular calyx with its deep seated annulus are some of 
the morphological attributes which are nearly identical in both taxa 


5. HEDEOMA APICULATUM Stewart, Rep. Spec. Nov., Beihefte 115: 25. 1939. 


Small, suffrutescent perennials, 10.0-15.0 cm (12.0 cm) high, forming 
mats. Stems numerous, unbranched, densely clothed in minute retrorsely 
curling hairs. Leaves coriaceous, somewhat folded upon drying, lanceolate- 
elliptic (occasionally ovate below), entire, sessile, 1.5-3.5 mm (2.5 mm) 
wide, 7.0-15.0 mm (10.0 mm) long, tapering to a conspicious but short 
apiculate apex, sparsely hispiduluse-tuberculate on veins and margins; nerva- 
tion indistinct; upper surface smooth, lower rugose and punctuate. Flowers sol- 
itary or in 2—3 flowered axillary cymules, crowded toward the apex; peduncles 
short, 1.0-1.5 mm long, pedice's 3.0-4.0 mm long, canescent, bracteoles 
narrowly lanceolate. Calyx 8.5-9.0 mm (8.8 mm) long, chartaceous; the 


240 


tube tubular-funnelform, flaring above, 5.5-6.5 mm (6.0 mm) long, 1.5—-1.8 
mm wide below to 2.0-3.0 mm wide above, minutely hispidulose-tuberculate 
below, glabrous above; the upper teeth connate for ca 1/4 or less of their 
length, forming an upper lip 2.0-2.2 mm long, the lobes broadly triangular 
or deltoid, apiculate at their apices, ca 1.0 mm wide at the base, 1.0-1.5 mm 
long, laterally spreading and reflexed, hirtellous and tuberculate-ciliate; the 
lower teeth ascending, broadly to narrowly triangular-aristate, ca 1.2 mm 
wide at the base, ca 3.0 mm long, conspicuously hirsute-ciliate; annulus dense, 
ca 1.2 mm wide, with long pilose hairs, seated ca 1.5 mm below the juncture 
of the upper and lower teeth. Corolla pink, showy, 19.0-20.0 mm long, 
pubescent in the throat, sparsely pubescent within the tube; the tube ca 10.0 
mm long conspicuously dilated upwardly, ca 4.0 mm wide above; the upper 
uP broadly obcordate in outline, ca 5.0 mm wide, ca 5.0 mm long, the lower 
lip broad, 9.0-10.0 mm wide, ca 8.5 mm long. Nutlets somewhat pointed at 
their apex, ca 2.0 mm long, 0.7-0.9 mm wide, the surface weakly glaucous. 
Chromosome number, 27 = 1 

Distribution (Fig. 2). Restricted to the Guadalupe Mountains of Texas 
and New Mexico, frequent among rocks of the streamway and canyon sides 
in the south fork of McKittrick Canyon and in the limestone crevices atop 
adjacent Pine Top Mountain. 5,000-8,000 ft. Jul-Aug. 

TYPE: TrExAs. Culberson Co., Guadalupe Mts., sunny crevices along 
stream, McKittrick Canyon, 22 Aug 1931, Moore & Steyer mark 3563 (Holo- 
type, LA!; isotypes, CAS! DS! GH! MICH! NY! PH!). 


Il. SUBGENUS POLIOMINTHOIDES Irving, subg. nov. 

Suffruticosum 15.0-100.0 cm altae incani ad basin valde lignosi; folia crassa 
membranacea, ae ovata vel elliptica, integra vel dentata, petiolata, 5.0-30.0 mm 
longa, 2.0-15.0 mm lata; cymulae axillares 3-31 flori, terminales densi interruptique, 
ageregatae; calyx tubularis, calycis dentes, subulati; series superae et interae minime 
distincti; corolla magna speciosa; mericarpum oblongum madefactum mucilaginum, 
chromosomatum numerous, 2” = 36. 


Suffrutescent perennials, or semi-shrubs, 15.0-100.0 cm tall. Shoots woody 
below, herbaceous above, ascending or shortly decumbent, arising from a 
woody caudex or short woody taproot, canescent. Leaves canescent and 
glandular punctuate, thick-membranous, rotund, ovate, or elliptical, 2.0-15.0 
mm wide, 5.0-30.0 mm long, margins entire or dentate; apex acute or 
obtuse; base narrowly attenuated and petiolate. Axillary cymules, 3—31- 
flowered, well-space along the upper 1/3-1/2 of the stems, or commonly 
aggregated into dense, interrupted terminal spikes. Calyx more or less radi- 
ally symmetrical and tubular, 5.0-8.0 mm long, pubescent evenly dis- 
tributed; calyx teeth only slightly differentiated into upper and lower sets 
(to be greater extent in H. patrinum), the upper teeth commonly subulate 
and scarcely united to form the upper lip; the lower teeth subulate; annulus 
dense and well-formed. Corolla white to lavender, large and showy, over 


241 


13.0 mm long, variously pubescent within, the tube dilating upwardly; upper 
lip ligulate to obcordate, emarginate; lower lip spreading. Nutlets oblong 
in outline, over twice as long as wide, brown at maturity with the surface 
areolate, glaucous and becoming mucilanginous when moistened. Chromo- 
some number 27 = 36. 

Type species: HEDEOMA PALMERI Hemsl. 


6. HEDEOMA PALMERI Hemsl., Biol. Cent. Am. 2: 548. 1881. 
ele rotundifolium Hemsl., Biol. Cent. Am. 2: 549. 1881. Type: HIDALGO: 

“Zimapan,” Coulter 1081 (Holotype. K!; photolypes: GH! LA! 7 PH! -US!). 

hie suffrutescent perennials 35.0-100.0 cm (58.8 cm) high. Shoots 
long, ascending or erect, arising from a woody caudex, branching sparse but 
occurring freely along the axis with the branches diverging at wide angles, 
densely pubescent or puberulent (in many representatives nearly cinerous) the 
hairs retrorsely curving or curling. Leaves variable, ovate or subrotund, 6.0- 
15.0 mm (11.5 mm) wide, 10.0-30.0 (20.0 mm) long; margins crenate or 
dentate, rarely entire; apex obtuse; base broadly or narrowly attenuated, 
petiolate with the petiole 2.5-6.0 mm long; pubescence highly variable, the 
under-surface varying from tomentose to simply pubescent, the margins and 
upper surface glabrate to pubescent; nerves not prominent. Axillary cymules 
15-31-flowered, occurring along the upper 1/3-1/2 of the stems and com- 
monly terminating in a dense interrupted spike; primary peduncles ca 1.5 
mm; pedicels ca 1.0 mm long, inconspicuous; bracteoles narrowly lanceolate 
usually longer than the pedicels they subtend and frequently as long or 
longer than the calyx tube (short in University of Illinois collection 994; 
Mueller 2707, 29956). Calyx 5.0-8.0 mm (6.3 mm) long, the tube radically 
symmetrical, 3.0-4.0 mm (3.5 mm) long, ca 1.1 mm wide in diameter, 
somewhat dilated upwardly, puberulent or densely long pilose (Irving 669, 
Shreve & Tinkham 97794, Cummins 23-63), calyx teeth only slightly differ- 
entiated into upper and lower lips; the upper teeth scarcely connate, the 
lobes long subulate (occasionally triangular ), commonly exceeding the lower 
in length, 1.9-3.7 mm (2.6 mm) long, erect and outwardly spreading; the 
lower teeth subulate, 2.0-3.3 mm (2.5 mm) long, erect and outwardly 
spreading; the annulus well-exserted, undulating, ca 0.7 mm wide and seated 
at the juncture of the upper and lower teeth. Corolla blue-lavender, 14.0— 
16.0 mm (15.0 mm) long, pubescent within but not annulate; the tube and 
throat together 9.0-10.0 mm (9.5 mm) long, conspicuously dilated upwardly, 
about 3.0 mm wide above; the upper lip broadly to narrowly ligulate, 
emarginate, 2.0-3.5 mm wide, 5.0-6.0 mm long, the margins and apex 
revolute; lower lip 5.0 mm wide, 5.0-7.0 mm long. Nutlets ca 0.5 mm wide, 
1.0 mm long, the surface minutely areolate and weakly gluacous. Chromosome 
number, 27 = 36. 

Distribution (Fig. 3). Abundant in a variety of habitats from Galeana, 
Nuevo Leon, south to Zimapan, Hidalgo. It is especially prominent near the 


— 


242 


city of San Luis Potosi. The habitats range from upper montane pine forests 
where the plants occur in open, rocky, exposed areas to the relatively xerophy- 
tic habitats at lower elevations, where the plants abound on open, rocky ex- 
posed areas (e.g., stabilized talus slopes), 5,000-10,000 ft. Jul-Sep 


a 
= 
= 
5) 
3 
ty 
a 


im 


il 


r0eeee 


CT 


aul 


Fig. 3. Distribution of Hedeoma palmeri (closed circles), H. molle (open circles), 
H. patrinum (closed squares) and H. montanum (open squares). 


243 


TYPE: SAN LuIs POTOSI: chiefly in the region of San Luis Potosi, 22° 
N. Lat., Alt. 6,000-8,000 ft., 1878, Parry & Palmer 728 (Holotype, K! iso- 
types, F! LA! MICH! NY! PH! US!; phototype, LA! ). 

Hedeoma palmeri is quite variable and its populations form a mosaic of 
morphological patterns. Because of this, no attempt has been made to sub- 
divide the species into subspecific categories. 

The densely tomentose populations growing in the Galeana area (the most 
northern part of H. palmeri’s range) are especially difficult to treat. In the 
words of Epling (unpublished notes), “these specimens [from Galeana] 
are such an extreme as to provoke a review of this species.” I.M. Johnston, 
according to information on herbarium labels (Shreve & Tinkham 9779), 
thought the plants represented an undescribed species. However, these 
workers examined only a few extreme specimens which gave a somewhat 
distorted picture. When many representatives are brought together, the 
extremes grade one into another. 

Hedeoma palmeri is quite variable and its populations form a mosaic of 
morphological patterns. Because of this, no attempt has been made to sub- 
divide the species into subspecific categories. 


7. HEDEOMA PATRINUM Stewart, Rep. Spec. Nov., Beihefte 115: 45. 1939. 


Spindly, suffruticose perennials or semi-shrubs, 15.0-62.0 cm (31.4 cm) 
high. Shoots ascending, or commonly shortly decumbent, rhizomatous pro- 
ducing adventitious roots at the lower nodes, arising from a slender taproot, 
branching freely along the axes with the branches divergent at wide angles, 
cinerous above with short retrorsely curling hairs (occasionally spreading), 
glabrous below with exfoliating bark. Leaves deciduous from the lower por- 
tion of the stems, spreading, ovate or elliptic, 3.0-5.5 mm (4.2 mm) wide, 
5.2-10.0 mm (7.6 mm) long; margins entire or occasionally obscurely den- 
tate in the upper 1/2 of the blade (Purpus 4588); apex broadly acute or 
obtuse; based broadly cuneate or rounded, petiole 1.0-1.7 mm long; upper 
surface densely puberulent, margins and undersurface cinerous and tomen- 
tulose; nervation inconspicuous. Axillary cymules 1-3 flowered, well-spaced 
along the upper 1/4 of the branches; primary peduncles (and also secon- 
daries) conspicuous, 1.0-2.5 mm long; pedicels 1.7—3.0 mm long, cinerous 
or simply pubescent; bracteoles subulate, 1.0-2.0 mm long, shorter than 
the pedicels they subtend. Calyx 5.5-8.0 mm (7.0 mm) long; the tube 
4.0-5.0 mm (4.9 mm) long, tubular or slightly gibbous below, sparsely or 
densely pubescent; the upper teeth variously united from scarcely at all 
(Mueller 3302) to forming an upper lip 1.0-1.2 mm long (McVaugh 
23685), the lobes triangular (deltoid in McVaugh 23685) ca 0.4 mm wide 
at the base, 0.4-0.9 mm long, spreading and outwardly curling at their apices, 
pubescent on their outer faces but not ciliate or only scarcely so; the lower 
teeth ascending, recurved, more or less subulate, narrowly triangular, ca 0.5 
mm wide at the base 1.5-3.0 mm long, hirsute ciliate; annulus dense, more 


244 


or less included, ca 0.7 mm wide and seated at the juncture of the upper and 
lower teeth. Corolla lavender or blue, 15.0-16.0 mm (15.1 mm) long, 
glabrous in the throat, hirtellous pubescent in the tube forming a loosely 
defined annulus ca 5.0 mm wide; the tube and throat together 12.0-13.0 mm 
(12.5 mm) long gradually dilated upwardly, ca 3.0 mm wide above; the 
upper lip ligulate flat, straight, 1.8 mm wide, ca 3.0 mm long, very shallowly 
emarginate; the lower lip considerably larger than the upper, ca 6.0 mm 
wide, ca 5.5 mm long. Nutlets ca 0.7 mm wide, 1.4 mm long, the surface 
minutely areolate, and weakly glaucous. Chromosome number, 2” = ; 

Distribution (Fig. 3). Common in the isolated mountains of the Chi- 
huahuan Desert of central Mexico, mostly in oak woodlands, 6,500-8,100 ft. 
Jun-—Sep. 

TYPE: ZACATECAS. Concepcion del Oro, Sierra Madre Oriental 2600- 
2700 m, 18-19 Jul 1934, Pennell 17432 (Holotype, LA!; isotypes, F! MICH! 
NY! PH! US!) 

Since the original description of H. patrinum in 1939, additional collec- 
tions have considerably expanded the known distribution of this relatively 
rare species. Notable collections have been made in the Sierra Mojada by 
Mueller and in the state of Aquascalientes by McVaugh. These have revealed 
considerable intra-specific variation. The southernmost collection of this species 
(McVaugh 23685) is quite different from the more northern “typical” 
material. Plants of the former are not cinerous with curling hairs; rather 
they are pubescent with straight, usually spreading hairs; the upper calyx 
teeth are well united; and their deltoid lobes approach the foliaceous upper 
lip found in H. jucundum and H. piperitum. 

The taxonomic position of H. patrinum is not certain. Its shrubby habit 
and entire cinerous leaves indicate an affinity with H. molle, H. montanum, 
and H. palmeri. | have therefore, like Epling and Stewart who placed H. 
patrinum between H, montanum and H. molle, included it in the subgenus 
Poliominthoides. 


8. HEDEOMA MONTANUM Brandegee, Univ. Calif. Publ. Bot. 4: 188. 1913. 


Robust canescent, suffruticose perennials, 20.0—30.0 cm (28.0 cm) high. 
Shoots prolific, ascending and arising from a woody caudex or thickened tap- 
root, branching primarily in the lowermost regions with the branches acsend- 
ing at close angles, canescent with dense, short, spreading or curling hairs. 
Leaves deciduous from the lower stem regions at maturity, narrowly elliptical, 
6.0-10.0 mm (8.0 mm) long, 2.0-3.1 mm (2.6 mm) wide; margins den- 
ticulate to dentate. occasionally entire; apex acute, base narrowly attenuated 
and petiolate, the petiole ca 1.0 mm long, both upper-and lower-leaf sur- 
faces densely cinerous; nerves somewhat elevated. Axillary cymules well- 
spaced along the upper 1/3-1/2 of the stems, occasionally somewhat con- 
gested, typically 1-5-flowered; primary peduncles inconspicuous less than 
0.5 mm long; pedicels ca 2.0 mm long, cinereous, bracteoles subulate, ca 


245 


3.0 mm long, longer than the pedicels they subtend. Calyx 6.0-8.0 mm (7.1 
mm.) long; the tube radially symmetrical or nearly so, ca 1.2 mm in diameter, 
cinereous with tightly curled puberulent hairs evenly distributed from the 
calyx base to the teeth apices; the calyx teeth only slightly differentiated into 
upper and lower sets and convergent at maturity; the upper teeth scarcely 
united for a 1/10 of their length, the upper lip 1.0-2.5 mm long, the lobes 
slightly shorter than the lower, subulate, ca 0.3 mm wide at the base 0.8—2.5 
mm long; lower teeth subulate, recurved but convergent with the upper at 
maturity, ca 0.5 mm wide at the base, 1.0-2.5 mm long, hirtellous-ciliate; 
annulus extremely dense, well-exserted, ca 1.0 mm wide, seated approximately 
at the juncture of the upper and lower teeth. Corolla blue-lavender, ca 15.0 
mm long, densely annulate at the juncture of the tube and throat; the tube 
and throat together ca 12.0 mm long, widening upwardly, ca 3.0 mm wide 
above; the upper lip ligulate, ca 2.5 mm wide, ca 3.0 mm long, straight, 
slightly emarginate and concave; the lower lip ca 4.0 mm long, 4.0 mm 
wide. Nulets ca 0.5 mm wide, ca 1.5 mm long, the surface minutely aerolate 
nearly smooth and glaucous. Chromosome number, 27” = 36. 

Distribution (Fig. 3). Arid mountains of southern Coahuila in the Chi- 
hauhuan Desert. Frequent on open, exposed, limestone ridges and hillsides, 
commonly associate with Agave lecheguilla. 7,000-8,000 ft. Jun; Sept-Oct. 

TYPE: COAHUILA. Sierra de la Paila, 7,000-8,000 ft. Oct 1910, Purpus 
4964 (Holotype, UC!; isotypes, BM! F! GH! LA! US! phototype, LA! ). 

Hedeoma montanum is a rather rare species being known by only a few 
collections from a relatively restricted area of southern Coahuila. It is most 
closely related to H. molle. 

9. HEDEOMA MOLLE Torr., Bot. U.S. Mex. Bound. Surv. 129. 1859. 

Poliomintha mollis (Torr.) A. Gray, Proc. Am. Acad. 8: 365. 1872. 

Robust, suffrutescent perennials or semi-shrubs, 16.0-55.0 cm (35.7 cm) 
high. Shoots numerous and long, ascending, tomentose-canescent or tomen- 
tulose-cinerous with short profusely branching hairs, arising from a woody 
caudex; branching almost completely confined to the basal region, the upper 
branches, when present, short and arising at close angles. Leaves narrowly 
to broadly elliptical to oval, highly variable, 3.0-10.0 mm (6.1 mm) wide, 
8.5-22.0 mm (14.1 mm) long; margins entire or obscurely crenate; apex 
obtuse to somewhat acute; the base broadly or narrowly attenuated, petiolate, 
the petiole 0.5-4.0 mm (1.8 mm) long; surfaces evenly tomentose or 
tomentulose with intertangled, profusely branching hairs; nerves elevated. 
Axillary cymules 3-15-flowered (typically 10), occurring along the upper 
1/3 of the stems, well-spaced or more commonly congested forming a dense 
but interrupted spike of cymes up to 7.0 cm long; primary peduncles promi- 
ent, 2.0-3.5 mm long, equal to or exceeding the length of the primary 
pedicel; pedicels 1.5-3.0 mm long, tomentose; bracteoles subulate as long as 
or longer than the pedicels they subtend. Calyx 5.2-7.0 mm (6.0 mm) long; 


246 


the tube nearly iene: symmetrical, or slightly upwardly dilating, ca 1.1 
mm wide in diameter, 4.2-6.0 mm long, occasionally slightly gibbous 
or curved (Correll 30389 and Hinckly & Warnock 46876, respectively ), 
tomentose with highly branched evenly distributed hairs; the calyx teeth 
only slightly differentiated into upper and lower sets; the upper calyx 
teeth connate for ca 1/3 of their length, the lobes subulate or very nar- 
rowly triangular, ca 0.7 mm wide at the base, 0.3-0.8 mm (0.5 mm) long, 
slightly laterally nadine or convergent, erect to sharply outwardly curling, 
the outer face and margins tomentose, the inner face covered with short pilose 
hairs as an extension of the prominent annulus; the lower teeth subulate, 
0.3-0.5 mm wide, at the base, 0.6-1.3 mm (0.9 mm) long, clothed in a 
branched hair tomentum; annulus exserted, extremely dense, with unbranched, 

ilose hairs, ca 0.7 mm wide, seated approximately at the juncture of the 
poorly defined upper and lower lips. Corolla lavender 13.2-14.0 mm (13.6 
mm) long, sparsely pubescent in the throat and in the lower tube but not 
forming a well-defined annulus; the tube and throat together 11.0-12.0 mm 
(11.5 mm) long conspicuously dilated upwardly, 3.0-3.5 mm wide above; 
the upper lip broadly ligulate, conspicuously emarginate, ca 1.9 wide, ca 
2.2 mm long; the lower lip 4.0-5.0 mm wide, 3.0-4.0 long. Nutlets narrow, 
ca 0.3 mm wide, ca 1.2 mm long, the surface minutely areolate, and weakly 
glaucous. Chromosome number, 27 = 36 

Distribution (Fig. 3). Restricted to rock slopes and ledges of the Davis, 
Glass, Chisos, Chinati, and Sierra Vieja mountains of Trans-Pecos, Texas. 
1,000-7,000 ft. Jul-Nov. 

TYPE: TEXAS. Presidio Co., w of Alpine, cliffs near Puerta de Paysano, 
Sep. Bigelow sn. (Holotype, NY! isotype, NY! ). 

Hedeoma molle occurs in widely separated populations, however, Judging 
from its representation in herbaria, it is abundant where it occurs. Specimens 
from the northern portion of its range (Davis Mountains: Madera Canyon, 
Cory 14042; Ajuga Canyon, Correll 33341) are unusually large-leaved and 
show only a slight differentiation between the upper and lower calyx teeth. 

ne specimen (Correll 33341) has gray-green leaves which measure 24.0 
mm long and 16.0 mm wide. To the west, in the more arid Chinati Mountains, 
the leaves are quite diminutive, measuring ca 7.5 mm long and 2.5 mm wide, 
and there is a marked differentiation of the upper and lower calyx teeth. 
(McVaugh 7453 

H, molle is most t closely related to H. montanum. 


III. SUBGENUS SATUREJOIDES, Irving, subg. nov. 


rbae annuae vel perennes raro suffrutescentes, 5.0-G60.0 cm altae; folia ovata, 
elliptica vel late elliptica, oblonga, lanceolata vel linearia, 3.0-28.0 mm longa, 1.0— 
16.0 mm lata, margine integra vel dentata, apice obtusa vel acuta, ad basin petiolata 
vel sessilia; cymulae 1—20-florae, remotae; calyx 4.0-10.0 mm longus, valde zygo- 
morphus; calycis tubus leviter gibbosus vel saccatus; corolla varia (minuta vel longi 
tubularis vel magna conspicuaque), 53.5-20.0 mm longa; mericarpum oblongum, 


247 


uplo longiorum quam latiorum, brunneum (ad maturitatem), areolatum glaucum 
mucilaginum (in stat humectato); chromosomatum numerus, 2” = 34, 36, 44, 72. 


Herbaceous annuals or perennials, occasionally suffrutescent, 5.0-60.0 cm 
high. Shoots solitary to numerous, strict, ascending or decumbent, sometimes 
rhizomatous sending out adventitious roots from the lower nodes, arising 
from a taproot, trailing rootstock, or woody caudex. Leaves membranous, 
oval, linear, 3.0-28.0 mm long, 1.0-16.0 mm wide, margins entire to coarsely 
dentate, apex obtuse or acute, base subsessile to petiolate. Cymules 1—20- 
flowered, usually well-spaced along the stems, occasionally ne toward 
a apices forming a short “spike.” Calyx zygomorphic; the tube slightly 

ous to saccate, 4.0-10.0 mm long, variable in pubescence; the calyx 
an well-differentiated into upper and lower sets; the upper teeth variously 
connate forming as upper lip; the lobes subulate, deltoid, variously ciliate, 
commonly spreading and reflexed; the lower teeth subulate and recurved, 
ciliate; annulus well developed. Corolla usually lavender-blue, variable from 
minute (H. hispidum) to long tubular (H. costatum) to large and showy 
(H. pulcherrimum. H. diffusum, and H. byssopifolium), 5.5-45.0 mm long. 
Nutlets oblong or ovate in outline usually twice as long as wide, occasion- 
ally orbicular, brown at maturity, the surface areolate or foveate, glaucous 
and mucilaginous when moistened. Chromosome numbers, 27 = 34. 36, 44, 
i 

Type species: HEDEOMA COSTATUM Gray. 

The subgenus Satwrjoides is subdivided into two sections, Alpine and 
Saturejoides. \t is the largest subgenus in Hedeoma and its species are 
reticulately related one to another 

e most homogeneous unit within the subgenus is the section Alpine. 
It is unified by a number of features. Section Saturejoides, on the other hand, 
is a more diversified group which presents greater difficulty in characteriza- 
tion and circumspection. 


A. SECTION ALPINE Irving, sect. nov. 


e perennes 1.0-5.0 dm altae; surculi multi ascendentes vel decumbentes; 
asi 


ee fere sphaeroideum vel ovoideum, diametro suo duplo longium vel 


brev 


Perennial herbs 10.0-50.0 cm tall. Shoots few to numerous, ascending or 
decumbent and rhizomatous, sending out adventitious roots from the lower 
nodes, arising f slender taproot. Leaves glandular, punctate on 


or obscurely serrate, apex obtuse, base typically tapered and_ subsessile. 
Cymules 3-7-flowered. Calyces variable between species, conspicuously 
zygomorphic, 4.0-8.0 mm long; the tube tubular or saccate; the upper teeth 
united to form a conspicuous upper lip, the lobes narrowly triangular or 


248 


deltoid; the lower teeth narrowly seo or subulate, recurved. Corolla 
showy, 5.0-17.0 mm long, the upper lip obcordate, commonly reflexed, 
the lower lip various, spreading. Nutlets ae or ovate in outline, nearly 
spherical, ovoid, ovate in cross section, ca as long as wide, brown at maturity, 
the surface areolate. Chromosome number, 27 = 44. 

ype species: HEDEOMA PIPERITUM Benth. 

A number of similarities, in addition to the morphic ones, unite the species 
of section Alpine: (1) they are found at the higher elevations, (2) they 
are the only representatives of the genus occurring in the Sierra ee 
Occidental of Mexico and, except for the South American species, H. ; 
galaefolium, they are all found in this western range; (3) they all oe 
a chromosome number of 27 = 44. 

From these diverse lines of evidence, it is difficult to deny the naturalness 
of section Alpine. The weakest evidence for the groups naturalness comes 
from morphology, since the combination of characters which unite them 
are subtle. 


10. HEDEOMA POLYGALAEFOLIUM Benth., Lab. Gen. et Sp. 367. 1834. 

Small wiry perennial herbs, ca 10.0 cm high. Shoots decumbent and rhizo- 
matous forming adventitious roots from the lower, buried nodes, branching 
freely along the axis; glabrate or with a few minute retrorsely curved hairs. 
Leaf pairs crowded with internodes uniformly ca 1.0 mm long. Leaves 
ascending, readily deciduous from the lower portions of the older stems, 
chartaceous becoming folded with drying, oblong to oblanceolate, 2.0—2.5 
mm (2.3 mm) wide, 6.5-7.0 mm (6.8 mm) long, margins entire, apex 
obtuse, base narrowly attenuate and subsessile to a decurrent base; leaves 
glabrous throughout; nervation consisting solely of a prominent midrib. 
Cymules 1-3-flowered, congested toward the branch apices; peduncles 2.0- 
2.5 mm long; pedicels 4.5-5.0 mm long, puberulent with retrorsely curved 
hairs; bracteoles lanceolate shorter than the pedicels they subtend. 
conspicuously zygomorphic, ca 8.0 mm long; the tube tubular—funnelform 
4.5 mm long not at all gibbous below, dilating and flaring upwardly, ca 2. 
mm in diameter above, glabrous; the upper teeth united for ca 2/3 of their 
length forming an upper lip ca 2.5 mm long, pubescent within, the lobes 
narrowly triangular, acuminate at the apex, ca 0.5 mm wide at the base, 
ca 1.5 mm long, slightly spreading and reflexed, the margins glabrous; the 
lower teeth triangular—subulate, recurved, ca 3.0 mm long, hirtellous—ciliate; 
annulus dense ca 1.2 mm wide, seated ca 1.0 mm below the juncture of 
the upper and lower teeth but extended into the faces of the upper teeth, 
more or less included. Corolla ca 10.0 mm long. Nutlets 0.8 mm wide, ca 
0.8 mm long, brown at maturity, the surface smooth and aerolate. Chromo- 
some number, unknown. 

aa ecaalner (Fig. 12). Brazil, province Rio Grande do Sul. 

ZIL. 1841, Sellow s.n. (Holotype, K! isotype LA!: phototypes, 
US! ). 


= 
“ 


LA! NY! a 


11. HEDEOMA BELLUM (Epl.) Irving, Brittonia 22: 345. 1970. 
Hesperozygis bella Epl., Bull. Torr. Bot. Club 67: 512. 1940. 


Small wiry perennial herbs, 15.0-30.0 cm in height. Shoots few to numer- 
ous, typically shortly decumbent (up to 7.0 cm) and rhizomatous sending 
out adventitious roots from the lower, buried nodes; sparsely branching; 
stems densely puberulent with the hairs spreading above and _ antrorsely 
curving below. Leaves spreading or cernuous, readily deciduous from the 
lower portions of the older stems, chartaceous, the margins often becoming 
revolute upon drying, oblanceolate-elliptical, 3.4-4.0 mm (3.6 mm) wide, 
10.0-17.0 mm (15.0 mm) long, margins entire, apex obtuse, base gradually 
attenuate and subsessile; glabrous above, strigulose on the margins and 
midrib; nerves consisting of a prominent midrib and inconspicuous second- 
daries. Cymules 2-3-flowered, congested in the upper 1/4 of the stem 
forming a short terminal “spike”, peduncles distinct but short, ca 1.5 mm 
long; pedicels 3.0-4.0 mm long, hirtellous with hairs spreading. Calyx 
conspicuously zygomorphic, ca 8.0 mm long; the tube ca 5.0 mm long not 
gibbous below, dilating upwardly, ca 2.5 mm wide above, sparsely hirtellous; 
upper teeth connate for approximately 2/3 of their length forming a con- 
spicuous upper lip, ca 2.5 mm long, pubescent within with short pilose 
hairs, the lobes more or less deltoid or semi-ovate, conspicuously apiculate 
at the apex, ca 1.0 mm wide at the base, 1.0 mm long, sharply reflexed, strig- 
ulose-ciliate; the lower teeth sharply recurved, broadly triangular and taper- 
ing into a long aristate apex, ca 14 mm wide at the base, 3.4-3.8 mm 
long, hirsute-ciliate; annulus dense ca 1.0 mm wide, seated ca 1.0 mm 
below the upper lip, included. Corolla ca 12.0 mm long, pink and red dotted 
in the throat, minutely hirtellous within; the tube ca 10.0 mm long, grad- 
ually dilating upwardly; the upper lip obcordate, emarginate, straight or 
slightly reflexed, ca 2.0 mm long; the lower lip ca 5.0 mm broad, ca 3.0 mm 
long. Nutlets ca 0.8 mm wide, ca 0.8 mm long. Chromosome number, un- 
known. 

Distribution (Fig. 4). Hedeoma bellum is known only from the type 
sae where, according to label data, it occurs on damp stream banks. i 
T : JALISCO. Sierra Madre Occidental, San Sebastian, east of Arro 
Santa ee 1500 m, 18 Jan 1927, Nee 1514 (Holotype, UC! oe 

F! NY!) 

Vegetatively, especially in the leaves and in the inflorescence, H. bellum is 
similar to H. patens. It might also be noted that the common name of this 
species “oreganum’, undoubtedly derived from its characteristic odor, is 
also the vernacular name given to H. patens. In the finer details of the calyx, 
one of the most elaborate in the genus Hedeoma, it is unique among the 
Nozth American representatives, but is similar to the equally restricted South 
American species, H. polygalaefolium. 


250 


12. HEDEOMA JUCUNDUM Greene, Pitt., 1: 156. 1888. 


Dense, caespitose perennial herbs, 10.0-15.0 cm in height. Shoots typi- 
cally numerous, widely ascending or shortly decumbent; branched profusely 
at the base, and frequently producing adventitious roots from the lower 


eee Sse 
fi 


ee 


l 


eenera 


Tl 


~ : UF 


‘ig. 4. Distribution of Hedeoma patens (open circles), H. floribundum (open 
square), H. jucundum (closed circles), H. bellum (open triangle) and H. piperi 
(closed squares). 


251 


nodes, unbranched above or sparsely so; stems moderately to densely hirtel- 
lous above with retrorsely curving or occasionally spreading hairs, becom- 
ing puberulent below. Leaves spreading or ascending, chartaceous with the 
margins frequently becoming revolute Ee drying, ovate to narrowly 
elliptical, 4.0-6.0 mm (5.3 mm) wide, 8.0-12.0 mm (11.0 mm) long, 
margins entire to ee serrate, apex “obtuse, base shortly attenuate 
and subsessile; glabrous above, strigulose on the margins and the midrib 
below; nerves not conspicuously raised. Cymules 3-5-flowered, well-spaced 
along the upper 1/2 of the stems; peduncles inconspicuous less than 1.0 
mm in length; pedicels 3.0-5.0 mm long, moderately hirtellous to puberu- 
lent. Calyx 6.0-7.0 mm (6.5 mm) long; the tube 3.5-4.5 mm (4.0 mm) 
oe slightly gibbous below for ca 1/2 of its length, the distended region 

1.3 mm wide, dilating upwardly to ca 1.5 mm wide, sparsely to moder- 
ere hirsute-hirtellous; the upper calyx teeth connate for ca 1/2 of their 
length forming a pronounced, laterally rounded upper lip ca 2.3 mm long, 
the margins outwardly extended, the lobes triangular, ca 0.8 mm wide at the 
base, ca 1.2 mm long, sharply reflexed and laterally spreading, hirsute— 
ciliate; the lower teeth recurved, narrowly triangular—subulate, ca 8.0 mm 
wide at the base, ca 3.0 mm long, ciliate; the annulus dense ca 1.0 mm wide 
and arched upwardly into the connate portion of the upper lip, included. 
Corolla 11.0-12.0 mm (11.5 mm) long, white or pink with violet mark- 
ings on the lower lip, glabrous within; the tube 9.0-10.0 mm long weakly 
dilating upwardly, ca 2.0 mm wide above; the upper lip obcordate in outline, 
ca 2.5 mm wide, 3.0 mm long, reflexed; the lower lip broad, ca 7.0 mm wide, 
ca 5.0 mm long. Nutlets ca 0.8 mm wide, 1.0 mm long, the surface minutely 
areolate, not glaucous. Chromosome number, 27 = 44 

Distribution (Fig. 4). Sierra Madre Occidental, common in the pine 
forests west of the village of El Salto, Durango. 7000-9000 ft. Aug—Oct. 
TYPE: DuRANGO. From the Sierra Madre, west of Durango, 8100 ft. Sep— 
Oct. 1881, Forrer sn. (Holotype, ND! isotypes, F! GH! LA! NMC! NY! 
PH! POM! UC! US!). 

Hedeoma jucundum is morphologically uniform and is most closely re- 
lated to H. piperitum. 


13. HEDEOMA PIPERITUM Benth., Lab. Gen. et Sp. 730. 1835. 
Cunila piperita Moc. et Sesse ex Benth., Lab. Gen. et Sp. 730. 1835. 


Loose, wiry, perennial herbs, somewhat caespitose, 10.0—20.0 cm tall. Shoots 
few to numerous, widely ascending or shortly decumbent; branching from 
the base, and frequently producing adventitious roots from the lower nodes, 
upper shoots typically unbranched or bearing a few short branches; stems 
densely hirsute with spreading hairs in the upper regions, hirtellous to 
puberulent below. Leaves spreading or drooping, membranous, broadly ovate 
to sub-orbicular, 5.0-13.0 mm (8.0 mm) wide, 7.0-18.0 mm (12.5 mm) 
long, margins entire or obscurely serrate, apex obtuse, base shortly attenuate 


222 


and subsessile; glabrous above, strigulose below on the nerves and margins; 
nerves not eae raised. Cymules 3—7-flowered well-spaced along 
the upper 1/2 of the stems; peduncles inconspicuous less than 1.0 mm in 
length; pedicles 2.0-5.0 mm_ long, hirsute. ee strongly ee reaga 
5.5-6.5 mm (6.0 mm) long; the tube 3.0-4.0 mm (3.5 mm) long, 
tinctly saccate below for ca 2/3 of the tube’s ne the distended region 
1.5-2.0 mm wide, sparsely to moderately hirsute; the upper calyx teeth 
united well over 1/2 of their length to form a reflexed, laterally rounded, 
upper lip ca 2.0 mm long, the margins pba flaring, hirtellous—ciliate, 
glabrous or pubescent on the inner face, the lobes semi-ovate to deltoid, 
ca 0.7 mm wide at the base, 0.7 mm ion. ciliate; the lower teeth recurved, 
narrowly triangular, ca 0.8 mm wide at the base, ca 2.3 mm long, hirsute— 
ciliate; annulus moderately dense, ca 0.5 mm wide and arched somewhat 
into the upper lip, included. Corolla, 7.0-8.5 mm (8.0 mm) long, white 
with maroon markings on the lower lip, puberulent to glabrous within; the 
tube 5.5-7.0 mm (6.5 mm) long gradually dilated upwardly, the upper lip 
obcordate in outline, ca 2.3 wide, 1.5-2.0 mm long, straight or very slightly 
reflexed; the lower lip broad, 3.5-4.5 mm wide, ca 3.0 mm long. Nutlets 
0.8 mm wide, ca 1.0 mm long, the surface minutely areolate, not glaucous. 
Chromosome number 22” = 44, 

Distribution (Fig. 4). Pine forests of central Mexico, 6,000-10,000 ft. 
Particularly abundant in sunny, open slopes associated with rocky lava or 
volcanic ash. Aug—Oct. 

YPE: Mexico. Mocino et Sesse (lectotype, here designated BM!; iso- 
types, F! MA, photoisotypes, LA! TEX! ). The specimen bearing the her- 
barium name C. piperita Moc. et Sesse was observed by Bentham in the 
Dunant herbarium which obtained the type from the herbarium of Pavon, 

Hedeoma piperitum is quite distinct and relatively homogeneous through- 
out its narrow range. The salient morphological features which distinguish 
this species are its caespitose, broadly ovate-suborbicular leaves, and saccate 
calyx with its well-developed, expanded, upper lip. It is closely related to 
H. jucundum. Examination of the westernmost populations of H. piperitum 
(Cheron, Michoacan) reveal several subtle morphological departures from 
the eastern populations around Mexico City. The western populations vary 
in the direction of H. juwcundum; that is, they have a more robust habit; 
thick, oes leaves ( Beal 4); and retrorsely curling stem pubescence 

13) 


(Beal 
14. HEDEOMA PATENS Jones, Contrib. West. Bot. 12: 70. 1908. 


Perennial herbs 20.0-55.0 cm high, averaging 30.0 cm. Shoots numerous, 
herbaceous above, decumbent (often one-sidedly so), woody below, branch- 
ing freely, densely puberulent or hirtellous with hairs retrorsely curling. 
Leaves variable, laxly spreading to drooping, membranous, broadly to narrowly 
elliptical to oblanceolate, 2.0-8.0 mm (4.0 mm) wide, 5.0-22.0 mm (12.0 


— 


253 


mm) long, margins entire or obscurely crenate, apex obtuse, base narrowly 
attenuate, short petiolate (seldom exceeding 1.5 mm) or subsessile; glabrous 
above, strigulose below and on the margins; nerves inconspicuous. Cymules 
well-spaced along the upper 2/3 of the stem with 3-7 flowers per cymule 
(typically 3-5), peduncles obscure to ca 2.0 mm long; pedicels 3.0-5.0 mm 
long, typically exceeding or equaling the length of the calyx tube, densely 
puberulent. Calyx 4.0-655 mm (6.0 mm) long, moderately zygomorphic; 
the tube 3.5-5.0 mm (4.0 mm) long, slightly gibbous below for 1/3-1/2 
its length, the distended region 0.7-1.6 mm wide, puberulent to hirtellous 
with hairs strongly antrorsely curling; the upper calyx teeth connate for 
1/3-1/4 of their length, forming a short upper lip 1.2 mm long, the 
lobes deltoid, ca 0.5 mm wide at the base, ca 0.5 mm long, laterally spread- 
ing, weakly reflexed, strigulose-ciliate; the lower calyx teeth recurved, sub- 
ulate highly variable as to length, ca 0.5 mm wide at the base, 1.2—2.3 mm 
(1.8 mm) long, typically hirsute—ciliate, annulus dense, ca 0.5 mm wide, 
typically seated ca 0.5 mm below the juncture of the upper and lower teeth 
and included, occasionally exserted (Knobloch 5407). Corolla 7.0-9.0 mm 
(8.0 mm) long, pink, glabrous within; the tube 5.0-7.0 mm (6.0 mm) 
long, the throat scarcely dilated upwardly (ca 2.0 mm wide above); the 
upper lip obcordate in outline, ca 2.0 mm broad at the widest point, ca 2.6 
mm long, reflexed; the lower lip, ca 5.0 mm wide, ca 4.5 mm long. Nutlets 
not quite twice as long as broad, 0.5 mm wide, ca 0.9 mm long, the surface 
minutely areolate, and weakly glaucous. Chromosome number 2” = 44. 

Distribution (Fig. 4). Pine forests of the Sierra Madre Occidental. 5000- 
7000 ft. Aug—Oct. 

TYPE: CHIHUAHUA. Sierra Madre Mts., “Guayanopa Canyon”, 5000 ft., 
23 Sep 1903, Jones s.n. (Holotype, POM! phototype, LA! ). 

Hedeoma patens is an exceedingly variable species. The specimens which 
I have examined seem to form a transition between H. hyssoptfolium and 
Hl. pulcherrimum (SECTION Saturejoides), and H. jucundum and H. piperi- 
tum (SECTION Alpine). 

Epling and Stewart (1939) placed H. patens close to H. pulcherrimum. 
Admittedly, they resemble each other in many respects, particularly in habit 
and vegetative morphology. They can be readily separated from each other, 
however, by the important details of the calyx and corolla. 


