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Mee PHYTOLOGIA 


Vol. 64 December 1987 No. 2 


CONTENTS 


__- ORZELL, S.L., & BRIDGES, E.L., Further additions and 
noteworthy collections in the flora of Arkansas, with 


historical, ecological and phytogeographical notes ........... 81 
TURNER, B.L., Taxonomy of Carphochaete (Asteraceae - 

Bo ES SE OS EE eS SSS EA Ole Cee ra i re ah a 145 
LOURTEIG, A., & SODERSTROM, T., Nomenclatura 

Plantarum Americanarum VI. Gramineae ...............+-- 163 


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FURTHER ADDITIONS AND NOTEWORTHY COLLECTIONS IN 
THE FLORA OF ARKANSAS, WITH HISTORICAL, 
ECOLOGICAL, AND PHYTOGEOGRAPHICAL NOTES 


Steve L. Orzell & Edwin L. Bridges 
The University of Texas Herbarium 
Austin, TX 78713 


ABSTRACT 


Sixty-four native or possibly native vascular plant taxa are 
reported as new to or reinstated in the flora of Arkansas, through 
integrated fieldwork, herbarium study, and literature surveys. 
Thirty of these taxa were apparently first collected in Arkansas 
by the authors from 1984 to 1986, seventeen were collected by the 
authors and also represented by previously misidentified or 
unreported material in herbaria, eight were collected by colleagues 
or uncovered in herbaria and determined or verified by the 
authors, and nine had previously been collected or known to occur 
in the state but were synonymized or overlooked in Smith (1978), 
the current authority on the Arkansas flora. A total of twenty- 
six taxa are reported here for more than one Arkansas county, 
indicating a broader distribution in the state than is usual for 
state record species. Noteworthy collections are presented 
extending the range of six plants which are extremely rare or 
narrowly endemic in Arkansas. Two species reported here have 
long been considered possibly extirpated from Arkansas. Six taxa 
are deleted from the flora of the state, as previous records were 
found to be misidentified. Brief ecological notes and a history of 
Arkansas reports are presented for each species. The 
phytogeographical discussion focuses on the distribution patterns 
relative to Arkansas expressed by these species, and their 
significance in the Arkansas flora. 


INTRODUCTION 


The Arkansas flora, after over 150 years of collecting, 
remains poorly known and little understood. The earliest Arkansas 
plant checklist (Nuttall 1835, 1836) was compiled when the state of 
Oklahoma was part of the Arkansas territory, thereby including 
many species clearly collected in present-day Oklahoma. Later 
checklists (Lesquereux 1860, Branner and Coville 1891, Demaree 
1943) generally accepted all previous reports as part of the flora, 
without examination of the identity and locality of the collections, 
thereby perpetuating numerous errors. Such misinterpretation of 
early "Arkansas" collections still occurs, as exemplified by Benson 
(1982), who maps Coryphantha missouriensis (Sweet) Britt. & Rose 
var. caespitosa (Engelm.) L. Benson for Miller County, Arkansas 


81 


82 PAH ¥Y T.0.LO:1¢G 1 A Vol. 64, No. 2 


based only on a collection by Pitcher in 1830, from "Red River, 
Arkansas," a common locality description for collections from 
southern Oklahoma during this period. A good account of some of 
Nuttall’s travels and collections in the part of "Arkansas" that is 
now Oklahoma is given by Geiser (1956). Numerous additional 
examples of these early "Arkansas" records were uncovered and 
excluded from the Arkansas flora by Smith (1978). In the history 
of Arkansas botany, the ultimate unworkable and untenable 
floristic list for the state is that by Demaree (1943), which 
includes every imaginable type of error in compilation and 
judgment. Based upon our efforts in uncovering the sources of 
his reports, this list was based on numerous misidentifications and 
includes plants collected in other states by Demaree, but not in 
Arkansas. In addition, the sources listed for the first state report 
of each species are not reliable. Demaree was certainly the most 
prolific general collector to work in Arkansas, and he provided 
much needed material of the common flora of the state. However, 
he rarely collected in less accessible areas or in specialized 
habitats, and therefore missed much of the floristic diversity of 
the state. The first carefully prepared list of Arkansas plants, 
citing voucher specimens, locations, and sources of reports is 
Buchholz and Palmer (1926). It was not, however, intended to be 
complete, but rather includes only those plants not listed by 
Branner and Coville (1891). The major, and current standard, 
floristic work for Arkansas is Smith’s (1978) An Atlas and 
Annotated List of the Vascular Plants of Arkansas. This volume is 
based primarily on vouchered material at UARK, and part of, but 
not necessarily all, Arkansas material at APCR, HDX, HSU, LTU, 
MO, NLU, SMS, SMU, and STAR. Species previously reported but 
not vouchered by material seen by Smith at these herbaria were 
usually relegated to a list of "Possible Additions” or "Excluded 
Names", or may appear in the text along with the source of the 
report. Due to the inaccuracies and discrepancies of earlier lists, 
we accept Smith (1978) as the sole authority on the flora of 
Arkansas up to the date of compilation. Smith’s Atlas was a 
valiant effort, and provided a much needed "starting point" for the 
floristic exploration of the state. Since 1978, five supplements to 
the Atlas (Smith 1979, 1980, 1981, 1982a, 1986) have been issued, 
each adding numerous new state and county records, revised 
nomenclature, and further documentation based upon new records 
at UARK and in the literature. In addition, numerous authors 
have reported new plant records for Arkansas since 1978 (Davis 
1981; Lipscomb 1980; Orzell and Peck 1985; Orzell et al. 1985; 

Peck et al. 1985; Rettig 1982; Richards 1982, 1985; Smith 1983; 
Sundell 1983, 1986; Taylor and Taylor 1981; Taylor 1984; Taylor 
and Johnson 1979; Thomas and Hooks 1985; Thomas et al. 1985; 
Werth and Taylor 1980). If a taxon does not appear in Smith 
(1978), Supplements I-V, (1979-1986) or in the above cited works, 
we are considering it as "unreported" for Arkansas, even though 
some of these taxa are listed in earlier checklists of the state’s 
flora. Some records reported in Smith’s most recent Atlas 


1987 Orzell & Bridges, Flora of Arkansas 83 


supplement (1986), issued while this manuscript was in preparation, 
and based upon our work for this paper, are included in order to 
more fully document their occurrence in the state. 

In addition, many of the species included in this paper were 
synonymized in Smith (1978). Our decision to recognize these 
taxa is based upon the opinions of experts in the genera involved. 
This situation is particularly true for the genus Carex, for which 
we are relying on the generous advice of Dr. Anton A. Reznicek 
at The University of Michigan. 

Despite the fact that many plant specimens have been 
collected in Arkansas, only the most abundant, non-habitat 
specific, and ruderal species are well-documented in the flora. 
Remote areas and specialized habitats generally have not been 
explored by botanists. Some notable exceptions have been the 
Arkansas collections of Ernest Palmer, Hugh IItis, and Paul 
Redfearn, and the recent work of Gary Tucker, Eric Sundell, and 
R. Dale Thomas. Since many areas of Arkansas are essentially 
unknown to botanists, collections from the state take on added 
significance. Many of the records reported here are not 
unexpected, and several are apparently fairly common in parts of 
the state. 

In considering the reasons for such floristic backwardness, 
we turn to the work of Fernald (1937), in one of his classic 
papers on countless new discoveries from the Coastal Plain of 
Virginia. In describing the occurrence of a fairly common species, 
previously collected several times but not recognized as occurring 
in the state by recent authors he states, 

"The fact that after such clear demonstrations of its 

abundance at the inner margin of the Coastal Plain in 

Virginia, the finding there of Silphium compositum 

should still be looked on as a new discovery speaks 

eloquently of the need for a trained taxonomist in the 

region (and by trained I mean one so familiar with 

vascular plants that he recognizes and promptly 

evaluates the insignificant and highly significant species 

when he sees them).” 

While we have by no means achieved this level of competence in 
Arkansas, it has been a goal of our work. Systematic collection 
efforts by highly trained botanists are essentially nonexistent in 
the state. It is only after such collections have been made, and 
detailed information compiled on the sites and habitats searched, 
that the Arkansas flora can be understood at the level taken for 
granted in most eastern and northern states. 

The large majority of the new records in this paper are 
based upon only 3000 collections made by the authors in 1985 and 
early 1986, with a deliberate eye towards significant records and 
taxonomically difficult groups. The vast majority of these 
collections were county records for Arkansas, with a large number 
being significant new records for rare species or range extensions 
within the state. Criteria for inclusion in this paper were that 
the taxon: 1) not be attributed to Arkansas as a distinct entity 


84 PUN YT. 0 EL OyG) TA Vol. 64, No. 2 


in Smith (1978), Supplements I-V (1979-1986), or previously cited 
reports of additions to the flora, or 2) be a significant range 
extension for a species which is quite rare or had been considered 
possibly extirpated from the state. A forthcoming paper will 
report many other significant records for the state not included 
here. 

After identifying our specimens as representing species new 
to Arkansas, we searched for additional collections of these 
species, which may have been previously misidentified or 
overlooked, at SMU, UARK, and APCR, and in the taxonomic 
literature. Most such collections were determined or verified by 
the authors; however, a few citations are included based on 
specimens we have not seen. In addition, we compiled county 
range maps for each species, based on the taxonomic literature, 
published floras, rare species publications, our collections from 
Arkansas and other states, and examination of specimens at APCR, 
ASTC, DUR, NLU, SMU, TEX, UARK, and VDB. These county 
range maps were used in analysis of the distribution patterns of 
each species, and to determine the significance of our Arkansas 
collections relative to the total range and nearest approach of the 
species to the state. A few species included are recorded for 
Arkansas in recent monographic works not yet assimilated into 
compilations of the Arkansas flora. The nomenclature used in this 
paper generally follows Kartesz and Kartesz (1980, 1985), differing 
primarily in our use of several recent nomenclatural changes found 
in works by Farmer and Bell (in prep.), Semple and Brouillet 
(1980), Thomas (1984) and Warnock (1981). 

The format of the species accounts includes abbreviated 
locality information in the form of standard legal locations to the 
section level (full documentation of localities and habitats are 
available on specimen labels and from the authors), description of 
habitats in Arkansas, and a history of previous knowledge of the 
species in the state. The phytogeographical discussion section 
treats the species distribution patterns, overall habitats, and 
significance of the Arkansas records, in groups of similar patterns. 


SPECIES ACCOUNTS 


ASCLEPIAS LONGIFOLIA Michx. ssp. LONGIFOLIA 
(Asclepiadaceae). Calhoun Co.: Sec 22, T14S, R13W, 30 Jun 1985, 
Orzell & Carl Amason 2293 (UARK); Sec 23, T14S, R13W, 6 Jul 
1985, Orzell and Bridges 2377 (SMU, UARK). 


Asclepias longifolia ssp. longifolia was found in a seasonally 
saturated open graminoid dominated area with Fuirena bushii Kral, 
Polygala cruciata L., Xyris ambigua Beyr. ex Kunth, X. torta Sm., 
Rhynchospora glomerata (L.) Vahl., R. inexpansa (Michx.) Vahl, R. 
globularis (Chapman) Small, Rhexia mariana L., Gelsemium 
sempervirens (L.) St. Hil., Sabatia gentianoides Ell., Gratiola pilosa 
Michx., Paspalum praecox Walt., Dichanthelium scoparium (Lam.) 
Gould., Ludwigia linearis Walt., Hypericum hypericoides (L.) Crantz, 


1987 Orzell & Bridges, Flora of Arkansas 85 


and Eupatorium rotundifolium L. 

Branner and Coville (1891) list Acerates longifolia Ell. for 
Arkansas, previous to the description of Acerates hirtella Pennell, 
to which this report can undoubtedly be referred. Small (1933) 
includes Arkansas in the range of A. floridana (Lam.) A. Hitche.. 
perhaps on the basis of the above report. Fernald (1950), 
Woodson (1954), Correll and Johnston (1970), and Godfrey and 
Wooten (1981) do not include Arkansas within the range of 
Asclepias longifolia, and Smith (1978) lists only Asclepias hirtella 
(Pennell) Woodson for Arkansas. Recent work for the Flora of 
the Southeastern United States indicated this taxon should be 
treated as Asclepias longifolia Michx. ssp. hirtella (Pennell) Farmer 
and Bell. Our specimens fit the characters used by Woodson 
(1954) in distinguishing A. longifolia from A. hirtella; however, our 
study of some specimens annotated by Farmer and Bell indicate 
that they may have a narrower concept of this subspecies which 
excludes many West Gulf Coastal Plain collections. Apparently, 
our collections are the first for ssp. longifolia in Arkansas. 


ASTER SHORTII LindL (Asteraceae). Montgomery Co.: Sec 
8, T4S, R27W, 8 Oct 1984, Orzel]l and Bridges 1495 (UARK). 
Newton Co.: "Lost Valley" a deeply entrenched narrow valley, 
with rich, moist, mesophytic woods (mainly Fagus), 2 mi NE of 
Boxley (49 mi E of Fayetteville), Alt. 1000 ft., 25 Oct 1953, H. H. 
Iltis 4456 (SMU); Sec. 29, TI6N, R22W, 23 Sep 1976, EF. B. Smith 
3095 (UARK). 


Our collection of Aster shortii is from a rocky dry to dry- 
mesic forest adjacent to a shale glade, at 1250 ft. elev. in the 
central Ouachita Mountains (Novaculite Uplift). Here, it is found 
under a canopy of Quercus shumardii Buckl., Pinus echinata P. 
Mill., and Carya tomentosa (Lam. ex Poir.) Nutt., with a subcanopy 
of Ostrya virginiana (P. Mill.) K. Koch, Amelanchier arborea 
(Michx. f.) Fern., and Acer rubrum L. Herbaceous layer associates 
include Antennaria plantaginifolia (L.) Richards, Arabis laevigata 
(Muhl. ex Willd.) Poir., Cunila origanoides (L.) Britt., Galium 
arkansanum Gray var. pubiflorum E. B. Smith, Porteranthus 
stipulatus (Muhl. ex Willd.) Britt., Liatris squarrosa (L.) Michx., 
and Solidago petiolaris Ait. 

Aster shortii is apparently close to A. azureus Lindl., which 
is common in the Ozarks and Ouachitas of Arkansas. No mention 
is made of A. shortii in Smith (1978). Cronquist (1980) includes 
Arkansas in the range of A. shortii, stating that it is apparently 
isolated in the Ouachita Mountains. The Iltis 4456 specimen was 
determined by Almut G. Jones in 1978. The Smith specimen was 
annotated in 1980 by Jones as "Not A. azureus; perhaps near A. 
anomalus Engelm. - If found in Illinois, I would call the plant A. 
shortii Lindl. in Hook." We found that this specimen had the 
pubescent involucral bracts of A. shortii, but the inflorescence was 
unusually compact, with short branches and few bracts, as also is 
true of the other specimens examined. As a group, we refer them 


86 PY 'T OL O61. A Vol. 64, No. 2 


to A. shortii, although they may be of hybrid origin or represent 
introgression from other species. 


ASTRAGALUS DISTORTUS T. & G. var. ENGELMANNII 
(Sheld.) Jones (Fabaceae). Garland Co.: Sec 25, T3S, R22W, 4 Apr 
1985, Bridges and Orzell 85-17 (NY, TEX, UARK). Miller Co.: Sec 
33, T18S, R28W, 31 Mar 1985, Orzell and Bridges 1617 (NY, SMU, 
UARK), 18 May 1986, Bridges 86-46 (TEX). Polk Co.: Sec 11, 
T2S, R32W, 30 Apr 1985, Orzell and M. Medley 1801 (TEX, UARK). 
Pulaski Co: Sec 34, TIN, R12W, 16 Apr 1984, Orzell 1267 
(UARK), 19 Mar 1985, Orzell and Bridges 1576 (SMU). Sharp Co.: 
Sec 7, TI8N, R4W, 16 Apr 1985, Orzell 1690 (TEX, UARK). 


This variety was found in a large number of habitats in 
Arkansas, including dolomite glades, nepheline syenite (igneous) 
glades, shale roadbanks and pits, and sandhill woodland openings. 
In general, it occupies more xeric, specialized habitats than var. 
distortus. Smith (1978) synonymizes this variety under A. 
distortus, but it is recognized as distinct by Barneby (1964), 
Correll and Johnston (1970), and Kartesz and Kartesz (1985). All 
the above specimens were verified by Dr. Rupert Barneby at the 
New York Botanical Garden. 


ASTRAGALUS LEPTOCARPUS T. & G. (Fabaceae). Miller 
Co.: Sec 17, T18S, R28W, 31 Mar 1985, Orzell and Bridges 1615 
(SMU, UARK), 18 Jun 1980, Tucker 19422 (APCR); 2.5 mi N of 
ARK 160 int W ARK 237, 31 Mar 1981, Davis and Rettig 2828 
(APCR); 1.4 mi N of int of ARK 237 and road leading to Atlanta 
TX, 25 May 1974, Tucker 14058 (APCR). Nevada Co.: Sec 24, 
T12S, R20W, 29 Mar 1985, Orzell and Bridges 1604 (NY, SMU, 
TEX); Sec ll, T11S, R20W, 5 May 1978, J. Roberts 728 (UARK). 
Ouachita Co.: Sec 35, T12S, R19W, 29 Mar 1985, Orzell and 
Bridges 1602 (NY, SMU, TEX); Near Chidester, 27 Jun 1956, D. M. 
Moore 56140 (UARK-2). 


Astragalus leptocarpus is locally abundant along the edges of 
sandhill woodlands and sandy roadsides. In Miller County 
associates include Quercus incana Bartr., Quercus margaretta Ashe 
ex Small, Hymenopappus artemisiifolius DC., Tradescantia 
reverchonii Bush, Astragalus soxmaniorum Lundell, Linaria 
canadensis (L.) Dum.-Cours., Krigia virginica (L.) Willd., 
Andropogon virginicus L., Viola rafinesquii Greene, and Triodanis 
perfoliata (L.) Nieuwl. 

We were surprised that the only annuai Astragalus we had 
collected in southwestern Arkansus was A. leptocarpus, since Smith 
(1978) listed only A. nuttallianus DC. for these counties. To 
investigate this problem, we examined the cited specimens at 
APCR and UARK. Smith had in 1984 annotated all UARK 
collections cited as A. leptocarpus, and we determined all APCR 
material also to be this species. Apparently, A. nuttallianus has 
only been collected once in Arkansas (Moore 400159, UARK) from 


1987 Orzell & Bridges, Flora of Arkansas 87 


Little River County. Arkansas is not included in the range of A. 
leptocarpus by Turner (1959), Barneby (1964), or Correll and 
Johnston (1970). Lasseigne (1973) reports it from six Louisiana 
parishes, and it has been collected by the authors and John and 
Connie Taylor in three counties in southeastern Oklahoma (Taylor 
and Taylor 1987). Despite having first been collected 30 years 
ago, ours is the first report of A. leptocarpus for Arkansas. 
Specimens 1602, 1604, and 1615 were verified by Dr. Rupert 
Barneby at the New York Botanical Garden. 


ASTRAGALUS SOXMANIORUM Lundell (Fabaceae). Miller 
Co.: Sec 17, T18S, R28W, 31 Mar 1985, Orzell and Bridges 1614 
(NY, UARK). Nevada Co.: Sec 28, T12S, R20W, 11 May 1982, 
Rettig 518 (APCR), 9 May 1983, Rettig 840 (pers. herb.); Sec 11, 
T12S, R20W, 9 May 1983, Rettig 855 (UARK); Sec. 24, T12S, R20wW, 
29 Mar 1985, Orzell and Bridges 1603 (NY, TEX); Sec. 11, T11S, 
R20W, 5 May 1978, J. Roberts 727 (UARK). Ouachita Co.: Sec 12, 
T12S, R19W, 9 May 1983, Rettig 832 (NY); Sec 33, T14S, R17W, 29 
Mar 1985, Orzell and Bridges 1599 (NY, SMU, TEX); Chidester, 1 
Apr 1951, D. M. Moore 510037 (UARK). Union Co.: Sec 13, T17S, 
R15W, 17 Mar 1985, Orzell, Bridges, and Carl Amason 1573 (TEX), 
3 Apr 1984, Orzell and Carl Amason 1258 (SMU), 16 Mar 1984, 
R.T. Huffman 35 (UARK). 


All of the Arkansas collections of this species are from areas 
of deep sandy soil, including relatively undisturbed sandhill 
woodlands, sandy old fields, and roadsides through these areas. 
The open canopy of the sandhill woodland sites includes Quercus 
arkansana Sarg., Quercus incana, Quercus margaretta, and Pinus 
taeda L. The herbaceous layer in both sandhill woodland, sandy 
fallow field, and sandy roadsides, includes Andropogon virginicus, 
Gymnopogon ambiguus (Michx.) B.S.P., Diodia teres Walt., Viola 
rafinesqulii, Monarda punctata L., Opuntia humifusa (Raf.) Raf., 
Cassia fasciculata Michx., Astragalus leptocarpus, Lithospermum 
carolinense (J.F. Gmel.) MacM., Lechea villosa Ell., Froelichia 
floridana (Nutt.) Moq., Hymenopappus artemisiifolius, Tradescantia 
reverchonii, Linaria canadensis, Krigia virginica, and Triodanis 
perfoliata. 

Barneby (1964) noted this species to be a local endemic of 
eastern Texas, reporting it from 14 sites in nine counties. Since 
then it has been found in three parishes in northern Louisiana 
(Lasseigne 1973), and now in southern Arkansas. In addition to 
the above cited specimens, the authors have observed this species 
at over 30 other Arkansas locations in Miller, Nevada, and 
Ouachita counties, in sandhill woodlands, on sandy roadsides, and 
particularly in cemeteries on sandy soil. This represents the first 
report of the species in Arkansas, and a northward range 
extension from northeast Texas and adjacent northwest Louisiana. 
Specimens 1258, 1573, 1599, 1603, and 1614 were verified by Dr. 
Rupert Barneby. 


88 PHY, 2: 0,L, 0 GT A Vol. 64, No. 2 


BERLANDIERA x BETONICIFOLIA (Hook.) Small, pro sp. 
(Asteraceae). Miller Co.: Sec 18, T18S, R28W, 17 May 1985, 
Orzell and Bridges 1879 (UARK); Sec 33, T18S, R28W, 18 May 
1986, Bridges 86-47 (SMU, TEX, UARK); Sec 7, T20S, R27W, 7 Oct 
1985, Orzell, Bridges, and Peacock 3281 (TEX); Sec 17, T18S, 
R28W, 18 May 1986, Bridges 86-54 (SMU, TEX, UARK). 


This hybrid complex is occasional along sandy roadsides in 
Miller County, associated with Hymenopappus artemisiifolius. 
Additionally, it is found in natural openings in sandhill woodlands 
with Tradescantia reverchonii, Thelesperma filifolium (Hook.) Gray, 
Selaginella arenicola Underwood ssp. riddellii (Van Eselt.) R. 
Tryon, and other sandhill species. Neither this hybrid nor one of 
its parents, B. pumila (Michx.) Nutt., has previously been known to 
occur in Arkansas, although both are known from adjacent Cass 
County, Texas (Pinkava 1967). Our specimens mostly fit the 
putative backcross hybrid of Berlandiera x betonicifolia with B. 
pumila, which is probably the most abundant form of this hybrid 
complex in northeastern Texas (Pinkava 1967). Although some of 
our specimens have dense, matted white-tomentose pubescense, and 
could be referred to pure B. pumila, we feel it is best to simply 
recognize the fact that this hybrid complex extends to the state, 
and that individual collections will express the characters of 
various forms of the complex, from pure forms of each parent to 
exact intermediates. 


CARDAMINE ANGUSTATA O. E. Schulz var. MULTIFIDA 
(Muhl.) Ahles (Brassicaceae). Polk Co.: Sec 21, T4S, R28W, 23 Mar 
1985, Orzell and Bridges 1577 (SMU, TEX, UARK), 1 Apr 1986, 
Orzell and Bridges 3824 (SMU, UARK); Sec. 20, T4S, R28W, 1 Apr 
1986, Orzell and Bridges 3826 (NLU, TEX); Sec. 20, T4S, R28W, 1 
Apr 1986, Orzell and Bridges 3830 (TEX, US); Sec. 7, T4S, R28W, 1 
Apr 1986, Orzell and Bridges 3831 (DUR, MO, NCU, NY, SMU, 
TENN, TEX, UARK). 


Cardamine angustata var. multifida was most often found on 
stable wet and sandy gravel streambanks covered with leaf litter 
along the spring-fed headwaters of Blaylock Creek. The nearby 
canopy consisted of Liquidambar styraciflua L. and Fagus 
grandifolia Ehrh., with a subcanopy of F. grandifolia, Quercus alba 
L. and Carpinus caroliniana Walt. Nearby herbaceous associates 
included Iris cristata Soland., Toxicodendron radicans (L.) Kuntze, 
Viola sororia Willd., Solidago caesia L., Epifagus virginiana (L.) 
Bart., Trillium pusillum Michx. var. ozarkanum (Palmer and 
Steyermark) Steyermark, Arabis laevigata, Erythronium rostratum 
W. Wolf, Thalictrum thalictroides (L.) Eames & Boivin, Tradescantia 
ernestiana Anderson & Woods., Athyrium filix-femina (L.) Roth var. 
asplenioides (Michx.) Farw., Uvularia sessilifolia L., Cardamine 
angustata var. ouachitana E. B. Smith, Luzula echinata (Small) F. 
J. Herm., L. acuminata Raf., and Carex sp. At one site this 
species was in a rocky seepage area at the headwaters of a small 


1987 ay Orzell & Bridges, Flora of Arkansas 89 


stream with many of the same species, and a particularly large 
population of Trillium pusillum var. ozarkanum. 