15. HEDEOMA FLORIBUNDUM Standley, Rep. Spec. Novy., Beihefte 115: 31. 
192). 

Perennial herbs 20.0—32.0 cm high. Shoots woody below, ascending or de- 
cumbent, branching freely along the axis with the branches ascending at 
close angles, clothed below in long spreading pubescence, above puberulent 
with hairs tightly retrorsely curling. Leaves laxly spreading to cernuous, 
membranous, narrowly elliptical or oblanceolate, 4.0-5.0 mm (4.7 mm) 
wide, 15.0-18.5 mm (15.9 mm) long, margins entire or obscurely crenate, 


254 


apex obtuse, base narrowly petiolate with the petiole ca 1.0 mm _ long, 
glabrate above, puberulent below; nerves inconspicuous. Cymules aggregated 
toward the branch apices forming a short terminal spike, ca 4.0 cm long, 
3-7-flowered (typically 3-5), peduncles and pedicels contorted; peduncles 
inconspicuous to ca 1.0 mm long; pedicels 2.0-2.5 mm long, shorter than 
the calyx tube, puberulent. Calyx 3.0-3.5 mm oe Uses Bel aaa 
puberlulent with antrorsely curling hairs, the tube 2.5-3.0 mm long. The 
tube weakly distended ca 1.0 mm wide; the upper and loca: teeth only 
slightly differentiated; the upper teeth united for ca 1/3 of their length 
forming an upper lip ca 0.5 mm long, the lobes erect, deltoid ca 0.4 mm 
wide at the base, ca 0.4 m long; the lower teeth ca 0.5 mm wide at the 
base, 0.5 mm long; annulus included, ca 0.5 mm wide and seated just below 
the juncture of the upper and lower lips. Corolla 5.0-6.0 mm long, glabrous 
within; the tube 4.0-5.0 mm long gradually widening upwardly, upper lip 
obcordate, 1.0-1.2 mm wide, 1.0-1.5 mm long; the lower lip ca 2.5 mm 
wide, 2.5 mm long. Nutlets 0.5 mm wide, 0.9 mm long, the surface minutely 
areolate, glaucous. Chromosome number, unknown. 
Distribution (Fig. 4). Pine forests along the Rio Mayo in the state of 

Chihuahua, ana . Oct 

YPE: CHIHUAHUA. Cans Quicorichi, Rio Mayo, 6 Oct 1935, Gentry 
1938 (Hol eae F!; isotype, 

he vegetative features of H. floribundum are similar to those of H. patens, 
and they have similar distributions. The salient differences between these 
two species reside in the inflorescence and calyx. Unlike H. patens, the 
inflorescence of H. floribundum is a dense terminal spike and the calyx 
shows a greater trend toward radical symmetry, both in the tube and in the 
disposition of its teeth. 


B. SECTION SATUREJOIDES, Irving, sect. nov. 


Herbae annuae vel cee: raro suffrutescentes, 5.0-60.0 cm altae; folia varia, 
margine integra vel aepe serrata vel dentata, ie basin petiolata vel subsessilia; 
4 


a ened : *s; coroll: 
vel magna aera ue, 53.5-20.0 mm ieee mericarpium oblongum, circa duplo 
longiorum quam latiorum brunneum (ad maturitatem), areolatum vel foveolatum. 
Herbaceous annuals or perennials, occasionally suffrutescent and woody 
below, 5.0-60.0 cm high. Shoots solitary to numerous ascending or decum- 
bent, occasionally rhizomatous and forming eee roots. Leaves glan- 
dular punctate, variously shaped. Margins entire, serrate, or dentate, apex 
acute or narrowly obtuse, base petiolate or subsessile. Ce 1—20-flowered. 
Calyx 4.0-10.0 mm long, zygomorphic; the tube slightly gibbous to distinctly 
saccate; the upper teeth variously connate, the lobes subulate or narrowly 
triangular; the lower teeth subulate and recurved; both upper and lower 
teeth variously ciliate. Corollas inconspicuous to large and showy, 5.5—45.0 
mm long; the upper lip ligulate, straight or reflexed, flat or sub-galeate; 


255 


the lower lip spreading. Nutlets oblong, at least twice as long as wide, ovate- 
triangular in cross section, brown at maturity, the surface areolate or foveate, 
glaucous. Chromosome numbers, 2” = 34, 36, 7 

TYPE: HEDEOMA COSTATUM Gray. 


16. HEDEOMA PULCHERRIMUM Woot. & Standl., Contr. U.S. Natl. Herb. 16: 
168. 1913. 

Robust, perennial herbs, 15.0-40.0 cm in height (typically between 20.0 
and 30.0 cm). Shoots few to numerous, ascending or very shortly decumbent, 
woody below, herbaceous above; branching frequently in the lower half, 
the branches long and widely ascending, stems below clothed in villous 
pubescence, above sparsely to densely puberulent with the hairs tightly 
retrorsely curling. Leaves thick membranous, elliptical-oblong, 8.0-18.0 mm 
(13.4 mm) long, 2.0-6.5 mm (4.2 mm) wide, margins entire to obscurely 
serrate, base attenuated, short petiolate or subsessile, apex obtuse; leaves 
glabrous above, strigulose below and on margins; nerves conspicuous. Cymules 
well-spaced along the upper 2/3 of the stems with 3-7 flowers per cymule 
(typically 3-5); peduncles short; 1.0-2.0 mm long; pedicels 2.0—3.5 mm 
long, minutely puberulent; bracteoles short. Calyx 6.0-7.0 mm (6.6 mm) 
long; the tube 3.0-5.0 mm (4.0 mm) long, slightly gibbous below for ca 
1/2 of the tube’s length, the distended region ca 1.2 mm wide, typically 
moderately hirsute; the upper calyx teeth connate for 1/3-1/4 of their 
length forming an upper lip, 1.5-2.0 mm long, the lobes very narrowly 
triangulate-lanceolate (nearly subulate), ca 0.3 mm wide at the base, ca 

mm long, laterally spreading and reflexed (with maturity becoming 
erect and convergent with the lower teeth), hirsute or hirtellous—ciliate, 
the lower calyx teeth recurved, subulate, ca 0.5 mm wide at the base, ca 
2.7 mm long; annulus dense, ca 1.0 mm wide, seated at the juncture of 
the upper and lower teeth, exserted. Corolla 10.0-14.0 mm (12.4 mm) long, 
blue, sparsely villous within the throat and densely short pubescent in the 
tube forming a well-defined annulus, 2.0-3.0 mm in width; the tube 8.0- 
10.0 mm (9.0 mm) long, the throat strongly dilated, ca 5.0 mm wide; the 
upper lip broadly ligulate, slightly emarginate, 3.0-3.3 mm broad, 3.0-4.0 
mm long, straight (not upwardly reflexed); the lower lip broad ca 7.0 mm 
wide, ca 5.0 mm long. Nutlets ca 0.6 mm wide, 1.2 mm long, the surface 
minutely areolate, weakly glaucous. Chromosome number 27 = 36 

Distribution (Fig. 5). Restricted to the Sacramento and White Mountains 
of south central New Mexico where it is common in Pine or Spruce-Fir 
forests, Poca) in disturbed areas; 5000-9000 ft. Jul-Sept. 

TYPE: New Mexico. Lincoln Co., collected in the White Mountains, 
6500 fet., 30 July 1897, Wooton 241 (Holotype, US! isotypes, DS!, LA! ND! 
NMC! NY! POM! UC! US!). 

H. pulcherrimum has been difficult taxonomically to place. In the original 
description, Wooton and Standley state: “This species [H. pulcherrimum] 
seems to be on the dividing line between the two principal groups of species: 


256 


its upper calyx lobes are not foliaceous expanded as in one group [Sect. 
Alpine} nor narrowly subulate as in the other but are triangulate-lanceolate 
and hispid-ciliate like those of the second group [Sect. Satwrejoides}". Al- 
though recognizing its transitional nature, they thought it closest to H. 
jucundum. Epling and Stewart however, aligned H. pulcherrimum with H. 


I have treated H. pulcherrimum as being most closely related to H. 
drummondii. Except for characters of the calyx, the western elements of the 
H. drummondiu complex are identical, or nearly so, to H. pulcherrimum. 
Moreover, natural hybridization occasionally occurs between these species 
with the F, hybrids vigorous and partially fertile (ca 20%). 


Tes, 


we poeoee eee 


Te meme 
im eae ~~"... 


° 


fe} 
ween 


lo 


7 


wet ee 
°. 


_- 
- 
-< 


fone) 


-— 
— 
= 


-<. 
-. 


4 


Wot, 


pay 


ea! 


Fig. 5. United States distribution of Hedeoma pulcherrimum (squares), H. 
drummondti (open circles), H. reverchonit var. serpyllifolium (triangles), and H.+r. 
var. reverchonii (closed circles) 


27 


17. HEDEOMA DRUMMONDJ Benth. Lab. Gen. et Sp. 1836. 
Hedeoma ciliatum Nutt., Journ. Acad. Nat. Sci. Phil., N.S. 1: 47. 
TYPE: NEW MEXICO: se Fe Co., Santa Fe, Gane 5M, eee GH! 
isotype, K!) 
Hedeoma ovatum A. Nels., Bull. Torr. Bot. Club 31: 245. 
TYPE: a Pole Creek, 30 Jun 1895, Nelson ae sees RM!; 
isotype, GH 
ie ee Rydb., Bull. Torr. Bot. Club 36: 695. 1909. 
TYPE: NEBRASKA. Banner Co., Cliffs of Canyons, Aug 1890, Rydberg x297 
(Holotype, NY!) 

Hedeoma Senee aie Flora Rocky Mts. 750. 1917. 
Based on H. longifloru 

Hedeoma drummondi Bal var. seta Irving, Britt. 22: 342. 1970 
TYPE: MEXICO, San Luis Pot 7.8 mi w of Guadalcazar, 10 ‘Ate 1965, 
Irving 632 (Holotype, TEX!; ae MONTU! ) 

Annuals or robust perennials, 15.0-45.0 cm high. Shoots few to numer- 
ous, shortly decumbent and/or frequently spreading upwardly at wide angles, 
branching freely along the lower 1/2 to 2/3 of the stems with the branches 
typically long, straight, and spreading at angles of ca 45°; stems densel 
pubescent or puberulent above with the hairs tightly retrorsely curling, 
glabrate below. Leaves bright green, cernuous, thick membranous, linear 
to elliptic-oblong, 1.2-4.0 mm (2.2 mm) wide, 5.0-11.0 mm (7.7 mm) 
long, entire, obtuse at apex, narrowly attenuated and short petiolate or sub- 
sessile, glabrous or sparsely pubescent above, densely pubescent below; 
nerves inconspicuous. Cymules somewhat crowded along the upper 1/2-3/4 
of the stems with 3-7 flowers/cymule,; peduncles inconspicuous to ca 1. 
mm long; pedicels 2.0-3.5 mm long, pubescent; bracteoles subulate ca 1/2 
as long as pedicels. Calyx finely hirsute, 5.0-6.0 mm (5.6 mm) long, con- 
spicuously saccate for ca 2/3 of its length, the distended region 1.2-1.3 
mm wide; the tube tapering to a slender neck ca 0.5 mm wide closing the 
orifice after anthesis; the upper calyx teeth slightly connate, aaa or very 
narrowly triangular, ca 0.2 mm wide at the base, 1.0-1. m long, erect 
and dorsiventrally convergent with the lower teeth to oe the orifice, 
hirsute-ciliate; the lower teeth subulate, recurved and arching over the upper 
teeth, 0.3 wide at the base, 1.5-2.0 mm long, ciliate; annulus dense ca 1.5 
mm wide and seated ca 1.5 mm below the juncture of the upper and lower 
teeth. Corolla blue, 7.0-9.0 mm (8.0 mm) long sparsely pubescent in the 
throat, densely to below forming an annulus 0.7-1.5 mm wide; the tube 
1.0 mm wide; the upper lip ligulate 1.0-1.5 mm long; the lower lip 2.5—3.0 
mm wide and 2.5-3.0 mm long. Nutlets ca 0.4 mm wide, ca 1.2. mm long. 
Chromosome numbers, 27 = 34, 36. 

Distribution (Fig. 5). Various habitats from Montana to San Luis Potosi, 
Mexico; and western California to western Nebraska. Jun—Sep. 

TYPE: Nuevo Leon. Near Monterrey, 1828, Berlandier s.n. (Holotype, 
isotypes, BM! GH! ). 

. drummondiw is a morphologically variable species and a number of 


258 


races can be discerned within it. In central Texas along the escarpment of 
the Edwards Plateau the plants take the form of annuals or biennials with 
linear leaves, sparsely pubescent calyx, and the teeth strongly converging. 
Many of these populations are aneuploid (27 = 34) and hybridize with 
both varieties of H. reverchonit. Towards the west and north, the plants 
take the form of tufted caespitose perennials with leaves more oblong- 
elliptic and the calyx to varying degrees sealed at the summit. In southern 
Nevada hybridization with H. nanum produces still another morphological 
grouping. To the west, mesic montane forms possess long cernuous elliptical- 
oblong leaves and calyx teeth only very loosely closed at the summit. This 
last form is common in the sha Mountains of west Texas and in the 
Sacramento Mountains of central New Mexico where it hybridizes with 

pulcherrimum. The eastern Mexico populations, occurring along the 
eastern edge of the Sierra Madre Oriental, vary in the direction of H. 
reverchonii var. serpyllifolium. 


18. HEDEOMA REVERCHONII Gray 


Key to the varieties of H. reverchonit 

1. Plants 25.0-60.0 cm tall branching usually restricted to the base and the upper 
1/2 = on shoots; leaves gray green, ca 3.7 mm wide, 11.7 mm long; calyx 6.0-7.0 
mm , very coarsely hirsute to hoary pilose; the upper ne widely oe 
ein ae 10.0-15.0 mm long, the tube conspicuously dilated _ plants 
restricted to the eastern edge of the Edwards Plateau and Lampa ys ut Plain of 
central Texas, and south central Oklahoma; lemon-scented . “pic = 
eee iauiav ee paras . Aor. var. aN 
1. Plants 15. 0-40. 0 cm tall branching freely one the 2h; of the shoots; leaves 
< green, ca 3.0 mm wide, 8.3 mm long; calyx 5.0—6. m long, coarsely hirsute, 

upper ee erect or only slightly spre Aue la a “oral small, 8.0-10 
mm long, the tube only very slightly dilated Uae plants from eastern — of 
Edwards eee south to the Central Coastal Plain Texas and west uth- 
eastern New Mexico; camphor-scented ... 8b. Hur. var. eh Sse 


18a, HEDEOMA REVERCHONII Gray var. REVERCHONII, Syn. Fl. ed 2, 2: 460. 
1886 


Hedeoma drummondii - reverchonit A. Gray, Syn. Fl. 2: 363, 8. 

Hedeoma latum Small, Fl. S.E. U.S. 1040, 1337, 1903. TYPE: a an Dallas Co. 
Rocky prairies, near Ses Texas, Jun—Sept. Reverchon (Curtis 2020) (Holo- 
type, NY! isotypes, BM! F! GH! ND! NY! SMU! US!) 

Robust, suffruticose perennials, 25.0-60.0 cm high, with the shoots of 
the previous season usually persistent. Shoots ascending or shortly decum- 
bent, arising from a stout woody taproot, branching more or less restricted 
to the very base, branches long and well-developed; stems densely pubescence, 
nearly canescent above with tightly retrorsely curling hairs, glabrous below. 
Leaves gray-green, eat erect, coriaceous, elliptic-oblong and highly vari- 
able in size, 3.0-5.0 mm (3.7 mm) wide, 9.0-14.0 mm (11.7 mm) long, 
margins entire, apex hae we mars attenuated and subsessile, sparsely 
pubescent above, densely so below and on the margins; nerves inconspicuous. 


299 


Cymules 2—4-flowered (occasionally more) well-spaced along the upper 1/2 
of the stems; peduncles 1.0-1.5 mm long; pedicels 3.0-4.0 mm long, clothed 
in tightly retrorsely curling hairs, bracteoles short. Calyx 6.0-8.0 mm 
(6.6 mm) long; the tube 4.0-5.5 mm (4.2) long, saccate below tapering 
to a narrow neck partially sealing the calyx orifice, hoary villous with the 
hairs antrorsely curling at their tips and up to 1.5 mm long; the upper 
calyx teeth not connate or slightly so, the lobes subulate, ca 0.2 mm wide 
at the base and ca 1.0 mm long, laterally spreading, slightly reflexed but 
with the base convergent to the lower teeth and closing the orifice, hirsute— 
ciliate; the lower teeth subulate, recurved 2.0-2.5 mm long, ciliate; annulus 
extremely dense ca 1.5 mm wide and seated ca 1.5 mm below the juncture 
of the upper and lower teeth. Corolla large and showy 10.0-15.0 mm (12.0 
mm) long, white or lavender, variously marked on the throat, sparsely 
pubescent in the throat, densely so in the tube forming a well-defined 
annulus; the tube dilating distally to ca 5.0 mm wide above; the upper lip 
ligulate and sub-galeate, ca 5.0 wide at the base, 4.0-5.0 mm long, the 
lower lip ca 4.0-5.0 mm wide, ca 5.0 mm long. Nutlets ca 0.6 mm wide, 
ca 1.2 mm long, areolate and glaucous. Chromosome number, 27 = 34. 

Distribution (Fig. 5). Open, exposed, calcareous outcrops in southern 
Oklahoma and central Texas along the fault escarpements of the Edwards 
Plateau and Lampasas Cut Plain. Also collected infrequently on calcareous 
outcrops in east Texas, Arkansas and Alabama, as yet however, it is un- 
reported in Louisiana and Mississippt. 

TYPE: Texas. Brown Co., Rocks, Blanket, 9 Aug 1877, Reverchon 755 
(Holotype, GH!; isotypes, F! NY! US!). 


18b. HEDEOMA REVERCHONI Gray var. SERPYLLIFOLIUM (Small) Irving, 
Britt. 22: 333-345. 1970. 
Hedeoma serpyllifolium Small, Bull. N.Y. Bot. Gard. 1: 287. 1896. 
deoma sanctum Small, Bull. N.Y. Bot. Gard. 1: 287. 1896. TYPE: TEXAS. Bexar 
Co., San Antonio, Apr 1853, Thurber s.n. (Holotype, NY!; isotype, GH!). 
Robust, suffruticose perennials, 15.0-40.0 cm high, the shoots of the 
previous season persistent. Shoots numerous, ascending or decumbent, 
branching freely along the lower 2/3 of the stems, the branches long and 
ascending, densely pubescent above with retrorsely curling hairs, puberulent 
below. Leaves dark green typically falling from the lower 1/2 of the shoots 
with maturity, ascending, thick membranous, elliptical-oblong, 2.24.0 mm 
(3.0 mm) wide, 6.0-10.0 mm (8.3 mm) long, margins entire, apex obtuse, 
base narrowly attenuated, petiolate or subsessile, glabrate above, densely 
pubescent below; nerves inconspicuous. Cymules usually 2—4-flowered, some- 
what crowded along the upper 2/3 of the stems, developing to one-side; 
peduncles 1.0-1.5 mm long; pedicels 2.0-3.0 mm long clothed in retrorsely 
curling puberulent hairs, bracteoles short. Calyx 5.0-6.0 mm (5.5 mm) 
long; the tube saccate below tapering distally and partially closing the tube, 


260 


3.0-4.0 mm (3.7 mm) long, densely hirsute pubescent with the hairs spread- 
ing or antrorsely curved; the upper calyx teeth slightly connate, subulate, 
ca 0.2 mm wide at the base, 1.0-1.2 mm long, erect or slightly laterally 
spreading, somewhat convergent with the lower and closing the orifice, 
hirsute-ciliate; the lower teeth subulate ca 0.2 mm wide at the base, 1.5—2.0 
mm long; annulus very dense, ca 1.5 mm wide and seated ca 1.5 mm below 
the juncture of the upper and lower teeth. Corolla chiefly white, occasionally 
lavender, 8.0-10.0 mm (9.0 mm) long, sparsely pubescent in the throat, 
densely so below forming an annulus; the tube dilating upwardly to ca 2.0 
mm wide; the upper lip ligulate ca 1.0 wide at the base, ca 1.5 mm long; 
the lower lip ca 3.2 mm wide, 3.2 mm long. Nutlets ca 0.5 mm wide, ca 
1.2 mm long, areolate and glaucous. Chromosome number, 27 = 34. 

Distribution (Fig. 5). Abundant in open calcareous outcrops and road- 
cuts from the Balcones fault escarpment of central Texas to southeastern 
ou bees and southward into the south Texas Coastal Plain. Jun—Sep. 

TEXAS. Kerr Co., Kerrville, 1600-2000 ft, 25-30 Apr ee 

Helle We ( Holoty pes, NY! isotypes, GH! IA! MICH! MCS! ND 
NY! PH! SMU! UC! US!) 


19. HEDEOMA MULTIFLORUM Benth., Lab., Gen. et Sp. 367. 1834. 


Hedeoma gilliesti Benth., Lab. Gen. et Sp., 367. 1834. TYPE: ARGENTINA. Cerro 
de Archiras ace _Mendosa, 1820, Gillies s.n. (Holotype, K!; isotype, LA!; phot 
types LA! NY! PH!). 

Micromeria ee, Fisch. et Meyer, Ind. Sem. Hort. Petrop. X: 56. 1845. 

not located. 


Satureia ites (Benth.) Brig., in Engler & Prantl. Nat. Pflanzanf. ed. 1, IV, 3a, 
300. 1897. Based on Hedeoma gilliesii Benth. 
— bonariensis (Fisch. et Meyer) Briq., in Engler & Prantl. Nat. svn ed. 
IV, 3a, 300. 1897. ae on eee bonariensis Fisch. et Mey 


ie caespitose perennials forming tufts, 5.0—25.0 cm in height. Shoots 
numerous densely pubescent or puberulent with retrorsely curling hairs on 
the upper herbaceous portions of the stems, becoming glabrate below on the 
lower woody portions, branching mostly confined to the lower half, the 
branches widely ascending to nearly 45° angles. Leaves deciduous from the 
lower stem regions with maturity, spreading, menage: linear-lanceolate, 
usually five times longer than wide, 1.0-1.5 n (1.2 mm) wide, 5.5-8.5 
mm (7.1 mm) long, margins entire, apex acutish, base narrowly attenuated 
and subsessile, upper and lower leaf surfaces and margins strigose; nerves 
indistinct. Axillary leaf buds develop on the immature stems yielding 2-3 
eaf pairs. Cymules 1-5-flowered (commonly 1-2-flowered ), well-spaced 
along almost the entire length of the shoots; peduncles inconspicuous, less 
than 0.5 mm long, pedicels 3.0-4.0 mm long, retrosely puberulent; bracteoles 
lanceloate, typically shorter than the pedicels they subtend. Calyx 5.0-6.0 

n (5.8 mm) long; the tube 3.9-4.5 mm (4.1 mm) long conspicuously 
saccate for ca 2/3 of its length, the distended region 1.2-1.5 mm wide, 


261 


the tube above the pouch constricted and tapered into a narrow upper neck, 
0.8-1.0 mm wide, partially closing the calyx orifice; calyx sparsely and 
minutely hirtellous with antrorsely curling hairs; the upper calyx teeth 
connate for 1/3-1/2 of their length forming an upper lip, 0.8-1.2 mm 
long, pubescent on its inner face, the lobes narrowly triangular, spreading 
and slightly reflexed, ca 0.3 mm wide at the base, 0.4-0.8 mm _ long, 
hirtellous-ciliate; the lower teeth subulate, 1.5-2.0 mm long; annulus dense, 
ca 0.7 mm wide and seated ca 1.0 mm below the juncture of the upper 
and lower teeth, extended upwardly into the inner face of the upper lip, 
included. Corolla ca 8.2 mm long, glabrous within; the tube ca 7.0 mm 
long gradually dilated upwardly, ca 2.0 mm wide; the upper lip ligulate, 
emarginate, 1.0-1.2 mm wide, ca 2.0 mm long; the lower lip ca 3.5 mm 
wide, ca 3.0 mm long. Nutlets ca 0.7 mm wide, 1.4 mm long, the surface 
foveate, brown at maturity, strongly glaucous. Chromosome number, 27 = 

Distribution (Fig. 12). Rocky calcareous outcrops of rivers and roadcuts 
in southern Brazil, Uruguay, and northern Argentina. Nov—Feb. 

TYPE: BRAZIL. 1830, Sello 1062 (Holotype, B destroyed; lectotypes here 
designated K! isotypes LA!; photolectotypes, LA! NY! PH! ). 

Hedeoma multiflorum is most closely allied to H. drummondii of the 
Northern Hemisphere and differs from it primarily in the details of the 
calyx. 


20. HEDEOMA DIFFUSUM Greene, Pittonia 3: 338. 1898. 
Hedeoma CE sal Greene, Pitt. 3: 339. 1898. TYPE: ARIZONA. Cocon 
Co. near Flagstaff, 29 Aug 1884, Jones 404 or 4104 (Holotype, US! ee 
BM! CAS! DS! GH! LA! MCS! NY! POM! UC! US!) 

Perennial herbs forming dense, circular, dwarfed mats with numerous 
decumbent shoots, 7.0—20.0 cm long; branching prolifically at the base and 
generally unbranched distally, stems puberulent above with retrorsely curl- 
ing or spreading hairs, pubescent with spreading hairs below. Leaves spread- 
ing, membranous, ovate, 2.0-455 mm (3.0 mm) wide, 3.5-0.0 mm (5.2 
mm) long, margins entire, apex narrowly obtuse or acute, base rounded or 
attenuated and petiolate, petioles 5.0-1.0 mm long; the blade glabrate or 
puberulent above, densely hirtellous-puberulent below and on the margins; 
nervation inconspicuous. Cymules 1—3-flowered, occurring well-spaced along 
nearly the full length of the numerous shoots; peduncles 0.5—-1.0 mm long; 
pedicels 2.0-3.0 mm long puberulent with retrorsely curving hairs, bracteoles 
subulate, shorter than the pedicels they subtend. Calyx 5.0-6.0 mm (5.3 
mm) long; the tube 3.0-4.0 mm (3.6 mm) long more or less tubular, 
slightly gibbous below, the distended portion ca 1.0 mm wide dilated up- 
wardly, hirtellous-hirsute with spreading hairs; the upper calyx pes united 
for ca 1/3 of their length or less, forming an upper lip 1.0-1.2 mm long, 
pubescent on its inner face, the lobes narrowly triangular, 06 1.3 mm 
long, laterally spreading and reflexed or erect, hirsute-ciliate; the lower teeth 


262 


subulate, recurved, 1.5—2.0 mm long, hirsute-ciliate; annulus dense, included 
or slightly exserted, 0.5-1.0 mm wide seated just below the juncture of the 
upper and lower teeth. Corolla ca. 9.5 mm long, blue, pubescent within on 
the throat, annulate below; the tube ca 7.5 mm long upwardly inflated, ca 
2.5 mm wide above; the upper lip ligulate, subgaleate, and straight but 
with the margins and apex revolute, ca 2.3 mm wide, ca 2.0 mm long, 
emarginate, the lower lip ca 5.0 mm wide, ca 5.0 mm long. Nutlets ca 0.5 
mm wide, 1.1 mm long, the surface areolate and strongly glaucous. Chromo- 
some number, 27 = 36. 

Distribution (Fig. 6). Pine forests in the vicinity of Flagstaff, Arizona; 
common on rocky outcrops and exposed areas, ca 7000 ft. Jul-Sep. 

TYPE: ARIZONA. Coconino Co. near Flagstaff, 1883. Rusby 786. (Holo- 
type, GH! isotypes, LA! MICH! NY! PH! UC! US!). 

Although I accept the name and type of H. diffusum designated by 
Greene, I cannot accept his morphological concept of the species. From 
his discussion Greene apparently had a broad concept of this taxon. 
he indicated, the range of the species was “from north to south through- 
out the state [Arizona}”. Thus, the distribution would extend into the range 
of H. nanum. H. diffusum, as it is realized here, is restricted to the Pinus 
ponderosa forest of the Flagstaff area. This distribution also concurs with 
that accorded H, diffusum by Epling and Stewart (1939). 

Hedeoma diffusum is most closely related to H. nanum. The ovate, entire, 
acute leaves, gibbous calyx, and the ad- and abaxially inflated corolla are 
all common morphological features. 


n 


21. HEDEOMA NANUM (Torr.) Briq. 


Key to the varieties of Hedeoma nanum. 

1. Coarse, sane Sipe with shoots few to numerous; leaves 5.0-16.0 mm wide, 

8. ong; cymules 1—7-flowered i well-spaced along the axis; calyces 

ieee, 6.0-8.0 mm long, slightly gibbous or saccate below; plants of central and 
western Arizona. 21c. 

. Plants not a 


H.n. var. macrocalyx 


— 


abov 
2. Caespitose ois als forming dense tufts with shoots extremely numerous, 
retrorsely puberulent-pubescent; leaves 2.1-4.5 mm _ wide S- n long; 
cymules 3—5-flowered, well-space along the axis; calyces 4.5-5.6 mm _ long, 
weakly saccate below; plants eee to southern-most Nevada and contiguous 
areas in California and Arizon lb. H.n. var. californicum 
2. Annual or perennial herbs of v various habits, shoots solitary to numerous, pube- 
scence spreading or seeps ae pilose or hirsute; leaves 2.6—-6.0 
wide, 5.5-13.0 mm long, cymules 3—15-flowered (tpically 8-10) well- Been or 
congested, forming a dense ‘ aie ; calyces 4.0-5.2 m oo distinctly saccate; 
plants widespread—Arizona, New Mexico, Texas and Mexi 


ve Hin. var. nanum 


21a. HEDEOMA NANUM (Torr.) Brig. var. NANUM, in Engler & Prantl, Die 
Naturlichen Pfl. EX, 3a, 294. 1897. 


Hedeoma dentatum var. nanum Torr., Bot. Mex. Bound. 130. 1859. 


263 


6. Distribution of Hedeoma diffusum (triangle), H. nanum va 


Fig. 
eae | ake H. 
(squ 


nanun. 
n. var. macrocalyx (open circles), and H. mn. vat. ie nin 


264 


Hedeoma sche ee Gray, Syn. Fl. N. Am. 2: 368. 1878. Based on H. dentatum 


var. nanum 
Hedeoma nanum ae Pittonia 3: 339. 1898. Based on H. dentatum var. nanum 


sacl nanum (Torr.) Brig. var. typicum pee ae Spec. Nov. Beihefte 115: 
. 1939. Based on H. dentatum var. nanum Tor 
cea var. mexicanum aan Rep. Spec. nov., Beihefte 
O LEON. Calcareous hills near Monterrey, 7 Apr 
CAS! F! GH! K! LA! MICH! 


Hon nanum (Torr.) 
15: 29. 1939. TYPE: Nu 
i508: Pringle 10203 (Holotype, US! isotypes, 

MSC! NY! PH! SI! SMU! UC! phototype, LA!) 


Plants annual or perennial herbs, 5.0-35.0 cm (14.1 cm) high. Shoots 
solitary to numerous, erect, ascending, or shortly decumbent up to 5.0 cm; 
branching primary at or near the base, typically unbranched above; shortly 
pilose or hirsute above, puberulent below, the hairs spreading or retrorsely 
curling. Leaves ascending to cernuous, ovate to oval, variable in dimension, 
2.5-6.0 mm (2.9 mm) wide, 5.5— n (7.4 mm) long, margins entire 
occasionally obscurely crenate, apex a acute to narrowly obtuse, base 
broadly rounded to narrowly attenuated, petiolate, the petiole 0.5-2.5 mm 
long; the leaf surface glabrous to sparsely pubescent above, margin and 
undersurface hirtellous pubescent; nerves inconspicuous. Cymules 3-15- 
flowered (usually between 8 and 10), well-spaced along shoots, or com- 
monly very congested forming a dense but interruped spike of cymes; pe- 
duncles inconspicuous to 3.0 mm long; pedicels long, typically exceeding 
the length of the calyx tube, 3.0-6.0 mm long, clothed in hirsute-hirtellous 
spreading hairs; bracteoles a shorter than the nae they subtend. 
Calyx 4.0-5.2 mm (4.9 m long; the tube 2.5—3.7 1 (3.3 mm) long, 
saccate or occasionaly eres below for 1/2 to 2/3 bE — tube's length, 
the distended region typically more than 1/2 the tube’s length, 1.0-1.5 
mm wide; the calyx tube densely hirsute to hirtellous with the hairs spread- 

ing; the upper calyx teeth connate for slightly less than 1/2 their length 
forming a conspicuous upper lip, 1.0-1.3 mm long, the lobes narrowly 

triangular-lanceolate, occasionally approaching subulate, ca 0.3 mm wide at 
the base, 0.4—0.7 mm long, laterally spreading, and reflexed (commonly at 
nearly right angles to the axis), hirtellous-ciliate, the inner face of the lip 
pubescent; the lower teeth subulate, recurved and not conspicuously exceed- 
ing the upper lip in length, 1.2-2.0 mm long, hirsute or hirtellous-ciliate; 
annulus dense, included, ca 0.7 mm wide, seated ca 0.8 mm below the junc- 
ture of the upper and lower calyx teeth, and continued upwardly into the 
inner face of the lower lip. Corolla 8.0-9.0 mm (8.9 mm) long, pubescent 
within the throat in two decurrent lines, the tube 6.0—7.0 mm long, strongly 
inflated, ca 3.0 mm wide above; the upper lip subgaleate, the margins and 

apex revolute, ca 2.0 mm wide, ca 2.0 mm long; the lower lip ca 4.0 mm 

wide, ca 3.5 mm long. Nutlets oblong, ca 0.5 mm wide, ca 1.1 mm long, 

areolate and weakly glaucous. Chromosome number, 2” = 36. 

Distribution (Fig. 6). Common on rocky, usually calcareous bluffs, ridges, 


— 


265 


roadcuts and streamways. Southern Nevada and adjacent California, southeast 
to southeastern Arizona, east to Trans-Pecos Texas and south to San Luis 
Potosi, Mexico. Apr—Jul. 

TYPE: TEXAS. El Paso Co., Rocky hills of the Rio Grande, near El Paso, 
4 May 1852, Bigelow, sm. (Holotype, NY!; isotypes, DS! K! US!). 

Hedeoma n. var. nanum is a variable taxon which harbours a number of 
more or less distinct races. One such race received formal recognition as 
var. mexicanum by Stewart and Epling. It is an annual with bristly, spread- 
ing stem pubescence and a somewhat shortened calyx and corolla. Repre- 
sentative collections include, Whiting 950 (LA); and Lundell 5030 (LA, 
MICH). Another well-marked race occurs in northern Arizona. It is char- 
acterized by long, trailing stems (up to 40 cm long), and exceptionally 
long internodes, peduncles and pedicels (the peduncles up to 3.0 mm long, 
and the pedicels up to 6.0 mm long). The following are representative: 
Howell 26454 and 26409; Bailey s.n., Clover 5134, Wooton 308, and Whit- 
ing 1089. 


21b. HEDEOMA NANUM (Torr.) Brig. var. CALIFORNICUM Stewart, Rep. 
Spec. Nov., Beihefte 115: 29. 1939. 


Dense, caespitose perennials forming tufts, 11.0-25.0 cm (17.1 cm) high. 
Shoots extremely numerous, ascending or shortly decumbent up to 4.0 cm 
long; branching almost exclusively at or near the base, densely pubescent- 
puberulent above with hairs retrorsely curling, puberulent below. Leaves 
small, spreading to cernuous, ovate, 2.0-4.5 mm (3.4 mm) wide, 3.4-8.5 
mm (5.0 mm) long, margins entire or obscurely cremate, apex acute, base 
broadly to narrowly rounded, shortly petiolate, the petiole 0.5-1.5 mm long; 
leaf surfaces puberulent above, margins and undersurface hirtellous- 
pubescent; nerves inconspicuous. Cymules 3—5-flowered well-spaced along 
the upper 2/3-3/4 of the shoots; peduncles 1.0-2.7 mm_ long; pedicels 
3.0-4.3 mm long; bracteoles subulate, shorter than the pedicels which they 
subtend. Calyx 4.5-5.6 mm (5.1 mm) long; the tube 3.0-4.0 mm (3.3 
mm) long, weakly saccate below for ca 2/3 of the tube’s length, the dis- 
tended region 1.2-1.5 mm wide, hirsute-hirtellous to puberulent with the 
hairs spreading, the upper calyx teeth connate for slightly less than 2 
of their length forming a conspicuous upper lip, 0.6-1.5 mm long, the lobes 
triangular, ca 0.4 mm wide at the base, 0.4-0.8 mm long, laterally spreading 
and slightly to sharply reflexed, hirtellous-ciliate, the inner face giabrate or 
pubescent; the lower teeth subulate, recurved, not conspicuously exceeding 
the lower in length, 1.5-2.0 mm (1.8 mm) long, hirsute-ciliate; annulus 
dense, included ca 0.7 mm wide and seated ca 0.7 mm below the juncture 
of the upper and lower teeth. Corolla as in H. nanum var. nanum ca 88 
mm long. Nutlets 0.6 mm wide, 1.2 mm long, strongly glaucous. Chromo- 
some number, 27 = 

Distribution (Fig. 6). Frequent on rocky limestone outcrops in the 


266 


mountains of the Mojave Desert of California and contiguous areas of 
southern Nevada and northwestern Arizona. Apr—June. 

TYPE: NEVADA, Clark Co., Good Springs, 1 May 1905, Jones s.n. (Holo- 
type, LA!; isotypes, CAS! Ds! F! GH! POM! US!). 

Hedeoma #. vat. californicum is a relatively uniform taxon separated 
from the varieties nanwm and macrocalyx by its multi-stemmed, tufted habit, 
finely puberulent stems and microphyllous leaves. 


2lc. HEDEOMA NANUM (Torr.) Brig. var. MACROCALYX Stewart, Rep. 
Spec. Nov., Beihefte 115: 29. 1939. 


Somewhat coarse, robust perennial herbs, 15.0-55.0 cm (27.0 cm) high. 
Shoots numerous, branches ascending at close angles, cansescent above, pub- 
erulent below, hairs retrosely curling. Leaves spreading or cernuous, ovate, 
oval, or elliptical, 5.0-16.0 mm (7.3 mm) wide, 8.0-25.0 mm (12.4 mm) 
long, margins obscurely crenate, apex broadly acute or obtuse, base broadly 
rounded to attenuated, petiole 1.5-5.0 mm in length; finely puberulent above, 
margins and undersurface hirtellous-puberulent, nerves inconspicuous. Cym- 
ules 5—7-flowered, well-spaced along the upper 2/3 of the shoots; peduncles 
0.5-1.5 mm long; pedicels 3.0-5.0 mm long and typically exceeding the 
length of the calyx tube; bracteoles subulate usually shorter than the pedicels 
they subtend. Calyx 6.1-8.0 (6.8 mm) long; the tube 4.0-5.0 mm (4.4 
mm) long, slightly gibbous or saccate below for ca 1/2 of the tube’s length, 
the distended region 1.3-1.8 mm wide, sparsely to densely hirsute-hirtellous 
with hairs spreading; the upper calyx teeth connate for slightly less than 
1/2 of their length to form a long and conspicuous upper lip 1.5—2.7 mm 
(2.0 mm) long, lobes narrowly lanceolate, ca 0.3 mm wide, ca 0.8-1.2 mm 
long, hirsute-hirtellous-ciliate; the lower teeth subulate, recurved and not 
conspicuously exceeding the upper lip in length, 2.0-3.0 mm (2.5 mm) 
long, hirsute-ciliate; annulus dense, included, ca 0.7 mm wide and seated 
ca 0.8 mm below the junction of the upper and lower calyx teeth. Corolla 
as in H. n. var. nanum, 8.0-9.0 mm long. Nutlets ca 0.6 mm wide and 1.4 
mm long, strongly glaucous. Chromosome number, 27 = 36. 

Distribution (Fig. 6). Rocky limestone outcrops and hills of arid central 
and southwestern Arizona. May—Aug. 

TYPE: ARIZONA. Pinal Co., Higley, Queens Creek, 19 May 1926, Peebles 
and Harrison 1916 (Holotype, LA! ). 


22. HEDEOMA MICROPHYLLUM Irving, Brittonia 22: 338-340. 1970. 


Tufted perennials, 10.0-17.0 cm high. Shoots numerous, decumbent or 
ascending with branching more or less restricted to the base, puberulent or 
hirtellous with hairs retrorsely curving. Leaves numerous and crowded, 
ovate, 3.0-5.0 mm (4.0 mm) wide, 4.0-6.5 mm (5.0 mm) long, margins 
obscurely serrate, apex acute, base rounded or weakly cuneate, petiolate with 
a petiole ca 1.0 mm long; surfaces glabrous above, minutely puberulent 


267 


below and on the margins; nerves conspicuously elevated. Cymules 1-3- 
flowered (occasionally up to 5-flowered), crowded along the upper 1/2 of 
the stems; peduncles inconspicuous, less than 1.0 mm long; pedicels ca 3.0 
mm long, clothed in hirtellous spreading hairs; bracteoles shorter than the 
pedicels they subtend. Calyx 5.5-6.5 mm (6.0 mm) long; the tube 4.0-4.5 
mm (4.1 mm) long, saccate below but not markedly so for ca 1/2 of the 
tube’s length, the distended region ca 1.2 mm wide, moderately hirsute 
with spreading or slightly recurving hairs; the upper teeth connate for ca 
1/3 of their length forming an upper lip, ca 1.5 mm long, the lobes reflexed 
and spreading, narrowly triangular, ca 0.4 mm wide at the base, ca 1.0 mm 
long, hirsute or hirtellous-ciliate; the lower teeth recurved, subulate, ca 0.5 
mm wide at the base, 2.0 mm long, hirsute or hirtellous, ciliate; annulus 
moderately dense, included, ca 0.5 mm wide and seated at the juncture of 
the upper and lower teeth. Corolla blue, ca 8.5 mm long, glabrate within; 
the tube ca 7.0 mm long, slender, only slightly dilated upwardly. ca 0.5 mm 
wide; the upper lip ligulate, slightly concave, reflexed and emarginate, ca 
1.0 mm wide, ca 1.5 mm long; the lower lip ca 2.5 mm wide, ca 3.0 mm 
long. Nutlets ca 0.5 mm wide, ca 1.0 mm long. Chromosome number, 2” 
= 36. 


Distribution (Fig. 7). Canyons near San Luis Potosi, Mexico, Jul-Sep. 
TYPE: SAN Luis Potosi. 10 km east of Guadalcazar, Aug 1966, Irving 
693 (Holotype, TEX! ). 


23. HEDEOMA MEDIUM Epl., Rep. Spec. Nov., Beihefte 115: 30. 1939. 


Caespitose perennials, 10.0-25.0 cm high. Shoots numerous, densely 
puberulent with retrorsely curling hairs on the upper herbaceous regions 
of the stems, glabrate below the woody portions; branching confined to 
the lower half of the plant with the branches ascending at close angles. 
Leaves deciduous from the lower stem regions with maturity, linear-elliptical, 
1.5-5.0 mm (2.4 mm) wide, 6.5-17.0 mm (8.0 mm) long, margins entire, 
apex acutish, base tapered and subsessile, upper surface glabrate to puber- 
ulent, lower surface and margins puberulent; nervation indistinct with only 
the midrib visible. Axillary buds developed on the immature stems yielding 
2-3 leaf pairs. Cymules 1—5-flowered, well-spaced along the upper 2/3 of 
the shoots, peduncles inconspicuous, less than 0.5 mm long, pedicels 4.0-5.0 
mm long, retrorsely puberulent, bracteoles lanceolate, typically shorter than 
the pedicels they subtend. Calyx 4.2-5.0 mm (4.5 mm) long; the tube 
2.5-3.0 mm (2.8 mm) long, conspicuously saccate for ca 2/3 of its length, 
the distended pouch ca 1.5 mm wide, constricted above; calyx tube glabrate- 
puberulent; the upper teeth connate for over 2/3 of their length to form 
a broad, upper lip, 1.0-1.2 mm long, pubescent on its inner face, the lobes 
deltoid, spreading and reflected, ca 0.3 mm wide at the base, ca 1.3 mm 
long, the margins glabrate or hirtellous-ciliate, the lower teeth subulate, 
recurved, 1.4—2.0 mm long, hirtellous or hirsute-ciliate; annulus dense, in- 


268 


cluded, ca 0.5 mm wide, seated approximately at the juncture of the upper 
and lower teeth, extended upwardly onto the inner face of the upper lip. 
Corolla ca 5.0 mm long glabrous within; the tube ca 4.5 mm long scarcely 
dilated upwardly, ca 1.2 mm wide; the upper lip ligulate, emarginate, 
straight, ca 1.3 mm wide, ca 1.5 mm long; the lower lip ca 3.0 mm wide, 
ca 2.0 mm long. Nutlets ca 0.5 mm wide, 1.0 mm long, the surface foveate, 
and glaucous. Chromosome number, unknown. 

Distribution (Fig. 12). Prairies and pasture lands of Uruguay and Argen- 
tina. Nov—F 

TYPE: UruGuay. Sorino, 1 Dec 1935, Gallinal sn. (Holotypes, LA!). 

Epling and Stewart comment that H. medium is ‘a species of uncertain 
quantity and disposition.” I, too, am in a quandry over its natural relation- 
ships and have placed it only tentatively with H. nanwm and its allies. The 
leaf morphology and foveate nutlet surface are very similar to those features 
of H. multiflorum and in habitat and general habit it resembles H. hispidum. 


24. HEDEOMA COSTATUM Gray. 


A number of species have been created from the variable Hedeoma 
costatum complex. Most of the species stem from an attempt, by many 
workers, to formalize the marked differences observed in morphology 
between mesophytic and xerophytic representatives of this complex. 
costatum, H. tenellum, H. quinquenervatum, H. pringeli, and H. ee 
are all names accorded by various authors to the mesophytes. H. pulchellum, 
H. albescentifolium and H. convisae are names which were applied to 
the xerophytes. The taxonomically significant variation between these two 
basic groups resides primarily in length of bracteoles, degree of leaf denta- 
tion, and overall pubescence pattern. However, after careful analysis of 
herbarium material, I have been unable to recognize these basic morpho- 
logical groups at the specific level. Instead, I have maintained the complex 
as a single species separated into two varieties: var. costatum, representing 
the mesophytic populations; var. pulchellum, the xerophytic populations. 

Representatives of the southern most populations of H. costatum (Chia- 
pas, Mexico) were tetraploid in chromosome number (22 = ; ese 
plants, however, displayed no discernable morphological differences accom- 
panying the increase in the ploidy level. 


Key to the varieties of Hedeoma costatum. 


. Plants ane pubescent but not canescent with hirsute-villous hairs; leaves with 
margins dentate, often obscurely so; bracteoles narrowly tria poe usually ap ter 
than the pie corolla long exserted; plants of central southern Mexic 

are 24a. He var. pera 

; Plants canescent throughout with stiff hirsute- villous hairs; leaves with margins 

coarsely dentate; bracteoles subulate, usually conspicuously longer than the eer sa 
corolla short Lacely exserted from the a t long exserted; plants of norther 

Mexico and southwestern United States... 24b. H.c. var. ealeh allan 


— 


—y 


269 


ig. 7. Distribution of Hedeoma costatum var. costatum (closed circles ) , H. cos- 
tatum vat. ee oe circles), H. microphyllum (closcd triangles), H. pilica- 
tum (closed squares), H. tenuipes (open squares), H. guercetorum (open Meaneles). 
and H. ee (asterisk). 


270 


24a. HEDEOMA COSTATUM Gray var. COSTATUM, Syn. Fl. 2: 363. 1878. 
Hedeoma costatum Hemsl., Biol. Centr. Am. Bot. 2: 547. 1881. 
TYPE: CHIAPAS. 1864-1870. Ghieshrecht 815. (Holotype, K!; isotypes, BM! GH! 
NY!; Cenaon GH! LA! 1). 
Hedeoma tenellum Hemsl., Biol. Centr. Am. Bot. 2: 549. 18 
NUEVO LEON. Guajuco, Feb-Oct 1890, Palmer ae (Holotype, K!; 
isotypes, GH! PH! US!; phototype, LA!). 
Hedeoma pringlei Briq., Ann. Conser. et Jardin Bot. Geneve 2: 182. 1898. 
EVO LEON. Wooded slopes of Sierra Madre near Monterrey, 13 Jul 
1888, Pringle 1911 (Holotype G!; isotypes, BM! GH! K! LA! MSC! ND! NY! 
PH! POM! UC! US!; phototype, LA!). 
Hedeoma permixtum Briq., Ann. Conserv. et Jardin Bot. Geneve 2: 183. 1898. 
Based on H. pringlei. 
Hedeoma quinquenervatum Bartlett, Proc. Am. Acad. 44: 634. 1908. TYPE: NUEVO 
adre above Monterrey, 3000 ft, 9 Apr 1906, Pringle 10241 
(Holotype GH!; isotypes, CAS! F! IA! LA! MICH! MSC! NMC! SMU! TEX! 
C! US!). 


Extremely variable, loose or tufted perennials, 4.0-36.0 cm (15.0 cm) 
high. Shoots few to numerous, ascending or shortly decumbent, branching 
copiously in the lower 1/4 of the plant, the upper branches sparse and 
short; moderately to densely hirsute in the upper stem regions, the hairs 
spreading or antrorsely curving, puberulent below. Leaves spreading, mem- 
branous, ovate, oval, or elliptical, 3.0-13.0 mm (6.0 mm) wide, 5.0—28.0 
mm (10.0 mm) long, margins strongly to obscurely serrate-dentate, apex 
acute, base broadly rounded or attenuated, the petiole 1.0-4.0 mm long; 
glabrate or pubescent below with the margin ciliate; nerves not conspicu- 
ously elevated. Cymules 3-7-flowered (typically 3-5), well-spaced along 
the upper 1/2 of the stems or becoming more or less congested toward the 
apex; peduncles 1.0—3.0 mm long; pedicels 4.0-5.0 mm long, hirtellous with 
the hairs spreading or retrorsely curving; bracteoles triangular, shorter than 
the pedicels they subtend. Calyx 5.0-10.0 mm (6.4 mm) long; the tube 
4.0-7.0 mm (5.0 mm) long, slightly gibbous for 1/3-1/2 of its length, 
the distended region, ca 1.1 mm wide, moderately or densely hirsute or 
hirtellous with the hairs spreading and/or antrorsely curling; the upper 
calyx teeth connate for 1/4-1/3 of their length forming an upper lip, the 
lobes narrowly triangular to subulate, 0.2-0.5 mm wide at the base, 1.5- 
2.5 mm long, laterally spreading and reflexed, hirsute or hirtellous-ciliate; 
the lower teeth recurved, subulate, ca 0.5 mm wide at the base, 2.0—3.0 mm 
long, hirsute or hirtellous-ciliate; annulus moderately dense, included, ca 
0.5 mm wide and seated at the juncture of the upper and lower calyx teeth. 
Corolla, well-exserted from the calyx, 9.0-20.0 mm (12.0 mm) _ long, 
pubescent in the throat and the tube but not forming a well-defined annulus; 
the tube 8.5-18.5 mm (11.5 mm) long, slightly dilated upwardly, ca 2.0 
mm wide; the upper lip flat, ligulate, slightly reflexed upwardly and emar- 
ginate, ca 1.5 mm wide, 1.5 mm long; the lower lip 3.0-4.0 mm wide, 
2.0-3.0 mm long. Nutlets 0.6 mm wide, ca 1.2 mm long, the surface 


eal 


minutely areolate, weakly glaucous. Chromosome numbers, 27 = 36, 72. 
Distribution (Fi oy Pine Oak Forest of the Sierra Madre Oriental, 
Nuevo Leon to Hidalgo and Chiapas. 8000-10,000 ft. Jun—Sept. 
TYPE: CHIAPAS. 1864-1870, Ghiesbreght 815 (Holotype, GH!; isotypes, 
BM! K! LA! NY!; phototypes, GH! LA! US!). 


24b, HEDEOMA COSTATUM Gray var. PULCHELLUM (Greene) Irving, Brit- 
tonia 22: 345. 1970 
Hedeoma pulchellum Greene, Leaflets Bot. Obs. 1: 213. 1906. Basionym. 
ee ma albescentifoliunm Bartlett, Proc. Am. Acad. 44: 633. 1908. TYPE: CHI- 
HUA. Santa Eulalia Mountains, Apr 1885, Pringle 133 (Holotype, GH!; 
aoe F! NY! POM! PH! UC! US!). 
Hedeoma convisae A. Nels., Am. Bot. 23: 270. 1936. TYPE: NEW MEXICO. Eddy 
Co., Carlsbad Caverns, chaparral, Apr 1930, Convise 6 (Holotype, RM!; isotype, 
LA!). 


Densely tufted perennials up to 15.0 cm high. Shoots numerous, ascend- 
ing, or shortly decumbent; branching in the lower half; canescent with 
retrorsely curling hirsute hairs. Leaves spreading, membranous, ovate or 
elliptical, 3.0-6.5 mm (48 mm) wide, 5.0-10.0 mm (7.8 mm) long, 
margins coarsely serrate, apex sharply acute, base rounded or shortly cuneate 
with a petiole ca 1.0 mm long; puberulent to nearly villous above, strigose 
to villous below and on the margins, nerves not conspicuously raised. 
Cymules 3—5-flowered, typically congested in the upper 1/2 of the stems; 
peduncles 1.0-2.0 mm long; pedicels ca 4.0 mm long, hoary-villous with 
the hairs spreading or pubescent with the hairs retrorsely curling; bracteoles 
subulate usually conspicuously longer than the pedicels they subtend. Calyx 
6.5-8.0 mm (7.2 mm) long; the tube 4.5-5.5 mm (5.0 mm) long, saccate- 
gibbous for ca 1/2 of its length, the distended region ca 1.3 mm wide, 
typically hoary-villous, occasionally pubescent, hairs spreading or antrorsely 
curving; the upper teeth connate for ca 1/4 of their length forming an upper 
lip 2.0-3.0 mm long, the lobes slightly reflexed and laterally spreading, 
subulate, ca 0.3 mm wide at the base, 2.0 mm long, hirsute-ciliate; the 
lower teeth recurved, subulate, ca 0.4 mm wide at the base, 2.0-3.0 mm 
long, hirsute-ciliate; annulus dense, included, ca 0.5 mm wide and seated at 
the juncture of the upper and lower teeth. Corolla dimorphic: in the first 
form white, slender and about equal to the length of the calyx, the lobes 
minute; in the second form blue, long and well-exserted from the calyx 
tube, 10.0-20.0 mm (14.0 mm) long, pubescent in the throat and tube 
but not annulate; the tube 8.5-18.5 mm (10.5 mm) long, slightly dilated 
upwardly ca 1.5 mm wide; the upper lip ligulate, slightly reflexed and 
emarginate ca 1.5 mm wide, ca 1.5 mm long; the lower lip 3.0-4.0 mm 
wide, 2.0-3.0 mm long. Nutlets ca 0.6 mm wide and 1.2 mm long, the 
surface minutely areolate, weakly glaucous. Chromosome number 2” = 36. 

Distribution (Fig. 7). Arid calcareous mountain regions of northern 
Mexico, western Texas and southern New Mexico. 3000-7000 ft. May—Jul. 


zie 


TYPE: New Mexico. Sierra Co., Kingston, 6600 ft, 19 May 1905, 
Metcalf 1599 (Holotype, ND!; isotype, NMC!; phototype, LA! ). 


25. HEDEOMA JOHNSTONII Irving, SIDA 7(2): 103-106. 1977. 


Wiry perennial herbs up to 30.0 cm tall; shoots ascending or decumbent, 
numerous, branching primarily from the base and rooting at the lower 
nodes, glabrous except for a minute tuft or retrorsely curling hairs at each 
node. Leaves spreading or ascending, stiff membranous in texture, glabrous, 
conspicuously glandular punctate on the lower surface, ovate-oblanceolate, 

0.0-20.0 mm long, 4.0-7.0 mm wide, dentate with 4-5 pairs of teeth 
terminating the costae, apex acute, base attenuate to a short petiole ca 1.5 
mm long, midrib and secondaries conspicuously elevated. Cymules 1—3- 
flowered, well-spaced along the upper one-third of the stem; primary 
peduncles ca. 1.0 mm long; primary pedicels ca. 3.0 mm long; bracteoles 
lanceolate, equal to or slightly longer than the pedicels. Calyx 10.0-15.0 
mm long (base to tip of upper teeth), tubular-funnelform, not gibbous, 
sparsely hirsute; upper teeth partially united forming a small but conspicu- 
ously reflected lip, the lobes triangular ca 1.0 mm long, sparsely ciliate; 
lower teeth deltoid below ca 1.0 mm wide, abruptly tapering to an aristate 
apex, ca 1.0-2.0 mm long, hirsute-ciliate, annulus located just below the 
teeth, included. Corolla showy, long and slender, up to 45 mm long from 
its base to the tip of the upper lip, pink-violet, broadly annulate within 
where seated in the calyx; upper lip short, ca 4.0 mm long, 3.0 mm wide, 
emarginate; the lower lip spreading, ca 5.0 mm long, ca 3.0 mm wide, 
Nutlets 2.0 mm long, 1.0 mm wide. Chromosome number unknown. 

Distribution (Fig. 7). Canyons and igneous slopes of the Sierra Del 
Jardin, Coahuila, Mexico. 1500-2250 m. Jul-Sep. 

TYPE: COAHUILA: Canyon Hundido on north side of Pico de Centinela, 
Sierra del Jardin, 8 km e. of Rancho El Jardin. “Steep canyon through 1 wee 
sierra, gravelly and sandy loam derived from extrusive igneous rocks, 15 
2250 m.” 27 July 1973: M. C. Johnston, F. Chiang, ‘i L, Wendt and D. 
Riskind 11803. 

Hedeoma johnstonii is a distinctive species apparently endemic to the 
steep igneous slopes of the Sierra del Jardin, Coahuila, Mexico. Its habit 1s 
very reminiscent of Hedeoma bellum, an endemic of the mountains of the 
western coast of Mexico, but, in the details of its calyx and corolla is more 
closely allied to Hedeoma costatum and its allies. It is readily separated from 
all species of Hedeoma by its exceptionally long and showy corolla. 


26. HEDEOMA PLICATUM Torr., Bot. U.S. Mex. Bound. Surv., part 2, 130. 
1859 
Perennial herbs, 15.0-37.0 cm high. Shoots typically numerous, ascend- 
ing or very shortly decumbent, copiously branched towards the base, sparsely 
so above with the branches short; hirtellous above with the hairs retrorsely 


273 


curving, puberulent below. Leaves spreading or ascending, membranous, 
elliptical or rhombic, 3.0-6.0 mm (4.4 mm) wide, 5.4—10.0 mm (7.9 mm) 
long, margins coarsely serrate, with the serrations typically confined to the 
upper 1/2 of the blade, apex acute, base broadly attenuated to a short petiole, 
ca 1.0 mm long; glabrous or puberulent above, puberulent or hirtellous below 
on the nerves and margins; nerves conspicuously elevated. Cymules 3-7- 
flowered (typically 3-5), well-spaced along the upper 2/3 of the stems; 
peduncles short ca 1.0 mm long; pedicels 3.0-4.0 mm_ long, hirtellous 
pubescent with retrorsely curling hairs; bracteoles short. Calyx 4.5-6.0 mm 
(5.9 mm) long the tube 3.4-4.2 mm (4.0 mm) long, pronouncedly gibbous 
for ca 1/2 of the tube’s length, the distended region ca 1.3 mm wide, the 
tube puberulent with short spreading hairs or with hairs antrorsely curling; 
the upper calyx teeth connate for 1/2 of their length forming an upper lip 
ca 3.0 mm long, the lobes narrowly triangular-lanceolate, ca 0.3 mm wide 
at the base, ca 1.4 mm long, laterally spreading and sharply reflexed, hirsute 
or hirtellous-ciliate; the lower calyx teeth subulate, recurved, ca 0.4 mm 
wide at the base, ca 2.0 mm long, hirsute-ciliate; the annulus moderately 
dense, included, ca 0.5 mm wide and seated at least 1.0 mm below the 
juncture of the upper and lower teeth. Corolla blue, 7.0-10.0 mm (9.0 mm) 
long, very sparsely pubescent in the throat in two decurrent lines, sparsely 
short pubescent in the tube; the tube 5.5-8.5 mm (7.5 mm) long, dilated 
upwardly, ca 2.5 mm wide above; the upper lip straight, concave and sub- 
galeate, slightly emarginate, ca 2.0 mm wide, ca 1.5 mm long, the lower 
lip 2.5-3.0 mm wide, 2.5-3.0 mm long. Nutlets ca 0.5 mm wide, 1.0 mm 
long, the surface minutely areolate, weakly glaucous. Chromosome number 
2n = 

Distribution (Fig. 7). Pine-oak woodlands of the mountainous areas of 
southern New Mexico, western “trans-Pecos” Texas, and the northern Mexico, 
5000-8000 . aes 

TYPE: AS. Jeff Davis Co., near Fort Davis in the Limpia creek area, 
dry ravines near the Limpia Mts, July 1849 Bigelow s.n. (Holotype, NY!; 
isotypes, K! US!) 

Hedeoma plicatum is a variable species which intergrades into H. costatum 
var. pulchellum. Hedeoma c. var. pulchellum is a xerophytic, spring flower- 
ing plant, whereas H. plicatwm is a mesophytic, late summer or fall flower- 
ing plant. Morphologically. two characters have emerged to distinguish the 
two species. The leaves o . plicatum are more or less rhombic with 
raised straight costae on the undersurface; the leaves of H.c. var. pulchellum, 
on the other hand, are ovate with the nerves unraised and usually curved 
and branched. The annulus of H. plicatuwm is seated at least 1.0 mm below 
the juncture of the upper and lower calyx while that of H.c. var. pulchellum 
is positioned directly at the juncture. 


27. HEDEOMA TENUIFLORUM Brandegee, Zoé 5: 254. 1908. 
Robust perennials, 30.0-35.0 cm high with numerous shoots long and 


274 


ascending, bristly-hirsute pubescent with spreading hairs above, puberulent 
below, branching basally from a woody caudex. Internodes elongated, up 
to 5 cm long. Leaves spreading or ascending, membranous, elliptical-ovate 
to somewhat rhomboid, 3.0-4.0 mm (3.9 mm) wide, 9.0-13.5 mm (11.0 
mm) long, margins entire, or obscurely crenate, apex acute, base narrowly 
attenuate petiolate, the petioles 2.0-4.0 mm long; margins and both sur- 
faces puberulent with short, stiff, spreading hairs; nervation indistinct, 
secondaries barely visible. Cymules 1-3-flowered, well-spaced along the 
upper 1/2 of the stems; peduncles ca 1.5 mm long; pedicels 3.54.0 mm 
long, densely hirsute with spreading hairs; bracteoles subulate shorter than 
the pedicels they subtend. Calyx ca 9.0 mm long; the tube ca 6.0 mm long, 
distinctly gibbous below, the distended region ca 1.0 mm wide, the tube 
dilated above, pubescent with short spreading hairs; the upper teeth con- 
nate for slightly less than 1/2 of their length forming a somewhat broad- 
ened, rounded upper lip ca 2.0 mm long, the margins more or less outwardly 
extended, the lobes reflexed, narrowly triangular-subulate, ca 0.3 mm wide 
at the base, ca 1.0 mm long, hirtellous-ciliate; the lower teeth triangular- 
subulate, slightly recurved, ca 0.4 mm wide at the base, ca 3.0 mm long, 
hirsute-ciliate; the annulus ca 0.7 mm wide but seated ca 1.0 mm below the 
juncture of the upper and lower teeth. Corolla 17.0-18.0 mm long, exserted, 
pubescent within but not annulate; the tube ca 15.0 mm long not dilated 
above. Nutlets ca 0.5 mm wide, ca 1.3 mm long, no mature nutlets seen. 
Chromosome number, unknown. 

sila Known only from the type locality. 

TYPE: BAJA CALIFORNIA. El Rancho Viejo, 30 Apr 1889, Brandegee 
SM. ease UC!; isotypes, PH! US! ). 

Several features of this rare species suggest an affinity with H. oblongi- 
folimm. However, until there is more material available for study H. tenwz- 
florum will be retained with H. costatum and its allies. 


28. HEDEOMA MARTIRENSE Moran, Trans. San Diego Soc. Nat. History 
Te 2127s 1909. 


Loose, wiry perennial herbs, 8.0-10.0 cm high forming mats or clumps. 
Shoots numerous, strongly decumbent and forming adventitious roots from 
the lower nodes; branching copiously throughout, the branches ascending 
at close angles; densely hispid-hirtellous above with spreading or retrorsely 
curving hairs up to 0.5 mm long, glabrate below with the lower portions 
becoming somewhat woody. Shoots of the previous season persistent. Leaf 
pairs crowded with the internodes 2.0-5.0 mm long, usually confined to 
the upper 1/4 of the shoots. Leaves membranous, spreading, ovate 2.0-3.0 
mm wide (2.7 mm), 5.0-6.0 mm long (5.4 mm), margins entire, apex 
obtuse, base petiolate, the petiole 1.0-1.5 mm long; leaf surfaces glabrous 
above and on the margins, strigulose below. Nerves raised. Flowers solitary 
in the upper leaf axils and restricted to the distil portions of the branches, 


pa) 


peduncles 2.0 mm long, pedicels 2.0-3.0 mm long, hirtellous with spread- 
ing hairs; bracteoles linear 1.0-2.0 mm long, shorter than the pedicels they 
subtend. Calyx 8.0-8.5 mm long, tubular, slightly sigmoid, saccate below 
for ca 1/3 of its length, ca 1.2 mm wide, flaring distally; the tube 5.5—6.5 
mm long, weakly strigulose below the hairs spreading; the upper lip con- 
nate for ca 1/3 of its length, the lobes triangular ca 0.5 mm wide at the 
base, ca 1.5 mm long, slightly reflexed, ciliate; lower teeth 0.7 mm wide 
at the base, 2.5 mm long, triangular-lanceolate, ciliate; annulus ca 0.5 mm 
wide, weakly developed, seated at the juncture of the upper and lower lips, 
included. Corolla white or pinkish, long and slender, 20.0—22.0 mm long, 
glabrous within; the tube ca 19.0 mm long dilated above, ca 2.5-3.0 mm 
wide; the upper lip obcordate, slightly galeate, ca 2.0 mm wide, 2.5-3.0 
mm long, shallowly notched; lower lip ca 4.0 mm wide, ca 3.0 mm long. 
Filaments quite short ca 3.0 mm long and arising just below the juncture 
of the upper and lower corolla lips (unique for Hedeoma). Nutlets brown, 
oblong, 0.5-0.7 mm wide, 1.4-1.6 mm long. Chromosome number, un- 
known. 

Distribution. North facing pine-fir covered slopes and on east facing open 
escarpments of N. Central Sierra San Pedro Martir, 7-9,000 ft. May—Aug. 

TYPE: BAJA CALIFORNIA. East rim of Sierra San Pedro Mare N. of 
Perba Buena, 1 June 1968, Moran 15069 (Holotype, SD 67322!; isotypes, 
K, MEXU, MONTU! UC, US). 

Hedeoma martirense is a rare and somewhat enigmatic taxon. In many of 
its morphic features, its large showy solitary flowers, its decumbent rhizo- 
matous habit, and its short leaf internodes, it shows strong phenetic affinities 
to H. diffuswm. In the remainder of its characters however, especially in its 
long, slender exserted corolla, and leaf structure it is most closely related 
to H. tenuiflorum, also endemic to Baja California. 


29. HEDEOMA QUERCETORUM Epl., Rep. Spec. Nov., Beihefte 115: 41. 
139. 

Lax perennial herbs, 15.0—30.0 cm high. Shoots few, ascending or shortly 
decumbent, branching basally with the branches widely ascending; puber- 
ulent-pubescent above with retrorsely curved hairs, glabrate below. Leaves 
spreading or ascending, membranous; the lower leaves ovate-elliptical, 
4.0-5.0 mm (4.5 mm) wide, 10.0-14.0 mm (12.0 mm) long, the upper 
leaves lanceolate or narrowly elliptic, at least four times as long as wide, 
2.0-3.0 mm (2.4 mm) wide, 12.0-30.0 (14.7 mm) long, margins entire 
or obscurely crenate, apex acute or narrowly obtuse, base narrowly attenuate 
and subsessile, the surfaces pubescent or glabrate above, sparsely pubescent 
below and on the margins; nerves more or less elevated. Cymules 1-—3- 
flowered, well-spaced along the upper 1/2 of the shoots, peduncles 0.5-1.0 
mm long; pedicels 3.0-6.0 mm long, pubescent or puberulent; bracteoles 
linear-lanceolate. Calyx 7.0-8.2 (7.7 mm) long; the tube 4.0-5.5 mm (4.9 


276 


mm) long, tubular-sigmoid and slightly gibbous below, the distended 
region 1.3-1.5 mm wide, densely hirsute pubescent with spreading hairs; 
the upper calyx teeth united less than 1/5 of their length forming an upper 
lip 1.5-2.0 mm long, the lobes narrowly triangular, 0.3-0.4 mm wide at 
the base, 1.5-1.8 mm long, spreading, erect or slightly reflexed, hirsute- 
ciliate; the lower teeth triangular-subulate, recurved, ca 0.5 mm wide at 
the base, 2.7—3.0 mm long, hirsute-ciliate; the annulus sparse, included, ca 
0.6 mm wide seated at the juncture of the upper and lower teeth. Corolla 
lavender, long exserted, ca 17.0 mm long, pubescent within the tube form- 
ing a broad annulus; the tube long and narrow, not at all dilated upwardly, 
15.0 mm long, 1.2 wide above; the upper lip obcordate-ligulate, emar- 
ginate, ca 1.0 mm wide, ca 1.5 mm long; the lower lip ca 5.0 mm wide, 
ca 3.5 mm long. Nutlets oblong-ovate, ca 0.6 mm wide, ca 1.2 mm long, 
the surface areolate and glaucous. Chromosome number, unknown. 

Distribution (Fig. 7). Oak woodland associations of the Sierra Madre 
Oriental of eastern Mexico in the states of Nuevo Leon and Tamaulipas. 
7000-9000 ft. Jul-Sep. 

TYPE: Nuevo LEON. Sierra Madre Oriental, San Francisco Canyon, ca 
16 miles SW of Pueblo Galeana, 7500-8000 ft, 16 May 1934, C.H. & MT. 
Mueller 411 (Holotype, LA!; isotypes, GH! TEX! ). 

Hedeoma quercetorum is very closely related to tenwipes. The morpho- 
logical differences reside principally in the number of flowers per cymule, 
the length of the calyx, and the length of the corolla. The variation in these 
characters as displayed by these two species, however, is no greater than 
that displayed in the single species H. costatum, but, until further field 
studies are carried out, they will be treated as distinct. 


30. HEDEOMAS TENUIPES Epl., Rep. Spec. Nov., Beihefte 115: 39. 1939. 


Lax perennials, 25.0—30.0 cm in height. Shoots strict or ascending, branch- 
ing primarily in the lower half with the branches widely ascending from 
the main axis; stems pubescent-puberulent with retrorsely curling hairs 
above, glabrate below. Leaves spreading, membranous; the lower leaves 
broadly ovate-elliptical 4.5-8.0 mm (6.1 mm) wide, 1.0-1.7 mm (1.3 mm) 
long; the upper leaves linear-lanceolate, at least five times longer than wide, 
1.8-3.0 mm (2.2 mm) wide, 12.0-17.0 mm (15.3 mm) long, margins en- 
tire or obscurely crenate, apex acute, base narrowly attenuated and sub- 
sessile, surfaces glabrate above, pubescent below with short hairs on the 
nerves and margins; nerves slightly raised. Cymules 3- over 20-flowered, 
well-spaced along the upper 1/2 of the stems, occasionally congested, 
peduncles ca 1.0 mm long; pedicels 4.5-6.7 mm long, sparsely pubescent; 
bracteoles linear, nearly equal to the pedicels they subtend. Calyx 5.0-6.0 
(5.6 mm) long; the tube 3.0-3.7 mm (3.6 mm) long, tubular and not 
at all gibbous below, or only slightly so, ca 1.0-1.2 mm wide, hirtellous 
pubescent with the hairs spreading or antrorsely curved; the upper calyx 


277 


teeth united less than 1/5 of their length forming an upper lip 1.0-1.5 
mm long, the lobes narrowly triangular, ca 0.3 mm wide at the base, 0.9- 
1.2 mm long, slightly reflexed, hirsute-ciliate; the lower teeth triangular- 
subulate, erect or slightly recurved, ca 0.4 mm wide at the base, 2.0-2.5 
mm long, hirsute-ciliate; the annulus, included, sparse, ca 0.5 mm wide, 
seated at the juncture of the upper and lower teeth. Corolla small, white, 
barely exserted from the calyx tube, glabrous within, ca 5.5 mm_ long; 
narrowly tubular, ca 1.0 mm wide above; the upper lip ligulate, ca 1.0 mm 
wide, ca 0.8 mm long, emarginate; the lower lip ca 2.0 mm wide, 1.2 mm 
long. Nutlets ca 0.6 mm wide, ca 1.3 mm long, the surface areolate, glau- 
cous. Chromosome number 27 = 36. 

Distribution (Fig. 7). Oak woodlands of the Sierra Madre Oriental, 
Tamaulipas, Mexico. 2000-5000 ft. Jul-Sep. 

TYPE: TAMAULIPAS. “Cerro Zamora near El Milagro,” 25 Aug 1930, 
Bartlett 1117 (Holotype, F!; isotypes, MICH! LA!; phototype LA! ). 


31. HEDEOMA HYSSOPIFOLIUM Gray, Proc. Am. Acad. L1: 96. 1876. 


Plants perennials forming dense, wiry tufts or large clumps, 15.0-47.0 
cm (29.1 cm) high. Shoots numerous, ascending, or decumbent, rhizomatous, 
producing adventitious roots at the nodes of the short subterranean por- 
tions of the stems; the upper regions glabrate or pubescent, but usually 
puberulent with retrorsely curved hairs, puberulent or hirtellous below; 
branching almost exclusively confined to the basal regions. Leaf pairs more 
or less crowded, equal to or exceeding the internodes. Leaves glabrate 
membranous, narrowly lanceolate to linear, at least five times longer than 
wide, and commonly more, 1.0-2.0 mm (1.8 mm) wide, 8.0-18.5 mm 
(13.4 mm) long, margins entire, the apex narrowly acute, the base tapered 
and sessile; nerves prominent but not strongly elevated. Axillary buds form 
a cluster of 2-3 axillary leaf pairs. Cymules 1—7-flowered (typically 3- 
flowered), well-spaced or somewhat congested along the upper 1/2 of the 
shoots; peduncles 0.5—-3.0 mm long; pedicels 3.0-5.0 mm long, puberulent 
to pubescent; bracteoles variable, subulate-lanceolate, typically exceeding 
the length of the pedicels they subtend. Calyx long and slender, 6.0-8.3 mm 
(7.5 mm) long, the calyx tube 4.9-5.5 mm (4.2 mm) long only slightly 
or not at all gibbous below, tubular-funnelform, dilated above, 1.0-1.5 
mm wide, typically glabrous or puberulent with short stiff spreading hairs 
in the proximal region, glabrous distally; the upper calyx teeth connate for 
ca 1/5 of their length forming an upper lip 1.0-1.6 mm long, the lobes 
narrowly triangular to subulate, ca 0.2-0.5 mm wide at the base, 0.9-1.6 
mm long, erect and parallel or slightly laterally spreading, glabrous or 
tuberculate on the margins; the lower teeth long subulate, recurved, ca 
0.2 mm wide at the base, 1.7—-3.0 mm long, hirsute or hirtellous-ciliate; 
annulus dense, 0.7-1.0 mm wide, exserted, seated approximately at the 
juncture of the upper and lower teeth and extended upwardly onto the in- 


278 


nerfaces of the teeth. Corolla lavender; 11.0-16.0 mm (14.1 mm) long, 
pubescent within the tube but not annulate; the tube dilating upwardly, 
but not abruptly so, 2.0-3.0 mm wide above; the upper lip broadly ligulate 
in Outline, emarginate, with margins outwardly flaring, 1.5-4.0 mm wide, 
2.2-5.0 mm long; the lower lip up to 7.0 mm wide, 5.0 mm long. Nutlets 
ca 0.6 mm wide, ca 1.3 mm long, the surface areolate, weakly glaucous. 
Chromosome number, 27 = 36 

Distribution (Fig. 8). A common member of the herbaceous understory 
flora of mature, pine stands in the mountains of southeastern Arizona, 
southwestern New Mexico and northern Mexico, lower montane to sub- 
alpine associations, 6000 to 10,000 ft. Most commonly a member of the 
upper montane association and found frequently in recently burned areas. 
Aug—Sep. 

TYPE: ARIZONA. Graham Co.: Mt. Graham, 9000 ft. Aug eee pie 
rock 418 (Holotype, GH!; Fragment, LA!; isotypes, F! K! 

US!; phototype LA! ). 

Hedeoma hyssopifolium is probably the most easily recognized taxon in 
the genus Hedeoma. This distinctiveness is due to its narrow, linear leaves, 
cylindrical, puberulent calyx, and large showy flowers. Although readily 
recognized, it nevertheless houses considerable variation. The most notable 
variation is revealed in the disjunct populations of northern Mexico (Fig. 
8). These populations are easily recognized and perhaps deserve varietal 
status. Their glabrous stems, subulate-filiform calyx teeth, and sparce annulus, 
all combine to set these plants apart from their more northwestern counter- 


arts. 

Hedeoma hyssopifolium intergrades into H. oblongifolium and H. denta- 
tum, most often through hybridization at the lower altitudes where these 
species come is contact. 


32. HEDEOMA OBLONGIFOLIUM (Gray) Heller 


Key to the varieties: 
1. Corolla glabrous within, leaves broadly a plants of U.S. 
sapcates 2a. H. oe var. oblongifolium 
1. Corolla annulate within, leaves eae elliptical, plants of Mexico. .....00..0......... 
32b. H. ea var. mexicanum m 


32a. HEDEOMA OBLONGIFOLIUM (Gray) Heller var. OBLONGIFOLIUM, 
Muhlenbergia 1: 1900. 
Hedeoma piperitum Benth. var. oblongifolium A. Gray, Proc. Am. Acad. 8: 366. 
1872. Basion 
rc thymoides A. Gray var. oblongifolinm A. Gray, Syn. Fl. N. Am. 2: 362. 


Stout, herbaceous perennials, 20.0-55.0 cm high, forming robust tufts. 
Shoots few to many, usually under 10, densely pubescent above with retrorsely 
curling hairs to hoary-pilose above with long, spreading hairs, puberulent 


below, stems ascending strictly from a slender rootstock; branching con- 
fined to the basal regions with secondary branches few and arising well 
after the maturity of the primaries. Leaves spreading, membranous, nar- 
rowly elliptic-ovate or linear-elliptic, at least twice as long as wide, 5.5 
mm (4.3 mm) wide, 8.5-14.0 mm (11.9 mm) long, margins denticulate, 
apex narrowly and sharply acute, base tapered and shortly petiolate with 
the petiole 0.9-1.5 mm long; the surfaces glabrate or pubescent above, 
undersurface and margins somewhat unevenly hirtellous; nerves more or 
less raised. Basal foliage distinct, broadly ovate, entire, glabrous above, 
minutely hirtellous below. Axillary buds arise early in the shoot develop- 
ment and form a cluster of 2—3 leaf pairs. Cymules 5—15-flowered, dense 
and congested along the upper 2/3 of the shoots; peduncles inconspicu- 
ous to 0.5 mm long; pedicels 4.0-7.0 mm long and commonly exceeding 
the length of the calyx tube, sparsely to densely hirtellous; bracteoles 
subulate-lanceolate ca equal to the length of the pedicels they subtend. 
Calyx 5.5-7.0 mm (6.2 mm) long; the tube 4.0-5.0 (4.5 mm) _ long, 
gibbous below for ca 1/3 of the tube’s length, the distended region 
1.0-1.2 mm wide, hirtellous of short spreading hairs to densely hirsute- 
pilose; the upper calyx teeth connate for ca 1/5 of their length forming an 
upper lip 1.0-1.3 mm long, the lobes narrowly triangular, 0.1-0.3 mm 
wide at the base, 0.7-1.0 mm long, laterally spreading and reflexed, ciliate; 
the lower teeth subulate, recurved, ca 0.2 mm wide at the base, 1.3—2.0 
mm long, hirsute-ciliate; the annulus dense somewhat exserted, ca 0.5 mm 
wide, seated approximately at the juncture of the upper and lower teeth. 
Corolla lavender, 10.0-11.0 mm (10.5 mm) long, glabrous within; the 
tube 8.0-9.0 mm (8.5 mm) long and only slightly dilated upwardly, ca 
1.5 mm wide above; the upper lip, flat, ligulate, ca 1.2 mm wide, ca 2.0 
mm long, emarginate, conspicuously shorter than the lower lip, and reflexed 
away from the axis; the lower lip ca 4.0 mm wide, ca 4.0 mm long. Nutlets 
ca 0.5 mm wide, ca 1.0 mm long, the surface areolate and weakly glaucous. 
Chromosome number, 27 = 36 

Distribution (Fig. 8). Common on the slopes of the oak-woodland, 
chaparral associations in mountainous regions of central and southern Ari- 
zona, New Mexico and adjacent areas of northern Sonora, Mexico. 5000- 
8000 fr. Aug—Sep. 

PE: New Mexico. 1851-52, Wright 1515 (Holotype, NY!; isotypes, 

BM! F! GH! K! PH! UC! US!) 

Hedeoma oblongifolium is more or less intermediate between the mesic 
montane species H. byssopifoliwm and the xerophytic H. dentatum. 


32b. HEDEOMA OBLONGIFOLIUM Ao Heller var. MEXICANUM Irving, 
Brittonia 22: 342-344. 1970 


Stout perennials 28.0-40.0 cm high, forming tufts. Shoots ascending, 
usually under 10, pubescence of tightly retrorsely curling hairs above, 


280 


puberulent below, branching mostly confined to the lower regions of the 
stem with the secondary branches long and ascending at close angles. Leaves 
spreading, thick membranous, narrowly elliptical or linear, at least three 
times as long as wide, 1.5-4.0 mm (2.8 mm) wide, 9.0-13.0 mm (10.8 


Fig. Distribution of Hedeoma hyssopifolium (open squares), H. oblongifolinm 
var. ae (closed circles), H. oblongifolium var. mexicanum (open circles), 
and H. dentatum (open triangles). 


Zed 


mm) long, the margins denticulate with the teeth confined to the upper 
regions of the blade, the apex sharply acute, the base tapered and sub- 
sessile; surfaces glabrous to glabrate above, glabrate below; nerves elevated. 
Basal foliage distinct, narrowly elliptic to ovate, entire, and evenly glab- 
rate. Axillary buds arise early in the shoot development and form a 
cluster of 2-3 leaf pairs. Cymules 5-10-flowered, dense and congested, 
well-spaced or crowded along the upper 2/3 of the shoots; peduncles in- 
conspicuous; pedicels, 4.0-5.0 mm long, puberulent; bracteoles lanceolate, 
ca equal to the length of the pedicels they subtend. Calyx 5.2-5.8 mm 
(5.5 mm) long; the tube 3.7-4.2 mm (4.0 mm) long, gibbous below for 
ca 1/2 of the tube’s length, the distended region 1.0-1.2 mm wide, slightly 
tapered above, hirtellous with short, stiff, spreading hairs; the upper calyx 
teeth only slightly united for ca 1/5 or their length forming an upper lip 
0.9-1.0 mm long, the lobes narrowly triangular, subulate, 0.1-0.2 mm wide 
at the base, 0.8-0.9 mm long, laterally spreading and reflexed; the lower 
teeth subulate, ca 0.3 mm wide at the base, 1.2-1.7 mm long, hirsute-ciliate; 
annulus dense partially exserted, ca 0.5 mm wide and seated approximately 
at the juncture of the upper and lower teeth. Corolla lavender, 8.0-9.0 mm 
long, annulate within of short pilose hairs seated at the juncture of the 
tube and throat; the tube 6.8-7.8 mm (7.0 mm) long, slightly dilated 
upwardly, ca 2.0 mm wide above; the upper lip ligulate, emarginate, ca 10 
mm wide, 1.2 mm long, conspicuously shorter than the lower lip, and re- 
flexed away from the axis; the lower lip ca 3.0 mm wide, ca 3.0 mm long. 
Nutlets ca 0.5 mm wide ca 1.0 mm long, the surface areolate and glaucous. 
Chromosome number, unknown. 

Distribution (Fig. 8). Common in the oak-woodland and chaparral asso- 
ciations in the mountains of northern Sonora and Chihuahua, Mexico. 
6000-9000 ft. Aug—Oct. 

TYPE: SONORA. Puerto del Cumarito, Sierra de la Cabellera, 5400 ft. 6 
Oct 1941, White 4638 (Holtoype, MICH!; isotypes, GH! US.). 


33. HEDEOMA DENTATUM Torr., Bot. U.S. Mex. Bound, Surv., part 2, 130. 


Tall herbaceous perennials, 29.0-40.0 cm tall, forming robust clumps. 
Shoots long, few to many, densely hirtellous pubescent with spreading or 
curling hairs; branching confined to the lower 1/2, the branches long, ascend- 
ing and arising well after the maturity of the primary shoots. Leaves 
spreading, ovate, elliptic, or rhombic, at least twice as long as wide, 3.0-5.0 
mm (4.1 mm) wide, 8.0-17.0 mm (10.3 mm) long, margins coarsely 
serrate-dentate, apex sharply acute, base tapered and subsessile, upper sur- 
face sparsely strigulose to glabrate, under-surface and margin hirtellous with 
short, stiff, spreading hairs; nerves raised. Basal foliage broadly ovate or 
obovate, coarsely serrate with both surfaces densely and evenly hirsute-pilose. 
Axillary buds develop early in the shoot development and form a cluster of 


282 


2-3 leaf pairs. Cymules 5-15-flowered (typically between 5-7), dense and 
congested along the upper 1/2 of the shoots; peduncles 0.2-1.1 mm long; 
pedicels 3.2-4.5 mm long and commonly exceeding the length of the calyx 
tube, pubescent; bracteoles lanceolate equal in length to the pedicels they 
subtend. Calyx 4.8-5.5 mm (5.2 mm) long; the tube 3440 mm (3.8 
mm) long, gibbous or slightly saccate for 1/3-1/2 of the tube’s length, 
0.9-1.2 mm wide below, slightly tapered above, hirtellous with short, stiff, 
spreading hairs; the upper teeth slightly connate for ca 1/5 of their length 
forming an upper lip 1.0-1.6 mm long, the lobes subulate, ca 0.2 mm 
wide at the base, 0.7-0.9 mm long, laterally spreading and reflexed, hirtellous- 
ciliate; the lower teeth subulate, recurved ca 1.5-1.7 long, hirsute-ciltate; 
annulus dense somewhat exserted, ca 0.5 wide, seated approximately at 
the juncture of the upper and lower teeth. Corolla lavender, 8.0-9.0 mm 
(8.5 mm) long, annulate within with short pilose hairs; the tube 6.5-7.5 
mm (7.4 mm) long, only slightly dilated upwardly, ca 1.5 mm wide above; 
the upper lip flat, ligulate, ca 1.2 mm wide, 1.5 mm long, emarginate, 
conspicuously shorter than the lower lip, and reflexed away from the axis; 
the lower lip ca 3.5 mm wide, ca 3.5 mm long. Nutlets ca 0.5 mm wide, 
1.0 mm long, the surface areolate and glaucous. Chromosome number, 27 


Distribution (Fig. 8). Common on open, well-drained soils in the oak 
woodland and chaparral associations in the mountains of central and south- 
eastern Arizona, and adjacent Mexico, SO00—10,000 ft. Aug—Oct. 