This taxon (as Dentaria multifida Muhl.) was first reported 
for Arkansas by the late Polk County botanist Aileen McWilliam 
(McWilliam 1966), who noted three distinct Dentaria species in the 
central Ouachita Mountains. Smith (1978) synonymized all previous 
reports of other taxa in Arkansas with Dentaria Iaciniata Muhl. 
Later (Smith 1982b) he recognized the uniqueness of one of Miss 
McWilliam’s 3 species, and described it as a new variety 
(Cardamine angustata O.E. Schulz var. ouachitana E.B. Smith). In 
1985 and 1986 we revisited the general area mentioned for D. 
multifida by Miss McWilliam and found it at several closely spaced 
stations. Miss McWilliam, being extremely conservation-minded, 
rarely collected vouchers, thus our collections probably represent 
the first of this taxon from Arkansas. Montgomery (1965), 
Fernald (1950), and Harriman (1965) do not include Arkansas in 
the range of this taxon. Harriman (1965) further notes its unusual 
discontinuous distribution pattern in the eastern states, and 
includes no records west of the Western Highland Rim of 
Tennessee and the Cumberland Plateau of Alabama. While 
examining specimens of Cardamine at DUR in May 1985, we 
annotated several collections, previously identified as D. laciniata, 
as C. angustata var. ouachitana and one as C. angustata var. 
multifida from the Ouachita Mountains of adjacent Oklahoma, both 
new for the state (Taylor and Taylor 1987). Our collections and 
that for Oklahoma are the first for this taxon west of the 
Mississippi River. 


Notes on the genus Carex 


Before discussing the occurrence of the following species of 
Carex in Arkansas, we feel that some general commentea on the 
taxonomy of the genus, particularly in the southern states, is in 
order. Carex is predominantly a genus of the northern latitudes, 
and it is there that most of its taxonomy originates. The 
taxonomy of southern Carex species, and of northern species 
extending to the southern United States, has largely been based 
on too little material from too few localities to be thought of as 
definitive. Many species which are inherently uncommon and 
restricted to specialized habitats in the southern states are known 
from only a very few collections in each state. There has been a 
tendency, when confronted with only a handful of collections 
supposedly representing several closely related species in their 
states, for southern authors to synonymize Carex species or their 
reports for the state of concern. This is particularly true of the 
Carex treatments of Correll and Johnston (1970) and Smith (1978), 
and to a lesser extent in Radford et al. (1968). Undoubtedly, 
critical taxonomic study including a good, representative, and 
thorough set of collections from the southern states would 
eventually uncover the intergradations between many of these 
species. However, in the absence of such studies for most 


90 PHHEYST: OnL7OrG) 17h Vol. 64, No. 2 


sections of the genus, it seems best to follow the traditional 
morphological species concepts, and leave the interpretations of 
which constitute biological species to critical taxonomic studies. 
Recognition of the occurrence of these traditional species in each 
state is important as it provides systematists with clues to their 
ranges and possible intergradations. Therefore we are attempting 
to follow the species concepts of northern workers such as 
Fernald (1950), Voss (1972), Mohlenbrock (1975), and Reznicek 
(pers. comm.) in our interpretations of Carex species. 


CAREX ARTITECTA Mackenzie (Cyperaceae). Jackson Co.: 
Sec. 16, T14N, R1W, 23 Apr 1987, Orzell and Bridges 5143 (TEX, 
UARK). Little River Co.: Sec. 1, T10S, R33W, 1 Apr 1985, Orzell 
and Bridges 1640 (MICH). Montgomery Co.: Sec. 8, T4S, R27W, 30 
Apr 1985, Orzell and M. Medley 1800 (MICH). Searcy Co.: Sec. 9, 
T16N, R17W, 11 Jul 1985, Orzell and Bridges 2518 (UARK). 


Smith (1978) includes C. artitecta for Arkansas, but in a later 
supplement (1982a) changes the name of this taxon to C. 
physorhyncha Liem. Most workers (Radford et al., 1968, 
Mohlenbrock 1975, Russell and Duncan 1972) think of these as two 
distinct taxa, both of which are in Arkansas according to 
Mackenzie (1935). We suspect that C. artitecta is the common 
member of this group in the Interior Highlands and south to the 
Cretaceous section of the Coastal Plain, and that C. physorhyncha 
is confined to the Coastal Plain in Arkansas. 


CAREX BICENELLII Britt. var. BICKNELLII (Cyperaceae). 
Franklin Co.: Sec. 3, T7N, R28W, 8 Jun 1978, G. Barber 861 
(UARK). 


This variety was found in a saline soil barrens developed on 
the Wing series (Aquic Natrustalfs). This unusual community type 
supports several plants which are rare in the state, such as 
Sporobolus pyramidatus (Lam.) A. S. Hitchc. and Geocarpon 
minimum Mackenzie. This species is listed by Barber (1979) in her 
flora of Franklin County, but apparently has not yet been reported 
in the published literature. The specimen was verified by the 
authors and by Dr. A. A. Reznicek. 

The other variety of thie species, Carex bicknellii var. opaca 
F. J. Herm., is endemic to the Grand Prairie region of eastern 
Arkansas (Hermann 1972). Carex bicknellii is listed for Arkansas 
by Mackenzie (1935) but this probably refers to the later described 
var. opaca. Smith (1978) does not mention the occurrence of var. 
bicknellii in Arkansas. 


CAREX BULBOSTYLIS Mackenzie (Cyperaceae). Hempstead 
Co.: Sec. 24, T12S, R27W, 27 Apr 1985, Orzell and M. Medley 1734 
(MICH). 


This species was found in a mesic sandy ravine forest on the 


1987 Orzell & Bridges, Flora of Arkansas i 


Coastal Plain, with some calcareous influence. Smith (1978) 
synonymizes this species under C. amphibola Steud., as do Correll 
and Johnston (1970). However, it is listed for Arkansas by 
Mackenzie (1935), and we consider it to be a distinct taxon. 

Carex bulbostylis ranges from western Arkansas and Louisiana 
south to central Texas. 


CAREX HYALINA Boott. (Cyperaceae). Miller Co.: Rich 
woods, 27 Apr 1905, B. F. Bush 2475 (NY); Texarkana, woods, 28 
Apr 1905, B. F. Bush 2500 (NY). County uncertain: near Fulton, 
W. M. Canby 186 (PH). 


This species is apparently a rare, or very undercollected, 
endemic of the upper West Gulf Coastal Plain. In addition to the 
above cited records, it is known only from one collection in 
McCurtain County, Oklahoma and from four counties in Texas 
(Correll and Johnston 1970). A collection originally identified as 
this species from Jackson Co., Arkansas (E. J. Palmer 35530, MO, 
NY) is apparently C. reniformis (Bailey) Small (Smith 1978), and is 
out of range for C. hyalina. Additional fieldwork is neccessary in 
order to understand the taxonomy and rarity of this little 
collected species of the section Ovales. 


CAREX GRISEA Wahl. (C. amphibola Steud. var. turgida 
Fern.) (Cyperaceae). Howard Co.: Sec 29, T7S, R27W, 27 Apr 
1985, Orzell and M. Medley 1746 (SMU). 


This species was found in a rich, dry-mesic calcareous slope 
forest developed on DeQueen Limestone (Cretaceous), with a 
canopy of Quercus shumardii, Q. muhlenbergii Engelm., Juglans 
nigra L., and Carya cordiformis (Wang.) K. Koch. Herbaceous 
associates include Cypripedium kentuckiense C. F. Reed, Senecio 
obovatus Muhl. ex Willd., Adiantum pedatum L., Phryma 
leptostachya L., Carex texensis (Torr.) L. H. Bailey, and Solidago 
auriculata Shuttlw. ex Blake. 

Mackenzie (1935) includes this species for Arkansas. Smith 
(1978) references this species under C. amphibola, stating that C. 
amphibola includes C. grisea of authors, not Wahl., as do Correll 
and Johnston (1970). Steyermark (1963) considered C. grisea to be 
dubiously distinct from C. amphibola. However, it seems to be 
recognized still by at least some workers (Russell and Duncan 
1972, Radford et al. 1968), and we reacknowledge its presence in 
Arkansas. 


CAREX INTERIOR L.H. Bailey (Cyperaceae). Fulton Co.: 
Sec. 7, T20N, R8W, 23 May 1985, Orzell and Bridges 1939 (MICH, 
MO, TEX, UARK). Sharp Co.: Sec 7, T18N, R4W, 18 Apr 1985, 
Orzell 1700 (MICH), 24 May 1985, Orzell and Bridges 1950 (MICH, 
UARK). 


In Sharp County, Carex interior is known from a series of 


92 P Horo bo Law Vol. 64, No. 2 


calcareous seep fens (Orzell et al. 1985), where it forms hummocks 
on a gravelly marly substrate constantly saturated by cold, 
minerotrophic seepage. Associates at the Sharp County station are 
Selaginella apoda (L.) Fern., Rudbeckia fulgida Ait. var. umbrosa 
(C.L. Boynt. and Beadle) Cronq., Parnassia grandifolia DC., 
Solidago riddellii Frank, Senecio aureus L., Carex suberecta 
(Olney) Britt., C. hystricina Muhl. ex Willd., C. leptalea Wahlenb., 
Castilleja coccinea (L.) Spreng., and Ludwigia microcarpa Michx. 

At the Fulton County site it is common on quaking sphagnous peat 
saturated by cold minerotrophic seepage, with Rudbeckia fulgida 
var. umbrosa, Pogonia ophioglossoides (L.) Juss., Senecio aureus, 
Carex suberecta, and Parnassia grandifolia. 

Smith (1978) reported C. interior in Arkansas, but in a later 
supplement (Smith 1981) he states that all Arkansas material 
annotated by A. Reznicek is either Carex atlantica Bailey subsp. 
atlantica or C. atlantica Bailey subsp. capillacea (Bailey) Reznicek, 
and thereby excludes C. interior from Arkansas. Our collections 
of Carex interior therefore represent the first authentic material 
from the state. Reznicek and Ball (1980) report this species as 
occurring south only to Pennsylvania, Ohio, Indiana, Illinois, and 
Missouri in eastern North America, although occurring sporadially 
south into Mexico in the west; therefore, our collection seems to 
be the first for the southeastern United States. Both specimens 
were verified by Dr. Anton Reznicek. 


CAREX LAXICULMIS Schwein. (Cyperaceae). Van Buren Co.: 
Sec 15 & 14, T12N, R16W, 9 Jul 1985, Orzell, Bridges, L. Peacock, 
and T. Foti 2462 (MICH). 


This species was found in a dry-mesic calcareous ravine 
forest in the Boston Mountains of the Ozark Plateaus. Other 
carices at this site include C. digitalis Willd., C. cephalophora 
Muhl. ex Willd., C. laxiflora Lam., and C. rosea Schkuhr ex Willd. 
Other associated herbaceous species include Uvularia sessilifolia, 
Brachyletrum erectum (Schreb. ex Spreng.) Beauv., Galium 
arkansanum, Taenidia integerrima (L.) Drude, Goodyera pubescens 
(L.) R. Br., Senecio obovatus, and Cypripedium pubescens Willd. 

Smith (1978) includes this species in his "possible additions", 
based on Steyermark (1963), as perhaps in northeast Arkansas. 
Bryson (1980) states that C. laxiculmis ranges southward to 
northern Arkansas, but cites no specimen nor indicates its 
presence on his distribution map. To our knowledge this is the 
first collection of this species for the state, and the southwestern 
most record for the species. 


CAREX LAXIFLORA Lam. (Cyperacese). Van Buren Co.: 
Sec 14 & 15, T12N, R16W, 9 Jul 1985, Orzell, Bridges, L. Peacock, 
and T. Foti 2471 (UARK). 


The habitat and associates for this species are the same as 
that for the previous one, C. laxiculmis. It was listed for 


1987 Orzell & Bridges, Flora of Arkansas 93 


Arkansas by Demaree (1943) and reported for Crittenden, Newton, 
and Stone counties by Wilcox (1973), Thompson (1975), and Browne 
(1974), respectively. Smith (1978) excludes this species from the 
flora, stating that he had not seen voucher material of authentic 
C. laxiflora var. laxiflora for Arkansas. We have not seen the 
specimens upon which these reports were based, so cannot judge if 
they are correctly identified. Bryson (1980) indicates that C. 
laxiflora ranges west only to Tennessee (to the Western Highland 
Rim) and Alabama (to the Cumberland Plateau). We present this 
record in order to have a citation for this species in Arkansas, 
perhaps the first from west of the Mississippi River and at its 
western limit. 


CAREX LONGII Mackenzie (Cyperaceae). Calhoun Co.: Sec. 
28, T14S, R13W, 16 Jun 1982, J. Roberts 1599 (MICH); Sec. 32, 
T13S, R12W , 16 Jun 1982, J. Roberts 1602 (MICH). Clark Co.: 
Sec 30, T9S, R21W, 15 Aug 1985, Orzell and Bridges 2821 (UARK), 
12 Oct 1985, Orzell, Bridges, and Peacock 3465(MICH). Pulaski 
Co.: Camp Robinson, north of Little Rock, 6 May 1939, G. M. 
Merrill 1896 (A). Saline Co.: low moist bottoms, P.O. Benton, 
elev. 300 ft., 16 May 1942, Demaree 22969 (SMU). 


Carex longii was collected in Clark County from a roadside 
ditch with seepage from an adjacent slope. Associated plants 
include Rhexia mariana, R. virginica L., Gratiola pilosa, Ludwigia 
alternifolia L., Lobelia puberula Michx., Eupatorium rotundifolium, 
Arthraxon hispidus (Thunb.) Makino, Rhynchospora inexpansa, R. 
Slomerata, Fimbristylis miliacea (L.) Vahl, Panicum verrucosum 
Muhl., Xyris difformis Chapman var. difformis, Juncus sp., 
Eleocharis sp. and Sphagnum moss. 

This species of the Ovales series was reported for Arkaneas 
in Demaree (1943). Smith (1978) excluded this species from the 
flora as probably not in the state. C. Jongii is closely related to 
C. albolutescens Schwein., with which it may previously have been 
confused in Arkansas. We have not seen the Merrill specimen 
from Pulaski County. 


CAREX LOUISIANICA L. H. Bailey (Cyperaceae). Hempstead 
Co.: Below Millwood Dam, bottoms, common, Beard Lake area, P. 
O. Columbus, Elev. 300, 25 Apr 1969, Demaree 59933 (SMU). 
Miller Co.: Low wet areas, common, P. O. Garland, Elev. 320, 23 
May 1960, Demaree 42410 (SMU). Union Co.: El Dorado, heavy 
soil, creek bottom, woods, 12 Jul 1953, A. J. Hoiberg 344 (SMU). 


_ Carex louisianica was first reported for Arkansas by Buchholz 
and Palmer (1926), and listed for the state by Mackenzie (1935). 
This species was placed in synonymy under C. Jupulina Muhl. ex 
Willd. by Smith (1978). Reznicek and Ball (1974) consider these 
taxa to be quite distinct, and we here reinstate C. louisianica in 
the Arkansas flora. 


94 PHY TO .b 0 GP A Vol. 64, No. 2 


CAREX MOLESTA Mack. ex Bright (Cyperacene). 
Washington Co.: Few in small seepage area just W. of the Art 
Dept. Annex (Parking Lot 19), U. of A. Campus in Fayetteville. 
Flowering stalks ascending and widely arching, 17 May 1982, E. B. 
Smith 3678 (UARK). 


Mackenzie (1935) does not list this species for Arkansas, 
however, it is listed for Arkansas by Steyermark (1963). Smith 
(1978) includes it in possible additions based on the Steyermark 
report. This collection, originally identified as C. bicknellii, may 
be the first for the state, and is the southernmost record for this 
species. The specimen was determined by A. A. Reznicek. 


CAREX MUHLENBERGII Willd. var. ENERVIS Boott. (C. plana 
Mackenzie) (Cyperaceae). Pulaski Co.: Sec 34, TIN, R12W, 10 
May 1985, Orzell and Bridges 1823 (MICH), Orzell and Bridges 1830 
(UARK). 


This variety was collected in dry, acidic woodlands within a 
nepheline syenite intrusive igneous glade complex on the south- 
facing slope of Granite Mountain, under an open canopy of 
Quercus stellata Wang., @. marilandica Muenchh., and Ulmus alata 
Michx. 

This variety was first reported for Arkansas by Buchholz and 
Palmer (1926) from Carroll and Logan counties, and listed for the 
state by Mackenzie (1935). It is synonymized under C. 
muhlenbergii by Smith (1978) and Radford et al. (1968), but 
maintained by Fernald (1950) and Steyermark (1963). It is perhaps 
dubiously distinct from typical C. muhlenbergii, but may have some 
merit. 


CAREX MUSKINGUMENSIS Schwein. (Cyperaceae). 
Crittenden Co.: Demaree 12959 (MICH). Lawrence Co.: Sec. 11, 
T17N, R2E, 24 Oct 1985, Orzell and Bridges 3598 (MICH). 


At the Lawrence County site, Carex muskingumenasis is 
scattered in open areas of a wet-mesic interstream flatwoods 
dominated by Quercus phellos L., with @. pagoda Raf., and @. 
lyrata Walt. Nearby associates include Lindera melissifolia (Walt.) 
Blume, Leitneria floridana Chapman, Scutellaria lateriflora L., and 
Ammannia coccinea Rottb. 

This species, perhaps the most distinctive member of the 
Ovales series, was first reported for Arkansas by Demaree (1941, 
1943). Hermann, who had determined the Demaree specimen, 
included it for Arkansas in his "Addenda to North American 
Carices" (Hermann 1954). Smith (1978) excludes this species from 
the flora of Arkansas, stating "this is another ‘minispecies’ of the 
Ovales series, perhaps possible in the state, but I have seen no 
evidence of it". Dr. A. A. Reznicek, who verified our 
determination, considers it "a thoroughly distinct species" (pers. 
comm.). 


1987 Orzell & Bridges, Flora of Arkansas 95 


CAREX OKLAHOMENSIS Mackenzie (Cyperacese). Arkansas 
Co.: Moist rice prairie habitats, P. O. DeWitt, Elev. 170, 12 May 
1940, Demaree 21094 (SMU). Drew Co.: Margins of small swamp, 
P. O. Monticello, Elev. 250, 2 May 1941, Demaree 22012 (SMU). 

Hot Spring Co.: Creek banks, P. O. Butterfield, Elev. 380, 23 Oct 
1958, Demaree 40786A (SMU). Lonoke Co.: Rice prairies, 
common, never plowed, P. O. Carlisle, Elev. 230, 31 Apr 1967, 
Demaree 55620 (SMU); Low wet areas, rice region, never plowed, 
river terraces, P. O. Carlisle, Elev. 230, 5 May 1968, Demaree 
57790 (SMU). Polk Co.: Small wet bottom, P. O. Cherry dill, Elev. 
850, 13 May 1962, Demaree 45575 (SMU). Prairie Co.: Rice 
prairies, never plowed, common, river terraces, low spots, P. O. 
Hazen, Elev. 215, 20 May 1970, Demaree 61903 (SMU). Pulaski 
Co.: Swampy areas 5 miles north of Little Rock on Highway 5, P. 
O. Little Rock, Elev. 400, 15 May 1963, Demaree 47672 (SMU). 


Carex oklahomensis seems to be the most common member of 
the series Vulpinae in low, open, wet areas throughout much of 
Arkansas. Correll and Johnston (1970) consider this species to be 
the product of past hybridization of "C. stipata Muhl. ex Willd. 
and C. muhlenbergii (especially the form "C. lunelliana 
Mackenzie")" but present no evidence for this claim. Smith (1978) 
includes this species in synonymy under C. stipata; however, many 
Carex experts consider it to be distinct as a variety or species. 
All of the cited specimens were determined or verified by A. A. 
Reznicek. 


CAREX OUACHITANA Kral, Manhart & Bryson (Cyperaceae). 
Montgomery Co.: Sec 8, T4S, R27W, 2 Jun 1985, Orzell and 
Bridges 2030 (MICH), 26 Apr 1986, Orzell 4284 (MICH, MO, NCU, 
SMU, TEX, UARK). Scott Co.: Sec 8, T3N, R32W, 21 Apr 1986, 
Orzell and Bridges 4212 (MO, NCU, TEX, UARK); Sec 30, T4N, 
R30W, 23 Apr 1986, Orzell and Bridges 4274 (SMU, TEX). 
Sebastian Co.: Sec 8, T3N, R32W, 21 Apr 1986, Orzell and Bridges 
4214 (MICH, TEX, UARK). 


This newly described species (Kral et al. 1987) provides a 
classic example of the underexplored and undercollected status of 
Arkansas botany. It is a common, sometimes dominant, species of 
the herbaceous layer of dry to dry-mesic acid, rocky ridgetop and 
slope forests throughout much of the Ouachita Mountains. This 
sedge is now known to occur in all three physiographic sections of 
the Ouachitas - in the Athens Piedmont Section on Stanley Shale 
in Howard and Polk counties, in the Novaculite Uplift Section on 
strongly faulted and folded Stanley Shale in Montgomery County, 
and in the Fourche Mountains Section on Jackfork Sandstone in 
Polk and Scott counties and LeFlore County, Oklahoma, and on 
the Savanna Formation of Poteau Mountain in Sebastian and Scott 
counties. At the Montgomery County site, C. ouachitana was 
common in a dry, steep, shaley slope forest dominated by Quercus 
shumardii, Ostrya virginiana, Pinus echinata, and Carya tomentosa, 


96 P..Hev eT 0) LuOrG Tek Vol. 64, No. 2 


with a subcanopy of Acer rubrum and Amelanchier arborea. Shrub 
and herb associates include Rhus aromatica Ait., Dioscorea villosa 
L., Vaccinium pallidum Ait., Galium arkansanum var. pubiflorum, 
Helianthus divaricatus L., Antennaria plantaginifolia, Liatris 
squarrosa, Solidago petiolaris, and Aster shortii. The Scott and 
Sebastian County sites are from stunted dry to dry-mesic oak 
forests on sandstone near the top of Poteau Mountain. 

We first collected this species in 1985, and were made aware 
of its unusual characteristics by A. A. Reznieck. We planned 
fieldwork in 1986 to collect more material in order to determine if 
it was a distinct species and to prepare a description. Meanwhile, 
we learned that Robert Kral was also working on a new Carex 
from the Ouachita Mountains. In June 1986, we sent a preliminary 
description of our specimens to Robert Kral, and offered our 
material from the above new sites for his study and use in 
describing the species. Despite his refusal of our offer, we 
refrained from publishing this species as a courtesy to its authors. 
Apparently, Kral felt that if one has "sufficient" material to 
prepare a description, then there is no need for additional 
specimens to study. Our collections represent significant new 
habitats and range extensions for this species, and may be 
important in understanding the variation present in this group. 


CAREX PLANOSTACHYS Kunze (Cyperaceae). Little River 
Co.: Sec 28, T12S, R32W, 1 Apr 1985, Orzell and Bridges 1626 
(MICH). 


This species was found under scattered, open-grown Juniperus 
virginiana L. within a chalk glade/outcrop complex developed on 
Annona Chalk (Cretaceous). Nearby herbaceous associates include 
Bouteloua rigidiseta (Steud.) A. S. Hitchc., Galium virgatum Nutt., 
Ophioglossum engelmannii Prantl, Carex cherokeensis Schwein., 
Dalea compacta Spreng. var. compacta, and Indigofera miniata 
Ortega var. leptosepala (Nutt.) B. L. Turner. 

This is the first collection of this species for Arkansas, and 
a disjunct from north central Texas, where it ranges south 
through the Edwards Plateau into Mexico. It is locally abundant 
and one of the most characteristic herbaceous species on limestone 
uplands and slopes dominated by a grassland - woodland vegetation 
mosaic in the Balcones region of the Edwards Plateau in central 
Texas (Riskind & Diamond 1986). Our specimen was determined by 
A. A. Reznicek. 


CAREX SPARGANIOIDES Willd. (Cyperaceae). Baxter Co.: 
low moist spots by railroad, Common, Below Cotter, P. O. Cotter, 
Elev. 450, 21 May 1951, Demaree 63552 (SMU). Van Buren Co.: 
Sec 7, T12N, R12W, 12 July 1985, Orgell and Bridges 2558 (MICH). 


Our collection is from a rich, mesic, north-facing limestone 
slope forest along the Middle Fork of the Little Red River. 
Among the herbaceous associates are Diarrhena americana Beauv., 


1987 Orzell & Bridges, Flora of Arkansas 97 


Galium triflorum Michx., G. circaezans Michx., and G. concinnum 
Torr. & Gray. Interestingly, the only Arkansas locality for the 
rare hybrid Dryopteris x leedsii Wherry is within one mile 
downstream from this site. 

Smith (1978) includes this species in "possible additions", 
stating that it may be rare in north or northwest Arkansas. 
Arkansas was not included in the range given for this species by 
Mackenzie (1935), Fernald (1950), or Steyermark (1963). Our 
collection seems to the the first for Arkansas, and extends the 
southwestern limit of the species. 


CAREX STRIATULA Michx. (Cyperaceae). Pope Co.: Nogo, 
rich woods, 24 Apr 1933, G. M. Merrill 215 (SMU). 


We found this sheet while studying Arkansas Carex 
collections at SMU. It was identified as and filed under C. 
laxiflora, and we determined it as C. striatula, later verified by A. 
A. Reznicek. Neither Smith (1978), Demaree (1943), Mackenzie 
(1935), or Bryson (1980) report C. striatula for Arkansas, although 
it was to be expected for the state based on its habitat and range 
in Mackenzie (1935) and Bryson (1980). Interestingly, the 
distribution given for this species in Bryson (1980) indicates that 
it would likely occur on the Coastal Plain of Arkansas, since it is 
cited for several Louisiana parishes and Texas counties bordering 
the state. This record seems to be the first for this species from 
Arkansas and from the Interior Highlands, and the 
northwesternmost record of the species. 