TYPE: SONORA. Santa Cruz, Sep 1851, Thurber s.n. (Holotype, NY!; 
isotypes, F! GH! US!). 

Hedeoma dentatum intergrades with both H. hyssopifolium and H. ob- 
longifolium. Hybridization probably accounts for the latter as there is ac 
least one documented case of gene exchange between H. dentatum and 
H. hyssopifolium in the Huachuca mountains. 

Epling and Stewart (1939) have postulated that H. dentatum is the 
result of past hybridization between H. oblongifolium and H. plicatum. 
However, it seems more likely that H. dentatum was selected out of 
peripheral populations of H. oblongifolium in response to xeric conditions. 


34, HEDEOMA ACINOIDES Scheele, Linnaea 22: 592. 1849. 


Delicate, ephemeral annuals, 5.0—30.0 cm (15.2 cm) high, arising from a 
very slender taproot. Shoots solitary to numerous, erect, ascending or very 
shortly decumbent for 1.0-2.0 cm; branching absent or restricted to the 
lower 1/2 of the shoots with the branches ascending from the main axis at 
wide angles; densely pubescent above, nearly lanate with the hairs strongly 
retrorsely curving or curling, puberulent below. Leaves spreading or cernu- 
ous, membranous, narrowly or broadly ovate, elliptical, or rhombic-elliptical, 
3.5-8.0 mm (5.8 mm) wide, 9.0-25.0 mm (14.2 mm) long, margins en- 
tire or obscurely serrate with the serrations restricted to the distil 1/2 of 


283 


the blade, apex obtuse, base rounded to narrowly attenuate, petiolate with 
the petiole 1.0-4.0 mm (2.6 mm) long; the surfaces glabrate or minutely 
strigulose on nerves and margins; nerves except for the midrib unelevated. 
Cymules 3—15-flowered and typically congested toward the apices forming 
a short dense spike of cymules; peduncles inconspicuous, less than 1.0 mm 
long; pedicels 2.0-5.0 mm long, shortly pubescent; bracteoles lanceolate- 
subulate, ciliate, the primary pair equal in length to the pedicels they sub- 
tend. Calyx 5.0-6.1 mm (5.3 mm) long, thin membranous; the tube rela- 
tively long and slender, 4.0-4.55 mm (4.1 mm) long, the lower 1/3 
distinctly saccate below, the distended region ca 1.2 mm wide, the tube 
giabrate or sparsely hirtellous on the lower 1/2 and with the hairs confined 
to the primary nerves; upper calyx teeth united for ca 1/3 of their length 
forming small but definite upper lip, 0.9-1.2 mm long, the lobes broadly 
to narrowly triangular, ca 0.2 mm wide at the base, 0.8-1.1 mm long, 
parallel or only slightly reflexed, densely hirtellous-ciliate; the lower teeth 
narrowly triangular to subuate, recurved, ca 0.4 mm wide at the base, 1.0-1.2 
mm long, hirtellous-ciliate; the upper and lower teeth partially closing the 
orifice through convergence; annulus very weakly represented, consisting of 
only a few short hairs, ca 0.4 mm wide, seated ca 0.4 mm below the junc- 
tion of the upper and lower teeth. Corolla pink, 9.0-13.0 mm (11.0 mm) 
long, the throat very slightly dilated upwardly ca 0.6 mm wide above; the 
upper lip ligulate and reflexed at right angles to the corolla tube, very slightly 
emarginate, 1.0-1.5 mm wide; 2.0-4.0 mm long; the lower lip 4.0-5.0 mm 
wide, 3.0-4.0 mm long. Nutlets small, 0.4 mm wide, ca 0.8 mm long, the 
surface areolate, not glaucous or only very slightly so. Chromosome number, 
2n = 34 

Distribution (Fig. 9). Common, in wet years, in the rocky limestone 
soil along the eastern periphery of the Edwards Plateau and Lampasas Cut 
Plain. It also occurs in limestone pockets of the South Coastal Plains of 
Texas and in the calcareous hills surrounding Monterrey, Mexico. Apr- 
May. 

TYPE: TExas. Comal Co., in the vicinity of New Braunfels, April, 
Roemer sn. (Holotype, not located ). 

Within this species a form can be recognized that is small, more heavily 
pigmented, glabrate and with entire leaves. Based on field observations 
in Travis Co., Texas, this form is apparently adapted to more xerophytic 
locations and often forms subpopulations several inches to several yards 
in, Size: 

In its interspecific relationships, H. acinoides is somewhat of an enigma. 
Its morphological attributes, in combination, are quite unique. Although 
Epling and Stewart (1939) have suggested an affinity with H. guercetorum, 
I can find no morphological or chemical substantiation for such an idea. 
In its overall habit and vegetative morphology, H. acimoides shows some 
arity with the eastern annual, H. pulegioides. The calyx, corolla, and 


=, 


simi 


284 


seed morphology, however, readily separate H. acinoides from this latter 
species. 

H. acinoides has the same chromosome number (2” = 34) as the par- 
tially sympatric taxa, H. reverchonii, Hr. var. serpyllifolium, H. drummondiu 


rT | | 
PP ty 
e 
Piet | 
r 
. e 
2 82 e 4 
ry 
e e 
* e e e 
e e 
: e ee ‘ 
~ e le ry 
“ye > , seen) ° 
8 © . e 
ae e e he 
\ e ; 
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ee . e 
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. eee WF : 


Fig. 9. Distribution of Hedeoma acinoides (closed circles). 


285 


(in part), and H. hispidum. As the latter 4 taxa are apparently closely 
related, it is quite possible that H. acinoides is a highly derived species 
stemming from the H. drummondii complex. Morphological support of 
this is seen in the convergence of the upper and lower calyx teeth, a feature 
also characteristic of H. drummondii and its allies. 


35. HEDEOMA HISPIDUM Pursh, Fl. Am. Sept. 2: 414. 1814. 

Hedeoma hirtum Nutt., Gen. 1: 16. 1818. Nom. superfl. TYPE: ARKANSAS 
{Terr.?}. On open alluvions of the Missouri, June. Nwétall s.n. (Lectotype. here 
designated GH!; isotype, K!). 

Ziziphora hispidum (Pursh) R. & S., Syst. Mant. 1: 179. 1822. Based on H. 
hispidum. 

Hedeoma hispidum Pursh forma simplex Lalonde, Nat. Canad. 85: 71. 1958. TYPE: 
QUEBEC. Pontiac Co., Vinton, 27 Aug 1956, P. Lowis-Marie 5614 (Holotype, 
CAN; isotype, GH! ). 

Coarse annuals, 9.0-40.0 cm (21.6 mm) in height, arising from a very 
slender taproot. Shoots slender, sharply quadrangular, usually solitary, occa- 
sionally numerous, erect or shortly decumbent for 2.0-3.0 cm; branches 
absent or consisting of 2-6 pairs, long ascending and usually confined to the 
lower 1/3 of the plant; shoots densely pubescent above with relatively long, 
retrorsely curling hairs, becoming puberulent below. Leaves typically spread- 
ing, coriaceous with the margins frequently becoming revolute, linear or 
linear-elliptic, usually over 5 times longer than broad, 1.0-3.0 mm (2.2 
mm) wide, 11.0-21.0 mm (16.4 mm) long, margins entire, apex obtuse, base 
narrowly attenuated and subsessile, surfaces glabrate above, strigulose below 
and on the margins; nerves not conspicuously elevated. Cymules born along 
the upper 2/3 of the stems, occasionally along its entire length, well-spaced 
or commonly congested forming a pronounced “spike”, 3-15-flowered but 
typically 5-8-flowered; peduncles inconspicuous less than 1.0 mm in length; 
pedicels 2.0-5.0 mm long, hirsute; bracteoles stiff and acute, narrowly 
lanceolate to subulate, hirsute-ciliate, longer than the pedicels they subtend. 
Calyx highly zygomorphic, 4.9-6.0 mm (5.2 mm) long; the tube 2.8-3.2 
mm (2.9 mm) long, markedly saccate below for ca 3/4 of its length, 
1.5-2.0 mm wide below, pubescent with hairs long hirsute and spreading; 
the upper calyx teeth connate for ca 1/2 of their length forming a broad, 
expanded upper lip, 1.2-2.0 mm long with margins outwardly flaring and 
exceeding the width of the tube below, the connate portion silky pubescent 
within on its inner face, the lobes narrowly triangular, laterally spreading, 
hirtellous-ciliate, ca 0.5 mm wide at the base, 0.5—0.9 mm long; the lower 
teeth recurved and subulate, 2.0-2.5 mm _ long, hirsute-hirtellous-ciltate; 
annulus dense, slightly exserted, ca 0.5 mm wide and seated at the juncture 
of the upper and lower teeth. Corolla dimorphic; the first form blue or 
white, small, disproportionately so in reference to the calyx, ca 5.3 mm 
long and scarcely exserted from the calyx tube, glabrous within. The second 
form blue, considerably larger, ca 6.0-7.0 mm long, and well exserted from 


286 


the calyx, glabrous within, the pe lip subgaleate, the lower spreading. 
utlets ca 0.6 mm wide, ca 1.2 mm long, the surface areolate and strongly 
glaucous. Chromosome number, on = =: 

Distribution (Fig. 10). Rich, loamy soil of prairie and pasture lands 
from Vermont west to Alberta, Canada, south to south-central Texas and 
eastern Florida, along the Gulf coastal plain. Apr—June 

PE: On the banks of the Missouri, New¢tall s.n. (Holotype, PH! ). 

Hedeoma ae is one of the most widespread of the taxa of Hedeoma, 
rivaled only by H. drummondii and H. puleg‘oides in total area of distri- 
bution, 

Because of its breeding behavior, its annual habit, and chromosome num- 
ber, H. hispidum is believed to be derived. Its morphology can be traced to 
two possible precursors—H. nanum ae H. drummondii, themselves closely 
related to one another. 

In past studies H. bispidum has always been associated with H. pulegioides 
to which it displays a close similarity in its overall morphology. However, 
differences in nutlet structure, fine details of the calyx, corolla, and chromo- 
some number clearly set H. hispidum apart from the latter (see discussion 
following treatment of H. pulegioides). 


IV. SUBGENUS HEDEOMA 


Loose, wiry perennials of robust annuals, 5.0-40.0 cm high. Shoots solitary 
to numerous, erect, ascending or shortly decumbent, arising from a slender 
taproot. Leaves well-spaced, membranous and spreading, ovate, or elliptical, 
3.5-11.0 mm wide, 5.0-32.0 mm long, margins dentate or crenate, base 
petiolate, the leaf surfaces strigose or glabrous, glandular dotted. Flowers 
in 1-15-flowered axillary cymules. Calyces strongly zygomorphic, 0 
mm long, the tube weakly to strongly saccate for over 1/2 of its length; 
the upper teeth united for 1/2 to 2/3 of their length to form a broad, ex- 
panded, upper lip, the lobes semi-ovate, deltoid, or triangular, erect or re- 
flexed; the lower teeth short scarcely exceeding the upper lip in length, 
ciliate. Corolla short, 3.0-8.5 mm long (in H. pulegioides and H. crenatum 
barely exserted from the calyx tube); the upper lip broadly ligulate, deeply 
emarginate (for ca 1/2 of its length); the lower lip not spreading, more 
or less held erect, nearly equally 3-lobed. Nutlets orbicular in outline or 
nearly so, black in color at maturity, the surface ruminate, not becoming 
mucilaginous when moistened, not glaucous. Chromosome number, 27 = 

e subgenus Hedeoma is composed o species which, despite their 
wide geographical separation, appear to form a natural unit. The salient 
morphological features which hold the group together are the highly 
zygomorphic calyx, small corolla, and orbicular, black, nonmucilaginous 
nutlets. 


Fig. 10. Distribution of Hedeoma hispidum. 


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36. HEDEOMA PULEGIOIDES (L.) Pers., Syn. 113. 1807. Based on Melissa 
pulegioides. 

Melissa pulegioides L., Sp. Pl. ed 1,593. 1753. 

Cuntla pulegioides L., Sp. Pl. ed. 2, 30. 1772. Based on Melissa pulegioides. 

Ziziphora pulegivides (L.) Desf., Table. ed. I: 55. 1804; ed. II: 66. 1815. Based on 
Melissa pulegioides. 

Hedeoma Met gee (L.) Pers. forma simplex Lalonde, LaRevue D’Oka 32: 151. 
958. TYPE: QUEBEC. Powe 10 Aug 1937. Desilets s.n. (Holotype, 


Robust annuals, 10.0-40.0 cm high, arising from a slender taproot. 
Shoots typically solitary, erect, straight, branching freely in the middle 
region with the branches long and ascending, densely puberulent with 
retrorsely curling hairs above, glabrate below. Leaves spreading or cernuous, 
deciduous from the lower 1/2 of the plant with maturity, membranous, 
narrowly to broadly elliptical or nearly oblong, 4.0-11.0 mm (5.9 mm) wide, 
12.0-32.0 mm (19.0 mm) long, margins obscurely to sharply dentate, apex 
narrowly obtuse, base attenuated and petiolate, the petiole 2.5-7.0 mm 
long; leaf surfaces glabrous or minutely strigulose on the nerves; nervation 
not prominent. Cymules 3-15-flowered, typically well-spaced along the 
upper branches, occasionally congested; peduncles 0.5-1.0 mm long; pedicels 
3.0-3.5 mm long, puberulent; bracteoles shorter than the pedicels. Calyx 
highly zygomorphic, 3.5-4.5 mm (4.1 mm) long; the tube 2.0-2.5 mm 
(2.2 mm) long, markedly saccate below for ca 2/3 of its length, the dis- 
tended region 1.3—1.6 mm wide, sparsely hirtellous or glabrate below with 
the hairs spreading; the upper calyx teeth connate for ca 2/3 of their length 
forming a broad, expanded, upper lip, 2.0-2.5 mm long with the margins 
rounded, and exceeding the width of the tube below, the connate portion of 
the lip glabrous on a inner face, the lobes narrowly triangular, ca 0.5 
mm wide at the base, 0.5-0.7 mm long, laterally spreading and reflexed, 
glabrous on the eee the lower teeth short, barely exceeding the upper 
lip in length, subulate and recurved, ca 0.5 mm wide at the base, 1.5-2.5 mm 
long, hirsute-ciliate; the annulus sparse, included, ca 0.5 mm wide and 
seated at the juncture of the upper and lower lips. Corolla minute, barely 
exserted from the calyx tube, ca 4.0 mm long, glabrous within; the tube 
ca 3.0 mm long, dilated core, ca 1.5 mm wide above; the upper lip 
ligulate, ca 0.8 mm wide at the base, ca 1.0 mm long, straight, deeply 
emarginate; the lower lip erect, eee 3-lobed, ca 2.0 mm wide, 1.0 mm 
long; nutlets 0.9 mm wide, 1.0 mm long, black, the surface ruminate, not 
glaucous. Chromosome number 27 = 36. 

Distribution (Fig. 11). Open woodlands and waste ground throughout 
central and eastern U. S., eastern Kansas and Oklahoma eastward to South 
Carolina and northward to Nova Scotia, Canada. Aug—Sep. 

TYPE: Plants of the Gronovian Herbarium of which Clayton 514 has 
been suggested as the standard (Epling 1939) (Lectotype, here designated 
BM). 


bo 
Cc 
\o 


Hedeoma pulegioides is a very widespread and distinct species which has, 
traditionally, been grouped with H. hispidum. Indeed, these two species form- 
ed a separate section in the revision by Epling and Stewart (1939). Although 
there are several morphological features shared by these species, notably 
the annual habit and the saccate calyx, their differences, as for example the 
features of the nutlet, outweigh their similarities. 

Hedeoma pulegioides is most closely related to H. crenatum of Brazil. 
Only habit and leaf shape distinguish these two taxa from one another. 


coor 


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Fig. 11. Distribution of Hedeoma pulegioides. 


290 


37. HEDEOMA CRENATUM Irving, Britt. 22: 344-345. 1970. Based on 
Pseudocunila montana Brade. 


Hedeoma Ee le Benth. var. montannm Dusén, Arch. Mus. Nac. XIII. 35. 
903. TYPE: BRAZIL. Serra do Itatiaia, March 1894. Ule 199 (Holotype, R). 

Pseudocunila montana Brade, Rodriguesia 7: 27. 1944. 

Cunila montana Brade ex Epl., Britt. 7: 140. 1951. TYPE: BRAZIL. Rio de Janeiro, 
Serra dos Orgaos, Estado do Rio Castello, 2100 m, 17 Jan 1945. Segadas Vianna 
675 (Holotype, LA!). 

Hedeoma montanum (Brade) Epl. & le Britt. 15: 369. 1963 (not Brandegee 
1913). Based on Cunila montana Brade 


jon 


Loose, wiry perennials, perhaps biennials, 15.0-30.0 cm high. Shoots 
numerous, typically decumbent and rhizomatous, producing adventitious 
roots from the lower nodes, densely pubescent with spreading or retrorsely 
curving hairs, branching mostly at or near the base. Leaves spreading, 
occurring all along the stems, membranous, ovate-spatulate, 4.0-5.0 mm 
(4.6 mm) wide, 9.0-10.0 mm (9.5 mm) long, glabrous, margins crenate 
especially along the upper 1/2 of the blade, apex obtuse, base rounded and 
petiolate with the petiole 2.5-3.0 mm _ long; nervation inconspicuous. 
Cymules 1-3-flowered, well-spaced along the upper 1/2 of the stems; 
peduncles inconspicuous; pedicels ca 2.5 mm long and puberulent; bracteoles 
shorter than the pedicels they subtend. Calyx highly zygomorphic, ca 4.0 
mm long; the tube ca 2.0 mm long, hirtellous, markedly saccate for over 
3/4 of its length, the distended region ca 1.0 mm wide; the upper calyx 
ag connate for ca 1/2 of their length rn an expanded, upper lip, 

0 mm long, with the margins rounded, the lobes semi-ovate, ca 0.7 
mm wide at the base, ca 1.0 mm long, more or less erect, hirtellous-ciliate; 
the lower teeth short barely exceeding the upper lip in length, narrowly 
triangular, ca 0.5 mm wide at the base, ca 2.0 mm long, hirtellous-ciliate, 
erect or slightly recurved; annulus dense, included, ca 0.5 mm wide, seated 
just below the juncture of the upper and lower teeth. Corolla minute, barely 
exserted from the calyx tube, ca 3.0 mm long, pubescent within; the tube 
ca 2.0 mm long, slightly dilated upwardly, ca 1.0 mm wide at the summit; 
the upper lip broadly ligulate ca 1.5 mm wide, 1.0 mm long, straight, flat, 
and deeply emarginate; the lower lip erect, nearly equally 3-lobed, ca 2.0 
mm wide, ca 1.0 mm long. Nutlets 0.8 mm wide, 1.0 mm long, the surface 
ruminate, and not glaucous. Chromosome number, unknown. 

Distribution (Fig. 12). Brazil, in the mountains of Minas Geraes and 
Rio de Janeiro. 6,000-7,000 ft. Nov—Jan. 

TYPE: BRAZIL. Rio de Janeiro, Serra dos Orgaos, 2200 m, A.C. Brade 
11,500 (Holotype, RF! ) 


38. HEDEOMA MANDONIANUM Wedd., Chlor. And. 2: 148. 1860. 


Hedeoma rev iflorum Griseb. in Lechler, Berb. Am. Aust. 58. 1857, nomen. TYPE: 
U gapata,”’ Jun 1854, Lechler 1848 (Holotype, K! 
Hedeoma adscendens Rusby, Phytologia 1: 74. 1934. TYPE: "BOL OVIA. Cordillera 


rap Al 


= 
a 6G 
on 
= 
| 
= 
e 
| a 
e a 
e 
e® e 
Ce of 
7 Ad 
x = : y 


He 12. Distribution of Hedeoma ee (closed circles), H. medium (open 
crenatum (open square), mandonianum (closed square) and H. 


ne 
ee (asterisk), 


292 


Real, top of the pass on the Tipuani-Ancoma-Sorata Trail, down to Sorata, 
13,000 ft, 20 Apr 1936, Tate 783 (Holotype, NY!; isotypes, GH! LA! PH! US!; 
phototype, LA!) 

Loose, densely branched perennials, 5.0-15.0 cm high. Shoots numerous, 
intertangled, ascending or shortly decumbent, rhizomatous, sending out ad- 
ventitious roots from the lower nodes, hirtellous above with short, stiff, 
spreading hairs, puberulent below. Leaves spreading or ascending, mem- 
branous, ovate to obovate 3.5-5.0 mm (4.6 mm) wide, 5.0-8.0 mm (6.4 
mm) long, margins in the upper half of the blade crenate, apex acute, the 
base rounded or cuneate with a small petiole, less than 0.5 mm long; surfaces 
glabrate to strigose above, strigose below and on the margins; nervation 
distinct. Cymules 1—4-flowered, occuring along the upper 1/2 of the stems 
but somewhat congested near the branch apices, peduncles inconspicuous; 
pedicels 2.0-3.5 mm long hirtellous; bracteoles subulate shorter than the 
pedicels they subtend, ca 1.5 mm long. Calyx 4.0-5.0 mm (4.7 mm) long; 
the tube 2.5-3.2 mm (3.0 mm) long, gibbous or saccate below for ca 1/2 
of its length, 1.1-1.5 mm wide, puberulent or hirtellous with fine spreading 
hairs; the upper calyx teeth connate for ca 1/2 of their length forming an 
upper lip 1.2-1.6 mm long, the lobes deltoid to Dearne erect or slightly 
reflexed, 0.3-0.5 mm wide at the base, 0.6-1.0 mm long, hirsute-ciliate; the 
lower teeth, recurved, subulate, scarcely eo the upper lip in length, 
ca 0.4 mm wide at the base, 1.3-2.0 mm long, hirsute-ciliate,; annulus in- 
cluded, very sparse, seated approximately at the juncture of the upper and 
ie! lips, ca 0.S mm wide. Corolla 6.0-8.5 mm long, glabrous within, the 

e 5.0-6.5 mm long gradually dilating upwardly, ca 1.5 mm wide at the 
summit; the upper lip obcordate, deeply emarginate, straight and flat, ca 2.0 
mm wide, 1.0-1.5 mm long; the lower lip not spreading, 3.0-4.0 mm wide, 
20-25 1 mm long, the central lobe broadened and well-exserted from the 
laterals. Nutlets 0.8 mm wide, ca 1.0 mm One black, the surface ruminate, 
not glaucous. Chromosome number, unknow 

Distribution (Fig. 12). Andes Mts. of ee Peru and northwestern 
Bolivia. 8000-14,000 ft. Jan—Jul. 

TYPE: BoLiviA. Province of La Paz, Cordillera de Sorata, 1858, Mandon 
5.2. (Holotype, P; phototype, LA! ) 

Hedeoma mandonianum is most closely related to H. crenatum of southern 
Brazil. 


EXCLUDED NAMES 
ng arkansana Nutt., Trans. Am. Phil. Soc. N.S. 5: 186. 1837. = Satureja glabra 


ee angulata Tharp. = ee pea ies — ) Epl 

Hedeoma bicolor (Wats) Briq., Engler & Prantl, Nat. Pflanzent, ed. 1, IV, 3a, 294. 
1897. = Poliomintha eeaions Gray. 

Hedeoma bracteolata Nutt., Gen. Am. 2: Add. 1818 = ? Conradina sp. (fide Epling). 

Hedeoma caerulescens Benth. D. C. Prodr. 12: 245. Nov. 1848 = Satureja caerulescens 
Epl. 


293 
yee a Benth. D. C. Prodr. 12: 


245. Nov. 1848. = Rhododon ciliatus 
(Ben 
Hedcome coccinea (Benth.) Briq., Engler & Prantl, Nat. Pflanzenf., ed. 1, IV, 3a, 
295. 


= Rhabdocaulon coccineus (Benth.) Epl. 

Hedeoma ee (Benth.) Briq., Engler & Prantl, Nat. Pflanzenf., ed. 1, IV, 3a, 

295. 1897. = Rhabdocaulon cease (Benth.) Epl. 

Hedeoma glabra Nutt., Gen. G: 16138 ar ee glabra Fern. 

Bria. Engler & Prantl, Nat. Pflanzenf., ed. 1, IV, 3a 

ray. 

Hedeoma glabrum Pers., Syn. 2: 131. 1807. =? Satureja glabrella Brig. (fide Epl- 
ing). 

Hedeoma oe Briq., Ann. Conserv. et Jard. Bot. Geneve 2: 185. 1898. = Cunila 
galioides Benth. 

Hedeoma wee (Benth.) Brig., Engler & Prantl, Nat 
295. 1897. = Rhabdocaulon gracilis ae Epl. 

Hedeoma gracilis var. eee (Benth. 

, IV, 3a, 295. 


. Pflanzenf., ed. 1, IV, 3a, 
, Engler & Prantl, Nat. Pflanzenf., ed. 

= Rhabdocaulon eras (Benth.) Epl. 

Peds: gracillima ean Contr. West. Bot. 15: 150. 1929. = Satureja brownet 
Briq. 

Hedeoma graveolens Chapman ex Gray, Proc. Am. Acad. 8: 1873. = Stachydeoma 
Sess ns Small. 

oma oe Briq., Ann. Conser. Jard. Bot. 

ae ‘flor 

Hedeoma ile Briq., Bull. Herb. Boiss 

Benth. 


Geneve 1. 1897. = Poliomintha 


. ser. 2, 7: 611. 1907. = Cunila spicata 
Be 
Hedeoma incana Torr., Bot. U.S. Mex. Bound. Surv. 130. 1859. = Poliomintha incana 
orr.) A. Gray. 
Hedeoma italtantae awra, Oestr. Bot. Zeitschr. 31: 70. 1881 
myrtoides (St. Hill ex Benth.) Epl. 


se ie longiflora (Gary) Brig., Engler & oo Nat. Pflanzenf. ed. 1, IV, 3a 
294, 7. = Poliomintha longiflora A. 

a the ee (Schauer) Briq., Engler ie Peat Nat. Pflanzenf., ed. 1, IV, 3a 
294. = Hesperozygis marifolia (Schauer) E 

Hedeoma oe Regel, Gartenfl. 357. 


. = Hesperozygis 


864, = = Mos ocymoides Hamilt. 

Lab. Gen. et Sp., 367. 1834. = Mosla ocymoides Hamilt. 

Briq., Engler & Prantl, Nat. Pflanzenf., ed. 1, IV, 3a. 295. 

= Hesperozygis nitda ae ) 

Ses pulegioides (L.) Pers. B-mexicanum, Lab. Gen. et Sp., 366. 1834. = 
As H. pulegioides is nareported from Mexico, Bentham’s §-mexicanum undoubt- 

edly refers to another species; however the type was not located. 

Hedeoma purpurea Kellogg. Proc. Calif. Acad. 5: 52. 1873. = Satureja sp. (fide 
Eplin 


2): 
Hedeoma saris (Benth.) Brig., Engler & Prantl, Nat. Pflanzenf., ed. 1, IV, 3a. 
294. 1897 = Eriothymus rubiacens (Benth.) Schmidt. 
Hedeoma schwackeana Glaziou, Bull. Soc. Bot. Fr. 48, Mem. III, 557. 1911 = 
Hesperozygis myrtoides (St. Hill ex Benth.) Epl 
Hedeoma scutellarioides (Benth.) Briq., Engler & a Nat. Pflanzenf., ed. 1, IV 
3a. 294. 18 = Hoehnea ere ieee ee 
ade eee var. pilosa (Schmidt) Bri “Eng - & Prantl, Nat. Pflanzenf., 
IV, 4-295. 1897. = ee sntelleroides (Benth.) Epl. 
Hedeome sappuliciaes Torr., Pac. R.R. Rep 23. 1856. = Peeoayne serpylloides 
Gray. 
eee stenodonta Briq., Bull. Herb. Boiss., ser. II, 7, 612. 1907. = Rhabdocaulon 
stenondontus (Briq.) Epl. 


294 


Hedeoma subequale Epl., Bull. Torr. Bot. Club 74: 513. 1947. = Satureja (Quar- 
doquia) micromiroides Briq. 

Hedeoma taxana Cory, Rhodora a8: 405. 1936. = Rhododon ciliatus (Benth.) Epl. 

oe thymoides Pers., Syn. 2: 131. 1806. = Thymus serpyllum L. (fide Bent- 
han 


Hedevm villosa (Benth.) Briq., Engler & Prantl, Nat. Pflanzenf., ed. 1, IV, 3a. 295. 
= Rhabdocanlon lavanduloides (Benth.) Fpl 

Pens villosa var. lavanduloides (Benth.) Briq., Bo ler & Prantl, Nat. Pflanzenf., 

ed. 1, IV, 3a. 295. 1897. = Rhabdocaulon lavanduloides (Benth.) Epl. 


ACKNOWLEDGEMENTS 


I should like to give special thanks for Drs. B. L. Turner and Tod F. 
Stuessy and to Joellen Henthorn for their support and Fear esi 
throughout this undertaking. Thanks are also owed to Dr. M. C. Johnston 
for his Latin diagnoses, to the late L. H. Shinners for his bibliographic 
assistance and to the curators of the various herbaria from which specimens 
were borrowed. Based on a dissertation submitted in partial fulfillment of 
the requirements for the degree of Doctor of Philosophy at the University 
of Texas at Austin 78712, much of the initial work was conducted under an 
NSF Traineeship GB-6914 to the Graduate School of the University of Texas 
at Austin. Subsequent work was assisted by NSF Grants GB 12910 and 
08559. Mrs. Elna Widman is gratefully acknowledged for her assistance in 
the preparation of the manuscript as is the New York Botanical Garden for 
granting permission to reprint Figure 1 (in part) from Irving (1970) as the 
frontispiece. Rita Paulaski did the original drawing. 


REFERENCES 
AXELROD, D. I. 1950. Evolution of desert vegetation in western North America. 
Carnegie Inst. Wash. Publ. 590: 215-360. 
AXELROD, D. L. 1958. Evolution of the Madro-Tertiary Geoflora. Bot. Rev. 24: 
433-509. 
BENTHAM, G. 1832--1836. Labiatarum Genera et Spec London 
BENTHAM, G. sae ee In A. P. de Candolle, cea. Scan Nautralis 
Regni oo 27-603. 
- D. ate 1876. Genera Plantarum 2: 1188-1189. 


s 
5 


BRIQUET,. . 1897. Labiatae. In A. Engler and E. K. Prantl, Die Natirlichen 
Manzoni 4(3a): 183-375. 
EPLING, 939. ale on the Linnean types of American Labiatae. J. Bot., Bric. 


& aha . = 
1945-19 ee SAG opsis of the South American Labiatae. I. Repert. Sp. Nov. 
o L. '85(1): 1- 06. 1935; Il. 85(2): 97-192. 1936. III. 85 (3): 193- 388. 1936; 
. 85(4): 289-341. 1937. (cf. 85(2): 132-134. 
. 1940. Supplementary notes on American Tabigae, Bull. Torr. Bot. Club 
67: 509-534, 
., and fe S. Stewart. 1939. A revision e Hedeoma with a review of allied 
genera. Report. Spec. Nov. Regi Veg., are 15: 1-49. 
GRAY, A. sg ec al Flom of Nort ay cee 7 Part I. New rk. 
HARDIN, J. W. 1957. A revision of the American eee meres eae 9: 
145-195. 


295 


IRVING, R. S. 1970. Novelties in Hedeoma (Labiatae). Brittonia 22: 338-345. 
1976. Chromosome and numbers of Hedeoma (Labiatae) and related 

Genera. Systematic Bot. 1: 46—56. 

, and R. P. Adams. 1973. Genetic and Biosynthetic Relationships of 

Monorecsence In Recent advances in Phytochemistry, Vol. 6, Academic Press, N.Y. 

KAPADIA, Z. J. 1964. Varieties and subspecies. A suggestion towards greater uni- 
formity. Taxon 12: 257-2 

LEWIS, H., and M. E. LEWIS. 1955. The Genus Clarkia. Univ. Calif. Publ. Bot. 


eee 

ORNDUEF, R. (ed.). 1967. Index to Plant Chromosome Numbers for 1965, Regnum 
Vegetabile 50: 1-128. 

PERSOON, C. H. 1805-1809. Synopsis Plantarum, Pari 

RAVEN, P. H. hee The Systematics of Oepoibers eee Chylismia, Calif. Pub. 
Bot. 34(1): 21. 

anEAEL, J.K. 1805. Flora of the Southeastern United States. Pub. by the author, New 
Yor 

TUR RNER , B. 1956. A real study of the genus Hymenopappus (Com- 
poste). ae 58: 163-308 


TAXONOMY OF TETRAGONOTHECA 
(ASTERACEAE-HELIANTHEAE) 


B. L. TURNER and DEBORAH DAWSON 
The University of Texas, Dept. of Botany, Austin, TX 78712 


Tetragonotheca was first proposed by Linnaeus in 1753 to accommodate 
a single epappose species, IT. helianthoides, which had been collected in 
the Carolinas by Gronovius. The genus receives its name from the four 
large, valvate, outer involucral bracts which house the florets and their 
associated receptacular scales or palea. 

Gray, in 1841, described the second known species, Tetragonotheca 
ludoviciana. Because it had somewhat different, epappose, achenes he 
originally placed this in a newly erected genus, Halea. This was subsequently 
transferred to Tetragonotheca. 

Engelmann and Gray, in 1848, added a third species, Tetragonotheca 
texana. Buckley, in 1861, added the fourth species, T. repanda, although Gray 
subsequently reduced this to varietal status under T. ludoviciana. The pres- 

ent treatment accepts the specific status of T. nies and notes its relation- 

ship to be with T. texana instead of T. ludoviciana. In short, we recognize 
four allopatric species for the genus. So far as known, they do not intergrade 
or hybridize in nature. 


CHROMOSOME COUNTS 

Turner (1959) was the first to report chromosome numbers of Tetrago- 
notheca, finding m= 17 pairs in three of the species examined. Shortly 
thereafter he counted the fourth species (T. helianthoides, n = 17). Voucher 
for the plants concerned are listed in Table 1. All counts were made from 
me‘otic material at late diakinesis. At this stage bivalents show, character- 
istically, two chiasmata (Fig. 1). Subsequent onan a 1) have also 
found these several taxa to be diploid with 2 = 17 pat 


GENERIC RELATIONSHIPS 

Most workers have placed Tetragonotheca in the tribe Heliantheae. Sub- 
tribal disposition has been somewhat more controversial. Both Bentham 
(1873) and Gray (1886) include it in the subtribe Verbesininae, as di 
Hoffmann (1893). Stuessy (1977), however, placed the genus in the sub- 
tribe Helianthinae. 

Species from genera of yet other tribes have been inadvertently (or 
ignorantly) placed in Tetragonotheca (cf., excluded species), the most 
notable being Guwizotia abyssinica (Coreopsideae) and Rumfordia (Melam- 


SIDA 8(3): 296-303. 1980. 


29) 


podineae). By inference, then, some relationship to these taxa may be 
postulated. It should be noted, however, that both Gwizotia (Baagde, 1974) 
and Rumfordia (Sanders, 1978) are but dubiously positioned in these 
tribes, having strong relationships with the Heliantheae. 

It is our feeling that Tetragonotheca is relatively remote from any 
extant genus. Were it to have unspecialized, multiseriated involucral bracts, 
it would probably be positioned in the subtribe Helianthinae, near Vigwiera. 


ACKNOWLEDGEMENTS 


n preparing distribution maps we have compiled records from those 
herbaria listed below (abbreviations according to Lanjouw and Stafleu, Index 
Herbariorum, Regnum Vegetabile 31: 1964). We are grateful to the insti- 
tutes concerned for the loan of this material: the number of sheets borrowed, 
upon which the distributional maps are based, is shown in parenthesis. 


L MO (94)* 
DUKE (15)* NC (G2) 
PM <90)* x (92) * 
Pow. 20" SMU (82) 
GH (100) * TENN (41) 
LAF (4) oe (100) 
LL (50) C (29)* 
MICH (27)* - (122) * 


This paper was originally begun in the early sixties by Mr. William L. 
McCart, graduate student at the University of Texas, Austin. Annotation 
of material from the above institutions marked with an asterisk (*) was 
undertaken at that time. Subsequent annotations have been made by the 
present authors 


Table 1. Chromosome counts in Tetragonotheca. 


Species Weucher* or sete dinee ” number 
(pairs ) 
T. belianthoides FLA. Alachua Co.: Pom a: et + al (1974) 17 
FLA. Columbia Co.: Turner 4927. aby 
FLA. Escambia Co.: Turner 4919 17 
GA. Harris Co.: Jones et al. ]: 17 
MISS. Lamar Co.: Jones et al. (1966) 17 
T. ludoviciana TEX. Montgomery Co.: Turner (1959) 17 
T. repanda TEX. Atascosa Co.: Turner 4979. 17 
TEX. Jim Hogg Co.: Strother (1972). 17 
TEX. Medina Co.: Turner (1959). 17 
T texana TEX. Travis Co.: Turner (1959). 17 
TEX. Travis Co.: Urbatsch (1974). 17 


* On file, Plant Resources Center, Austin. 


xs 
8 oo 
YT 


Fig. 1. Meiotic configuration at diakinesis in Tetragonotheca helianthoides (X 


2600). 


TETRAGONOTHECA L. 


Tetragonotheca L., Sp. Pl. 2: 903. 1753. 

Bikera Adans., Fam. 2: 130. 176: 

Gonotheca Raf., im Med. Repos. N. ren 5: 352. 1808. 

Halea T. & G., Fl. N. Amer. 2: 304. 1842. 

Tetragonosperma Scheele, Linnaea 22: ee. 1849. 

Coarse, erect, perennial, caulescent herbs with strong tap roots. Leaves 
Opposite, simple, varying from nearly entire to coarsely-toothed to some- 
what pinnatifid, their sessile bases often connate-perfoliate. Heads large, 
solitary, the disk and ray flowers yellow. Involucre double, the outer of 4 
large, ovate phyllaries which are valvate in bud and remain united below 
forming a 4-angled or winged cup; the inner phyllaries membranous, as 
many as the ray flowers, and partly enclosing their achenes. Receptacles 
convex or conical, with narrow, membraneous chaff. Ray flowers pistillate, 
fertile. Disk flowers perfect, fertile. Style branches flattened, with marginal, 
stigmatic lines, the apices with an elongate hispid appendage. Achenes 


27 


quadrangular to sub-terete, truncate or rounded at the summit; pappus 
wanting or of several to numerous small, short scales. Basic chromosome 
number, x = 17 pairs. Type species: Tetragonotheca helianthoides L. 


Key to species of Tetragonotheca 


1. Pappus of 16 or more well-developed scales; plants of deep sandy soils in eastern 


S. 

2; me mostly ete ae on the main stem few, remote, and an reduced; 
T. repanda 
2. Leaves not mostly basal, those on main stem well-developed, markedly 
perfoliate, the ae usually pore peduncles 2-12(15) cm _ long; 
plants of eastern Tex 2. T. ludovictana 

. Pappus absent or a aor developed fringe of ciliate scales (rarely comprised o 
15 or fewer lacerate or fimbriate scales up to 1.0 mm long); plants of the south- 


— 


3. Mid-stem leaves (3)4-8 cm wide, not perfoliate; involucral Beas mecde 15 
mm long or more; plants of the southeastern U.S. (east of the Mississippi 
River ) 1. T. helianthoides 


1. TETRAGONOTHECA HELIANTHOIDES L. Sp. Pl. 2: 903. 1753. Type: 
VIRGINIA. Gronovins 301. Phototype (GH! ) 


Gonotheca helianthoides (L.) Raf., Med. Repos. N. York 5: 352. 1808. 


Perennial herbs from stout tap roots. Stems commonly tufted, 3-10 cm 
tall, densely pubescent with crisp, spreading hairs (rarely glabrate or villous). 
Leaves broadly ovate to elliptical, sessile, mid-stem blades 5-20 cm long, 
4-10 cm wide, acute or acuminate at the apex, remotely but sharply toothed 
or merely crenulate to nearly entire. Heads, excluding the rays, 2-4 cm 
across. Peduncles 5-8 cm long. Involucral ee broadly ovate, 2—3 cm 
long, 1-2 cm wide. Ray florets 8-13, the ligules yellow, 2-4 cm long, 
0.5-1.0 cm wide; tube 6-8 mm long, pubescent with spreading hairs. Disc 
florets greenish yellow; tube 1.5—2.0 mm long, sparsely to densely, crisp- 
spreading pubescent; throat 4-5 mm long. Achenes broadly clavate, turgid, 
subterete to somewhat 5(4)-sided, 4—6 mm long, striate, glabrous, epappose. 
Chromosome number, #7 = 17 II 

DISTRIBUTION AND ECOLOGY (Fig. 2). Mostly in sandy soils of cut-over 
pine and oak woodlands; southeastern United States, not known west of 
the Louisiana River. Flowering Apr—Aug. 

Except for its vestiture, the species is remarkably uniform considering its 
wide distribution. On both morphological and chemical grounds (Urbatsch, 
pers. comm.), T. elianthoides seems closest to T. Iudoviciana, ancestral 
populations from which it was presumably derived through regional isola- 
tion, loss of pappus and numerous other small but, in consort, significant 
diagnostic features. 


Pes * 
as aves ye oy . 
Sere sf 


ae 


> 
Cok 
ak 


Ks 


ws 
< 


% 
yr 


yf 
5% 


ie 
S 


Ley 
Wan 


a 


yee 
uae! 


> 9 
va 


relays 

reAee ay, w 
AC RS 
gave senne 7 


SIT 


Fig. 2. Distribution, by county, of Tetragonotheca helianthoides. 


301 


2. TETRAGONOTHECA LUDOVICIANA (T. & G.) Gray ex Hall, Pl. Texanae 


Halea ludoviciana T. & G., Fl. N. Amer. 2: 304. 1841. Type: LOUISIANA. “Wes- 
tern Louisiana.” Dr. Pen ae. $2. (apparently collected before Jun 6, 1837, 
according to notation on the holonpe): Holotype (N 


Perennial herbs from stout, elongate tap roots. Stems mostly single but 
sometimes tufted, especially following injury, densely white villous to glab- 
rous. Leaves broadly triangular-ovate to somewhat repand, especially below, 
the mid-stem leaves nearly always connate or perfoliate, irregularly coarsely 
dentate to crenulate. Heads, excluding the rays, 1.5—3.5 cm across. Peduncles 
mostly short, slender, 2-8(12) cm long. Involucral bracts ovate, 1.0-1.8 cm 
long, 0.8-1.2 cm wide. Ray florets mostly 13, the ligules yellow, 1.0-1.8 
cm long, 0.5-0.7 cm wide; tube 4-5 mm long, glabrous to sparsely pubescent. 
Disc florets yellow; tube ca 1.5 mm long, sparsely pubescent; throat 4—5 
mm long. Achenes 4.5-6.0 cm long, 5(6)-sided, sparsely appressed pube- 
scent; pappus a crown of 20-30 short, a entire or erose, scales, 
0.5-1.5 mm long. Chromosome number, 2 = 

DISTRIBUTION AND ECOLOGY (Fig. 3). Deep oe usually sterile, white 
soils of southeastern Texas and adjacent Louisiana. Flowering, Apr—Aug. 


3, TTETRAGONOTHECA TEXANA Gray & Engelm., Proc. Amer. Acad. 48. 1848. 
Type: TEXAS. Guadalupe Co.: Hills between the Guadalupe and Cibolo 
rivers, 12 mi W of New Braunfels, according to label data (MA), Apr 
1846, F. Lindheimer 431. (Holotype, GH!; isotypes MO!, NY!, UC!, 
US!). 


Halea texana (Gray & Engelm.) Gray, Proc. Amer. Acad. 83. 1849. 
Te ees lyratifolium Scheele, Linnaca 22: 166. 1849. Type: TEXAS. 
Guadalupe Co.: Rocky soil, Cibolo River, Apr-May 1846? Roemer s.n. (Holo- 
type, B 
Perennial herbs from tough tap roots. Stems mostly slender, solitary or 
tufted, only moderately branched above, 3-6(7) dm tall, frequently tinged 
with maroon, finely pubescent to glabrate. Leaves oblong, oval or elliptic 
in outline, pinnatifid, incised or repand, the lower-most merely connate, 
the upper connate-auriculate, 3-10 cm long, 2-4 cm wide. Heads, excluding 
the rays, 1-2 cm across. Peduncles elongate, slender, (8)10-25 cm long. 
Involucral bracts 1.0-1.5 cm long, 1.0-14 cm wide. Ray florets 8-13, the 
ligules yellow, 4-5(6) mm wide; tube ca 4 mm long. Disc florets yellow; 
tube ca 1 mm long, glabrous; throat ca 4 mm long. Ac cr obpyramidal, 
3-4 mm long, pubescent, eppapose or sometimes with 4-12 small, ciliate 
scales. Chromosome number, 2 = 17 I 

DISTRIBUTION AND ECOLOGY (Fig. 3). Calcareous, rocky soils, or some- 
times in mixed gravelly stream bottoms of south central Texas, extending 
into north central Mexico. Flowering Apr—Oct., depending on rains. 


302 


4, 


TETRAGONOTHECA REPANDA (Buckl.) Small, Fl. Southeastern US. p 


1250, 1340. 1 
Helea rep ene Buckl., Proc. Acad. Nat. Sci. Phila. 458. 1861. Type: TEXAS. 
ve orpus Christi:’’ May w/o year, eae s.n. (Holotype, PH! ). 
. N. Amer. 2: 256. 1884. 


Nueces Co.: “ne 
Halea Indoviciana var. repanda (Buckl.) Gray, Syn. F 
Perennial herbs from stout, elongate, tap roots. Stems 1-5 dm tall at 

first single but soon forming a short bushy habit through the production of 
stout laterals from the root-crown, minu soft pubescent, particularly at 
the base and below the nodes. Leaves irregularly dentate, 3-12 cm wide, 


[a 4]r. ludoviciana 
[*.]7. repanda 
[|r texana 


Fig. 3. Distribution of Tetragonotheca species. 


303 


12-24 cm long; blades of the rosette triangular with slender petioles; stem- 
leaves ovate, lanceolate to repand, narrowed into petiolar-like bases. Heads, 
excluding the rays, mostly 1.5-2.5 cm across. Peduncles stout, (5)10—40 
cm long. Involucral bracts ovate to triangular-ovate, 1.1-2.0 cm long, 1.0- 
1.8 cm wide. Ray florets 16-24, golden-yellow, the ligules 2-3 cm long, 
0.5-0.8 cm wide; tubes 5-6 mm long. Disc florets yellowish-green; tube 
glabrous to sparsely pubescent, 1-2 mm long; throat 4-5 mm long. Achenes 
4.7-5.5 mm long, turgid, 4(5) sided, sparsely hispid-pubescent, especially 
along the angles; pappus a crown of 20-30 entire or erose scales, 0.3-1.5 
mm long, those of the ray about 1/3 as long as those of the disc. Chromo- 
some number, # = 17. IL. 

DISTRIBUTION AND ECOLOGY. (Fig. 3). Deep sandy soils of southern 
Texas and possibly adjacent Mexico. Flowering Mar—Nov., depending on 
rains, 

Gray treated the species as a variety of Tetragonotheca ludoviciana but, 
as noted by Urbatsch e¢ al. (1978; unpubl.), on chemical grounds it is 
closest to T’. texana. It is also similar to the latter morphologically. In fact, 
occasional roseate or mowed plants of 7. texana may be taken for T. repanda, 
but these can be readily identified by their epappose or fimbriate scales of the 
achenes, by their smaller leaves and generally slender stems. 


Excluded Species 


Saks a abyssinica Ledeb, Ind. Sem. Hort. Dorpat Suppl. 1824. = Gwizotia 
74). 
eee parviflora Jacq., Enum. Pl. Carib. p. 28, 1760 (from description and 


Tetragonotheca guatemalensis Coulter, Bot. Gz 16: 99. 1891. = Rumfordia ee 
malensis (Coult.) Blake, Wash. Acad. Sci. 18: 25. 1928 (Cf. Sanders, 1978 


REFERENCES 
BAAG@E, J. 1974. The genus Gwizotia (Compositae). A taxonomic revision, Bot. 
39): 


BENTHAM, G. (1873). Notes on the cer aeeey see and geographical dis- 
tribution of Compositae. J. Linn. as (Bot.). 13: 335-577 

GRAY, A. 1886. Compositae. In Ed. 2, Fl. N. Aer ie 48— 480. 

HOFFMANN, O. 1893. Compositae. : Die natiirlichen Pflanzenfamilien 4: 87-387. 

POWELL, A. M., D. W. KYHOS and P. RAVEN. 1974. Chromosome numbers 
in Compositae. X. Amer. J. Bot. 61: 909—< 

SANDERS, R. 1978. Taxonomy of arse Vteraceae Brittonia 2: 302-316. 

STROTHER, J. 1972. Chromosome numbers in western North American Compositae. 

Amer. J. Bot. 59: 242-247. 

STUESSY, T. 1977. Heliantheae-systematic review. In The Biology and Chemistry 
of See II [Heywood ef al., eds.}] Academic Press, London. 

TUR , B. L. 1959. Meiotic chromosome counts for 12 species of Texas Com- 
ote ‘Busse 11: 173-17 

URBATSCH, L., N. FISHER and J. BACON (1978). Flavonoids, sesquiterpene, 
lactones, and polyacetylenes from Tetragonotheca. In Bot. Soc. Amer. Misc. Ser. 
Pbl, 156: 6. 


NEW SPECIES OF 
CLITORIA SUBGENUS BRACTEARIA 
SECTION BRACHYCALYX (LEGUMINOSAE) 
FROM COLOMBIA, VENEZUELA, AND BRAZIL’ 


PAUL R. FANTZ 


Fairchild Tropical Garden, Miami, Florida 33156 
Present address: Dept. of Horticultural Science, Box 5216, 
North Carolina State University, Raleigh, NC 27650 


Members of Clitoria L. subgenus Bractearia (Mart. ex Benth.) Fantz section 
Brachycalyx Fantz are characterized by short tubular to cup-shaped calices with 

minute lobes, inflorescences emerging from denuded nodes cere than the sane 
of the leaves, fruits which become slightly convex around the seeds and are co 
spicuously depressed between the seeds at maturity, i dae leaves, and esas 
an erect habit (trees or tall shrubs). Section Brachycalyx includes seven species 
(Fantz, 1979): Clitoria brachystegia Benth., Clitoria brachycalyx Harms, Clitoria 
glaberrima Pittier, Clitoria dendrina Pittier, one the three species newly described 
below. 


1. CLITORIA hermannii Fantz, sp. nov. 
Frutex ad 2 m altus. Folia trifoliata, foliola suborbiculata vel late ovalia, obtusa 


et abrupte minute acuminata, supra trichomatibus brevibus mollibus appressis, subter 
velutinis. Inflorescentia 132 cm longa, axillaris a nodis denudatis ante foliis 


anic 
4 mm longis. Flores resupinati, albi, 4.8-5.2 cm. Calyx brevitubularis, dense appresso- 
pubescens; tabus 11-13 mm _ longis; lobi breves, 3-4 mm longi. Vexillum extus 
fulvo-sericeum. Tubus staminalis 24-26 mm, apice trichomatibus uncinatis. Legumen 
incognitum. 


Erect shrub to 2 m tall. Branches solid, pubescence moderately dense with 
short, appressed to suberect trichomes; axillary buds 3-3.5 mm. Leaves 
3-foliate, subcoriaceous, leaflets suborbicular becoming broadly oval when 
expanded, apex obtuse, abruptly minutely acuminate, base weakly cor- 
date to rotund, midrib impressed above, primary nerves of 8-10 pairs, 
upper surface soft pubescent, trichomes short (ca 0.5 mm), appressed, 
lower surface velutinous, juvenile leaflets 4-7 cm in diameter, lamina 
expanding to 10-12 cm long by 7-9 cm wide. Petiole elongate, 6.5-15 cm, 
pubescence dense, appressed; rachis (1.5) 2—2.5 cm; petiolules quadrangular, 
tomentose, 5—8 mm. Stipules caducous (?— not observed); stipels caducous, 

‘Taxonomic studies on Clitoria accomplished at the IFAS Herbarium, Dept. of 
Botany, The University of Florida, Gainesville, Florida. Florida Agricultural Boek. 

1346. 


ment Station Journal Series No. 


SIDA 8(3): 304-311. 1980. 


305 


linear, acute, 3-nerved, 2-2.5 mm long, 0.5 mm wide, sparsely pubescent. 
Inflorescence axillary, multiflowered, at denuded nodes appearing earlier 
than the leaves, subpaniculate with primary lateral branches which bear 
the pedicels subsessile to 4 mm long; axes crowded toward branch apices, 
pubescence dense; central axis 1.5-2 cm long; pedicels 5-7 mm. Bracts 
persistent, 1.5-2 mm long, 1 mm _ wide, pubescence dense, appressed. 
Bracteoles minute, ovate to suborbicular, obtuse, 2-3 mm long, 2 mm wide, 
pubescence appressed, dense. Flowers resupinate, white, 4.8-5.2 cm. Calyx 
short-tubular, pubescence dense, appressed, tube 11-13 mm long, 5-G mr 
wide at base expanding to 8-9 mm wide at throat, lobes minute, broadly 
ovate-deltoid, 3-4 mm long, 2-2.5 mm wide, ventral lobe 4-5 mm long. 
Vexillum with indumentum on dorsal surface ae appressed, tawny, 


blade ca 3 cm wide, claw broadly cuneate, 8- ong. Alae extended 
well beyond carina by 6-9 mm, blade 19-23 mm av 6-10 mm wide, 
2-15 mm. Carina blade falcate, 10-12 mm long, 4 mm wide, claw 


21-24 mm. Staminal tube 24-26 mm long, uncinate- spube escent near apex, 
vexillary stamen coherent near middle of tube, free above and below, free 
filaments 2-5 mm long; anthers lanceolate, 2-2.5 mm long, 0.5 mm wide, 
connective apiculate. Gynophore 5 mm, with a few spreading white tri- 
chomes near apex; ovary 14-15 mm long, 1.3 mm wide, pubescence white 
slightly tinged yellowish, dense; style bearded, 16-17 mm, genicula te 7-8 
mm; stigma subcapitate, 0.5 mm in diameter. Legwme unknown. Fig 


TYPE COLLECTIONS: COLOMBIA. Vichada: shrub 18 in., ae leafless, fl. white, 
grassy llanos, along Rio ea ae José . Ocuné, 100 m, 19 Jan 1944, Hermann 
10994 (HOLOTYPE: NY. Iso GH). PARATYPES: an Vichada: 6 km 
NE of Nee on Ilanos ine Rio a ca 36 km NE of San José de Ocuné, 
100 m, 21 Jan 1944, Hermann 11019 (NY, US 1879110). 


Clitoria hermannii appears similar to C. dendrina and C, brachycalyx in 
its vegetative aspects, but the larger flowers, smaller stipels, and other 
morphological comparisons indicate closer affinities with Clitoria brachy- 
stegia of Ecuador, which is easily distinguished from C. hermannii by the 
elongated inflorescences (5—22 cm) and acuminate leaflets which are pilose 
below. All known collections of this species are from localities near San 
José de Ocuné, Colombia 


2. CLITORIA canescens Pittier ex Fantz, sp. nov. 


Clitoria canescens Pittier, nom. in sched. 
Frutex. Folia trifoliata, taliola lanceolate elliptica, acuminata, supra glabra, subter 
u entia 1.5—3.5 cm vel longi 


c race 
toris lignosis pee vel ica O super ramis primariis lateralibus ad 6-7 mm longis 
portati. Flores parva, resupinati, 13.5 cm. Calyx brevitubularis, canescens, tubus 13-15 
mm, lobi 3-4 mm. Vexillum ex ts canescens, praesertum in albastro. Twbus staminalis 

Eee 28 mm once Legumen incognitum. 


Shrub. Branches subterete, solid, internodes in weak zigzag pattern, ju- 


306 


venile branches with pubescence uncinate (vidi 30X) plus scattered, short, 
appessed trichomes (vidi 10X), becoming glabrous, axillary buds 3 mm. 
Leaves 3-foliolate, coriaceous, leaflets lanceolate-elliptic, apex acuminate, 
base broadly cuneate, midrib impressed above appearing as if it is in a 
shallow groove, primary nerves of 9-10 pairs, upper surface faintly glossy, 


PUTT : Ha AS GENS cae NRE SOE 
ae Hee SUAS 


SS GP CURRMRE 
tis ded Nitec 
194425 Sliteria co SG Suter 
Sh 


Set, a . aes is 
ye ‘ . a Sheed Us de et JaefSenrs Mla, white, 
¥Ou Gresky Lisnes, me 1 a. 
< abong the Kio Viehada 
CANCELLED“S Berenice - San sead ue ocunt 
zat SORES i TS Sys 
7 


Figure 1. Holotype of Clitoria hermannit (Hermann 10994, NY). 


307 


glabrous, lower surface dull, pubescence on nerves, inconspicuous, short, 
appressed, lamina 15-17 cm long, 6-7 cm wide. Petioles subquadrangular- 
terete, 9 cm; rachis 3.5 cm; petioles subquadrangular, rugose, glabrate, 
8-10 mm. Inflorescence terminal, racemose-nodose to subpaniculate, pedice!s 
borne on central axis at lignose knobs or occasionally on a primary lateral 
branch of 6-7 mm; axes compressed laterally, subquadrangular, twisting, 
pubescence moderately dense, appressed; central axis 1.5-3.5 cm long or 
possibly longer (apex broken); pedicels strigose, 6 mm. Bracts minute, 
ovate, acute, strigose, 1-2 mm long, 0.5-1 mm wide. Bracteoles minute, 
ovate, acute to short-acuminate, 2-3 mm long, 1-1.5 mm wide, strigose. 
Flowers resupinate, small, 3.5 cm, purple (?-in dry state the appearance 
is similar to C. dendrina which has purple flowers). Calyx dark-colored, 
with short-appressed trichomes (buds densely pubescent, canescent), tube 
short, 13-15 mm long, 4-5 mm wide at base expanding to 7 mm at the 
throat, lobes minute, broadly deltoid, short-acuminate, 3-4 mm long, 2.5-3 
mm wide at base, ventral lobe linear, 6 mm long, 0.5-0.7 mm wide. Vexil- 
lum with indumentum on dorsal surface dense, canescent, appressed, blade 
obovate, 1.5 cm wide, claw 6-7 mm long, cuneate. Alae extended beyond 
the carina by 4-5 mm, blade 17 mm long, 3-5 mm wide, claw 16 mm. 
Carina blade falcate, acute, 16 mm long, 4-5 mm wide, claw 20 mm. 
Staminal tube glabrous, 28 mm long, free filaments 3-5 mm; anthers 3 mm 
long, 0.5 mm wide. Gynophore 3 mm long, brownish-black, moderately 
densely sericeous, white trichomes; ovary 13 mm long, | mm wide, blac 
with very dense pubescence, trichomes white, sericeous; style black near 
base and sericeous, bearded above, 18 mm long, geniculate 3-4 mm; stigma 
compressed, 0.5 mm in diameter. Legwme unknown. Fig. 2 

TYPE COLLECTION: VENEZUELA. Amazonas: sitios arenoso de la boca del Gahama, 
Cano §. Miquel, ae alto Rio Negro, 26 Mar 1942, Williams 14893 (HOLO- 

TYPE: VEN 403 

The type specimen bears the name “Clitoria canescens Pittier, sp. n.” typed 
on the label and is placed in a separate type folder bearing this name. No 
record of the publication of Pittier’s name has been found. Pittier (1944) 
did not include this species in his key to the species of Clitoria in Venezuela. 

Dried flowers of this species morphologically suggest Clitoria gies 
more than any other species in the genus, although C. canescens can 
cee easily by the canescent vexillum and calyx tube pacer 
during the bud stage), longer calyx tubes, longer style, and the sparse 
i ie of the leaves. In addition, the inflorescences are less crowded with 
flowers and occasionally bear primary lateral branches, the ovary lacks any 
yellowish pigmentation, and other floral structures differ in their measure- 
ments. Clitoria canescens is known only from the type locality. 


3. CLITORIA froesii Fantz, sp. nov. 


Frutex caulibus scandentibus. Folia trifoliata, foliola lata ovata vel late ovalia vel 


308 


HERBARIO NACIONAL DE VENEZUELA 


> itoria panencans Pittior BP. Ne 
oe Ameo oe SS RS. 


Pear cf 


_ Hy iy cal 


Figure 2 


Ls 


Holotype of Clitorta canescens (Williams 14893, VEN 4038) 


= ceostece Figtha PSR A 


de Agriciturs y Cris 


309 


late ee acuminata, supra hirsuta, subter pilosa. Inflorescentia 1.5—3 cm, pauci- 
flora. Flores resupinati, parvi, 3.5-4 cm. alyx peas tubus 12-13 mm, 

i 3-4 mm. Vexillum extus sericeum. Tubus staminalis 
glaber, 24-28 mm, antherae 2.4-3 mm, connectivo —. apiculato. Legumen 
incognitum. 


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Liana. Branches 3-9 mm in diameter, solid becoming hollow in larger 
diameter branches, juvenile branches longitudinally striated, angular-terete, 
pubescence densely uncinate (vidi 30X) beneath the erect to appressed 
macrotrichomes (vidi 10X), the latter deciduous with age, mature branches 
becoming glabrate; axillary buds 5 mm. Leaves 3-foliolate, thick- 
membranaceous, conspicuously pubescent, leaflets slightly asymmetrical, 
broadly ovate to broadly oval or broadly oblong, apex acuminate, acumen 
to 1.5 cm nae base rotund, midrib and some of the basal primary nerves 
weakly raised above, pcb densely pubescent, primary nerves of 7-11 
pairs, upper surface dark green, minutely hirsute, texture similar to fine 
sandpaper, lower surface light green, pilose, nerves densely pilose-hirsute, 
lamina 10-25.5 cm long, 7-16.5 cm wide. Petioles elongate, 13-24 cm, 
subterete, trichomes dense, erect; rachis 4-6.5 cm; petiolules 5-7 mm, 
quadrangular, trichomes dense, erect, tawny. Stipules caducous, lanceloate, 
acute, 4-7 mm long, 2 mm wide on juvenile branches to 10 mm x 3 mm 
on more robust branches; stipels caducous, linear, acute, weakly 3-nerved, 
3-6 mm long, 0.5-1 mm wide. Wnflorescenee axillary and terminal, racemose- 
nodose; axes solitary, occasionally branched near base, few flowered, 1.5—3 
cm, trichomes dense, erect; pedicels 6-7 mm. Bracts minute, ovate, acute, 
2-3 mm long, 2 mm wide, densely pubescent. Bracteoles ovate, acute, 

mm long, 2 mm wide, densely pubescent. Flowers resupinate, small, 3.5 
cm. Calyx pubescence dense, appressed, bigs short, 12-13 mm long, 4-6 
mm. wide at the base expanding to 7-9 mm wide at the throat, lobes minute, 
deltoid, acute, (3) 4 mm long, 2 mm ade ventral lobe subequal, 1-1.5 
mm wide Vexillum with indumentum sericeous on the dorsal surface, ae 
2.5 cm wide, claw 4-6 mm. Alae extended beyond the carina by 4-6 m 
blade 18-21 mm long, 4-8 mm wide, claw short, 7-9 mm. Carina ae 
9-13 mm long, 4 mm wide, claw 12-16 mm. Staminal tube glabrous, 
24-28 mm long, vexillary stamen coherent below to near the middle, 
free filaments 2-4 mm long; anthers lanceolate, 2.4—-3 mm long, 0.5-0.8 mm 
wide, connective long-apiculate. Gynophore 2-3 mm; ovary 10-11 mm 
long, 1.2-1.3 mm wide, pubescence dense, white; style 14-15 mm, bearded, 
geniculate 6 mm; stigma capitate, 0.5 mm in diameter. Legume unknown. 


TYPE COLLECTIONS: BRAZIL. Amazonas: terra fir rma, Ma oe on Rio Maria, 
basin Rio Negro, a Jan — noe 12441/185 (HOLOTYPE: Y-Hb. Krukoff. 
SOTYPE: A). PARATYPE: BRAZIL. Amazonas: Pirapuca on Rio oe 1941, Froés 
12431/175 (S-Hb. at Wall). 


Clitoria froesi is the only species of section Brachycalyx with a climbing 


310 


habit, but the short calyx with minute lobes, the pubescence on the upper 
surface of the leaflets, and the lack of cauliflorous flowers suggest that this 
species should be assigned to section Brachycalyx rather than section 
Cauliflorae which is characteristically composed of species with a climbing 


Figure 3. Holotype of Clitoria froesti (Froes 12441/185, NY). 


onal 


habit. Clitoria froesi has close affinities with Clitoria dendrina which is dis- 
tinguished by the arboreal habit, smaller calyx, smaller anthers, more 
crowded inflorescences, and leaflets which are velutinous below and broadly 
ovate to rhombic-ovate to orbicular. Clitoria froesii is known only from 
western Amazonas, Brazil. 

Froes noted that the local Indians call this plant “Darume Vine” and 
plant it in fair quantities, considering it to be the most toxic plant in the 
region of the Rio Icana. Frées does not indicate for what purpose the 
toxin is used by the Indians. Other South American species of Clitoria are 
known to produce toxic seeds which are used locally as a fish poison. 

Acknowledgement is made to Helen Correll for the Latin descriptions, 
and to Daniel B. Ward, Dana Griffin II, and Donovan Correll for reviewing 
the manuscript and providing constructive criticisms. 


REFERENCES 
FANTZ, PAUL R. 1979. Taxonomic notes and new sections of Clitoria subgenus 
Bractearia (Leguminosae). Sida 8: 90-94. 
PITTIER, HENRY. 1944. Leguminosas de Venezuela I: Papilionaceae. Bol. Tech. 
Min. Agric. Caracas 5: 49. 


NOTES 


A CHROMOSOME COUNT FOR JUNIPERUS ASHEI (CUPRES- 
SACEAE) AND ADDITIONAL CHROMOSOME NUMBERS FOR 
HEDEOMA (LABIATAE)—lIn a_ study of possible hybridization in 
Juniperus, some years ago, I was able to establish a chromosome record for 
Juniperus asher Buchholtz. To my knowledge this is the first documented 
chromosome report for this species. The count was taken from root tip 
material using excised embryos grown on nutrient enriched agar. With this 
technique no pretreatment (stratification) of the seed material was necessary. 

During recent biosystematic studies of Hedeoma and allied genera (Irving 
et al., 1979) additional chromosome numbers were established for three 
previously uncounted taxa. These counts, derived from root tips, sup- 
plement Hedeoma chromosome numbers reported earlier (Irving, 1976). 
The numbers for H. montanum and H. nanum var. macrocalyx (2n = 36) 
are consistent with those of related taxa. H. multiflorum of Uruguay and 
Argentina, however, was tetraploid (22 = 72) but is closely related to H. 
drummondi (2n = 36) of Mexico and western USS. 


JUNIPERUS ASHEI oo tz (Fig. 1) 2” = 22. USA, Texas, Hays Co.: 
R. S. Irving s.n. (MONTU). 

HEDEOMA MONTANUM Brandegee 27 = 36. MEXICO, Coahuila, Sierra de 
Parras: R. S. Irving 77-7 (TEX). 

HEDEOMA NANUM (Torr.) Brig. var. MACROCALYX Stewart, 27 = 36 U.S.A, 
Ariz., Yavapi Co.: R. B. Oxford 428 (ASU). 

HEDEOMA MULTIFLORUM Benth. 2” = 72. URUGUAY, Mercedes: R. S. 


FIGURE 1. Chromosomes of Juniperus ashei. Tracing from photomicrograph. 


SIDA 8(3): 312. 1980. 


313 


Irving 7715 (TEX).—Robert S. Irving, 1422 Summit, Little Rock, AR 
72202. - 


REFERENCES 
IRVING, R. S. 1976. Chromosome numbers in Hedeoma (Labiatae) and related 
genera. Syst. oe 1: 46-56. 
oe, RENHOLTS, & D. D. IRVING. 1979. Artificial hybridization in 
Hedeoma Gablatae): Syst. Bot. 4: 1-15. 


NEW AND RE-INSTITUTED COMBINATIONS IN GUTIERREZIA 
(COMPOSITAE: ASTEREAE)—In connection with monographic work 
on what is commonly known as the Xanthocephalum complex, and treat- 
ment of these genera for the Flora of the Chihuahuan Desert Region (M. C. 
Johnston, and collaborators, in preparation), the combinations listed below 
are necessary. Justification of such dispositions and full synonymy will be 
provided in a forthcoming doctoral dissertation on the group. 

There is strong morphological and chromosomal evidence supporting the 
close alliance of the species of the former genus Greenella with those of 
Gutierrezia, reflected here by the transfer of Greenella arizonica and G. 
amulosa into Gutierrezia. The third species of Greenella, G. discoidea Gray 
(Proc. Amer. Acad. Arts 19: 2. 1883), which is known only by the type 
material, has been found by this author to be a rayless form of Nanthocepha 
lum wrightii (Gray) Gray. Since I have here returned this latter species to 
Gutierrezia also, the genus Greenella as a whole is reduced to synonymy. 
The only objection that Gray himself had to this placement was ray-floret 
color, and since there are several South American gutierrezias with white 
rays, maintenance of a distinct genus on that basis is meaningless. 

As a result of these transfers and the changes of status of two of the taxa 
involved, there are now 14 North American species of Gutierrezia. Gymno- 
sperma glutinosum is closely related to this group and may be transferred 
to Gutierrezia in the future; this taxonomic decision must await additional 
study. 

1. GUTIERREZIA conoidea (Hemsley) Lane, comb. nov. Based o 
Xanthocephalum conoideum Hemsley, Biol. Centr. Amer. 2: 109-112. 1882. 

2, GUTIERREZIA ALAMANI Gray, Smithsonian Contr. Knowl. 3(5): 91 
(Pl. Wright. 1: 91). 1850. This species has been known most recently as 
Xanthocephalum linearifolium (IDC) Greenman (Publ. Field Mus. Nat. 
Hist. Bot. series 2: 345. 1912), based on Keerlia linearifolia DC (Prod. 5: 
309-310. 1836). However, when placed in Gutierrezia, it must take Gray's 
epithet because of the pre-existence of G. linearifolia Lag. (Gen. et Sp. Nov. 
30. 1816) 

3. GUTIERREZIA ALAMANI Gray var. megalocephala (Fernald) Lane, 
comb. & stat. nov. Based on: Xanthocephalum megalocephalum Fernald, 


SIDA 8(3); 313. 1980, 


314 


Proc. Amer. Acad. Arts 36: 505. 1901. 

4. GUTIERREZIA sericocarpa (Gray) Lane, comb. nov. Based on: 
Xanthocephalum sericocarpum Gray, Proc. Amer. Acad. Arts 15: 31-32. 
1880. 


5. GUTIERREZIA WRIGHTI Gray, Smithsonian Contr. Knowl. 5(6): 78 
(Pl. Wright. 2: 78). 1883. Gray himself later transferred this species to 
Nanthocephalum (Proc. Amer. Acad. Arts 8: 632. 1873), but my work has 
convinced me that the original placement is more correct. 

6. GUTIERREZIA arizonica (Gray) Lane, comb. nov. Based on: Greenella 
arizonica Gray, Proc. Amer. Acad. Arts 16: 81-82. 1880. 

7. GUTIERREZIA ramulosa (Greene) Lane, comb. nov. Based on: Green- 
ella ramulosa Greene, Pittonia 1: 302-303. 1887. 

8. GUTIERREZIA SPHAEROCEPHALA Gray, Mem. Amer. Acad. Arts ns. 
4(1): 73-74 (Pl. Fendl. 73-74). 1849. This species was transferred by 
Shinners to Xanthocephalum and considered by Solbrig to be a synonym of 
Gutierrezia glutinosa, however, in my treatment it will receive specific status 
in Gutierrezid. 

9, GUTIERREZIA TEXANA (DC) T. & G. var. glutinosa (S. Schauer ) 
Lane, comb. & stat. nov. Based on: Hemiachyris glutinosa S. Schauer, 
Linnaea 19: 724. 1847.—Meredith A. Lane, Department of Botany, Univer- 
sity of Texas, Austin, TX 78712. 


HESPERALOE FUNIFERA (AGAVACEAE) IN TEXAS.—Hesperaloe 
funifera (Koch) Trelease was collected in the course of field work for a 
Texas Natural Area Survey of the Devil’s River—Dolen Falls area in Val 
Verde Co. (Smith, J. M. and M. Butterwick. 1975. A Vegetational Survey 
of the Devil's River—Dolan Creek Area. In A Natural Area Survey Part VI 
of VUI. Division of Natural Resources and Environment, The University 
of Texas, Austin, Texas. Available at the Texas State Library ). This species 
is known from northern Mexico from eastern and central Coahuila to Nuevo 
Leon (one station) and San Luis Potosi (several stations) (R. Engard, 
Desert Botanical Garden, Phoenix, pers. comm.). A small Texas population 
was observed in shallow limestone soils of rocky upper slopes, ca. 5 km 
ENE of the Finnegan Ranchhouse (29° 56’ N, 100° 55° W). An additional 
individual has been transplanted at the entrance of the Finnegan ranch. 
Hesperaloe funifera with its large (to 5 cm wide and 2 m long), rigid leaves 
and purplish-green, glaucous flowers is easily distinguished from H. parvi- 
flora (Torr.) Coult. which has linear, arcuately spreading leaves (2.5 cm 
wide and 1.2 m long), and rosey-red to salmon colored flowers. Collection 
data: In rocky limestone soil at the entrance to the Finnegan Ranch (29° 
56’ N, 100° 58’ W), ca 65 km NW of Del Rio, Val Verde Co., 2 Aug 


SIDA 8(3): 314. 1980. 


gD 


1979, Deal sn. (TEX-LL)—Mary Butterwick, Bureau of Land Manage- 
ment, Phoenix, AZ 85017, and Jackie M. Poole, Department of Botany, Uni- 
versity of Texas, Austin, TX 78712, 


CUCURBITA DIGITATA (CUCURBITACEAE) IN TEXAS.—A speci- 
men of Cucurbita, collected during field studies for a Texas Natural Area 
Survey of Fresno Creek, proved to be a new state record. Cucurbita 
digitata Gray was previously known from southern New Mexico, Arizona, 
and California with Mexican localities in Sonora and Chihuahua. 
Cucurbita digitata has palmately 5-parted leaves which are narrowly and 
deeply lobed, a character that easily distinguishes it from the other Texas 
species. Collection data: Infrequent in gravelly alluvium along the banks of 
Fresno Creek, ca 4.0 km N of the Smith House of the Big Bend Ranch 
(29° 23’ 30” N, 103° 51’ ’0” W/), Presidio Co., 30 Sep 1975, Butterwick 
1726 (TEX-LL) —Mary Butterwick, Bureau of Land Management, Phoenix, 
AZ S017; 


PHYLLANTHUS POLYGONOIDES (EUPHORBIACEAE) NEW TO 
ARKANSAS.—On June 21, 1979, Phyllanthus polygonoides Nutt. was dis- 
covered 414 miles northeast of Eureka Springs, Carroll County, Arkansas. 
The suffruticose plants are characterized by ascending whiplike branches 
clustered on a caudex, obovate leaves with acute or mucronulate tips, and 
fruiting calyx lobes 1.5-2.5 mm long. Hundreds of the plants were scattered 
in glades surrounded by oak-juniper woodlands and in an old overgrown 
roadbed. They were growing in exposed shallow, gravelly soil underlain by 
limestone. 

Webster (A revision of Phyllanthus (Euphorbiaceae) in the Continental 
United States. Brittonia 22: 44-76, 1970) listed the range of P. polygonides 
as “Oklahoma and Texas south to the Mexican plateau, with outliers in 
New Mexico, Missouri, and Louisiana.” The discovery of this station in 
northwest Arkansas suggests that the Missouri plants (known only from 
Stone County in southwest Missouri) are not disjunct but rather are at 
the northeast periphery of the species range which is likely continuous from 
Oklahoma to southwest Missouri. Dr. Edwin Smith, University of Arkansas, 
Fayetteville (pers. comm.) indicated to me that the flora of northwest 
Arkansas has been less than adequately worked and habitats similar to that 
described here are common in the region. Specimens (Brooks 14254) are on 
deposit at KANU, MO, NY and UARK.—Ralph E. Brooks, University of 
Kansas Herbarium and State Biological Survey of Kansas, 2045 Avenue A, 
Campus West, Lawrence, KS 66044. 


SIDA 8(3): 315. 1980. 


SIDA CONTRIBUTIONS 
TO BOTANY 
VOLUMES = ——SNUMBER 4 DECEMBER 1980 


CONTENTS 


A new Bahamian Euphorbia and some new combinations. 


Donovan S. Correll. aly 
Cyperus difformis L. (Cyperaceae) in North America. 

Barney L. Lipscomb. 320 
The distribution of Habranthus tubispathus (L’Her.) Traub in 

South America and North America—Texas and Louisiana. 

W. C. Holmes and Christopher J. Wells. 328 

“> The migration and establishment of Juncus gerardii (Juncaceae ) 

in the interior of North America. Ronald L. Stuckey. 334 

Notes on the flora of the Chinati Mountains, Presidio County, Texas. 
348 


Emily J. Lott and Mary L. Butterwick. 


NOTES. Distribution records for Digitaria bicornis in eastern United States. 352.— 
Najas marina: new to the Indiana flora. 353—Gaura odorata Lag. (Onagraccae ) 
in Louisiana. 354—A note on flower color of Wahlenbergia linarioides (Lam.) A. 
DC. in northwest Florida. 355—Brachyeletrum erectum and Talinum rugospermum, 
new species to Texas and notes on Schoenolirion wrightit. 355—Holosteum um- 
bellatum (Caryophyllaceae) in Ontario. 356. 


INDEX TO VOLUME 8. Barney L. Lipscomb and Jessica Procter. 359 


US ISSN 0036-1488 


SIDA, CONTRIBUTIONS TO BOTANY 
Founded by Lloyd H. Shinners, 1962 


Editor & Publisher 


Dallas, Texas, 75275 


Associate Editor Assistant Editor 
John W. Thieret Barney L. Lipscomb 
Northern Kentucky University SMU Herbarium 
Highland Heights, Kentucky, 41076 Dallas, Texas, 75275 


Guidelines for contributors are available upon request beginning with 
volume 8. 


Subscription: $6.00 (U.S.) per year; numbers issued twice a year. 


Sida, Contributions to Botany, Volume 8, Number 4, pages 317-372. 
Copyright 1980 
by Wm. F. Mahler 


A NEW BAHAMIAN EUPHORBIA AND 
SOME NEW COMBINATIONS 
DONOVAN S. CORRELL 
Fairchild Tropical Garden, Miami, Florida 33156 


With the project on the Flora of the Bahamian Archipelago nearing com- 
pletion we find a few loose ends that still need attention. This is one of them. 


EUPHORBIA abbreviata Correll, sp. nov. 
Frutex crassus brevis —— ad 70 cm altus, aliquancum cupreus, dense et grosse 
osus, ramis glabris, typice cum textura dura et rigida. Nodi annui instructi vaginis 
aes sdawialive isles Cilia primaria 2-3 mm longa. Folia opposita 


saepe contiguae, sine appendiculis. Capsula glabra, circa 2 mm _ longa, era crasso 
circa 1 mm longo; semina lactea, levia in uno extremo acuta, circa 1.3 mm _ longa, 
ecarunculata angulis rotundatis. 


Stout short bushy shrub to about 70 cm tall, somewhat copper-colored, 
densely and coarsely branched with the branches typically hard-tissued and 
rigid, glabrous. Nodes annual, provided with stipular sheaths that are ciliolate 
adaxially; primary internodes 2-3 mm long. Leaves opposite and solitary, 
subsessile, fleshy, entire, glabrous, often clustered at the nodes, globose- 
orbicular or shortly oblong-rectangular or narrowly cuneate-obovate, 1-3 mm 
long. Cyathia solitary in the axils, very shortly pedunculate, about 1 mm long; 
glands rather large and prominent for the cyathium, blood-red or blackish, 
often contiguous, without appendages. Capsule glabrous, about 2 mm long, 
supported by a stout pedicel about 1 mm long; seeds white, pointed at one 
end, smooth, about 1.3 mm long, with rounded angles, ecarunculate. 


YPE COLLECTION: Bahama Islands, Little Inagua, on sandy thinly coppiced slope 
about saline lake in southwest sector of island, plants 0.7 m tall, coppery-colored, 
August 12-16, 1975, D. S. Correll 45998 (holotype, NY; isotypes, F, FTG, GH, JJ 
K, MO 

Additional collecti tions: Caicos Islands, Long Cay, northern end, March 7, 1911 
F. & C.M. Millspaugh 9237 (NY, US); Long Cay, July 28, 1930, Paul Bartsch 
s. w. (US). 


Grisebach, in his Flora of British West Indies, p. 52. 1864, described 
Euphorbia vaginulata from Turks Islands, and it is to this taxon that the 
presently described species has previously been referred. In my explorations 
I have collected the rather widespread E. vaginulata on Grand Turks Island, 
throughout Great Inagua, in the Caicos Islands (Pine Cay, Providenciales ), 


SIDA 8(4): 317-319. 1980. 


318 


Fig. 1, a, branchlet of Exphorbia vaginulata; b, branchlet of Exphorbia abbreviata. 
Drawn by Priscilla Fawcett. 


319 


on Crooked Island and at its extreme northern limit on San Salvador Island. 
It is a rather slender plant with flexible, “rubbery” branches that produce 
elongate internodes and linear-oblong leaves. 

The presently described species is local in its occurrence. It has a short, 
stout trunk from which arise rigid thick branches that have much-abbreviated 
internodes and suborbicular-quadrate to shortly elliptic leaves. 

The following transfers are necessary for these plants to be coordinated in 
our Flora. They mainly represent a difference in taxonomic interpretation. 


EUPHORBIA exumensis . Isp.) Correll, comb. nov. 


aa syce exumensis Millsp., Field Mus. Bot. 2: 301. 1909. 
. lechioides var. poets (Millsp.) Burch, Ann. Missouri Bot. Gard. 53: 92, 
9. 1966 


EUPHORBIA proctorii (Burch) Correll, comb. nov. 
Chamaesyce proctorii Burch, Ann. Missouri Bot. Gard. 63: 378. 1976. 


EUPHORBIA wilsonii (Millsp.) Correll, comb. nov. 


Chamaesyce wilsonit Millsp., Field Mus. Bot. 2: 301. 
C. lechioides var. wilsonii (Millsp.) Burch, Ann. iVissourh Bot. Gard. 53: 91, 99. 
1966. 


CYPERUS DIFFORMIS L. (CYPERACEAE) 
IN NORTH AMERICA 
BARNEY L. LIPSCOMB 


Herbarium, Southern Methodist University 
Dallas, TX 75275 


Cyperus difformis is a widespread species of southern Europe and the 
tropics of the eastern hemisphere (Kiikenthal, 1936; Koyama, 1961; Ohwi, 
1965). It is spreading and becoming established in North America like 
many other oriental introductions, e.g., C. haspan, C. iria, and C. rotundus. 
In North America, it was first noted in México, then in the United States, 
and now can be reported in Nicaragua, Central America. 

Early American collections and literature on the species date back to the 
1850's. In his monograph on North American Cyperaceae, Torrey (1836) 
did not list or mention C. difformis. Later Torrey (1859) reported a collec- 
tion (1851-1852) as C. lateriflorus from México. Britton (1886) noted the 
presence of C. J/ateriflorus in North America as being nearly allied to C. 
difformis of the old world and may be the same. Howell (1934) reported 
C. difformis in the United States from California; Fernald (1935), from 
Virginia. Numerous other collections have expanded the known range into 
Alabama, Arizona, Louisiana, North Carolina, and Oklahoma. The known 
distribution of C. difformis is given in Figure 1. 

Cyperus difformis is native to the tropics of the eastern hemisphere spe- 
cifically tropical and western Asia. It is a common weed in a number of 
habitats but most commonly found in rice paddies. The annual plant, with 
culms 15-60 cm tall, is similar in habit to C. haspan but can be distin- 
guished from it by its subglobose spikes and obovate scales (Fig. 2). Correll 
& Correll (1972) has a description of the species. C. difformis was placed in 
section Fusci by Kiikenthal (1936) and there it is most closely related to 
C. fuscus of Europe and western Asia which also has been introduced into 
North America. 


HISTORY AND LITERATURE 


The earliest known American collection of C. difformis is that of Charles 
Wright from Sonora, México, in 1851. Wright's specimen 1950 was des- 
cribed as a new species, C. lateriflorus, by Torrey (1859). Ayers (1946) 
did not mention the Wright specimen in his treatment of the genus in 
México citing only those previously cited by McGivney (1938) who cites 
Wright “1960” from “New Mexico.” Wright actually collected the species in 
Sonora, and McGivney’s citation of “1960” was apparently a misprint or was 
miscopied because the label clearly shows 1950. 


SIDA 8(4): 320-327. 1980, 


400 MILES. 


600 KILOMETERS 


400 


8 
8 
8 


2 


1 = Distnto Federal 


MCE x We 


NA flacotecas” 


Fig. 1. Distribution of Cyperus difformis in North America. Long curved arrows 
indicate probable points of introduction and short straight arrows possible migration 
of the species. 