CAREX SUBERECTA (Olney) Britt. (Cyperacese). Fulton 
Co.: Sec. 7, T2Z0N, R8W, 23 May 1985, Orzell and Bridges 1941 
(MICH, UARK, TEX), Orzell and Bridges 1942 (MICH, MO, SMU). 
Sharp Co.: Sec. 7, T18N, R4W, 24 May 1985, Orzell and Bridges 
1955 (UARK). 


At the Fulton County station this species is common on 
quaking sphagnous peat saturated by cold minerotrophic seepage 
with Rudbeckia fulgida var. umbrosa, Pogonia ophioglossoides, 
Senecio aureus, Carex interior, and Parnassia grandifolia. In 
Sharp County, Carex suberecta occurs in calcareous seep fens with 
Selaginella apoda, Rudbeckia fulgida var. umbrosa, Parnassia 
grandifolia, Solidago riddellii, Senecio aureus, Carex interior, C. 
hystricina, C. leptalea, Castilleja coccinea, and Ludwigia 
microcarpa. 

Smith (1978) includes this species in his list of possible 
additions, stating “based on its distribution in Steyermark (1963) 
perhaps in NE or N-central Arkansas.” Our specimens are the 
first collections of this species from the state, and were verified 
by Dr. A. A. Reznicek. 


CAREX TEXENSIS (Torr.) L. H. Bailey (Cyperaceae). 
Howard Co.: marshy meadow and edge of pond, 4 mi SW of 


98 PeHOYT O8L/ Ore LA Vol. 64, No. 2 


Nashville, 14 Apr 1954, H. H. Iltis and D. M. Moore 666 (SMU); 
Sec 29, T7S, R27W, 27 Apr 1985, Orzell and M. Medley 1748 
(MICH). Jackson Co.: Sec. 16, T14N, R1W, 23 Apr 1987, Orzell 
and Bridges 5144 (MICH, MO, NCU, NLU, SMU, TEX, UARK). 


Our Howard County site is a dry-mesic calcareous slope 
forest (Bridges, in prep.) developed on DeQueen Limestone 
(Cretaceous), with a canopy of Quercus shumardii, Q. muhlenbergii, 
Juglans nigra, and Carya cordiformis. Herbaceous associates 
include Cypripedium kentuckiense, Phyrma leptostachya, Solidago 
auriculata, Carex grisea, and Osmorhiza longistylis (Torr.) DC. 

The Jackson County site is a flatwoods forest in an aeolian 
dune/depression area, with a canopy of Quercus alba, @. phellos, 
@. pagoda, Carya tomentosa, and Nyssa sylvatica Marsh. 

Arkansas was included in the range of this species by 
Mackenzie (1935). It is synonymized under C. retroflexa Muhl. ex 
Willd. by Smith (1978), Radford et al. (1968), and Correll and 
Johnston (1970). This species was recognized as distinct by 
Fernald (1950), in Georgia by Russell and Duncan (1972), in Illinois 
by Evans (1976), and as a variety of C. retroflexa by Steyermark 
(1963). We are considering it as at least of some merit, though 
further taxonomic study is necessary in order to understand at 
what level it should be recognized. 


CAREX WILLDENOWII Schkuhr. (Cyperaceae). Ashley Co.: 
Sec. 7, T18S, R5W, 11 Apr 1986, Orzell and Bridges 4021 (NLU, 
SMU, UARK); Sec. 19, T18S, R5W, 11 Apr 1986, Orzell and Bridges 
4032 (MO, TEX). Bradley Co.: Sec. 34, T14S, ROW, 22 Mar 1986, 
Orzell and Bridges 3713 (SMU), Orzell and Bridges 3714 (UARK). 
Hempstead Co.: Sec. 24, T12S, R27W, 27 Apr 1985, Orzell and M. 
Medley 1733 (MICH), Orzell and M. Medley 1735b (MICH). 
Montgomery Co.: Sec 17, T4S, R24W, 29 Apr 1986, Orzell 4310 
(MO, SMU, TEX, UARK); 18 May 1982, Rettig 532A (pers. herb.). 
Ouachita Co.: Sec. 12, T12S, R18W, 23 Apr 1987, Orzell and 
Bridges 5141 (MICH, SMU, TEX, UARK). Union Co.: Sec. 13, 
T17S, R14W, 12 Apr 1986, Orzell and Bridges 4060 (UARK). 


Carex willdenowii is found in a variety of mesic, well- 
drained, rather neutral soil situations throughout much of the 
Arkansas Coastal Plain and into the Ouachita Mountains. Where 
found in the Coastal Plain it occupies sites with a richer vernal 
flora than most Arkansas Coastal Plain forests, with associates at 
some sites including Obolaria virginica L., Styrax grandifolia Ait., 
Lindera benzoin (L.) Blume, and Polygonatum biflorum (Walt.) Ell. 
The two other members of the Section Phyllostachyae in Arkansas 
are’ found in quite different habitats; Carex jamesii Schwein. is 
found in more alkaline, calcareous clay soils, and Carex 
latebracteata Waterfall is found on well-drained dry to dry-mesic 
slopes in the Ouachita Mountains. In Bradley, Ashley, Hempstead, 
and Ouachita counties, C. willdenowli was found on lower slopes 
of ravine forests with sandy surface layers but with some 


1987 Orzell & Bridges, Flora of Arkansas 99 


calcareous influence. ‘Ihe Union County site was on a mesic 
stream terrace surrounded by sandy uplands. The Montgomery 
County site was also on a stream terrace, but surrounded by 
siliceous rocky slopes. 

Carex willdenowii is not mentioned in Smith (1978), nor is 
Arkansas included within its range by Mackenzie (1935), or Correll 
and Johnston (1970), the latter having seen no material of this 
species from Texas. Interestingly, we have collected C. 
willdenowii from nine Texas counties in 1987 alone. We believe 
these collections to be the first for Arkansas of this relatively 
common and widespread species in the Coastal Plain and Ouachita 
Mountains of the state. 


CLADIUM JAMAICENSE Crantz (Cyperaceae). Bradley Co.: 
miry bog in hard pan prairie, P.O. Warren, elev. 170 ft., 9 Jun 
1939, Demaree 19433 (SMU, MO). 


Typically, Demaree specimens with this locality information 
were collected in what is now known as Warren Prairie, an 
extremely unusual open, herbaceous-dominated saline soil barrens. 
The water table in these areas is very high in winter, and some 
areas are marshy almost year round. It is here we suspect that 
Cladium once occurred. The hydrology of this site has been 
altered, and despite the fact that this area has been investigated 
countless times by many botanists, Cladium jamaicense, as well as 
several other rare plants, have not been relocated and should be 
considered extirpated from Arkansas. 

Demaree (1943) reported this species for Arkansas, and it was 
in searching for the source of this report that we uncovered these 
specimens. Smith (1978) justifiably excluded this species from the 
Arkansas flora, stating it "does not seem very likely for Arkansas, 
based on its habitat and distribution in Correll and Johnston 
(1970) and Radford et al. (1968)". In Louisiana, this species 
occurs north only to Tangipahoa and Calcasieu parishes (Joyce 
1974). Additional specimens from Union Co. labelled by Demaree 
as C. jamaicense (Demaree 19423, SMU, MO) were determined by 
the authors to be immature material of a Rhynchospora or Scirpus. 


CONVOLVULUS EQUITANS Benth. (Convolvulaceae). Little 
River Co.: Sec. 21, T12S, R32W, 18 Jun 1985, Orzell and Bridges 
2173 (UARK). 


This species was occasional in gravelly blackland prairie 
openings, often climbing on Engelmannia pinnatifida Gray ex Nutt. 
Associates of C. equitans include Croton monanthogynus Michx., 
Heliotropium tenellum (Nutt.) Torr., Dalea compacta var. compacta, 
Phyla nodiflora (L.) Greene var. incisa (Small) Moldenke, Sabatia 
campestris Nutt., Hedyotis nigricans (Lam.) Fosberg, Bouteloua 
curtipendula (Michx.) Torr. and Juniperus virginiana. 

This species was listed for Arkansas as C. incanus Vahl by 
Branner and Coville (1891), on the basis of its report by Nuttall 


100 PH YeTAONEL O} GT A Vol. 64, No. 2 


(1837) as C. hastatus Nutt., probably based on an Oklahoma 
collection. It was also listed for Arkansas by Demaree (1943). 
Smith (1978) excluded this species from the Arkansas flora, stating 
that it is "unlikely for Arkansas, based on its distribution in 
Correll and Johnston (1970)." We believe our collection to be the 
first from within Arkansas’ current borders. 


CYPRIPEDIUM KENTUCKIENSE C. L. Reed (Orchidaceae). 
Clark Co.: 27 Apr 1985, Orzell and M. Medley 1710 (UARK). 
Crawford Co.: 30 Apr 1985, Orzell and M. Medley 1802 (UARK). 
Franklin Co.: 30 Apr 1977, G. Barber 417 (UARK); 31 Apr 1967, G. 
Tucker 4104 (NCU, SMU). Garland Co.: May 1970, D. Rhodes 
7818 (LTU); 26 Apr 1975, L. Adams s.n. (ATU); 12 May 1984, 
Orzell 1325 (NYS); 30 Apr 1942, D. M. Moore 420073 (UARK); 14 
Apr 1946, D. M. Moore 460031 (UARK); 30 May 1967, Grahams 747 
(NCU)?; 11 May 1984, J. & E. Price 8038 (SEL). Grant Co.: 19 
Apr 1981, M. Locke 4182 (UARK). Hempstead Co.: 18 May 1924, 
E. J. Palmer 24993 (UARK); 27 Apr 1985, Orzell and M. Medley 
1743 (UARK). 4 Oct 1923, J. T. Buchholz 401 (UARK); Howard 
Co.: 1 Oct 1974, D. Lawson 2447 (NLU); 27 Apr 1985, Orzell and 
M. Medley 1744 (UARK). Jefferson Co.: 20 Apr 1976, M. Locke 
1638 (UARK); 24 May 1975, R. D. Thomas et al. 44018 (NLU). 
Lincoln Co.: 24 Apr 1984, D. Flugrad 35 (UARK); 2 May 1985, M. 
Medley and Orzell 12626-85 (pers. herb.) [possible hybrid 
specimen]. Madison Co.: 3 May 1977, T. Fargue s.n. (UARK); 1 
May 1985, Orzell and M. Medley 1803 (UARK). Montgomery Co.: 
28 Apr 1985, Orzell and M. Medley 1758 (UARK); 29 Apr 1985, 
Orzell and M. Medley 1781 (NYS); 30 Apr 1985, Orzell and M. 
Medley 1785 (SEL); 29 Apr 1985, Orzell and M. Medley 1778 
(MO); 8 May 1932, Gardner 265 (OKLA); 20 Aug 1964, J. W. 
Thieret 18194 (USLA); 4 Apr 1979, Plant Taxonomy Class s.n. 
(ATU); 16 May 1966, Demaree 53626 (NLU); 21 May 1977, F. Sloan 
s.n. (ATU). Ouachita Co.: 23 Apr 1987, Orzell and Bridges 5139 
(UARK). Perry Co.: 26 Apr 1951, J. E. Moore 3335 (UCA). Pike 
Co.: 3 Oct 1985, Orzell, Bridges, and A. Higgenbottom 3150 
(UARK); 13 Oct 1977, D. Lawson s.n. (NLU). Polk Co.: 2 May 
1982, L. Magrath 12832 (OCLA - 2); 23 Apr 1955, A. McWilliam 
s.n. (UARK); Apr 1962, D. M. Moore 62-022 (APCR); 2 May 1964, 
D. M. Moore s.n. (APCR). Pulaski Co.: 9 May 1985, Orzell, 
Bridges, and L. Peacock 1804 (NYS); 28 Apr 1985, F. Lane 101 
(UARK). Saline Co.: 28 Apr 1985, F. Lane 103 (UARK). Sevier 
Co.: 11 Oct 1985, Orzell, Bridges, and L. Peacock 3437 (UARK). 
Yell Co.: 6 May 1951, J. E. Moore 510242 (UARK). 

In addition to the above the following specimens have been 
annotated by Max Medley (DHL) and Dr. John Atwood (SEL) as 
Cypripedium kentuckiense: St. Francis Co., 19 Sept 1979, F. 
Deneke 1443 (MEM); Newton Co., 6 Aug 1974, R. L. Thompson 692 
(NLU, UARK); however we believe these collections to represent 
Cypripedium pubescens Willd. 


This southeastern member of the Cypripedium pubescens 


1987 Orzell & Bridges, Flora of Arkansas 101 


complex was recently described by Reed (1981) based on 
collections from Kentucky. Over the next few years, it was 
recognized that this plant also occurred in Tennessee, Alabama, 
Mississippi, Louisiana, Texas, Oklahoma, and Arkansas (Atwood 
1985). Atwood also notes that the number of herbarium specimens 
of this species indicate that it may be most abundant in Arkansas. 
The senior author first noted the occurrence of this plant in 
Arkangas when first visiting one of the Garland County sites in 
May 1984. Independently, in the same month, the junior author 
learned of the occurrence of this plant in Arkansas through Larry 
Lowman of Wynne, Arkansas, who had 4 cultivated specimen taken 
from the Crawford County population. At this time, the only 
states to which this species had been definitely attributed were 
Kentucky and Tennessee. By 1985 we had discovered several 
additional localities for C. kentuckiense in Arkansas, and in April 
of 1985 the senior author led Max Medley on a systematic survey 
of the Arkansas sites. Full documentation of the ecology and 
status of C. kentuckiense in Arkansas will be provided in a 
manuscript in preparation by the authors. Two other yellow lady’s 
slippers are still recognized as occurring in parts of Arkanaas, C. 
pubescens Willd. and C. parviflorum Salisb. 

It is important to note that this species, although newly 
described as such, represents an entity long known to 
orchidologists. Correll (1940, 1950) knew of this plant and 
included its distinctive characters within his description of C. 
calceolus L. var. pubescens (Willd.) Correll. In Correll (1950), 
Wherry considered it the "southeastern" ecological entity of the C. 
calceolus complex. Published photographs of yellow lady’s slippers 
from Louisiana (Brown 1972; Dorman 1959; Duffy 1970) and Texas 
(Ajilvsgi 1979, 1983; Loughmiller 1984; Peacock 1980) have all been 
what is now called C. kentuckiense. We present these citations in 
order to unequivocally document the presence of C. kentuckiense 
in Arkansas, where it is more common than in any other part of 
its range, and more common than any other Cypripedium in 
Arkansas. 


DALEA COMPACTA Spreng. var. PUBESCENS (Gray) Barneby 
(Fabaceae) Hempstead Co.: low ridges, open woods, P.O. McNab, 
elev 340 ft., 20 Jul 1961, Demaree 44664 (SMU). 


This variety was listed for Little River County, Arkansas by 
Smith (1978) as Petalostemum pulcherrimum (Heller) Heller. The 
specimen upon which this record was based (Moore and Iltis 442, 
UARK, WIS), was cited by Barneby (1977) as representing D. 
compacta var. compacta. Smith, in an atlas supplement (1980), 
therefore deleted P. pulcherrimum from the Arkansas flora. We 
uncovered the Demaree specimen while annotating Arkansas 
collections at SMU, and determined it to be correctly identified. 
Specimens originally identified as this taxon at APCR from Little 
River County (Tucker 16113, APCR) were found to be var. 
compacta. Arkansas was not included in the range given for this 


102 Pil YT O27 Li07G 1's Vol. 64, No. 2 
taxon by Wemple (1970) or Barneby (1977). 


DALEA VILLOSA (Nutt.) Spreng. var. GRISEA (T. & G.) 
Barneby (Fabaceae). Miller Co.: Sec. 33, T18S, R28W, 17 Jun 
1985, Orzell and Bridges 2146 (APCR, NY, SMU, UARK); Sec. 17, 
T18S, R28W, 17 Jun 1985, Orzell and Bridges 2148 (TEX, UARK), 
17 Jun 1980, R. Davis and G. Tucker 2570 (APCR). 


Both of our collections are from areas of deep Briley loamy 
fine sand (Arenic Paleudults). Dalea villosa var. grisea is 
occasional in a sandhill woodland dominated by Quercus incana, Q@. 
margaretta, and Pinus echinata, with Thelesperma filifolium, Liatrie 
elegans (Walt.) Michx., Cassia fasciculata, Helianthemum 
georgianum Chapman, and Selaginella arenicola ssp. riddellii. 
Associates along a sandy roadside through a partially cleared 
sandhill woodland include Astragalus soxmaniorum, Chrysopsis 
pilosa Nutt., Heterotheca latifolia Buckl., Clematis reticulata Walt., 
Haplopappus divaricatus (Nutt.) Gray, Asclepias amplexicaulis Sm., 
Eragrostis secundiflora Presl., Pediomelum digitatum (Nutt. ex T. & 
G.) Isley, and Rhus copallina L. 

Demaree (1943) lists Petalostemon villosum Nutt. for 
Arkansas, apparently based on the earlier report by Buchholz and 
Palmer (1926) of a collection (Palmer 5994, MO) near Cotter in 
Marion County. We have not seen this specimen, but it would 
represent a significant disjunction of var. villosa to Arkansas. 
Smith (1978) excludes P. villosum from the Arkansas flora, and 
this species as well as P. griseum T. & G. are not recorded for 
Arkansas by Wemple (1970) or Barneby (1977). The Davis and 
Tucker specimen was in bud, and uncovered in the unidentified 
Dalea collections at APCR annotated by the authors. These 
collections are apparently the first of this variety for Arkansas. 


DELPHINIUM CAROLINIANUM Walt. subsp. PENARDII (Huth) 
Warnock (Ranunculaceae). Hempstead Co.: Sec. 5, T12S, R27W, 15 
May 1985, Orzell and Bridges 1855 (SMU, UARK), 19 May 1986, 
Bridges 86-55 (MO, TEX). Little River Co.: White Cliffs, 29 May 
1940, E. Brinkley 715 (UARK); Sec. 32, T12S, R32W, 16 May 1980, 
R. Davis and G. Tucker 2085 (APCR), 28 May 1976, G. Tucker 
16174 (APCR). Sevier Co.: Sec. 25, T11S, R29W, 28 Apr 1981, R. 
Davis and J. Rettig 2969 (APCR). 


Delphinium carolinianum subsp. penardii is occasional on a 
dry hillside blackland prairie dominated by Schizachyrium 
scoparium (Michx.) Nash. Other associates include Sorghastrum 
nutans (L.) Nash, Hedyotis nigricans, Dalea purpurea Vent., 
Grindelia lanceolata Nutt., Bothriochloa saccharoides (Sw.) Rydb., 
Brickellia eupatorioides (L.) Shinners, and Penstemon cobaea Nutt. 

This taxon was first reported for Arkansas (as D. penardii 
Huth) by Buchholz and Palmer (1926), from Carroll County (Palmer 
5585, MO). Demaree (1943) lists D. virescens Nutt., now 
considered to be a later synonym of D. penardii (Warnock 1981), 


1987 Orzell & Bridges, Flora of Arkansas 103 


for Arkansas. Smith (1978) excludes D. virescens from Arkansas, 
stating "Arkansas reports probably based on pale flowered forms of 
D. carolinianum". This is undoubtedly true for most reports, and 
no Arkansas collections of this subspecies were noted by Warnock 
(1981). However, he did not examine the specimens cited at 

UARK or APCR. We found that specimens of the D. carolinianum 
complex from the dry blackland prairies and chalk outcrops of the 
Cretaceous region of southwestern Arkansas match the vegetative 
characters noted by Warnock (1981) as distinguishing this 
subspecies, although their flower color can vary from white to 
pale blue. Specimens from elsewhere in the state lack the 
distinctive foliage of this subspecies. 


DELPHINIUM CAROLINIANUM Walt. subsp. VIMINEUM (D. 
Don) Warnock (Ranunculaceae). Miller Co.: Sec. 17, T18S, R28W, 
17 May 1985, Orzell and Bridges 1896 (MO, SMU, UARK); Sec 36, 
T18S, R28W, 18 May 1986, Bridges 86-43 (SMU, UARK); Sec 3, 
T19S, R28W, 18 May 1986, Bridges 86-44 (MO, TEX). Nevada Co.: 
Sec. 11, T11S, R20W, 14 Jun 1977, J. Roberts 342 (UARK). 
Ouachita Co.: Sec. 25, T11S, R19W, 20 Jun 1985, Orzell and 
Bridges 2215 (UARK). 


All Arkansas collections are on deep sand in sandhill 
woodlands dominated by Quercus incana, with Selaginella arenicola 
ssp. riddellii, Helianthemum georgianum, and Cnidoscolus texanus 
(Muell. - Arg.) Small. Other Ouachita County associates include 
Cassia fasciculata, and Eriogonum multiflorum Benth. in a disturbed 
sandhill, while Thelesperma filifolium, Bouteloua hirsuta Lag. and 
Streptanthus hyacinthoides Hook. are associates at the relatively 
undisturbed Miller County sandhill woodland. 

Smith (1978) includes this taxon (as D. vimineum D. Don) in 
his possible additions to the flora of Arkansas, on the basis of its 
having been listed as "probably in La. and s.w. Ark." by Correll 
and Johnston (1970). Warnock (1981) did not include Arkansas in 
the range of this subspecies; however, he did not consult 
collections at UARK. These collections are the first of this taxon 
for Arkansas, and represent a slight range extension from adjacent 
northeast Texas. 


DELPHINIUM NEWTONIANUM D. M. Moore (Ranunculaceae). 
Pike Co.: Sec. 10, T5S, R25W, 30 Jun 1985, Albert Higgenbottom 
s.n. (UARK). 


This very distinctive Delphinium has long been considered as 
narrowly endemic to an area of about 1500 sq. km. in four 
contiguous counties (Newton, Searcy, Johnson, and Pope) in the 
Boston Mountains of the Ozark Plateau (Moore 1939, Tucker 1984b, 
Kral 1983, Orzell and Bridges, in prep.). This collection extends 
its range about 160 km south to the southern edge of the central 
Ouachita Mountains. Here it occurs in dry-mesic rocky forests 
similar to those supporting the largest Ozark populations. The 


104 P) HyWeT. OF (0-G AT A Vol. 64, No. 2 


canopy is dominated by Quercus alba, with Carya tomentosa, 
Quercus rubra L., @. stellata, Acer rubrum, Liquidambar 
styraciflua, and Pinus echinata. The subcanopy is dominated by 
Cornus florida L., and the most common species in the herb layer 
are Toxicodendron radicans and Desmodium nudiflorum (L.) DC. 
Small chert-like fragments of novaculite rubble cover 20-30% of 
the ground surface. Mr. Higgenbottom, who so kindly sent us the 
specimen, indicated that the species is quite common in several of 
the nearby ravines, and visible from logging roads in late June. 


DROSERA CAPILLARIS Poir. (Droseraceae). Calhoun Co.: 
Sec. 23, T14S, R13W, 30 Jun 1985, Orzell and Carl Amason 2300 
(SMU); Sec. 1, T14S, R15W, 7 Jul 1985, Orzell, Bridges, and Car! 
Amason 2402 (UARK); Sec. 23, T14S, R13W, 7 Jul 1985, Orzell, 
Bridges, and Carl Amason 2412 (MO, TEX, UARK). 


All the Arkansas collections are from the nearly level 
Quaternary terrace deposits east of the Ouachita River in southern 
Calhoun County. Drosera capillaris may be locally abundant on 
exposed seasonally saturated fine sandy loam where the common 
associates are Lycopodium appressum (Chapman) Lloyd and 
Underwood, Sphagnum moss, and Polypremum procumbens L. In 
one graminoid dominated area adjoining a seep forest, associates 
were Eriocaulon decangulare L., Cynoctonum sessilifolium (Walt.) 
Jaume St. Hil, Polygala cruciata, Xyris difformis Chapman var. 
curtissii (Malme) Kral, X. torta, and X. jupicai L.C. Rich. 

This species was first reported for Arkansas by Branner and 
Coville (1891); however, they did not report D. brevifolia Pursh (or 
any synonym of this species), which is much more common in the 
state. Demaree (1943) and Correll and Johnston (1970) also list 
this species for Arkansas, presumably based on the previous 
report. Arkansas is not included in the range of D. capillaris 
given by Wynne (1944), Wood (1960, 1966), or Shinners (1962). 
Smith (1978) excluded this species, since "Wood (1960) does not 
mention Arkansas in the range of this species, and I believe that 
reports of it for the state are probably based on D. brevifolia.” 
We agree with Smith and believe that our collections are the first 
authentic material of this species for the state, which differs from 
D. brevifolia in its glabrous scapes and papillose seeds. 


ECHINACEA SANGUINEA Nutt. (Asteraceae). Miller Co.: 
Sec. 6, T20S, R27W, 18 May 1985, Orzell and Bridges 1911 (SMU, 
UARK), 17 Jun 1985, Orzell and Bridges 2142 (TEX). 


This species was listed for Arkansas by McGregor (1968) and 
Smith (1978) based only on a specimen from Miller County (Eggert 
s.n., MO) collected in 1898. We found it to be locally common in 
the southern part of Miller County, in both sandy and gravelly- 
clayey soils. Apparently no collections were made in Arkansas in 
the 85 years between the Eggert collection and ours, and this 
species had been considered as possibly extirpated from Arkansas. 


1987 Orzell & Bridges, Flora of Arkansas 105 


GEOCARPON MINIMUM Mackenzie (Caryophylilacese). 
Franklin Co.: Sec. 10, T7N, R28W, 20 Apr 1986, Orzell and 
Bridges 4135 (MO, UARK). 


This federally listed threatened species (Norquist 1986) was 
found on a saline soil barrens developed on soile of the Wing 
series (Aquic Natrustalfs). The entire area is dominated by low 
herbs, with microtopographic variation from high, better drained 
mounds, flats of intermediate elevation, to low, wet, almost barren 
"slick spots" of the exposed natric subsoil horizon. Geocarpon 
occurs near the lower edges of the flats of intermediate elevation, 
often with high cover of lichens and Nostoc sp. Associates of 
Geocarpon include Coreopsis grandiflora Hogg ex Sweet, Ambrosia 
bidentata Michx., Croton capitatus Michx., Nothoscordum bivalve 
(L.) Britt., Diodia teres, Oenothera linifolia Nutt., Juncus tenuis 
Willd., Talinum parviflorum Nutt. ex Torr. & Gray, Sagina 
decumbens (Ell.) Torr. & Gray, Hedyotis australis W. H. Lewis & D. 
M. Moore, Erigeron tenuis Torr. & Gray, Astragalus distortus var. 
distortus, Neptunia lutea (Leavenworth) Benth., Aristida sp. and 
Sporobolus sp. 

Geocarpon minimum has long been known to occur in 
Arkansas only in the southeastern Coastal Plain, in Bradley, 
Cleveland, and Drew Counties (Smith 1978, Rettig 1983). It also 
has been previously reported on Channel Sands (Pennsylvanian age) 
glades in six southwestern Missouri counties (Morgan 1980). This 
collection is the first for the Arkansas River valley section, of the 
Ouachita Province. Previous to this collection, the Arkansas and 
Missouri ranges of this species were separated by 390 km; thie 
intervening site is 180 km south of the nearest Missouri sites and 
230 km northwest of the nearest Arkansas sites. A complete 
inventory of all areas mapped as Wing soil in western Arkansas 
(with the exception of one area in the target area for heavy 
artillery training at Fort Chaffee, in which no human has set foot 
for 40 years!), conducted from April 20-23, 1986, revealed no 
additional sites for Geocarpon. This inventory should be repeated 
at an earlier date in the growing season in a wetter year, as most 
of the plants seen at the Franklin County site were dead and 
drying by this date due probably in part to droughty weather 
conditions in 1986. 


JUNCUS SUBCAUDATUS (Engelm.) Coville and Blake 
(Juncaceae). Sharp Co.: Sec. 7, T18N, R4W, 23 Oct 1984, Orzell 
and Bridges 1559 (UARK). 


Juncus subcaudatus is scattered along narrow streamsides of 
Rock Creek, a calcareous spring-fed stream, and in adjacent 
calcareous streamside seep fens. Associates include Lysimachia 
quadriflora Sims, Parnassia grandifolia, Pycnanthemum virginianum 
(L.) Durand and Jackson, Rhynchospora capillacea Torr., Scleria 
verticillata Muhl. ex Willd., Solidago riddellii, Aster novae—angliae 
L., Polypremum procumbens, and Hypericum sphaerocarpon Michx. 


106 PREAAOT (00 12 OPGF TA Vol. 64, No. 2 


Smith (1978) includes this species in his possible additions, 
stating "based on its distribution in Steyermark (1963), perhaps 
rare in extreme N-central Arkansas". Our collection confirms this 
prediction, and this species is probably restricted to fens in 
Arkansas. 


LASALLEA SERICEA (Vent.) Greene subsp. SERICEA (Aster 
sericeus Vent.) (Asteraceae). Newton Co.: Sec 29, T17N, R20W, 
28 Jun 1985, Orzell and Bridges 2233 (UARK). 


This species (as Aster sericeus) was first reported for 
Arkansas by Demaree (1943), but listed as a “possible addition" by 
Smith (1978). We found the Demaree specimen on which his 
report was apparently based (16 Oct 1943, Demaree 24786, SMU), 
originally identified as A. sericeus, but annotated as A. pratensis 
Raf. by B. Lipscomb in 1979 and J. C. Semple in 1980; therefore 
Demaree first collected in Arkansas the taxon we are considering 
as L. sericea subsp. pratensis (Raf.) Semple & Brouillet. This 
taxon was reinstated as new to the Arkansas flora by Sundell 
(1983) as A. pratensis, and it is locally abundant on saline soil 
barrens in southeastern Arkansas. Lasallea sericea ssp. sericea 
was first discovered in Arkansas by Paul Nelson of the Missouri 
Department of Natural Resources while on a natural area workshop 
field trip to Devil’s Knob-Devil’s Backbone State Natural Area in 
Izard County, and this record was reported by Rettig (1982). Our 
site for this taxon is the second for the state, and indicates the 
potential for many additional sites on calcareous glades in the 
intervening and adjacent counties. Our nomenclature for this 
group is following Semple and Brouillet (1980). 


LILIUM SUPERBUM L. (Liliaceae). Logan Co.: Magazine 
Mountain, near top - south end, 29 Jun 1952, D. M. Moore 520773 
(SMU, UARK). Pope Co.: Sec. 5 & 6, TION, R20W, 24 Jun 1985, 
Orzell and Bridges 2224 (MO). Stone Co.: Vicinity of Cole 
Fork/Stewarts confluence (on Stewarts Fork). Very rare. 14 Apr 
1981, R. Davis 2914 (APCR). 


At the Pope County station a colony of robust plants occurs 
in a canopy opening along a lower north-facing slope dominated 
by Fraxinus americana L., Acer saccharum Marsh, Carya 
cordiformis, Juglans nigra, and Liquidambar styraciflua. The 
understory and shrub layers include Acer saccharum, Cornus 
florida, Hamamelis virginiana L., Lindera benzoin, Dirca palustris 
L., and Halesia carolina L. Herb layer associates include 
Delphinium newtonianum, Carex albursina Sheldon, Asarum 
canadense L., Phryma leptostachya, Adiantum pedatum, and 
Desmodium glutinosum (Muhl. ex Willd.) Wood. 

This species was first reported for Arkansas by Nuttall (1835) 
and listed by Branner and Coville (1891) on the basis of this 
report. Buchholz and Palmer (1926) report this species from 
Washington Co., but no specimen was cited. Demaree (1943) also 


1987 Orzell & Bridges, Flora of Arkansas 107 


listed this species for the state, but Smith (1978) includes this 
taxon under L. michiganense Farw. as "L. superbum” of auth., not 
L." The history of knowledge of these taxa in the midwestern 
United States is complicated by the relatively late date of 
description of L. michiganense (1915) and L. canadense L. ssp. 
editorum (Fern.) Wherry (1943), previous to which these taxa were 
commonly called L. superbum, and the slow and inconsistent 
recognition of these as distinct taxa (cf. Hull 1942, 1943, Gleason 
1952, Boivin and Cody 1956). More recent works (Wherry 1947, 
Mohlenbrock 1962, Adams and Dress 1982) maintain all of these as 
distinct taxa, but report only L. michiganense as occurring in 
Arkansas. We have critically examined the above cited specimens 
and conclude that true L. superbum does occur very rarely in 
Arkansas, often in very mesic areas known to support other 
primarily Appalachian species. Our specimen has the glabrous leaf 
margins and veins, ribbed sepal midribs, and large medifixed 
anthers characteristic of this species, and the other specimens 
(the Davis specimen is vegetative and agrees with L. superbum in 
those characters) examined have characters consistent with these. 


LITHOSPERMUM TUBEROSUM Rugel ex DC. (Boraginaceae). 
Hempstead Co.: Sec. 24, T12S, R27W, 2 Apr 1985, Orzell and 
Bridges 1652 (UARK, SMU). Howard Co.: Sec. 12., T8S, R29W, 2 
Apr 1985, Orzell and Bridges 1655 (UARK). Sevier Co.: Sec. 14, 
T10S, R29W, 3 Apr 1985, Orzell and Bridges 1658 (TEX, UARK); 
Sec. 32, T10S, R29W, 8 Oct 1985, Orzell, Bridges, L. Peacock, and 
T. Foti 3314 (MO). 


Lithospermum tuberosum is a characteristic plant of dry- 
mesic calcareous ravine forests on Cretaceous strata in the 
southwestern Coastal Plain of the state. These forests are 
dominated by Quercus muhilenberglii, Q@. shumardii, Juglans nigra, 
Acer saccharum, Carya ovata (P. Mill.) K. Koch, and C. 
myristiciformis (Michx. f.) Nutt., in contrast to the surrounding 
mixed pine-hardwood forests with Pinus taeda. These calcareous 
ravine forests have a lush, species-rich vernal flora, in a region 
intrinsically depauperate in this floristic element. Herbaceous 
associates of L. tuberosum include Phlox pilosa L., Osmorhiza 
longistylis, Sanguinaria canadensis L., Trillium recurvatum Beck, 
Uvularia sessilifolia, Solidago auriculata, and Phryma leptostachya. 

This species is not listed in Smith (1978) nor in any other 
work on the flora of Arkansas. We believe our collections to be 
the first for the state, representing a short range disjunction from 
northcentral and northeast Louisiana. 


LUDWIGIA MICROCARPA Michx. (Onagraceae). Sharp Co.: 
Sec. 7, T18N, R4W, 25 Oct 1985, Orzel] and Bridges 3628 (UARK, 
SMU). 


Ludwigia microcarpa is common in calcareous seep fens on a 
gravelly marly substrate saturated by minerotrophic seepage. 


108 PeHaYATIO0E ONGIT A Vol. 64, No. 2 


Associates include Scleria verticillata, Rhynchospora capillacea, R. 
capitellata (Michx.) Vahl., Parnassia grandifolia, Lysimachia 
quadiflora, Selaginella apoda, Solidago riddellii, Cynoctonum 
mitreola (L.) Britt., and Fuirena simplex Vahl. 

Smith (1978) includes this species in his possible additions, 
stating "based on the distribution given in Steyermark (1963), 
should be in Arkansas." Actually, this species is known only from 
two sites in Missouri (Sharon Morgan, pers comm), and is very 
rare in Louisiana (Peng 1982), Tennessee (Peng 1982; Milo Guthrie, 
pers. comm.), and Texas (Orzell and Bridges, in prep.). 


LYCOPODIUM CAROLINIANUM L. (Lycopodiaceae). Calhoun 
Co.: Sec 27, T14S, R13W, 1 Nov 1985, Orzell and Bridges 3648 
(MIL, MO, UARK). 


Lycopodium carolinianum has been found at a single station 
on wet exposed fine sandy loam with Aletris farinosa L., 
Cynoctonum sessilifolium, scattered Sphagnum moss, RAynchospora 
inexpansa, R. rariflora (Michx.) Ell., R. gracilenta Gray, 

Lycopodium appressum, Anthaenantia rufa (Ell.) Schultes, Xyris 
baldwiniana Schultes, Aristida longespica Poir., and Drosera 
capillaris. Nearby associates include Eryngium intergrifolium Walt., 
Platanthera ciliaris (L.) Lindl., Rhynchospora plumosa Ell., Myrica 
cerifera L., and Eriocaulon decangulare. 

This species is not listed for Arkansas by Smith (1978) or 
Taylor (1984), and we believe our collection to be the first for the 
state. This record represents a disjunct of about 220 km from the 
more continuous range in the longleaf pine belt of central 
Louisiana, with only one intervening record in Jackson Parish, 
Louisiana (Thieret 1980). Peck et al. (1987) have also reported 
this new record for Arkansas. Our determination was verified by 
Dr. James Bruce. 


LYCOPODIUM x COPELANDII Eiger (Lycopodiaceae). 
Calhoun Co.: Sec. 1, T14S, R15W, 7 Jul 1985, Orzell, Bridges, and 
Carl Amason 2399 (MIL, MO, TEX, UARK). 


This natural hybrid was found with Platanthera ciliaris, 
Drosera capillaris, and Lycopodium appressum in a shallow roadside 
ditch adjacent to a seep forest. When we made this collection, we 
could not locate one of the putative parents, L. alopecuroides L., 
at this site or from the state of Arkansas. The nearest 
documented sites for L. alopecuroides are 270 km south in Rapides 
Parish, Louisiana and 270 km east in Attala County, Mississippi 
(Thieret 1980; Evans 1978). This hybrid is known to occur in the 
absence of L. alopecuroides in Kentucky (Cranfill 1980, 1981). 
Hybridization between L. appressum and L. alopecuroides is 
common in mixed populations on the outer Coastal Plain (Thieret 
1980). Later searches of this and nearby sites by Peck et al. 
(1987) resulted in the discovery of not only this hybrid, but also 
L. alopecuroides, L. prostratum, L. x bruceii (L. appressum x L. 


1987) 16 Orzell & Bridges, Flora of Arkansas 109 


prostratum), and L. alopecuroides x L. prostratum. The result is a 
remarkable assemblage of Lycopodium species and hybrids for an 
area about 420 km from the Gulf coast. 


LYGODESMIA JUNCEA (Pursh) Hook. (Asteraceae). Hot 
Springs Co.: Igneous intrusive area, bottoms and foothills, P.O. 
Magnet Cove, elev. 600 ft., 18 Sept 1937, Demaree 16264 (SMU). 


Demaree (1943) reported this species from Arkansas, and 
Vuilleumier (1973) noted its occurrence in the southeastern United 
States "only in the region of Little Rock, Arkansas (fide Tomb)." 
Smith (1978) excluded this species from Arkansas, stating "reported 
from Pulaski Co. by Vuilleumier (1973), probably on the basis of 
waifs." Tomb (1980) cites and maps the above Demaree specimen 
for Arkansas, as long-disjunct from the main range of the species. 
We believe that L. juncea naturally occurred on open, very xeric 
granitic outcrops at Magnet Cove, now almost completely 
destroyed by mining activities, and that this species has probably 
been extirpated from the state. 


OENOTHERA HETEROPHYLLA Spach subsp. HETEROPHYLLA 
(Onagraceae). Miller Co.: Sec 17, T18S, R28W, 18 May 1986, 
Bridges 86-53 (UARK), 17 Jun 1980, G. Tucker 19400 (APCR). 


This species is common in natural openings in sandhiil 
woodlands dominated by Quercus incana, with Dalea phleoides 
(Torr. & Gray) Shinners var. microphylla (Torr. & Gray) Barneby, 
Opuntia humifusa, Eriogonum longifolium Nutt., Helianthemum 
georgianum, Streptanthus hyacinthoides, and Selaginella arenicola 
ssp. riddellii. 

The description of O. heterophylla subsp. orientalis Dietrich, 
Raven & W. L. Wagner (Wagner 1983) from Arkansas and Alabama 
left subsp. heterophylla as occurring from central and eastern 
Texas to eastern Louisiana. Subsp. orientalis occurs in Calhoun, 
Nevada, and Ouachita counties in Arkansas, whereas we have 
found subsp. heterophylla only west of the Red River in Miller 
County, where it is contiguous with the range of this subspecies. 
Our specimen has the spreading, longer sepal tips and red 
pustulate hairs characteristic of this subspecies (Wagner 1983). 
We have not seen the Tucker specimen, but presume it to be the 
same taxon as it was collected from the same site. 


OENOTHERA PILOSELLA Raf. subsp. SESSILIS (Pennell) 
Straley (Onagraceae). Lafayette Co.: Sec. 25, T16S, R24W, 19 
May 1985, Orzell and Bridges 1925 (MO, UARK). 


The Lafayette County site for this taxon is a fragipan 
flatwoods forest dominated by Quercus phellos. Herbaceous 
associates include Tradescantia occidentalis (Britt.) Smyth and 


Carex spp. 
Almost all known extant sites for this subspecies are on 


110 Poy TO> BOF G TA Vol. 64, No. 2 


prairie remnants in the Grand Prairie region of the Mississippi 
Alluvial Plain of eastern Arkansas, in Arkansas, Prairie, and 
Lonoke counties (Tucker 1984a), and recently St. Francis County. 
It is also known from single historical collections in Louisiana and 
Texas. This collection represents a very different habitat for this 
taxon, and greatly increases our knowledge of its area and habitat 
of potential occurrence. 


PASPALUM PRAECOX Walter (Poaceae). Calhoun Co.: Sec. 
22, T14S, R13W, 30 Jun 1985, Orzell and Carl Amason 2292 (TEX); 
1 Nov 1985, Orzell and Bridges 3646A (SMU, UARK). 


This species is common in a open graminoid dominated area 
with Fuirena bushii, Polygala cruciata, Xyris ambigua, X. torta, 
Rhynchospora glomerata, R. inexpansa, R. globularis, Rhexie 
mariana, Sphagnum moss, Gelsemium sempervirens, Asclepias 
longifolia ssp. longifolia, Gratiola pilosa, Sabatia gentianoides, 
Dichanthelium scoparium, Ludwigia linearis, Hypericum lobocarpum 
Gattinger, and EFupatorium rotundifolium. 

Smith and Lipscomb (1975) reported this species from Chicot 
County, but Smith (1978) later excluded this species, stating it to 
be based on a misidentification of P. pubiflorum Rupr. ex Fourn. 
We have not seen the specimen for this report. Our collections 
represent the first authentic material of this species from 
Arkansas, and a slight range extension northward from Louisiana. 


PEDIOMELUM DIGITATUM (Nutt. ex T. & G.) Isley 
(Psoralea digitata Nutt. ex T. & G.) (Fabaceae). Miller Co.: Sec. 
33, T18S, R28W, 17 May 1985, Orzell and Bridges 1902 (SMU, 
UARK), 25 Jun 1980, Davis and Kral 2689 (APCR), Kral 65502 
(VDB). County uncertain: "Arkansas, common,” 23 Jul 1894, B. 
Bush 67 (NY, US). 


Pediomelum digitatum is occasional along a sandy roadside, 
through a former sandhill woodland that is now a cleared field. 
Associates include Astragalus soxmaniorum, Chrysopsis. pilosa, 
Heterotheca latifolia, Clematis reticulata, Haplopappus divaricatus, 
Asclepias amplexicaulis, Eragrostis secundiflora, and Dalea villosa 
var. grisea. 

Pediomelum digitatum was first reported for Arkansas by 
Branner and Coville (1891), and included in the Arkansas flora by 
Demaree (1943). An early collection of P. digitatum was examined 
by the authors, and based on what we know of Bush’s itinerary, 
could have been collected in Miller County. A Leavenworth 
collection from Arkansas at NY could well have been collected in 
Oklahoma. Smith (1978) listed Psoralea digitata var. digitata as a 
possible addition to the flora of Arkansas, stating "reported for 
Arkansas by Demaree (1943) and Correll and Johnston (1970), 
perhaps in west or southwest Arkansas". Davis (1981) reported P. 
digitata var. digitata from Arkansas, based on his collection cited 
above. Shinners (1951) described Psoralea digitata var. parvifolia 


1987 Orzell & Bridges, Flora of Arkansas 111 


Shinners as a narrow-leaved variety occurring only in eastern 
Texas, therefore necessitating the above references to var. 
digitata. Smith (1978) does not mention this variety, but by 
giving a variety name alludes to its existence. Our collection and 
the specimens cited by Davis (1981) are actually all referable to 
var. parvifolia. James Grimes (pers. comm.) believes var. parvifolia 
to intergrade with var. digitata, and perhaps not worthy of 
recognition. However, the specimens we have seen from areas of 
sandy soil in eastern Texas, adjacent Louisiana and southwestern 
Arkansas mostly retain the characters Shinners used to distinguish 
this variety. We agree that it probably does not represent a valid 
subspecific taxon, but, nevertheless is the ecological form of this 
species in Arkansas. 


PEDIOMELUM HYPOGAEUM (T. & G.) Rydberg subsp. 
SCAPOSUM (Gray) Ockendon (Psoralea hypogaea Nutt. ex T. & G. 
var. scaposa Gray) (Fabaceae). Miller Co.: Sec 17, T18N, R28W, 
18 May 1986, Bridges 86-48 (UARK). 


This species was rare in openings in a sandhill woodland 
dominated by Quercus incana. Herbaceous associates include 
Selaginella arenicola ssp. riddellii, Thelesperma filifolium, Bouteloua 
hirsuta, Scutellaria cardiophylla Engelm. & Gray, and Berlandiera x 
betonicifolia. Our specimen was determined by James Grimes at 
TEX, who found that it had the glands, size, and leaflet shape of 
P. hypogaeum but the inflorescence and pubescence of P. 
subulatum (Bush) Rydberg. 

This unusual record was not listed for Arkansas by Smith 
(1978) or any other list for the state. P. hypogaeum ssp. 
scaposum ranges from central to north-central Texas, where it 
commonly occurs on limestone outcrops. It was considered as 
endemic to Texas by Turner (1959) and Ockendon (1965). Our 
collection is unusual both in being disjunct 300 km from the 
nearest locality, and in occurring on deep sand, a habitat more 
commonly associated with the more western ssp. hypogaeum. This 
collection, and several additional collections we have made of this 
variety in eastern Texas, suggest that P. hypogaeum ssp. 
hypogaeum, P. hypogaeum ssp. scaposum, and P. subulatum may all 
be part of the same wide-ranging species complex, with the 
variation within the group tied to habitat conditions. 


PEDIOMELUM SUBULATUM (Bush) Rydberg (Fabaceae) 
(Psoralea subulata Bush). Miller Co.: Sec. 7, T20S, R27W, 18 May 
1985, Orzell and Bridges 1910 (MO, SMU, TEX, UARK). Nevada 
Co.: Near Bluff City, in a field, 12 May 1984, H. D. Moore s.n. 
(UARK). 


At our collection site, this species was rare in a sandhill 
woodland on Sparta sand overlying the Weches formation (Eocene). 
The canopy is dominated by Quercus velutina Lam., Q. margaretta, 
and @. incana. Associates of Pediomelum subulatum include 


112 PH NenvOrL OnG St A Vol. 64, No. 2 


Pteridium aquilinum (L.) Kuhn, Tradescantia reverchonii, Rhus 
aromatica, Cnidoscolus texanus, Aristolochia reticulata Jacq., 
Callicarpa americana L., and Castanea pumila (L.) P. Mill. 

This species was not included in the Arkansas flora by Smith 
(1978) or any other source. Psoralea subulata was considered as 
endemic to Texas by Ockendon (1965), although Turner (1959) 
noted that it occurred in Louisiana and possibly adjacent 
Oklahoma. Waterfall (1972) reports it from two counties in 
central Oklahoma, and we have collected this species on deep sand 
in Atoka County in southeastern Oklahoma. Lagseigne (1973) 
reports P. subulata from only Natchitoches Parish in Louisiana. 
The Nevada Co. specimen at UARK had been identified as P. 
esculenta Pursh until our annotation. 


POGONIA OPHIOGLOSSOIDES (L.) Ker. (Orchidaceae). Fulton 
Co.: Sec. 7, T20N, R8W, 23 May 1985, Orzell and Bridges 1938 
(MO, SMU, TEX, UARK). 


This species was common in two open graminoid dominated 
fens in this section, particularly on Osmunda regalia L. hummocks 
within the fens. Herbaceous associates include Sphagnum sp., 
Dulichium arundinaceum (L.) Britt., Juncus effusus L., Eupatorium 
perfoliatum L., Andropogon gerardii Vitman, Rhexia mariana, 
Ludwigia alternifolia, Dichanthelium scoparium, Aster lateriflorus 
(L.) Britt., Senecio aureus, Coreopsis tripteris L., Pedicularis 
lanceolata Michx., Carex suberecta, C. lurida Wahlenb., and C. 
interior. Over 200 individuals were counted at one of the sites, 
several with two flowers per plant. 

These two colonies represent the second and third extant 
sites for this species in Arkansas, the other being in Jefferson 
County, with a historical record from Saline County. This also is 
the second reported site for the Interior Highlands, and the first 
in this region of Arkansas. The only other site for P. 
ophioglossoides in the Interior Highlands is a fen on the Salem 
Plateau, in Reynolds County, Missouri (Steyermark 1963; Orzell 
1983). 


PORTULACA UMBRATICOLA H.B.K. (Portulacaceae). Monroe 
Co.: Sec. 18, TIN, R2W, 6 Nov 1985, Orzell 3653 (UARK), 7 Nov 
1985, Orzell 3663 (MO, NCU, NLU, SMU, TEX, UARK). 


Portulaca umbraticola was common along the edge of a 
harvested soybean field on a moist deep sandy upland, with Cassia 
obtusifolia L., Chamaesyce maculata (L.) Small, Eleusine indica (L.) 
Gaertn., Campsis radicans (L.) Seem. ex Bureau, and Digitaria 
sanguinalis (L.) Scop. 

This species has not been previously reported for Arkansas 
by Smith (1978) or any other source. It has long been known 
from the southeastern United States as P. coronata Small, 
considered to be synonymous with the widespread southwestern P. 
umbraticola (Matthews and Levins 1985). We consider this species 


$9677) Orzell & Bridges, Flora of Arkansas BES 


to be questionably native in Arkansas; it does occur in a region 
naturally having somewhat droughty deep eolian sand deposits, and 
it could have been a part of the native flora. However, ample 
opportunity has existed for it to have been introduced from 
further southwest in this highly disturbed region. 


PRUNUS GRACILIS Engelm. and Gray (Rosaceae). Miller 
Co.: Sec. 6, T20S, R27W, 18 May 1985, Orzell and Bridges 1919 
(MO, NLU, SMU, TEX, UARK). 


Prunus gracilis is locally abundant at the edge of a cleared 
sandhill woodland (now planted in pines). Associates include 
Quercus margaretta, Pteridium aquilinum, Tradescantia reverchonii, 
Rhus copallina, Cnidoscolus texanus, Alophia drummondii (Graham) 
R.C. Foster, Chionanthus virginicus L., Astragalus soxmaniorum, 
Stillingia sylvatica Garden ex L., Hymenopappus artemisiifolius, 
Tetragonotheca Iudoviciana (Torr. and Gray) Gray, and Tephrosia 
virginiana (L.) Pers. 