Wright’s collecting was done in association with the U.S. Mexican Boun- 
dary Commission in 1851. He accompanied Colonel Graham of the Topo- 
graphical Engineers between 2 Sep-4 Oct on a trip from Santa Rita, New 
Mexico, to Santa Cruz, Sonora. During the latter part of this trip (29 Sep), 
Wright collected the Cyperus just east of Santa Cruz. Wright's (1949-52) 


Fig. 2 
tis) 


a 


" Ie 


ut Ky a 
: FAN wi 
ae ats 


N 
NP 

\ : 

NM ~~ 

Mh ae hh | 
iH Ne Db ae 

cy \ Ye Vy ig ; 

: d if 


rity 


in ¥ sh iyi 
ANS Ae Bi 
WN: a oe 4 i) 

feats wa a fy. i oy 

Sn 


Pad, 


. Cyperus difformis L. A. habit. B. achene, C & D. scale (from Delahoussaye 


523 


field notes give the following data: along mountain torrents east of Santa 
Cruz, 29 Sep 1851, Wright 674. Wright's field number of 674 is the same 
s “Wright 1950” assigned by Gray for distribution. 

Wright's collections of 1849-1852 from southwestern United States cre- 
ate a problem for botanist who try to work with his plants. Gray, who 
received and processed the specimens, sent them out under a new set of 
numbers and not under Wright's own field numbers. Wooton (1906) noted 
that the specimens of the 1851 and 1852 collections were under the same 
printed label and this made it difficult, if not impossible, to fix localities 
and exact dates of collections. Most of the printed labels do not show the 
habitat or exact locality and date as given in Wright's field notes (Wooton, 
1906). 

Since Wright’s collection, additional collections of Cyperus difformis in 
1890 and 1909 were reported by McGivney (1938) in southern Mexico. It 
was 25 years later that the species was first collected in the United States. 
Howell (1934, 1949, 1958) first reported it from California followed by 
Fernald (1935) from Virginia. It was then reported from Arizona by Kear- 
ney and Peebles (1951). Additional reports from the United States are 
Thieret (1964) from Louisiana, Correll and Correll (1972) from Oklahoma, 
Tyndall et al. (1977) from North Carolina, and Godfrey and Wooton (1979) 
from Alabama. 


INTRODUCTION AND DISTRIBUTION 


The data on introduction and distribution of Cyperus difformis follows a 
chronological order based on label data taken from specimens examined. 
The data of the earliest collections, as well as others, does not necessarily 
indicate the time of introduction, establishment or the direction of migration. 

Cyperus difformis was first collected on the North American continent in 
northern México. However, this first collection does not establish this locality 
and date as the time or place C. difformis was introduced. The earliest refer- 
ence to C. difformis in North America was Torrey (1859). A search of 
earlier literature [Bentham, (1839-1857); Humboldt et al. (1815); and 
Kunth (1822)} yielded no mention of C. difformis. However, since the 
plant was probably present (maybe even several years) before collection, it 
would be reasonable to assume that the approximate time of introduction 
was sometime between 1815-1851. The first collection was reported by 
Kiikenthal (1936) who cites Wright 1950, Santa Cruz, New Mexico. Mc- 
Givney (1938) also cited it from New Mexico and it has since been errone- 
ously reported from New Mexico in various treatments, e.g. Correll & Cor- 
rell (1972), Godfrey & Wooton (1979), Kearney & Peebles (1951, 1960), 
and Tidestrom & Kittell (1941). 

Additional specimens examined: MEXICO: Guanajuato: 1 mi NW of Salamanca, 
Waterfall & Wallis 13901 (SMU). Jalisco: near La Barca, Pringle 3261 (ENCB). 
Michoacan: valley of the Lerma, Pringle 3261 (MO); vicinity of Morelia, Arsene 


324 


3173 (MO); vicinity of Morelia, Arsene 3092 (MO). 


Based on herbarium specimens from México and the United States C. 

difformis appeared in the United States flora some 65 years later than 

ico. It was first collected in California in 1915: Butte Co.: very abundant 

in canals, ditches, roads, and fields, Jul 1915, Johnston W132] (NY). Howell 

(1934) published the first United States report of C. difformis based on a 

1921 collection from California: Butte Co.: weed abundant in rice fields, 
Nelson, 25 Nov 1921, McLearn 15 (CAS) 


Additional specimens examined: CALIFORIA: Alameda Co.: 16th Street Statin, 
Oakland, 8 Jul 1935, Jussel sn. (CAS). Butte Co.: rice fields, 9-10 Aug 1921, 
Kennedy s.n. (UC); Sanford slough, SW corner of Butte Co., Mason & Grant 12636 

UC); Gray Hodge Game Refuge NW of Pennington, Mason 12737 (CAS, UC); 
Sacramento Outing Club, Mason & Grant 13024 (DS, UC); rice plots, Biggs Rice 
Experiment Station, Tucker 2680 (CAS, SMU, UC); eastern outskirts of Oroville, 
Howell 34637 (NY). Colusa Co.: Colusa rice fields, 18 Sep 1925, ye SM. 
(UC); rice fields, Dunshee 14 (UC); rice fields, 3 mi W of Williams, Howell 
19838 (CAS, DS, MO, NY, SMU, UC); 8 mi N of Colusa, Mason & Grant 12983 
(UC); 2 mi W of Colusa on Williams ee Mason & Grant 13011 (UC); 1.6 mi 
W of Williams, Mason & Grant 13116 (UC); rice field, 1 i 
Nobs & Smith 1877 (UC). Fresno Co.: es River E of Burrel, Mason & Smith 
8341 (DC, UC). Glen Co.: edge of pond S of Willows Migratory Wildfowl Refuge, 
Beetle 3061 (UC). Kern Co.: Tupan hwy 1/4 mi S of Adohr Rd., Twisselmann 
6580 (CAS, NY); Lake Barak near Delano, Twisselmann 8957 
Madera Co.: Crocker-Hoffman Ranch near Merced, Wagnon 4a (UC); San Joaquin 
River just E of Firebaugh, Si aie 9131 (CAS). Marin Co.: Mt. Tamalpais, 
camp above Marin Woods, Howell 20771 (CAS). Merced Co.: 0.5 mi S . Lake 
Yosemite, 27 Jul 1938, Wagnon sv. (UC); Crane Ranch S of jct of M 
an Joaquin rivers, eee & Smith 8328 (UC); Los Banos Bird ne Jussel 90 


Ve 
io) 
nae 
5 
5 
* 


(CAS); San Joaquin River 0.25 mi S of hwy 152 2.9 mi E of Santa Rita Park, 
Wiggins & Wiggins 20701 (DS, ENCB, NY). Nevada Co.: Wolf oo 1 mi N of 
Nevada-Placer Co. line, Nobs & Smith 1534 (UC). Plumas Co.: near Virgilia, 


Howell 35064 (NY). pene Co.: American River at Fair Oaks, Crampton 
1713 (CAS). San Diego Co.: Balboa Park, Gander 8893 (DS, Oy: San Francisco 
Co.: Lake Merced, 7 Aug 1953, Bush s.n. (CAS). Santa Barbara Co.: Las Positas Rd. 
near Fox Canyon, 12 Oct 1962, Pollard s.n. (CAS, ENCB); Las Positas Rd., near jet 
of Veronica Springs = 3 Nov 1962, Pollard sn. (MO, NY, SMU); 3 Nov 1963, 
Pollard s.n. (CAS). Sonoma Co.: Guerneville, Rabtzoff 2919 (CAS). Stanislaus Co.: 
4 mi S of Oakdale, Hoover 57 (CAS); Claus, Hoover 1628 (UC). Sutter Co.: 6.3 
mi E ot Bue Landing, 6 Jul 1938, Cantelow sm. (CAS); Sutter by-pass on 

Yuba Knight’s Landing hwy, Mason 12705 (NY, UC); 1.7 mi S of Robbins, 
7 Bee (NY). Tehama Co.: Pacific hwy 0.5 mi N of Butte Co. line, Heller 
14804 (MO, NY, UC). Tulare Co.: Pixley Nat. Wildlife Refuge, Tip ebaone 
11590 (CAS). 


KNOWN RECORDS FROM OTHER STATES 
VIRGINIA. The collection of C. dfformis from Virginia in 1934 (Fernald, 
1935) is apparently the only one known from the state. This probably repre- 


sents a separate and direct introduction from Asia than from the west coast. 
Its most probable source of introduction is in rice-straw used in packing, 


325 


as suggested by Fernald. 

ARIZONA. Cyperus difformis was first reported as likely to occur in 
Arizona by Kearney & Peebles (1942). “Not known definitely to occur in 
Arizona but has been collected at Santa Cruz, Sonora, about 20 miles south 
of the international boundary.” The following specimens are known from 
Arizona. Maricopa Co.: swamp area below dam of Lake Pleasant Regional 
Park, 8 Oct 1964, Lehto 5461 (NCU). Mohave Co.: Goose Lake of Havasu 
Lake Refuge near Toprock, 27 Oct 1948, Monson s.n. (CAS). These collec- 
tions probably represent introductions from México or California. 

LOUISIANA. Thieret’s report of C. difformis in Louisiana is based on 
the following collection: Lafayette Parish: rice field 13 mi W of Lafayette, 
23 Oct 1963, Delahoussaye 71 (SMU). Through correspondence, Thieret 
indicated he had later collected the species in Lafayette Parish but at a 
different locality: Lafayette Parish: rice fields, 2 mi NW of Indian Bayou, 
10 Sep 1977, Thieret 50236 (KNK). These collections are probably inde- 
pendent introductions from Asia. 

HOMA. The Oklahoma report was made by Correll and Correll 
(1972): Leflore Co.: gravel bars along spillway of Lake Wister, 17 Sep 1970, 
Correll & Correll 39765 (TEX). One additional specimen can be reported 
from Johnston Co.: moist soil along Blue River, 24 Sep 1977, Puckett 36 
(SMU). This specimen was a gift from Dr. Mickey Cooper of Cameron Uni- 
versity and was collected on a class field trip through southeastern Oklahoma. 
The source and origin of introduction of these are questionable. It is possible 
that they have originated from the populations in Louisiana. 

ALABAMA. Limestone Co.: bottoms of Tennessee River of Wheeler Na- 
tional Wildlife Refuge N of Dekatur, 22 Sep 1970, Kral 41221 (ENCB, 
NCU, SMU). C. difformis has only recently been reported from the state by 
Godfrey & Wooten (1979). 

NORTH CAROLINA. Dare Co.: ca 1.5 km N of Duck, along the shore- 
line of Currituck Sound, with Scirpus americanus, 2 Oct 1978 Tyndall 791 
(ODU, SMU). It was first collected 14 Oct 1974 at the same location, 
Tyndall sn. (ODU) and has only recently been reported from North Caro- 
lina by Tnydall et al. (1977). 


DISTRIBUTION IN CENTRAL AMERICA 


Early studies—e.g., Clarke (1908), Hemsley (1882-1886), and Standley 
(1930)—do not list C. difformis from Central America. Hemsely’s report 
of the species is that of C. Jateriflorus from Sonora. 

The only collection examined was collected in 1969 in Nicaragua. Mata- 
galpa: beside pond, Rte 1, S of Dario, Calabazas, 25 Dec 1969, Atwood 
2566 (SMU). This has recently been reported from Nicaragua by Seymour 
(1980) as C. haspan with duplicates at BM, ENAG, F, GH, NY, WDP. The 
nearest known localities are reported in this paper and are about 1000- 
1100 miles to the northwest in Michoacan. It is unknown whether or not 


326 


C. difformis has become established in Nicaragua. The specimen was mature 
when collected and had already dropped “seed,” thus probably establishing 
the original source or increasing the existing population. 


SUMMARY AND DISCUSSION 


Cyperus difformis was originally described by Linnaeus in 1756. The 
habitat noted by Linnaeus was in India. It is suspected and generally accepted 
that, by means of man, especially with the introduction of rice, C. difformis 
has spread to the new world tropics and subtropics of Central and North 
America, Since the introduction of it into the United States, probably around 
1910-1915, it has become established through the southeast and southwest 
from Virginia to California. Since being introduced in California it has 
spread over much of the state as a weed and has become a potential problem 
in certain rice-growing regions. 


ACKNOWLEDGEMENTS 


Thank you John Thieret for taking time to read and review this paper. 
Your comments and suggested changes are much appreciated. I thank the 
herbaria, CAS (24), DS (6), ENCB (4), KNK (1), MO (6), NCU (2), 
NY (12), SMU (9), and UC (24), who kindly loaned specimens. A total 
of 88 sheets were borrowed. The numbers in parenthesis are numbers of 
sheets borrowed. 


REFERENCES 
cae B.B. 1946. The genus Cyperus in México. Catholic Univ. Amer., Biol. Stud. 


BRITTON, N.L. 1886. A preliminary list of North ae er of Cyperus, 
with descriptions of new forms. Torrey Bot. Club. Bull. 13: —216. 

CLARKE, C. B. 1908. The Cyperaceae of Costa Rica. a ts "Natl. Herb. 10: 
443-471. 

CORRELL, D.S. & H.B. CORRELL. 1972. Aquatic a wetland plants of pig 
western United States. Environmental Protection Agency. Washington, D.C. 

FERNALD, ae 1935. Midsummer vascular plants of oe Virginia. —. 
dora 37: 378-4 

ee REY, R. K & J. W. WOOTEN. 1979. Aquatic and wetland plants of south- 

astern United States. The University of Georgia Press, Athens. 272. 

HEMSLEY, W. B. 1882-1886. Biologia Centrali-Americana. Vol. 3: 449. Porter and 
Dulau & Co., Lon 

HOWELL, J. T: 1934, Tepes difformis L. in California. Leafl. West. Bot. 1(10) : 
104. 


_ . 1949, ee Flora. Univ. of California Press, Berkeley. 88. 
, P. H. VEN, & P. RUBTZOFF. 1958. A flora of ‘San Francisco, Cali- 
focaia: The ae - San Francisco, California. 47. 


KEARNEY, T. H. & R. H. PEEBLES. 1942. eee plants and ferns of Arizona. 
S.D.A. _ Publ. 423. Washington, D.C. 15 

1951. Arizona flora. Univ. of California Press, Berkeley. 149. 
. Arizona flora (2nd ed, with suppl. by T. J. Howell 

& x E =F es ene Tey. ar California Press, Berkeley. 149. 


aed. 


KOYAMA, T. 1966. Classification of the family Cyperaceae (3). Quart. J. Taiwan 
Mus. 14: 159-194. 

KUKENTHAL, G. 1936. In Engler, Das Pflanzenreich. 4(20): 1-315. 

MC GIVNEY, SISTER M.V.D.P. 1938. A revision of the eae Eucyperus found 
in the United States. Catholic Univ. Amer., Biol. Ser. No. 26. 

OHWI, J. 1965. Flora of Japan (in Faglich): Smithsonian Institution, Washington, 
D.C. 


OLE 
eee F. C. 1980. A check list of the vascular plants of Nicaragua. Phytologia 
p. 66. 


Mem 

Ge P. C. 1930. The Cyperaceae of Central America. Publ. Field Mus. Nat. 
Hist., Bot. Ser. 8(4) : 237-292. 

THIERET, J. W. 1964. More additions to the Louisiana flora, Sida 1(5): 

TIDESTROM, I. & S. T. KITTELL. 1941. A flora of Arizona and — Mexico. 
Catholic Univ. of America Press, Washington, D.C. 766. 

TORREY, J. 1836. ae of North American Cyperaceae. Ann. Lyceum Nat. 
Hist. a Aa oe 

= 859. In cot on ac United States and Mexican Boundary Survey. 2(1): 

TYNDALL, R. W., J. W. USHER, & G. F. LEVY. 1977. Cyperus difformis L. a 
North Carolina state as and additions to the flora of Dare County. J. Elisha 
Mitchell Sci. Soc. 93: 

WOOTEN, E. O. oe Souther localities visited by Charles Wright. Bull. Torrey 
Bot. Club. 33: 561- 

WRIGHT, C. 1849- ede Wright's field notes and collection numbers. L. H. 
Shinners’ copy at 


THE DISTRIBUTION OF HABRANTHUS 
TUBISPATHUS (L’Her.) TRAUB IN SOUTH 
AMERICA AND NORTH AMERICA— 
TEXAS AND LOUISIANA 
W. C. HOLMES 
Biology Department, Northwestern State University, Natchitoches, LA 71457 
and 
Institute for Botanical Exploration, Mississippi State, MS 39762 
and 
CHRISTOPHER J. WELLS 
Biology Department, Mississippi State University, Mississippi, MS 39762 
and 


Institute for Botanical Exploration 


ABSTRACT 


The plant known as Habranthus texanus (Herb.) Steud. is considered synonymous 
with H. tubispathus (L'Her.) Traub. Its distribution includes warm temperate South 


seventeenth or early eighteenth centuries possibly by Spanish missionaries. 


The plant known as Habranthus texanus (Herb.) Steud. is cited by Sealy 
(1937) as the only species of the genus to occur in the wild outside of 
South America. Despite being considered endemic to Texas by Correll and 
Johnston (1970), this plant has been reported in Louisiana as early as 1934 
by Dormon. It has a long and confusing nomenclatural history. This appears 
to be caused, in part, by the plants of the United States never having re- 
ceived clear distinction from certain warm temperate South American ex- 
pressions of the genus. This paper is intended to clarify the nomenclature 
and distribution of this interesting plant and provide information on its 
ecology and occurrence in the United States. 

The plant, known locally as the “Copper Lily,” is a scapose perennial 
arising from a brown ovoid bulb of about 1-2 cm in diameter. The apex 
of the bulb is abruptly narrowed into a tubular-like sheath composed of 
the remains of the leaves. The bulb bears 2-6 linear, deep green grass-like 
leaves that are 3-5 mm wide and up to 20-25 cm in length. Leaves normally 
wither prior to flowering. The scape is 10-20 cm tall and bears one flower. 
The bulb may flower more than once per year, occasionally possessing two 
scapes, each with a flower or fruit in various stages of development. Borne 


SIDA 8(4): 328-333. 1980. 


a2) 


slightly beneath the flower is a tubular scarous bract ending with a bifid, 
long attenuated tip. The pedicel is about twice the length of the bract. The 
flower is 2-3 cm long, slightly zygomorphic, and inclined. Tepals are obovate 
with rounded apices ending in an apiculate claw-like tip, this more pro- 
nounced on the outer whorl. The perianth is bright yellow, with the outer 
surfaces tinged with burgundy, the color deepening toward the base. The 
six stamens are slightly declinate and in fascicles of four different lengths 
and apparently attached to the base of the perianth. At the point of attach- 
ment are scarous, lacinate enations. The style is about two-thirds the length 
of the corolla (about the length of the longest stamen) and ends in a trifid 
stigma. The inferior ovary is somewhat tubular, three-loculate, and possesses 
very narrow, sub-herbaceous wings that extended onto the pedicel. At ma- 
turity the fruit is ovoid, three-lobed, and contains numerous black wafer-like 
seeds about 3—5 mm in diameter and 0.55 mm thick. 

Field observations and data from herbarium specimens indicate the Copper 
Lily proliferates in dryer, well-drained, but disturbed areas, of low vegetation. 
Maximum flowering occurs from July through October following periods 
of heavy rainfall, At such times it may exhibit complete aspect dominance. 
Sporadic, but very light flowering may occur at other times during this 
period. Flory (1939), in reviewing Paces (1913) embryological work on 
this plant, pointed out that this was apparently the first known incidence of 
diploid parthenogenesis, while at the same time having a sexually produced 
endosperm. Crane (1978) refers to this condition as semigamy. This appears 
to be the common condition in the Zephyrantheae tribe of the Amaryllidaceae 
(“rain lillies’). Ic gives a special advantage in allowing self pollination 
without genetic penalty in this group whose erratic and unpredictable flower- 
ing may lower the chance of insect pollination (Charles Crane, pers. comm. ) 

The synonymy of this species reflects not only its uncertain specific status 
but also the close affinities that exist between Habranthus and Zephyranthes. 

he plant of the United States was first designated Zephyranthes texana by 
Herbert. Upon proposing the genus Habranthus, Herbert himself gave it 
varietal status as H. andersonianus var. texanus. Steudel reinstated the plant 
to specific status as H. texanus. Later, Green created the combination, 
Atamasco texana. Direct comparison of the plants of the United States 
(known as Habranthus texanus) with those of South America (part of the 
H. andersonii complex) showed no differences that warrant retention of 
specific status for H. texanus. In fact, no differences could be detected and 
correlated with geography to give a definite geographical race or varlety 
worth recognizing. Our opinion is that the name H. texanus rests solely on 
its disjunct distribution rather than an actual morphological distinction. 
This point of view is shared by Sealy (1937), Alexander (1939), and 
Ravenna (1970). Ravenna also showed that the type of H. andersonii (of 
which H. texanus is here considered synonymous) is identical to the type 
of H. tubispathus (’Her.) Traub, the basionym being Amaryllis tubispatha. 


— 


330 


Therefore, the correct name for the only Habranthus outside of cultivation 
in the United States is H. inca pies synonymy is as follows: 


Zephyranthes andersonii (Herb.) Steud., Nom. Bot. - - A 70. 1840. 


m : 9, 
Zephyranthes andersonii (Herb.) Penis & Hooker a on Pl. Ae 724, 1883. 
Zephyranthes andersonii (Herb.) Baker, Handbook Amaryll. 37. 1888. 

Atamasco texana Green, Pittonia 3: 187. 

Atamasco Wie G. M., Rev. Fac. Let. & Cisne. 19: 227. 1914. 


The distribution of the species includes southern Brasil, Uruguay, Argen- 
tina,, central and southern Chile, and the United States (east Texas and 
northwest Louisiana). The species has, under various of the synonyms cited 
above, been reported in much of the Antilles and northern South America 
(see Pulle, 1966; Grisebach, 1864; Adams, 1972; Moscoso, 1943; and Urban, 

1903). However, the plant of those areas has a white flower and is referable 
to Zephyranthes puertoricensis Traub. 

Herbaria abbreviations follow that of Index Herbariorum (ed. 6). except 
for Institute for Botanical Exploration (IBE), Mississippi State, Mississippi. 


Specimens examined: NORTH AMERICA: United States: Texas. Anderson Co.: 


Wilcox, near Palestine 1 Aug 1943, Barkley s.n. (TE astrop Co.: Texas Hwy. 71 
on the first ll E of Garfield, 5 Oct 1966, Mears 1017 (TEX); Bexar Co.: San 
Antonio, Oct Thurber (F); Brazos Co ollege Station, 7 Sep 1969, 


n 
ee 1108 (SMU): Caldwell Co.: 4.65 mi N of Lockhart on Texas Hwy. 183, 
1966, Mears 680 (TEX); Calhoun Co.: Port Lavaca, Gentry 46 (F); Colo- 

oes ee Eagle Lake, Oct 1930, Biology Class sn. (TEX): Comal Co.: New 
Braunfels, Oct 1850, Lindheimer 1207 (F, SMU); DeWitt Co.: without further 
location, 18 Jul 1942, Riedel s.n. (TEX); Fayette Co.: Muldoon, 20 Jul 1950, 
Ripple 51-713A (TEX); Jim Wells Co.: 15.2 mi S of Alice, 13 Sep 1955, John- 
ston 2781A (SMU); Gonzales Co.: 4.5 mi S of Belmont, 27 Sep 1958, Correll 
20466 (LL); Karnes Co.: Karnes City, 29 Aug 1953, Johnson 1302 (SMU); Mc- 
Lennan Co.: M-K-T RR N of Gapshead, Jul 1946, Smith 56 (TEX); Milam Co.: 
U.S. Hwy. 190, ca 10 mi NE of Cameron, 26 Oct 1963, epee 63-1833 (SMU, 
TEX); Nacogdoches Co.: E. Austin St., Nacogdoches, 7 Jul 1955, Lacey 32 (SMU); 
San Patrico Co.: ca : mi NW of Sinton, 24 Oct 1948, ae 6643 (TEX); Travis 
0.: mi S of Victoria, Cory sae eel Waller Co.: Hempstead, 10 Jun 
Ble Hall 654 (F); “Willia amson Co.: 1 SW of Georgetown, 2 Oct 1944, Wolcott 
7 (TEX); Walker Co.: Near eRe Jun 1914, Young s.n. (TEX); Louisiana. 
oo. Parish: Columbia Park, Shreveport, 31 Jul 1977, MacRoberts 262 (LSUS); 


(IBE, NATC); Sabine Parish: ca 3/4 mi W of Natchitoches-Sabine Parish line 
on La. — 6, 18 Sep 1978, Holmes 3344 (IBE, NATC). SOUTH AMERICA. a 
tina. Buenos Aires. La Casada, Troucoso 1296 (F):; Brasil: Minas Gerais. Serra d 

Se ee 11 Nov 1938, “Bopreto 8809 (F); Chile: Cordillera de Chile, 1(90, 


Bene 


Hanenke 2122 (F); Paraguay: Cordillera de Altor, Mar 1903, Frebrig 1017 (F); 
Uru Montevideo. Sayago, 3 Jan 1930, Herter 85498 (F) and 85494 (F); 
Rahal pa: Dec 1925, Herter 79892 (F) and 79891 (F) 


Since the plant formerly known as Habranthus texanus is considered 
synonymous with H. twbispathus, it is doubtful it is native to both southern 
South America and Texas and Louisiana. Several points make it probable 
that the plant is native to southern South America and its disjunct distribu- 
tion of well over 10,000 kilometers is better explained in another way. 
These include the following: 


. Most other species of Habranthus (ca 23, Ravenna, 1970) are native to Sout 
Fev which undoubtedly is the center of distribution for the genus. Two ae 
nae, since 1937, been described fro 

Although plants pospelgiclly identical to the plants of the United States 
exist, considerably more variation is present in the South American populations 
this species. The variation is well documented by Ravenna (1970) who proposed 
eee infraspecific names based on flower size and minor color differences. If in- 
troduced into the United States, it would almost certainly have come from one 
population and not exhibit the total range of variation present in the species 
(Founders Principle). eta coupled with its a habit would explain 
and insure a great amount of morphological uniform eee of herbarium 
specimens, field ae and cultivation support thi is argumen 

. The plants of the United States inhabit disturbed places of ee usually grassy 
wevetuion (lawns, roadsides, railroads, etc.). 

he plant is very abundant in west-central Louisiana only in the areas settled 
or developed at the same time Texas was being settled by the Spanish. These in- 
clude the City of Natchitoches, founded in 171 14, and the El Camino Real (now 
roughly following Louisi ana hway 6) a trail Baa in the early 1700's con- 


abundant at Los aes, nee easternmost Spanish mission Pande in L/L, which is 
15 miles west of Natchitoches on 1 Camino Real, miles from the 

state line. It is abundant on y 6, westward from Los Adaes for eo 
five miles. Th t has not been oe om any other place in Louisiana, 


of this plant in these, the oldest inhabitated areas of Louisiana, and with | 
es that existed with Spanish Texas, (also being settled at the same time) apparently 
indicates that it may have been introduced into nai in the same mann 


same time as it appears it was introduced in Texa 


In considering the possible ways in which Habranthus tubispathus could 
be introduced into the United States, natural methods can hls be dis- 
counted because the pl ant does 1 not appear to possess either | ong or moderate 


through the digestive tract seems doubtful. 


332 


The introduction of Habranthus tubispathus appears best explained by 
human activity. This was first proposed by Sealy (1937) at the suggestion 
of H. H. Hume of the University of Florida. He proposed that Spanish 
missionaries transported the plant between 1690 and 1703 during the estab- 
lishment of the various missions in Texas. It should be noted, as pointed 
out by Flory (1938), that the only records available concerning the estab- 
lishment of Spanish missions in Texas cite that they were founded by 
Spaniards coming out of Mexico and H. texanus (H. tubispathus) has not 
been reported in that country. This would certainly preclude chance intro- 
duction from Mexico, but not intentional introduction if the plant were 
cultivated there and could not become naturalized due to unfavorable con- 
ditions. The plant's slow rate of spread favors intentional human transport. 
Documented in Natchitoches some 50 years ago, and probably being present 
there for a much greater period of time, the plant has failed to spread out- 
side of the city, as cited above. When its distribution area in the United 
States of well over 125,000 square miles is considered, it is obvious that 
either the plant arrived here much earlier than the Europeans (which we 
feel the evidence does not support), or was a direct, intentional introduc- 
tion as an ornamental or possibly for some other use not known. today. 
In no other way could the plant have spread to occupy its present distribu- 
tion area. We suggest ic was brought into Texas from South America, pos- 
sibly via Mexico. Once established at one site, it was carried to other settle- 
ments. Under favorable conditions it escaped, became established, which 
further hastened its spread. These we feel are the only conditions that 
explain the plants’ present distribution in the United States. It appears to 
be only a matter of time before the plant spreads to other favorable habitats 
in Louisiana and possibly the southeast United States. 

ACKNOWLEDGEMENTS 

We wish to thank the curators of the hebaria for their loan of specimens 
and Sidney McDaniel of the Institute for Botanical Exploration for his 
review and comments on the manuscript. Appreciation is also extended to 
W. S. Flory of Wake Forest University and Charles Crane of the University 
of Missouri, Columbia, for their suggestions and comments. 


REFERENCES 

ADAMS, C. D. 1972. Flowering plants of Jamaica. Univ. West Indies Press, p. 
8-79 

ALEXANDER, E. J. 1939. Habranthus andersonii. Native of Texas, Argentina, 
Uruguay, and Chile. Addisonia 22: 45-46. 

CORRELL, D. C. and M. C. JOHNSTON. 1970. Manual of the vascular plants of 
Texas, nae Research Foundation, Renner, Texas. p. 418-419. 

CRANE, C. F. Apomixis anc crossing mi eue a in some Zephyrantheae. Un- 
pubeoned oa Dissertation, University of Texas, Austin, 1978. 
RMON, C. 1934. Wildflower of eee Doudleday, Doran, & Co. Garden 
City, New York. p. 


jos 


222 


FLORY, W. S. 1938. Cytotaxonomic notes on the genus Habranthus. Herbertia 5: 
51-153 


GRISEBACH, A. H. R. ies Flora of the British West Indian Islands, Lovell Reeve 
and Co., London p. 584 

MOSCOSO, R. M. 1943. Calon Floraes Domingensis. p. 83. 

PACE, L. 1913. Apogamy in Atamasco. Bot. ees 56: 376-396. 

PULLE, A. 1966. Flora of Surinam, Vol. I, Pt. 1. E. S. Brill, Leiden, p. 444-445 

REVENNA, P. F. 1970. Contibutions to the ee American oe ia 
Plant Life 26: 73-103. 

SEALY, J. R. 1937. se Pyrolirion, Habranthus, and Hippeastrum, Journ. 

Royal Hort. Soc. 63: 195- 
URBAN, I. 1903. Symb. fae 4(1): 


THE MIGRATION AND ESTABLISHMENT OF 
JUNCUS GERARDII (JUNCACEAE) IN THE 
INTERIOR OF NORTH AMERICA 


RONALD L. STUCKEY 


Department of Botany, College of Biological Sciences, 
The Ohio State University, Columbus 43210 


Juncus gerardii Loisel (Juncaceae), a member of the rush family occurs 
along the Atlantic and Pacific coasts of North America and locally inland 
on the continent. On the Atlantic coast, the plant is often a dominant mem- 
ber of the upper littoral zone where it may form extensive stands in. salt 
marsh meadows at the upper limit of ordinary tides (Nichols, 1920). Ap- 
parently once confined to the coastal salt marshes, J. gerardii has evidently 
migrated inland where it has been reported from ten midwestern and Great 
Lakes states and the provinces of Manitoba and Ontario in the past 120 
years. More recently it has invaded Colorado and Utah in western United 
States. This paper presents the documentation for the migration and estab- 
lishment of J. gerardii as isolated occurrences in the interior of North 
America. The progressive range extension over a period of approximately 
120 years and the present known North American distribution of J. gerardii 
based on herbarium data and published accounts are mapped in figure 1. 

Although a true rush, J. gerardii is commonly called black grass. Other 
common names that have been applied to the species are given in Table 1. 
In earlier North American scientific literature, the species was reported 
under the name J. bulbosus L. 


EAST COAST DISTRIBUTION AND EARLY OCCURRENCES IN THE INTERIOR 


In eastern North America, the apparent native distribution of J. gerardii 
is in the Atlantic coastal salt marshes. Its range is usually stated as extend- 
ing from Newfoundland, Quebec, Nova Scotia, New Brunswick, the coastal 
states of the United States to Virginia and sometimes to Florida. I have 
verified herbarium specimens of its coastal range from Newfoundland to 
Virginia. However, the southern limit of its Atlantic coastal range is not 
clear from the literature. In his classic revision of the North American 
species of Juncus, Engelmann (1866) reported the southern limit of J. 
gerardit as North Carolina and Florida and cited as authorities, Curtis and 
Ware, respectively. This range to Florida has been repeated by Fassett 
(1940), Fernald (1950), Muenscher (1944), Small (1933), Wiegand 
(1900), and mapped by Hultén (1958). However, its range is given as 
far south as Virginia in Gleason (1952), Gleason and Cronquist (1963), 


SIDA 8(4): 334-347. 1980. 


ope) 


TABLE 1. 


COMMON NAMES OF Juncus gerardii LOISEL OTHER THAN BLACK GRASS 


REGION OF THE 
COMMON NAME UNITED STATES WHERE USED REFERENCES 
Black-grass Rush Florida Ward (1968) 
Gerard’s Rush New York; Hanmer (1940); 

Ohio Schaffner (1914, 1932) 
Hog-rush Southeastern United States Small (1933) 
Nut grass Maryland McAtee (1933) 
Salt-marsh Rush New England Seymour (1969a) 
Saltmeadow Rush British Columbia Taylor and MacBryde 

(1977) 


and only as far south as Delaware by Potter (1932). It is not reported for 
the Carolinas by Radford, Ahles, and Bell (1968) or from North Carolina 
by Beal (1977), but is listed for Florida by Ward (1968). According to 
Eleuterius (1976), J. gerardii is replaced by J. roemerianus beginning in 
New Jersey and extending southward in the east coastal salt marshes to 
Florida and on the Gulf Coastal Plain to Texas. 

What is believed to be the earliest record of occurrence of J. gerardit 
inland in the United States appears in the North American Botany by 
Eaton and Wright (1840), where they reported the species for Michigan. 
No specimen has been located to verify this record. A second inland occur- 
rence in the Great Lakes region was reported as “near Chicago .. . 
Vasey,’ by Engelmann (1866). This second record was apparently a 
on specimens obtained in 1862 in vacant lots at Chicago, Illinois, by George 
Vasey (GH, US). Vasey’s specimens substantiate the statement in the 5th 
edition of Gray’s Manual that J. gerardii was “rare along the Great Lakes” 
(Gray, 1867). The previous edition of the Manwal noted that the species was 
“common along the coast from New Jersey northward” with no mention of 
locations in the interior on the continent (Gray, 1863). Gray’s statement 
in the Sth edition of the Manual served as the basis for the record cited in 
three early catalogues of the flora of Michigan (Wheeler and Smith, 1881, 
Beal and Wheeler, 1892; Beal, 1905), rather than the citation of the pee 
in Eaton and Wright (1840). 

The beginning of the inland spread of J. gerardii is documented in New 
Jersey and Pennsylvania where the species was first reported as “Not fre- 
quent” from a specimen obtained in 1866 on ballast ground at Petty’s Isl land 


Be 


near Philadelphia in the Delaware River (Smith, 1867). Herbarium 2a 
mens (MO, PENN, PH, US) from ballast habitats in Camden and Phila- 
delphia date from 1863 to 1876 and document this early introduction. The 
species has spread northward into Leigh, Montgomery, and Northampton 
Counties, Pennsylvania, where it its known from alluvial flats and wet 
meadows along railroads, as documented by specimens dating from 1924 
into the 1960’s (PENN, PH). Schaeffer (1949) reported that the popula- 
tions in Northampton County “probably represent plants which were intro- 
duced by salt hay.” 


ESTABLISHMENT IN THE MIDWESTERN AND GREAT LAKES STATES 


Expans‘on of the range of J. gerardii in New York evidently came in 
the mid-to late-nineteenth century. Torrey (1824) gave its distribution 
in that state as “common in the vicinity of New-York,” and later Torrey 
(1843) wrote “Borders of creeks and ditches in salt marshes: common on 
Long Island and Staten Island.” Gray (1863) in the 4th edition of his 
Manual noted its distribution as “Salt marshes; common along the coast 
from New Jersey northward.” However, in the 5th edition, Gray (1867) 
first noted it as “also in saline marshes of W. New York.” This apparent 
range extension was probably based on the specimens obtained in 186 
and 1865 at Salina, Onondago County, by George W. Clinton (MO, PH) 
which also was the basis for the citation in Engelmann’s revision (1866). 
In a catalogue of plants of that a Paine (1865) noted J. gerardi 
from “Rivuletsides and wet banks, east Salina and west of Onondaga 
Lake, in thick patches. Common there.” ees years later, the species was 
listed as “sparingly, on the brackish marshes E. of Montezuma” in the 
Cayuga Lake Basin in western New York (Dudley, 1886). In central New 
York, additional locations are known from Fall Creek, Myers Point, the 
Union Springs (Wiegand and Eames, 1926), at Silver Springs near Gaines- 
ville in Wyoming County, and at Greigsville near York in Livingston 
County (Zenkert, 1934). At the Silver Springs locality, J. gerardii was noted 
on 30 August 1926 as common in a number of places along Wolf Creek 
where salt and brine from the Remington Sale Factory was being dumped 
into the stream (Muenscher, 1927a, b). Additional localities in central and 


Figure 1. Distribution of Juncus gerardit in North America, with emphasis on its 
spread into the interior of the continent. Large symbols show locations of cited speci- 
men records and periods of ae they were obtained, according to the following key: 


Triangles in circles (1860-1 884), squares in circles (1885-1919), dots in circles 
(1920-1949), stars in ioe Ae 1979). Small symbols indicate the general range 
of the species along the coast. Dots show the apparent native range along the east 


coast; squares show the ns on the west coast. In addition to herbarium records, the 
east coast range is derived from maps by ee (1960), Hultén (1958), Potter 
(1932), Roland (1945), and Rousseau (1974 


338 


western New York include Hamburg, Ithaca, Lansing, McLean, Piffard, 
Retsof, Springport, and Syracuse, as derived from the list of cited specimens. 
The species has apparently spread in the man-created saline habitats of 
central and western New York, although some writers have doubted its 
being non-indigenous to the area (Wiegand and Eames, 1926). A century 
later, J. gerardii continues to persist in Onondago County, the same county 
where it was initially discovered in the interior of the state of New York 
(Faust, 1961). In the eastern part of the state, the species has been located 
on rocks in the bed of Kinderhook Creek at Stuyuesant Falls in Columbia 
County (McVaugh, 1958) and in moist depressions on dredgings from the 
Hudson River at Rensselaer (BH, CU) indicating that it had moved inland 
along the Hudson River in the 1930's. 

In Ohio, J. gerardii has always been considered native, including the most 
recent treatment by Braun (1967), although she allowed that the species 
was “pethaps introduced along railroad tracks.” First recorded by Beardslee 
(1874) as “rare” from “Lake [Erie] Shore,” this record was repeated by 
Kellerman and Werner (1893). The earliest known specimens are those 
obtained in 1891 and 1895 at Cleveland by Edo Claassen (OS), which 
served as the basis for the record in his Cuyahoga County list of plants 
(Claassen, 1895). These Cuyahoga County specimens also served as the 
basis for Schaffner’s only record in his two catalogs of Ohio vascular plants 
(Schaffner, 1914, 1932). A second record was taken 34 years later at Phalanx 
in Trumbull ene by Almon N. Rood (OS). The plants were “abundant 
in low, wet swale,” but were erroneously reported as J. greenei (Schaffner, 
1933), and later coueeiel (Schaffner, 1935). Additional specimens obtained 
by Rood in 1936 were labeled as “Phlanax Station, wet swale by R.R., 
supporting the hypothesis that the species had probably invaded via thé 
railway. More recent records have come from along a railroad at Crestline 
in Crawford County by Floyd Bartley (OS) and from a ditch at the Inter- 
national Salt Company at the mouth of the Cuyahoga River in Cuyahoga 
County by Allison Cusick (KE). 

The only known ane of J. gerardii for Michigan are those obtained 
by C. K. Dodge in 1911 and 1915 at Port Huron in St. Clair County and 

y O. A. Farwell in 1931 and 1932 at Wayne in Wayne County (Voss, 
1972). The 1911 specimens came from railroad yards. The 1931 plants 
were from wet ditches, and as reported by Farwell (1941) were then be- 
lieved to be the first record for Michigan. Voss (1972) has stated that 
this species is “quite probably not indigenous in Michigan.” 

In Indiana, Peattie (1930) first reported J. gerardii as very rare from 
brackish wet sand in Indiana Harbor, Lake County, but attempts to locate 
a specimen from this locality have been unsuccessful. In 1935, however, the 
species was first found to be well established on dry open ground along the 
Nickle Plate Railroad four miles east of Kokomo, Howard County by C. M, 
Ek (Hermann, 1938, 1940). In 1942, the species was obtained farther 


52) 


north in the next county, Miami County, along the Pennsylvania Railroad, 
three miles west of Bunker Hill by C. M. Ek (GH, MO) 

Vassey’s record of J. gerardii from near Chicago has long been disputed. 
Jones and Fuller (1955) excluded it from the flora of Illinois, because the 
species was “not attributed to Illinois, or indeed even mentioned, in any of 
Vasey’s published reports on the flora of Illinois, . . .” Their conclusion was 
accepted until more recent studies have verified its occurrence in the Chicago 
area, based on specimens collected in 1900 and 1956 (DeFilipps, 1964, 
1966; Mohlenbrock, 1970; Swink, 1969, 1974). 


KNOWN RECORDS FROM OTHER STATES 


VERMONT: First reported “on the margin of a marshy meadow very near 
the railway station at New Haven Junction: where it was believed to have 
been imported in “saltmarsh hay often used in packing” (Eggleston, 1904). 
Not reported earlier in the flora of Vermont (Brainerd, Jones and Eggleston, 
1900), nor in the most recent flora of Vermont (Seymour, 1969b). 
WISCONSIN: Reported from near railroad yards in Milwaukee and She- 
boygan Counties (McIntosh, 1950), based on specimens obtained in the 
1930's, and in Racine County as early as 1900 (Swink, 1969, 1974). The 
most recent specimen, taken in 1977, is from a wet area along railroad tracks 
in Fox Point, Milwaukee County (OSH). 
MINNESOTA: Stated by Hultén (1958) as a weed along railways in 
Minnesota, with a mapped location in the southern part of the state evi- 
dently based on the record from Martin County obtained in 1950. The 
species is also cited for Clay and Kittson Counties (McGregor and Barkley, 
1977) based on records from the 1960's. The specimen from Clay County 
is J. alpinus (shoulder of roadside ditch, Muskoda, 8 Jul 1960, O. A. Stevens 
2231, DAO, US), and the specimen from Kittson County is J. compressus 
(roadside at Caribou, 28 Jun 1962, J. W. Moore 26083, TRT). 
NORTH DAKOTA: Listed for Cass and Richland Counties (Stevens, 1961; 
McGregor and Barkley, 1977). The record from the latter county is dated 
C 
MISSOURI: Observed during the years 1964 through 1966 in the southern 
part of the Ranken yard of the Terminal Railroad Association, east of 
Compton Avenue, in St. Louis (Muehlenbach, 1969). 
KENTUCKY: Obtained in 1978. This specimen is the most recently known 
record and is from a population of plants in diesel sludge mud of railroad 
yard near tracks, near junction of West 19th Street and Augusta Street, Cov- 
ington, Kenton County (OS). 