This species was listed for Arkansas by Correll and Johnston 
(1970) and Robertson (1974). Tucker (1976) did not include this 
species in the flora, but noted that it was very likely in the 
southwest corner of the state. Smith (1978) included it in possible 
additions, based on the above reports. We believe our collection 
to be the first of this species from Arkansas, and that the 
previous reports for Arkansas were likely based on specimens 
collected in present-day Oklahoma. 


RHYNCHOSPORA COLORATA (L.) Pfeiffer (Dichromena 
colorata (L.) A. H. Hitchc.) (Cyperaceae). Little River Co.: Sec. 
4, T13S, R32W, 13 Aug 1985, Orzell and Bridges 2726 (APCR, MO, 
NCU, SMU, TEX, UARK). 


Our new site is a vigorous and dense stand of thousands of 
plants in a disturbed, artificially open wet swale in a shallow 
drainage. Associated species include Rhynchospora caduca Ell., 
Cynoctonum mitreola, Polypremum procumbens, Lythrum alatum 
Pursh var. lanceolatum (Eli.) Torr. & Gray, Eupatorium perfoliatum, 
Desmanthus illinoensis (Michx.) MacM. ex B. L. Robins. & Fern., 
Diodia virginiana L., Setaria geniculata (Lam.) Beauv., and 
Axonopus furcatus (Flugge) A. S. Hitchc. 

This species has long been known to occur in Arkansas only 
on the basis of specimens collected by Demaree in Bradley County 
between 1937 and 1939 [Demaree 15044 (F, MO, SMU, NY), 

Demaree 19264 (GH-2, MO, NY, SMU, UARK), Demaree 19268 

(SMU), Demaree 19269 (MICH, NLU, NY-2, TEX-5, UARK)]. 

Despite the relatively specific site information and the efforts of 
numerous botanists to relocate this conspicuous species, it has not 
been seen at the Bradley County site since 1939. Many of the 
labels of the Demaree collections state that the species was "not 
common", and each sheet contains numerous entire plants. We 
feel that R. colorata was overcollected to the point of local 


114 PHY eT Oe OuGat 2A Vol. 64, No. 2 


extirpation by 1939, and until our recent collection, was generally 
accepted as extirpated from Arkansas. 


RHYNCHOSPORA MICROCARPA Baldw. ex Gray 
(Cyperacese). Calhoun Co.: Sec. 34, T14S, R13W, 14 Jun 1985, 
Orzell and Bridges 2118 (SMU, TEX, UARK). 


Rhynchospora microcarpa is common in a wet cutover Pinus 
taeda flatwoods where it grows in association with Rhynchospora 
inexpansa, R. corniculata (Lam.) Gray, R. glomerata, R. globularis, 
Xyris laxifolia Mart. var. iridifolia (Chapman) Kral ined., Diodia 
virginiana, and Myrica heterophylla Raf. 

This remarkable find has not been previously reported for 
Arkansas, and is disjunct over 220 km from the nearest isolated 
locality in LaSalle Parish and over 420 km from the more 
continuous range in extreme southern Louisiana (Joyce 1974). We 
have found this species to occur typically near the boundary 
between the coastal prairie and freshwater marsh regions of 
southwestern Louisiana. This is by far the most inland record for 
this outer Coastal Plain species. Our determination was verified 
by Dr. Wm. Wayt Thomas at the New York Botanical Garden. 


RHYNCHOSPORA PLUMOSA Ell. (Cyperaceae). Calhoun Co.: 
Sec. 27, T14S, R13W, 3 Aug 1985, Orzell and Carl Amason 2628 
(UARK). 


Rhynchospora plumosa is common in a wet swale with Aster 
umbellatus P. Mill., Eupatorium rotundifolium, Rhynchospora 
inexpansa, R. globularis, R. glomerata, R. gracilenta, Lycopodium 
appressum, Helianthus angustifolius L., Pluchea foetida (L.) DC., 
Eryngium integrifolium, Crotonopsis elliptica Willd., Gelsemium 
sempervirens, and Eriocaulon decangulare. 

This very distinctive sedge was not listed for Arkansas by 
Smith (1978) or any other source. Our collection is the first for 
Arkansas and is disjunct 210 km from the nearest locations in the 
longleaf pine belt of central Louisiana (Joyce 1974). 


SABATIA GENTIANOIDES Ell. (Gentianaceae). Calhoun Co.: 
Sec. 22, T14S, R13W, 30 Jun 1985, Orzell and Carl Amason 2281 
(APCR); Sec. 23, T14S, R13W, 6 Jul 1985, Orzell and Bridges 2378 
(UARK, SMU). 


At both Calhoun County stations, Sabatia gentianoides thrives 
in open areas with Fuirena bushii, Polygala cruciata, Rhynchospora 
inexpansa, R. glomerata, Eupatorium rotundifolium, and 
Dichanthelium scoparium. Other associates which occur at one of 
the two sites include Xyris ambigua, X. torta, Paspalum praecox, 
Ludwigia linearis, Hypericum lobocarpum, Polypremum procumbens, 
Cynoctonum sessilifolium, Rhynchospora rariflora, Hypericum 
hypericoides, Crotonopsis elliptica, and Myrica cerifera. 

Sabatia gentianoides was first found in Arkansas by Carl 


1987 Orzell & Bridges, Flora of Arkansas 115 


Amason, who called the authors when it flowered in 1985 and led 
the senior author to the site that day. This species was not 
listed for Arkansas by Smith (1978) and the state was not included 
in its range by Wilbur (1955) or Perry (1969). We believe our 
collections to be the first for the state, and disjunct over i170 km 
from the more continuous range in the longleaf pine belt of 
central Louisiana. 


SANGUISORBA ANNUA Nutt. (Rosaceae). Sebastian Co.: 
Sec. 16, T4N, R32W, 21 Apr 1986, Orzell and Bridges 4202 (MO, 
NCU, SMU, TEX, UARK). 


This species was found in a disturbed saline soil barrens 
(Wing soil series: Aquic Natrustalfs) in the prairie region of the 
western Arkansas River valley near Hartford. Associated species 
included Nemophila phacelioides Nutt. ex W. Bart., Delphinium 
carolinianum, Solanum elaeagnifolium Cav., and Camassia scilloides 
(Raf.) Cory. 

This species has long been inwii.ded in the Arkansas flora 
(Lesquereux 1860, Branner and Coville 1891, Demaree 1943, Correll 
and Johnston 1970, Robertson 1974), likely all on the basis of the 
original Nuttall collections probably made in present-day Oklahoma. 
A specimen at GH, collected by Leavenworth in Arkansas, is part 
of a sheet which Gray designated as the type, and another 
specimen at GH is labeled "Ark. Mts., Douglas". Again, both of 
these were likely to have been collected in Oklahoma. We are 
considering our collection to be the first from within the present 
borders of Arkansas. 


SCLERIA RETICULARIS Michx. (Cyperaceae). Calhoun Co.: 
Sec. 23, T14S, R13W, 6 Jul 1985, Orzgell and Bridges 2380 (TEX, 
UARK). Fulton Co.: Sec. 7, T20N, R8W, 2 Oct 1984, Orzell 1434 
(UARK), Orzell 1439 (SMU, TEX). 


In Fulton County, S. reticularis was found in two 
minerotrophic seeps, one with quaking sphagnous peat 
accumulation, on the Salem Plateau, associated with Rudbeckia 
fulgida var. umbrosa, Dichanthelium scoparium, Parnassia 
grandifolia, Pedicularis lanceolata, Carex lurida, Rhynchospora 
capitellata, Senecio aureus, and Xyris torta. The Calhoun County 
site is in an open clearcut recently planted in Pinus taeda on the 
almost level Quaternary terrace desposits east of the Ouachita 
River, in the Gulf Coastal Plain, associated with Aletris farinosa, 
Polygala cruciata, Cynoctonum sessilifolium, Polypremum 
procumbens, Rhynchospora rariflora, Fuirena bushii, and Sabatia 
gentianoides. 

Scleria reticularis was listed for Arkansas by Nuttall (1835), 
Branner and Coville (1891), and Demaree (1943), but was excluded 
from the Arkansas flora by Smith (1978), based upon its 
distribution as shown by Fairey (1967). Our specimens have the 
characteristics of var. pubescens Britt. (S. muhlenbergii Steud., S. 


116 PHY, 2.0) LO, G TCA Vol. 64, No. 2 


setacea Poir.), but we believe this form to be of little taxonomic 
merit. 


SEBASTIANA FRUTICOSA (Bartr.) Fern. (Euphorbiaceae). 
Lafayette Co.: Sec. 23, T17S, R24W, 7 Oct 1985, Orzell, Bridges, 
and Peacock 3218 (NCU, SMU, UARK); Sec. 14, T17S, R24W, 7 Oct 
1985, Orzell, Bridges, and Peacock 3247 (TEX). 


This shrub was first discovered in Arkansas by Lance 
Peacock, who brought it to our attention. It is rather abundant 
along Bodcau Creek on the lower slopes of the uplands bordering 
the stream floodplain, often adjacent to seepage areas, and 
occasionally on infrequently flooded stream terraces. Typical 
overstory associates include Liquidambar styraciflua and Quercus 
nigra L., sometimes also with Nyssa sylvatica or Taxodium 
distichum (L.) L.C. Rich. 

This species was not listed for Arkansas by Smith (1978) or 
any other source. It is a typical species of the Coastal Plain west 
to southeast Texas, and we believe our collections to be the first 
for Arkansas. 


SOLIDAGO BICOLOR L. (Asteraceae). Garland Co.: 
Weyerhauser Forest, rocky disturbed woods, P.O. Mountain Pine, 
elev. 650 ft., 26 Oct 1971, Demaree 64307 (SMU). 


This species was reported for Arkansas by Branner and 
Coville (1891), but they did not list its much more common 
relative, S. hispida Muhl. ex Willd. It was listed as occurring in 
the Ouachita mountains of Arkansas by Cronquist (1980), but Smith 
(1978) referred Arkansas reports of S. bicolor to S. hispida. We 
have examined the Demaree specimen and found it to be true S. 
bicolor. Cronquist (1980) states that this species hybridizes 
extensively with S. hispida and S. erecta Pursh, but retains its 
populational identity over large areas. Perhaps in the western 
part of its range, pure S. bicolor is quite rare, since S. hispida is 
quite abundant in the Interior Highlands. 


SOLIDAGO ULMIFOLIA Muhl. var. MICROPHYLLA Gray. 
(Asteraceae). Garland Co.: 11 Aug 1937, Demaree 15622A (SMU), 
Demaree 16662A (sic) (SMU). Logan Co.: 25 Aug 1937, Demaree 
16052 (SMU). Polk Co.: 12 Aug 1937, Demaree 15712 (SMU); 19 
Aug 1937, Demaree 15877 (SMU-2); 10 Sep 1953, Demaree 34229 
(SMU); 26 Oct 1966, Demaree 54970 (SMU); 18 Sep 1954, A. 
McWilliam s.n. (UARK-2); 12 Oct 1952, D. M. Moore and H. H. IItis 
520823 (UARK); 8 Sep 1954, D. M. Moore 54266 (SMU); 19 Aug 
1969, Gary Morton 3988 (SMU). 


Although Solidago ulmifolia is quite common throughout 
Arkansas, confusion exists as to the varieties present in the state. 
Smith (1978) includes S. delicatula Small in synonymy under S. 
ulmifolia var. palmeri Cronq., and S. ulmifolia var. microphylla in 


1987 Orzell & Bridges, Flora of Arkansas 117 


synonymy under S. ulmifolia var. ulmifolia. In contrast, Cronquist 
(1980) lists as occurring in Arkansas the taxa S. delicatula, S. 
ulmifolia var. palmeri, and S. ulmifolia var. ulmifolia, with no 
additional synonymy. Taylor and Taylor (1984) list S. ulmifolia 
var. microphylla as the common variety of the species in Oklahoma 
and Texas, and include S. delicatula and S. helleri Small in its 
synonymy. We concur with the nomenclature and synonymy of 
Taylor and Taylor (1984) for this group, and have reexamined all 
Arkansas specimens of S. ulmifolia at SMU and UARK (103 sheets) 
using several published descriptions and keys to determine the 
taxa. We found specimens of var. microphylla, var. palmeri, and 
var. ulmifolia in the state, with distinct geographic separation. 
Solidago ulmifolia var. microphylla reaches its eastern limit in the 
Ouachita Mountains of west-central Arkansas, from the above 

cited counties. Solidago ulmifolia var. palmeri, described twice 
from Arkansas and Alabama specimens by Cronquist (1947, 1955), is 
common throughout all but the extreme western part of the 
Ouachita Mountains, and occurs occasionally in the Ozark Plateaus. 
Solidago ulmifolia var. ulmifolia is the only variety in the Coastal 
Plain and Mississippi Alluvial Plain sections of the state, and 
occasionally reaches slightly into the Interior Highlands. Some of 
the specimens cited were first determined as S. ulmifolia var. 
microphylla by C. E. Taylor. 


SPIRANTHES ODORATA (Nutt.) Lindl. (Spiranthes cernua (L.) 
L.C. Rich. var. odorata (Nutt.) Correll) (Orchidaceae). 
Cleveland Co.: Sec. 10 and 11, T11S, R12W, 19 Oct 1985, Orzell, 
Bridges, and Peacock 3480 (MO, NCU, NLU, NYS, SMU, TEX, 
UARK). 


This species was first found in Arkansas by Lance Peacock, 
who brought it to our attention. The population consists of 
several hundred plants, forming stoloniferous colonies over a five 
acre area, growing in shallow standing water of a backswamp 
dominated by scattered Taxodium distichum, Liquidambar 
styraciflua, and Quercus lyrata over a dense subcanopy of Fraxinus 
caroliniana P. Mill. Common associates of S. odorata include 
Panicum gymnocarpon Ell., Aster vimineus Lam. var. subdumosus 
Wieg., Justicia ovata (Walt.) Lindau var. lanceolata (Champman) 
R.W. Long, Rhynchospora corniculata, Carex joorii Bailey, C. 
gigantea Rudge, Saururus cernuus L., and Tradescantia occidentalis. 

Our specimen was verified by Dr. Charles Sheviak at the New 
York State Museum. He states that two other Arkansas specimens 
we sent him for verification [Hot Springs Co.: 20 Oct 1923, Palmer 
24239 (UARK); Union Co.: n.d., D. M. Moore s.n. (UARK)] can be 
referred to S. cernua, but with a genetic contribution from S. 
odorata, which is to be expected. An additional collection, Orzell, 
Bridges, and Peacock 3277 from Miller County, was stated by 
Sheviak to perhaps be depauperate S. odorata, but material was 
insufficient for proper determination. This species was not listed 
for Arkansas by Smith (1978) or Sheviak (1982), and our collection 


118 POHUY  TiOrLrONG I yA Vol. 64, No. 2 


seems to be the first for the state, and a slight range extension 
from northern Louisiana. 


SPIRANTHES PRAECOX (Walt.) S. Wats. (Orchidaceae). 
Calhoun Co.: Sec. 32, T14S, R13W, 20 May 1985, Orzell and 
Bridges 1932 (UARK); 26 May 1978, J. Roberts 800 (UARK). Hot 
Spring Co.: low places, 9 Jul 1896, H. Eggert s.n. (MO). 


Scattered plants of Spiranthes praecox were found growing in 
a saline soil barrens, with Schoenolirion wrightii Sherman, 
Ambrosia bidentata, and Fimbristylis puberula (Michx.) Vahl ex 
Small & Britt. The particular barren developed on the Bonn soil 
series (fine-silty, mixed, thermic Glossic Natraqualf). 

Spiranthes praecox was first reported for Arkansas by 
Branner and Coville (1891), on the basis of a Grand Prairie report 
by Harvey; however, they did not include the much more common 
S. vernalis Engelm. & Gray. Spiranthes praecox had been reported 
for Arkansas by Moore (1965) on basis of misidentified Spiranthes 
vernalis. This orchid was listed for Arkansas by Demaree (1943), 
and Arkansas was included in the ranges given by Correll and 
Johnston (1970) and Correll (1950). Smith (1978) considered it 
under possible additions, "perhaps in south Arkansas.” We have 
not seen the Eggert specimen from Hot Spring County. Our 
specimens and those of J. Roberts were determined by Dr. Charles 
Sheviak at the New York State Museum. 


TETRAGONOTHECA LUDOVICIANA (Torr. & Gray) Gray 
(Asteraceae). Miller Co.: Sec. 6 and 7, T20S, R27W, 18 May 1985, 
Orzell and Bridges 1914 (UARK, SMU, MO, TEX). 


Tetragonotheca ludoviciana is locally abundant at the edge of 
a cleared sandhill woodland (now planted in pine) and occassional 
in openings in more intact sandhill woodland. Associates include 
Quercus incana, Pteridium aquilinum, Tradescantia reverchonil, 
Rhus copallina, Cnidoscolus texanus, Aristolochia reticulata, 
Chionanthus virginicus, Schizachyrium scoparium, Stillingia 
sylvatica, Hymenopappus artemisiifolius, Rhynchosia latifolia Nutt. 
ex Torr. and Gray, and Tephrosia virginiana. 

Tetragonotheca ludoviciana was not included in the Arkansas 
flora by Smith (1978) or any previous checklist. Lesquereux (1860) 
reported T. helianthoides L., a more eastern species, for Arkansas, 
but Coville thought this was probably a mistake, and had no idea 
of the true identity of this report (Branner and Coville 1891). fT. 
ludoviciana was known only from eastern Texas and Caddo Parish, 
Louisiana by Turner and Dawson (1980). It has since been found 
in Allen and Sabine parishes in Louisiana (Nelwyn Gilmore, pers. 
comm.). 


TRADESCANTIA BRACTEATA Small (Commelinaceae). Scott 
Co.: Sec. 30, T4N, R30W, 23 Apr 1986, Orzell and Bridges 4268 
(TEX, UARK). Sebastian Co.: Sec. 34, T4N, R32W, 21 Apr 1986, 


1987 Orzell & Bridges, Flora of Arkansas 119 


Orzell and Bridges 4217 (TEX, UARK); Sec. 35, T4N, R32W, 21 Apr 
1986, Orzell and Bridges 4228 (MO, TEX, UARK). 


Tradescantia bracteata was found in rocky, siliceous, dry- 
mesic to mesic north-facing upper slope and ravine forests near 
the top of Poteau Mountain, at elevations from 500 to 800 meters. 
Associated species include Trillium viridescens Nutt., Smilacina 
racemosa (L.) Desf., Uvularia grandiflora Sm., Thelypteris 
hexagonoptera (Michx.) Weatherby, Camassia scilloides, 
Porteranthus stipulatus, Erythronium rostratum, and Carex 
ouachitana. 

No mention is made of this species in Smith (1978) or any 
Atlas supplements. Anderson and Woodson (1935) show it as 
occurring south to central Missouri and southern Kansas, primarily 
in prairies. McGregor et al. (1977) extend the range to 
northeastern Oklahoma, including Delaware, Adair, and Sequoyah 
counties on the Arkansas border. MacRoberts (1980a) reports 
having seen specimens of T. bracteata from McCurtain County, 
Oklahoma at DUR and TAES, and having collected plants 
"resembling" this species in central Texas. All of our specimens 
are from within 40 miles of Sequoyah County, and seem to be the 
first verified reports of this species in Arkansas, and perhaps at 
the southeastern limit and the highest elevations for the species. 


TRADESCANTIA VIRGINIANA L. (Commelinaceae). Arkansas 
Co.: Sec. 4, T5S, R3W, 14 Apr 1986, Orzell and Bridges 4076 
(LSUS, MO, NCU, SMU, TEX, UARK). 


This remarkable record was found in a very rare and 
significant habitat, a rich, mesic hardwood slope forest along a 
small stream well within the Mississippi Alluvial Plain. The 
canopy trees include Carya ovata, C. cordiformis, C. laciniosa 
(Michx. f.) Loud., Fraxinus americana, Quercus falcata Michx., q. 
nigra, Q. texana Buckl., Q. michauxii Nutt., Celtis laevigata Willd., 
and Ulmus americana L. Much of the site has a dense subcanopy 
of Asimina triloba (L.) Dunal, Cercis canadensis L., and Staphylea 
trifolia L. The herbaceous layer is extremely rich, including the 
first records of Solidago auriculata, Uvularia sessilifolia, and 
Smilax lasioneuron Hook. for the Mississippi Alluvial Plain of 
Arkansas, as well as other species rare in the region, such as 
Dentaria laciniata, Erythronium albidum Nutt., Sanguinaria 
canadensis, Trillium recurvatum, and Viola pubescens Ait. 

This species was listed by Smith (1978) as a "possible 
addition," and he states that “much Arkansas material has been 
determined this. Most such material is misidentified and 
represents putative hybrids." Deneke (Deneke 1981; Deneke and 
Browne 1987) listed this species for St. Francis County, and Smith 
(1982) included this report in Atlas Supplement IV. However, in 
Atlas Supplement V, Smith (1986) again deleted 7. virginiana from 
the Arkansas flora, stating that "all material at UARK that had 
been determined this apparently represents T. occidentalis or 


120 PHY Poms OGL A Vol. 64, No. 2 


hybrid material." Since Deneke had sent a duplicate to UARK 
(Deneke 534), his report was apparently erroneous. 

The taxonomy and identification of Tradescantia in the 
south-central United States is notoriously difficult, and it is with 
great trepidation that we report our records of this genus. As far 
as T. virginiana is concerned, MacRoberts (1980a) shows its clear 
distinction from T. hirsutiflora Bush, and does not include it is 
the Louisiana flora (MacRoberts 1980b). Our specimens have the 
very sparse pubescence and thicker roots characteristic of T. 
virginiana. The nearest approach to Arkansas of this species of 
the northeastern and north-central states is in southeastern 
Missouri and middle Tennessee. These two new Tradescantia 
species for Arkansas bring the total number of taxa recognized for 
this genus in the state to twelve, equaling the number in Texas 
and more than any other state. 


TRAGIA SMALLIT Shinners (Euphorbiaceae). Miller Co.: 
Sec. 17, T18S, R28W, 17 May 1985, Orzell and Bridges 1897 
(TEX, UARK); 17 Jun 1985, Orzell and Bridges 2149 (APCR), 12 
Aug 1985, Orzell and Bridges 2690 (MO, SMU). 


In Miller County this plant is common in a sandhill woodland 
on Briley loamy fine sand dominated by Quercus incana, Q. 
margaretta, Pinus echinata, and Cornus florida. Associates of 
Tragia smallii include Pteridium aquilinum, Vitis rotundifolia 
Michx., Clematis reticulata, Cnidoscolus texanus, Opuntia humifusa, 
and Aristolochia reticulata. Tragia smallii is algo occassional in 
natural sandhill openings with Matelea cynanchoides (Engelm.) 
Woods. and Tradescantia reverchonii. 

This species was not included in the Arkansas flora by Smith 
(1978) or any other source. The ranges given for this species by 
Shinners (1956) and Miller and Webster (1967) confine it to the 
longleaf pine belt of the Gulf Coastal Plain. These collections are 
the northernmost for the species, and disjunct about 170 km from 
the nearest locations in central Louisiana. We have found this 
species to be rather common in the dry longleaf pine savannahs of 
southeastern Texas. Our determinations were verified by Robert 
Kral at Vanderbilt University. 


XYRIS DIFFORMIS Chapm. var. CURTISSII (Malme) Kral 
(Xyridaceae). Calhoun Co.: Sec 29, T14S, R13W, 11 Jun 1985, 
Orzell and Bridges 2060 (UARK, VDB), 17 Jul 1981, R. Kral 67522 
(VDB); Sec 23, T14S, R13W, 30 Jun 1985, Orzell and Carl Amason 
2298 (VDB). 


Xyris difformis var. curtissii and X. baldwiniana are the 
rarest of the seven Xyris species now known from Arkansas. All 
the Arkansas collections of Xyris difformis var. curtissii are from 
nearly level Quaternary deposits east of the Ouachita River in the 
West Gulf Coastal Plain of Calhoun County. Both sites are in 
open graminoid dominated areas adjacent to seep forests with 


1987 Orzell & Bridges, Flora of Arkansas 12 


Magnolia virginiana L., Pinus taeda, and Acer rubrum. Associated 
species at these sites include Rhynchospora rariflora, R. inexpansa, 
Eriocaulon decangulare, Utricularia subulata L., Lycopodium 
appressum, Drosera capillaris, and at one site Eriocaulon 
kornickianum van Huerck and Muell.-Arg. 

This species was not listed for Arkansas by Smith (1978), and 
Arkansas was not included in its range by Kral (1966). The above 
collections are the first for the state. Specimen 2060 was verified 
by Robert Kral at Vanderbilt University. 


XYRIS DIFFORMIS Chapm. var. DIFFORMIS (Cyperaceae). 
Clark Co.: Sec. 30, T9S, R21W, 15 Aug 1985, Orzell and Bridges 
2820 (VDB), 12 Oct 1985, Orzell, Bridges, and Peacock 3464 (SMU, 
TEX, UARK). Hot Springs Co.: 3.9 mi NW Malvern on US 67, 
sphagnous sandy seepage area by highway, 3 Sep 1967, R. Kral 
29174 (VDB). Ouachita Co.: Margins of Bragg Lake, Bragg City, 
P.O. Chidester, 13 Sept 1964, Demaree 51311 (VDB). Perry Co.: 
Common in gravel and brush of river bed, Big Maumelle River, 
near the bridge on Hwy 9, 0.35 mi S of Williams Junction (ject. 
Hwys 9 & 10), 27 Jul 1977, E. B. Smith 3227 (LSU). 


Our collection of Xyris difformis var. difformia is from a 
roadside ditch saturated by seepage from an adjacent slope. 
Associated plants include Carex longii, Rhexia mariana, R. 
virginica, Gratiola pilosa, Ludwigia alternifolia, Lobelia puberula, 
Eupatorium rotundifolium, Arthraxon hispidus, Rhynchospora 
inexpansa, Panicum verrucosum, Juncus sp., Eleocharis sp. and 
Sphagnum moss. 

The taxonomy and nomenclature of this species was quite 
confused before the revision of the genus by Kral (1966); 
therefore it is difficult to interpret the nature of earlier reports. 
Branner and Coville (1891) and Demaree (1943) report X. torte and 
X. caroliniana Walt. for Arkansas. The name X. caroliniana was 
often misapplied during this period to the species now known as 
X. difformis (e.g. Small 1933), but we have not seen a specimen 
upon which this report could have been based. Xyris difformis is 
not listed for Arkansas by Smith (1978) or Kral (1966), and we 
believe the above cited specimens to be the first ‘or the state. 
All the above specimens were determined or verified by Robert 
Kral. 

EXCLUDED SPECIES 


In addition to the new and additional records reported for 
Arkansas in the above accounts, some deletions of species 
previously reported for the Arkansas flora and nomenclatural 
changes need to be made. Smith (1983) reported Cypripedium 
candidum Muhl. ex Willd. (S. L. Timme 2434, UARK) from Benton 
County. The specimen upon which this record was based has been 
determined to be a white-lipped specimen of a normally yellow- 
lipped taxon, probably C. parviflorum Salisb., by the authors and 
Dr. Charles Sheviak. We have also collected small white-lipped 


122 PHYTOLOGIA Vol. 64, No. 2 


Cypripediums similar to this one in Sharp County. 

Lithospermum ultiflorum Torr. ex Gray in Arkansas was 
based in Smith (1978) on an Jltis 5156 specimen (UARK), which 
when examined by the authors proved to be an unusually small, 
late season, fruiting specimen of L. incisum Lehm. Later 
collections originally identified as L. multiflorum at APCR (Davis 
s.n., Davis and Rettig 2843) were all found to be misidentifications 
of L. incisum and L. carolinense. Lithospermum multiflorum is a 
far western species very unlikely for Arkansas, and should be 
deleted from the flora. 

Rhynchospora chalarocephala Fern. & Gale was included in 
Smith’s Atlas Supplement V (1986) on the basis of a collection 
from Cleveland County (MW. M. Shepherd and M. Schiffl 243, 

UARK). The specimen was tentatively determined by E. B. Smith 
and subsequently verfied by the authors and by W. W. Thomas 
(NY). However, as part of a critical study of Rhynchospora in the 
West Gulf Coastal Plain, we have reexamined this specimen and 
determined it to be an immature R. glomerata. The specimen has 
underdeveloped second florets in each spikelet, which rules out R. 
chalarocephala. We find specimens of the highly variable R. 
glomerate to often be misidentified as the one-fruited species R. 
chalarocephala and R. cephalanthe Gray, or as the more northern 
R. capitellate (Michx.) Vahl. 

Several subspecific taxa in Smith (1978) were examined by 
the authors to see if they were in fact good representations of 
these taxa in the state. All Phlox pilosa L. subsp. fulgida 
(Wherry) Wherry collections for Arkansas at UARK had been 
determined by J. R. Gibson at UNC in 1983 to represent subsp. 
pilosa, the most common subspecies in the state, and we 
concurred. We found the Hite 164 and Smith 3568 collections of 
Teucrium canadense L. var. occidentale (Gray) McClintock & Epling 
at UARK to differ in several characters from good var. occidentale 
and best referred to var. virginianum (L.) Eat., the only variety of 
this species in the state. Despite much taxonomic and 
nomenclatural confusion, we found no good distinctive characters 
for the presence of the northern Viola pubescens Ait. var. 
pubescens in Arkansas, the Moore 410149 specimen at UARK is 
var. eriocarpa (Schwein.) Russell. This taxon is poorly 
differentiated from var. lJeiocarpa (Fern. & Wieg.) Seymour, and it 
seems best to us to refer all stemmed yellow violets in Arkansas 
to one taxon, leaving the choice of the proper name to rangewide 
population studies, with the possibility that only one good taxon 
may emerge from this complex. 


PHYTOGEOGRAPHICAL PATTERNS 


In reporting native species new or rediscovered in a state, it 
is important to assess the significance of the new records relative 
to the total range of the species, patterns of habitat distribution, 
and statue of the species in adjacent states. By analysis of this 
information in relation to distribution patterns of the known flora, 


1987 Orzell & Bridges, Flora of Arkansas 123 


hypotheses can be made as to the origin and relative significance 
of these new records. Since many of the species considered in 
this paper exemplify several recurring general distribution 
patterns, they will be discussed in groups of related patterns. 

Very little has been written on the general phytogeography 
of Arkansas, and some background is needed before discussing the 
rather unusual distribution patterns of the species reported here. 
The state lies near the western boundary of the Eastern Deciduous 
Forest, and includes several of the sections described by Braun 
(1950). The northwestern half of the state is within the Interior 
Highlands section of the Southern Division of the Oak-Hickory 
Forest Region. This area is characterized by sedimentary rocks of 
Pennsylvanian to Ordovician age, and includes both siliceous and 
calcareous strata, with calcareous strata more common in the 
Salem and Springfield Plateaus and siliceous strata in the Boston 
and Ouachita Mountains, although some outcrops of almost all rock 
types of the region can be found in most sections. The eastern 
part of the state is within the Mississippi Alluvial Plain section of 
the Southeastern Evergreen Forest Region. Actually, this is 4 
misnomer since this section has very few vegetation types 
dominated by evergreen trees (a few outlying stands of loblolly 
pine do occur). The natural vegetation is a mosaic of wetland 
deciduous hardwood forests, sandy upland deciduous forests, 
graminoid dominated upland fragipan barrens (=Grand Prairie of 
Arkansas), and floodplain swamps. This entire region (with the 
exception of Crowley’s Ridge, a Tertiary Coastal Plain outlier) is 
the Quaternary floodplain of the Mississippi River and its major 
tributaries. The southern part of the state is within the Gulf 
Slope Section of the Oak-Pine Forest Region. This region is 
underlain by sands and sandy clays of Eocene age, and generally 
has a characteristic Upper Coastal Plain vegetation and flora, with 
mixed pine-hardwood forests (primarily Pinus taeda, Liquidambar 
styraciflua, and Quercus falcata) dominating most of the uplands. 
A small area in the southwestern part of the state is included in 
the Forest-Prairie Transition Area of the Southern Division of the 
Oak-Hickory Forest Region. This area is developed on heavy clays 
interspersed with gravelly sands of Cretaceous age, and had a 
natural vegetation of bands of blackland prairie and calcareous 
ravine forests alternating with forests more like those of the 
Eocene region of the Coastal Plain. 

Major floristic elements in the state include the general 
Eastern Deciduous Forest element, found more or less throughout 
the state, the Northern or Upland Eastern Deciduous Forest 
element, centered to the north and northeast of Arkansas, and 
generally most common in the Interior Highlands within the state, 
and the Southeastern Coastal Plain element, most common in the 
Mississippi Alluvial Plain and Coastal Plain. All of these could be 
considered as intraneous elements, that is, consisting of species 
found throughout much of the state and tending not to reach their 
range limits within its borders. The patterns in the following 
discussion tend to represent extraneous elements, that is, those 


124 PHHOYeT OP Ly OFG? TA Vol. 64, No. 2 


including species which are disjunct to or reach their range limits 
in some direction within the state. Rare species and new state 
records tend to exhibit extraneous distribution patterns, and 
understanding these is critical to comprehending the significance 
of their records. The remainder of this section will discuss these 
extraneous patterns in relation to the species reported in this 
paper. References will be given to published detailed range 
descriptions or maps for these species, but the bulk of the 
patterns presented are based on unpublished range maps compiled 
by the authors. Other rare or restricted species in Arkansas 
exhibiting the same general patterns are noted. No attempt is 
made to provide a full historical phytogeographic interpretation of 
these patterns; rather, they are presented in a descriptive manner 
as they are currently known, to stimulate ideas on their origin 
and historical phytogeographical significance. 

Some records included here are new, shortly disjunct, records 
for endemic or near-endemic species of the physiographic sections 
represented in Arkansas. These include Delphinium newtonianum, 
Carex ouachitana, Oenothera pilosella ssp. sessilis, and Geocarpon 
minimum (Figure 1). While the distance between the previous 
records and those reported here is generally less than 250 km, 
these species have such narrow ranges that these are extremely 
important records which may be significant in understanding their 
origin and migration patterns. 

A few of the species reported in this paper generally range 
to the north of Arkansas, being most common in the northeastern 
to north central United States and adjacent Canada. Two of 
these, Carex interior (Reznicek and Ball 1980) and C. suberecta 
(Figure 2), are found in fens or minerotrophic seepage areas 
resembling fens in Arkansas and Missouri (Orzell and Kurz 1986) 
and throughout much of their range. Both of these species are 
rare south of the glacial boundary, with the exception of a 
secondary center of distribution in the Salem Plateau of the Ozark 
Uplift. Several other species rare in Arkansas exhibit this same 
pattern, notably Spiranthes lucida (H. H. Eaton) Ames, 
Rhynchospora capillacea, Lysimachia quadriflora, Solidago riddeliii, 
and Pedicularis lanceolata (Orzell et al. 1985), all of which are 
found with or nearby these species. The Arkansas locations are 
the southernmost known for these carices, and are contiguous with 
the disjunct secondary center of distribution in south central 
Missouri. 

Several of the species reported here are most common in the 
prairie and prairie-forest border regions of the north central 
United States, and reach their southeastern range limit in 
northwestern Arkansas, although they may extend south to 
Oklahoma and Texas at their southwestern limits. Among these 
are Carex bicknellii var. bicknellii (Figure 3), Carex molesta, and 
Tradescantia bracteata. Other species which are rare in Arkansas 
and tend to display this pattern include Helianthus rigidus (Cass.) 
Desf., Muhlenbergia bushii Pohl, Amorpha canescens Pursh, 
Androsace occidentalis Pursh, Gentiana puberulenta J. Pringle, and 


1987 


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Figure 1. Range of Geocarpon minimum. Figure 2. Range of 
Carex suberecta. Figure 3. Southwestern part of range of Carex 
bicknellii var. bicknellii (dots) and total range of Carex bicknellii 
var. opaca (triangles). Scales of all figures in paper are expressed 
in kilometers. 


126 POH XY 1.0L OrG TA Vol. 64, No. 2 


Sagittaria rigida Pursh. In Arkansas, all of these species have the 
majority of their occurrences in counties which at least 
historically supported some tallgrass prairie vegetation, although 
now many survive only on rocky glades and disturbed areas within 
their former range in the state. 

A generally distributed north central wetland species reaching 
its southern limit in the Mississippi Alluvial Plain of northeastern 
Arkansas is Carex muskingumensis (Figure 4). This sedge tends to 
reach its range limits in the alluvial plains of major rivers, 
including the Mississippi, Des Moines (Gilly 1946), and Cumberland. 
The only other southeastern state where this species occurs is 
Tennessee, where it is listed by the Tennessee Heritage Program 
as a species of special concern. 

Some species reported here are generally found in the forest 
regions of the north-central states east to the Appalachian 
provinces. These include Aster shortii and Carex sparganioides 
(Figure 5), both of which reach their southwestern limit in 
Arkansas. Numerous rare species in Arkansas display similar 
patterns, including many of more restricted occurrence or longer 
disjunctions in the state. Examples with related patterns include 
Orbexilum onobrychis (Nutt.) Rydb., Trillium flexipes Raf., 
Stylophorum diphyilum (Michx.) Nutt., Hydrophyllum macrophyllum 
Nutt., Euonymus obovatus Nutt., Carex careyana Torr. ex Dewey, 
Collinsia verna Nutt., Carex hitchcockiana Dewey, and Cacalia 
muhlenbergii (Schultz-Bip.) Fern. Many of these are calciphilic 
species which are more common in the Interior Low Plateaus and 
glaciated Central Lowlands than in the more siliceous regions of 
the Appalachians. 

Several species are common in both the northeastern and 
north central United States, becoming less common in the western 
and southern portions of their range and reaching their 
southwestern limit in Arkansas. Among these are Carex laxiculmis 
(Figure 6), Solidago bicolor, and Tradescantia virginiana. Other 
species exhibiting this pattern in Arkansas include Dennstaedtia 
punctilobula (Michx.) T. Moore, Thelypteris noveboracensis (L.) 
Nieuwl., Gaylussacia baccata (Wang.) K. Koch, and Waldsteinia 
fragarioides (Michx.) Tratt. These tend to be rare in adjoining 
regions of other states and disjunct to Arkansas. An unusual 
related pattern is exhibited by Juncus subcaudatus (Figure 7), 
which is most common in the Appalachian Mountains from 
Pennsylvania south to North Carolina. It occurs in a few counties 
of Georgia, Tennessee, Kentucky, and Ohio, and then has a 
disjunct secondary center of distribution in the Salem Plateau of 
south central Missouri and adjacent north central Arkansas. Here 
it occurs in similar habitats to those of the northern Carex 
interior, Carex suberecta, and previously mentioned associates 
which are rare or absent from the central and southern 
Appalachians. This disjunct center is 650 km from the main range 
of the species and 320 km from the nearest isolated localities. 

True Appalachian disjunctions are rather rare in Arkansas, 
since most such species can find at least some areas of suitable 


“a 
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vs 


128 PuH,Y>T OL 0sGyI A Vol. 64, No. 2 


habitat in the Interior Low Plateaus or even in the East Gulf 
Coastal Plain. One species which exemplifies this pattern is 
Lilium superbum (Adams and Dress 1982), which is most common in 
the central and southern Appalachians from Pennsylvania to 
Georgia, but extends well onto the Coastal Plain in Georgia, 
Alabama, and Misisisippi, and to the Interior Low Plateaus in 
southern Indiana and Illinois. Other mostly Appalachian disjuncts 
in Arkansas include Fothergilla major (Sims) Lodd. (Tucker, in 
prep.), Melanthium hybridum Walt., Disporum lanuginosum (Michx.) 
Nichols., and Iris verna L., all of which have their only localities 
west of the Mississippi River in Arkansas. 

Few species of restricted southern distribution are found in 
both the southeastern states and in Arkansas. These tend to have 
unusual patterns, but generally are found west and south of the 
Appalachians, sometimes also including areas in the Piedmont, or 
the Appalachian provinces themselves. One of these is Cardamine 
angustata var. multifida (Montgomery 1955), which is centered on 
the Cumberland Plateau and Interior Low Plateaus from southern 
Indiana through Kentucky and Tennessee to Alabama, and extends 
into the Blue Ridge and Piedmont of Georgia with a disjunct area 
in the central Piedmont of North Carolina. The Arkansas and 
Oklahoma localities are at least 600 km from the nearest 
occurrences in Alabama. Another unusual pattern included here is 
that of Cypripedium kentuckiense (Figure 8). This newly 
described, yet long known variant of the yellow lady slipper 
complex, seems to occur in three distinct but isolated areas of the 
southeastern United States. One is part of the Cumberland 
Plateau in Kentucky and Tennessee, where it occurs on sandy 
stream floodplains. The second is within the East Gulf Coastal 
Plain in Alabama and Mississippi, where it is now very rare near 
seepage areas, but was perhaps somewhat more common before 
extensive conversion to monoculture pine plantations. The third 
region includes the Boston Mountains, Ouachita Mountains, and 


‘eastern part of the West Gulf Coastal Plain in southern Arkansas, 


southeastern Oklahoma, Louisiana, and eastern Texas. In this 
region it is almost always found adjacent to seepage areas, and is 
much more abundant in Arkansas than elsewhere in the range, 
although still declining due to land alteration. Another unusual 
southern distribution pattern is that of southern calciphiles, an 
example of which is Lithospermum tuberosum (Figure 9). This 
species is most common in the Interior Low Plateaus and East Gulf 
Coastal Plain, and extends west on calcareous sediments in the 
Mississippi Alluvial Plain and West Gulf Coastal Plain. A similar 
pattern is shown by Solidago auriculata. Other rare or restricted 
species in Arkansas exhibiting unusual southern distribution 

| Patterns include Heuchera villosa Michx. (s.]. incl. H. villosa var. 
“arkansana (Rydb.) E. B. Smith), Philadelphus hirsutus Nutt., Halesia 
searolina, and Schizandra glabra (Bickn.) Rehd. 

A few species reported here are generally species of the 
“southern part of the Eastern Deciduous Forest region. This is a 


~<*“@ommon pattern in the Arkansas flora, and the fact that these are 


1987 Orzell & Bridges, Flora of Arkansas 129 


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Figure 8. Range of Cypripedium kentuckiense. 


130 PH ¥ TO L0G TA Vol. 64, Nora 
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Figure 9. Range of Lithospermum tuberosum. 


"new" for Arkansas indicates that they have previously been 
subsumed into other taxa or overlooked by collectors, rather than 
being rare or restricted in the state. Among these are several of 
the species of Carex reported here, such as C. striatula and C. 
willdenowii. Some other sedges are similarly intraneous, though 
centered more in the south central states, including Carex texensis 
and C. oklahomensis. It is predicted that these species would be 
found, with dilligent collecting, to be rather common and 
widespread in the southern two-thirds of the state. 

Several species reported here are most common on the 
Atlantic and Gulf Coastal Plains, but also occur inland in other 
provinces, having some localities as far north as the Great Lakes 
region. These are part of the Coastal Plain element in the flora 
of the Great Lakes region discussed by Peattie (1922) and others. 
Those reported here include Carex longi, Xyris difformis var. 
difformis (Kral 1966), and Scleria reticularis (Figure 10). All of 
these are rare or unknown in the upland provinces of the 
southeast and midwestern states between the Coastal Plain and the 
Great Lakes region. Another rare species in Arkansas displaying 
this pattern is Eleocharis equisetoides (Ell.) Torr. Other species, 
such as Pogonia ophioglossoides (Figure 11), are common in both 
the Coastal Plain and the boreal forest region south to the Great 
Lakes states, and are rare in the interior states. 


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Figure 10. Western part of range of Scleria reticularis. 
Figure 11. Southwestern part of range of Pogonia ophioglossoides. 


A few species primarily found on the southeastern Coastal 
Plain tend to have ranges extending northward in the Mississippi 
valley of the central United States, or have disjunct localities in 
this region. Among those reported here are Carex louisianica and 
Ludwigia microcarpa. Carex louisianica can be expected to occur 
throughout the Coastal Plain and Mississippi Alluvial Plain sections 
of Arkansas, whereas Ludwigia microcarpa (Figure 12) is disjunct 
to seepage areas near the eastern margin of the Salem Plateau in 
Arkansas and Missouri. Other rare species in Arkansas exhibiting 
variations on this pattern include Carex decomposita Muhl., Scirpus 
divaricatus Ell., and Cynoctonum mitreola. 

The largest number of species reported in this paper have 
ranges which are centered on part or all of the Coastal Plain of 
the southeastern United States, extending sometimes north on the 
Atlantic Coastal Plain, inland into the upland provinces of the 
southeast, or west into the southern Great Plains and adjacent 
regions. These patterns will be discussed in relation to their 
degree and direction of disjunction to Arkansas. The first of 


132 Piney *TaOsLVOuG7 Pea Vol..'64,, ilo. ae 


these includes characteristic species of the Coastal Plain east of 
the Mississippi River which reach their northern limit west of the 
Mississippi in the southern counties of Arkansas, and sometimes 
also southeastern Oklahoma. These include Sebastiana fruticosa, 
Spiranthes odorata (Luer 1975, Sheviak 1982), and Spiranthes 
praecox (Luer 1975). Numerous rare species in Arkansas exhibit 
variations on this pattern, including Sporobolus junceus (Michx.) 
Kunth, Platanthera cristata (Michx.) Lindl., Lycopodium appressum, 
Smilax walteri Pursh, Seymeria cassioides (Walt.) Blake, Polygala 
nana (Michx.) DC., Erynigium integrifolium, Asimina parviflora 
(Michx.) Dunal, Coelorachis rugosa (Nutt.) Nash, Cynoctonum 
sessilifolium, Aletris aurea Walt., Habenaria repens Nutt., Quercus 
incana, Solidago patula Muhl. var. strictula Torr. & Gray, 
Tillandsia usneoides L., Helianthemum georgianum, and 
Helianthemum rosmarinifolium Pursh. 

More significantly, many Coastal Plain species reported here 
for southern Arkansas are apparently disjunct from the longleaf 
pine belt of central Louisiana and the open seepage-bog habitats 
of southeastern and east central Texas. These species often occur 
inland to the fall line in the Atlantic Coastal Plain, and even in 
the upland provinces. They become progressively more restricted 
westward to the outer part of the East Gulf Coastal Plain, and 
west of the Missisippi River are generally absent from the Inner 
Coastal Plain, that area either north of 32 degrees latitude or 
developed on Eocene or earlier strata. Typically, these reach 
their northern limit in Louisiana in Grant and Natchitoches 
parishes, and are disjunct about 200 km to a small area of 
Calhoun County, Arkansas. Among those reported here are 
Asclepias longifolia ssp. longifolia (Woodson 1954), Rhynchospora 
Plumosa (Figure 13), Drosera capillaris (Figure 14), Lycopodium 
carolinianum (Figure 15), L. x copelandii, Paspalum praecox, Xyris 
difformis var. curtissii (Figure 16), and Sabatia gentianoides 
(Wilbur 1955). Other species with similar disjunctions to Calhoun 
County include Xyris baldwiniana, Rhynchospora rariflora, R. 
gracilenta, and Eriocaulon decangulare. Rhynchospora colorata 
(Thomas 1984) is known historically from Bradley County, and now 
is reported for Little River County. One species of the sandy 
uplands of the Longleaf Pine Belt, Tragia smallii (Figure i7), is 
reported here as disjunct to Miller County, Arkansas. Longleaf 
pine belt species disjunct to other southern Arkangas counties 
include Anthaenantia rufa, Dryopteris ludoviciana (Kunze) Small, 
Stewartia malachodendron L., Halesia diptera Ellis, Platanthera 
nivea (Nutt.) Luer, and Persea borbonia (L.) Spreng., the last three 
known only from historical records in Arkansas. 

A few species in Arkansas are disjunct from the extreme 
Outer Coastal Plain, generally isolated inland 400 km from the 
more continuous range. These include Cladium jamaciense (Figure 
18), found only as far inland as Iberville and Calcasieu parishes in 
Louisiana, but occurring far inland in the Edwards Plateau and 
Chihuahuan Desert in Texas, and Rhynchospora microcarpa (Gale 
1944), most common in southern Georgia and Florida, but having 


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Figures 12-17. Western part of ranges of Coastal Plain species. 
12. Ludwigia microcarpa. 13. Rhynchospora plumosa. 
14. Drosera capillaris. 15. Lycopodium carolinianum. 
16. Xyris difformis var. curtissii. 17. Tragia smallii. 


134 Pn Y TO E1006 Tz Vol. 64, No. 2 


isolated localities west to extreme southern Louisiana and the 
coastal bend of Texas, as well as inland in the Carolinas and 
Georgia. Few other rare species in Arkansas exhibit far southern 
patterns, one of which, Lilaeopsis carolinensis Coult. & Rose, is 
known only from a single historical record. 

Numerous records presented here are for taxa more or less 
restricted to the West Gulf Coastal Plain. Most of these species 
were described from specimens collected in east Texas and until 
recent years most were thought to be endemic to Texas. 

Recently, many have been found in a few specialized localities in 
Louisiana, southeastern Oklahoma, and/or southern Arkansas. One 
rare species collected only twice in Arkansas is Carex hyalina, 
otherwise known only from four collections in eastern Texas and 
one in southeastern Oklahoma. This species of the section Ovales 
is in need of field study to determine its status, relationships, and 
overall distribution. Other facultative wetland species restricted to 
the same region include Crataegus brachyacantha Sarg. & Engelm., 
Rudbeckia maxima Nutt., and Amorpha paniculata Torr. & Gray. 

Most species reported here having a restricted West Gulf 
Coastal Plain pattern are found on deep sandy soils, primarily in 
east Texas, and are reported as new to Arkansas from Miller 
County. A large number of such endemics exist, and many have 
been found in adjacent Louisiana, Oklahoma, and Arkansas, with a 
few extending into the southern Great Plains in Oklahoma and 
Texas. Among these are Echinacea sanguinea (McGregor 1968), 
Oenothera heterophylla ssp. heterophylla (Figure 19), 
Tetragonotheca ludoviciana (Turner and Dawson 1980), Dalea 
villosa var. grisea (Wemple 1970), Berlandiera x betonicifolia 
(Pinkava 1967), Pediomelum subulatum (Ockendon 1965), and Prunus 
gracilis (Figure 20). Other species with this pattern are here 
reported for Miller County as well as other counties in Arkansas. 
These include Astragalus distortus var. engelmannii, A. leptocarpus, 
A. soxmaniorum (Barneby 1964), and Delphinium carolinianum ssp. 
vimineum (Warnock 1981). Other rare or restricted species in 
Arkansas with this pattern found presently only in Miller County 
include Dalea phleoides var. microphylla, Aphanostephus 
skirrhobasis (DC.) Trel., and Matelea cynanchoides. Several 
species with this pattern are found in Nevada and Ouachita 
counties, as well as Miller County, including Tradescantia 
reverchonii, Eriogonum multiflorum, Hymenopappus artemisiifolius, 
and Streptanthus hyacinthoides. Others are found in Nevada 
and/or Ouachita counties but not in Miller County, and include 
Coreopsis basalis (A. Dietr.) Blake, Aristida desmantha Trin. & 
Rupr., Penstemon murrayanus Hook., Stylisma pickeringii (Torr. ex 
M. A. Curtis) Gray var. pattersonii (Fern. & Schub.) Myint, and 
Polanisia erosa (Nutt.) Iitis. Selaginella arenicola ssp. riddellii, 
like Astragalus distortus var. engelmannii, is found in xeric 
habitats in several regions of Arkansas. 