Selected Specimens: ILLINOIS: [Cook Co.}: Vacant lots, Chicago, 1862, Vasey 5.n. 
GH, US); railroad and roadside, Chicago, 1956, Glassman 3788 (CHI, ILLS, NP). 
INDIANA: Howard Co.: Railroad, Kokomo, 1935, E& s.n. (GH, IND), 1936, Deam 
57045 (NY). Miami Ca Railroad, Bunker Hill, 1942, Ek s.n. (GH, MO). KEN- 
TUCKY: Kenton Co.: Railroad yard, Covington, 1978, Applegarth 465 (OS). MICHI- 


340 


GAN: [St. Clair Co.]: Railroad freight yards, Tunnel Station, 1911, Dodge s.n., 
(MICH); damp ground, Port Huron, 1915, Dodge s.n. (US). [Wayne Co.}: Diche: 
Wayne 1931, Farwell sn. (GH), Farwell 8897 (MICH). MINNESOTA: Martin Co.: 
Railroad, oe 1950, Moore 20689 (GH, MIN). Mrssour!: St. Louis Co.: Rail- 
road, St. Louis, 1964, Muehlenbach 2356 (MO, US): 1965, Muehlenbach 2505, 2522, 
2476 (MO); 1966, Muehlenbach 2625 (MO). NEW JERSEY: [Camden Co.} : Camden, 
1863, Canby s.n. (MO): ballast. Petty’s Island in Delaware River, 1866, Burk s.n. (MO), 
Diffenbach s.n. (MO, PH), Parker s.n. (GH, MO, PH); ballast, Kaighns Point, 
1876, Martindale sn. (PH, US). NEW YorK: Cayuga Co.: Brackish meadow east 
of Salt Creek, Montezuma, 1885, Dudley sn. (CU); brackish place by railroad  sta- 
tion, Union Springs, Springport, 1919, Eames & Wiegand 11730 (CU, GH); salty 
meadows, Montezuma, 1919, Wiegand, Randolph, & Eames 11729 (CU). Columbia 
Co.: Bank of creck, Stuyvesant Falls, 1933, McVaugh 997 (NY, PENN); rocks in 
Kinder Creek, Stuyvesant Falls, 1945, McVaugh 3126 (CU). Erie Co.: Meadow, 
Hamburg, 1939, Glowenke 3868 (PENN). Livingston Co.: Ditches along roadside, 
salt factory, Piffard, 1926, Muenscher 16495 (CU, GH); stream bank, salt factory, 
Retsof, 1926, Muenscher 16495 (CU). [Onondaga Co.}: Salina, 1864, Clinton s.n. 
(MO), 1865 (PH); salt marshes, Syracuse, 1880, paar 2973 axe salt marshes, 
Onondaga Lake, 1883, Habertr 954 (CAN, US), (MO). Rensselaer Co.: Moist 
depressions in pees Hudson River, Sees 1939, House 27070 (BH, CU). 
Tompkins Co.: Railroad, Myers, Lansing, 1918, Eames 9559 (CU, GH); railroad 
ditch, McLean, 1920, Wiegand & Muenscher 13448 (CU, GH). Wyoming Co.: 
Salt Sek Silver Cee 1926, Muenscher & Burkholder 16498 (CU, GH). NoRTH 
a KOTA: Cass Co.: Railroad, Fargo, 1954, Stevens 1497 (DAO, NY, US). Richland 
ane soil, Colfax, ie Ball 694 (US). OHIO: Crawford Co.: Railroad, Crest- 
me 1960, Bartley s.n. (NY, OS). Cuyahoga Co.: Cleveland, 1891, 1895, Claassen 
sm. (OS); ditch, Tatecnioaal Salt Company, mouth of Cuyahoga River, Cleveland, 
oe Cusick 13723 (KS). Trumbull Co.: Wet swale, Phalanx Station, ee Rood 
. (OS); wet swale, railroad, Phalanx Station, 1936, Rood 1077 (KS, NY, P N). 
eee LVANIA: Lackawanna Co.: Roadside ditch, Carbondale, 1946, ira 6812 
(GH, PENN). Leigh Co.: Raileond. Laury’s Station, 1924, Pretz 12155 (PH); 
meadow, Allentown, 1960, Schaeffer 6095 (PH). a es Co.: Railroad, Hat- 
boro, 1943, oe 59871 (PH). Northampton Co.: adow, Butztown, 1946, 
Schaeffer 22738 (PH); railroad, Northampton, 1947, oe 25947 (PH); shore, 
Siegried, 1961, haeffer 63226 (PH). {Philadelphia Co.]: — ground below 
the navy yard, Phil piskg 1866, Parker Ly Aas PH). VERMONT: [Addison Co.}: 
Railroad, New Haven Junction, 1897, B era sn. (GH). ae NSIN: [Racine 
Co.}: Lake beach, i habor, 1900, Wadmond 2740 (CU, MIN, WIS). Milwaukee 
Co.: Railroad Estabrook Park, Shorewood, 1939, Shinners 1069 (PENN, WIS); 
railroad, Fox Point, 1977, Harriman 14125 (OSH). Sheboygan Co.: Railroad yard, 
Sheboygan, 1933, Goess! s.n. (WIS). 


o 


DISTRIBUTION IN THE INTERIOR OF CANADA 


In Canada, J. gerardii was early reported by Macoun (1888) from only 
coastal marshes in Nova Scotia, New Brunswick, Quebec, and Vancouver 
Island. In the interior of Canada, in Ontario, J. gerardii was not included 
among the introduced plants of that province (Montgomery, 1956). How- 
ever, the species has been known from the province as early as 1903, based 
on specimens obtained by William Scott (CAN, CU, DAO, TRT) from a 
railroad roundhouse at Niagra Falls. J. gerardii has been located in Ontario 
at other isolated sites along railways, usually at stations were salt has been 


341 


sprinkled to melt snow in the winter. The most recent records seen are 
from 1974, 1975, and 1976, (CAN, DAO, TRT). In the Toronto region, 
Catling and McKay (1975) have described the plant communities at two 
saline sites, one a salt storage depot on the west side of the Don River 
near the Bloor Street viaduct and the other a snow dumping site on land- 
fill at the foot of Leslie Street, where J. gerardii is plentiful with other 
halophytes. Boivin (1952) reported the species as a weed along railway 
embankments at Cochrane et Hope i in Ontario and at Brandon in Manitoba. 


Selected Specimens: MANITOBA: Weed along railway embankments, Brandon (cited 
by Boivin, 1952). ONTARIO: Carleton Co.: Slightly saline soil, at Mississippi River 
bridge east of Arnprior, 27 miles west of Ottawa, 1959, Dore & Cody 17425 (MSC); 
river, Rockcliff Park, Ottawa, 1947, Calder, Dore, & Cody 1641 (MO). [Cochrane 
Co.}: Railway yards, ae 1952. Baldwin 3912 (CAN, GH). Durham Co.: 
Abandoned railway, Hope Township, 1948, Reeve 87 (DAO). Essex Co.: 1913, 
Malte sn. (CAN); oe open saline ground, Windsor o Factory, Windsor, 1975, 
Catling & McKay sn. (CAN, TRT). Huron Co.: Saline soil, south side of Maitland 
River, below Sifto Salt beta operation, Goderich, 1975, Catling & McKay 5.n. 
(CAN, TRT). Welland Co.: Railroad peondaouse: Niagra Falls, 1903, Scott s.n. 
(CAN, CU, DAO, TRT); Montrose railway yard, Niagra Falls, 1976, Catling & 
Mcintosh s.n. (DAO, TRT); ditch between two rail embankments, Bertie Township, 
Niagra, 1976, Catling & Riley sm. (DAO, TRT). York Co.: Dump heap, Ashbridge’s 
Bay, Toronto, 1904, Scott s.n. (CU, TRT); wet depression receiving run off from 
rock salt storage depot, opposite Don Valley Brickyard, Don Walley, Toronto, 197 4, 
Catling & McKay sn. (CAN, TRT); in snow dumping area, foot of Leslie Street, 
Toronto, 1974, Catling & McKay s.n. (TRT). 


DISTRIBUTION IN WESTERN NORTH AMERICA 


On the west coast, J. gerardii is known from the coastal salt marshes in 
the Puget Sound area in the state of Washington, on Vancouver Island, and 
in southern British Columbia. Some question has been raised as to whether 
or not the species is native there because the morphology of the plants 
more closely resemble those from Europe, whence they may have been intro- 
duced at an early date (Hitchcock, Cronquist, and Ownbey, 1969). The 
species was first reported from salt marshes at Victoria, Vancouver Island 
by Macoun (1838), based on his collection of 26 Jun 1887 (GH). How- 
ever, the species was not included in the Flora of Washington by Piper 
(1906). The earliest known specimen from Washington is dated 1917 and 
comes from Lopez on San Juan Island where it was obtained by S. M. and 
Mrs. E. B. Zeller (GH). Additional records were added in the 1920's and 
1930’s. Muenscher (1941) wrote the J. gerardii was local in salt marshes 
and cited two localities, South Bellingham and Marietta in Whatcom County. 
St. John (1928) considered the plants on the west coast as a native species 
and proposed the name J. fucensts. 

Inland in the Rocky Mountains of western United States, J. gerardii has 
been located in two states: 


342 
RADO: Known from wetlands at one site in Adams County and two 
locations in Boulder County, as reported by Hermann (1975). The earliest 
record is from the latter county, dated 1952. 
UTAH: Reported as he reg and established in salt marshes near Salt 
Lake City” (Tidestrom, 1925). This record is apparently based on a speci- 
men from a salt grass pasture near Salt Lake Say (Jun bree, G. QO. Bateman 
2., US). The species was not included in the Intermountain Flora by Cron- 
ae (1977): 


SOME GENERAL CONSIDERATIONS AND CORRECTIONS 


It has become apparent that the spread of J. gerardii inlard into the Great 
Lakes region and midwestern United States has come about by either migra- 
tion of the species via railroads or by establishment in inland saline marshes, 
as documented in the above specimen citations and summarized in Table 2. 

f the 58 known inland stations east of the Great Plains, 26 of them (44% ) 
are from along railroads and 13 of them (22%) are from inland saline habi- 
tats in New York, Ohio, and Ontario. Most, if not all, of the inland saline 
habitats cited here have been artificially created since the coming of Euro- 
pean man. Other significant localities are man-made roadside ditches and 
ballast ground, which account for the habitat at eight localities (14%). The 
occurrence in isolated sites and especially in man-created habitats which 
account for 80% of the known localities, strongly suggests that J. gerardii 
is a non-indigenous member of the wetland flora of the Great Lakes region, 
mid-western, and certain western areas in the United States. Furthermore, 
the occurrence of J. gerardii in the Finger Lakes region of New York, for 
example, conforms to Svenson’s idea that human agencies have been perhaps 
the most effective means of distributing these halophytic species into that 
region, rather than surviving as remnants from a_ post-Pleistocene marine 
submergence (Svenson, 1927). 

As pointed out by Eggleston (1904) in Vermont and by Schaeffer (1949) 
in Pennsylvania, J. gerardii has been transported as a salt hay in packing. 
In Massachusetts, Deane (1915) noted that black grass was cut in the sum- 
mer and fall and used for packing on celery beds to keep the stocks from 
freezing during the winter. The plants also can form “balls” or “pebbles” 
from matted sods which are thrown upon beaches during ditching opera- 
tions (Ganong, 1905). These sods are rolled about, worn down, and rounded 
by action of the waves and may even be transported 

Potter (1932, 1934) cited J. gerardii as an example of a halophytic species 
whose distribution supported the idea of a post-Pleistocene marine connec- 
tion between Hudson Bay and the St. Lawrence River basin. Those species 
which he used as evidence occurred in the southern region of Hudson Bay 
at James Bay, were disjunct to the Gulf of St. Lawrence, and then ranged 
southward along the Atlantic coast. J. gerardii, however, showed certain 
irregularities by having isolated occurrences in the Finger Lakes region of 


343 


TABLE 2. 


SUMMARY OF INLAND HABITAT TYPES OF J. gerardii IN RELATION TO THE 
NUMBER OF LOCALITIES IN THE MIDWESTERN AND GREAT LAKES STATES 
AND IN MANITOBA AND ONTARIO 


HABITAT NUMBER PERCENTAGE OF TOTAL 


OF LOCALITIES NUMBER OF LOCALITIES 

Railroad 26 _ : 4A 
Salt marsh; saline 

or brackish marsh 13 22 
Roadside ditch 4 7 
Ballast ground and dredgings 4 7 
Other habitats 11 20 
Total 58 100% 


New York, at the southern end of Lake Michigan, and on the Pacific coast. 
This irregularity has now been explained. Earlier, Peattie (1922) had cor- 
rectly omitted J. gerardii as an example of a natural member of the Atlantic 
Coastal Plain element in the Great Lakes region. Moreover, as reported and 
mapped by Rousseau (1974), J. gerardii is not known at James Bay, and 
he has stated that Potter’s specimen at the Gray Herbarium is J. alpinus, 
as determined by L. Cingq-Mars. The report from a James Bay by Macoun 
(1888), which Rousseau could not verify, apparently is from a local James 
Bay at Vancouver Island, British Columbia. Macoun’s published report is 
clear, “Salt marsh at the head of James Bay, Victoria, Vancouver Island, 
1887. (Macoun.),” and agrees with the date on the label of his specimen 
(24 June 1887, Macown 27,869, CAN). 


ACKNOWLEDGEMENTS 


My thanks are extended to Mr. Mark Davis who assisted with this research 
during the summer of 1977. My thanks are also extended to the curators of 
the herbaria who made specimens and records available. Those herbaria in 
which I have examined specimens are: BH, CAN, : : : ; 
MO, MSC, NYS, OS, OSH, PENN, PH, US, Those herbaria from which 
only records were seen are: GH, MIN, NY, TRT, WIS. 

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- 665 + pls. 1-15. (Reprinted in the series, Reprints of US-Floras, Vol. 3. Verlag 

n J. Cramer, Lehre. 

TORREY. J. 1824. Flora of the northern and middle sections of the United States; 
or, A systematic arrangement and description of all the plants hitherto discovered 

in the United States north of Virginia. T. and S. Swords, New York. 

_ «1843. A Flora of the state of New-York, comprising full descriptions of 
all the ‘indigenous and naturalized plants hitherto discovered in the state; with 


remarks on their economical and medical properties. Vol. II D. Appleton & Co., 


New York. 572 p 
VOSS, E. G. 1972. MMichieee Flora: A guide to the ee: fication and occurrence of 
the native and naturalized seed-plants of the state Part I gymnosperms and monocots. 
Cranbrook Inst. Sci., ae Hills, Bull. 55, and Univ. Michigan Herb., Ann 
Arbor. xviii + 488 

WARD, D. B. 1968. Checklist of the vase) @y noe of Florida Part 1. Agric. Exp. Sta., 
Univ. Florida, ne Tech. Bull. 726. 72 pp. 

WHEELER, C. nd E. F. SMITH. 1881, cues Flora. Ann. Rep. State Hort. 
Soc. Michigan Eciaan 10: 427-529. (Reprinted with title page and repaged 1-105, 
as catalogue of the phaenogamous and vascular cryptogamous plants of Michigan, 
indigenous, a eaees: and adventive. ) 

WIEGAND, K. M. 1900. aed tenuis Willd. 


allies. Bull Torrey oe Club 27: 511-527. 
nd A. J. EAMES. ice The flora has the Cayuga Lake Basin, New York. 
1 pp 


i 


and some of its North American 


~ Cornell Univ. Agric. Exp. Sta. Mem. 92. 
ZENKERT, C. A. 1934. The flora of the 


Niagara frontier region. Ferns and flower- 
ing plants of Buffalo, N.Y., and vicinity. Buffalo Soc. Nat. Sa. Bull. 1 


6: 1-x, 1-328. 


NOTES ON THE FLORA 
OF THE CHINATI MOUNTAINS, 
PRESIDIO COUNTY, TEXAS 


EMILY J. LOTT 


Herbario Nacional, Instituto de Biologia, UNAM, 
Apartado Postal 70-233, México 20, D.F., México 


MARY L. BUTTERWICK 


Bureau of Land Management, Phoenix District Office, 
2929 West Clarendon Avenue, Phoenix, AZ 85017 


The first definite records of botanical observations in the Chinati Moun- 
tains are those of V. Havard (1885). G. C. Nealley (1888) collected grasses 
and forage plants and made general observations in western Texas, visiting 
the “Chenate” Mountains in late October, 1887. M. S. Young made general 
collections in the foothills of the Chinatis in September, 1914, but probably 
never reached the main mass of the mountains (Tharp 1962). The most 
extensive botanical explorations of the Chinatis were begun by L. C. Hinck- 
ley in August, 1936. Collecting trips were made to Chinati Peak in 1945 
by C. H. Muller and Rogers McVaugh. B. H. Warnock began working in the 
area both alone and with Hinckley during the 1940's; specimens from these 
and subsequent trips are deposited at TEX-LL and SRSC herbaria. 

The present authors made collecting trips into the area in June and 
October of 1977. Voucher specimens will be deposited in the University 
of Texas at Austin Herbaria (TEX-LL). Nomenclature for species previously 
reported from Texas is in accordance with Correll and Johnston (1970). 

The rugged summits of the Chinati Mountains rise over 1400 m above 
the surrounding pediments, dominating the 30 km expanse between Pinto 
Canyon and Shafter in south central Presidio Co. The Chinati Mountains 
are one of three major mountain ranges in the Big Bend area of Trans-Pecos 
Texas that are of igneous origin, the others being the Chisos and Davis 
Mountains. The considerable erosive force of tributaries to the Rio Grande, 
such as Pinto and Dead Horse canyons, has uncovered various sedimentary 
rocks of Permian and Cretaceous age. These localized outcrops of limestone, 
sandstone, and shale, primarily along the margins of the intrusive and volcanic 
rocks, are the substrates for plant assemblages that are noticeably different 
from those found on neighboring igneous slopes. 

n the limestone hills just N of Chinati Peak, associated with Agave 
lecheguilla, Dasylirion leiophyllum, Quercus pungens, Mortonia scabrella, 
Leucophyllum frutescens, Ayenia pilosa, and Bouteloua ramosa, we found two 
species of particular interest. 


SIDA 8(4): 348-351. 1980. 


349 


POLYGALA NUDATA Brand. On limestone hills N of Chinati Peak between 
Dead Horse and Pinto Canyons; Butterwick 3807B; 9 Jun 1977. This is the 
fourth known United States locality for this perennial Polygala, which until 
recently has been placed in synonymy with the Mexican P. minutifolia Rose 
(Wendt, 1978). Polygala nudata is characterized by its upright broom-like 
incurved-puberulous stems and short seed pubescence. 

PETERIA SCOPARIA Gray. Infrequent on limestone hills N of Chinati Peak; 
Butterwick & Lott 3829; 11 Jun 1977. This species is previously unreported 
from the Chinatis but its presence there could be expected. It is poorly repre- 
sented in herbaria and generally occurs in small, scattered populations. 

Indian Cave Canyon, a side canyon to Dead Horse Canyon, on the N side 
of the Chinatis, provides a sheltered, more mesic habitat for Prunus serotina, 
Fraxinus cuspidata, F. arizonica, Quercus gambelii, Q. grisea, Symphoricarpos 
sp., Heuchera rubescens, Aquilegia sp., and Pericome caudata in its upper 
reaches, which lead to Chinati Peak. In this canyon we collected a species 
of fern reported for the first time from the Chinatis. 

POLYPODIUM THYSSANOLEPIS KI. Uncommon in crevices of cliffs and 
boulders, lower Indian Cave Canyon, with Garrya ovata, Selaginella rupincola, 
Bommeria hispida, Eupatorium rothrockii, Ungnadia speciosa, Silene laciniata, 
Sedum wrightii, and Cheilanthes wrightii; Butterwick & Lott Oct 
1977. According to T. L. Wendt,’ who is preparing a treatment of Chihuahuan 
Desert Polypodium, this collection is of an undescribed variety of the south- 
western United States and northern Mexico. It was previously known in 
Texas only from the Davis and Chisos Mountains. 

The summit of Chinati Peak is surprisingly level, resembling a high 
mesa top. Stipa tenwissima occurs beneath the numerous gray oaks, and 
Bouteloua gracilis covers the more open areas. Sedum wrightu, Talinum 
parviflorum, Heuchera rubescens, Aletes acaulis, and Triodanis biflora are a 
few herbaceous species collected on or near the peak. 

OPUNTIA POLYACANTHA Haw. var. TRICHOPHORA (Engelm. & Bigel.) 
Coult. Locally common among grasses on the summit of Chinati Peak; A. M. 
Powell et al. 3109; 7 Jun 1977. This collection represents the southernmost 
known population of this widespread species, which ranges as far north as 
Peace River, Alberta. The variety occurs in Trans-Pecos Texas and the Pan- 
handle, to Colorado and Arizona through western Oklahoma. 

GAURA HEXANDRA Gomez Ortega subsp. GRACILIS (Woot. & Standl.) 
Raven & Gregory. Rare in rocky igneous soils near summit of Chinati Peak, 
ca 2313 m elev; Butterwick & Lott 3774; 7 Jun 1977. Scarlet flowers, red- 
dish stems with white spreading hairs, and truncate fruit bases distinguish 
this species, a new state record for Texas and an extension of its range 


1The undescribed variety of Polypodium mentioned above is now published as 
P. thyssanolepis Kl. var. riograndense Wendt (Amer. Fern J. 70: 5-11. 1980). 


350 


from Arizona and New Mexico to Sonora, Chihuahua, and Durango, and 
now to western Texas. 

Two large canyons on the S§ side of the Chinati Mountains yielded note- 
worthy collections. Both are relatively closed drainages whose walls provide 
some protection from the southwesterly winds. Of particular interest was 
Tinaja Prieta (Palillos) Canyon, with its numerous tinajas and seep springs. 

PLUMBAGO SCANDENS L. On low gravelly terrace of lower Tinaja Prieta 
Canyon, just above falls, and med the upper reaches of the canyon; 
Butterwick & Lott 3872; 16 Jun 1977. The range in Texas is now extended 
from Cameron and Hidalgo counties in extreme southern Texas. Known 
also from Pima Co., southern Arizona, and from northern Mexico and south- 
ern Florida, this species is widespread in the tropical Americas. 

SIPHONOGLOSSA LONGIFLORA (Torr.) A. Gray. Rare among boulders in 
lower part of E branch of San Antonio Canyon, § side of Chinati Mountains; 
Butterwick & Lott 3864B; 15 Jun 1977. The essentially glabrous S. longiflora 
has petioled leaves and clusters of white tubular flowers in the upper axils 
which easily distinguish it from the coarsely pubescent S. pilosella. The 
Chinati Mountains locality for this species extends its range eastward from 
southern Arizona and Sonora and is apparently the first documented report 
of the species from Texas. This species is not included in the Manual of 
the vascular plants of Texas (Correll & Johnston 1970), nor are any previ- 
ous Texas collections on deposit in TEX-LL, SRSC or GH. Rickett (1970) 
cited the species as “reported in western Texas” but stated that he had seen 
no specimens of it from Texas. 


ACKNOWLEDGMENTS 


We thank P. H. Raven for determination of the Gawra specimen and 
T. L. Wendt for his verification of our pteridophyte and Polygala determi- 
nations. J. Henrickson helped locate specimens of Siphonoglossa longiflora, 
and R. A. Hilsenbeck verified our determination and graciously provided 
useful information on this species. We are grateful to A. M. Powell for 
reviewing the manuscript, and to D. E. Deal for arranging permission with 
the landowners. Texas Natural Area Survey is gratefully acknowledged for 
financial and logistical support of field work in the area. 


REFERENCES 
CORRELL, D. S. and M. C. JOHNSTON. 1970. ines of the vascular plants of 
Texas. Texas Research Foundation, Renner, Texa 
HAVARD. V. 1885. Report on the flora of western and southern Texas. Proc. U. S. 
533. 


NEALLEY, G. C. 1888. Report of an investigation on the forage plants of western 
Texas. U. . Dep. Agr. Div. Bor Bull. 6. U. S. Government Printing Office, Wash- 
ington, 

RICKETT, - W. 1970. Wild flowers of the United States. Vol. 3: Part 2 (Texas). 
McGraw-Hill, New York (publication of the New York Botanical Garden) 


peak 


THARP, B. C. and C. V. KIELMAN, eds. 1962. Mary S. Young's journal of botanical 
explorations in Trans-Pecos Texas, August- eis 1914. Southwest. Hist 
65: 366-393; 512- a 
WENDT, T. L. 1978 systematic study of cae. section Rhinotropis (Poly- 
galaceae). Ph.D. .. University of Texas, Aust 


NOTES 


DISTRIBUTION RECORDS FOR DIGITARIA BICORNIS IN EAST- 
ERN UNITED STATES—Henrard’s Monograph of the genus se 
(1950) restricted the distribution of Digitaria bicornis (Lam.) R ; 
tropical crabgrass, to tropical Asia. Swallen (1963) described D. dieters 
a later synonym of D. bicornis, based on type material from Jamaica and 
gave the distribution in the United States as Florida and Texas. Gould 

5) reported D. bicornis in Texas as occurring in the southeastern 
prairies and coastal marshes. Correll & Johnston (1970) stated that D. 
diversiflora (ie. D. bicornis) is probably indigenous to and is common on 
the Rio Grande Plains. 

Recent collections of D. bicornis | have made in eastern United States 
have shown that this taxon is much more widespread than previously be- 
lived. It was found to be common and abundant on the sandy coastal plain 
of the southeastern states. The range of the species is now known to include 
eastern North and South Carolina, and eastern Texas (Fig. 1). The distribu- 
tion of this species seems to be limited to areas of coarse-textured soils and 
moderate to high rainfall. Populations of D. bicornis and D. ciliaris (Retz. ) 
Koeler, southern crabgrass, frequently grow intermingled, which possibly 
accounts for D, bicornis being frequently mistaken for the latter, better- 

nown species. Morphological distinctions between D. ciliaris and D. bicornis 
are given by Gould (1975) 

Voucher specimens (TAES) of D. bicornis were collected at locations in 
six states from which the species has not previously been reported: North 
Carolina, South Carolina, Georgia, Alabama, Mississippi, and Louisiana. This 
crabgrass was found to be an important invading species of cultivated soils 
throughout the southeastern coastal plain. Specimens were not collected 
north of North Carolina; however, there seems to be no environmental 
reason why this species should not occur in sandy coastal areas of Virginia 
and Maryland. Texas collections I have made extended the range of D, 
bicornis into the pineywoods and post oak savanna vegetational regions of 
the eastern portion of that state. The northernmost collection site in Texas 
was only 30 miles from the Arkansas border and 60 miles from the Okla- 
homa border; future collections extend the range of the species north 
into Avlainsas and Oklahoma. t D. Webster, Department of Range 
Science, Texas AEM University, Colleve Station, TX 77843 


REFERENCES 
CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the vascular plants of 
Texas. Texas Research me Renner, Texas. 1881 pp. 
GOULD, F. W. 1975. Grasses of Texas. Texas A&M University Press, Col 
tion, Texas. 653 pp. 


ege Sta- 


SIDA 8(4): 352. 1980. 


eee) 


Figure 1. Distribution of Digitaria bicornis in southeastern United States. 


HENRARD, J. T. 1950. Monograph of the genus D¢gitaria. Leyden, Universitare 
Per Leiden. 999 pp. 

SWALLEN, J. R. 1963. New species of Digitaria and Trichachne. Rhodora 65: 
22-307. 


NAJAS MARINA: NEW TO THE INDIANA FLORA—On a recent 
plant collecting trip through the Midwest, two collections were made of 
Najas marina L., the prickly naiad, in northeastern Indiana. After consult- 
ing the herbaria at IND and ND (acronyms follow Holmgren & Keuken, 
1974), I have determined that this species is new to the Indiana flora. Col- 
lecting data for N. marina in Indiana are as follows: Steuben Co., off hwy 
120 near I-69 at Green’s Lake, ca 7 mi N of Angola, 20 Jul 1979, Davenport 
1424; Noble Co., on hwy 3 at W side of Cree Lake, S of South Milford, 
20 Jul 1979, Davenport 1427. Voucher specimens for both collections are 
deposited at UNA. 


SIDA 8(4): 353. 1980. 


354 


In North America, Najas marina is found infrequently in brackish or 
highly alkaline waters of the western United States, Texas, the Great Lakes 
states, and Florida (Haynes, 1979). Whether it is a native or a naturalized 
J is under question. After studying its distributional history, Wentz 

& Stuckey (1971) concluded that N. marina was introduced into the Ohio 
region, and fairly recent collections in Michigan (Near & Belcher, 1974), 
Wisconsin (Ross & Calhoun, 1951), and Illinois (Winterringer, 1966) 
seem to bear out this conclusion. Its occurrence in Indiana, therefore, is to 
be expected and indicates the further spread of this species in the Great 
Lakes region —L, J, Davenport, Ecology and Systematics Section, Department 
of Biology, The University of Alabama, University, AL 35486. 


REFERENCES 
—— vi R. 1979. Revision of North and Central American Najas (Najadaceae) . 
—56 


nae P. K. and W. KEUKEN. 1974. Index herbariorum, part 1: The 
eae of the world. Utrecht, Netherlands. 397 pp. 

NEAR, K. A. and R. O. BELCHER. 1974. New _— for Najas minor and 
marina in eee Michigan. Michigan Bot. 1-185. 

ROSS, J. G. and B. M. CALHOUN. 1951. ee reports on flora of 
Wisconsin. XX XIII. Naladaae Trans. Wisconsin Acad. Sci. 40: 93-1 

WENTZ, W. A. and R. L. STUCKEY. 1971. Se changing distribution . ae genus 
Najas CNiadacess) <1 in aoe Ohio J. Sci. 71: 292-302. 

WINTERRINGER, G. S. 1966. Aquatic note plants new for Illinois. Rhodora 68 
221-222. 


on 


GAURA ODORATA LAG, (ONAGRACEAE) IN LOUISIANA.—Speci- 
mens from a small population of Gawra found in Lafayette were identified 
through the keys of Munz (1965) and Correll and Johnston (1970). Com- 
parison with specimens borrowed from SMU confirms identification as Gaura 
odorata Lag., which can now be recorded from Louisiana. Lafayette Parish: 
rhizomatous perennial growing aggressively with Cynodon dactylon on un- 
cemented square around telephone pole in parking lot on N. University Ave. 
near corner of Louise St. Lafayette, Vincent 2310 (LAF), 18 Jun 1979. 

The plants were observed to flower and produce fruit sporadically from 
March to October 1979 in spite of several mowings. They were vigorously 
producing new shoots in March 1980 and thus may persist and disperse in 
the area. Their presence in Lafayette marks a significant eastward range 
extension for the species, a native of Texas and adjacent Mexico that is 
rare in east Texas. Previously unreported from Louisiana, G. odorata was 
not included in the preliminary report on Louisiana Onagraceae by Ellis 
and UrBatsch (1979). 

Duplicates are being sent to AC, GA, NCU, NLU, NO, SMU, and VDB. 


SIDA 8(4): 354. 1980. 


355 


I thank Dr. William F. Mahler for the loan of specimens—Karl A. Vincent, 
University of Southwestern Louisiana, Lafayette, LA 70504 


REFERENCES 
CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the vascular plants of 
Texas. Texas Research see Renner, Texas. 
ELLIS, R. R. and L. E. URBATSCH. ar Preliminary reports on the flora of 
Soar IV. Onagraceae. ee 44: 28-38. 
UNZ, P. A. 1965. Onagraceae. North American Flora. New York Botanical Garden. 
Series II, part 5: 1-231. 


A NOTE ON FLOWER COLOR OF WAHLENBERGIA LINARIOIDES 
(LAM.) A. DC. IN NORTHWEST FLORIDA—A recent article by D. B. 
Ward (Phytologia 39: 1-12, 1978) includes a key for distinguishing the 
two known species of Wahblenbergia which occur within the state of Florida: 
W. marginata (Thunb.) A. DC. and W. linarioides (Lam.) A. DC. The 
key was adapted and expanded from a previous key and descriptions pre- 
sented in an article by R. K. Godfrey (SIDA 1: 185, 1963). In both of 
these papers the flower color of W’. /inarioides is stated as blue 

During more than ten years of botanizing in the Pensacola area I have 
encountered W’. linarioides at numerous sites along roadsides and in sandy 
pinewoods. All of the observed specimens have had white flowers. I have 
recently discussed this with R. K. Godfrey, and he, too, has seen primarily 
white-flowered specimens of W’. linarioides in recent years. 1 conclude that 
current populations of W.. linarioides in northwest Florida are predominantly 
white-flowered, and that if blue-flowered individuals exist they are rare 
indeed. 

The fact that W. linarioides has white flowers makes it easy to distinguish 
in the field from W’. marginata, which has blue flowers, and botanists 
collecting in northwest Florida should take note of this. The following 
white-flowered specimens of W. linarioides may be examined in the herbaria 
indicated: Burkhalter 3908 (UWFP), 6374 (UWFP, CAS) —James R. 
Burkhalter, University of West Florida, Pensacola, FL 32504. 


BRACHYELETRUM ERECTUM AND TALINUM RUGOSPERMUM, 
NEW SPECIES TO TEXAS AND NOTES ON SCHOENOLIRION 
WRIGHTIU—tThe grass Brachyeletrum erectum (Schreb.) Beauv. is gener- 
ally distributed throughout the eastern United States, including Oklahoma 
and Louisiana (Hitchcock, 1950; Gleason, 1968). It is not surprising there- 
fore, that it occurs in eastern Texas. Brachyeletrum erectum was initially 
found in 1971 (McCrary 171 ASTC) and has since been collected from 
the same location (Kyle 40, Jun 1976 ASTC; Nixon 8723, Aug 1978 
ASTC). Generally this species occurs in dry to moist wooded areas. We 


SIDA 8(4): 355. 1980. 


356 


found it within Nacogdoches County along a branch of Naconiche Creek 
about 15 miles northeast of Nacogdoches, Texas. Several small populations 
were present. It was within a forest dominated by Fagus grandifolia, 
Carpinus caroliniana and Liquidambar styraciflua (Nixon and Raines, 1976). 

Talinum rugospermum Holz. was collected from the extreme northern 
portion of Nacogdoches County along FM 1087 (Nixon 7752, Aug 1977 
ASTC; Nixon and Marietta 9492, Jul 1979 ASTC). The species occurred 
occasionally in open, rather disturbed areas, on deep sandy soils. The reported 
distribution of T. rvgospermum was the north central United States—lIa., 
Minn., Wis., Ill, and Ind. (Rickett, 1969; Gleason, 1968). It grows in 
sandy soils or on sandstone in that area. It is somewhat amazing, therefore, 
that this species is encountered so far south. 

Page wrightii Sherman (S. texanum (Scheele) Gray; Sherman, 

79) is a rare small white-flowered species that grows on moist prairies 

in nee Alabama, and East Texas (Hardin, Austin, Brazos, and Walker 
counties). It has been proposed as an endangered species (U.S. Dept. of 
the Interior, 1976) and was last collected in Texas in 1963 in Walker County 
(Sherman 560, 15 Apr 1963 VDB). Schoenolirion wrightii was collected 
in Apr 1978 (Marietta and Nixon 61 ASCT) in an open prairie like area 
in Jasper County. This collection extends the known range of S. wrighti 
somewhat and verifies its present existence in East Texas—Elray S. Nixon 
and Kay L. Marietta, Biology Department, Stephen F, Austin State Univer- 
sity, Nacogdoches, TX 75961; Michael McCrary, National Plant Materials 
Center, Beltsville, MD 20705. 


REFERENCES 

GLEASON, H. A. 1968. New Britton and Brown illustrated flora of the northeastern 
United States and adjacent Canada. Hafner Publishing Company, Inc. New York. 

HITCHCOCK, A. S. 1950. Manual of the grasses of the United Sie. United States 
Dept. of Agr. Misc. Pub. No. 200, Washington D.C. 

NIXON, E. S. and J. A. RAINES. 1976. so creekside vegetation of Nacogdoches 
County, Texas. Texas J. Sci. 27: 

RICKETT, H. W. 1969. Wild Rowers - a United States: Volume 3, Texas. Mc- 
Graw-Hill Book Company, New Y 

SHERMAN, H. L. 1979. Evidence “of misapplication of the name Schoenolirion 
texanum (Scheele) Gray (Liliaceae). Southw. Nat. 24: 123-126. 

U. S. DEPARTMENT OF THE INTERIOR. 1976. Endangered and threatened species 
—plants. Federal Register 41: 24524-24572. 


HOLOSTEUM UMBELLATUM (CARYOPHYLLACEAE) IN ON- 
10.—The spread of Holosteum umbellatum L., jagged chickweed, in 

the United States has been discussed by Shinners (1965). This species, evi- 
dently native to Europe and/or western Asia, is now found throughout the 
United States, but the colonies tend to be widely separated and in some 


SIDA 8(4): 356. 1980. 


357 


cases not long persistent. It is most common in the warmer regions, and 
records from Michigan and New England remain few. 

The only recent collections of H. wmbellatum from Canada have been 
from southwestern British Columbia (Scoggan, 1978). Campbell (1895 ) 
reported having found H. wmbellatwm at Kettle Point, Lambton County, 
Ontario, in 1894. This record, if correct, represented a considerable westward 
expansion of the known range of H. wmbellatum in North America at the 
time (see quotations in Shinners 1965). Later authors, however, have found 
no specimens documenting this report and have cited it with a question 
mark (e.g. Scoggan 1978) or omitted it. Specimens from Campbell's her- 
barium, including many from Kettle Point, are in M MG, but, according 
to information kindly provided by Marcia J. Waterway, there is no speci- 
men among them now or formerly identified as H. #mbellatwm, either under 
Holosteum or in a genus that might likely have been confused with 
Holosteum (other Alsinoideae; Androsace). It seems probable, therefore, 
that this report was erroneous, and that the specimen was reidentified soon 
thereafter, especially if, as the title of Campbell’s paper impies, the plants 
in question flowered in summer. (Holostewm wmbellatum is in mature fruit 
by mid-spring, and the remains of the plants disappear in summer.) There 
have been no other reports of H. wmbellatum from eastern Canada. 

Flowering stems of H. wmbellatum were brought to me for identification 
on 18 April 1980 by J. A. Runnells of Stoney Creek, Ontario. On 2) spit 
following Mr. Runnells’ directions, I found a population of H, umbellatum 
comprising thousands of plants along the Canadian National tracks near the 
Royal Botanical Gardens’ Rock Garden in Hamilton, Hamilton- Wentworth 
Region (43° 17’ 17” N, 79° 53’ 32” W). Here the railroad runs through 
a cut in a sandbar of Lake Iroquois age, with H. wmbellatum growing on a 
sunny south-facing slope of the cut, in dry sandy soil, intermixed with 
Bromus tectorum (Pringle 1936, DAO, HAM, OAC, MO). On 5 May 1980, 
I found numerous plants of H. wmbellatum in Burlington, Halton Region, 
growing as weeds in crushed-stone walks in the R.B.G.’s Hendrie Park 
(43° 177 30” N, 79° 52’ 30” W, 148 km ENE of the Hamilton popula- 
tion), on nearly level terrain, in full sun, associated with Erophila verna 
and Sedum acre (Pringle 1937, HAM). Some plants were also found in 
nearby lawns where the soil is gravelly, derived from Lake Iroquois beach 
deposits (Pringle 1938, CAN, HAM, UWO). 

Holosteum umbellatum can readily be distinguished from related plants 
of the subfamily Alsinoideae by its umbellate inflorescences and by its 
petals, which are jaggedly few-toothed at the apex but not regularly bifid. 
Following Walters’ (1964) treatment of Holostewim, these Ontario popula- 
tions represent H. wmbellatum ssp. umbellatum—Iames S. Pringle, Royal 


SIDA 8(4): 357. 1980. 


358 


Botanical Gardens, Box 399, Hamilton, Ontario, Canada L8N 3H8. (Con- 
tribution No. 40 from the R.B.G.) 


REFERENCES 


CAMPBELL, R. 1895. Some of the rarer summer flowers of Canada. Canad. Rec. Sci. 
6: 342-351, 

SCOGGAN, H. J. 1978. Satente In: The Flora of Canada 3: 690. Natl. Mus. 
Nat. Sci. Publ. Bot. No. 

SHINNERS, L. H. 1965. | ven eases go Ra vaee population ex- 
plosion and fractionated suicide. Sida 2: 119-1 

oo S. M. 1964. Holosteum L. In: ae a G., et al., eds. Flora Europaea 
1: 136. Cambridge: University Press. 


SIDA 8(4): 358. 1980. 


SID 


CONTRIBUTIONS 
TO BOTANY 


INDEX TO VOLUME 8 


Names of contributing authors are in capital letters. New scientific names 


are in boldface. Synonyms are in italics. 


Abutilon theophrasti 84 
Ac alypha gracilens 80, 145; rhomboidea 


Acer ooiado 142; pensylvanicum 142; 
rubrum 137, 142; saccharinum 75, 


138, 14 
Achillea millefolium 77 
Acorus calamus 


Aegilops cylindrica 81 

Aesculus glabra ae Rar 138, 146; 
pavia 102, 107, 
Agalinus aa a ie 

Agavaceae 314 

Agave lecheguilla 245, 348; virginica 
142 


Agrimonia parviflora 86, 149; pubescens 
149; rostellata 14 
Agropyron cristatum 168; repens 81 
Agrostis hyemalis 81; perennans 148; 
81 


Ailanthus altissima 150 


Aizoaceae 75 
Albizi oer 145 
eee acaulis 349; farinosa 147 
lisma HRS -aquatica 73, 75; sub- 


cordatum 142; triviale 119 

Alismataceae 75, 142 

Allium canadense 84, 147; sativum 84; 

vineale 84 

Alnus surrulata 144 

Althaea rosea 84 

Amaranthaceae 75 

Amaranthus hybridus 75; spinosus 108 
42 


bidentata 77; trifida 77 
Amelanchier arborea 74, 86; canadensis 
149 


. 