A species primarily found in the southwestern United States 
with an isolated occurrence in the Mississippi Alluvial Plain of 
Arkansas is Portulaca umbraticola. This species is also found on 


1987 Orzell & Bridges, Flora of Arkansas 135 


granitic cutcrops in Georgia and South Carolina, and an isolated 
station on a sandy hill in southern Misisisippi (Matthews and 
Levins 1985). Otherwise, it is widespread and rather weedy in 
Texas, Oklahoma, and westward. The Arkansas record is 400 km 
from the nearest locality, and must be considered as possibly 
adventive. 

Several species reported here have ranges centering in the 
southern Great Plains, western and southern parts of the West 
Gulf Coastal Plain, Edwards Plateau, and other areas generally 
south and west of Arkansas. These tend to be found in Arkansas 
in the Cretaceous region of southwestern Arkansas, primarily in 
Little River County. Among these are Dalea compacta var. 
pubescens (Wemple 1970, Barneby 1977), Carex planostachys (Figure 
21), and Convolvulus equitans (Figure 22), all disjunct from north 
central Texas or adjacent south central Oklahoma. Other rare 
species in Arkansas with similar patterns include Eriogonum 
annuum Nutt. and Bouteloua rigidigeta. Some species with the 
same general range occur more continuously through southeastern 
Oklahoma to the Cretaceous region of Arkansas, including 
Delphinium carolinianum ssp. penardii (Warnock 1981). Other rare 
species in Arkansas with similar patterns include Dalea compacta 
var. compacta, Astragalus nuttallianus, Carex microdonta Torr. & 
Hook., Engelmannia pinnatifida, Pyrrhopappus multicaulis DC., 
Lindheimera texana Gray, Ruellia humilis Nutt. var. depauperata 
Tharp & Barkl., Lesquerella gracilis (Hook.) S. Wats., Indigofera 
miniata var. leptosepala, Hedeoma drummondii Benth., Allium 
drummondii Regel, Astragalus crassicarpus Nutt. var. crassicarpua, 
and Eryngium leavenworthii Torr. & Gray. Two species reported 
for Miller County are more commonly found further west. 
Pediomelum digitatum ranges throughout the Great Plains and into 
the West Gulf Coastal Plain, where it occurs in adjacent Louisiana 
and Texas. Pediomelum hypogaeum ssp. scaposum (Ockendon 1965) 
is unusually disjunct from north central Texas to Miller County, 
and with a distinct difference in habitat. 

Two species reported here are Great Plains species which 
reach their eastern range limits in the Ouachita Province of west- 
central Arkansas. One of these, Solidago ulmifolia var. 
microphylla (Taylor and Taylor 1984), is found in dry forests with 
rocky outcrops. Other species with similar patterns include 
Marsikallia caespitosa Nutt., Paronychia virginica Spreng. var. 
scoparia (Small) Cory, and Galium texense Gray. Others, such as 
Sanguisorba annua (Figure 23), are primarily prairie species, and 
reach their eastern limit in the prairies of the western Arkansas 
River valley in Arkansas. Other species with this pattern include 
Castilleja indivisa Engelm., Croton lindheimerianus Scheele, Rosa 
foliolosa Nutt. ex Torr. & Gray, and Sporobolus pyramidatus. The 
farthest western species reported here is Lygodesmia juncea (Tomb 
1980), generally found only east to west-central Oklahoma, central 
Kansas, and on loess hill prairies in northwestern Missouri. The 
Arkansas record is 380 km from the nearest isolated locality, and 
about 470 km from the limit of the continuous range. 


136 PHY TOLG G2 A Vol. 64, No. 2 


ro 100 200 300 400 $00 ga ael Snares J 
SRR ee pee ys antes ageenra de 
Bits Om yeas CCU 
Pane eeeee eT eau serine 


| I hx 1 
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Figure 18. North central part of range of Cladium jamaciense. 
Figure 19. Range of Oenothera heterophylla ssp. heterophylla 
(dots) and ssp. orientalis (triangles). 

Figure 20. Range of Prunus gracilis. 

Figure 21. Northern part of range of Carex planostachys. 


The large number of new records reported in this paper, both 
those based on new discoveries and on reinterpretation or 
relocation of historical collections, indicate that much more work 
is needed in order to document and understand the flora of 
Arkansas and its phytogeographical relationships. It is hoped that 
the records and interpretations presented here will spur further 
work on the flora of Arkansas, particularly more fieldwork in the 


1987 Orzell & Bridges, Flora of Arkansas 137 
aa PP PPS 
mail 
ei 


Saale 


J 


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Ane ARE Oe Oe | 
° 100 200 300 400 500 600 


Figure 22. Eastern part of range of Convolvulus equitans. 
Figure 23. Range of Sanguisorba annua. 


unusual and specialized habitats of the state and among 
taxonomically complex groups. Although, as shown in many 
instances in this paper, the state does have a botanical history on 
which to draw, much of it is inconsistent and difficult to interpret 
correctly. We hope that our presentation of the field, herbarium, 
and literature study that has gone into this paper will be useful in 
understanding the importance of all of these sources in the 
compilation of new records towards a state flora. 


ACKNOWLEDGMENTS 


In a paper drawing from as many diverse sources and fields 
of specialization as this one, it seems appropriate to acknowledge 
those workers who directly or indirectly have influenced the 
approach taken by the authors in floristic exploration and 
documentation. Among past botanists the diligent explorations of 
M. L. Fernald and the phytogeographic interpretations of E. L. 
Braun have been an inspiration and challenge. The viewpoints and 
approaches taken have been directly influenced by the brilliant 
work of contemporary scholars who have transcended modern 
boundaries of specialization, particularly J. A. Steyermark and A. 
E. Radford. Special thanks is given to Jerry Roberts, Albert 
Higgenbottom, and Lance Peacock, astute observers of the 
Arkansas landscape and its flora who first led us to the sites of 
some of these records, and provided additional information on 
their collections and observations. In particular, we are especially 
grateful to Carl Amason of Calion, Union County, Arkansas for 
introducing the senior author to the wonders of the Coastal Plain 


138 PoHOY!TAGIL OrGrl A Vol. 64, No. 2 


flora and his lifetime of observation of the local flora of south 
central Arkansas. Publication of this paper would be impossible 
without this encouragement and generous financial assistance. Our 
specimens of Carex were determined or verified by Anton A. 
Reznicek, who also generously shared his extensive knowledge of 
the genus and helped us sort out the taxonomy and nomenclature 
of Arkansas Carex species. Thanks is due to Robert Kral, Charles 
Sheviak, James Bruce, James Grimes, Rupert Barneby, and Wm. 
Wayt Thomas who determined or verified many of the cited 
collections. Our compilation of distributional records was 
facilitated by the cooperation of curators of several herbaria, 
including E. B. Smith (UARK), Gary Tucker (APCR), Barney 
Lipscomb (SMU), Guy Nesom (TEX), John and Connie Taylor 

(DUR), Elray Nixon (ASTC), Robert Kral (VDB), and R. D. Thomas 
(NLU), who also gave us access to his unpublished compilation of 
Louisiana county records at NLU, LSU, and USL. 


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140 PHYTOLOGIA Vol. 64, No. 2 


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2 


TAXONOMY OF CARPHOCHAETE (ASTERACEAB-EUPATORIEAE) 
B.L. Turner 
Department of Botany, Univ. of Texas, Austin, TX 78713 
ABSTRACT 


A taxonomic treatment of Carphochaete is rendered. 
Seven species are included in the genus: C. bigelovii, 
Cc. durangensis, C. grahamii, C. macrocephala, C. 
pringlei, C. schaffneri and C. wislizeni. These include 
the monotypic genera Crongquistia King, and Revealia King 
& H. Robinson. One new species, C. durangensis, and one 
new varietal combination, C. pringlei var. simulans, is 
proposed; in addition C. gummifera McVaugh is reduced to 
synonymy under D. grahamii. Descriptions, keys, 
distribution maps, and a complete synonymy are provided. 


The genus Carphochaete was first proposed by Gray 
in 1849 with his description of C. wislizeni. Shortly 
thereafter (1852) he added two additional species, C. 
bigelovii and C. grahamii. Greenman, in 1901, added C. 
schaffneri, B.L. Robinson in 1906 added C. simulans and 
McVaugh in 1972 proposed C. gummifera. The most recent 
additions, C. durangensis and C. macrocephala, were first 
proposed by the late Dr. Jerold Grashoff, who was engaged 
with a revisionary study of the group at the time of his 
early death. 


I have accepted seven species in the genus 
including all those proposed by Gray, Greenman, and 
Grashoff. Robinson's C. simulans has been reduced to 
varietal status under C. bigelovii and McVaugh's C. 
gqummifera has been placed in synonymy under C. _grahamii. 


It should be noted that King (1968) excluded C. 
pringlei from the complex, creating the monotypic genus 
Cronquistia, and King and Robinson (1976) subsequently 
described a new monotypic genus Revealia, based upon 
their R. stevioides. This was soon found to be a synonym 
of the earlier Oxylobus macrocephala Paray, which name 
was transferred to Revealia, replacing R. stevioides. In 
my opinion, neither of the two monotypic genera are 
worthy of recognition and I follow Grashoff in reducing 
them here. 


CHROMOSOME COUNTS 
Relatively few chromosome counts are published for 


Carphochaete. Those available to date are listed below. 
The genus would appear to be dibasic with x=1ll or 12; two 


145 


146 PHY TO G06 "1A Vol. 64, No. 2 


of the species C. bigelovii and C. grahamii, possess x=11 
and C. _durangensis has x=12. The latter was reported as 


Cronquistia pringlei by King et al. (1976). 


Taxon Chromosqme count Reference 

C. bigelovii 2n=22 Gaiser (1953) 

C. bigelovii n=1l1 pairs Powell and Powell 
(1978) 

C. bigelovii n=ll pairs Turner (1959) 

C. durangensis* n=12 pairs King et al. (1976) 

C. grahamii 2n=22 Grasshoff et al. 
(1972) 


A base chromosome number of x=1l or 12 would 
suggest a relationship with Stevia which is essentially 
tribasic with x=11,12 and 17. On morphological grounds 


Carphochaete appears closer to those taxa of Stevia 
possessing base numbers of x=l]] or 12. 


GENERIC RELATIONSHIPS 


The species of Carphochaete superficially resemble 
certain species of Stevia (e.g., S. pelophila Blake) as 
noted by Grashoff (1972) in his monumental treatment of 
Stevia for North America. Carphochaete, however, is 
readily distinguished by its style branches, and yet 
other characters of the head and florets. 


Robinson and King (1976) place Carphochaete and 
their monotypic generic segregates, Crongquistia and 
Revealia, next to each other near Metastevia and Stevia, 
which is about where I would place the groups, as would, 
presumably, Grashoff (1975) to judge from his remarks as 
to the relationships of Metastevia. That is, the latter 
genus is closer to Stevia, on morphological grounds, than 
it is to Carphochaete (indeed, on phyletic grounds I 
would include Metastevia within Stevia, as presently 
constituted); but Carphochaete has characters of both 
Stevia and Metastevia and is perhaps ancestral to both. 
Certainly the semipaleate, large heads with numerous 
florets, and bristly pappus of C. durangensis makes that 
species a likely candidate. But these are matters for 
the future; any resolution of the problem will require 
new insights into the groups, especially using 
macromolecular data. 


SPECIES RELATIONSHIPS 
As I view the species they fell into four groups as 


follows: Group I) C. pringlei and C. durangensis; Group 
II) C. wizlizeni, C. grahamii and C. schaffneri; Group 


1987 


character 


species 


Turner, Taxonomy of Carphochaete 


I6 |} 17} 18 


IS 


13 


l2 


lOj Il 


14 
ololololojolo| jo 
jt fofofofofo]i fo 

0 

0 | 

0 

[oO 

oe 

0] 

bd 


i as 

SG ols Bie Sa) 

yi ls a ila 

a Sa i orchestra 

ny oes Bees Bis oad 
i) 

a ie 

Sia 

5 Cc 

wo 

Elo 

n Cc 

(er) 

Ss Nee 

=) 

= a= 


bigelovii 


Metastevia Foe 


pringlei 
wizlizenii 
grahami 
schaffneri 
macrocephala 


Carphochaete and Metastevia. 


Character states among species of 


Table |. 


147 


148 Pohe¥YAtnOnLeO Galva Vol. 64, No. 2 


III C. bigelovii; and Group IV C. macrocephala. The 
relationships among these are shown in Fig. l. 


Construction of the diagram was based upon the following 
assumptions as regards primitive (0) or advanced states 
(1) of the characters concerned: 


Character Primitive Advanced 
State (0) State (1) 
1.Habit suffruticose herb shrub 
2.Stems from woody crown rhizomatous 
3.Leaf arrangement opposite alternate 
throughout above 
4.Leaf venation 3-nervate l-nervate 
5.Involucral bracts(I.B) subequal graduate 
6.1.B. vestiture not glandular glandular 
7.1I1.B. margins not scarious scarious 
8.1.B., number 9 or more 5-9 
9.Receptacle chaffy(in part) not chaffy 
10.Florets/head T=15 3=5 
11.Corolla pubescent absent present 
without 
12.Corolla pubescent absent present 
within 
13.Style node abrupt not so yes 
14.Achenes 8-9 ribbed 4-5 ribbed 
15.Pappus with mid-rib w/o mid-rib 
16.Pappus bristle no. 4-5 8-16 
17.Anthers bifid not so yes 
18.Heads single and not so yes 
sessile 


Character states for the various species of 
Carphochaete are presented in Table 1 and these were used 
to construct the cladistic analysis shown in Fig. l. In 
this I have used a hypothetical outgroup (HOG), for 
cladistic purposes. Other workers might have proposed 
their own ‘HOG bwt. fo7...me,..£0m. ny analyses eo 
Carphochaete, I like the HOG proposed here. Cladistic 
purists might wish to have an appropriate “real" outgroup 
selected for such analysis, but to what avail? I mean, 
how does one recognize an outgroup where relationships 
are obscured by reticulate divergence, or whatever. 
Nevertheless, to this end I have provided such an 
analysis using the genus Metastevia as an outgroup, since 
Grashoff (1975) felt that the latter genus "developed 
from S. elatior-like ancestors during the ecariy 
colonization of Mexico by members of this group". Stevia 
elatior belongs to a group of species that share many 
characters with Carphochaete, thus the selection of 
Metastevia as an appropriate outgroup is not too far- 
fetched. Besides, Metastevia is monatypic, making the 


1987 Turner, Taxonomy of Carphochaete 149 


Fig; J.) /a=-d. Cladograms showing relationships among 
Carphochaete species. a. Consensus tree 1; b. Consensus 
Second-level consensus tree; d. Subjective 


tree 2; ©. 
(derived) cladogram-anc=ancestor; met=Metastevia; big=C. 


bigelovii; dur=C. durangensis; gra=C. grahamii; mac=C. 
macrocephala; pri=C. pringlei var pringlei; sim=c.p. var. 


Simulans; sch=C. schaffneri; wis=C. wislizeni. 
Additional explanation in text. 


wis sch mac 


big 


— synapomorphy 
= parallelism 
V reversal 


150 PUHyY TeOsksO Griva Vol. 64, No. 2 


calculations relatively simple. Results of the 
calculation are shown in Fig.i. 


The hypothesis of phylogeny presented here is based 
on a cladistic analysis using Wagner parsimony. The 
computer program PAUP written by David Swofford was used 
to analyze the data. Two outgroups were used to polarize 
the character states: (1) the monotypic genus 
Metastevia, and (2) a hypothetical ancestor (HOG) that 
best represents my ideas regarding primitiveness in 


Carphochaete. 


Using Metastevia as the outgroup, six trees of 3l 
steps each and a consistency index of 0.581 were found. 
They are represented in Figure laby a strict consensus 
tree that summarizes the topologically stable areas of 
the six trees. Using the HOG, seven trees of 27 steps 
each and a consistency index of 0.667 were found. They 


are represented in Figureib. by a second strict consensus 
tree. In these two analyses, two lineages are clearly 


represented: C. pringlei-simulans-durangense, defined by 
alternate leaves and C.. ‘schaffneri-wislizeni-grahamii- 
macrocephala-bigelovii, “defined by characters 4, 5, 8, 9, 
10 and 13. If a consensus of the trees in Fig. la and 
Fig. lb is constructed, the resulting tree (Fig. lc) 
provides only this minimal level of resolution. Finally, 
I offer an admittedly somewhat subjective cladogram (Fig. 
ld) constructed from the two consensus trees that 
displaces unresolved topology from one tree with resolved 
portions from the other and also that best fits my 
personal view of evolution in Carphochaete. This tree, 
however, exactly matches one of the seven trees obtained 
using the HOG as outgroup. Character state changes have 
been appended to the tree in Fig. ld, using HOG to 
establish the polarities. 


It should be noted that the cladistic branch 
bearing C. durangensis C. pringlei and its variety 
simulans (Fig. ld) is at variance with the nomenclature 
adopted here. That is, the var. simulans on cladistic 
grounds, using the data adopted here would more logically 
be tyeated as a variety of C. durangensis. However, I 
have not let my putative phylogeny affect the 
nomenclature in this instance, for if so treated the 
correct specific name would be C. simulans,with C. 
durangensis becoming a variety of the latter. Of course 
both of these could be treated at the specific level, but 
lacking new experimental data I have maintained the 
existing nomenclature so far as possible. 


ACKNOWLEDGEMENTS 


1987 Turner, Taxonomy of Carphochaete V5 


This study is based upon the examination of 
approximately 465 specimens, as follows: ARI2Z(79), 
ASU(37.) 4° CAS=DS (22) 7+ -F:G24)'7,,GH(76), LL(30), ,8O( 32)’, 
MICH(38), MSC(20), TEX(60), UC(44). I am grateful to the 
Directors concerned for these loans. Guy Nesom provided 
the Latin diagnosis and assisted with the cladistic 
analyses. 


CHARPHOCHAETE A. Gray 


Perennial suffruticose herbs or shrubs to 3 m high. 
Stems arising from short rhizomes or ligneous root 
crowns. Leaves opposite throughout or markedly 
alternate, sessile or nearly so, l-nerved or with 3 
parallel nerves, markedly glandular-punctate. Heads 
large, cylindrical or turbinate, borne in l1-numerous 
terminal cymes. Involucres 2-4 seriate, graduate to 
eximbricate, persistent. Receptacle convex or plane, 
epaleate or rarely partially paleate. Corollas tubular, 
white, pink or lavender, the throat cylindrical, glabrous 
or hirtellous without, pubescent or glabrous within; the 
lobes linear, of differing lengths. Anthers with well- 
developed appendages, these often with a central rib, or 
bifid. Style branches filiform with smooth, linear, 
narrowly-oblanceolate, appendages, the shaft with a 
gradually or abruptly swollen basal node. Achenes linear 
to narrowly obpyramidal, 4-5, or more often, 8-9 ribbed, 
the pappus scales ribless or ribbed, or both. Base 
chromosome number, x=ll or 12. 

Type species, Carphochaete wislizeni A.Gray. 


A genus of seven species, all of which are confined 
to Mexico, mostly from Guerrero northward, where they 
usually occur in pine-oak woodlands from 1500-3800 m. 


Key to Species 


is Leaves predominantly alternate along the upper 
stems; blades at least faintly 3-nerved. 


2s Leaves 5-8 cm long; blades strongly 3-nerved------- 
SS a C. durangensis 
2. Leaves 1-3 cm long; blades faintly 3-nerved-------- 
a SE SS SS Cs. pringle 
ihs Leaves predominantly opposite along the upper 


stems; blades with a single mid-vein. 


3. Sprawling shrubs to 3 m high; pappus a laciniate 
clown, Cue. ce. oe C. macrocephala 


152 PAH PY) TAO 8Ih'0). Gel A Vol. 64, No. 2 


ce Suffruticose erect herbs; pappus of prominent 
scales; Central Plateau of Mexico. 
4. Heads 3-4 cm high, sessile or nearly so; Chihuahua, 
Coahuila and adjacent U.S.A.---------- C. bigelovii 


4. Heads 2-3 cm high, pedunculate, arranged in a 
terminal capitulescence. 


aye Involucral bracts densely covered with minute, 
stipitate-glandular, trichomes; San Luis 
a a C. shaffneri 


Sig Involucral bracts glabrous or merely ciliate, 
sometimes gummy or viscid 


6. Pappus bristles _§8-16----------— C. grahamii 
6. Pappus bristles 4-6----------- C. wislizeni 


CARPHOCHAETE BIGELOVII A. Gray, Smithson. Contr. Knowl. 
3.989 21852), TYPE:U.S.A. (MEXICO?): “On the boundary 
between Mexico and New Mexico", Mimbres" (on type sheet), 
w/o date, I. J. Bigelow s.n. (holotype GH!) 


Perennial suffruticose herbs or subshrubs 0.3-1.0 m 
high. Stems stiffly erect, densely short-puberulous to 
glabrate, tan or reddish. Leaves opposite throughout, 
mostly in axillary fascicles, 1-3 cm long, 3-10 mm wide, 
sessile or nearly so, l-nerved, glabrous, linear- 
oblanceolate to somewhat elliptic (very rarely 3-nervate 
and somewhat denticulate). Heads large, single, terminal 
or axillary, mostly 3.0-3.5 cm high, the capitulescence a 
spike-like or locse corymbose panicle. Involucres 4-5 
seriate, graduate, mostly 15-20 mm long, the bracts 10- 
14, linear oblanceolate, puberulent, glandular-punctate, 
the apices acute or apiculate. Florets mostly 4 per 
head; corollas white or pinkish-white, 15-18 mm long, 
glandular or hirtellous without, pubescent within along 
the lower part. Achenes 11-12 mm long, 8-9 ribbed, 
hispidulous; pappus of 10-12 linear-lanceolate scales 12- 
15 mm long, the mid-rib extending into barbellate 
bristles 2-4 mm long, the outer pappus of 1-4, ribless, 
linear scales, 1-3 mm long, or absent; chromosome number, 
2n=22. 


DISTRIBUTION (Fig. 2): Southern Arizona, New Mexico and 
trans-Pecos Texas in the U.S.A. and adjacent Son, Chi and 
Coa in Mexico; pine-oak-juniper woodlands in mostly 
igneous soils from 1700-2500 M; Sep-Jul, depending open 
rains. 


REPRESENTATIVE SPECIMENTS: U.S.A. ARIZONA: Cochise Co.: 
Chiracahua Natl. Monument, Bonita Canyon, below Shake 
Springs, Cupressus forest, 5760 ft, 18 Apr 1975, Reeves 


1987 Turner, Taxonomy of Carphochaete 153 


R2519 (ARIZ, ASU). Gila Co.: Tonto Forest, Parker creek, 
5750 ft; 9 Apr 1935, Johnson 104 (ASU). Graham Co.: 
Pinaleno Mts., Frye Mesa Reservoir, 5000 ft. 17 Apr 1985, 
Johnson 11456 (ASU). Pima Co.: Santa Catalina Mts, Sabino 
Canyon, “The Horse", 9 Apr 1905, Thornber & Terrell s.n. 
(ASU, TEX). Pinal Co.: Superstition Mts., Top of ridge, §S 
side, 4000 ft 26 Mar 1932, Gillespie 5468 (GH). Santa 
Cruz Co.: Cobre Ridge, 10 mi SSE of Arivaca, 4200 ft, 17 
Apr 1973, Holmgren & Holmgren 6834 (ASU). Yavapa: Co.: 
Happy Valley, 13 Mar 1966, Hesselberg s.n. (ARIZ). NEW 
MEXICO. Dona Ana Co.: Organ Mts, Apr 1852, Wright (GH, 
TEX). Grant Co.: Emory Point, 6600 ft, 1905, Blumer 189 
(GH). Greenlee Co.: Clifton, Apr 1987, Traphagen s.n. 
(GH). Hidalgo Co.: Animas Mts., 23 mi S of Animas, 6500 
ft, 2 May 1976, Hess & Stickney 3760 (ARIZ). TEXAS. 
Brewster Co.: Chisos Mts., above Lost Mine Peak Trail, 2 
Apr 1959, Correll 20692 (LL). Jeff Davis Co.: upper 
canyon of Limpia Creek, 10 Jun 1926, Palmer 30669 
(A, TEX); Presidio Co.: NE slope of Chinati Peak, Horse 
Creek Canyon, ca 6500 ft, 21 Jun 1942, Hinckley 2516 
(ARIZ, GH). 


MEXICO. CHIHUAHUA: Sierra Charuco, 17-25 Apr 1948, 
Gentry 8016 (ARIZ, MICH, UC); Cascada de Basaseachic, ca 
2150 m, 27 Apr 1986, Nesom 5457 (TEX). COAHUILA: Sierra 
de Hechiceros, 17-19 Sep 1940, Johnston & Muller 1311 
(GH,LL,MICH,MSC); Sierra Maderos del Carmen, 2100 m, 1 
Apr 1974, Wendt et al. 121 (LL); Serranias del Burro, 12 
Apr 1976, Riskind & Patterson 1977 (TEX). SONORA: 5 mi E 
of Esqueda, 27 Mar 1970, McGill & Pinkava 6429 (ASU); 17 
mi SE of Magdalena, Palm Canyon, 10 Mar 1979, Steadman & 
Schmidt s.n. (ARIZ). 