Amorpha canescens 83 


76, 142 
Andropogon gerardii 72, 81, 148; sco- 
parius 148; ternarius 190, 191; vir- 
, 148, 190, 191 


Anemone quinquefolia 149; virginiana 
Angelica venenosa 88, 142 
Anisostichus capreolata 144 
Annonaceae 
Antennaria plantaginifolia 77, 142; soli- 
taria 142 
Anthemis cotula 77 
Apiaceae 142 
pios americana 74, 83, 145 
Aplectrum hyemale 147 
Apocynaceae 76 
Apocynum cannabinum 76 
Aquifoliaceae 
Aquilegia 349; canadensis 14 
Arabis canadensis 144; laevigata 144 


5 

Aralia racemosa 142; spinosa 142 

Arctium minus 77 

Arisaema macrospathum 119; triphyllum 
42 


Aristida dichotoma 81, 148; longespica 
90; ee 

Ri ciclochiae ae 14 

Aristolochia an 138, 142; ser- 

pentaria 142 

Arnica wilsonii 344 

Aruncus dioica 149 

gee gigantea 148 


Asarum can 

Meclepiadacens 6, 142 

Ascleptias emplestcau 142; hirtella 76; 
incarnata 76; purpurascens 76; quad- 


rifolia 142; syriaca 76; ated 76, 


360 


142; variegata 142; verticillata 76 


Ascyrum hypericoides var. hypericoides 
190 


Asimina triloba 142 
Asparagus officinalis 84 
Aspleniaceae 
Asplenium montanum 141; platyneuron 
a 141; resiliens 213; ghizophy liom 


> 
— 


14 
Aster 35; cordifolius 142; divaricatus 
142; dumosus 143; infirmus 143; 


laevis 143; lateriflorus linarii- 
folius 143; SOwaTEE ANS 14 eeu 
folius 77; pate : > paternus 
143; pilosus var. demotus 77; praeal- 


simplex 143; so lidagineus 143; sur- 
culosus 143; umbellatus 137, 143 

Asteraceae 

Astilbe biternata 150 

Atamasco texana 329, 330; tubispatha 
330 

Athyrium asplenioides 141; pyenorat pon 
138, 141; thelypterioides 141, 213 

Aureolaria ievieats 150; wiewinies 150 

Austrobassia 57 

Avena sativa 81 

Axyris amaranthoides 168 

Ayenia pilosa 348 

Baccharis halimifolia re 198 

Balsaminaceae 76, 14 

Baptisia leucantha 83. leucophaca 83; 
tinctoria 14 

Barbarea vulgaris var. arcuata 79 

Bartonia virginica 146 

Bassia 57; hirsuta 57, 59, 60; hyssopi- 
folia 57, 58, 60, ap 161 


— 


Betula lutea 144; nigra 144 
Betulaceae 76, 144 
ae bipinnata 77, 143; frondosa 77 
; polylepis 77, 143 

a 76, 144 
— era 29 

BLAC Sih eee H.. JRa D7; 
Blechnaceae 
Bochmeria Te 88, 150 
Bommeria hispida 349 


~~ 


ch 


7, 10 
eke dissectum 14 1; virginianum 
85 


ea gracilis 349; ramosa 348 


ae eae erectum 148, 354 
ractearla amazonica 92 

Bele, juncea 79; coe var. pinnati- 
fida 79 

Brassicaceae 144 

Brazoria pulcherrima 211 

Bromus commutatus 81; japonicus 81; 
ee ns 148; secalinus 148; tectorum 


tS 


sD 

a RALPH E. 
aoe ves ae 8: anne 150 
BURK , JAMES R. 354 
Seni Wee 315, 348 
Buxaceae 144 
Cacalia atriplicifolia 143 
Calamogrostis cinnoides148 
Callicarpa americana 188, 190 
Callirhoe digitata 84 
Callitrichaceae 7 
Call ses 34, 35, 56; heterophylla 76; 

terrestris 76 
C he ccae 144 
Calycanthus floridus 137, 
Calystegia aa iy, spithamaca 144 


Comsanida cee a 144 

Campanulaccae 76, 

Campsis radicans i. en 109, 144 

Capparidaceae 76 

Caprifoliaceae 76, oe 

Capsclla bursa-pastori 

Cardamine hirsuta 144: 
arenicola 79 

Carduus nutans 77 

Carex 73; aestivalis 144; albolutescens 
144; albursina 144; amphibola 144; 


~! 
\o 


; parviflora var. 


meadii 79; molesta 79; muhlenbergii 
79; pedunculata 139, 145;  pensyl- 
vanica var. distans 145; plantaginea 
145; purpurifera 145; retroflexa 145; 
rosea 145; scoparia 73, 79; squarrosa 


79: swanii 145; tribuloides 79; vire- 
scens 145; vulpinoidea 73, 79 
Caroliana 35 
Carpinus caroliniana 139, 144, 210, 355 
Carya cordiformis 146; glabra 146; 


146; pallida 146; texana 74, 82; 
tomentosa 74, 82, 138, 146 
Guia 77, 144, 355 
Cassia marilandica 145; nictitans 83, 
145 


Castanea dentata 146 

Cathartolinum elongatum 182 

CATLING, P. M. 188 

Caulinia 35, 36; alagensis 45; flexilis 
40; fragilis 51; guadalupensis 43; 
indica 51; minor 51; tenella 

Caulophyllum thalictroides 144 

Ceanothus americanus 149; var. pitcheri 
86 


Celastraceae 77, 144 
Celastrus scandens 77 
Celtis laevigata 107, 108; occidentalis 


Ceachnae incertus 205; longispinus 81 

Centaurea maculosa 77 

Centrosema 152; angustifolium 152, 
155; galeotii 155: grandiflorum 155; 


Cephalanthus occidentalis 73, 87, 14 

Ceras ae Le ratum 77; holosteoides 
var. vulgare 144; vulgatum 

Cercidium 216 

Cercis ae, 83, 14 

Chaero phyllum procumbens 88; taintu- 


142 
Ghanaians luteum 138, 147 
ee exumensis 319; lechioides 
ar. exumens is 319; var. wilsonii 319; 
ems z 319 
c eR latifolium 148; laxum 


Cheilanthes alabamensis 314; lanosa 
141; wrighti1 3 
Chelone glabra 150; lyonii 138, 150 
Chenolea 58 
Chenopodiaceae 57, 77, 157 
Chenopodium album 77 
imaphila maculata 145 
Chondrophylla 14 


361 


Chrysanthemum leucanthemum 77, 143 
Chrysopsis pilosa 

Cichorium Hes vi: 

Cicuta maculata 142 

Cimicifuga mre 140, 149 
Ciminalis 16 

Cinna arundinacea 73, 148 

Circaea lutetiana ssp. ome 147 
Cirsium altissimum 3; muticum 


143; vulgare 77 
Cissus incissa 108, 109 
Cistaceae 77, 144 


Cladrastis lutea 138, 145, 216 

Clarkia 295 

Claytonia caroliniana 138, 149; virgin- 
ica 86 

Clematis oe dictyota mace integrifolia 
119; pitcheri 194; var. dictyota a 194, 
195; var. anes ie 195; reticulata 
8; virginia 

Clintonia osalis 

Clitoria 90; gee a 90, 91; amazo- 


7; 
arborescens 91, 94; brachycalyx 93, 
94, 304, 305; brachystegia 94, 304; 


subgenus bractearia 90, > sect. 
brachycalyx 93, 304; sect. ibractearia 
5; sect. caulifl > sect 


flexuosa 93; froesil 
311; glaberrima 94, 304; Peione 
155; hermannii 304, 305, 306; 
hoffmannseggii 91; javitensis A 


juninensis 95, 96, , 99, 1; 
leptostachya 94; mariana 145; ee 
bambensis , 100, 101; nervosa 
3, 95; obidensis 94; poitaei 90, 91; 
pozuzoensis 93; rac $a ; 
sagotii 94; selloi 94 


COLLINS, SCOTT L 

Collinsonia emnadengs io 146, 216 

Comandra richardsiana 87 

Commelina communis 77; erecta var. 
angustifolia 

Commelinaceae 77, 144 

Compositae 65, 77, 216, 313 

Conopholis americana 14 

Convolvulaceae 78, 144 

Convolvulus arvensis 78 


362 


Corallorhiza odontorhiza 147 

Coreopsis auriculata 143; major var. 
stellata 143; palmata 77 

Cornaceae 79, 144 

Cornus alternifolia 144; amomum 139, 
144; florida 74, 79, 144 

CORRELL, DONOVAN S. 317 
“orylus americana 76, 144 

Crassulaceae 144 

Crataegus crusgalli 86, 149; flabellata 
149; flava 149; viridis 149 

Crotalaria sagittalis 83 


< 


Croton capitatus 80; glandulosa var. 
septentrionalis 80; monanthogynus 
80, 145 


Cruciferae 79 
“ryptotaenia eter 142 

Cucurbita digita 

Cucurbitaceae 135 

Cunila 220; galioides 293: montana 
290; origanoides 82; piperita 251: 
pulegioides 288 spicata 293 

Cuphea carthagenensis 114, 115; petio- 
lata 84 

Cupressaceae 79, 142, 312 

Cuscuta cam pestr ris 144; compacta 144; 

pentagona 78, 


’ 


Co. be 350, 321, 322, 323, 
324, 325, 326, 327: fees 79; 
3? 


praclongatus 145; pseudovegetus 80; 
retrofractus 145; rotundus 320; strig- 
osus 80, 145 

yee eae, ee a oe calceolus 
var. pubesc s 138, 

Cyrtomium otn Dr 

Cystopteris bulbifera 141; protrusa 141 

48 


? 


Datura stramonium var. tatula 88 
Daucus carota 88, 142 
DAVENPORT, L. J. 1, 353 
DAWSON, DEBORAH 296 
Delphinium ajacis 86; 149, 216 
Dendrocyamus 90 


Dennstaediaceae 141 

Dennstaedtia punctilobula 141 

Dentaria laciniata 79, 144 

Deschampsia flexuosa 148 

Descurainea pinnata var. brachycarpa 79 

Desmanthus illinoensis 83 

Desmodium ciliare 83; cuspidatum 145; 
glutinosum 145; iinoense 83; mari 


Diarrhena americana 148 

Dicentra cucullaria 146, 214 

Digitaria bicornis 351, 352; ciliaris 351; 
diversiflora 351; ischaemum 81, 148: 


, > 


Dioscorea batatas 145; villosa 80, 145 

Dioscoreaceae 80, 145 

Diospyros 72; virginiana 74, 80, 107, 
8, 145 

Dipsacaceae 80 

Dipsacus sylvestris oi 

Dirca palustris 139, 

Disporum ee ve maculatum 
38, 147 


Distichlis spicata 107 

Dodecatheon meadia 85, 216 

Dolichos regularis 173 

Draba verna 79 

Dracocephalum virginianum 146 

Drosera annua 190 

Dryopteris goldiana 138, 141: inter- 
media 138, 141; marginalis 141 

Dulichium erundmaceuin 73s 

DUNCAN, Meee H. 116, 170 

Ebenaceae 80, 

Echinacea ne 77, 78; purpurea 143 

Echinochloa crusgalli 81; muricata 148 


s 57 

Eleocharis 73; acicu aa 80; obtusa 80, 
45; i ema a 73, 80; smallii 
80; tenuis var. verrucosa 80 

Hee hoe carolinianus 143; tomen- 
tosus 14 

Eleusine indica 148 

ELEUTERIUS, ee N. 10 

Elymus canadensis 72, 81; virginicus 73, 
81, 


Epifagus virginiana 148 

Epigaea r 

Eragrostis cilianensis 81; oxylepis 205; 
spectabilis 72, 81, 148 

Erechtites hieracifolia 143 

Erianthus alopecuroides 148 

Ericaceae 145 

Ericala sedifolia 18 

Erigenia bulbosa 142 

Erigeron 65; annuus 78, 143; bigelovii 
70; breweri 70; canadensis 78, 143; 
chiangii 65, 67, 68, 70; foliosus 70; 
var. confinis 69; var. hartwegil 69; 
hessii 70; kuschei 70; p hiladelphicus 
108; pulchellus 78, 143; strigosus 78, 
143 


Eriodictyon californicum 7 
Eriothymus 220; rubiaceus 293 


Erythronium albidum 84; americanum 
147 


Euonymus americanus 138, 144; atro- 
purpureus 144, 21 


Eupatorium album 43; aromaticum 
43; compositifolium 189, 190; hys- 
sopifolium ; incarnatum 143; 
perfoliatum ; purpureum 143; 
rothrockii 349; rotundifolium 143; 
rugosum 143; serotinum 78, , 143 
Euphorbia 317; abbreviata 317, 318; 
corollata var. zinniiflora 145, 


dentata 80; exum mensis 319; maculata 
80, 108; mercurialina 145; proctorii 
19; supina 80; vaginulata 318; 


Fagus oe 138, 146, 210, 355 
FANTZ, PAUL R. 90, 95, 152, 304 
ates elatior 148; obtusa 81, 148; 

pratensis 81 

ee autumnalis 73, 80 
Fluvialis 35; flexilis 40; minor 51 
Forestiera eee 190 

216 


Fouqui 

sone vitginiane a 86 

raxinus americana 85; var. biltmoreana 
47; arizonica 349; ie 210; 


1 49; 
cuspidata 349; pe eee 147, 
216 


363 


Froelichia gracilis 75 

FRYXELL, PAUL A. 123 

Fumariaceae 1 

Galactia 170; elliottii 170, 171; erecta 
70, 171; floridana 170, 179; var. 
microphylla 172; glabella 170, 171, 
73, 174, 175, 176, 179; macreei 
170, 178, 179; minor 170, 172, 175, 
176, 179; méssissippiensis 173; mollis 
170, 171, 172, 173, 179; pilosa 173; 


173, 175, 176, 178, 179; villosa 173: 
volubilis 145, 170, 171, 173, 177, 
178, 179; var. txtermedia 179; var. 
i aes 13 

Galium >; aparine oie 149; circaezans 


florum 150; virgatum 87 

Garrya ovata 349 

Gaura biennis 85; hexandra ssp. gracilis 
349; odorata 353 

Gentiana 14; acaulis 16; aquatica 14; 
beamanii 16, 17, 19, 21, 22, 23, 25, 
27, 28; cespitosa 17; chimboracensis 
18; edifolia var. casapaltensis 18; 
fieldiana 16, 17, 19, 23, 24, 26, 27, 
28; perpusilla 15, 16, 17, 23, 26, 28, 
a. 32; podocarpa 14: prostrata 15, 
. 2420, ey Soe puberulenta 80; 

milio 17, 23, 26, 27 6295013 

see 14; qu rae 146; 
saponaria 146; sedifolia 15, 17, 19, 
20, 21, 23, 27, 28, 32; var. B grandi- 


Geranium oe 80, 146; macu- 
latum 80, 14 

Geum ue 149; var. camporum 
86 


Gillenia stipulata 86, 149; trifoliata 
14 


Glechon 

Gleditsia tes 8, 74, 83, 108 

Glyceria striata 73, 81 

Gnaphalium helen! 143; oan 
78, 143; purpureum 78, 143 

Cancthen 298; helianthoides 299 

Goodyera pubescens 14 

GRAHAM, SHIRLEY A. 115 

Gramineae 81 


364 


Gratiola virginiana 150 
Greenella 313; arizonica 314; ramulosa 
314 


Guizotia abyssinica 296, 303 
Gutierrezia 3; alamani 313; var. 
ieeaiicenheld 313; arizonica 314, 
conoidea 313; slutinosa 314; ramu- 
losa ericocarpa 314; 
a ee 314; texana var. 
glutinosa 314, wrightii 314 


He byanthus gndeiionienas 330; var. 
texanus 330; andersonii 332; texanus 
328, 330, 331, 332; tubispathus 328, 
330, 331, 

Hackelia virginiana 76 

Halea 398: sadoviciatia 301; var. re- 

eA : 

eee 219, 312; acinoides 223, 225, 

85; 


27, 234, 282, 283, 
adscendens 290; albescentifolinm 268, 


292; bellum 230, 249, 250 


we 


costatum 223, 229, os 247, 255, 
268, 272, 276; var. costatum 268, 
269, 270; costatum var. pulchellum 
268, 269, 271; crenatum 234, 286, 
289, 290, 291, 292; dentatum 234, 
278, 279, 280, 281, 282; var. 
nanum 262, 264; denudata 293: dif- 
fusum 229, 233, 247, 261, 262, 263, 
275; drummondii 223, 233, 256, 


greggii — oe ae eat 


247, 268, 285, 286, 280: Tena 
simplex 285; hyssopifolium 223, 224, 


229, 229, 255, 247. 255. 271 OT 8, 
279, 280, 282; incana 293; italianiae 
293; johnstonii 234, 269, 272; jucun 
dum 232, 244, 250, 251, 252, 253; 
latum 258; longiflora 293; pia 


66, 
molle 228, 229, 230, 232, 242, 244, 
245, 246; montanum 228, 232, 242, 
244, 245, 247, 312; ele Herden 223, 


macrocalyx 262, 263, 266, 312; var 
mexicanum 2 var. 
62, 263, 265, var. typicum 264; 
napalensts oo nitda 293; oblongi- 
folium 234, 274 at 279, 282; var. 
mexicanum 778. 
longifolium 278, en noatiien 957; 
palmerit 225, 227, 230, 232, 241, 
ee 243, 244; patens 232, 249, 252, 
254; patrinum 228, 232, 240, 
244; permistun 268, 270; 
pilosum 228, 231 ; 238; piperi- 
tum 230, 232, re ae 250, 251, 
252, 253; var. en 278; 
plicatum 229, 234, 269, 272, 273; 
subgenus Poliominthoides 231, 240; 
polygalaefolium 231, 248, 249, 291; 

montanum 290; pringeli 268, 
270; pulchellum 268, 271; niilehenr 
mum. 225,229) 233,247,299, 25), 
256, 258; pulegioides 219, 223, 230, 
231, 234, 283, 286, 288, 289; B- 
mexicanum 293; forma simplex 288; 
purpurea 293; pusillum 229, 231, 
236, 237, 238; quercetorum 234, 269, 


293; serpyllifolium 259; serpylloides 
293, stenodonta 293; subequale 294; 
tenellum 268, 270; tenuiflorum 234, 


273, 274, 275; tenuipes 234, 269, 
276; texana 294; thymoides 219, 264, 
294; var. oblongifolium 278; todsenii 
227, 228, 232, 238, 239; villosa 294; 


var. lavanduloides 294 


; glomeratus 

161, 162, i 165, 168, ee sativus 
160, 161; souda 160, 161, 

Haloragidaceae 82, 1 

Hamamelidaceae 1 

Hamamelis virginiana 146 

Hedyotis crassifolia 87; longifolia 87; 
nigricans 


Hedysarum volubile 173, 179 


hirsutus 78, 143. microcephalus 143. 
mollis 78 
Heliopsis helianthoides 78, 143 


Hesperaloe funifera 314; parviflora 314 

Hesperozygis 220; bella 249; ciliolata 
35; marifolia 223, 293; myrtoides 
293; nitda 293; pusilla 236 

Heterotheca graminifolia 143; mariana 


Heuchera americana 150; parviflora 
150; richardsonii 87; rubescenes 349; 
villosa 150; var. macrorhiza 21 

Hexalectris spicata 1 

Hexastylis arifolia 138, 142 

Hibiscus lasiocarpos 74, 84; syriacus 
14 


Hickoria olivaeformis 8; pecan 8 
3 
Hieracium gronovii 143; paniculatum 
4 


Hippocastanaceae 146 


(C328 

Holosteum umbellatum 355, 356, 357 
Holubia 14; pyrenaica 14 

Holubia 14; pyrenaica 14 
Heleboueniie 14 
HOPKINS, CHRISTINE O. 157 
Hordeum vulgare 
Houstonia caerulea 150; purpurea 150 
Hyas 35 


365 


Hybanthus concolor 150 


Hypericum denticulatum var. recog- 
i 46; drummondii 82; gentia- 
noides 82, 146; hypericoides 107, 
108; multilum 82, 146; fardtdocain 
146; ee 82; prolificum 146; 
punctatum 82; stans 146; stragalum 
146 


Hypoxis hirsuta 75, 142 
Hyptis mutabilis 202, 203, 204, 205, 
206, 207, 208, 209; spicata 203, 209 
A 


toria 107, 
Impatiens cece 76, 143; pallida 143 
Ipomoea hederacea 79; lacunosa 79; 
pandurata 79; purpurea 79, 144: 
sagittata 108 


6 
VING, ROBERT S&S. 219, 313 
a brachiatus 82 
Isotria verticillata 137, 147 
Itea virginica 137, 
Ittnera 35; major 37; minor 513 najas 


3 
Iva annua 78; frutescens 107, 108 
Juglandaceae 82 
Juglans cinerea 146: nigra 82, 146 
Juncaceae 82, 146, 334 
Juncus 73; acuminatus 74, 82; brachy- 


337, 338, 339, 340, 341, 342, 343; 
saab 338; interior 74, 82; margina- 
s 146; nodatus 82; repens 137, 146; 

roemerianus 107, 335, 344; tenuis 
74, 82, 146, 347 

Juniperus ashei 312; silicola 102, 107, 
108, 111s virginiana 74.79, 

Kalmia latifolia 137, 145 


366 


KEENER, ‘aa S. 114, 216 

Keithta 22 

Kochia Sais 214 

Koeleria pyramidata 72 

Krigia biflora 78, 143; oppositifolia 78 
Kuhnia eupetotioides 7§ 


Lactuca Rene 78, 143; floridana 
143; serriola 78 

famincere 146, 
jum amplexicaule 83; purpureum 


Laportea canadensis 150 
Larrea 216 

Lauraceae 83, 

LAWSON, CHERYL A. 1 

Lechea racemulosa 144; tenuiiela 77 
1; virginica 148 
304 


tH 
oO 
we 
“eR 
a 
5 
fs) 
n 
D 
= 
0 


ee campestre 7 79; virginicum 79, 


Leptochioa filiformis 81 
bicolor 145; capitata 83; 
cuneata 83, 145; hirta 145; & inter- 
media 145; intermedia 145; proc 
bens 83; repens 146; striata 
stipulacea 83 steuvei 146; wiolacea 
ica 8 


3; vir 
Leucophyllum frutescens 348 
Liatris aspera 78, 143; microcephala 
143; peal 78; spicata 143 
Ligusticum canaden 42 


Lilium michiganense 137, 147 

Linaceae 84, 147 

Lindera benzoin 147 

Lindernia dubia 87, 150 

Linum elongatum 181, 182, 183, 185, 
186; lundellii 183, 184, 185, 186; 


pactum 181; filifolium 181, 182, 183, 
185, 186; striatum 147; subteres 185; 
sulcatum 84; virginianum var. fori: 
danum 147 

LIPSCOMB, BARNEY L. 118, 320 

Liquidambar  styraciflua 8, 137, 139, 
146, 355 

Liriodendron tulipifera 138, 147 

Lithospermum arvense 76; canescens 76 
tuberosum 144 

Loasaceae 84 


Lobelia cardinalis 144; inflata 144; 
puberula 144; spicata 76 
ee 147 
Lonicera japonica 144; sempervirens -44 
81 


Lolium multiflorum 148; perenne 
Loranthaceae 14 

LOTT, E J. 348 

Ludwigia paige 85, 147; palustris 


var. americana 8 
Luzula acuminata 146; bulbosa 82, 146 
Lycopodiaceae 141 
Lycopodium lucidulum 141; obscurum 

141; porophilum 141;  tristachyum 

1 
Lycopus rubellus 83; oo 83, 146 
Lygodium palmatum 


Lysimachia hybrida 86; lanceolata 86, 
149; quadrifolia 149; tonsa 149 
Lythraccae 84 
Lythrum carthagenense 114 
ie cae acuminata 138, 147; grandi- 
flora 210; tripetala 147 
tees 147 
, WM. F. 211, 212 
Saeed pean 138 
Malaxis unifolia 138, 147 
Malvaceae 84, 147 
Manisuris rugosa 10 
MANSFIELD ee A. 196 
MARIETTA KAY L. 355 
Marshallia cinetwie 139, 143 
Matelea carolinensis 108, 142 
McDANIEL ee 210 
McINTOSH, K. 
Medeola virginiana oie 
Medicago lupuli ina — sativa 83 
Melanthium virgin 4 
Melastomataccac M7 
Melica mutica 1 
Melilotus alba on officinalis 83 


Menzclia oligosperma 
se ogee 203; spicatum 203 
Micromeria 222; bonariensis 260 
Nicest vimineum 148 
imulus alatus 150; ringens 87 
Mirabilis nyctaginea 84 
Mitchella repens 139, 150 
Mitella diphylla 138, 150 
Mitracarpus hirtus 117, 118; véllosus 
117, 118 


Mollugo verticillata 75 


Monarda clinopodia 146; fistulosa 83 
Monotropa hypopithys 145; uniflora 
145 


Monotropsis odorata 138, 145 

Moraceae 84, 147 

MORGAN, DAVID 210 

Mortonia scabrella 248 

Morus alba 84; rubra 107, 108, 111, 
147 


Mosla 220; ocymoides 293; dianthera 
146 


Muhlenbergia capillaris 190, 191; schre- 
beri 81, 148; sylvatica 

MUSSLEMAN, LYTTON J. 196 

Myosotis virginica 76 

Myrica cerifera 107, 108 

Najadaceae 34 

Najas 34; alagensis 45; ancistrocarpa 


40; dichotoma 51; flexilis 37, 40, 41, 
42, 43; ssp. os O, 41; var. 
congesta 40, var. curassavica 43; 
var. lees a var. guadalupensis 
var. busta 4 ly 

poe 37; gracillima 47, 49, 50, 54: 
graminea 37, 43, 45, 47, 48, 52; 
guadalupensis 41, 43; var. floridana 
7, 45, 46, 52; var. guadalupensis 44, 
52; var. muenscheri 37, 45, 46, 52; 
var. Olivacea 37, 43, 46 
51; var. gracillima 47; latifolia 37; 
latior 37; major 28; var. angustifolia 


a 

Le 38; marina 35, 36, 39, 
52 353; var. angustifolia 37; var. 
as ma 37; var. bollei 37; var. 


var. genwina 37; var. gracilis 37; var. 
latifolia 38; var. latior 38; var. mexi- 
cana 38; var. multidentata 38; var. 
muricata 38; var. paucidentata 38; 
var. recurvata 38; microdon var. 
curassavica 43; as guadalupensis 43; 
, 54, 56; var. setacea 
51; muewschen a 4); piuusesied 


nuda 495; 51; SH ulate 51; 


multidentata 54. 


307 


Nelumbo lutea 73, 84 

Nepetha multabilis 202 

NELSON, GUY L. 65 

Neviusia alabamensis 215 

NICKRENT, DANIEL L. 196 

Nicolletia occidentalis 216; trifida 216 

NIXON, ELRAY 355 

Nothoscordum bivalve 84 

Nyctaginaceae 84; Nymphaea 34, 56, 
84 


Nyssaceae 85, 147 

Nyssa sylvatica 74, 85, 137, 147 

Oenothera 295; biennis 85, 147; 
laciniata 85; tetragona 147 

OGBORN, JAMES E. A. 196 

Oleaceae 85, 147 

OLSEN, JOHN 128 

Onagraceae 85, 147, 353 

Ophioglossaceae 85, 141 

Ophioglossum 113;  crotalophoroides 
113; nudicaule var. tenerum oe 
petiolatum 113; vulgatum var. pyc- 
nostichum 113 

Opuntia compressa 109; polyacantha 
var. ae ie a stricta 109 

Orchidaceae 85, 

Orchis specail es 147 

Orobanche 

een 148 

Osmorhiza claytonii 142 

Osmundaceae 141 

Osmunda cinnamomea 137, 141; regalis 
137, 14 

Ostrya virginiana 144 

OTVOS, ERVIN C., Jr. 102 

Oxalidaceae 85, 148 

Oxalis dillenii 85, 109, 148; stricta 


85, 148 

Oxydendrum ee 137, 145 

Oxypolis rigidior 

Pachysandra eaten te se 

Panax quinquefolium 140, 

Panicum amarum 109; 2 an 148; 
boscii 148; brachyanthum 190; capil- 
lare 81; clandestinum 81, 148; 
dichotomiflorum 81; dichotomum 


12 Ol: philadelphicum 148; poly- 
anthes 148;  rigidulum 7 81; 
sphaerocarpon 148; virgatum 73, 81, 
109 


368 


Papaveraceae 148 
Parnassia asarifolia 137, 150 
Parthenium integrifolium 143; hispid- 


um 7 
Parthenocissus quinquefolia 89, 138, 
150 


Paspalum 205; laeve 81, 148;  pubi- 
flo var. glabrum 81; setaceum 


ar. 

Passifloraceae 85, 

Passiflora lutea ne. 148; var. glabri- 
flora 85 

Pastinaca sativa 


PATRICK, [ous 5:13) 


7, ee 
Pellaea atropurpurea 141, 214, 
Penstemon canescens 150: eee 87; 
tubaeflorus 87 
Pennisetum typhoides 196 


85; hydropiper- 

: lapathifolia 85; pensylvanica 
86; punctata 

Petalostemum candidum 83; purpureum 
83 


Peteria scoparia 349 
Phacelia bipinnatifida 138, 146 
Phaethusa oe 13° 


PHILLIPPE, LOY R. 135 

Phleum pratense 82 

Phlox amoena 149; carolina 149; eee 
cata 138, 149; glaberrima 139, 
maculata eee ssp. pyramidalis i 
pilosa 85, 139 

Phoradendron serotinum 147 

Phragmites communis 109 

Phryma Cee 148 

Phrymaceae 148 

Phyla oe 88 

Phyllanthus polygonoides 315 

Physalis angustifolia er heterophylla 
88, 150; subglabrata 8 

Physocarpus opulifolius 2 


Phytolacca americana 85; rigida 109 
Phytolaccaceae 85 
Pilea pumila 88, 150 


Pinaceae 142 

inus Here 137, 139, 142; glabra 
8, taeda 142: virgiana 138, 139, 

Plantaginaceae 85, 148 

an fae aristata 85, 148; lanceolata 89; 
148; pusilla ae rugelii 85, 
ee 85, 

Platanaceae vas 

Platanthera ciliaris 137, 147; clavellata 

48; yee as 48; integ- 

ale 137, 139, 

Platanus Paci , 148 

Sone andens 

Poa ua 

ae 148 

Poa compressa 148; languida 139, 1 
pratensis 82; 148 

Podophyllum peltatum 76, 144 

Pogogyne 331; serpylloides 293 

Polanisia dodecandra 76 

Polemoniaceae 85, 14 

Poliomintha 220; glabrescens 236, 293; 
incana 293; longiflora 223, 236, 292, 


as 
- 


Polygala curtissii 149; incarnata 85, 
149; minutifolia 349; nudata 349; 
sanguinea 85; senega 85, 149; verti- 
49 


Polygonaceae 85, 149 

Polygonatum biotin 84, 147; pubes- 
cens 138, 

Polygonum aviculare 86; cespitosum 
var. longisetum 149; persicaria 149; 
punctatum 149; scandens 74, 86, 
149; virginianum 8 

Polymnia canadensis 143; uvedalia 143 

Polypodiaceae 86, 141 

eeu lone 141; thyssano- 

lepis; virginianum 141 

Polystichum scrontichides 141 

POOLE, JA M. 315 

Polytaenia ae 88 

Portulacaceae 86, 149 

Potamogeton 35, 55; diversifolius 73, 
86 


Potamogetonaceae = 

Potentilla canaden 149; aa 
149; recta 86. ae 86, 
Prenanthes altissima 143, aspera 78; 


149 
PRINGLE, JAMES S. 14, 119, 356 


Prosperpinaca palustris 82; pectinata 


> 


Prunella vulgarus 146; var. lanceolata 
83 


Prunus americana 149; hortulana 87; 
maheleb 87; munsoniana 87; persica 
74, 87; serotina 74, 87, 149, 349 

Psendocunila montana 290 

Psoralea psoralioides 146; var. eglandu- 
losa 84 

Preridium aquilinum 86, 141 

Pycnanthemum incanum 147; muticum 
147; tenuifolium 83; linifolium 7 

Pyrolirion 333 

Pyrrhopappus carolinianus 78, 143 

Pyrularia pubera 1 

Pyrus angustifolia 149; arbutifolia 149; 
ioensis 87; malus 74, 87; melano- 


sae ae 74, 80, 138, 139, 146; 
coccinea 146; daraddii 7; falcata 139, 


188, 190, 191; velutina 74, 80, 146; 
virginiana 108 

Ranunculaceae 86, 149 

Ranunculus 35, 55; abortivus 86, 149; 
fascicularis 114; var. apricus 86; 
laxicaulis 86; ete 86; recurva- 
tus 149; trilobus 114 

Ratibida pinnata 78 

Rhabdocaulon 22 coccineus 293, 
denudatus 293; eraalis 293; iavond 
loides 294; wae 2 

Rhamnaceae 86, 149 

Rhamnus caroliniana 149 

Rhexia mariana 147; virginica 147 

Rhododendron alabamense 145; arbore- 
scens 139, 145; calendulaceum 145; 
cumberlandense 145; maximum 145; 
nudiflorum 137, 145 

Rhododon 220; ciliatus 221, 223, 292, 
293, 294 
Rhus 72; aromatica 74, 76; e 
74, 76, 107, 108, 142: cotinoides 7; 
glabra 74, ae 142; radicans 74, 76, 
109, 138, 

Breck oe ee 145; cornicu- 
lata 137, 145 

Ribes missouriense 74, 87 


369 


Robinia 146; pseudo-acacia 146 
ROGERS, C. M. 181 

ROGERS, KEN 215, 216 

Rosa carolina 87, 149; multiflora 87, 


; betulifolius 149; 
“igs 211; flagellaris 87; hispidus 
149; occidental: 87, 149; pennsyl- 
vanicus 74, 87 phoenicolasius 149; 
trivialis var. duplaris 211, 212; var. 
trivialis 211 

Rudbeckia fulgida var. umbrosa 143; 
hirta 78, 143; laciniata 143; triloba 
78 


Ruellia caroliniensis 142; humilis 75 

Rumex acetosella 86, 149; crispus 86, 
149 

Rumfordia 296, 297; aaa 303 

Sabal palmetto 107, 

Sabatia angularis 80; fide 146 

Sageretia minutiflora 102, 107, 108, 
109 


Sagittaria cuneata 119; “aie rama 
142; lat hope yo Fee OF aet ie ola 

Salicaceae 87, 

Salix eine HE at humilis 87, 
150; nigra 73, 

Salsola kali 159; ie on ae 161 

Salvia lyrata 147; anientoles 14 

Sambucus canadensis 74, 76, un 108, 
144; pubens 144 

Samolus parviflorus 109, 210 

Sanguinaria canadensis 148 

Sanicula canadensis 88, 142; gregaria 
142; smallii 142; trifoliata 142 

Santalaceae 87, 150 

Saponaria officinalis 77 

Sapotaceae 15 

Sassafras 72; albidum 74, 83, 147 

Satureja 222; bonariensis 260; brownei 
293; caerulescens 292; georgiana 205; 
glabra 292, 293; glabrella 293; 
gillie sii 260; _micromoroides 29 

0 


Schizachyrium scoparium 72, 82; var. 
0 


Schoenolirion wrightii 354, 355 
Schrankia microphylla 146; uncinata 84 
Scirpus 73; americanus 325; atrovirens 


a7 


80; var. georgianus 145; californicus 
117; cyperinus 145; pendulus 80; 
validus var. creber 80 

Scleria oligantha 145; triglomerata 145 

Scrophularia marilandica 87 

Scrophulariaceae 87, 180, 196 

Scutellaria elliptica 147; lateriflora 83; 
ovata 83, 109; var. versicolor 215; 
parvula 83; —— rrata 147 

Secale cereale 8 

Sedum ternatum 1 138, 144; eas 349 

Selaginella apoda 142; rupincola 349; 
ncinata 214 

Selaginellaceae 142 

Senecio aureus 143; obovatus 143; 
smallii 143 

Serenoa repens 108 

Setaria faberii 82, 149; geniculata 149; 
slauca 82; viridis 82 


Sida anomala 123; elliotit 123, 124, 


124, be 12 otosina 124 
rzedowskii 123, 
eee 84; texana 
io turner 125 
Siegesbeckia laciniat 132 
Silene antirrhi 77; laciniata 349; 
rotundifolia ivr stellata 77, 144; 
virginica 77, 144 
ia integrifolium 78; trifoliatum 
43 


eee rad 

Simmondsia 2 

Siphonoglossa longiflora 350; pilosella 
0 


Sisymbrium officinale 

Sisyrinchium ees 146; angusti- 
folium 82, 146, 210; campestre 82 

Smilacina racemosa 84, 147 

Smilax bona-nox 83, 108, 109, 147; 
var. hastata 190; ecirrhata var. hugeri 
147; glauca 147; hispida 84, 147; 
rotundifolia 2 

Solanaceae 88, 

Solanum americanum 88; carolinense 


ree arguta 143; Res var. boottit 

A; var. eumees 14 caesia a 
canadensis 78; 
scabra 143; curtisit 143: erecta "143. 


flexicaulis 143; gigantea 143; gr 
minifolia var. nuttallii 143; hispida 


laris 78 riddellii 9; rigida 78; 

roanensis 143; r semper- 

virens 109; sphacelata 216; “ulmifolia 
8, 143 


Sorter avenaceum 72, 82; nutans 
49 


ae bicolor 82, 196; halepense 82 


Spartina alterniflora cynosuroides 
109; patens 345; pectinata 73, 82; 
spartinae 109 


Specularia perfol | ie 144 

Spermacoce glabra 

Sphenopholis ere ue 

Spigelia semi 147 

Spiraea japonica 

Spiranthes cernua 35, 193; gracilis 
85; aoe 188; ae Bory 189, 190, 
191, 

era junceus 189, 190; vagini- 
florus 82, 149 

Stachydeoma 220; graveolens 221, 223, 
2¢ 


dy fy 
Xo 


Stellaria media 77, 109; pubera 144 
Stewartia ovata 150 
Stipa avenacea 149; tenuissima 349 
Streptopus amplexifolius 13 

a angu ale 196, 201; 
asiatica 198, 199, 201; aspera 196, 
197, 198, 199; euphasioides 196; 
gesnerioides 196, 199, 200, 201; 
hermonthica 196, ot 198, 199, 
200; klingii 199, 200, 201; /utea 
196; macrantha Aas 200, 201 


Strophostyles helvola 8 


’ 


Serlophiorum diphyllum 138 148 
Stylosanthes biflora 84, 146 
Styracaceae 213 
Styrax americana 213 
Suaeda hyssopifolta 58 
wertia caroliniensis 146 
Symphoricarpos 349; ee 76 
Tacnidia integerrima 14 15 
Talinum parviflorum ve rugosper- 
mum 354, 35 
Taraxacum officinale 78, 14° 
Tephrosia pedicellata 199; 
146 


84, 


virginiana 


Tetragonosperma 2 5 ie 301 
si oa ee abyssinica 303; 
temalensis 303; Pel nietes 296, 

oe. 298, 299, 300; ludoviciana 296, 
297, 299, 301, 302, 303; parviflora 
303; repanda 296, 297, 299, 302, 
303; texana 296, 297, 299, 302, 303 


149; dasycarpum 
114; debile 114; pubescens 149; revo- 
lutum 86; thalictroides 149 

Thaspium barbinode 142 

Theaceae 150 
helypteris yeaa 141; nove- 
boracensis 1 

THIERET, Sonne 

ile arvense 79; pestliscm 79 

THOMAS, R. DALE 113 

TOE On RALPH L. 70 

Thymelaecae 150 

Thymus serpyllum 294 

Tiarella cordifolia 150 

Tilia eee 138, 156 

Tiliac 

Tillandsi sia usneoi oy oe 

Tipularia disc olor 


9 
Tradescantia ernestiana 215; ohiensis 
77; subaspera 144 
Tragia ln tn 80 
Tragopogon dubiu 
‘Trautvetteria i ae 215 
Trichachne 352 
richomanes petersii 139, 141 


Trillium cuneatum ia. erectum 138, 
147; grandiflorum 147; oo 209, 
210; recurvatum 138, 

Triodanis biflora 76, 349: ae - 
riosteum angustifolium 77;  perfoli 


tu 
Triphora trianthophora 148 
Tripsacum dactyloides "53, 82 
Triticum aestivum 82 
Tsuga eee 137, 138, 142 
TURNER 296 
Typha latifolia 73, 88, 150 
Typhaceae 88, 150 
Ulmaceae 88, 150 
Ulmus alata 88, 150; americana 88, 


371 


0, 210; Lee 74, 88, 150 


Uvularia grandiflora 147; perfoliata 


aboreum 145; constablaei 
145; corymbosum 145; stamineum 


Wate nace eae 88 
Valerianella radiata 88 


, 14 
Verbascum blattaria F 150; thapsus 


Me Wenacese 88, 150 
erbena bracteata 88; canadensis 88; 
simplex 88, 150; stricta 88; urticifolia 


50 
Verbesina alternifolia 143; helian- 
thoides 78; laciniata 130; microptera 
128... 130, Tol ae loa: 
Mol Gine 133; occidentalis 109; 
polycephala 132; rumicifolia 134; 


128, 129, 130, 132, 
laris 133; var. laciniata 128, 

132; var. palmeri 134; virginica 128, 
131 


Se altiss 143;  arkansana 
; baldwintt 78: missurica 78 
ere arvensis 87; peregrina 87 
Veronicastrum virginicum 33 
Viburnum acerifolium 137, 144; cassi- 
noides 144; dentatum a nudum 
137, 144; rufidulum 74, , 14 
Vicia caroliniana 146; eee 84; 


Viola affinis 150; blanda 138, 150; 
canadensis 150; conspersa 138, 150; 
hastata 150; hirsutula 150; missouri- 
ensis 88; oa 88, 150; pensyl- 
vanica 150, 15 
primulifolia i rafinesquii 88; 
rostrata 150; sagittata 88; triloba 
1 


50 

Violaceae 88, 150 

Vitaceae 89 

Vitis aestivalis 89, 150; cinerea 89, 150; 
aay lia 150; vulpina 8 
ttaria 142 

oe 142 


ae 


Vulpia octoflora 82 

Wahlenbergia linarioides 354; margin- 
ata 

WEBB, DAVID H. 135 

WEBSTER, ROBERT D. 351 

WELLS, CHRISTOPHER J. 328 

Wisteria frutescens 108, 109 

WOFFORD, B. EUGENE 135 

Woodsia obtusa 216 

Woodwardia areolata 137, 141; virgin- 
ica 137, 138, 139, 141 
Xanthium strumarium 

Xanthocephalum 313; conoidenm 313; 
lineartfolium 313; megalocephalum 
313; sericocarpum 314; wrightii 313 


Barney Lipscomb and Jessica Procter 


Herbarium 
Southern Methodist University 
Dallas, Texas 75275 


Xanthorhiza simplicissima 139, 149 
Xyridaceae 150 

Xyris torta 137, 150 

a i 107, 108 

Zam 

ee clava-herculis 107, 108 


Zephyranthes 329; andersonit 330; 
commersoniana 330; SNE ER 
330; texana 229, 330 

Zigadenus leimanthoides 137, 139, 147 

Zizia aptera 142; aurea 

Ziziphora 220; hispidum 285; pule- 
gioides 288