CARPHOCHAETE DURANGENSIS Grasshoff ex B. Turner, sp. nov. 


C. pringlei var. simulans simile sed foliis multum 
grandioribus laminis valde 3-nervatis et capitulis 
grandioribus flosculis numerosioribus. 


Perennial, somewhat suffrutiose, herbs 30-60 cm high. 
Stems glandular-pubescent or puberulous, reddish, stiffly 
erect, 1-8 arising from a short, fibrous-rooted, 
rhizomatous caudex. Leaves opposite for the first 
several nodes then markedly alternate thereafter, 4-8 cm 
long, 7-14 mm wide, gradually reduced upwards, sessile or 
nearly so, the blades linear-elliptic, strongly 3- 
nervate from the base, with fine reticulate-nerves 
between the major veins, glabrous or nearly so. Heads 2- 
15, bright pink-lavender to purple, arranged in stiffly 
erect, terminal, flat-topped cymes, the ultimate 
peduncles glandular-pubescent, mostly 1-3 cm long. 
Involucres turbinate, eximbricate or nearly so, 10-13 mm 


154 : PR VT) Gv 0 G. BA Vol. 64, No. 2 


O p. var pringlei 


<> p. var simulans 


A durangense 


@ bigelovii 


Fig.2 Distribution of Carphochaete spp. 


1987 Turner, Taxonomy of Carphochaete 155 


high; bracts densely glandular-pubsecent, broadly 
lanceolate, 3-5 nervate, the apices acute. Receptacle 
somewhat convex, epaleate or with well-developed chaff. 
Florets 15-20 per head; corollas tubular, 6-8 mm long, 
pubescent without, glabrous within, the lobes 2-3 mm 
long. Achenes 4.0-4.5 mm long, 8-9 ribbed, the faces 4 
or 5, pubescent; pappus of 3-5 lanceolate scales, 2-5 mm 
long, with their mid-ribs extending into short bristles, 
these alternating with 3-5 ribless scales; chromosome 
number, 2n=22. 


DISTRIBU EON (Craigie t2y)i: Known only from southern 
Durango in pine-oak woodlands, 2400-2700 m; Aug-Sep. 


TYPE: MEXICO. DURANGO: ca 30 mi W of Durango, ca 8500 ft, 
"In rocky, rhyolitic soil in oak-pine woodland". 28 Sep 
1962, A. Cronguist 9539 (holotype TEX; isotypes GH!, 
MICH!, NY). 


ADDITIONAL SPECIMENS EXAMINED: DURANGO: 50 km W of 
Durango along highway 40 (23°52'N x 105°00W), area with 
much exposed rock, 2500m, 12 Sep 1984, Barrie 1003 
(MEXU;TEX); Parque El Teivan, 58 km al ESE de Durango, 4 
Sep 1984, Casillas et al. 6 (TEX); Jarocho, railroad W of 


Durango, 2400-2500 m, 27 Aug 1934, Pennell 18242 (GH). 


This taxon superficially looks like a robust form of 
C. bigelovii var simulans but differs by a number of 
characters, the most notable being the large, strongly 
paralleli-nerved, leaves. King (1968) included the type 
and only collection known to him, within his concept of 
Cronquistia pringlei, but subsequent collections reveal 
the taxon to be fairly uniform and common to the west and 
south of Durango City. It does not ,intergrade with’ Ga 
pringlei and is remarkably distinct, as surmized by the 
late J. Grashoff, who first annointed the species and 
designated its type. 


Some of the specimens (Casillas et al. 6, TEX) have 
well-developed chaff on the receptacles; occasionally, 
chaff also occurs among the peripheral florets of C. 


pringlei, contrary to the observations of King (1968). 


CARPHOCHAETE GRAHAMII A. Gray, Smithson. Contr. Knowl. 
32895591852)... TYPE: MEXICO. MEXICO SSTATE:.or “MICHOACAN: 
according to McVaugh (1984), who examined type material, 
the type was probably collected about the villages of 
Tlalpujahua and Angangueo, in NW Mexico State or adjacent 
Michoacan, 1830, G. J. Graham 81 (holotype kK). 


Carphochaete gummifera McVaugh, Contr. Univ. Michigan 
Herb. 9:385.1972. TYPE: MEXICO. ZACATECAS: between Jalapa 


156 PSHYaTROrL0? Galva Vol. 64, No. 2 


and Tlaltenango, 2300-2500 m, 22 Dec 1970, McVaugh 25617 
(MICH! ).= 


Perennial suffruticose herb or shrublets, 30-70 cm 
high. Stems sparsely puberulent to glabrate, reddish. 
Leaves opposite throughout, linear-oblanceolate to 
oblanceolate or somewhat spatulate, 2-4 cm long, 3-7 mm 
wide, sessile, glabrous to sparsely pubescent on both 
surfaces, markedly glandular-punctate, l-nerved, the 
apices usually obtuse or rounded, but rarely acute. 
Heads 1-3 at the apices of stems, the ultimate peduncles 
5-20 mm long, pubescent to glabrate. Involucres 14-17 mm 
high, 3-4 seriate, puberulent, ciliate; bracts 8-10, the 
apices usually rounded and apiculate but sometimes 
gradually narrowed and acute. Style shaft with basal 
node. Florets usually 4 to a head; corollas lavender- 
pink, 15-20 mm long, glabrous within and without, the 
lobes 5-6 mm long. Achenes ca 1 cm long, with 8-9 ribs, 
sparsely hispid to glandular-hirtellous; pappus of 8-10 
linear-lanceolate scales 9-14 mm long, the apical 
barbellate extensions mostly 3-4 mm long, an outer series 
of short ribless scales may be present or absent. 
Chromosome number, n=1l pairs (Grasshoff 533, TEX). 


DISTRIBUTION (Fig. 3): Southern Durango to Mexico 
State, mostly along the western Central Plateau in pine- 
oak woodlands, 1400-2700 m; Sep-Nov. 


REPRESENTATIVE SPECIMENS: MEXICO. AGUASCALIENTES: 
ca 10 mi SE Calvillo, 2000-2300 m, 4 Nov 1959, McVaugh & 
Koelz 179 (LL,MICH). DURANGO: Mcpio. El Mezquital, 22 km 
NE Los Charcos, 2750 m, 1 Nov 1982, Gonzales & Rzedowski 
2347 (CAS, TEX); 74 km WNW Huejuquilla El Alto, 2720 m, 
22 Oct 1983, Breedlove 59187 (CAS, TEX); Mepie. Suen, 
San Juan de Michis, 21 Nov 1985, Alvarado 608 (TEX). 
GUANAJUATO: 30 km WSW Dolores Hidalgo, 2300 m, 29 Dec 
1967, Rzedowski 25935 (DS,LL,MICH,MSC). JALISCO: summit 
of mountains above Etzatlan, 27 Oct 1903, Pringle 8772. 
(F,GH,LL,MO,MSC,UC). MEXICO: Bluffs, Flor de Maria, 18 
Oct 1890, Pringle 3315 MORELIA: Lake Maria, 9 Oct 1911, 
Arsene s.n. (CAS); ZACATECAS: Sierra de los Huicholes, 5 
mi N of Tepetates, 2400-2600m, 13 Jan 1975, McVaugh 25772 
(MICH). 


Collections from Durango generally have broader 
more oblanceolate blades which are more puberulent than 
is typical, but otherwise differ but little from material 
to the south. 


I take C. gummifera to be a somewhat, narrow- 
leaved, gummy, form of C. grahamii. In nearly all other 
characters it is like the latter and falls within the 


1987 Turner, Taxonomy of Carphochaete 157 
geographic range of that species. 


CARPHOCHAETE MACROCEPHALA (Paray) Grashoff ex B. Turner & 
Kerr, Pl. Syst. Evol. 151:86.1985. 


Oxylobus macrocephalus Paray, Bol. Soc. Bot. Mex. 
22:1.1958. TYPE: MEXICO. GUERRERO: Cerro Teotepec, NE of 


Chilpancingo, 3500-3600m, 27 Dec 1946, Paray 973 (MEXU; 
photoholotype TEX!) 


Revealia stevioides King & H. Rob., Phytologia 
33:277.1976. TYPE: MEXICO. GUERRERO: ca 60.5 mi NE of 
Atoyac and 67.5 mi NE of Puerto del Gallo, 10,500 ft, 19 
Oct 1975, Reveal et al. 4319 (holotype US). 


Revealia macrocephala (Paray) King & H. Rob., 
Phytologia 23:376.1976. 


Sprawling semi-succulent shrubs to 3 m high. Stems 
puberulent ot glabrate, reddish, the nodes numerous and 
mostly shorter than the leaves. Leaves opposite 
throughout, 1-2 cm long, 2-5 mm wide, sessile, l-nerved, 
glabrous, oblanceolate, entire or with a few minute 
serrations. Heads lavender or purple, single or 2-5 in 
terminal cymes, the ultimate peduncles mostly 2-8 mm 
long. Involucre campanulate, 2-3 seriate, subimbricate; 
bracts elliptic with scarious margins, the apices 
rounded. Receptacles somewhat convex, glabrous, 
epaleate. Florets 10-14 per head; corollas 13-15 mm 
long, lavender, tubular, pubescent without and within, 
the lobes 3-6 mm long. Achenes 6-8 mm long, with 4-5 
sides, the faces occasionally with weaker ribs, glabrous 
or faintly pubescent above; pappus a lacerate crown ca l 
mm high. 


DISTRIBUTION (Fig. 4): Known only from Guerrero in 
the region of Cerro Teotepec in pine-fir forests from 
2900-3500 m; Sep-Dec (Apr). 


ADDITIONAL SPECIMENS EXAMINED: MEXICO. GUERRERO: 
Summit of Teotepec, 3100 M, 12 Nov 1973 Breedlove 36075 
(CAS); Cerro Teotepec, ca 40 mi N Coyuca de Benitez, 
Feddema 2931 (CAS,MICH,TEX); 19.5 km al NE de Puerto del 
Gallo, 23 Nov 1983, Martinez & Barrie 5659 (TEX); Cerro 
Teotepec, 3300 m, 11 Apr 1963, Rzedowski 16494 (F, MICH, 
TEX); Cerro Teotepec, 3350m, 5 Dec 1963, Rzedowski 18156 
(DS,LL,MICH,TEX); ca 8 km NE de Puerto del Gallo, 7 Sep 
1983, Villasenor Rios 558 (TEX). 


King and Robinson (1976) thought that this species 
diverged (as Revealia!) "from between Carphochaete and 
Cronquistia [=C. pringlei]..." They contend that the 


158 Pub ¥) TeOnknO0 Galvé Vol. 64, No. 2 


most important difference between these two taxa is that 
of hairs-on the inner surface of the corolla in Revealia. 
Actually the inner surface of the corolla of Carphochaete 
bigelovii is pubescent like Revealia and I can find 
little merit in the recognition of their monotypic 
proposal, nor did Grashoff, to judge from his 
annotations. 


Nevertheless, the species is perhaps the most 
distinct member of Carphochaete, possessing a well- 
defined, semi-succulent, shrubby, habit and 4-5 ribbed 
achenes, characters which suggest a remote position 
within the genus. 


CARPHOCHAETE PRINGLEI (S. Wats) Grashoff ex B. Turner, 
comb. nov. Based upon Stevia pringlei S. Wats., Proc. 
Amer. Acad. Arts 23:276.1888. 


Perennial suffructicose herbs 30-70 cm high. Stems 
purplish, hirtellous to puberulous, but soon glabrate, 
arising from a ligneous root-stock. Leaves opposite for 
the first several nodes but thereafter markedly 
alternate, mostly 2-4 cm long, 2-4 mm wide, gradually 
reduced upwards, 3-nerved, linear-lanceolate and often 
somewhat falcate, the apices acute. Heads lavender-pink, 
turbinate, borne in 1-10, rather flat-topped, terminal 
cymes, the ultimate peduncles mostly 1-4 cm long. 
Involucres 2-3 seriate, subimbricate; bracts 14-16, 
lanceolate, 7-10 mm long, puberulent to glabrate, the 
apices acute. Florets 3-9 per head; corollas tubular, 6- 
7 mm long, glabrous or rarely pubescent without, glabrous 
within, the lobes 2-3 mm long. Achenes 8-9 ribbed, 4-5 
sided, densely hispidulous, 4-5 mm long; pappus of 3-5, 
awned, scales alternating with 4-5 short awnless scales, 
or of 10 awnless scales 1-2 mm long, these often united 
into a crown. ‘ 


Two varieties are recognized: 


Involucral bracts and corollas densely pubescent 
with glandular trichomes; Chi and Dur (Fig. 73)-———=-——— 
i ee Se as a ibs oe a re var. simulans 


Involucral bracts and corollas without glandular 
trichomes. 63557-5-4 en ee aS ee var. pringlei 


C. PRINGLEI (S. Wats) Grashoff ex. B. Turner var. 
PRINGLEI 


Stevia pringlei S. Wats., Proc. Amer. Acad. Arts 
23:276.1888. TYPE: MEXICO. CHIHUAHUA: foothills of the 
Sierra Madre, Sep 1887, Pringle 3101 (holotype GH!; 


1987 Turner, Taxonomy of Carphochaete 159 


isotypes F!,NY,UC!,US!). 
Cronguistia pringlei (S. Wats) R.M. King, Brittonia 
20:12.1968. 


DISTRIBUTION (Fig. 3). Sierra Madre Occidental of 
Chihuahua and possibly adjacent Sonora, in pine-oak 
woodlands from 2000-2500m; Aug-Oct. 


REPRESENTATIVE SPECIMENS: MEXICO. CHIHUAHUA: SW of 
Tomochi, ca 2100 m, 25 Sep 1980, Cronquist 11718 (CAS, F, 
GH, MICH, MO, TEX); 10 mi SE Madera, 22 Sep 1939, 
Muller 3414 (GH,MICH,TEX,UC). 


A large number of additional specimens are cited by 
King (1968) all of which belong to this variety except 
for the two collections from Durango which serve as the 
types of the following variety, and Carphochaete 
durangensis, described above. 


C. PRINGLEI var. SIMULANS (B.L. Rob.) B. Turner, comb. 
nov. 


Stevia simulans B.L. Rob., Proc. Amer. Acad. Arts 
42:34. 1906. TYPE: MEXICO. DURANGO: on Mesa de Sandia, 
3050m, 14 Oct 1905, C. G. Pringle 10144 (holotype GH!; 
isotypes F!, NY!,UC!,US!). 


This taxon can be distinguished by its copious 
glandular-trichomes on the upper stems, involucral 
bracts, and usually the corollas; the latter, if not 
pubescent, will take on a viscid or gummy sheen. The 
var. pringlei is usually without glandular trichomes, or 
these are relatively few and confined to the peduncles. 
I agree with King (1968) that the pappus characters 
emphasized by Robinson in his recognition of Stevia 
Simulans are not valid, but the glandularity appears to 
hold for populations in southern-most Chihuahua and 
adjacent Durango. 


ADDITIONAL SPECIMENS EXAMINED: MEXICO. CHIHUAHUA: 
17.6 km NNE of El Vergel, open woods of pine-oak- 
manzanita, 2450 m, 24 Aug 1983, Nesom 4912 (TEX); 20 km 
WNW of Santiago Papasquiaro (25° 04' N x 105°47'W), mixed 
pine, fir and oak woodland, 2800 m, 25 Aug 1983, Diaz 660 


[Worthington 11406] (TEX). 


CARPHOCHAETE SCHAFFNERI Greenm., Proc. Amer. Acad. Arts 
40:34,1904. TYPE: MEXICO. SAN LUIS POTOSI: Sierra de San 
Miguelito, valley of San Luis Potosi, Sep 1986, J.G. 
Schaffner 241 (lectotype GH!, selected by King and 
Robinson, by annotation, 1984; isolectotype F!, UC!). 


160 P Hy XD. OL, O»G) I A Vol. 64, No. 2 


Suffruticose erect, rhizomatous, perennials, 25-45 
cm high. Stems minutely glandular-pubescent to glabrate, 
reddish. Leaves opposite throughout, sessile, linear- 
lanceolate, 2-4 cm long, 1-3 mm wide, glabrous, l-nerved, 
markedly glandular-punctate, the apices acute. Heads l, 
or rarely 2, on terminal peduncles 5-20 mm long, the 
whole arranged in an open, 3-15-headed, capitulescence 
with ascending branches. Involucres 10-15 mm high, 2-3 
seriate; bracts 5-7, gradually tapering to an acute apex, 
or abruptly obtuse and apiculate, densely short 
glandular-hirtellous or merely glandular-punctate, not at 
all ciliate. Florets mostly 4 per head; corollas pinkish 
to purplish, 15-18 mm long, glabrous without, very 
sparsely pubescent within near orfice, the lobes 3-4 mm 
long. Achenes with 8-9 ribs, ca 1 cm long, minutely 
glandular-hirtellous; pappus dimorphic, an inner series 
of 6-8 linear-lanceolate, l-ribbed scales, 14-16 mm long, 
the mid-rib extending into well-defined awns, 6-8 mm 
long, the outer series of 2-6, short, ribless scales, 
oS) JOM Lond. 


DISTRIBUTION (Fig. 3): Mountainous regions about 
San Luis Potosi in oak woodlands from 2300-2500 m; Oct- 
Jan. 


ADDITIONAL SPECIMENS EXAMINED: MEXICO. SAN LUIS 
POTOSI: region of San Luis Potosi, 1850-2465 m, 1878-79, 
Parry & Palmer 329, (GH,MO); Sierra de San Miguelito, ca 
Cueva del Mezquite, "chaparral de encino", 2300 m, 9 Nov 
1954, Rzedowski 5456 (MICH,MSC); Sierra de San Miguelito, 
"parte superior de la Canada de San Antonio," 2350 m, 5 
Jan 1955, Rzedowski 5671 (MSC). 


The taxon is closely related to C. grahamii but can 
be distinguished by its glandular-hirtellous involucral 
bracts and a few other minor characters. Ultimately it 
may be reduced to varietal rank under that species. 


Collections by Parry & Palmer 329, Cite@ 2h the 
protologue by Greenman, in part at least, are apparantly 
mixed, for sheets at F and MO are clearly C. grahamii, 
possessing the ciliate eglandular involucral bracts and 
pubescent achenes of the latter. 


CARPHOCHAETE WISLIZENI A. Gray, Mem. Amer. Acad. Arts 
4:65.1849. TYPE: MEXICO. CHIHUAHUA: mountains W of 
Chihuahua, ca Cosiquiriachi, 19 Sep 1846, Wislizenus 175 
(holotype MO!; fragment GH!) 


Perennial, basally suffruticose, herbs 20-40 cm 
high. Stems glabrous, or nearly so, reddish, arising 
from slender rhizomes, forming small colonies. Leaves 


1987 Turner, Taxonomy of Carphochaete 161 


opposite throughout, 2-5 cm long, 1-2 mm wide, sessile, 
linear, glabrous, l-nerved, markedly glandular-punctate, 
the apices acute. Heads 1-5 in rather congested terminal 
corymbs, often numerous-headed, with lateral branches and 
associated stems producing a flat-topped capitulescence. 
Involucre 10-12 mm high, 2-3 seriate; bracts 6-8, linear- 
lanceolate, reddish, ciliate or nearly glabrous, 
gradually, or rarely abruptly, tapered into an acute 
apex. Florets usually 4 per head; corollas pinkish- 
purple to lavender, 13-15 mm long, glabrous within and 
without, the lobes ca 4 mm long. Achenes 8-9 mm long, 8- 
9 ribbed, hispidulous; pappus dimorphic, an inner series 
of 4 or 5, linear-lanceolate, scales, 11-12 mm long, the 
mid-ribs extending into bristles 2-4 mm long, the inner 
series of 4 or 5 alternating ribless scales, 1.0-1.5 mm 
long. 


DISTRIBUTION (Fig. 3): Chihuahua, Durango and 
Zacatecas, pine-oak woodland in mostly rocky igneous 
soils, 2000-2500 m; Aug-Nov. 


REPRESENTATIVE SPECIMENS: MEXICO. CHIHUAHUA: 
Mountains near Chihuahua, 16 Oct 1886, Pringle 765 (ARIZ, 
PGi wai, MCh, 2 MO, MSG, TEx, Were Site one 7 
Basaseachic, ca 2000 m, 4 Oct 1982, Tenorio eye y AL 
(TEX). DURANGO: ca 50 mi W of Durango, ca 8000 re r soe 
1962, Cronquist 9579 (GH, MICH,MO,TEX); 49 mi W of 
Parral, ca 8400 ft, 13 Sep 1972, Reveal & Hess 3058 (GH, 
MO, TEX, UC). ZACATECAS: ca Sombrerete, ca 2400 m, 26 Sep 
1948, Gentry 4876 (ARIZ,GH,MICH,UC). 


LITERATURE CITED 


Gaiser, L. 1953. Chromosome studies in Kuhniinae 
(Eupatorieae). I. Brickellia. Rhodora 55:253-267. 


Grashoff, J. 1972. A systematic study of the North and 
Central American species of Stevia. Doctoral Thesis, 
The Univ. of Texas, Austin. 


- 1975. Metastevia (Compositae: Eupatorieae): 
a new genus from Mexico. Brittonia 69:73. 


King, R. 1968. Studies in the Eupatorieae (Compositae) 
VI. Brittonia 20:11-12. 1968. 


King, R. et al. 1976. Chromosome numbers in Compositae. 
XII. Eupatorieae. Ann. Missouri Bot. Gard. 63:862-888. 


King, R. and H. Robinson. 1976. A new genus Revealia 
from Mexico. Phytologia 33:270-280. 


162 PHY T0.L OG 2 A Vol. 64, No. 2 


Powell, A. and S. Powell. 1978. Chromosome numbers in 
Asteraceae. Madrono 25:160-169. 


Turner, B. 1959. Meiotic chromosomes for 12 species of 
Texas Compositae. Brittonia Phe L79=S177 3 


@ wislizeni 
O grahamii 
* schaffneri 


4 macrocephala 


Fig. 3. Distribution of Carphochaete spp. 


NOMENCLATURA PLANTARUM AMERICANARUM VI. GRAMINEAE 
A. Lourteig and Th. Soderstrom 


This note should had been published in collaboration with Thomas 
Soderstrom (ft 1st. sept. 1987). His untimely death has deprived the 
World of an outstanding agrostologist; the Bamboos, particularly,would 
weep for this lose. 

As soon as I discovered this problem I explained it to Thomas, and 
an exchange of letters followed up. We agreed from the beginning on the 
necessity of a new combination, and we came to the last fine details. 
However, it is my turn to put our paper out with: Arthrostylidium farc- 
tum (Aublet) Soderstrom et Lourteig n.c. 


PLUMIER described in MSS 5: ic. 92 : 

Arundo farcta tenuis,altissimique scandens 
which was put in binomial nomenclature by Aublet who cited Plumier’s 
protologue and the corresponding illustration. At the same time he 
quoted P. Browne, Jamaica 139. 1756. 


Plumier’s description excels in perfection : placing the species 
in Arundinaceae, starting with the roots in details; then, ... "culmi 
seu stipitis arundinacei and farcti seu omnino pleni, exiles, politis- 
simi, flexiles seu fractu contumaces...." &c. It is remarkable to learn 
that the culms are solid. No doubt the leaves so characteristic are al- 
so well interpreted as: ".... nude e nodis subtiles longissimique depen- 
dent funiculi arbores ipsas penitus onustantes ac per quaedam interval- 
lia verticillatim radiatimque emittentes innumera foliola angustissime 
acutissima, virentia duos aut tres aut quatuor pollices longa ex eadem 
vaginula squamata quaterna aut quina simul exorientia"., The icon corres- 
ponds exactly with the description. 


No specimen related to Browne’s protologue has been found (Doctor 
Ch. Jarvis” personnal communication, to whom we are much obliged). 


According with Soderstrom who wrote : “Aublet (1775, vol. 1, pe 
52), under "arundo (farcta) refers to the Plumier manuscript and illus- 
tration. His Latin description, taken from Browne, certainly refers to 
Arthrostylidium capillifolium. This name "farcta"and Latin description 
are correct for this species of bamboo and represent the earliest name" 
the nomenclature is as follows ; 

Arthros 
Arthrostylidium farctum (Aublet) Soderstrom et Lourteig n.c. 


Arundo farcta Aublet, Fl. Guiana Franc. 1: 52. 1775 quoad protol. et 
icon. Plumer., MSS 5: 92 excluding all reference to French Guiana). 
Calamagrostis farcta Gmelin, Systema 2 : 172. 1791 quoad synon. Plum. 
excl, Guiana. 


Arthrostylidium capillifolium Grisebach, Plantae Wrightianae 2. Mém. 


163 


164 Pei Y clPORLVOUG:; Iva Vol. 64, No. 2 


Acad. Sci. Arts n.s. 8: 531. 1862. Type: Cuba, in sylvis densis, fruti- 
ces vel arbores scandens, Wright 738 a. 1860. Isotype P. 

Arundinaria capillifolia (Griseb.) Hackel, Osterr. bot. Zeitschr. 53; 
69. 1903. Urban, Symb. Antil. 8: 51. 1920; Repert. Spec. Nov. Beihefte 
5: 108. 1920. 


Type: MSS 5: 92 by Plumier, based on a plant from Saint Thomas Island, 
found also in Insula Dominicana: La Bande su Sud, Grand Cul de Sac. 


This species is known as the "slender climbing reed". 


After Soderstrom, who was among this plant in the Dominican Repu- 
blic early in 1987, this species is known only from the West Indies, 
not South America. 


eo 


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